A icle h ps://doi.o g/10.1038/s41467-023-36240-6
Clima e- ai ela ionships exhibi s ong
habi a specifici y in plan communi ies
ac oss Eu ope
S ephan Kambach
1,2
, F ancesco Ma ia Saba ini
1,2,3,4
, Fabio A o e
5
,
Idoia Biu un
6
, Ge ha d Boenisch
7
, Gianma ia Bona i
8
,And ažČa ni
9,10
,
Ma ia Lau a Ca anza
11
, Alessand o Chia ucci
3
,MilanChy ý
12
,
Jü gen Dengle
13,14
, Emmanuel Ga bolino
15
,Valen inGolub
16
, Behlül Güle
17
,
U e Jand
1,2
,JanJansen
18
, Anni Jaško á
12
, Bo ja Jiménez-Al a o
19
,
Di k Nikolaus Ka ge
20
, Jens Ka ge
2,7
, Ilona Knollo á
12
,Gab ieleMidolo
12
,
Jespe E enskjold Moeslund
21
, Remigiusz Pielech
22
,
Vale ijus Rašoma ičius
23
,Sol i aRūsiņa
24
,Joze Šibík
25
,
Z jezdana S ančić
26
, Angela S anisci
11
, Jens-Ch is ian S enning
27
,
Se gey Yamalo
28
, Niklaus E. Zimme mann
20
& Helge B uelheide
1,2
Ecological heo y p edic s close ela ionships be ween mac oclima e and
unc ional ai s. Ye , global clima ic g adien s co ela e only weakly wi h he
ai composi ion o local plan communi ies, sugges ing ha impo an ac-
o s ha e been igno ed. He e, we in es iga e he consis ency o clima e- ai
ela ionships o plan communi ies in Eu opean habi a s. Assuming ha local
ac o s a e be e accoun ed o in mo e na owly defined habi a s, we
assigned > 300,000 ege a ion plo s o hie a chically classified habi a s and
modelled he e ec s o clima e on he communi y-weigh ed means o ou key
unc ional ai s using gene alized addi i e models. We ound ha he p e-
dic i e powe o clima e inc eased om b oadly o na owly defined habi a s
o specific lea a ea and oo leng h, bu no o plan heigh and seed mass.
Al hough mac oclima e gene ally p edic ed he dis ibu ion o all ai s, i s
e ec s a ied, wi h habi a -specifici y inc easing owa d mo e na owly
defined habi a s. We conclude ha mac oclima e is an impo an de e minan
o e es ial plan communi ies, bu u u e p edic ions o clima ic e ec s
mus conside how habi a s a e defined.
P edic ing he e ec s o a changing clima e on he di e si y and unc-
ioning o he ecosphe e equi es an unde s anding o how clima e
d i es he dis ibu ion o plan species and ecosys em p ope ies1,2.
Ecosys em unc ioning, such as p oduc i i y and nu ien cycling, is
s ongly de e mined by he unc ional composi ion o he plan
communi y3–6. Func ional ai s ep esen species’li e-his o y
s a egies7, and a e o en summa ized wi h a ew main, la gely inde-
penden , axes o a ia ion, such as he as -slow con inuum8,as
eflec ed in he lea -economics spec um9, hespecies’ ep oduc i e
s a egy10, he plan size spec um7, and he con inuum o collabo a ion
wi h myco hizal ungi11. A ounda ional, ye globally weakly suppo ed,
assump ion in ai -based ecology is ha he geog aphical dis ibu ion
o dominan unc ional ai s in plan communi ies is shaped by mac o-
en i onmen al g adien s, independen ly o axonomy12–14. He e, we
add essed his assump ion by s udying he consis ency o
mac oclima e- ai ela ionships among Eu opean plan communi ies.
Recei ed: 11 Ap il 2022
Accep ed: 20 Janua y 2023
Check o upda es
A ull lis o a filia ions appea s a he end o he pape . e-mail: s [email protected]
Na u e Communica ions | (2023) 14:712 1
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The mos popula app oach o s udying clima e- ai ela ionships
in ol es eg ession analysis be ween clima ic g adien s and he
communi y-weigh ed mean o plan unc ional ai s (CWMs15). A he
global scale, howe e , CWMs ha e been weakly p edic ed by linea
g adien s o clima ic condi ions13. Ye , in Aus alia, which can be con-
side ed a ela i ely homogeneous e olu iona y uni , clima e p edic ed
43% o he a ia ion in CWMs16, sugges ing ha global clima e- ai
ela ionships migh be blu ed by di e en phylogene ic his o ies
when modelled ac oss biogeog aphic ealms. Fu he mo e, hese
ela ionships could also be significan ly a ec ed by habi a -specific
ac o s17. When conside ing local plan communi ies, CWMs a e likely
shaped by soil condi ions18,19, mic oclima e20, human land use21,
dis u bance22, bio ic in e ac ions23 (e.g. he bi o y o compe i ion24)
and e olu iona y his o y ha cons ains he a ailable species pool25.All
o hese ac o s likely in e ac wi h mac oclima ic g adien s. Hence,
hey may mask he ac ual e ec s o clima e on local plan communi ies.
