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Connec ing he mul iple dimensions o global soil ungal di e si y
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Mik yuko , Vladimi ; Dulya, Olesya; Zizka, Alexande ; Bah am, Mohammad;
Hagh-Dous , Nilou a ; Anslan, S en; P ylu skyi, Oleh; Delgado-Baque izo, Manuel;
Maes e, Fe nando T.; Nilsson, Hen ik; Pä n, Jaan; Öpik, Maa ja; Moo a, Ma i;
Zobel, Ma in; Espenbe g, Mikk; Mande , Ülo; Khalid, Abdul Nasi ; Co ales,
Ad iana; Agan, Ah o; Vasco-Palacios, Aída-M.; Sai a, Alessand o; Rinaldi, And ea;
Ve beken, Annemieke; Sulis yo, Bobby; Tamgnoue, Bo is; Fu neaux, B endan;
Dua e Ri e , Camila; Nyamukondiwa, Caspe ; Sha p, Ca hy; Ma ín, Césa ;
Goha , Daniyal; Kla ina, Da a; Sha mah, Dipon; Dai, Dong-Qin; Nouh a, Edua do;
Bie sma, Elisabe h Mach eld; Rähn, Elisabe h; Came on, E in; De C op, Eske;
O sing, E eli; Da ydo , E geny; Albo noz, Felipe; B ea ley, F ancis; Buegge ,
F anz; Zahn, Geo ey; Boni o G ego y; Hiiesalu, Inga; Ba io, Isabel; Heilmann-
Clausen, Jacob; Ankuda, Jelena; Doležal, Ji i; Kupagme, John; Maciá-Vicen e, Jose;
Djeugap Fo o, Joseph; Geml, Józse ; Ala alo, Juha; Al a ez-Manja ez, Julie a;
Põldmaa, Kad i; Runnel, Kad i; Adamson, Kale ; B å hen, Ka i-Anne; P i sch,
Ka in; Tchan, Issi ou Kassim; A molai is, Kęs u is; Hyde, Ke in; Newsham, Ke in
K.; Panksep, K is el; La ee , Adebola Azeez; Hansson, Linda; Lami , Louis; Saba,
Malka; Tuomi, Ma ia; G yzenhou , Ma ieka; Bau e s, Ma ijn; Piepenb ing, Meike;
Wijayawa dene, Nalin N.; Yo ou, Nou ou; Ku ina, Ola i; Mo ime , Pe e ; Meidl,
Pe e ; Kohou , Pe ; Puusepp, Rasmus; D enkhan, Rein; Ga ibay-O ijel, Robe o;
Godoy, Robe o; Alkah ani, Saad; Rahimlou, Saleh; Dudo , Se gey; Põlme, Se gei;
Ghosh, Soumya; Mund a, Sunil; Ahmed, Talaa ; Ne he way, Ta quin; Henkel,
Te y; Roslin, Tomas; N ezi yayo, Vincen ; Fedoso , Vladimi ; Onipchenko,
Vladimi ; Yasan hika, Wee agalle, A achchillage E andi; Lim, Young; Van Nuland,
Michael; Soudzilo skaia, Nadejda; An onelli, Alexand e; Kõljalg, U mas;
Aba enko , Kessy; Tede soo, Leho
Mik yuko , V., Dulya, O., Zizka, A., Bah am, M., Hagh-Dous , N., Anslan, S., P ylu skyi, O.,
Delgado-Baque izo, M., Maes e, F. T., Nilsson, H., Pä n, J., Öpik, M., Moo a, M., Zobel, M.,
Espenbe g, M., Mande , Ü., Khalid, A. N., Co ales, A., Agan, A., . . . Tede soo, L. (2023).
