What is a tree in the mediterranean basin hotspot? A critical analysis

RESEARCH Open Access Wha is a ee in he Medi e anean Basin ho spo ? A c i ical analysis F édé ic Médail 1* , Anne-Ch is ine Monne 1 , Daniel Pa on 1 , Toni Nikolic 2 , Panayo is Dimopoulos 3 , Gianluigi Bacche a 4 , Juan A oyo 5 , Zol án Ba ina 6 , Ma wan Cheikh Albassa neh 7 , Giannian onio Domina 8 , B uno Fady 9 , Vlado Ma e ski 10 , S ephen Mi sud 11 and Aga he Le iche 1 Abs ac Backg ound: T ee species ep esen 20% o he ascula plan species wo ldwide and hey play a c ucial ole in he global unc ioning o he biosphe e. The Medi e anean Basin is one o he 36 wo ld biodi e si y ho spo s, and i is es ima ed ha o es s co e ed 82% o he landscape be o e he i s human impac s, housands o yea s ago. Howe e , he spa ial dis ibu ion o he Medi e anean biodi e si y is s ill impe ec ly known, and a ocus on ee species cons i u es a key issue o unde s anding o es unc ioning and de elop conse a ion s a egies. Me hods: We p o ide he i s comp ehensi e checklis o all na i e ee axa (species and subspecies) p esen in he Medi e anean-Eu opean egion ( om Po ugal o Cyp us). We iden i ied some cases o woody species di icul o ca ego ize as ees ha we u he called “c yp ic ees”. We collec ed he occu ences o ee axa by “adminis a i e egions”, i.e. coun y o la ge island, and by biogeog aphical p o inces. We s udied he species-a ea ela ionship, and e alua ed he conse a ion issues o h ea ened axa ollowing IUCN c i e ia. Resul s: We iden i ied 245 ee axa ha included 210 species and 35 subspecies, belonging o 33 amilies and 64 gene a. I included 46 endemic ee axa (30 species and 16 subspecies), mainly dis ibu ed wi hin a single biogeog aphical uni . The coun ies wi h he highes ee ichness a e G eece (146 axa), I aly (133), Albania (122), Spain (155), Macedonia (116), and C oa ia (110). The species-a ea ela ionship clea ly disc imina ed he iches cen al-eas e n (Balkans) and no he n (Alpine and Ce enno-Py enean) biogeog aphical p o inces, agains he i e wes e n p o inces in he Ibe ian Peninsula. We iden i ied 44 un ecognized “c yp ic ees”, ep esen ing 21% o he o al ees. Among he 245 axa iden i ied, 19 a e conside ed o be h ea ened (15 CR + EN + VU) o nea h ea ened (4 NT) by IUCN. Conclusions: The Medi e anean-Eu opean egion includes an unsuspec edly high numbe o ee axa, almos 200 ee axa mo e han in he cen al Eu opean egion. This ee di e si y is no dis ibu ed e enly and culmina es in he cen al-eas e n pa o he Medi e anean egion, whe eas some la ge Ty henian islands shel e se e al na ow endemic ee axa. Few axa a e ecognized as h ea ened in he IUCN Red lis , and he ulne abili y o hese species is p obably unde es ima ed. Keywo ds: Plan biogeog aphy, Medi e anean egion, Th ea ened ees, T ee de ini ion, T ee dis ibu ions * Co espondence: [email p o ec ed] 1 Aix Ma seille Uni , A ignon Uni , CNRS, IRD, IMBE. Technopôle de l’A bois-Médi e anée, cedex 4, BP 80, 13 545 Aix-en-P o ence, F ance Full lis o au ho in o ma ion is a ailable a he end o he a icle © The Au ho (s). 2019 Open Access This a icle is dis ibu ed unde he e ms o he C ea i e Commons A ibu ion 4.0 In e na ional License (h p://c ea i ecommons.o g/licenses/by/4.0/), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided you gi e app op ia e c edi o he o iginal au ho (s) and he sou ce, p o ide a link o he C ea i e Commons license, and indica e i changes we e made. Médail e al. Fo es Ecosys ems (2019) 6:17 h ps://doi.o g/10.1186/s40663-019-0170-6 Backg ound A ecen census o he biomass dis ibu ion on Ea h in- dica es ha plan s, mos ly e es ial species, ep esen abou 80% (i.e. ≈450 G C) o he sum o he biomass ac oss all axa wo ldwide, and ha plan biomass in- cludes abou 70% s ems and ee unks (Ba -On e al. 2018). This esul unde lines he c ucial ole o ees in he global unc ioning o he biosphe e. Wi h ca. 60,000 ee species ep esen ing 20% o he ascula plan spe- cies wo ldwide acco ding o GlobalT eeSea ch (Beech e al. 2017), ees cons i u e a majo componen o he s uc u e and unc ion o o es ecosys ems loca ed in empe a e and opical biomes. Highe le els o ecosys- em se ices, such as biomass p oduc ion, wa e supply, nu ien cycling and soil ca bon s o age, cha ac e ize o es s wi h a highe ee di e si y (Gam eld e al. 2013). Since o es s play a c ucial ole in he conse a- ion o e es ial biodi e si y by p o iding habi a s and ecological niches o a majo i y o plan s and animals (www.global ees.o g), i is necessa y ha assessmen s o he di e en ee axa in ol ed in o he composi ion o ecosys ems should be comp ehensi e and obus by in- cluding a biogeog aphical pe spec i e. Ne e heless, he global knowledge o he numbe and dis ibu ion o ee species emains su p isingly low bo h a he in e na ional le el (Beech e al. 2017) and o la ge biogeog aphic a eas ( egions and p o - inces). While h ea s on he opical o es s u ge he scien is s o ill he gap in knowledge on hei di e si y (Se a-Diaz e al. 2017), he i e Medi e anean eco e- gions (Medi e anean Basin, Cali o nia, cen al Chile, Cape Region o Sou h A ica and sou h-wes e n Aus alia) s ill equi e u he s udies. Wi h only 2% o he wo ld’s e es- ial su ace, Medi e anean eco egions con ain nea ly 20% o he Ea h’s o al plan di e si y, making e y signi ican biodi e si y ho spo s, second only a e opical ones (Cowling e al. 2005). The Medi e anean Basin is one o hese 36 cu en biodi e si y ho spo s wo ldwide (Médail and Mye s 2004), including bo h egional (Médail and Quézel 1997) and local ho spo s, i.e. mi- c o- and nano-ho spo s sensu Cañadas e al. (2014). I is es ima ed ha Medi e anean o es s co e ed 82% o he landscape be o e he beginnings o human impac s on ecosys ems housands o yea s ago (WWF 2001). Bu he whole Medi e anean Basin s ill comp ises a high ee ichness es ima ed o 290 indigenous axa (species and subspecies) including 201 endemics (Quézel and Médail 2003;Médail2008). The spa ial dis ibu ion o his woody plan biodi e si y