The classifica ion o plan communi ies in o hie a chically a an-
ged habi a ypes (he ea e habi a s) o e s a p omising app oach o
s udying clima e- ai ela ionships because i accoun s o in e e ing
local ac o s ha define di e en habi a s. Res ic ing analyses o
habi a s wi h simila flo is ic composi ion, sha ed e olu iona y his o y
and compa able en i onmen al condi ions could disen angle he
e ec s o in e ac ing ac o s and esul in igh e clima e- ai ela-
ionships, as has been obse ed in analyses ocused on specifichabi-
a s (e.g. o es and und a habi a s20,26). Ye , i is unclea whe he he
s eng h and exp ession o clima e- ai ela ionships depend on he
hema ic esolu ion (he ea e “na owness”)o hehabi a defini ion.
Analysing clima e- ai ela ionships a di e en le els o habi a na -
owness could help us unde s and whe he hese ela ionships a e
consis en ac oss bo h le els ( esolu ion-in a ian ) and habi a s (i.e.,
habi a -specific). This unde s anding migh be impo an o mo e
accu a e p edic ions o he e ec s o a changing clima e on he dis-
ibu ion and unc ioning o na u al plan communi ies.
We used >300,000 geo- e e enced ege a ion plo s (Supple-
men a y Fig. S1) om he Eu opean Vege a ion A chi e (EVA27), 19
high- esolu ion bioclima ic a iables om he CHELSA da abase28,29
and species-le el plan ai in o ma ion om he TRY da abase30,31.
Each plo was cha ac e ized by he CWM o ou key unc ional ai s:
plan heigh , specific lea a ea (SLA), seed mass and specific oo leng h
(SRL) (Table 1). P e ious esea ch has shown ha hese ou ai s can
cap u e he majo g adien s in he global spec um o plan o m and
unc ion (plan heigh , SLA and seed mass7) and in he oo economic
spec um (SRL11). Each plo was classified o h ee hie a chical le els o
he EUNIS habi a classifica ion32,33 which include eigh b oadly defined
le el 1 habi a s (he ea e b oad habi a s), 40 mo e na owly defined
le el 2 habi a s (in e media e habi a s) and 216 o he mos na owly
defined le el 3 habi a s (na ow habi a s)33. We applied gene alized
addi i e mixed-e ec s models ha accoun ed o spa ial dependence
among plo s o de e mine he linea e ec s o he p incipal compo-
nen s o he Eu opean clima ic g adien s (clima e PCs) on he dis-
ibu ion o he ou CWMs. We hypo hesized ha (i) clima e has a
gene al e ec on CWMs ac oss habi a s (consis en wi h p e ious
findings, see Table 1), bu (ii) classi ying plo s in o mo e na owly
defined habi a s inc eases hep opo ion o CWM a ia ion ha can be
explained by clima e. Fu he mo e, we examined whe he na owly
defined habi a s exhibi clima e- ai ela ionships simila o hose
obse ed in he mo e b oadly defined habi a s. We show ha he
obse ed e ec s o mac oclima e on he dis ibu ion o plan unc-
ional ai s a e sensi i e o he ype and na owness o he plan
communi ies s udied.
Resul s
Clima e- ai ela ionships ac oss b oad habi a s
We summa ized he in o ma ion om he 19 bioclima ic a iables in o
ou p incipal componen s (PCs) using p incipal componen analysis.
These ou g adien s join ly cap u ed 88.4% o he a ia ion in mac-
oclima ic condi ions (Supplemen a y Fig. S1) and ep esen ed he
Eu opean g adien s om suba c ic o d y-summe Medi e anean cli-
ma es (PC1, he ea e Medi e anean g adien ), om colde con-
inen al o wa me coas al clima es (PC2, empe a u e g adien ), om
colde o wa me summe clima es (PC3, summe - empe a u e g a-
dien ) and a g adien o inc easing seasonali y o p ecipi a ion and
empe a u e egimes (PC4, seasonali y g adien ). All ou clima ic
g adien s we e significan p edic o s o he ou CWMs ac oss all b oad
habi a s (Fig. 1and Supplemen a y Fig. S2). An inc ease in he Medi-
e anean g adien was associa ed wi h highe communi y-weigh ed
plan heigh , seed mass and SRL and lowe SLA. An inc ease in em-
pe a u e g adien was also associa ed wi h highe communi y-
weigh ed plan heigh , seed mass and SRL and lowe SLA. The posi-
ions o he b oad and in e media e habi a ypes along he ou cli-
ma ic PCs a e shown in Supplemen a y Figs. S3–S6.
Clima e- ai ela ionships along he hie a chy o habi a s
Wi hin he b oad habi a s, we obse ed se e al de ia ions om he
gene al clima e- ai s ela ionships obse ed ac oss habi a s. Along he
Medi e anean g adien , communi y plan heigh dec eased in o es
and coas al habi a s; SLA inc eased in we lands and SRL dec eased in
coas al sal ma shes, o es s, and man-made habi a s. Along he em-
pe a u e g adien , plan heigh dec eased in coas al habi a s; SLA
inc eased in ou o he eigh b oad habi a s and SRL dec eased in
hea hland and o es habi a s. The e ec s o clima e on communi y
seed mass we e gene ally mo e consis en han o he o he h ee
ai s (Fig. 1and Supplemen a y Fig. S2).
