Past, present and future of the ecosystem services provided by cetacean carcasses

Ecosys em Se ices 54 (2022) 101406 A ailable online 3 Feb ua y 2022 2212-0416/© 2022 The Au ho s. Published by Else ie B.V. This is an open access a icle unde he CC BY-NC-ND license (h p://c ea i ecommons.o g/licenses/by- nc-nd/4.0/). Re iew Pape Pas , p esen and u u e o he ecosys em se ices p o ided by ce acean ca casses M.-Ma ina Quaggio o a , * , Jos´ e A. S´ anchez-Zapa a b , Da id M. Bailey c , Ana Payo-Payo d , Joan Na a o e , And ew B ownlow c , Rob Dea ille , Se gio A. Lambe ucci g , Nu ia Sel a h , Aina a Co ´ es-A izanda i , j , Fe nando Hi aldo i , Jos´ e A. Don´ aza i , Ma cos Mole´ on k a Biological and En i onmen al Sciences, Uni e si y o S i ling, S i ling, UK b Depa men o Applied Biology, Uni e si y Miguel He n´ andez, Elche, Alican e, Spain c Ins i u e o Biodi e si y, Animal Heal h and Compa a i e Medicine, College o Medical, Ve e ina y and Li e Sciences, Uni e si y o Glasgow, Glasgow, UK d School o Biological Sciences Zoology Building, Tillyd one A e, Abe deen, Sco land e Ins i u de Ci` encies del Ma (ICM-CSIC), Ba celona, Spain Ins i u e o Zoology, Zoological Socie y o London, London, UK g G upo de In es igaciones en Biología de la Conse aci´ on, INIBIOMA-CONICET, Uni e sidad Nacional del Comahue, Ba iloche, A gen ina h Ins i u e o Na u e Conse a ion, Polish Academy o Sciences, K akow, Poland i Depa men o Conse a ion Biology, Do˜ nana Biological S a ion (EBD-CSIC), Se ille, Spain j Applied Func ional Ecology G oup, Depa men o Plan Biology and Ecology Facul y o Biology, A enida de la Reina Me cedes, 6, 41012 Se ille, Spain k Depa men o Zoology, Uni e si y o G anada, G anada, Spain ARTICLE INFO Keywo ds: Ca ion Dolphin Ecosys em unc ion Managemen S anding Whale ABSTRACT Ecosys em se ices associa ed wi h ce acean s andings ha e been al e ed by humans h ough exploi a ion o wild popula ions du ing he whaling e a and mo e ecen ly by egula ions on ca cass managemen and disposal o abide by en i onmen al heal h equi emen s. He e, we sys ema ically e iew he scien i ic li e a u e and ga he da a on ce acean s andings wo ldwide o: 1) iden i y he ecosys em se ices p o ided by s anded ce acean ca casses in he pas and p esen ; 2) es ima e he densi y o ce acean s andings cu en ly occu ing in selec ed coas al a eas a ound he globe, and analyse i s associa ion wi h human popula ion densi y and egu- la ions; and 3) iden i y and discuss he egula ions and me hods conce ned wi h whale ca cass disposal in speci ic egions o he wo ld. Ou li e a u e e iew e ealed ha s anded ce acean ca casses ha e p o ided a ich and a ied a ay o p o isioning, egula ing, cul u al, and suppo ing ecosys em se ices o ancien and mode n ci ilisa ions wo ldwide. Also, we ound ha he cu en densi y o s anded ca casses (mean: 0.090 s andings • yea −1 •km −1 ; ange: 0.001–0.978) and he disposal me hods widely a ied ac oss he s udied egions and coun ies. In addi ion, nei he human popula ion densi y no he exis ence o egula ions we e good p edic o s o s anding densi ies. Finally, we p o ide ecommenda ions o he u u e managemen o s anded ce acean ca casses, by iden i ying hose disposal me hods ha minimize cos s and maximize ecosys em unc ions and se ices. In pa icula , we encou age na u al decomposi ion in si u whene e possible; o he wise, he p esen coas al managemen s a egies could be imp o ed by including zoning, seasonal use limi a ion and educa ional ou each depending upon he local scena io. O e all, u he socio-ecological esea ch is s ongly needed o guide s anded ce acean ca cass managemen owa ds enhancing he ne bene i s ha humans and ecosys ems gain om ca casses, especially conside ing ha coas al a eas become mo e popula ed, new disposal egula ions a e app o ed, and ce acean popula ions a e eco e ing – and hus s andings may become mo e equen . 1. In oduc ion Mode n socie ies a e inc easingly disconnec ed om na u e (Dijks a, 2016). One o he many mani es a ions o such disconnec ion is he g owing implemen a ion o “asep ic” s a egies in na u al esou ce managemen . Fo ins ance, ca casses o bo h domes ic (Don´ aza e al., * Co esponding au ho a : Co ell Building, Biological and En i onmen al Sciences, Facul y o Na u al Sciences, Uni e si y o S i ling S i ling, FK9 4LA, UK. E-mail add ess: [email p o ec ed] (M.-M. Quaggio o). Con en s lis s a ailable a ScienceDi ec Ecosys em Se ices jou nal homepage: www.else ie .com/loca e/ecose h ps://doi.o g/10.1016/j.ecose .2022.101406 Recei ed 9 Feb ua y 2021; Recei ed in e ised o m 27 Decembe 2021; Accep ed 12 Janua y 2022 Ecosys em Se ices 54 (2022) 101406 2 2009) and wild animals (Ma galida and Mole´ on, 2016) a e equen ly emo ed om he ield o minimize disease ansmission isk in many egions, o en manda ed by egional, na ional o in e na ional egula- ions. This managemen scena io la gely igno es he pi o al ecological ole o ca ion (DeVaul e al., 2003; Wilson and Wolko ich, 2011) and he many con ibu ions ha dead animals can o e o humans (Mole´ on e al., 2014). Ce acean ca casses a e a na u ally occu ing elemen o he coas and humans ha e been exploi ing hem o millennia (See sholm e al., 2016), p o iding coas al inhabi an s wi h goods such as ood, oil and bones, bu also wi h knowledge o he sea while obse ing hem s anded on he sho e (Ellis, 1991). Also, s anded ce aceans a e a key sou ce o ood o sca enge s, and nu ien s o he sedimen s (e.g. Laid e e al., 2018; Tucke e al., 2019a), hus being an in eg al pa o coas al ecosys ems (Panels 1 and 2, Figs. 1 and 2). Howe e , abou o y pe cen o he human popula ion cu en ly li es wi hin 100 km o he coas , and as a esul , many coas al a eas ha e been ans o med (C ossland e al., 2005). The changes include no only he in ense u baniza ion o he coas and subsequen al e a ion o ecological p