This article isThis article by protected copyright. All rights reserved. 10.1111/1365doi: diff to lead copyediting, paginationbeen throughthe andproofreadingtypesetting, process,may which This article acceptedhas been for publication andundergone fullpeer review buthasnot UniversitSapienza 4 3 QLD 40 2 Australia & 1 C.SibaraniMarsya Author details phylogenetic andfunctionaldiversity thesurrogacyMeasuring of potential charismatic Title Handling Editor: Johandu Toit type Article :Research Article MS MARSYACHRISTYANTIID:0000 SIBARANI (Orcid AcceptedGlobal CSIRO &Water, QLD Land Brisbane, 4001, Australia forCentre Biodiversity andConservation The Biodiversity Article

72, Australia 72, Mammal Assessment

erences betweenerences the of this version cite this article and asVersion Record.Please

Conservation - Research Group, à 2664.13360 1

, Moreno Di Journal of Applied Ecology di Roma, RomeI

Science, Program

Marco

The Schoolof Sciences, Biological

on a megadiverse on amegadiverse island - The University ofQueensland, Brisbane,QLDThe University 4072, 00185, Italy , Department, andBiotechnologies, ofBiology 2 , 3 , Carlo , Carlo Science, TheUniversityofQueensland, Brisbane,

Rondinini

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and Kark Salit

- 0003 - 0942 across taxonomic,

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This article isThis article by protected copyright. All rights reserved. Abstract [email protected] Australia School ofBiological SibaraniMarsya Corresponding author

Accepted3. 2. Article1.

charismatic charismatic megafau representation levelsmarginally compared only to the using . Among four the = 2% (SD mammalother species the We foundthat identified priorityconservation spatial areas the prioritisation using mammal species, We suitability habitat models used torepresent the distribution of 184terrestrial biodiversity (taxonomic, phylogeneti island, and thepriority areas forthe facetsconservation ofthree ofmammalian representing distribution co the ofall Dicerorhinus sumatrensis maximus sumatranus Sumatra Four charismat potential, i.e. as “flagships” formarketing campaign surrogates Conservation organisations andgovernments charismatic oftenuse megafauna

:

the Sumatran tiger 2%

to represent ,

) for the ) forthe rhino. Combining four the together species the extent the theirprotectionto which other biodiversity benefits elements. ic megafauna speciesic used are as surrogatesinthe Sciences, The ofUniversity Queensland, Brisbane, QLD 4072, 160

habitat betweeneachof overlaps n ,

a species, theextent of suitablea species, habitat of Sumatran tiger phylogenetic groups

Sumatran

varied broader biodiversity.broader While speciesthese areprimarily selected . W .

Panthera tigris sumatrae Panthera tigris , ranging frommean of52% a(SD e examined how well each ofthese performede examinedhowwelleach species in orangutan c and functional). c andfunctional). - occurring occurring mammalterrestrial onthe species s , it is, it important evaluate theirsurrogacy to

and Pongo abelii 74 functional

the fourspecies the charismatic , Sumatran

and

Sumatran trait groupstrait

= elephant megadiverse

only improved the 27%) for27%) the tiger to software Zonation. rhinoceros . We then

Elephas Elephas

island of as covered

and the

This article isThis article by protected copyright. All rights reserved. Abstrak

Accepted2. 1. Article5. 4.

Kami menggunakanKami pemodelan habitat kesesuaian untuk merepresentasikan distribusi keanekaragaman hayati mamalia mengevaluasi bagaimana sumatera tigrisPanthera sumatrae sebagai digunakan memberikan keuntungan kampanye, penting pengganti Walaupun spesies sebagai Organ areaprotected network is required. resources in surrogacy analyses are determineimportant in to order ofconservation allocation the a importance. ofsystematic Wecombination that suggest spatial prioriti mammalian biodiversity, maymiss butthey with somehighbiodiversity keyareas Synthesis andapplications. charismatic species phylogenetic (8%) andfunctional(19%) didnotoverlap diversity with any ofthe someWe top foundthat ofthe outperformedmost mammal with ofother species similarrange sizes. highestproportion ofthe conservation priority areas.

isasi konservasi danpemerin spesies te Pongo abelii restrial restrial biodiversity pengganti untukmerepresentasikan biodiversitas lebih yang luas. lainnya

spesies ’ suitable habitat juga juga untuk mengetahui apakah perlindungan

dan (taksonomik, filogenetik, dan fungsional). - , bagi rich keempat gaj di Sumatera danareap

pengganti Sumatera, diPulau yaituharimau sumatera badak sumatera W biasanya memilik utama peran

ah sumatera

areas such Sumatra, as of wherean expansion the ide aspek aspek Empatbiodiversitas lainnya. megafauna karismatik - ranked for conservation areas taxonomic (28%), - ranging charismatic canrepresent species broader spesies spesies tersebut merepresentasikan distribusi spesies tah seringkali menggunakanmegafauna seringkali tah karismatik

.