is s ill impe ec ly known e en in Medi e anean-Eu opean egion. A biogeo- g aphical ocus on ee species dis ibu ion is needed o conse a ion o he Medi e anean o es s which ha e unde gone apid changes in s uc u e and com- posi ion in ecen decades (Mazzoleni e al. 2004), e en i o es su ace emains s able on he global scale (FAO and Plan Bleu 2018). Among he woody di e si y, he de ini ion o a ee pe se migh be p oblema ic and equi es some ag ee- men (e.g. Gschwan ne e al. 2009). This is especially he case o his o ically dis u bed ee species ha o en appea s as sh ubs and display as ue ees only a e a long pe iod wi hou dis u bance o in cul i a ion in ga - dens. This aspec is pa icula ly impo an in he Medi- e anean egion whe e he impac s o na u al and human dis u bances a e bo h e y old and se e e. In his s udy, we p o ide he i s checklis o all ee axa (species and subspecies) p esen in he no he n pa o he Medi e anean eco egion (i.e. he Medi e anean-Eu opean egion), since Medi e anean o es s occu mainly in his a ea (in Eu ope, o es s co e ca. 33% o o al land a ea: see Albe di Asensio e al. 2015) and he a ailable da a a e mo e obus and eadily a ailable he e han in he sou he n and sou h-eas e n Medi e anean. The e o e, he main objec i es o his s udy a e: (i) o examine whe he an ex ensi e su ey o he whole pu a i e ee axa challenge he de ini ion o wha is gene ally conside ed as a Medi e anean ee; (ii) o p o ide a comp ehensi e checklis o ee axa o he Medi e anean-Eu opean egion,i.e. om Po ugal o Cyp us; (iii) o examine he global spa ial dis ibu ion o ee axonomic di e si y (na i es and endemics) a he biogeog aphical and adminis a i e scales. Finally, we compa ed he dis ibu ion o ee di e si y in he Medi e anean-Eu opean egion om p e ious assess- men made in he whole Medi e anean Basin and he Eu opean con inen and discussed he implica ions o conse a ion. Me hods S udy a ea We de ined he Medi e anean-Eu opean egion as he No h Medi e anean e es ial eco egion ollowing he de ini ion o e es ial eco egions o he wo ld om Olson e al. (2001) (Fig. 1). They p o ide a biogeog aphic scheme de ined by exis ing global maps o lo is ic p o inces and by egional maps o uni s based on he dis ibu ion o selec ed g oups o plan s and on pub- lished egional classi ica ion sys ems o e es ial habi- a s. The biogeog aphical limi s o he Medi e anean egion a e simila o hose p oposed by Médail and Quézel (1997), excep o some a eas o he moun ain idge o he I alian peninsula and o he Balkans. Indeed, clima e indices a e no s ic ly ele an o de ine he bio- geog aphical limi s pe se, no ably o he uppe pa s o he Medi e anean moun ains. To al co e ed a ea, including islands, is 1,610,200 km 2 , wi h coas line o app oxima i ely 45,200 km. The o o- g aphic a ia ion in he a ea is la ge, and he al i udes Médail e al. Fo es Ecosys ems (2019) 6:17 Page 2 o 19 ange om 0 m a.s.l. up o 3482 m a.s.l. (Mulhacén, Sie a Ne ada –Spain). The s udy a ea co e s he ollowing coun ies (as admin- is a i e uni s) om wes o eas : Po ugal, Spain, F ance, I aly, Mal a, Slo enia, C oa ia, Mon eneg o, Albania, Macedonia, G eece and Cyp us. La ge Medi e anean islands (in addi ion o Mal a and Cyp us) co esponding o he Balea ic a chipelago, Co sica, Sa dinia, Sicily and C e e, a e ea ed as independen uni s o achie e a be e spa ial esolu ion o ee axa and island speci ici ies. The s udy a ea does no include he non-Eu opean Medi e anean a ea, i.e. No h A ica and Nea Eas . The a ea is cha ac e ized by a Medi e anean clima e ypes ma ked by mild and humid win e s, ho and d y summe s and a ela i e long pe iod o a idi y each yea which impose an e ec i e and se e e d ough on he plan s. Rain all pa e ns and he oceani y a e ex emely a ied, and his explain why he Medi e anean Basin is cha ac e ized by he lowes ain all eliabili y (quan i ied as in e annual a ia ions in mon hly and seasonal ain- alls, and as he equency o indi idual ain alls) o he i e Medi e anean-clima e ecosys ems o he wo ld (Cowling e al. 2005). In gene al, annual ain alls a e g ea e in he no he n han in he sou he n a eas o he Medi e anean Basin and inc ease wi h ele a ion in he moun ain anges. Du ing mois season he ain all can be unp edic able and o en in ense om au umn h ough sp ing. Medi e anean ege a ion ypes a y widely wi h en i - onmen al cons ain s (clima e, geology, geomo phology, soil ype), dominan plan species, and dis u bances in- duced by man and i s he ds. The woody ege a ion is cons i u ed by di e se ypes o ma o als o sh ublands (maquis, ga igue, ph ygana) and o es s. In ela ion o hebioclima e ypeand heal i ude,se e almajo ypeso his ege a ion can be ecognized in he Medi e anean- Eu opean egion (Quézel and Médail 2003; Ri as-Ma ínez e al. 2007; Médail 2008): (i) a he mo-Medi e anean bel om sea le el o ~ 200–500(800) m a.s.l., domina ed by scle ophyllous communi ies mos o en as a na ow s ip along he coas (wi h Olea eu opaea,Ce a onia siliqua, Chamae ops humilis,Pis acia len iscus,Pinus halepen- sis,Pinus b u ia, e c.); (ii) a meso-Medi e anean bel , be ween ~ 100–500(1000) m a.s.l., mainly wi h scle - ophyllous o es s (Que cus ilex,Que cus sube and Q. cocci e a)o Pinus halepensis/P. b u ia o es s in a eas wi h low ain all o o me ly dis u bed by man; (iii) a sup a-Medi e anean bel , be ween ~ 500–1500(1800) m a.s.l., wi h di e se deciduous oaks o es s (including Ace , Ca pinus,Os ya,Que cus and So bus) in he mo e humid omb o ypes; (i ) a moun ain-Medi e anean le el (~ 1500– 2000 m a.s.l.) including Fagus and deciduous and semi-de- ciduous Que cus o es s and no ably coni e ous o es s wi h Pinus nig a, Pinus syl es is and i s (Abies alba and mos o he Medi e anean Abies spp.) and e en ceda (Ced us libani s.l.)onCyp us. An ex emely impo an ea u e o his woody Medi e anean ege a ion is ha i has been d ama ic- ally in luenced by human ac i i ies o housands o yea s (e.g. Thi good 1981; Blondel e al. 2010). Fo es log- ging, i es, g azing, ag icul