ocesses (e.g. Huijbe s e al., 2013), bu also he applica ion o policies u ging he emo al o ca casses om he coas al a eas o p o ec human heal h (Tucke e al., 2018). Combined wi h he pas deple ion o ce acean popula ions (Hacquebo d, 2001; Panel 3), he exis ing egula ions on emo al o s anded ca casses has led o adical changes in he abundance and a ailabili y o la ge ma ine biomass inpu s, which has g ea ly al e ed he ecosys em unc ions and se ices associa ed wi h ma ine ca ion o e ime. In his s udy, we e iew he scien i ic li e a u e and ga he da a on ce acean s andings o: 1) iden i y he ecosys em se ices p o ided by s anded ce acean ca casses. He e, ecosys em se ices a e de ined as he bene i s people ob ain om ecosys ems (Millennium Ecosys em Assessmen , 2005). These include p o isioning, egula ing, and cul u al se ices ha di ec ly in luence people, as well as suppo ing se ices (o ecosys em unc ions) necessa y o main ain he o he se ices. While he ecen ly de eloped Na u e’s Con ibu ions o People amewo k (NCP) includes bo h he bene icial (i.e. ecosys em se ices) and de imen al (i. e. disse ices, con lic s o damages) con ibu ions o na u e o socie ies’ quali y o li e (see Díaz e al., 2018 o an o e iew o he e olu ion o he concep ), he e we delibe a ely ocus on he bene icial side o s anded ce acean ca casses; 2) es ima e he densi y o ce acean s andings occu ing in selec ed coas al a eas a ound he wo ld and analyse whe he human popula ion densi y and he p esence o egu- la ions a ec hese eco dings, in o de o gain unde s anding on he spa ial dis ibu ion and he ac o s a ec ing he po en ial p o ision o ecosys em se ices by s anded ce aceans; and 3) enume a e he egu- la ions and me hods conce ned wi h whale ca cass disposal in speci ic egions o he wo ld, and discuss he p os and cons associa ed wi h each me hod in e ms o ecosys em se ice p o ision. Based on he indings o he pape we p o ide ecommenda ions o u u e esea ch and he Fig. 1. Ce acean ca casses suppo a wide ange o sca enge s, bo h e eb a es and in e eb a es, in di e en habi a s. See pho o c edi s in Appendix A. Fig. 2. Andean condo s (Vul u g yphus) used o in ensi ely exploi s anded whale ca casses. Howe e , he cu en sca ci y o s anded whales has induced impo an changes in condo s’ beha iou (see Panel 2). Adul Andean condo soa ing abo e a li ing whale in Pe u (a); se e al Andean condo s o di e en age eeding upon a s anded whale ca cass in Pa agonia, A gen ina (b). See pho o c edi s in Appendix A. M.-M. Quaggio o e al. Ecosys em Se ices 54 (2022) 101406 3 managemen o ce acean s andings in an inc easingly asep ically managed wo ld. 2. Ma e ials and me hods 2.1. Li e a u e e iew We conduc ed a sys ema ic li e a u e e iew acco ding o he PSALSAR me hod (Mengis e al., 2020), which is an ex ension o he SALSA amewo k ha has equen ly been used in sys ema ic e iews (G an and Boo h, 2009) and ha gua an ees concep ualiza ion, accu- acy and ep oducibili y. In he i s s ep, we de ined ou s udy scope: ecosys em se ices p o ided by s anded ce acean ca casses. Secondly, we de ined ou sea ch s ing (see Table B1) and used Web o Science o sea ch o a icles (published un il 6 h Augus 2020) con aining hese keywo ds in he en i e a icle. In summa y, ou sea ch p o ocol used he e ms “humans” and “ecosys em se ices” in combina ion wi h a ious synonyms o ce acean ca casses (i.e. “s anded whale”, “whale ca cass”, “s anded ce acean”). This yielded 484 a icles (Table B1), including epea ed a icles esul ing om di e en sea ches. Thi dly, we es ic ed he sea ch o pee - e iewed a icles. Then, we sc eened i les and ab- s ac s o iden i y hose a icles ha deal wi h he ecosys em se ices p o ided by ce acean ca casses, and we ully ead he selec ed a icles. This led o a inal selec ion o 27 a icles dealing wi h se ices p o ided by ce acean ca casses (Table B1). Fou hly, we ex ac ed he ollowing in o ma ion om he selec ed a icles: 1) ecosys em se ices men ioned (sepa a ing hem in o p o isioning, egula ing, cul u al and suppo ing se ices), 2) ce acean species o g oup o which he ecosys em se ices we e ela ed, 3) s udy a ea, and 4) s udy pe iod (p ehis o ic, his o ic –i. e. be o e he s a o he 19 h cen u y when he onse o comme cial whaling occu ed– and mode n –i.e., a e he s a o 19 h cen u y–). Fi hly, we ca ego ised he selec ed a icles acco ding o hese o me ac o s (ecosys em se ice ype, ce acean g oup, a ea and pe iod). Spe- cies we e g ouped acco ding o amilies and pa o de s o Ce acea: a) baleen whales, b) oo hed whales (excluding small- oo hed whales), and c) dolphins and po poises, which we e sepa a ed om he o he oo hed whales because o hei smalle body size and gene ally highe abundance. 2.2. Da a collec ion on s andings and egula ions We e ie ed da a on ce acean s anding e en s om 29 coun ies o egions (Table B3) and egula ions om 23 o hem (Table B4), om an ini ial lis o 67 coas al coun ies in all con inen s ha we e conside ed and asked o in o ma ion. Da a on s andings and egula ions we e di ec ly sha ed ei he by he ma ine mammal ne wo k es ablished in hese coun ies o by scien is s wo king on ce acean s andings in di e en academic o ganisa ions, go e nmen al ins i u ions o inde- penden socie ies. In addi ion, we ex ac ed complemen a y in o ma ion om local g ey and p ima y scien i ic li e a u e. We selec ed s anding da ase s ha we e ep esen a i e o he species occu ing in each s udy a ea and co e ed mul iple yea s. Reco ds o ce aceans ha s anded as ca casses, died a e s anding o we e unsuccess ully e- loa ed ( e- s anded and died) we e g ouped as men ioned abo e: a) baleen whales, b) oo hed whales (excluding small- oo hed whales), c) dolphins and po poises, and d) uniden i ied ce aceans. Fo each g oup, he o al numbe o s andings was di ided by he numbe o yea s co e ed by he da ase and he coas line leng h (km) o he e i o y o coun y (see mo e de ails in Table B3). Few da ase s also p o ided in o ma ion ela ed o he a e o he s anded ca casses in e ms o disposal, which will be summa ised in he Resul s. 2.3. Analysis o s anding da a To unde s and i s anding epo ing a e in each egion could be explained by he popula ion densi y and exis ence o s anding managemen policy o egula ions, we used he 23 egions o which we we e able o ga he in o ma ion on he exis ence o managemen policy o egula ions (Table B4), coas leng h and human popula ion densi y. The exis ence o s anding managemen policy o egula ions was coded as: “yes”, i p esen and known; “no”, i no p esen and known; and “na” o unknown ( i e egions). We used he decimal loga i hm o human popula ion by coas al km 2 as a p oxy o human coas al popula ion densi y. We pe o med model selec ion using he Akaike’s in o ma ion c i e ion (AIC; Bu nham and Ande son, 2004). We conside ed models o be equi alen when he di e ence in AIC wi h he bes model was ΔAIC <2 (Bu nham and Ande son, 2004). All we e Gene alized Linea Mixed models (GLMMs) buil in R (R Co e Team, 2013) using a Gaussian link unc ion. 3. Resul s 3.1. Ecosys em se ices p o ided by ce acean ca casses Ou li e a u e e iew showed ha ce acean ca casses ha e mainly supplied p o isioning se ices, ollowed by cul u al, suppo ing and egula ing se ices (Fig. 3; Table B2). Mos a icles e e ed o mode n imes, hough hose men ioning p o isioning se ices mos ly ela ed o p ehis o ic imes (Fig. 3). In ou sea ch, he keywo d “human” was included in mo e a icles han “ecosys em se ice”. This e eals ha , despi e he scien i ic li e a u e acknowledging he associa ion be ween humans and ce acean s andings, li le o his associa ion is explained in e ms o ecosys em se ices. He e, we p esen an o e iew o he his- o ical and cu en ole o ce acean s andings as p o ide s o ecosys em se ices o humans by compiling he examples p oduced by ou li e - a u e sea ch and u he pe inen a icles (see also Panel 1). Ce acean ca casses p o ided ood and aw ma e ial o ea ly ci ili- sa ions wo ldwide (Fig. 4). Fo ins ance, emains o wo ba nacle spe- cies, o iginally a ached o he body o sou he n igh whales (Eubalena aus alis), we e ound in a ca e a a p ehis o ic si e in sou he n Spain, sugges ing ha p imi i e humans used o consume s anded whale mea (´ Al a ez-Fe n´ andez e al., 2014). Bone agmen s belonging o ce acean specimens we e also iden i ied in he Mesoli hic deposi s o o he p e- his o ic ca es (e.g., G o a dell’Uzzo, I aly; Mannino e al., 2015). The disco e y, in Angola, o nume ous ools made om whale bone, such as choppe s and lakes, indica ed ha many componen s o ca casses we e used in he Lowe Palaeoli hic (Gu ie ez e al., 2001). In he p e-Columbian e a, indigenous people, such as A awak and Ca ib g oups om Ba bados (Ca ibbean), also consumed s anded dol- phins and spe m whales (Physe e mac ocephalus; Rome o and C eswell, 2010). S anded ca casses ha e been o c ucial impo ance o he su - i al and cul u al de elopmen o o he adi ional coas al communi ies, such as he Inui li ing in he A c ic egion (Ka ona and Whi ehead, 1988), he “canoe people”, and he Fuegians, inhabi ing he sou he n- mos Pa agonian coas . The la e conside ed s anded ce aceans a “g ea Fig. 3. Resul s o he sys ema ic li e a u e e iew on he ecosys em se ices associa ed wi h ce acean ca casses, acco ding o se ice ype and pe iod o ime. Ring size is p opo ional o he numbe o a icles ound in he e iew. Fo mo e de ails, see Table B2. M.-M. Quaggio o e al. Ecosys em Se ices 54 (2022) 101406 4 gi o na u e” o be sha ed wi h he en i e communi y (Qui oz e al., 2016), whe eas Aus alian Abo igines and islande s om he To es S ai belie ed ha s andings could connec hem o ances al lands, seas, and hei ances o s (Ha cou e al., 2014). Simila ly, in New Zealand, M¯ ao i ega ded whale s andings as a symbol o abundance and ichness because hey we e g ea sou ces o p o ein and ma e ial: hese esou ces we e ex ac ed ollowing s ic p o ocols o espec hei spi i ual impo ance (Rodge s, 2017). In he la e medie al pe iod, bo h Anglo-Saxons and No dic coas al communi ies used o exploi s anded ma ine mammals (B i o e al., 2011; Pa sons and Monaghan-B own, 2017). Since he ele en h cen u y, ce aceans s anded on English sho es ha e been known as “Royal Fish”, ollowing a s a u e enac ed by Edwa d II (Ga dine , 1997). The ce a- ceans’ lesh was consumed by weal hy ci izens whe eas he emains o he ca cass we e ende ed o p oduce oil and he bones used as u ensils o a is ically ca ed (e.g. he F anks Caske which is a small deco a ed box made o whale bone om he 8 h cen u y, ep esen ing Anglo-Saxon a ). Fo he medie al English, whales we e “sea-mons e s” and ac- co ding o he popula supe s i ion, hei s andings p edic ed he imminen occu ence o ex ao dina y e en s (Ga dine , 1997; Fig. 4). As humans de eloped owa ds mode n socie y, hei ela ionship wi h ma ine mammals also changed. P o isioning ecosys em se ices a ibu able o s andings we e slowly subs i u ed o cul u al se ices (Fig. 4). S a ing om he 18 h cen u y, s anded ca casses we e s udied and La in names we e gi en o ce acean species. S anded ca casses on beaches, pa icula ly o la ge ce aceans, also equen ly gene a ed high public in e es (Sousa and B i o, 2012). F om he 20 h cen u y, i became common o scien is s o collec s anded specimens o imp o e hei unde s anding o hese aqua ic mammals (Fig. 4). The skele ons we e collec ed by museums o public exhibi ion, educa ion (Wei and Pie ce, 2013) o u u e esea ch (Bo - ella and Bo e o, 2017). Ce aceans and hei ca casses a e now iewed as indica o s o ocean heal h, and s andings p o ide impo an in o ma- ion ha enhances ou scien i ic unde s anding o ma ine ecosys ems and he e ec s o an h opogenic impac s (Bossa , 2011). Recei ing much a en ion om he media and he public, s andings o la ge ce acean specimens became sou ces o sus ainable en i onmen al edu- ca ion. In addi ion, he wo k o olun ee –based o ganisa ions esponding o ma ine mammal s andings is o g ea alue in aising public awa eness, which can po en ially enhance animal wel a e and conse a ion (Roman e al., 2018). Whale ca casses a e also p o ide s o suppo ing se ices, such as subsidies o sca enge s. They g ea ly con ibu e o nu ien cycling and ca bon seques a ion, ep esen ing – in he case o whale alls – a po- en ial sink o an h opogenic ca bon ha may be impo an o mi i- ga ing global clima e change (Pe shing e al., 2010; see Panel 1). 