Elephas maximussumatranus

Dicerorhinus sumatrensis Dicerorhinus rioritas untukkonservasitiga aspek

The Sumatran sebagai spesies tersebut dapat tersebut spesies

tiger also tiger also “ . flagship

Kami , sation and orangutan ” untuk This article isThis article by protected copyright. All rights reserved. Sumatranelephant, Sumatran orangutan, Functional Keywords

Accepted5. Article4. 3.

biodiversitas, Sumatera, seperti mana di untukpenting menentukan sumber alokasi dayak untuk menentukan area merepresentasikan dan Sintesis aplikasi karismatik mana pun. (8%), danfungsional filogenetik (19%) tidakbertumpang areaBeberapa konservasi mengungguli besar sebagian mamalia yang lain memiliki luashabitatyang serupa. melingkupi keempatDi antara karismatik, yangbagi spesies habitat sesuai harima meningkatkan = bervariasi,lainnya berkisardarirerata Tumpang menggunakan perangkat Zonation.lunak karakter 184 spesies mamalia 2% yang

) diversity, phylogeneticdiversity,spatial conservation prioritisation, Sumatran untuk badak istik - penting untukkonservasi tindih habitat tindih habitat keempatdi antara spesies karismatik dan spesies mamalia area area

fungsional. Kami mengidentifikasi kemudian areakonservasi prioritas tumpang konservasi prioritas . prioritas keanekaragaman Menggabungkan . Spesies karismatik dengan yangdistribusi dapat luas ter

- estrial, 160 estrial, tindih prioritas konservasi

habitat untuk keanekaragaman taksonomik (28%),

kelompok rhinoceros, Sumatran tiger,

biodiversitas

mamalia, tetapidapat dengan proporsi tertinggidengan proporsi distribusi distribusi dibandingkan 52% (SD =2 52% (SD dan analisis surogasi merupakan yang hal perluasan

filogenetik, danfilogenetik, 74 keempat sedikit hanya spesies . Kombinasi antara onservasi di areayangkayaakan

kawasan konservasi diperlukan. diperlukan. kawasan konservasi

7%) 7%) jika jika untuk harimau hingga hanya menggunakan harimau. - tindih dengan dengan habitat spesiestindih juga . Harimau juga sumatera surrogate species surrogate melewatkan kelompo analisis u sumatera u sumatera k

beberapa spasial spasial 2% (SD 2% (SD

This article isThis article by protected copyright. All rights reserved. mammals Americarepresentin Latin substantial for proportion of habitats highquality other terrestrial from do Williams ( al. et charismatic species, reported areeffectivemegafauna species surrogates 2010) Angelstam, 2004) conservation large conferprotection to a number co of biodiversity species in theirnatural vastconservation andthe ofbiodiversity range biodiversity primarily seekfunds for directly the and rarely flagships are aimed benefit to audience” marketing of possession campaign onits based Aflagshipspecies flagships. employ Maloney,&Possingham,Joseph, 2009) conservationeffortshence must strategically planned be andprioritised Conserving Introduction

Accepted Article not perform better in not perform in representing better the random . However

While actasumbrella species do notnecessarily a species orgroupa species of species, little (Thornton etal.,2016) (Thornton (Verissimo, MacMillan, & Smith, 2011) , when all biodiversityall features the at same time (Smith, 2012) &Jones, Veríssimo, Isaac, ly selected

evidence for the effectivenessevidence surrogate forthe ofvarious schemes 2000) , p

revious workhas mixed shown as results to charismatic whether properly planned properly

for species regional charismatic megafauna both mayas serve flagship and umbrella

found areas that identified onthe based habitat potential hasa great to

species. species. In contrast, conservationnetworks for jaguar

is

“a speciesusedas focus the ofconservation abroader ,

and giant the panda .

in many and large cases charismatic . One conservation s . Oneconservation

conservation diversity birds ofmammals and than for biodiversity. that one more or that appealto target traits the . require protection However, most

is not is not . Considering limited resources for contribute to persistenceof the in the in the United States. Similarly, ‐ Ailuropoda melanoleuca occurring species occurring

feasible Andelman Fagan( & trategy trategy that is often isto applied distribution offlagship species due to limited resources of the , conservin (Bottrill al., et 2008; — , including ,

(Roberge & campaigns species species whose other

Panthera onca areas identified

i s g 2000)

, an flagship as effective

(Caro, (Caro,

,

This article isThis article by protected copyright. All rights reserved. the Sumatranmegafauna species: tiger C 11 theto achieve under commitments set theConvention Diversity’s onBiological AichiTarget The protectedcoverage area ofrequires Sumatra from ~11%an expansion current the to17% etal.,2007; Soehartono Susilo,e Soehartono, megafauna conservationefforts is toguide (Devictor 2010) et al., protectedunderrepresented in areas,taxonomicwhereas diversity facets overlap among toppriority the conservation areas mammalfor conservation across the three arediversity notspatially congruent example, facets different because of often biodiversity show non 2006) multidimensionalin a spacedefined niche bytheir life evolutionarytheir history biodivers However,diversity. taxonomic does diversity capturealone not otherimportant facetsof invertebrates andplants for theother mammals, provide yet for poor representations amphib conservationpriority areas African “Big forthe species represent priority Five” highly areas forendemicmammal species surrogate inChina andbird onservation efforts have inSumatra largelytargetedtowards been