u e de elopmen , soil dis u - bances, excessi e up ake o wa e o he de imen o ipa ian ees, changes in he dis ibu ion o na i e species due o la ge e o es a ions, a o es a ions, and in oduc ions o alien axa led o massi e landscape ans o ma ions and changes in he s uc u e and dynamics o hese na u al o - es ecosys ems (Quézel and Médail 2003;Blondel2006). Fig. 1 Numbe o na i e ee axa (species o subspecies –including endemic and pu a i e na i e) in each con inen al adminis a i e a ea and main islands in he Medi e anean-Eu opean egion. Da ke g eens indica e highe ichness Médail e al. Fo es Ecosys ems (2019) 6:17 Page 3 o 19 Medi e anean islands a e in his espec o pa icula conce n, due o he highe h ea hey su e , as a com- bina ion o highe ypical sensi i i y o dis u bances, and long his o y o human occupancy (e.g. Médail 2017). Fo his eason, we conside ed la ge islands as sepa a e uni s in ou analysis. T ee de ini ion The IUCN’s Global T ee Specialis G oup (GTSG) de ines a eeas“a woody plan wi h usually a single s em g owing o a heigh o a leas wo me e s, o i mul i-s emmed, hen a leas one e ical s em i e cen ime e s in diame e a b eas heigh ”(Beech e al. 2017). Ou de ini ion is sligh ly di e en and we de ine as ee he plan s wi h he ollowing biological ea u es: (i) seconda y g ow h (Fahn 1990); (ii) pe ennials ( hey li e many yea s, mos ly o de- cades o cen u ies); (iii) ypically (a leas in some en i on- men al condi ions o loca ions) ha ing a single s em o unk, g owing o ho opic wi h monopodial o mo e o en sympodial b anching sys ems, and bea ing la e al b anches a some dis ance om he g ound; and (i ) he heigh o he ma u e indi idual is a leas h ee me e s. Such plan s co espond o he g ow h- o m “phane o- phy es”acco ding o Raunkiæ (1934) and Ellenbe g e al. (1991) (o iginally P, excluding NP = nanophane ophy es, heigh 0, 5–5 m), o phane ophy es “scapose”(o iginally Pscap), phane ophy es “caespi ose”(o iginally Pcaesp) and phane ophy es “s iscian i”(o iginally P ep ) acco ding o Pigna i (1982). By s ic ly ollowing hose c i e ia, we iden i ied some cases o woody species di icul o ca ego ize ha we u - he called “c yp ic ees”. Indeed, his de ini ion includes some axa ha gene ally occu as sh ubs wi h poo ly de- ined mul icolous s ems bu can o m ue ees unde ce - ain en i onmen al condi ions o in si ua ions whe e he dis u bances ha e been absen o e y educed o a leas se e al decades. Su p isingly, i u ned ou ha hese cases we e no a e in he s udy a ea. This is he case o ins ance o Spa ium junceum o Genis a e nensis (Fabaceae) which usually g ows in he o m o a bush (nanophane ophy e) o less han 5 m high. We included hem in he checklis be- cause in some locali ies o Co sica, Sa dinia and Sicily (Aeolian islands) hese axa mee he abo e c i e ia o ee. Many o he ee species epo ed in his s udy may e y equen ly appea as sh ubs o e en p os a e sc ubs due o in ense dis u bance, pa icula ly he bi o y. Occu ences and s a us by adminis a i e o by biogeog aphical a eas Occu ences o each ee axa we e compiled (i) by coun y (n= 12) and la ge island (n= 5) (he ea e , “ad- minis a i e egions”), and (ii) by biogeog aphical p o - inces. To da e, a ine spa ial app oach is no a ailable o all axa pe coun ies h oughou he s udy a ea. The de ini ion o he biogeog aphical p o inces ollowed he Biogeog aphic map o Eu ope p o ided by Ri as-Ma í- nez e al. (2004). Hence, hi een biogeog aphic p o inces occu in he s udy a ea (Fig. 3a). Th ee o hem a e in- cluded in o he Eu osibe ian biogeog aphical egion bu a e ep esen a i e o ansi ion zones be ween he Medi- e anean and he Eu osibe ian egions (Ce enno-Py e- nean, Alpine, and Apennino-Balkan); he o he en a e included in he Medi e anean egion. We s udied he species-a ea ela ionship wi hin he adminis a i e o biogeog aphical egions by plo ing he o al na i e ee axa ichness o each geog aphic uni as a unc ion o hei su ace, a e log ans o ma ion. Fo each species o subspecies, he p esence in each adminis a i e o biogeog aphical uni , was de ined as na i e (N) o in oduced (I). We conside endemic ee axa as ange- es ic ed axa in and ou side o he s udy a ea, in gene al loca ed wi hin a unique biogeog aphical p o ince sensu Ri as-Ma ínez e al. (2004). We ma ked axa wi h unclea au och honous o alloch- honous o igin as pu a i e na i e (i.e. “N?”, e.g. Te aclinis a icula a in Mal a), o pu a i e in oduced (i.e. “I?”). Taxa o which hep esenceisno con i medbu possiblewe e quo ed as pu a i e p esence (i.e.“?”,e.g.C a aegus pen a- gyna in Mon eneg o). Many cul i a ed and some ime na u- alized ee species ha e no been conside ed. We ocused only o a small numbe o axa ha ha e been cul i a ed o a long pe iod (a cheophy es, i.e. p esen be o e he clas- sical da e o he “disco e y”o Ame ica in 1492) and, apa om cul u e, a e widely na u alized in some pa s o he s udied a ea bu also na i e in some a eas o he Medi e anean-Eu opean egion i.e. Cas anea sa i a Mill., Ce a onia siliqua L., Cup essus sempe i ens L., Ficus ca ica L., Juglans egia L., e c. (see Addi ional ile 1). We conside ed h ea ened ee axa using he IUCN (2018) classi ica ion sys em, as axa included in o he h ee classes: c i ically endange ed (CR), endange ed (EN), o ulne able (VU). The o he ca ego ies included nea h ea ened axa (NT), leas conce n axa (LC), and da a de- icien axa (DD) a e also conside ed in he p esen wo k. Da a sou ces The da a we e collec ed and agg ega ed o each coun y and la ge island. The p ima y sou ces o da a we e he in e na ional da abase Eu o + Med Plan Base (h p://www.emplan base.o g/home.h ml), Flo a Eu o- paea (Tu in e al. 1964–1993), and he olume se ies Cho ology o ees and sh ubs in sou h-wes Asia and adjacen egions (B owicz 1982–1996). These comp e- hensi e da a we e ca e ully comple ed and checked wi h a ailable na ional lo as, na ional da abases, egional da abases, o he publica ions, and ou own expe ise, when he a o emen ioned sou ces we e no a ailable: Albania (Ba ina e al. 2017; Z. Ba ina pe s. comm.), Médail e al. Fo es Ecosys ems (2019) 6:17 Page 4 o 19 Co sica (Jeanmonod and Gamisans 2013; CBN Co se pe s. comm.), C e e (Chil on and Tu land 1997;h p:// www.c e an lo a.com/;S id2016), C oa ia (Nikolić 2004–onwa d), Cyp us (Hand e al. 2011–onwa d), con- inen al F ance (Silene-Flo e da abase: h p:// lo e.silene.eu, IGN In en ai e Fo es ie : h p://in en ai e- o es ie .ign. /, Tison e al. 2014), G eece (Dimopoulos e al. 2013,2016; S id 2016), Ibe ian Peninsula i.e. Balea ic Islands, Spain and Po ugal (An hos: h p://www.an hos.es;Cas o iejo 1986–2015; Sociedade Po uguesa de Bo ânica 2014), I aly (Ba olucci e al. 2018; Galasso e al. 2018; Vegi aly: h p:// www. egi aly.i /), Macedonia (V. Ma e ski ined.), Mal a (Mazzocchi 1969; Baldacchino and S e ens 2000;Mi sud 2002–2014;Casha2015; Baldacchino 2018;Calleja2018); Mon eneg o (F. Médail & A.-C. Monne obs. pe s.), Sa - dinia (A igoni 2006–2015; Bacche a e al. 2012;G.Bac- che a obs. pe s.), Sicily (Gia dina e al. 2007), Slo enia (Jogan e al. 2001). In addi ion, some o he mo e hema ic and comp e- hensi e da a sou ces we e used such as he Eu opean A las o Fo es (San-Miguel-Ayanz e al. 2016) and com- ple ed by some ecen pape s conce ning he desc ip ion o some new ee axa, such as Tama ix minoa in C e e (Villa e al. 2015) o wo c yp ic species in he Alnus glu inosa g oup (Ví e al. 2017). Da a syn hesis was led by D. Pa on & F. Médail be ween 2014 and 2018. Taxonomy and nomencla u e The p esen checklis o Medi e anean-Eu opean ees includes only he axa a he species and subspecies le els which mee bo h c i e ia: “s udy a ea”and “ ee de ini ion”. This lis does no con ain o he sub-speci ic le els, such as a ie ies, o ms, o hyb ids, because o hei axonomical and nomencla u al ins abili ies. We ollowed he classi ica ion o he Angiospe m Phylogeny G oup o he de ini ion o amilies (APG III 2009;APGIV 2016). The nomencla u e o Medi e anean and Eu opean ees is o en luc ua ing and con o e sial be ween he di e en majo axonomic wo ks. We conside ed in p io - i y he Eu o+Med da abase (h p://www.emplan base.o g/ home.h ml) and he Plan Lis da abase (h ps://www. he- plan lis .o g), bu also he majo na ional lo as (see e - e ences in he “da a sou ces”pa ag aph). In case o axonomic-le el disag eemen , we e ained he mos consensual axonomic le el based also on ecen phylo- gene ic s udies o sys ema ic e ision o a gi en g oup, i a ailable, adding consul ing local expe s o ecen lo as. Resul s Medi e anean-Eu opean ees in a nu shell Ou analysis o all he axa ha can po en ially be ee species in he Medi e anean-Eu opean egion p o ides a checklis o 245 ee axa (i.e. species and subspecies), in- cluding 210 species and 35 subspecies (see Addi ional ile 1). These axa belong o 33 di e en amilies; he mos ep esen ed amilies a e Rosaceae (42 axa), Fagaceae (33 axa), Pinaceae (21 axa), Salicaceae (20 axa), Sapindaceae (19 axa), Be ulaceae (16 axa), Tama ica- ceae (16 axa), and Cup essaceae (14 axa). These ee axa a e included in 64 di e en gene a which ep esen s 86% o he o al ee gene a (n= 74) ound in he whole Medi e anean biogeog aphic egion (Quézel and Médail 2003). Fou o hese gene a (Cha- mae ops,Philly ea,Spa ium,Te aclinis) ha e hei dis- ibu ion cen e ed in he Medi e anean egion and can be conside ed as Medi e anean endemics sensu la o (a he biogeog aphical egion le el), whe eas h ee axa ha e a Te hysian o igin (Ce a onia, My us,Ne ium) and occu in adjacen biogeog aphic egions as well. The o he gene a ha e la ge dis ibu ions, in he Palea c ic ealm o beyond. Among his checklis , 46 a e endemic ees (30 spe- cies and 16 subspecies), i.e. mainly loca ed wi hin a sin- gle biogeog aphic p o ince sensu Ri as-Ma ínez e al. (2004). This means ha he a e o endemism is equal o 18.9% i we conside he species and subspecies le els, o 14.3% a he species le el. The gene a wi h hemo eendemic axaa eQue cus (6 axa), Abies, Ace , and Pinus (4 axa each), Alnus,Salix and So bus (3 axa each). The p esence o h ee na ow endemic species included in wo elic gene a (Liquidamba and Zelko a) and cha ac e is ic o he Cenozoic paleo lo a dis ibu ed in he Palea c ic ealm should also be no ed (Palama e 1989). E en hough we only ook in o accoun axa consid- e ed as indigenous o a gi en coun y o a biogeog aph- ical a ea, he na i e s a us o some ee species can some imes be di icul o es ima e, no ably when hey a e equen ly cul i a ed o used o a o es a ion, o when hei cen e o o igin o na i e ange a e s ill unclea (e.g. Cas anea,Ce a onia,Ficus,Juglans, Olea: see discussion). O e all, i conce ns 24 axa (ca. 10% o he o al checklis ) ha a e widely cul i a ed o selec ed o a o es a ion pu posed by o es se - ices. The dis ibu ion o hose na i e bu locally do- mes ica ed Medi e anean ee axa will be discussed in de ail below since i ep esen s a c ucial issue o he conse a ion o local gene ic esou ces. This checklis also includes 44 un ecognized eal ees o “c yp ic ees”, i.e. usually wi h a sh ubby habi bu which o m a ee habi unde ce ain en i onmen al condi ions (Table 1).These axabelong o ou main amilies (Rosaceae: 11 ee species, Cup essaceae: 5 ee species, Anaca diaceae and Faba- ceae: 4 ee species each), and o wo main gene a (9 ee species in C a aegus and 5 in Junipe us). These c yp ic ees ep esen he imp essi e and unexpec ed a e o ≈18% o he Medi e anean-Eu opean ees. Médail e al. Fo es Ecosys ems (2019) 6:17 Page 5 o 19 Table 1 The 44 un ecognized eal ees o “c yp ic ees”, i.e. usually wi h a sh ubby o m, occu ing in he Medi e anean-Eu opean egion, wi h indica ions abou hei dis ibu ion and ecology Taxa Family Global dis ibu ion a ea Ecology Alnus i idis (Chaix) DC. Be ulaceae A c ico-Alpine Subalpine and o o-medi e anean sc ublands Buxus balea ica Lam. Buxaceae SW Medi e anean, S Ana olia Sc ublands on p edominan ly ca bona e subs a es Buxus sempe i ens L. Buxaceae Medi e anean-Eu opean Submedi e anean sc ublands Chamae