Habi a was gene ally a be e p edic o o CWMs han he com-
bined fixed and habi a -specific e ec s o clima e. In models ha did
no accoun o he habi a -specifici y o clima e- ai ela ionships (Cl
+b oad in Fig. 2), communi y plan heigh was he ai bes explained
by b oad habi a s (wi h a maximum o 77.2% o explained a ia ion),
ollowed by seed mass (67.1%), SLA (53.3%) and SRL (45.6%). When
accoun ing o habi a -specifici y in na ow habi a s (see Cl*na ow in
Fig. 2), hese alues inc eased o 92% o plan heigh , 73.9% o seed
mass, 63.7% o SLA, and 53.6% o he a ia ion in SRL. Descending he
classifica ion hie a chy owa ds mo e na owly defined habi a s, we
ound ha he p opo ion o a ia ion in CWMs explained by clima ic
condi ions inc eased o wo o he ou plan ai s ( om Cl*b oad o
CI*na ow in Fig. 2). Fo SLA and SRL, he p opo ion o explained
a ia ion inc eased owa d mo e na owly defined habi a s (wi h a
maximum o 10.3% and 8.0% o explained a iance, espec i ely)
whe easplan heigh andseedmassshowednosuchpa e n(wi ha
maximum o 5.7% and 6.8% o explained a iance, espec i ely). Fo
plan heigh , SLA and seed mass, we ound ha clima e- ela ed a ia-
ion in CWMs was maximized when habi a ype was no accoun ed o
(see Cl in Fig. 2). Ne e heless, he habi a -specific in e ac ion e ms in
ou models explained a significan p opo ion o CWM a iabili y o
all ai s and all le els o he classifica ion hie a chy (Supplemen a y
Table S1).
Clima e- ai ela ionships in in e media e and na ow habi a s
In he in e media e-le el habi a s, he obse ed clima e- ai ela ion-
ships mos ly ma ched hose in he mo e b oadly defined habi a s,
albei wi h an inc eased p opo ion o nonsignifican and some imes
con as ing ela ionships (see Fig. 3 o he exempla y ela ionships o
he fi s and second p incipal componen s wi h plan heigh , Supple-
men a y Fig. S7 o he hi d and ou p incipal componen s and
Supplemen a y Figs. S8–S13 o he o he ai s wi h he ou p incipal
componen s). Wi hin na ow habi a s, we obse ed ha clima e- ai
ela ionships we e equen ly habi a -specific, o en wi h con as ing
ela ionships among habi a s ha we epa o hesamesupe io
b oad habi a (Fig. 4). This habi a -specifici y o clima e- ai ela ion-
ships was obse ed o all ai s and in all b oad habi a s.
A icle h ps://doi.o g/10.1038/s41467-023-36240-6
Na u e Communica ions | (2023) 14:712 2
Discussion
Unde s anding whe he clima e- ai ela ionships a e habi a -specific
o gene al, and whe he hese ela ionships a e in a ian o he na -
owness o he applied habi a defini ion could imp o e ou p edic-
ions o how plan communi ies will be a ec ed by u u e clima e
change. In his s udy, we summa ized he e ec s o clima e on ou key
unc ional plan ai s—plan heigh , SLA, seed mass and SRL– om
b oadly o mo e na owly defined habi a s in Eu ope. While clima e
was ound o be a significan p edic o o CWMs o all ou plan ai s,
bo h wi hin and ac oss b oad habi a s, ou esul s e ealed h ee key
findings. Fi s , habi a ype was gene ally a be e p edic o o ai
dis ibu ions han clima e pe se. Ye , o plan heigh , SLA and seed
mass, clima e and habi a sha ed some o he explained a ia ion in
ai dis ibu ion, sugges ing ha clima e also s ongly influences he
b oad dis ibu ion o habi a ypes (see Supplemen a y Figs. S3–S6).
Second, decomposing he analysis in o inc easingly na ow habi a s
e ealed s onge clima e- ai ela ionships o SLA and SRL bu no
o plan heigh and seed mass. Thi d, in he mos na owly defined
habi a s, he e ec s o clima e on he exp ession o all ou ai s we e
s ongly habi a -specific, ega dless o which b oad habi a ype o
clima ic g adien was conside ed. By using inc easingly na ow habi a
defini ions, we we e able o educe he e ec s o local ac o s on
CWMs and hus be e un eil he e ec s o mac oclima e on he
unc ional ai composi ion o plan communi ies.
The habi a classifica ion we used is based on flo is ic and bio-
geog aphic cha ac e is ics. As a esul , plan communi ies we e ana-
lysed wi hin homogeneous g oups ha eflec common adap a ions o
local en i onmen al ea u es ha a e ei he independen o global
clima ic g adien s34 o di ficul o cap u e wi h global da ase s on cli-
ma e and soil condi ions. We ound ha he e ec s o clima e on plan
communi y composi ion in b oad habi a s a e no necessa ily eflec ed
in simila e ec s in mo e na owly defined habi a s35. Fo ins ance,
communi y-weigh ed mean plan heigh inc eased o e all in g assland
habi a s along he Medi e anean g adien , bu his pa e n e e sed
when ocusing on seasonally we and we g asslands (habi a R3).
Simila ly, we ound ha highe empe a u es led o highe communi y
plan heigh in we land habi a s, excep o pe iodically exposed
sho es (Q6), whe e empe a u e was nega i ely ela ed o plan heigh .
Ou esul s on he habi a -specifici y o clima ic e ec s a e in acco -
dance wi h p e ious findings on he dis inc e ec s o p ecipi a ion on
ege a ion p oduc i i y in g assland e sus o es habi a s in China36.