Accepted, assumingproportio Article . This globallimits understanding potential the of surrogacy for charismatic species (Brum 2017) etal., Sumatra is a part ofthe Sumatra isapart biodiversitySundaland hotspot evaluations ofsurrogatespeciesfocused representationMost ontheir of taxonomic ity, phylogenetic suchas diversity global global mammal hotspots defined using taxonomic, phylogenetic andfunctional . nal allocation area toland across the major inIndonesia islands

(Di Minin (Di Minin &Moilanen,2014) . On a country scale,funct. Onacountry (Faith, 1992) (Faith, (Mazel e (Mazel — Panthera Panthera sumatrae tigris an a — d functional diversity common t al., 2014) t Wibisono,al., al., 2007;Soehartono, et 2007) the distinctiveness based among on species ional diversityofbirds inFrance practice practice . - - history traits

co . Similarly,there4.6% is only

( ngruent spatial For patterns. Li &Pimm , whereof use the charismatic (MoF, 2007;(MoF, PHKA, 2015; , — the the i s overrepresented the the distribution of species ians, reptiles, reptiles, ians, four (Petchey &Gaston, Sumatran elephant 2016) charismatic charismatic . InAfrica,the i s . .

This article isThis article by protected copyright. All rights reserved. Acceptedmill2.3 1990 settlements (35.6%). Otherproduction landuses ofthe(40.8% include island) agriculture landand biodiversity oftheconservation (10.6% system island), water protection and (13%) mainon the island 2009) Sumatra islands isoneof largest inIndonesiawith atotal the of430,000area km Study region andMaterials protectionthe ofthefour charismatic species. phylogenetic andfunctionaldiversity, assesshowmuch andto these areas wo theto identify priority areas formammal inSumatra conservation basedontaxonomic, Articleof other Sumatra andthe native mammalterrestrial on species island.Thethe aim second was taxonomic, andphylogeneticmammal functional (hereafter diversity “surrogacy potential”). thewhether phylogenetic diversity when comesespecially it multiple to ofbiodiversity,facets such as occu mammals rhinoceros maximussum Elephas r  ring species. . Theislandissurroundedbymany satellite focused smaller islands,yet only this study 2010, an estimated 7.5 million2010, anestimated 7.5 ofprimaryhectares forest measureOur firstaim the wasto habitatoverlap thefour between charismatic species ion havehectares degraded, been leaving of only 30% Sumatra byprimary covered have an Dicerorhinus sumatrensis (Margono, Turubanova, Potapov, &Tyukavina,Zhuravleva, 2012) available habitatfor species inSumatraavailable fourflagship ofis representative the

Methods umbrella However, (Fig. 1). Forest (Fig. 1).Forest land usezonesaredivided three functions:into atranus . importantIn this work, this we addressed shortfall and evaluated

effect, and that this proclaimed , the the Sumatran orangutan . Conservationists

saving them alsosave will umbrella effectumbrella

Pongo abelii often

remains untested, largely propose in Sumatra functional and

and

that these large a number of the the were cleared and Sumatran uld benefit from . Between 2 (GADM, other co other -

This article isThis article by protected copyright. All rights reserved. classes ofhabitat suitability range The elevation deductive features. andhydrological resulting models c habitat preferencesspecies for three environmental variables we al.2016) et been models have suitability in used a numberstudies ofother (e.g., onaregional distributions represent species’ models to onthe island wedistribution species, models forall deductive used (expert Information). listed,being including thespecies foursurrogate ( (excluding on species onlythat islands). occur resulted satellite This in 193mammal spe information website onthe IUCN pages List thatoccurSumatra Redtofilterspecies in di geographic distribution forterrestrialranges mammals ofThreatenedon IUCN List Specieswebsite(www.iucnredlist.org) and Red the (ii) IUCN We ofnative list obtainedthe mammalterrestrial species in Sumatra usingthesearch (i) tool Mammal distribution data million, withagrowthrateof14.5% 2000 between and2010 in2010 forest (ii) (i) d notreturnthesame wemerged number ofspecies, them thespecies and checked

Accepted Rondinini et unavailable. re al. (2011) assessed species’habitatsuitability byconsidering Article

found, yet cannot persistfound, cannot the yet without primary and habitat; medium high habitat representing suitability, primarythe where habitat speciescanpersist; the Given the lackGiven the ofcomprehensiveoccurrence for data developing statistical

and global and global scale (e.g.,

habitat suitabilityhabitat (Margono al.,2012) (Margono et

(Rondinini al.,2011) et , repre Brum 2017) etal. . senting habitatthe secondary where species canbe The h uman ofpopulation Sumatra in2010was50.6 see Appendixsee S1in : , wherecomprehensive occurrence data

(IUCN, 2016) (Rondinini al.,et 2011)

: -

(BPS 2012) the typesofthe landcover, driven) habitat s

. Because the two lists . Becausethetwo lists Supporting Supporting .