ops humilis L. A ecaceae CW Medi e anean The mophilous sc ublands, coas al sc ee and cli s Co inus coggyg ia Scop. Anaca diaceae S Eu opean, I ano-Tu anian Eumedi e anean and submedi e anean d y sc ublands, wa m and mesophilous habi a s o sou he n exposu e inland Co oneas e g ana ensis Boiss. Rosaceae Endemic o S Spain (Andalusia) Rocky ma o als, deciduous and pine o es s C a aegus aza olus L. Rosaceae E Medi e anean Open sc ublands C a aegus held eichii Boiss. Rosaceae Balkan Open coni e ous o es s, mixed sc ubs, d y ocky meadows C a aegus lacinia a Uc ia Rosaceae CW Medi e anean Clea ings o moun ain woods C a aegus lae iga a (Poi .) DC. Rosaceae CW Medi e anean Mesophilic shingles and hedges o he hilly and moun ainous ege a ion bel , oak o es s ou side he lood, moun ain beech o es s C a aegus monogyna Jacq. Rosaceae Medi e anean-Eu opean Coas al d y sc ublands, Inland mesophilic shingles and hedges C a aegus ne adensis K.I.Ch . Rosaceae Endemic o S Spain (Andalusia) Clea ings and edges o moun ain woods C a aegus o ien alis Pall. ex M.Bieb. Rosaceae E Medi e anean and SW Asia Rocky places in meadows and open woodlands C a aegus pen agyna Willd. Rosaceae Balkan Open o es , mixed sc ubs, d y ocky meadows C a aegus pycnoloba Boiss & Held . Rosaceae Endemic o G eece (Peloponnese) Rocky limes one idhes and slope, d y g assy pla eau, sc ubs and open Abies cephalonica o es s Cy isus aeolicus Guss. Fabaceae Endemic o Sicily (Aeolian Islands) D y slopes and open sc ublands on olcanic subs a es E ica a bo ea L. E icaceae Medi e anean, Maca onesian, E. A ica, Saha a The mophilous sc ublands in non-ca bona ic subs a es Fon anesia philli aeoides Labill. Oleaceae E Medi e anean D y maquis F angula alnus Mill. Rhamnaceae C Eu opean, Medi e anean Ripa ian o es s o alde and ash Genis a e nensis (Ra .) DC. Fabaceae Endemic o Sicily, Sa dinia and Co sica A bo escen ma o als on non-ca bona ic subs a es Genis a y hena Valsecchi Fabaceae Endemic o Sicily (Aeolian Islands) and Lazio (Pon ian Islands) A id olcanic slopes on ocky and sandy subs a es Junipe us communis L. Cup essaceae Eu asia ic Rocky meadows, moun ain pas u es, mesophilic hedges and sh ubs, o es s edges in submon ane a eas Junipe us del oides R.P.Adams Cup essaceae CE Medi e anean Open woodlands and sc ublands on mid-al i udes Junipe us na icula is Gand. Cup essaceae Endemic o SW Ibe ian Peninsula A bo escen ma o al on ma i ime sands Junipe us oxyced us L. Cup essaceae Medi e anean-Pon ic Eumedi e anean and submedi e anean d y a eas in open o es s and ma o als Junipe us phoenicea L. Cup essaceae Medi e anean, Maca onesian, Red Sea Eumedi e anean d y and wa m a eas in e e g een oak and pine o es s, coas al ma o als My ica aya Ai on My icaceae Maca onesia, S Po ugal Unde wood o pinewoods on siliceous My us communis L. My aceae Medi e anean, Maca onesian, I ano-Tu anian The mophilous sc ublands on non-ca bona ic subs a es, small swampy o es s ands Ne ium oleande L. Apocynaceae Medi e anean, Saha a The mophilous and empo a y i e beds, we lowlands Médail e al. Fo es Ecosys ems (2019) 6:17 Page 6 o 19 Medi e anean-Eu opean ee di e si y by adminis a i e egions Mapping he p esence o ee species in each Medi e anean-Eu opean coun y is impo an o es- ima e conse a ion p io i ies a he adminis a i e le el. Ou analysis shows ha he coun ies (con inen al pa s) wi h he highes o al ichness o na i e ees (> 110 axa) a e G eece (146 axa), I aly (133 axa), Albania (122 axa), Spain (115 axa), Macedonia (116 axa), and C oa ia (110 axa) (Fig. 1a, Table 2). The e is a s ong dispa i y in o al ichness be ween he la ge Medi e anean islands: Sicily has he highes ichness (88 axa), and he Balea ic Islands he lowes one (35 axa), whe eas Co sica and Sa dinia in he wes e n Medi e anean Basin exhibi a simila ichness ( espec i ely 70 and 76 axa) and Cyp us and C e e in he eas e n basin sha e qui e simila le el o ee ichness ( espec i ely 42 and 52 axa). While he o al ee ichness is concen a ed mainly in he Balkans, endemic ees mos ly occu ed in he la ge coun ies o he h ee peninsula in wes e n, cen al and eas e n Medi e anean a ea (Table 2). G eece (11 endemic axa), Spain (9 endemic axa), and I aly (7 endemic axa) exhibi ed he highes numbe o endemic axa. Emphasis should be gi en o he place o some la ge Ty henian islands ha shel e a signi ican numbe o endemic ees (Sicily: 10 axa, Sa dinia: 7 axa). The species-a ea ela ionships show con as ing e- sul s be ween adminis a i e egions and la ge Medi e - anean islands (Fig. 2). The ela ionship is signi ican o he islands (log ichness islands: −1.89841 + 0.25788 log su ace islands, p= 0.01) bu no o he con inen al a eas o he coun ies (log ichness con inen s: 3.8459 + 0.03493 log su ace con inen s, p= 0.41). The con in- en al a eas o he cen al and eas e n Medi e anean coun ies, om I aly o G eece (i.e. mainly he Balkans), ha e a signi ican ly highe ee axonomic ichness han hose loca ed u he wes o on he la ge islands. Thus, he islands always ha e a lowe ichness pe uni a ea han con inen s. This is pa icula ly he case o Mal a and he Balea ic Islands. Ne e heless, in Sicily and Sa dinia, he a io is close o some con inen al si ua- ions (e.g. F ance, Slo enia) and is e en highe ha Po ugal which shows he lowes a ea-species a io o a con inen al coun y. Table 1 The 44 un ecognized eal ees o “c yp ic ees”, i.e. usually wi h a sh ubby o m, occu ing in he Medi e anean-Eu opean egion, wi h indica ions abou hei dis ibu ion and ecology (Con inued) Taxa Family Global dis ibu ion a ea Ecology Philly ea angus i olia L. Oleaceae W Medi e anean E e g een oak and alepo pine o es s, d y and wa m habi a s o en s ony, ma o als Philly ea la i olia L. Oleaceae Medi e anean E e g een oak and alepo pine o es s, on d y and wa m habi a s o en s ony, ma o als Pis acia len iscus L. Anaca diaceae Medi e anean, Maca onesian, E e g een oak o es s and ma o als Pis acia e ebin hus L. Anaca diaceae Medi e anean, S