3.2. S andings a ound he wo ld Mean densi y o s anding pe coun y/ egion was 0.09 s andings • yea −1 •km −1 (SD =0.21). No he n Te i o y (Aus alia) had he lowes annual densi y o s anding e en s (0.001 s andings •yea −1 • km −1 ), whe eas Belgium he highes (0.978 s andings •yea −1 •km −1 ; Fig. 5). In gene al, as expec ed om hei ela i e abundances (Ham- mond e al., 2013), dolphins and po poises we e he mos abundan g oup ac oss all coun ies (Fig. 5), wi h he excep ion o A gen ina and Chile, whe e baleen whales domina ed he eco ded s anding e en s (Figu e B.1). Rega ding he GLMMs explo ing i he s anding densi ies eco ded could be p edic ed by he p esence o managemen policy o human popula ion densi y ac oss he s udied a eas, he se o bes compe ing models (see Table B5) comp ised he model including he e ec o human densi y (M 1 ) and he model including he e ec o legisla ion p esence (M 2 ). Howe e , hese models we e no signi ican ly di e en om he cons an model, and hei e ec s we e no signi ican (i.e. none o he be as was signi ican ly di e en om ze o; see Table B5 and Figu e B.2). Thus, nei he o he abo emen ioned ac o s we e good p edic o s o eco ded s anding densi ies. 3.3. Regula ions a ound he wo ld The implemen a ion o egula ions o ca cass managemen is a Fig. 4. Examples o ecosys em se ices associa ed wi h ce acean ca casses. P o isioning se ices: p oduc ion o ools (baleen pla es used as b oom; a), mea (b), e ilize s (c) and oil (oil dish o whale oil; d). Regula ing se ices: CO 2 seques a ion in he o m o loa ing ca casses be o e sinking (e) be o e unde going nu ien ecycling h ough ac ion o sca enge s and decompose s ( ). Cul u al se ices: scien i ic ad ancemen in he pas (g) and p esen (h); p oduc ion o a (F ank Caske , an Anglo-saxon box made o whalebone; i); educa ional alue in ou doo (j) and indoo museums (m); eco ou ism (k); and eligious signi icance (l). See pho o c edi s in Appendix A. M.-M. Quaggio o e al. Ecosys em Se ices 54 (2022) 101406 5 ecen and ongoing phenomenon. Ou o he 23 s udy coun ies/ egions, i e did no ha e egula ions ela ed o s andings (22%; Table B4). The lack o legisla ion was jus i ied by he ins i u ions in ol ed in s anding managemen because he a eas we e emo e and had low human pop- ula ion densi y (e.g. New oundland and Lab ado in Canada and No he n Te i o y). The absence o a s anding esponse ne wo k and a compe en na ional au ho i y also de e mined he lack o egula ions (e. g. I eland, which, howe e , has an e icien olun ee ne wo k; IWC, 2016). In some coun ies/ egions, o icial pe missions a e needed o handle s anded ce aceans in dis ess, o hei ca casses, and speci ic s akeholde s a e esponsible o he in e en ion and he ca casses’ a e. Se e al s akeholde s a e in ol ed in hese p ocesses. These may include local go e nmen s, such as he ci y council and he municipali y, and/o land manage s, academic ins i u ions, s anding ne wo ks, local NGOs and museums. Thei ole in he managemen o ca casses a ies om coun y o coun y. In some coun ies, like New Zealand, indigenous people a e also in ol ed in he p ocess: o ins ance, M¯ ao i would p ay o he dead ce aceans be o e hei disposal (Bu e wo h, 2017). P o- ocols may be implemen ed o guide and acili a e he p ocess o ca cass disposal (Table B4). This is he case in New Sou h Wales (Aus alia), whe e he Na ional Pa ks and Wildli e Se ices de eloped a checklis o guide land manage s when disposing o whale ca casses (NPWS, 2020). A wide spec um o ca cass disposal me hods exis s, om comple ely o ela i ely na u al p ocedu es, such as na u al decomposi ion in si u o he whole ca cass o in pieces, bu ial o he ca cass, owing ou o sea (o sho e), o land (insho e) o compos ing, o me hods ha in ol e Fig. 5. Ce acean s anding densi y map (N s andings •yea −1 •km −1 ) and absolu e abundance o s andings pe g oup o ce aceans (Baleen whales o Mys ice i; Too hed whales o Odon oce i, excluding small oo hed whales like dolphins and po poises; Dolphins and Po poises; and Uniden i ied ce aceans) o all s udy coun ies/ egions. Fig. 6. Di e en me hods o s anded ce acean ca cass disposal and associa ed ecosys em se ices and cos s, o de ed acco ding o hei equency in egula ions o coun ies/ egions conside ed in his s udy. M.-M. Quaggio o e al. Ecosys em Se ices 54 (2022) 101406 6 mo e in e en ion, such as disposal a a local land ill, incine a ion o ende ing (Fig. 6). In Wes e n Aus alia, mos ca casses a e bu ied (30%), and only 8% o hem a e le in si u. Excluding he ca casses whose a e is unknown, in New Zealand and USA mos o he ca casses a e le on si e o bu ied (NZ: 39% and 38%; USA: 28% and 21%, espec i ely). Also, in New Zealand, mass s andings ha e o ced au- ho i ies o use na u al ide d i en decomposi ion. Ca cass ans- po a ion o land ills o incine a ion plan s is common in many Eu opean coun ies. Fo ins ance, in Belgium and F ance, which a e cha ac e ised by high densi ies o s andings, he only op ion is o des oy ce acean ca casses in app op ia e acili ies. Mul iple disposal me hods a e allowed in o he Eu opean coun ies (e.g. Spain). In USA, he hi d mos used me hod is he land ill (18%). Specimens ha will be used o scien i ic