onsist ofthree onsist uitability . Habitat Thornton cies cies

This article isThis article by protected copyright. All rights reserved. lowestthe value forconservation based occurrence level onthe ofbiodiversity features fromstarts and fullextent ofinterest, the oflandscape then iterat species and habitat types) whichrepresenttheirspatial distribution. The Zonation algorithm 2014) We identified theAssessing representationofpriority areas 3.4.1version ‘sp’packages charismatic Wemegafauna species. spatialdataprocessingR the performed using all ofproportion aspecies’ ofsuitable habitatextent overlapping withth ofsuitable habitatextent other withthat ofall mammals. was Representation calculated asthe We surrogacy the assessed potential of megafauna charismatic by species overlaying their theAssessing bythecharismatic representationofmammal megafauna species th (including of 184species werespecies notmodelled split persist.likely O to the aremostintended conservationareasystemsareaswherespecies to include resulting represent the occurrence(hereafter“extent ofthespecies habitat”) ofsuitable because we Accepted Article (iii)

orangutan unlikely to occur.unlikely to low . The input data for. Theinput analysis data Zonation mapsare raster biodiversity of features (e.g., In the In the

habitat suitability (R DevelopmentCoreTeam, 2008) spatial prioritisation analy conservation (Pebesm

population f the193 mammal native terrestrial species inSumatra, two the a & Bivand, a & 2005)

s

by Rondinini etby Rondinini al. (2011) , representing areas where , representingareaswhere the aregenerallynotfound species or priority areas usingtheZonationv.4softwarepriority e fourcharismatic species). (Nater (Nater et 2017) al., sis, we ofhigh used only areas habitatsuitabilityto

and ‘raster’ and ‘raster’ based on

were considered asonespecies,andeight .

; therefore, analysis our includeda total (Hijmans vanEtten, & 2012)

multiple facetsof biodiversity

ively ively removes with areas e habitatof each (Moilanen et al.,

recently

in R

This article isThis article by protected copyright. All rights reserved. Accepted species to repr island. endemicthe We 14species twice weighted toSumatra as a compromisecomputational between time for andrelevance managementconservation on mapssuitability from th taxonomic We diversity. performed area speciesbiodiversityWe individual the used as features for the prioritisationspatial based on Biodiversity prioritisation for features spatial detailstechnical prioritisation); andan basedanalysis onfunctional groups (functional prioritisation). analysis (taxonomic prioritisation);an analysis basedonphylogeneticgroups (phylogenetic to analyses identify prioritiesseparate different forthree f biodiversity (BLP) toproduce more solutions compactreserve We connectivity incorporated considerations also theBoundary LengthPenalty byusing Article couldidentify offeatureshence core areas in biodiversity the ranks cells across the landscapebased onthe most important occurrenceof and afeature, &2017;Thuiller, Jetz, Olden,Whittier,Strecker, Paukert, & 2011) congruency ofof areasformultiple facets priority biodiversity umbrellaassessing effects ofsurrogate species zoned mapfor toppriority conservation. areas has of beeZonation already 0(lowestbetween priority) 1(highest be and that can hierarchical visualised as a 2009) ofconservationranking priorities throughout the landscape 200 et(Moilanen al., . Each cellofraster output the map ofconservation arank contains prioritiesranging W e used Core Are Core e used esent higheresent the global of theirconservationvalue island.We onthe also

are providedare 5) . This iterative generates removalmap a process hierarchic showinga e original 300m original e resolution to1km.We aresolutionof 1km used as a Zonation (CAZ)

in Appendix S - weighted up resampling toscale habitat the

2 as the cell as the removal Zonation in rules .

(Di & Minin Moilanen, 2014)

(Moilanen (Moilanen & Wintle, 2007)

- poor areas (Moilanen, Kujala, & Leathwick, (Brum Pollock, 2017; etal., much as .

(Moilanen acets: aspecies acets: the non the n usedfor

and examining . We three ran

et al., 2014) , which - endemic Further Further - level . This article isThis article by protected copyright. All rights reserved. Acceptedoutpu mass,maximum reproductive speed (weaningage,gestationlength), longevity, reproduction fiveselected lifehistoryandecological categories,represented traits: bynine adultbody prioritisation,based us spatialgenerated maps groupdistribution ofphylogenetic from thismatrix. contain themultiplied twomatricestoobtaingrid habitatnarrow distributions) reducesthe threshold risk ofin maps 1kmresolution into at werereclassified a binarymapsofthreshold 0.05 with awe created matrix binary of species had fewer number ofspecies To in Sumatra. represe variables.Ourquantiles were binary than wider those in used Article 10%into split quantiles altogetherThese eigenvectors accounted variation. for69%ofthetotal was Each eigenvector distance matrix super species in Sumatra, afterexcluding three species mammalianwhich were inthe not available by updated We themammalian used phylogenetic supertree created by followinggroups framework the by developed differencesshowed nosubstantial ( on analysis performedsensitivity weightingalternative ofendemic a the but results species, - t (neonate bodymass, littersize per andlitters year) and resource use(trophiclevel and tree. We performed then analysis(PCoA)coordinate phylogenetic principal onthe ed the numberbelonging aparticular phylogenetic ofspecies to We group. then We mappedWemammalian with functional traits similar with phylogeny procedure prioritisation For spatial based Fritz al.Fritz et (2009) and selected the firstand selectedthe 16 eigenvectorsexplained which >1% ofvariation. total ing life

(see (see , and then were converted , andthenwere converted gridinto Appendix S4

to create aphylogenetic distance matrix among 181mammal - history asopposed traits phylogenetic to eigenvectors. We troducing commission errors troducing commission errors penalising specieswithout with see × phylogenetic group× phylogenetic

on phylogenetic diversity, we defined phylogenetic Appendix S4

× for phylogenetic groups matrix, in whichagridcell the Strecker etal. and (2011) Brum (2017) etal.

sensitivity test) sensitivity nt phylogenetic groups inspatial maps, ).

