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
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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.
Recei ed: 17 Decembe 2018 Accep ed: 28 Feb ua y 2019
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