On he global scale, he p edic i e powe o clima e o he dis ibu ion
o communi y lea ai composi ion inc eased when 14 a he han
ou b oad habi a ypes we e conside ed37.Thisfinding p o ides an
explana ion o why s udies in mo e na owly defined habi a s
e ealed mo e p onounced e ec s o clima e on plan ai s (e.g. as
shown in und a habi a s26) han s udies ac oss habi a s13.The wo
communi y ai s ha we e less well p edic ed by habi a alone
(namely SLA and SRL) we e inc easingly be e p edic ed by clima e
when habi a s we e mo e na owly defined. The unde lying eason
migh be ha SLA depends mo e s ongly on soil han on clima ic
condi ions19,38 and ha he ela ionship be ween SRL and en i on-
men al g adien s also s ongly depends on he habi a conside ed39.
Taken oge he , hese obse a ions sugges ha b oad habi a s
include species wi h di e en li e-his o y s a egies (in e ms o com-
pe i ion o ligh and collabo a ion wi h myco hizal ungi), while mo e
na owly defined habi a s include species wi h mo e simila li e-his o y
s a egies. Consequen ly, b oad habi a s migh allow o a wide ange
o al e na i e clima ic esponses, while in na ow habi a s, species end
o sha e simila clima ic esponses. Thus, he e ec o clima e on he
p opo ion o species wi h high o low SLA (o SRL) becomes clea e .
The wo ai s ha did no ollow his pa e n, i.e. communi y plan
heigh andseedmass,a es onglyco ela edwi hin heglobalspec-
um o plan o m and unc ion7,40 and mos s ongly de e mined by
habi a ype. Communi y plan heigh is ela ed o he occu ence o
Table 1 | Desc ip ion o he analysed unc ional ai s
T ai Ecological unc ion Repo ed ela ionships wi h Re e ences
p ecipi a ion empe a u e
Plan heigh Cen al o species’ca bon alloca ion s a egy and abili y o compe e o ligh . Posi i e Posi i e 16,53,67
Specific lea a ea (SLA) = a io o lea a ea o
d y mass
Cen al o species’acquisi ion s a egy along he lea economic spec um. Nonsignifican o posi i e Nega i e o posi i e 9,16,37,53
Seed mass De e mines dispe sal and seedling su i al unde un a ou able condi ions. Nonsignifican o posi i e Nonsignifican o posi i e 53,68–70
Specific oo leng h (SRL) = a io o oo leng h o
d y mass
Cen al o species’collabo a i e s a egy as species wi h myco hizal associa ions end o ha e
lowe SRL ( hicke oo s).
Nonsignifican o posi i e Nega i e 39,71,72
Re e enced s udies we e conduc ed ac oss la ge clima ic g adien s and mul iple habi a s.
A icle h ps://doi.o g/10.1038/s41467-023-36240-6
Na u e Communica ions | (2023) 14:712 3
woody species, which ma kedly depends on land use and is also mo e
s ongly de e mined by soil mois u e and nu ien s han by b oad
clima ic condi ions41.
Al hough he in es iga ed clima e- ai ela ionships we e
mos ly consis en ac oss b oadly defined habi a s, we ound se e al
habi a -specific ela ionships. Fo ins ance, he inc ease in plan heigh
along he Medi e anean and empe a u e g adien s could be
explained by he longe g owing season and highe ligh a ailabili y,
which bo h inc ease compe i ion o ligh in wa me egions42,43.In
o es habi a s, he dec ease in plan heigh along he Medi e anean
g adien could be a ibu ed o he limi ing e ec s o highe
empe a u es and lowe wa e a ailabili y on he g ow h o canopy
ees. The dis ibu ion o o es unde s o y ai s, howe e , is ai ly
independen o p ecipi a ion g adien s in Eu ope20. Communi y heigh
in coas al habi a s could be limi ed by p ecipi a ion because dune soils
a e highly pe meable (and hus edaphically d y) and he g oundwa e
is no a ailable o mos plan s44. Fo SLA, we obse ed a decline along
he Medi e anean g adien (i.e., SLA dec eased wi h inc easing
d ough s ess), co esponding o he global13 and in a-specific
educ ion o SLA unde d ough 45. The same a gumen s apply o he
e ec s o empe a u e. We land communi ies, o ins ance, migh be
able o main ain a highe SLA, e en unde highe empe a u es,
PC1: Medi e anean g adien
dec easing ↔inc easing
PC2: empe a u e g adien
dec easing ↔inc easing
Plan Heigh SLA
Seed MassSRL
−10 0 10 −10 −5 0 5
−2
−1
0
1
2
−2
−1
0
1
2
−2
−1
0
1
2
−2
−1
0
1
2
P incipal componen sco es
Habi a ype
Coas al sal ma shes (MA)
Cos al habi a s (N)
We lands (Q)
G asslands (R)
Hea hlands (S)
Fo es s (T)
Spa sely ege a ed (U)
Man−made habi a s (V)
Signi icance
signi ican
nonsigni ican
Fig. 1 | E ec s o clima e on ou plan ai s in b oad habi a s. The g aphs show
he communi y-weigh edmeans o ou plan unc ional ai s as linea unc ions o
he fi s and second p incipal componen s (PCs) o he 19 CHELSA bioclima ic
a iables, ob ained wi h gene alized addi i e mixed-e ec s models. Slopes show
he ela ionships ac oss all (black) and wi hin he mos b oadly defined habi a s o
he EUNIS classifica ion (colou s). Solid lines indica e significan ela ionships a
p< 0.05 (based on sepa a e wo-sided es s). G ey hexagons show he dis ibu ion
o plo -le el obse a ions. SLA specific lea a ea, SRL specific oo leng h.