. Area Bininda Brum etal. (2017) × , and then conver -

species matrix. we Finally, weighted habitatsuitability - Emonds al. et (2007)

because we ted into

(this -

and . This article isThis article by protected copyright. All rights reserved. Accepted weremuchspecies for (SD 18% thefororangutan elephant (SD lower: =12%),9% the = habitat ofother Sumatranmammals charismatic2). Therepresentationlevelsfor(Fig. other mammalother sp megafaunaAmong charismatic four species, the tiger hadthe highest overlap the spatial with forterrestrialSurrogacy mammal species Results species to compare performance ofthefourcharismatic the m In addition, calculated we also surrogacy potentialsthe ofothermammal species inSumatra Sumatra that overlapped extentwith the of habitatof suitable the surrogacypotentialofthe megafaunaproportion charismatic asthe them extenthabitat ofsuitable withthe ofcharismatic megafauna species. Wemeasured 10%, and top 5%, areas 17% 25% ofpriority forscenarios prioritisation andoverlaideach Articleprioritisation andlevels of traitsintrait functional intaxonomyinputs ofprocedure preparationisprovidedin data matrix ofgridresulting multiplied Next, wethe grid 10%quantiles, and into ofeachwe assigned presence/absence same level ofcategorical each traits binary matrixin a of species etal.,(Jones 2009) al.(2013) et habitat breadth). We global the for used dataset mammal terrestrial traits compiled by . We th ran

and Di Marco &Santini (2015) ecies, representing a mean =27%)oftheextentsuitable (SD of52% e prioritisation analysis

- and AnAge based prioritisation, groupbased phylogenetic maps phylogeny in ×

trait trait spatial mapsinto offunctionaltrait levels. The detailed × species (Tacutu et al.,2013) et (Tacutu matrix withspecies

separately

Appendix S2 from sources,various suchas PanTHERIA - based prioritisation. We extrac then

. We presence/absence the assigned of using species’ habitatsuitability maps as e gafauna species with species gafauna

.

× trait ×

trait. Wetrait. continuous split traits species under consideration matrix. We the converted of priority acrossareas - size trait level trait level to spec size

that of that - based other Pacifici Pacifici ted the the ted ies. ies.

. This article isThis article by protected copyright. All rights reserved. Accepted halfofwestern areas island.Priority forconserving the functionaltraitswere m solution basedspatial forprioritisation groups onphylogenetic wasmoreclumped inthe were Sumatra andmountain western ranges. taxonomy In sharedareas amongscenarios mostly prioritisation three in the ofoccurred highlands northern 21,595 km priority areastop 5% for biodiversityfacets,each anareawhich altogether encompassed of phylogenetic andfunctionaldiversity prioritisations 3).Whenlooking scenariosthe (Fig. at We substantial spatial discovered mismatches areas forpriority basedonSurrogacy multiplefacetsof biodiversity = 0 surrog betweenoverlap range tiger their and range ranging from89%. to Combining four the 40% Article of12%. 11 werelevel species well Theother endangered represented by the withthe tiger underrepresented bythe pangolin habitatsuitable fortigeroverlapped substantially with (52%) thecritically endangered Sunda 41% but only oftheelephant’s treeshrew tailed withoverlap orangutan the (i.e.,81% forbig the horseshoe bat eared for elephant93% the (i.e., large the lesser an 11%),  12%) 12%) compared the to alone. performance ofthetiger more out spread acrossthe island, including lowland coasts. central andeastern The ate species only improved therepresentation levelsby a mean =6%, (SD range of5% Most ofthe Most habitatsuitable fororangutan andrhinoceros overlappedwith tiger, the d 2% rhinoceros for the (SD Only highspatialoverlap =2%). twospecieshada with Manis javanica 2 , we found an , wefound an Ptilocercus lowii Rhinolopus macrotis tiger, i.e., tiger, i.e., the hairy . species Of (Fig. 12endangered the 2), overlap ofonlyoverlap 3,227km extent of with suitable habitatoverlapped tiger’s. the The ).