and educa ional pu poses a e ei he collec ed, bu ied o consumed by sca enge s p io o eco e y o he skele on. In Wes e n Aus alia and New Zealand, only a small numbe o ca casses a e collec ed o scien i ic (WA: 0.7%; NZ: 1.3%) o educa ional (WA: 1.2%) pu poses. When ca casses a e emo ed, hey a e o en nec opsied, so hey s ill may p o ide a scien i ic and e e ina y se ice. Howe e , in some cases, he nec opsy pe se is he ul ima e goal, which usually means mo ing he ca cass o a wildli e escue cen e, whe e i is elimina ed o cleaned o ob ain he skele on. Nec opsies o la ge whales a e no mally done in si u. The me hod o ca cass disposal is con ex -dependen in 45% o coun ies/ egions, as i may change depending on he exac loca ion whe e he animal s ands (Table B4). Gene ally, when he ca cass is ound in a emo e o di icul o access a ea, he common p ocedu e is o le i decompose na u ally, whe eas i s emo al is conside ed p e e able when occu ing in a popula ed o ou is ic a ea. This is he case in New Sou h Wales and Wes e n Aus alia (Aus alia; NPWS, 2020; DPAW). Howe e , his choice is no con empla ed when he emo al o he ca cass is compulso y (F ance and Belgium), o he e a e no egula ions in place because hey a e conside ed unnecessa y (New oundland and Lab ado , and Falkland Islands). Regula ions a y when s andings occu in p o ec ed a eas, depending on he po en ial con amina ion and impac on he exis ing lo a and auna. Special conce ns a ise when s anded la ge whales a e eu hanized, as oxic chemicals used in some me hods may pose a possible isk o sca enge s and he su ounding en i onmen ( elay oxici y), po en ially a ec ing humans nega i ely (Ha ms e al., 2014). In hese cases, ca casses a e always emo ed o app op ia e disposal (e.g. UK). In he case o Denma k, emo al o he ca cass is no loca ion-, bu species-dependen , as only ha bou po - poises a e le a si e o decompose. 4. Discussion Ancien and mode n ci ilisa ions wo ldwide ha e bene i ed om he ich and a ied a ay o p o isioning, egula ing, cul u al, and sup- po ing ecosys em se ices associa ed wi h s anded ce acean ca casses. F om Palaeoli hic A icans and Eu opeans o p e-Columbian Ame icans, and om bo eal Inui s and Icelande s o Paci ic M¯ ao i, p e-indus ial humans li ing in o nea he coas iewed s andings as a gi om na- u e (e.g. h al eki in Icelandic, meaning “wind all”, Fielding, 2018), and ce acean ca casses we e widely used o ood and as sou ce o aw ma- e ial o hun ing and bu che ing ools and o namen s (e.g. Ga dine , 1997). Ce aceans also p oduced a deep spi i ual and a is ic inspi a ion (e.g. Ha cou e al., 2014). Mo e ecen ly, cul u al se ices became he dominan bene i s associa ed wi h s anded ce aceans (as happened wi h o he aunal g oups; e.g. Mole´ on e al., 2014), including scien i ic dis- co e y, in si u and ex si u educa ion, public awa eness aising, and eco ou ism. O e all, o many ancien and mode n people, ce acean ca casses signi y an appa en mani es a ion o na u e’s magni icence. All his e eals he impo an cul u al he i age ha humani y has con- s uc ed a ound s anded ce aceans. In addi ion, h ough he key ecological e ec s o ce acean ca casses, which pe ade well beyond he coas al sys ems and may las o yea s, s andings ha e also led o ele an suppo ing se ices o pas and p esen ecosys ems. Howe e , cu en ly, ecosys em se ices p o ided by s anded ce aceans a e g ea ly h ea ened by bo h he esul s o educ ion in ce acean popula ions due o hun ing (see Panel 3) and mode n policies ega ding he disposal o hese ca casses. F om he In e na ional Whaling Commission 1982 mo a o ium on comme cial whaling, he numbe o coun ies adhe ing o he In e na ional Con en ion o he Regula ion o Whaling (1946) inc eased yea a e yea (IWC). In addi ion o his in e na ional ag eemen , many coun ies ha e pu o wa d na ional legisla ion o p o ec ma ine mammals, including hei emains. As a esul , he managemen o any ype o ope a ion in ol ing ma ine mammals ( om i s in e en ion o disposal o he ca cass) is o en egula ed a na ional o in a-na ional le els. Because o he na u al ecu ence o s anded animals on he coas , hese egula ions a e indispensable o no only he conse a ion o ce aceans and hei ull ecological ole, bu also human well-being. Decomposing ce acean ca casses a e a po en ial sa e y haza d o human heal h, and a nuisance o some locals and beach use s when occu ing in popula ed a eas (Tucke e al., 2018). Thus, he sys ema ic emo al o ca casses is o en applied o a oid heal h- ela ed isks and/o people’s malcon en . The la e could be conside ed he main disse ices ha s anded ce acean ca casses p o ide o humans. A oiding hese de imen s, howe e , o en equi es complex logis ics and impo an economic cos s, which could also be iewed as a so o disse ice. Fo ins ance, in Aus alia, he beach bu ial o a whale can each up o app oxima ely $18,000 (USD) when including all he cos s o he ma- chine y, ope a ions and consul a ion wi h s akeholde s. The anspo and des uc ion o a small whale a a dedica ed acili y, ins ead, is a ound $15,000 (USD), wi h cos s a ying acco ding o ca cass size (Tucke e al., 2018). Mo eo e , as he coas al a eas become “cleaned” and he coas al ecosys ems “asep ic” o ul il hese human necessi ies, he s uc u e o na u al communi ies is des abilised and hei ecological unc ionali y los (Huijbe s e al., 2013), and he p o ision o o he human demands such as aes he ic enjoymen , educa ion and o he cul- u al se ices is comp omised. Thus, unde s anding he ecosys em se - ice po en ial associa ed wi h ce acean s andings is he i s s ep o ully ecognise and es ablish he di ec and indi ec bene i s o human socie y and i s needs (Malinauskai e e al., 2021a). 