A icle h ps://doi.o g/10.1038/s41467-023-36240-6
Na u e Communica ions | (2023) 14:712 4
because hey a e no cons ained by wa e a ailabili y. G assland
species migh benefi om highe SLA in wa me empe a u es i hey
a e able o compensa e o na u al o human-induced dis u bances by
eg owing ela i ely quickly. In man-made habi a s, he esponse o
SLA migh depend on he ela i e abundance o di e en li e o ms,
wi h he ophy es being ela i ely mo e abundan a highe
empe a u es46. Communi y seed mass showed he g ea es con-
sis ency in clima e- ai ela ionships ac oss all b oad habi a s. Ye , he
inc ease in seed mass along he Medi e anean g adien was qui e
unexpec ed, as seed mass ends o inc ease wi h p ecipi a ion and
dec ease wi h a idi y on he global scale13 o is only weakly ela ed o
p ecipi a ion in Mongolia, China47 and Aus alia48. In habi a s whe e
wa e a ailabili y depends on g oundwa e , such as coas al sal ma -
shes, coas al habi a s and we lands, communi y seed mass was inde-
penden o he Medi e anean g adien . An explana ion migh be ha
la ge seeds, which can bu e he su i al o seedlings and saplings49,
a e mos impo an du ing ho and d y summe s. Fo communi y SRL,
he obse ed inc ease along he Medi e anean and empe a u e g a-
dien s e u ed ou expec a ions. In coas al sal ma shes, hea hland and
o es habi a s, howe e , we also obse ed dec eases in SRL along bo h
g adien s, sugges ing ha he e ec s o mac oclima e may be s ongly
habi a -specific o masked by local ac o s, such as local soil
condi ions19. These local condi ions we e inc easingly accoun ed o
by including mo e na owly defined habi a s in he models o clima e-
SRL ela ionships.
In o de o syn hesize he e ec s o mac oclima e ac oss a la ge-
scale da ase , we had o accep some sho comings in ou analysis. We
ocused only on he linea e ec s o clima e and igno ed in e ac ions
among he di e en dimensions o clima e, al hough we can assume
ha he dis ibu ion o he ou plan ai s is also shaped by nonlinea
and in e ac ion e ec s (c. . ee heigh 50 and lea ai s20). We pa ially
accoun ed o non-linea i y by using log- ans o med CWMs. S ill, ou
models iden ified s onge linea e ec s o clima e on communi y
ai s compa ed o global-scale analyses13. Whe he he classifica ion
in o na owe habi a s migh ha e s eng hened o alle ia ed any
nonlinea o in e ac ion e ec s has, o ou knowledge, no been es ed
ye . By ocusing exclusi ely on clima e, we did no accoun o
impo an in e ac ions wi h edaphic condi ions (such as e apo-
anspi a ion), which a e he bes p edic o o lea a ea and maximum
plan heigh dis ibu ions ac oss Aus alia16. Fu he mo e, i is con-
cei able ha he e ec s o clima e on communi y ai composi ion
could change a he end o clima ic g adien s. This assump ion could
be es ed by compa ing ai -en i onmen ela ionships unde benign
e sus ex eme condi ions39. In o es s, we did no di e en ia e
be ween he canopy and unde s o y laye s, al hough hese laye s
migh show con as ing esponses o clima e20. Finally, we calcula ed
CWMs based on species-le el ai a e ages a he han in si u mea-
su emen s, which p ecluded any analysis o in a-specific ai a ia-
ion. These sho comings may ha e dilu ed he e ec s o clima e on
CWMs, bu we e una oidable gi en he la ge geog aphic ex en o
ou s udy.
Ou esul s show ha mac oclima e is a consis en p edic o o
he CWMs o plan ai s in all habi a s and ac oss all clima ic g adien s
in Eu ope. These e ec s o clima e a y ac oss habi a s, depending on
he hema ic esolu ion o he habi a defini ion and he iden i y o he
habi a . We hus an icipa e ha clima e-change e ec s will no be
limi ed o pa icula ly sensi i e communi ies51, bu will ha e di e se
e ec s on a wide ange o plan communi ies52. To accu a ely p edic
he e ec s o a changing clima e on he composi ion and unc ioning
o he ecosphe e, we ecommend conside ing he habi a -specifici y o
clima e- ai ela ionships.