). Likewise, two had only species a h - footed myotis - nosed otter

among theareasidentified taxonomic, under - eared horseshoe batand71%foreared the pen 2

- (15%) across the the Prioritythree facets. across (15%) based prioritisation, prioritisation, based the priority areas Lutra sumatrana Myotis hasseltii

only

one was

, and 85%forthebig

with representation with representation igh spatial ore congruent congruent ore - - This article isThis article by protected copyright. All rights reserved. s hadopposite, potential anunderperforming than sizewas predicted byits surrogacy of compared to represented varyi potential surrogacy ( representing areas priority revealed that the extent ofsuitable of habitat was a goodpredictor were outside extentofhabitat the suitable ofthecharismatic species. areas,priority (taxonomy 28% covered were not smallonly bya amount comparedtiger (Fig. the to We 5). identified also priority areas that tiger. Combiningthe all charismatic species improved representation the of areas,but priority suitabletiger’s habitat,represented less than represented phylogenetic particularly well. priorities Elephant, approximately having half of the coveragehabitat had highest areal areas ofpriority forthe threebiodiversityfacets, and it areas,priority the given of distribution restricted species 5).The these (Fig. t ofhighareas conservation priority, these onlyaccounted for amountssmall oftop areal 0.05 (complete summary statistics in prioritisation, IQRelephant hadelephant most the in representing variability e.g.,conservation priorities, for taxonomic higherconservationrelatively priority ranks thanthetigerandelephant. Among four, the conservation priority (Fig. 4).Theorangutan andrhinoceroscovered to phylogeny Accepted habitat.uitable Article Evaluation on the While mostofhabitatsfor suitable orangutan the the andrhinoceros overlappedwith habitatIn general, ofsuitable most of extent the ofthecharismatic was species ofhigh most - based (20% thanbased overlap) taxonomy

ng proportion by any ofthefour charismatic species’ distributions, e.g.,for top5% the other species with species other similarsize habitat.of Theelephant, on suitable the Fig. 6

potential oftheother180mammal species inSumatra in ). However, species ). However,species with similar ofsuitable size habitat

= 0.44,IQRtiger - s based priority),based 8%(phylogeny

of priority areas. The surrogacy potential tigerhadahigher Appendix S4

= half of amount the ofpriority areas covered by 0.23, IQRorangutan - ). based prioritisation prioritisation based (17% overlap).

- based) based) and 18%(trait

= 0.10, IQRrhinoceros

areas inSumatra with iger’s suitableiger’s - based)

= This article isThis article by protected copyright. All rights reserved. Accepted offlagshipchoice species depends on broad the of conservation goals developing people within areas charismaticthe spec insufficient for Sumatran biodiversity the given restricted distribution of the two species. protectioncontribution tothe ofbroaderbiodiversit areasprotect within the rangeof the andorangutan rhino the would result high ina encompassed top restrictedtheir distributionHowever,almost range. the entire representedrelatively lowerrhinoceros priority amountsoftotal areas, which mountain western ranges most where ofthepriorityareas of expectedproportion areasthan priority byitsrangesize becauseitdidnotocc Sumatra interms oftherepresentation ofco mammals finding fromaprevious study that large predators forother haveimportantsurrogateroles Article potential surrogacy than b predicted compared othercharismatic the species to inSumatra. However, the tiger alsohadhigher phylogenetic andfunctionaldiversity). is This related to itsrelatively large distribution covered high it amountsof areasforthe priority mostbecause co ofthe Sumatra charismatic megafauna assurrogate We recentlydeveloped used techniques evaluate to the conservationeffectiveness ofusing of surrogatesThe role the megafaunainSumatraasbiodiversity charismatic Discussion The combination of The combination The Sumatran elephant secondhad the as potential highest asurrogatespecies in n mammals (Thornton etal.,2016) (Thornton

- ranked areas. priority Thisimplies allocating conservation resourcesto . We. found that theSumatran tigerhad surrogacyhighest potential ‐ occurring mammals represented well withinits were distribution, and spatial spatial ies’ lowhabitats with priorities. highand conservation .

prioritisation surrogacy and allowed analysis identifyus to s ased onitsdistribution Thisased sizealone. supportsthe

for taxonomic,phylogenet - occurring species. However, covered it three biodiversity facets (taxonomic, y, butthis alone strategy will likely be

were found

ranges oftheseranges twospecies ic ic and functional diversity . The orangutan and. Theorangutan

match ur in most of ed a lower

with The

of

This article isThis article by protected copyright. All rights reserved. Accepted that factors tobeconsidered tomaximiseneeds conservation effectivenessthe ofselected (and important relate areasforbiodiversity how these tochar planning tonecessary mammalian duetodataavailability diversity species, whichconsidered not are currently ava featuresinput for study.This future speciesfor all study consideredinthe ordeveloping inductivemodels distribution species as addresseddisturbances. Thisissue canbe byperforming suitabi habitat presence Possingham, 2006) excluded fromhave been thedistribution distributions compared ranges geographic (IUCN to ra Articlewell biodiversity that rangeoutside the ofcharismatic species. and policy makers should inSumatra protect important areas forbiodive areas,priority withallspeciescombined, even suggestthat conservation managers ourresults charismaticConsidering that megafauna notrepresent did the of wholeextent charismatic thatalsomegafauna species provide tothebenefits broader biodiversity conservationprioritisationspatial techniques areasto identify the within of ranges use not be marketingthe campaign preferences andthe of audiences,so their target our analysisshould - targeting surveyed taxonomic and groups types habitat Habitat Habitat suitability models inthis used studyareamore depiction accurate ofspecies ,