Eu opean E e g een oak o es s and ma o als P unus mahaleb L. Rosaceae S Eu opean-Pon ic The mophilous o es s o pubescen oak and ma o als Que cus cocci e a L. Fagaceae Medi e anean E e g een oak o es s and ma o als Rhamnus ala e nus L. Rhamnaceae Medi e anean Mixed e e g een and deciduous o es s, ma o als domina ed by e e g een oaks Rhododend on pon icum L. E icaceae S Ibe ian Peninsula, Lebanon, Pon ic egion Mixed woodlands in humid a ines on siliceous Sambucus nig a L. Adoxaceae Eu opean-Caucasian Mesophilous unde b ushs and hedges in hilly and moun ains a eas, deciduous o es s Sea sia ipa i a (Uc ia) Mo e Anaca diaceae S Medi e anean, Saha a Open ocky ma o als in a id and dese ic a eas, empo a y i e banks Spa ium junceum L. Fabaceae Medi e anean, Eu opean Eumedi e anean d y, wa m and s ony a eas, o en in mo e o less pu e s ands S y ax o icinalis L. S y acaceae CE Medi e anean Rocky and d y g asslands, he mophilous ma o als wi h e e g een oaks and alepo pine Vi ex agnus-cas us L. Lamiaceae Medi e anean, I ano-Tu anian Ri e beds, we lowlands, he mophilous damp o loodes unde b ushes Zelko a abelicea (Lam.) Boiss. Ulmaceae Endemic o C e e Rocky slopes o la alley bo oms in he moun ains Zelko a sicula Di Pasquale, Ga i & Quézel Ulmaceae Endemic o Sicily Sc ublands on basal ic soil, i e beds Médail e al. Fo es Ecosys ems (2019) 6:17 Page 7 o 19 Medi e anean-Eu opean ee di e si y by biogeog aphical a eas To go beyond hei dis ibu ion in he adminis a i e e- gions, we ex ac ed he numbe o na i e ee axa (species and subspecies) o each o he 13 biogeog aphic p o inces sensu Ri as-Ma ínez e al. (2004)(Fig.3a). The aw numbe s o ee ichness a e culmina ing in he cen al-eas e n p o inces o he Medi e anean-Eu opean egion: Apennino-Balkan (142 axa), G aeco-Aegean (137 axa), Ad ia ic (135), and I alo-Ty henian (128 axa). The lowes ichness (40 axa) is ound in he Cilicio-Phoenician p o ince which co esponds only (in ou s udy a ea) o he island o Cyp us (Fig. 3b). The highes axonomic ich- ness in endemic ees occu s in he I alo-Ty henian (n= 16 axa) and in he G aeco-Aegean (n=12 axa) p o inces, hen in he Be ican (n=9 axa) and Ad i- a ic (n= 8 axa) p o inces (Fig. 3c). The lowes en- demic ichness (n=1 axa) is ound in sou he n F ance, and in he sou he n pa s o he Alpine and Ce enno-Py enean p o inces. The species-a ea ela ionship o biogeog aphic p o - inces clea ly disc imina ed he cen al-eas e n (mainly Balkans) and no he n (Alpine and Ce enno-Py enean) p o inces, agains he i e wes e n p o inces in he Ibe - ian Peninsula (Fig. 4). Th ea ened ees Among he 245 ee axa o he checklis , 15 cons i u e h ea ened axa sensu IUCN (2 CR, 7 EN, 6 VU) and 4 a e classi ied as nea h ea ened axa (NT) (Table 3). Only 154 ee axa (145 species and 9 subspecies) a e classi ied as “leas conce n”(LC), and 23 species wi h da a gap o assessmen (DD). Wha is mos wo ying is ha 84 axa (19 species and 65 subspecies) included in he checklis a e no conside ed a all in he global IUCN assessmen . Ele en o hese h ea ened ee axa, including some e y na ow endemic ees such as Abies neb odensis,Cy isus aeolicus,Rhamnus pe sici olia,Zel- ko a abelicea and Zelko a sicula a e only dis ibu ed on some la ge (no ably Sicily, C e e, Cyp us and Sa dinia) o medium ( ew C oa ian islands shel e ing Pinus nig a subsp. dalma ica popula ions) sized islands o he Medi e anean Sea. Discussion Dis ibu ion o ee di e si y in he Medi e anean- Eu opean egion Ou comp ehensi e analysis o ee axonomic di e si y in he Medi e anean-Eu opean egion iden i ied 210 ee species o 245 species and subspecies, including 46 endemic ees o he s udy a ea. This is o cou se a iny po ion o he es ima ed ee di e si y wo ldwide (ca. 60,065 ee species cu en ly known o science, see Beech e al. 2017), bu an impo an numbe ega ding o he Eu opean o es s. T ee axonomic di e si y was es ima ed a 135 species and subspecies wi hin he medio-Eu opean egion (Quézel and Médail 2003), whe eas only 42 ee species om 11 amilies occu in Cen al Eu ope (Leuschne and Meie 2018). I ep e- sen s also o high ee di e si y compa ed o he one in he en i e Medi e anean biogeog aphical egion which was es ima ed a 290 species and subspecies (Quézel and Médail 2003), i.e. 85% o he ees o he whole Medi e - anean egion occu in he Eu opean pa . This key e- sul shows he impo ance o he sou he n pa o Eu ope in p ese ing his ich biological he i age o a empe a e egion. The axonomic ichness o he Medi e anean-Eu opean egion is also ound in ee gene a, since 86% o he o al gene a (n= 74) o he whole Medi e anean biogeog aphic egion a e p esen in his Eu opean pa . Only 11 gene a (Acacia,Amygdalus,A gania,Balani es,Calo opis,Dios- py os,D acaena,Mo inga,P ospis,P e oca ya,Ziziphus) a e es ic ed o he sou he n o eas e n pa s o he Medi e anean egion. These esul s a e no su p ising since Medi e anean o es s a e e y di e se and he e ogeneous ega ding hei s uc u e, hei speci ic composi ion and hei dy- namics (Quézel and Médail 2003), and his explains he exis ence o 35 e es ial sub-eco egions including Table 2 Numbe o o al na i e ee axa (including endemic and pu a i e na i e axa), endemic ee axa, and pu a i e na i e ee axa) by adminis a i e a eas and main islands o he Medi e anean-Eu opean egion. “Taxa”include bo h species and sub-species le el Coun ies To al na i e axa ichness Endemic axa ichness Pu a i e na i e axa ichness P esence unce ain y Albania 122 2 3 0 Balea ic 33 1 0 0 Co sica 70 5 0 0 C e e 52 4 0 0 C oa ia 110 2 0 0 Cyp us 42 3 0 0 F ance 102 1 3 0 G eece 146 11 0 0 I aly 133 7 0 0 Macedonia 116 3 3 2 Mal a 25 0 5 0 Mon eneg o 108 1 0 1 Po ugal 69 3 3 0 Sa dinia 76 7 0 0 Sicily 88 10 0 0 Slo enia 85 0 1 1 Spain 115 9 0 0 Médail e al. Fo es Ecosys ems (2019) 6:17 Page 8 o 19 o es s (WWF 2001). The main o es ypes a e: (i) scle ophyllous o es s (no ably wi h he oaks Que cus ilex, Q. sube ,Q. cocci e a); (ii) d y coni e (Pinus halepensis,P. b u ia) o es s; (iii) deciduous