Me hods
Vege a ion da a
The aw ege a ion su ey da a consis ed o 1,741,856 plo s wi h
37,318,600 species eco ds om 107 da abases, colla ed and cu a ed
by he Eu opean Vege a ion A chi e (EVA)27 and accessed on May 12 h,
2021. Indi idual da abases a e lis ed in Supplemen a y Da a 1. Species
abundances we e measu ed o con e ed o pe cen age co e . Be o e
he analysis, we emo ed all plo s (1) wi h only p esence/absence da a,
(2) wi hou geog aphic coo dina es, (3) si ua ed in G eenland, (4) wi h
la i udinal coo dina es lowe han he sou he nmos poin o 34° N o
highe han he no he nmos poin o 82° N, (5) wi h ai in o ma ion
o less han 80% o he o al plan co e and (6) o which clima ic
a iables could no be calcula ed due o emp y pixels a ound he ocal
plo . To educe he e ec s o spa ially clus e ed and epea ed cen-
suses, we only e ained he mos ecen census o each unique loca-
ion and andomly s a ified he da ase o include only one plo pe
Plan Heigh SLA Seed Mass SRL
Clima eHabi a
Cl
Cl + b oad
Cl * b oad
Cl * in e media e
Cl * na ow
Cl
Cl + b oad
Cl * b oad
Cl * in e media e
Cl * na ow
Cl
Cl + b oad
Cl * b oad
Cl * in e media e
Cl * na ow
Cl
Cl + b oad
Cl * b oad
Cl * in e media e
Cl * na ow
0
5
10
15
75
50
25
0
Explained a ia ion (%)
Fig. 2 | P opo ion o a ia ion in plan ai s explained by clima e and
inc easingly na ow habi a defini ions. The g aphs show ma ginal R² alues
om gene alized addi i e mixed-e ec s models o he linea dependence o he
communi y-weigh ed means o ou plan unc ional ai s on habi a ( ed ba s) and
he gene al and habi a -specific e ec s o clima e (blue ba s). Model complexi y
inc eases om le o igh . Cl: e ec s o clima e ac oss all plo s o habi a s,
modelled wi h fixed e ec s o he ou p incipal componen s o he 19 bioclima ic
a iables. CL + b oad: fixed e ec s o clima e plus andom in e cep e ec s o
b oad habi a s. Cl*b oad/in e media e/na ow: fixed e ec s o clima e plus he
andom e ec s o b oad, in e media e, o na owly defined habi a s plus he
habi a -specific e ec s o clima e, modelled wi h andom-slope e ec s be ween he
clima ic p incipal componen s and he espec i e habi a s. SLA specificlea a ea,
SRL specific oo leng h.
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MA22
MA24
MA25
N1
N2
N3
Q1 Q2
Q4
Q5
Q6
R1
R2 R3
R4
R5
R6
S1
S2
S3 S4
S5
S6
S7
S9
Sa
Sb
T1
T2
T3
U2
U3
U7
V1 V3
MA22
MA24
MA25
N1
N2
N3
Q1
Q2
Q4
Q5
Q6
R1
R2
R3
R4
R5
R6
S1
S2
S3
S4 S5
S6
S7
S9 Sa
Sb
T1
T2
T3
U2
U3
U7
V1
V3
PC1: Medi e anean g adien
dec easing ↔inc easing
PC2: empe a u e g adien
dec easing ↔inc easing
Coas al
sal ma shes (MA)
Cos al
habi a s (N)
We lands (Q)
G asslands (R)
Hea hlands (S)
Fo es s (T)
Spa sely
ege a ed (U)
Man−made
habi a s (V)
−10 0 10 −5 0 5
−0.8
−0.6
−0.4
−0.8
−0.4
0.0
−1.0
−0.5
0.0
0.5
−1.5
−1.0
−0.5
0.0
−1
0
1
1.0
1.2
1.4
−1.5
−1.0
−0.5
0.0
−0.7
−0.6
P incipal componen
Plan Heigh
Indi idual slopes
posi i e
nega i e
nonsigni ican
Fig. 3 | E ec s o clima e on plan heigh in in e media e-le el habi a s. The
g aphs show he communi y-weigh ed means o plan heigh as a linea unc ion o
he fi s and second p incipal componen s (PCs) o he 19 CHELSA bioclima ic
a iables, ob ained om gene alized addi i e mixed-e ec s models. Significance
was de e mined a p< 0.05 (based on sepa a e wo-sided es s). Slope es ima es
om habi a s wi h ewe han 100 plo obse a ions we e omi ed. Codes wi hin
boxes e e o habi a ypes (as lis ed in Supplemen a y Da a 2).
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Na u e Communica ions | (2023) 14:712 6
1
5
1
1
11
1
17
9
5
22
11
3
8
12
2
5
6
6
2
14
4
1
3
3
6
7
3
12
16
5
20
11
2
10
10
2
2
7
1
5
2
5
9
4
1
6
2
10
1
19
6
6
18
16
2
11
11
6
5
4
4
6
11
1
5
2
8
5
6
10
15
8
21
7
1
13
8
1
10
1
5
2
7
9
2
Plan Heigh SLA Seed Mass SRL
Coas al sal ma shes (MA)
Cos al habi a s (N)
Fo es s (T)
G asslands (R)
Hea hlands (S)
Man−made habi a s (V)
Spa sely ege a ed (U)
We lands (Q)
Rela ionships wi h PC1: Medi e anean g adien
2
5
3
2
8
12
12
7
15
16
5
9
12
1
5
4
2
3
5
11
7
1
5
1
9
4
5
9
17
10
21
5
3
10
9
2
6
3
1
7
6
8
4
2
4
1
4
9
14
14
3
13
22
1
13
9
4
6
1
8
6
11
1
7
2
9
2
1
11
19
11
21
4
11
11
7
4
1
7
4
11
3
Plan Heigh SLA Seed Mass SRL
Coas al sal ma shes (MA)
Cos al habi a s (N)
Fo es s (T)
G asslands (R)
Hea hlands (S)
Man−made habi a s (V)
Spa sely ege a ed (U)
We lands (Q)
Rela ionships wi h PC2: empe a u e g adien
Rela ionships
posi i e
nonsigni ican
nega i e
Fig. 4 | E ec s o clima e on plan ai s in na owly defined habi a s.The g aphs
show he numbe o slope es ima es be ween he fi s and second p incipal com-
ponen s (PCs) o he 19 CHELSA bioclima ic a iables and he communi y-weigh ed
mean o ou unc ional ai s, calcula ed wi h gene alized addi i e mixed-e ec s
models. Colou ing indica es he exp ession and significance o he ela ionship.