(Di etal., Marco 2017) as species mayas species beextirpated fromhabitat suitable tohuman due its d to prioritiseconservation effort flagships. between Instead,

conservation targets incorporate other vertebrate, other incorporate invertebrate and planttaxa spatial in conservation charismatic megafauna . However, suitable habitatisnot necessarily translated species into . Moreover, , yet of benefit pr the their ,

however in conservation Our findings withp agree (Rondinini, Wilson, Gra Boitani,

while was ourstudy aimed only identifying at along facets biodiversity all ,

requires collecting for occurrence all data ilable. (Di Minin & Moilanen, 2014) is not is not enough to In nge maps) because habitats unsuitable nge this studythis

otection can beimproved byusing ismatic thereare species), other , revious work sugges

we only considered achieve broader litymodel validation we the propose use of rsity rsity that occur . Future workFuture isstill ntham, & - caused identified .

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This article isThis article by protected copyright. All rights reserved. Accepted functioning. theprotection guarantee of historyand themaintenance ofecosystem evolutionary 2004) Amundsen, Species aretheaspects. of products andecologicalevolutionary processes biodiversity conservationplanningcould in facet in result the failure otherto maintain the represent top areas priority underallbiodiversity facets phylogenetic andfunctionaldiversity Sumatra, w species questioned andregarded as inadequatebecause it distinctivenessfails to represent among ofbiodi rest Article protecting the of habitats charismatic willalsoprovidemegafauna species co protectio species that vital ecosystemplay orare functions but evolutionarydistinct, oftenreceive little endangered charismaticby the there are species, also non many threatened foractions.Whilefunding anddefinepriorities it is importan Charismatic havespecies beenthefocusmany ofconservation organisations biodiversitymultiple facetsof What conservation actions. areas,priority should we The useoftaxonomicaloneforpriorities diversity identifyingconservation hasbeen (Vane n (Isaac, Turvey,Collen,(Isaac, Waterman, 2007) &Baillie, e found that onlye foundthat (3,227 15% km versity can versity lead tothe extinctionofunderstudied species.

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species

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t extinctionto reduce risks faced . Simply assuming that , 000 the costthe of undertaking km

2

would be required would berequired to ufficient to ufficient to - (Bøhn & charismatic , both - benefits to the

the taxonomic,the

to allocate This article isThis article by protected copyright. All rights reserved. occurri used conservation decades journals thanit to inpast Southeast(i.e., Asia) todayreceivingproportionally that is species is a ofkey driver conservationaction there. Thisisespecially relevantwithin aregion concentrations and ofterrestrial threatened mammals, distribution andthe ofcharismatic formulating marketingthe strategy amoreThis needs thorough assessment systemsarea funds andraise flagship species or flagship from fleet list overlookedspecies topromote newprotected the taxonomic,the phylogenetic and functional diversity ofSumatran mammals. Choosing new would result much ina coverageofmore complete for toppriority areas an area assmall 1,734 as im boththegoal protection of speciesand protectionofareas charismatic the of highbiodiversity audience,target so and campaigns create that increaseslightly the spatial ofpriority representation basedareas, on elements. ourresults suggestedthat combining Although the four charismatic species only charismatic species’ habitats thatalso t give and cost the distributionHowever,entire protecting of athreatened charismatic species biodiversity an importance couldbe effective strategy forprotec

Accepted distribution outside the ofportance species. Our these resultsthat evenprotecting showed Article ng with charismatic species We study Sumatraasour chose case areabecausehosts some it ofthehighest itis followacomplementaryWeimportant suggest to also approach, integrating asa a Selecting wide - inefficient. Therefore, wesuggest prioritising protectionthe ofwide a ‘ increase -  fleetflagship ranging charismaticspeciestodelineateprotected areas ofhigh 5,949 km (Smith al.,2012) et potential funding support and

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protected area establishment area protected and

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the conservation of .

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- 2. 1. This article isThis article by protected copyright. All rights reserved. island: the managers that inSumatra may advanc help fragmented rate conse usingcharismatic species spatialprioritisation techniques. charismatic species, and identified biodiversityareas important outsidethedistribution of benefits framework conservationstrategies plan in whichtheprotection to o as planning makers also consider broader should for benefits aspectsin other ofconservation biodiversity megafauna primarily are forsake species,protected the ofthese conservationdecision enforcedareas are bymeans Sumatra al.,(Linkie et 2015) enforcement protected patrolling efforts, the areasfrom suchas tosafeguard activities illegal Accepted Article raise funds andraise gainpo unprotected top priority areas identified study in this byusing Sumatran the four” “big to ofExpansion protected current areas or establishment areas protected tocover ofnew with the Mountain Ranges,Barisan especially facets wherethe three of over also biodiversity taxonomic, phylogenetic andfunctionaldiversity, Leuser suchas LandscapeandBukit Enforcement ofprotected management area where highthere is inplaces between overlap habitat ofthe four”, “big be newflagship species/fleetshould identified

of human population ishigh, rvation While

broader aspects of biodiversity; where (i) where (i) conservation ofthe charismatic megafauna protected isprominent(ii) and