mixed o es s wi h b oad-lea ed oaks (Que cus pubescens,Q. aginea,Q. i ha- bu ensis) and o he gene a (Ace ,Ca pinus,F axinus, So bus); (i ) moun ain coni e o es s wi h Abies, Ced us,Junipe us,Pinus nig a. T ee di e si y and he ecological complexi y in o es ypes can be explained bo h by he complex paleogeog aphy and his o ical bio- geog aphy o he Medi e anean egion, and also by he high he e ogenei y in he clima e (Médail 2008;Blondel e al. 2010). This woody biodi e si y is une enly dis ibu ed wi hin he Medi e anean-Eu opean egion. Ou analysis, pe - o med ei he a he adminis a i e o biogeog aphic le el, indica e ha he con inen al a eas o he cen al and eas e n Medi e anean coun ies, om I aly o G eece (i.e. mainly he Balkans), ha e a signi ican ly highe ee ichness compa ed o hose loca ed u he wes (Ibe ian Peninsula) o on la ge Medi e anean islands. The highe axonomic di e si y in he eas e n Medi e anean Basin is ound in se e al o he s udies and his pa e n is p obably ela ed o he key biogeog aphical c oss oad o his a ea and o he exis ence o mo e sui able condi ions du ing he LGM (no ably wa me summe empe a u es) inducing la ge biodi e si y e ugia (Médail and Diadema 2009; Fady and Cono d 2010). Ou esul s a e also consis en wi h hose ob ained om wo s udies ha included a mo e educed sampling o ees in Eu ope: (i) he dis ibu ion o 55 Eu opean ee species sug- ges s ha he ee ichness is maximal in he Balkans, especially o ees wi h a es ic ed dis ibu ion (S enning and Sko 2007); (ii) he dis ibu ion o ee ichness ( o al n= 187 species) o Eu ope shows he signi ican concen- a ion o ee di e si y along a la ge coas al a ea ex ended om C oa ia o no h-wes e n G eece (Mon oya and Rod íguez 2007). Bo h s udies concluded ha he main d i e o he a ia ion o ee ichness is he cu en clima e, and ha he clima e du ing he Las Glacial Maximum (LGM, ca. 20 ky ago) o he pe iod since an a ea became ee o ice ha e a signi ican in luence, no ably o explain he concen a ion o ange- es ic ed ee axa. Some s udies also e ealed ha olde clima ic e en s h oughou he Miocene we e also c i ical o ex- inc ion in he wes e n pa o he Medi e anean Basin ( he Ibe ian Peninsula) and may pa ly explain he ela i e lowe ichness o ee axa he e (Pos igo Mija a e al. 2009). The species-a ea ela ionships indica e ha Medi- e anean islands always ha e a lowe ichness pe uni a ea, and only he wo la ges islands, Sicily and Sa dinia, a e g ouped wi h he con inen al e i o ies o highe ee Fig. 2 Linea eg essions be ween he o al na i e ee axa ichness and he su ace o con inen al adminis a i e a eas (black eg ession line and black ci cles) o main islands (g ey eg ession line and black iangles) in he Medi e anean-Eu opean egion wi h hei espec i e 95% con idence in e al. Abscissa and o dina e axes ha e a loga i hmic scale. Do s abo e each line indica e highe ichness han p edic ed by he species-a ea ela ionship, and do s below each line indica e lowe ichness han expec ed Médail e al. Fo es Ecosys ems (2019) 6:17 Page 9 o 19 minoa, occu ing in a unique locali y o NW C e e on sandy i e banks close o he sea (Villa e al. 2015), is quo ed Da a De icien (DD). Among he o he DD axa, a leas wo cha ac e is ic axa (Pla anus o ien alis,Vi ex agnus-cas us) cu en ly expe ience a se e e educ ion o hei popula ions due o he a i icializa ion o hei habi a s, e en i hey s ill coun a qui e la ge numbe o popula ions. Ano he unde es ima ed h ea is in og ession be- ween phylogene ically ela ed ee axa. This is he case o he g oup o black pines (Pinus nig a) whe e in o- g ession isk is likely be ween Pinus nig a subsp. nig a, equen ly plan ed o e o es a ion, and wo o he sub- species ha ing a mo e es ic ed dis ibu ion ange: subsp. dalma ica in C oa ia, and subsp. salzmannii in F ance and Spain. Al hough in equen and po en ially bene icial o some ai s, such gene low om plan ed o na i e ees o could con ibu e o wo sen local adap- a ion, o example o diseases such as Do his oma sp. (Sco i-Sain agne e al. 2018). F om a geog aphical poin o iew, we mus emphasize he ole o he la ge Medi e anean islands, which in- clude 11 o he 19 ee axa cu en ly p esen in he IUCN da abase. This shows he impo ance o hese is- land ecosys ems o he di e en ia ion and pe sis ence o axonomically singula ees, bu also he se e e h ea s in hese spa ially es ic ed sys ems. This is he case o Sicily which includes h ee a e na ow endemics (Abies neb odensis,Cy isus aeolicus,Zelko a sicula) among which he wo la e a e only p esen on some small islands o he Aeolian a chipelago (T oia 2012). Some o he na ow endemic ees (Genis a y hena om Aeolian and Pon ine a chipelagos in I aly, and Que cus alni olia es ic ed o he T oodos ange in Cyp us) a e classi ied as leas conce n (LC), bu due o hei es ic ed ange and popula ion size hese e alua- ions a e p obably oo op imis ic. A mo e p ecise isk analysis by egion would also p obably highligh he high le el o h ea o locally se e al Medi e anean ees. Fo example, he y henian insula endemic Genis a e nensis which is anked globally as LC is classi ied as c i ically endange ed (CR) in Co sica. This is also he case o se - e al ees wi h a pa chy dis ibu ion in specialized moun- ain habi a s such as Junipe us d upacea,J. oe idissima, J. hu i e a, and Pinus held eichii which a e classi ied as LC as well. Some o he locally h ea ened axa (Buxus balea ica,Rhododend on pon icum) wi h a highly e- s ic ed ange in he N. Medi e anean egion a e no e en conside ed in he IUCN da abase. Conclusion This s udy p o ides he i s comp ehensi e assessmen o ee dis ibu ions in he Medi e anean-Eu opean e- gion. The p esen checklis includes an unsuspec edly high numbe o ee axa (245 species and subspecies), wi h a leas 44 axa ha a e gene ally bushy bu can exp ess a ue ee a chi ec u e unde ce ain condi ions (“c yp ic ees”). This means ha he Medi e anean-Eu opean e- gion has almos 200 ees mo e han he Cen al Eu opean egion. This axonomic biodi e si y is highes in he cen