Significance was de e mined a p< 0.05 (based on sepa a e wo-sided es s). Slope
es ima es om habi a s wi h ewe han 100 plo obse a ions we e omi ed. SLA
specific lea a ea, SRL specific oo leng h.
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0.01° g id cell (app oxima ely 1.11 × 0.69 km in Cen al Eu ope). The
esul ing da ase con ained 300,021 ege a ion plo s wi h 7,320,239
indi idual species occu ence obse a ions ha we e conduc ed
be ween 1873 and 2020 and dis ibu ed be ween 34.8° N o 80° N
la i ude and 10° W o 59.3° E longi ude wi h a ocus in Cen al Eu ope
(Supplemen a y Fig. S1).
Taxonomy
We ha monized species names om he Eu opean Vege a ion A chi e
using he Taxonomic Name Resolu ion Se ice 5.0 ( n s.bienda a.o g).
Subspecies and a ie ies we e me ged a he species le el. Species
names we e ma ched o he axonomic backbone 3.0 o he sPlo
global ege a ion da abase53 o maximize compliance wi h species
names om he TRY plan ai da abase30. Fungi, b yophy es, algae,
and lichens we e emo ed om he da a, esul ing in 13,758 ascula
plan species.
Communi y-weigh ed ai means
We ex ac ed species- and genus-le el a e age alues o plan heigh ,
SLA, seed mass and SRL om he TRY plan ai da abase30 e sion 5,
which co e ed species-le el mean alues o 33 ai s o 50,404 spe-
cies, de i ed wi h Bayesian Hie a chical P obabilis ic Ma ix Fac o -
iza ion. This algo i hm p edic s indi idual-le el ai alues based on
he obse ed ai eco ds, obse ed ai - ai co ela ions, and he
axonomic hie a chy eflec ing phylogeny54,55. The oo -mean-squa e
e o o he z- ans o med p edic ed e sus a ailable ai alues an-
ged om 0.07 (seed mass) o 0.2 (SLA) (see Supplemen a y Fig. S14).
F om his gap-filled da ase , we assigned species-le el mean ai
alues o 6,025 species om ou da ase and o an addi ional
690 species, we assigned genus-le el mean ai alues om he gap-
filled da ase . The ela ionships be ween he ou ocal ai s and 24
ai s om he gap-filled da ase a e shown in Supplemen a y Fig. S15.
The o iginal publica ions o he da a used a e lis ed in Supplemen a y
Table S2. Wi hou impu ed alues, wewe eable o assign ai alues o
45%(plan heigh ), 28.4% (SLA), 34.4% (seed mass) and 5.4% (SRL) o all
species in he da ase used o he analysis, which accoun ed o an
a e age o 95.6%, 91.4%, 90.8% and 47.4% o plan co e a he plo
le el, espec i ely. Fo each plo , we calcula ed he CWM o he ou
plan ai s acco ding o he equa ion in13.
Habi a classifica ion
Each plo was assigned o a habi a ype based on he classifica ion
expe sys em EUNIS-ESy (Eu opean Na u e In o ma ion Sys em32,33),
upda ed on Oc obe 25 h, 2021. We used he fi s h ee le els o he
classifica ion hie a chy, anging om he highes le el 1 (b oad habi a
ypes, e.g. o es s—habi a code T) h ough he in e media e le el 2
(in e media e ypes, e.g. b oadlea ed deciduous o es s –T1) o he
lowes le el 3 (na ow ypes, e.g. empe a e Salix and Populus ipa ian
o es –T11). This classifica ion sys em included he ollowing b oadly
defined habi a ypes a le el 1: coas al sal ma shes (MA, 3,085 plo s),
coas al habi a s (N, 5,045 plo s), we lands (Q, 28,931 plo s), g asslands
(R, including lands domina ed by o bs, mosses o lichens, 107,457
plo s), hea hlands (S, including sh ubland, sc ub and und a, 25,866
plo s), o es s (T, including o he wooded land, 98,311 plo s), inland
spa sely ege a ed habi a s (U, 2,932 plo s) and ege a ed man-made
habi a s (V, 28,394 plo s). All in e media e and na ow habi a s a e
lis ed in Supplemen a y Da a 2. Fac shee s on he flo is ic composi ion
o EUNIS habi a s a e lis ed in33 and56.
Clima ic da a
Fo each plo , we ex ac ed he 19 bioclima ic a iables om he
CHELSA Clima ologies e sion 1.228,29 a a pixel esolu ion o 30 a c sec
(~1 km) using he cloud-based Google Ea h Engine pla o m. Biocli-
ma ic alues we e calcula ed as he a e age alue om all pixels wi hin
500 m om he plo coo dina es. Plo s wi h missing clima e da a in
adjacen pixels we e omi ed om u he analyses. Co a ia ion
among he 19 bioclima ic a iables was emo ed by a p incipal com-
ponen analysis. The fi s ou p incipal componen s (PCs) join ly
explained 88.4% o he clima ic a ia ion (Supplemen a y Fig. S1). We
used he loadings o he 19 bioclima ic a iables o ex ac he posi-
ions o all plo s along hese ou PCs, which we e used as p edic o s in
he subsequen analyses.