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we facerateof increasing species worldwide. B

planning is planning challenging it toprotectit allaspectsofis ideal implementing biodiversity, s ased on our findings, we provide several weprovide onourfindings, ased recommendations forconserva . Therefore, our . Therefore, recommendation is charismatic megafaunacharismatic species. litical and public that exclude any harmful illegal that excludeanyharmfulWhile charismaticactivities. illegal

land tenure conflicts areprominent ,

especially especially in growth developingcountries where the

we showedthe‘umbrella’ effectiveness ofdi ing support

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also relevant

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Our study revealedOur study a

in regions and wildlife habitatsand wildlife are

ystematic beyond fferent fferent -

tion tion lap

3. This article isThis article by protected copyright. All rights reserved. Rondinini, (Sibarani, DiMarco, 2019) & Kark, Data Data Accessibility online. other testsand ofspatial sensitivity prioritisation available results (AppendixS4)analysis are (A ofmammalList preparation species inSumatra S1),data (Appendix for prioritisation analysis Supporting Information Research Council. Australian was supportedbythe Endowment ofFinance(LPDP) Fundfor oftheMinistry Education fundingtoMCS.SK and the We A.Residefortheadvice thank in Zonationusing Acknowledgements forapproval publication. writingofthe themanuscript. the writing,andall M.C.S. andM.D.M.analysed data.ledthe authors contributed in globalC.R. andprovidedthe developed habitatsuitabilitymodels forterrestrial mammals. theideas, M.C.S,M.D.M.,andS.K.M.C.S. conceived the and study designed Authors’ Contribution

Accepted S2),representationppendix ofnativemammal terrestrial (Appendix and S3) species Article species distributionspecies modelling andspatial conservation planning. Collation and digitisation recordsofall ofspecies occurrence to taxa enablemore robust available via available via the

E ditor

for constructive

Dryad Digital Reposito s

All contributed criti authors

commentsmanuscript onthe ry .

https://doi.org/10.5061/dryad.6n0b28n

;

Bob Smith, ananonymous

cally draftsfinal andgave tothe .We theIndonesian thank

.

M.D.M. and reviewer

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- – - This article isThis article by protected copyright. All rights reserved. Accepted islandinthe Indonesia. Figure Article Figures Williams, Flagship ecological H.,Burgess,C.(2000). species, N.D.,&Rahbek, P. Africa. andconservingcomplementarity the of diversity mammals in sub andbirds 1

. Map ofthestudySumatra region, Island. Animal Conservation

, 3 (3), 249 – 260. doi:10.1017/S1367943000000974 Small map insert location of showsthe

- Saharan Saharan

This article isThis article by protected copyright. All rights reserved. fouracross the charismatic CR: CriticallyEN: Endangered; Endangered. Note scalesofy that the DD:categories. LC: DataDeficient; Concern;Least NT:Near Threatened; VU:Vulnerable; of megafauna: charismatic (a)overalland(b) Figure (b) Accepted Article (a)

2 . Therepresentationofthe co megafauna species. - occurring mammaloccurring within suitable the species habitat categorised according totheirIUCN List Red

- axis ofdifferent are (b)

This article isThis article by protected copyright. All rights reserved. Acceptedmaps re ofandtop25% top10% Top 5%(values >0.95), (>0.90), 17%(>0.83) (>0.75). Theinset importance). We highpriority intoareas categorised fourgroups ofnestedpriority rankings: thethroughout la top maps theprioritisation results, spatial theof report reporting ranking conservation priority Figure Article 3 port the spatialdistributionof the port charismatic megafauna species. .

Priority conservation areas in Sumatra onmultiple based facets of The biodiversity. ndscape 0meaning (with lowest importance and1meaning highest

This article isThis article by protected copyright. All rights reserved. Accepted fromranging 0(lowest 1priority) priority). to (highest priority ranks werecomputed that based oniterative removal cell process of Zonation, habitatsuitable ofcharismaticmegafauna species. They Figure Article 4 . Boxplots showingvariationofconservationpriority the theranks within extentof

- axis represents the conservation

This article isThis article by protected copyright. All rights reserved. Accepted megafauna.four charismatic size greater ofarea is protected ofprioritylandscape andthehighest25% sc protected.Forexample,to be top the 25%priorityarea refersofthe ofthe to25% size entire charismatic species. Figure Article 5 . Representation oftoppriority withinareas the extent of A decrease inpriority correspondswith increasing sizeofareasthat need

less represented thewithin extentofsuitable habitat of the ores from algorithm. Therefore, Zonation a

suitable habitatof

This article isThis article by protected copyright. All rights reserved. Accepted models.additive species). Blue linesrepresent fitted ofpriority values area represen Sumatran rhinoceros. charismatic T=tiger,megafauna E=Asian species: elephant, O =Sumatran orangutan, R= to megafauna relative their areas priority and (c), top 25%priorityareas (d) within the suitable habitat of charismatic Figure Article 6 .

The representationoftop 5% priority topareas (a), 10%priorityareas (b), top17%

Grey point extent s

of habitat.suitable Letters in plot the indicate

the otherthe mammal species inSumatra (

tation ongeneralised based indicate the 180