al-eas e n pa o he Medi e anean egion, no - ably in he Apennino-Balkan p o ince. We mus also emphasize he ole o some la ge Ty henian islands (especially Sicily and Sa dinia) ha shel e a signi ican numbe o endemic ees. These a eas a e pa icula ly impo an om he poin o iew o unc ional conse - a ion because ee species ichness is posi i ely linked o highe le els o mul iple ecosys em se ices (Gam eld e al. 2013), and a e species –like mos o he Medi e a- nean endemic ees –suppo o iginal and ulne able unc ion (Mouillo e al. 2013). Bu h ea s o his woody biodi e si y a e o en signi ican and may especially a ec ees wi h na ow dis ibu ions. Howe e , he IUCN Red lis includes so a only 19 Medi e anean ees conside ed o be h ea ened (CR + EN + VU) o nea h ea ened (NT). The e o e, he ulne abili y o se e al o hese ees is p obably unde es ima ed and u he assessmen s a e needed in he nea u u e. Fo Medi e anean ees a o ed o cul i a ed by man, some imes o millennia, i is necessa y o de- elop app oaches combining phylogeog aphy and paleoecology, a he scale o he dis ibu ion a ea spe- ci ic o each axon. This will allow a be e es ima e o he na i e s a us o each biogeog aphical a ea, and o conside he e olu iona y dimension in he p ese - a ion o hese ees. This checklis ep esen s he i s s ep owa ds a be e unde s anding o he di e si y o ees in he Medi e a- nean egion. The nex s ep should be o imp o e he co e age in occu ences eco ds o hese ee axa in he egion in he exis ing ee species da abase (Noce e al. 2016; Se a-Diaz e al. 2017). These da a will help o explo e he key scien i ic ques ions associa ed o he d i e s o he dis ibu ion pa e n and o he shape o he species-a ea ela ionship. Quan i a i e in es iga ions o ecological (e.g., clima e, soil, dis u bance) e sus his- o ical (e.g., pas clima e, gene ic o iginali y) d i e s o cu en ee biodi e si y can he e o e be conside ed. This checklis will cons i u e also an essen ial back- g ound o unc ional and phylogene ic analyses o his key biological con ingen o e es ial ecosys ems. The p ac ical issues conce n a be e assessmen o he h ea ened s a us o some hese unapp ecia ed ees wi hin a highly human-modi ied egion. We also hope such wo k will ca alyze u he assessmen o ee di e - si y a he scale o he whole Medi e anean egion (i.e. also No h A ica and Middle-Eas ) e en i he da a a e s ill e y complex o syn hesize. Médail e al. Fo es Ecosys ems (2019) 6:17 Page 16 o 19 Addi ional ile Addi ional ile 1: Table S1. Checklis o he ee axa (species and subspecies) occu ing in heMedi e anean-Eu opean egion, om Po ugal o Cyp us. The de ini ion o he amilies ollows he Angiospe m Phylogeny G oup (APG III 2009; APG IV 2016). The e ained nomencla u e o he ee axa is compa ed wi h hose o he Eu o+Med da abase (h p://www.emplan base.o g/home.h ml), he olume se ies Cho ology o ees and sh ubs in sou h-wes Asia and adjacen egions (B owicz 1982– 1996) and he Wo ld Checklis Kew. Fo he occu ence o each ee axa in a coun y o a la ge island, we indica e he endemism s a us, he cul i a ed s a us, heau och honous (na i e: N) o alloch honous (in oduced: I) s a us. Taxa o which he p esence is no con i med bu possible a e quo ed as "pu a i e p esence », i.e.“?”. (XLSX 71 kb) Acknowledgmen s The au ho s hank Aggeliki Doxa (Ins i u e o Applied and Compu a ional Ma hema ics, He aklion), Lae i ia Hugo and Paula Spinosi (Conse a oi e bo anique na ional de Co se / O ice de l’en i onnemen de la Co se), Vi gile Noble (Conse a oi e bo anique na ional médi e anéen), Sal a o e Pas a (CNR Pale mo), Es e anía San os Ba ea (Uni e sidad de Se illa) o hei help in acqui ing he da a o o hei commen s on he dis ibu ion o s a us o he di e en ee species. Funding This wo k was unded by he F ench Founda ion o Resea ch on Biodi e si y (FRB) h ough i s Cen e o Syn hesis and Analysis o Biodi e si y da a (CESAB) p og amme, as pa o he WOODIV esea ch p ojec . ACM and MCB we e unded by Labex OT-Med (n° ANR-11-LABX-0061). The unde s had no ole in s udy design, da a collec ion and analysis, decision o publish, o p epa a ion o he manusc ip . A ailabili y o da a and ma e ials A e accep ance da a will be made publicly a ailable as an addi ional ile. Au ho s’con ibu ions Concei ed and designed he da abase: FM, DP, ACM, GB, PD, TN and AL. Analyzed he da a: ACM and FM. W o e he pape : FM, ACM and TN. All au ho s pa icipa ed o he discussions leading o he aming o he manusc ip du ing WOODIV wo kshops. All au ho s ead, con ibu ed sugges ions and app o ed he inal manusc ip . E hics app o al and consen o pa icipa e No applicable. Consen o publica ion No applicable. Compe ing in e es s The au ho s decla e ha hey ha e no compe ing in e es s. Au ho de ails 1 Aix Ma seille Uni , A ignon Uni , CNRS, IRD, IMBE. Technopôle de l’A bois-Médi e anée, cedex 4, BP 80, 13 545 Aix-en-P o ence, F ance. 2 Facul y o Science, Uni e si y o Zag eb, Ho a o ac 102a, HR-10000 Zag eb, C oa ia. 3 Depa men o Biology, Di ision o Plan Biology, Labo a o y o Bo any, Uni e si y o Pa as, Uni e si y Campus, 26504 Rio, G eece. 4 Cen o Conse azione Biodi e si à, Dipa imen o di Scienze della Vi a e dell’Ambien e, Uni e si à degli S udi di Caglia i, Viale S. Ignazio da Laconi, 13, 09123 Caglia i, I aly. 5 Depa amen o de Biologia Vege al y Ecologia, Uni e sidad de Se illa, Apa ado 1095, 41080 Se illa, Spain. 6 Depa men o Bo any, Hunga ian Na u al His o y Museum, P . 137, Budapes 1431, Hunga y. 7 So bonne Uni e si y, IEES Bâ 44-34, cc 237. 4, Place Jussieu, Pa is, F ance. 8 Depa men o Ag icul u e, Food and Fo es Sciences, Uni e si y o Pale mo, Viale delle Scienze bldg. 4, 90128 Pale mo, I aly. 9 INRA, UR629, Ecologie des o ê s médi e anéennes, A ignon, F ance. 10 Macedonian Academy o Sciences and A s, K s e Misi ko 2, 1000 Skopje, Republic o Macedonia. 11 EcoGozo, Regional De elopmen Di ec o a e - Minis y o Gozo, Fla 6, Sunse Cou B, T iq Ma sal o n, Xagh a, Gozo, Mal a. 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