S a is ical analysis
P io o he ollowing analyses, all plo -le el CWMs we e log ans-
o med o app oxima e no mali y. We es ima ed he e ec s o he
ou clima ic PCs on he CWMs o he ou ai s using sepa a e
gene alized addi i e mixed-e ec s models ha always included a
spline-on- he-sphe e smoo hing e m (based on la i ude and
longi ude57) o accoun o spa ial dependence among plo s58. Fo
each o he ou ai s, we c ea ed he ollowing fi e di e en
models: Model 1 included only he fixed-e ec s o he ou clima ic
PCs wi hou any in e ac ions. Only in his model, we accoun ed o
he di e en numbe o obse a ions pe habi a ype by including
only he same numbe o andomly selec ed plo s om each o he
b oad habi a s (based on he minimum numbe o 2,932 plo s in U –
Spa sely ege a ed habi a s). Model 2 included he fixed-e ec s o
he ou clima ic PCs plus he andom e ec s o he b oad le el 1
habi a ypes. Model 3 included he fixed-e ec s o he ou clima ic
PCs, he andom e ec s o he b oad le el 1 habi a ypes plus
andom in e ac ion e ms be ween he fixed e ec s o he clima ic
PCs and he b oad le el 1 habi a ypes. Models 4–5 we e simila o
model 3 excep ha he b oad le el 1 habi a ypes and hei andom
in e ac ion e ms we e eplaced wi h he in e media e le el 2 o he
na ow le el 3 habi a ypes and hei espec i e in e ac ions wi h
he clima ic PCs. The implemen a ion o all models is documen ed
on Gi Hub. Model esiduals we e isually checked and a e shown in
Supplemen a y Figs. S16-S20.
Fo models 3–5, we quan ified he e ec s o clima e on ai
CWMs, bo h ac oss and wi hin habi a s, wi h p edic ed ma ginal
mean eg ession slopes. All eg ession slopes we e assigned an
app oxima e confidence in e al (slope es ima e ± 1.96 * s anda d
e o ) and confidence in e als ha did no include ze o we e con-
side ed significan a app oxima ely p< 0.05. We did no es
eg ession slopes ha we e based on ewe han 100 plo obse a-
ions. Fo each model, we calcula ed he p opo ion o a ia ion in
CWMs ha could be explained by clima ic g adien s (fixed e ec s o
clima ic PCs and in e ac ions be ween clima ic PCs and habi a ypes)
and habi a ype ( andom e ec s). The p opo ion o explained a -
ia ion was quan ified using he ma ginal and condi ional R²59, whose
calcula ion o andom-slope models was based on he a iance
componen s o he fixed and andom e ec s and whose imple-
men a ion we adop ed om he R sc ip p o ided by60. To check
whe he he ob ained esul s depended on he p opo ion o species
wi h ac ual ai measu emen s, we epea ed all analyses wi h models
in which he con ibu ion o each plo was weigh ed acco ding o he
summed co e o species wi h ai alues di ided by he o al plan
co e . The esul s o hese weigh ed analyses we e quali a i ely
simila o he esul s o he unweigh ed analyses and a e shown in
Figs. Supplemen a y S21–S25. The code implemen ed o calcula e he
ma ginal and condi ional R² o gene alized addi i e mixed-e ec s
models is documen ed on Gi Hub.
All analyses we e conduc ed in R61 using he package mgc o
gene alized addi i e mixed-e ec s models57,58,emmeans and gge ec s
o ma ginal mean eg ession slopes62,63,dismo o geog aphical s a-
ifica ion o plo s64 and ggplo 2 o da a isualiza ion65.
Repo ing summa y
Fu he in o ma ion on esea ch design is a ailable in he Na u e
Po olio Repo ing Summa y linked o his a icle.
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Na u e Communica ions | (2023) 14:712 8
Da a a ailabili y
The da a gene a ed in his s udy (plo -le el in o ma ion on plo coo -
dina es, su ey yea , CWMs and clima ic PCs) we e deposi ed in he
da a eposi o y o he Ge man Cen e o In eg a ed Biodi e si y
Resea ch (iDi ) Halle-Jena-Leipzig (h ps://doi.o g/10.25829/idi .3527-
g89e k). Raw ege a ion da a a e a ailable unde es ic ed access
because hey belong o he owne s o each ege a ion da abase, bu
can be eques ed a eu o eg.o g/e a-da abase-ob aining-da a. The
bioclima ic and plan ai da a used in his s udy can be downloaded
om chelsa-clima e.o g/bioclim and www. y-db.o g/T yWeb/dp.php,
espec i ely.
Code a ailabili y
Thecodeused oconduc he ollowings epso heanalysesis
a ailable a gi hub.com66 (h ps://doi.o g/10.5281/zenodo.7404176): (i)
ex ac ing he plo -le el bioclima ic a iables, (ii) unning and analys-
ing he gene alized addi i e mixed-e ec s models, (iii) c ea ing he
p esen ed figu es.
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