4.1. Policy and managemen ecommenda ions S anded ce acean ca cass managemen will be an inc easing global ma e as coas al a eas become mo e popula ed, new egula ions a e app o ed, and ce acean popula ions eco e (Clapham e al., 1999; see Panel 3). In addi ion, clima e change and popula ion shi s may po en- ially lead o s andings in egions which may no ha e p e iously encoun e ed hem, and s andings may in ol e di e en species han p e iously seen (Williamson e al., 2021). Thus, managemen s a egies may need o adap in esponse. One way o maximise he bene i s de i ed by ce acean ca casses is iden i ying hose disposal me hods ha can p ese e and/o enhance ecosys em unc ions and se ices. All he exis ing disposal me hods can suppo ecosys em se ices, bu some be e han o he s (Fig. 6). Man- agemen o ce acean ca casses needs o balance hei ecological impo - ance, he ecosys em se ices hey may p o ide, cos s o managemen ac ions, biodi e si y conse a ion egula ions and legal and public heal h equi emen s (Fig. 6). When le in si u, a ce acean ca cass can p o ide all he ypes o ecosys em se ices, wi h e y low managemen cos s, bu social discon en may ise. Bu ial is a cos -e ec i e op ion; bu i he ca cass is bu ied close o he sho eline, he nu ien en ichmen o he su ounding en i onmen may con amina e he g ound wa e s and po en ially a ac sha ks o he sho e, which may diminish he ec ea- ional use o he beach (Tucke e al., 2018; Tucke e al., 2019a; Heiss, 2020). Re u ning he ca cass o he sea by owing i would allow i s na u al decomposi ion and p o ision o egula ing and suppo ing se - ices. Howe e , a loa ing ca cass could also become a sa e y haza d in M.-M. Quaggio o e al. Ecosys em Se ices 54 (2022) 101406 7 a eas o high ma i ime a ic (Tucke e al., 2018). The ull emo al o s anded ce aceans om he coas al a ea leads o he loss o ecosys em se ices and inc ease o he cos s ela ed o hei anspo and disposal. In pa icula , incine a ion is mo e expensi e han o he me hods and is a sou ce o g eenhouse gas emissions (Mo ales-Reyes e al., 2015). Among he op ions equi ing he emo al and anspo o ca casses, compos - ing cos s (pe uni weigh ) hal o he p ice o he disposal in a land ill o incine a ion plan (King e al., 2018). Mo eo e , his op ion comes wi h en i onmen al bene i s oo, e.g. imp o ed soil quali y and b eak-down o con aminan s and pollu an s (King e al., 2018). Gi en he p ominen impo ance o s anded ce acean ca casses o endange ed sca enge s in some a eas/ egions (Chambe lain e al., 2005; Lambe ucci e al., 2018, see Panel 2) and ecosys em unc ions (Smi h and Baco, 2003; Bui, 2009; Smi h e al., 2015), we encou age na u al decomposi ion in si u, p o ided ha legal and sa e y needs a e ul illed. This measu e, which is he cheapes one, can ob iously be easily applied in emo e coas s and/o p o ec ed a eas, as some coun ies al eady do. In mo e popula ed and ou is ic a eas lea ing ca casses in si u may aise p oblems o sigh , smell, changes in sedimen and g oundwa e chem- is y, and sha k and e al dog a ac ion (Tucke e al., 2018). In hese cases, managemen may be season- and/o loca ion-dependen . When s andings ake place du ing he low- ou ism season, ca casses may be le in si u. When s andings occu in he high- ou ism season, a empo al es ic ion, co e ing he main decomposi ion pe iod, could be applied o he icini y o he ca cass. The implemen a ion o zona ion maps iden- i ying emo e and u ban beaches could also be implemen ed in o de o acili a e he decision making. Al e na i ely, ca casses could be chopped in o smalle pa s o accele a e hei consump ion o decomposi ion. Ou analysis o s andings a ound he wo ld e ealed ha no all coun ies o egions will ace he same challenges, as bo h he densi y and size o he s anded ce aceans we e highly spa ially he e ogeneous. In gene al, hose coun ies wi h he highes densi ies o s anded ce a- ceans and mo e es ic i e egula ions o ca cass emo al (e.g. F ance, Belgium and he Ne he lands) mos ly ecei ed small-sized ce aceans. A sensible policy migh ake in o accoun bo h he size and numbe o ca casses. The disposal and anspo o smalle ca casses gene ally cos less, bo h in e ms o manpowe , logis ics and o e all inancial cos . In a coun y whe e s andings a e domina ed by small ce aceans such as dolphins and po poises, a p opo ion o ca casses may, he e o e, be chosen o be le in si u, o anspo ed o a mo e emo e a ea wi h limi ed cos s. I bo h small and la ge whales occu a a simila ex en , hen logis ics may be eased and cos s educed by emo ing he small size ca casses, whe eas la ge ca casses could be chopped in si u a e nec- opsy a oiding anspo cos s, bu po en ially inc easing s a e o s. Finally, we ecommend ha he pas and p esen alue o s andings, and ce acean ca casses in gene al, a e included in educa ional ini ia- i es. In emo e a eas, like in he A c ic egions, in o ma ion on whale ca casses can be inco po a ed as an educa ional esou ce o he locals and used o p omo e he inc easing ou is indus y alongside wi h he one exis ing on li ing whales (Ge aci and Lounsbu y, 1993; Mali- nauskai e e al., 2021a). This, and o he eco ou ism expe iences, could be enhanced and expo ed o o he places wo ldwide, as sca enge s p o ide impo an ec ea ional, aes he ic, lea ning and inspi a ion ex- pe iences o people (Aguile a-Alcal´ a e al., 2020). An inc eased public awa eness o he ecological and cul u al ole o ce acean s andings could help educe nega i e a i udes owa ds hese phenomena and hus balancing ecosys em se ices and disse ices. 5. Conclusions and u u e esea ch We ha e p o ided a comp ehensi e o e iew ha could be used o aise public awa eness o he bene i s ha ce acean ca casses ha e p o ided o ou ances o s, and he se ices om which mode n humans could bene i . Thus, u u e s eps should iden i y he main social ac o s in ol ed and e alua e he public pe cep ion o s andings and associa ed se ices, which could e en ually imp o e cu en egula ions on ca cass disposal (Malinauskai e e al., 2021b). Fu he , we encou age he implemen a ion o inno a i e science ou each ia ci izen science and educa ion o os e enhanced local ecological knowledge and app ecia- ion o ecosys em se ices p o ided by ce acean ca casses, which may lead owa ds mo e sus ainable p ac ices. Finally, bo h use and non-use alues o ce acean ca casses should be es ima ed in di e en local so- cial con ex s by in eg a ing ma ke and non-mone a y alua ions o each ecosys em se ice p o ided (Cook e al., 2020). We ecommend enhancing he p esen coas al managemen s a e- gies by widening hei ecosys em pe spec i e by including zoning, sea- sonal use limi a ion and science ou each. Undoub edly, hese managemen ac ions need o be suppo ed by a be e scien i ic knowl- edge o ce acean s andings – and popula ions. Long- e m moni o ing o s anding e en s, especially i coo dina ed a he sup ana ional le el, should be a basic componen o he conse a ion o ma ine mega auna and he ecosys em se ices hey p o ide. These p og ams may supply de ailed da a on ce acean s anding numbe s and hei spa io empo al dis ibu ion, which is ele an o e ine ou unde s anding o he ac o s a ec ing s anding dynamics and o imp o e he e iciency o ca cass de ec ion and decision making. These da a and knowledge a e also essen ial o cla i ying he ela ionship be ween s anding e en s and mul iple explana o y ac o s in addi ion o human densi y and p esence o egula ions he e explo ed. Impo an ly, his in o ma ion should be made easily accessible o esea che s. In addi ion, u he e alua ion o he ecological impac o he di e en managemen echniques o s andings and compa ing hei cos s and bene i s is essen ial (Tucke e al., 2018). Only h ough a deepe scien i ic knowledge o he socio- ecological con ex ha cha ac e izes he ca casses o s anded ce a- ceans, we will be able o delinea e he mos e icien managemen s a egies in coas al a eas in an inc easingly popula ed (by bo h humans and ce aceans) wo ld. Panel 1. Ecological e ec s o ce acean ca casses. Whale ca casses a e a signi ican sou ce o ca ion in he ma ine sys em (Baco and Smi h, 2003), ep esen ing a huge ood supply o sca enge s and decompose s, bo h ma ine and e es ial (Fig. 1). “Whale alls” a ac and sus ain a di e se and abundan deep-sea communi y o e eb a e and in e eb a e sca enge s. By p o iding in e media e habi a s o be colonised, whale ca casses link chemosyn- he ic communi ies oge he , enhance he s uc u al complexi y o he seabed o yea s (Smi h and Baco, 2003) and p omo e adap i e adia ion o ce ain o ganisms such as he bone-ea ing wo m (Osedax spp.; Smi h e al., 2015). Mos dead whales sink o he bo om, bu abou 10% may become buoyan because o gas p oduced du ing decomposi ion (Pyenson, 2010). While loa ing, ca casses become an easy and ene gy- ich sou ce o ood o o cas and sha ks (Whi ehead and Ree es, 2005; Lecle c e al., 2011; Tucke e al., 2019a). O hese, some ca casses e en ually become s anded on he sho e (Roman e al., 2014). The ecological impac o hese s anded ca casses on he coas al sca enging communi y is subs an ial, a ac ing a wide ange o ma ine and e es ial sca enge s and elici ing nume ical and/o unc ional esponse in consume s (Polis e al., 1996; Fig. 1). Gian pe els (Wa ham 1990), skuas (L¨ onnbe g 1906) and pola bea s all sca enge on s anded ce aceans, p o iding he a and p o ein necessa y o cope wi h ex en- si e pe iods o as ing (Laid e e al., 2018). Whale ca casses may ha e suppo ed pola bea s in pas wa me in e glacial pe iods, when seal a ailabili y was limi ed (Laid e e al., 2018), and hus, may play a c ucial ole in hei adap a ion o ongoing clima e change. A mo e empe a e la i udes, e es ial sca enge s such as b own bea s, wol es (Lewis and La e y, 2014), oxes (Ka ona and Whi ehead 1988), hyenas and jackals (Skinne e al., 1995) also mo e o he coas o eed on dead ma ine mammals. The ex inc Cali o nia g izzly bea is epo ed o ha e ed upon salmonids and s anded ma ine mammals (Laid e e al., 2018). In addi ion, condo s, which a e obliga e sca enge s, la gely specialised on s anded whales (Chambe lain e al., 2005; Lambe ucci e al., 2018; o mo e de ails, see Panel 2, Fig. 2). Finally, simila ly o whale alls, s anding e en s a e likely o en ich subs a a wi h nu ien s (as happens M.-M. Quaggio o e al. Ecosys em Se ices 54 (2022) 101406 8 wi h bu ied ce aceans; Bui, 2009) and be discha ged in o coas al wa e s po en ially a ac ing sha ks (Tucke e al., 2019b; Heiss, 2020). These nu ien s may also p o ide supplemen s o algae and plan s in he su - ounding en i onmen , as well as o inc ease landscape he e ogenei y. Panel 2. Ecological and conse a ion consequences o ma ine ca cass deple ion: he case o condo s. Ce acean s andings p o ide long-las ing pulses o ood, which may de e mine local changes in dis ibu ion and abundance o sca enging o ganisms and popula ions, as well as changes in he ene gy lux o ood webs. Whales in pa icula we e an impo an ca ion sou ce o sca - enge s in his o ic and p ehis o ic imes. Pleis ocene mega aunal ex inc ion s ongly impac ed e es ial mammals and, o a lesse ex en , ma ine mammal popula ions. A e he la e Pleis ocene e es ial mega aunal ex inc ion in No h Ame ica, he Cali o nia condo (Gym- nogyps cali o nianus) became ex inc in he a eas whe e hey ed mos ly on e es ial animals. The condo ’s dis ibu ion ange was es ic ed o he Paci ic coas al a ea, whe e hey exploi ed he abundan ca ion e- sou ces p o ided by s andings (Chambe lain e al., 2005). The se e e deple ion o ma ine mammals du ing he whaling e a o ced condo s o shi back o e es ial ood esou ces in he la e 1700s, which has caused a se ious conse a ion p oblem due o lead poisoning om bul- le s ound in he e es ial ca casses (Chambe lain e al., 2005). In Pa agonia, Sou h Ame ica, he Andean condo (Vul u g yphus) has adi ionally nes ed on he Paci ic slope o he Andes, allowing e icien exploi a ion o he ce acean ca ion ha was once abundan on he coas (Lambe ucci e al., 2018; Fig. 2). Those b eeding a eas close o he Paci ic Ocean a e main ained e en oday, when ew whale ca casses a e a ailable on he wes e n Pa agonian coas . Thus, condo s a e o ced o ou inely c oss he Andes o he eas o sca enge on ungula e ca casses om he Pa agonian s eppe, inc easing he ene gy expendi u e and po en ial isks ha h ea en his species (Lambe ucci e al., 2018). Popula ions o bo h condo species dec eased un il almos he ex inc ion in high isk a eas associa ed wi h human impac s. These h ea s include sa e na i e wild ood educ ions, and he inc easing use o an h opogenic ood sou ces (Plaza and Lambe ucci, 2020). In he pas , be o e he se e e declines in whale popula ions, s andings could ha e been so common ha coas al ecosys em s uc u e and unc ion may ha e been di e en om hose known oday, and hei e ec s could ha e e en pe mea ed inland. Unde his scena io, he his o ic lack o whale s andings has played an impo an ole in he loss o sa e abun- dan ood sou ces o condo s, and po en ially many o he sca enge s. Panel 3. The global decline o ce acean popula ions. Hun ing o whales, especially baleen whales (subo de Mys ice i) and he spe m whale (Physe e mac ocephalus; subo de Odono oce i), has been p ac iced o cen u ies by some ci ilisa ions such as he Jap- anese and Inui (Shoemake , 2005). Mo e ecen ly, whaling s a ed o be p ac iced also ac oss di e en popula ions such as he Basques in he 11 h cen u y, No semen in he ea ly medie al pe iod, and Du ch, Ca- ibbeans and Eu opean colonis s o Ame ica in he 15 h cen u y (Rome o and C eswell, 2010; Pa sons and Monaghan-B own, 2017). Whaling began in S alba d in 1611 and las ed abou h ee hund ed yea s. By a ound 1850, he G eenland igh whale was comple ely emo ed om ma ine ecosys ems a ound S alba d (Hacquebo d, 2001). The ad en o mode n, comme cial whaling wi h mo e sophis ica ed essels and he use o explosi es (la e 1800; Ellis, 1991) pe mi ed whale s o ca ch la ge and as e ce acean species (Pa sons and Monaghan-B own, 2017). Wi h he expansion in o An a c ic wa e s and he es ablishmen o whaling s a ions, whaling eached i s his o ical peak in he ea ly 20 h cen u y. The lack o sus ainabili y o his indus y, which d as ically educed whale s ocks a a global scale, led i s o he signing o he Con en ion o he Regula ion o Whaling (Sep . 24, 1931, 155 L.N.T.S 349) in Gene a o An a c ic whale ishe y in 1931, and e en ually he In e na ional Whaling Commission (IWC) o ake ac ion and egula e whale ca ches as ea ly as 1946 (IWC). In 1982, a en-yea mo a o ium was decided and hen applied in 1986 o pe mi whale popula ions o eco e , bu a he end o i , he pause in comme cial whaling was ex ended o inde ini e ime (IWC). Cu en ly he e a e 88 pa icipan membe go e nmen s, as Japan wi hd ew i s membe ship in 2013 and esumed whaling in 2019. In addi ion, No way is s ill hun ing minke whales (Balaenop e a acu o os a a) o in e nal consump ion and expo o Japan (Lecle c e al., 2011). Iceland and Fa oe Islands also con inue o p ac ice whaling. Abo iginal subsis ence whaling is also cu en ly allowed in G eenland, Chuko ka, Bequia and Alaska, al hough i s e ec on whale popula ions is conside ed negligible (Clapham e al., 1999). Wi h millions o ce aceans killed (Clapham e al., 1999), la ge-scale comme cial whaling is one o he g ea es episodes o unsus ainable wildli e exploi a ion in human his o y (Clapham e al., 2008). As a esul , se e al species we e deple ed almos o ex inc ion, al hough some sign o eco e y ha e been obse ed in some popula ions in ecen de- cades (Clapham e al., 1999). Because o he slow popula ion g ow h a e cha ac e ising hese K-s a egy animals, some whale species (B. musculus, B. physalus and Eubalena aus alis) will ake mo e han a cen u y o eco e o hal o hei o iginal p e-exploi a ion numbe s, al hough o he s may eco e soone (e.g. Megap e a no aeangliae; Tul- loch e al., 2018). The cascading e ec s o he huge dec ease in whale popula ions ha e p o oundly al e ed he unc ioning o whole ecosys- ems (e.g. Hacquebo d, 2001). Al hough i has no been in es iga ed, i can be assumed ha he numbe o s anded ce aceans has educed in p opo ion o he dec ease in li ing whales. The diminished a ailabili y o ce acean ca casses has almos ce ainly also a ec ed he p o ision o ecosys em se ices and unc ions. The ca bon seques a ion h ough sinking ca casses, o ins ance, would be an o de o magni ude g ea e i whale popula ions we e es o ed o p e-whaling numbe s (Pe shing e al., 2010). Decla a ion o Compe ing In e es The au ho s decla e ha hey ha e no known compe ing inancial in e es s o pe sonal ela ionships ha could ha e appea ed o in luence he wo k epo ed in his pape . Acknowledgemen s This s udy was suppo ed by he Spanish Minis y o Economy, In- dus y and Compe i i eness and EU ERDF unds h ough he p ojec s CGL2015-66966-C2-1-2-R and CGL2017-89905-R. M.M. was suppo ed by a esea ch con ac Ram´ on y Cajal om he MINECO (RYC-2015- 19231), and A.CA. by a con ac Juan de la Cie a Inco po aci´ on (IJCI- 2014-20744; Spanish Minis y o Economy and Compe i i eness, Spain) and a Pos Doc con ac P og ama Viçen Mu o Go e n Balea , Spain (PD/039/2017). NS was suppo ed by he Bekke p og amme o he Polish Na ional Agency o Academic Exchange (NAWA). Acknowl- edgemen s o speci ic ins i u ions and people sha ing hei aluable s andings da abase is in Table B3. Appendix A. Supplemen a y da a Supplemen a y da a o his a icle can be ound online a h ps://doi. o g/10.1016/j.ecose .2022.101406. Re e ences Aguile a-Alcal´ a, N., Mo ales-Reyes, Z., Ma ín-L´ opez, B., Mole´ on, M., S´ anchez-Zapa a, J. A., 2020. Role o sca enge s in p o iding non-ma e ial con ibu ions o people. Ecol. 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