Connec ing he mul iple dimensions o global soil ungal di e si y. Science Ad ances, 9(48),
A icle eadj801. h ps://doi.o g/10.1126/sciad .adj8016
2023
ECOLOGY
Connec ing he mul iple dimensions o global soil
ungal di e si y
Vladimi Mik yuko
1
*†, Olesya Dulya
1
†, Alexande Zizka
2
, Mohammad Bah am
3
,
Nilou a Hagh-Dous
1
, S en Anslan
1
, Oleh P ylu skyi
4
, Manuel Delgado-Baque izo
5
,
Fe nando T. Maes e
6
, Hen ik Nilsson
7
, Jaan Pä n
1
, Maa ja Öpik
1
, Ma i Moo a
1
, Ma in Zobel
1
,
Mikk Espenbe g
1
, Ülo Mande
1
, Abdul Nasi Khalid
8
, Ad iana Co ales
9
, Ah o Agan
10
,
Aída-M. Vasco-Palacios
11
, Alessand o Sai a
12
, And ea Rinaldi
13
, Annemieke Ve beken
14
,
Bobby Sulis yo
14
, Bo is Tamgnoue
15
, B endan Fu neaux
16
, Camila Dua e Ri e
17
,
Caspe Nyamukondiwa
18
, Ca hy Sha p
19
, Césa Ma ín
20
, Daniyal Goha
21
, Da a Kla ina
22
,
Dipon Sha mah
23
, Dong-Qin Dai
24
, Edua do Nouh a
25
, Elisabe h Mach eld Bie sma
26,27
,
Elisabe h Rähn
10
, E in Came on
28
, Eske De C op
14
, E eli O sing
21
, E geny Da ydo
29
,
Felipe Albo noz
30
, F ancis B ea ley
31
, F anz Buegge
32
, Geo ey Zahn
33
, G ego y Boni o
34
,
Inga Hiiesalu
1
, Isabel Ba io
35
, Jacob Heilmann-Clausen
36
, Jelena Ankuda
37
, Ji i Doležal
38
,
John Kupagme
21
, Jose Maciá-Vicen e
39
, Joseph Djeugap Fo o
15
, Józse Geml
40
, Juha Ala alo
41
,
Julie a Al a ez-Manja ez
42
, Kad i Põldmaa
1,43
, Kad i Runnel
1
, Kale Adamson
10
,
Ka i-Anne B å hen
44
, Ka in P i sch
32
, Kassim Tchan Issi ou
45
, Kęs u is A molai is
46
, Ke in Hyde
47
,
Ke in K. Newsham
27
, K is el Panksep
48
, Adebola Azeez La ee
49,50
, Linda Hansson
51
,
Louis Lami
52
, Malka Saba
53
, Ma ia Tuomi
44
, Ma ieka G yzenhou
54
, Ma ijn Bau e s
55
,
Meike Piepenb ing
56
, Nalin N. Wijayawa dene
57
, Nou ou Yo ou
45
, Ola i Ku ina
58
,
Pe e Mo ime
59
, Pe e Meidl
60
, Pe Kohou
61
, Rasmus Puusepp
21
, Rein D enkhan
10
,
Robe o Ga ibay-O ijel
42
, Robe o Godoy
62
, Saad Alkah ani
63
, Saleh Rahimlou
21
, Se gey Dudo
64
,
Se gei Põlme
21,43
, Soumya Ghosh
54
, Sunil Mund a
65
, Talaa Ahmed
41
, Ta quin Ne he way
3
,
Te y Henkel
66
, Tomas Roslin
3
, Vincen N ezi yayo
67
, Vladimi Fedoso
64
, Vladimi Onipchenko
64
,
Wee agalle A achchillage E andi Yasan hika
47
, Young Lim
68
, Michael Van Nuland
69
,
Nadejda Soudzilo skaia
70
, Alexand e An onelli
71
, U mas Kõljalg
1,43
, Kessy Aba enko
43
,
Leho Tede soo
21,63
*
How he mul iple ace s o soil ungal di e si y a y wo ldwide emains i ually unknown, hinde ing he man-
agemen o his essen ial species- ich g oup. By sequencing high- esolu ion DNA ma ke s in o e 4000 opsoil
samples om na u al and human-al e ed ecosys ems ac oss all con inen s, we illus a e he dis ibu ions and
d i e s o di e en le els o axonomic and phylogene ic di e si y o ungi and hei ecological g oups. We show
he impac o p ecipi a ion and empe a u e in e ac ions on local ungal species ichness (alpha di e si y) ac oss
di e en clima es. Ou indings e eal how empe a u e d i es ungal composi ional u no e (be a di e si y)
and phylogene ic di e si y, linking hem wi h egional species ichness (gamma di e si y). We in eg a e ungi
in o he p inciples o global biodi e si y dis ibu ion and p esen de ailed maps o biodi e si y conse a ion
and modeling o global ecological p ocesses.
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The Au ho s, some
igh s ese ed;
exclusi e licensee
Ame ican Associa ion
o he Ad ancemen
o Science. No claim o
o iginal U.S. Go e nmen
Wo ks. Dis ibu ed
unde a C ea i e
Commons A ibu ion
License 4.0 (CC BY).
INTRODUCTION
As he la ges con ibu o o global biomass a e plan s and bac e ia
(1), soil-inhabi ing ungi play c ucial oles in main aining he
heal h, p oduc i i y, and nu ien cycling o e es ial ecosys ems
(2–4). Ye , ou unde s anding o ungal di e si y dis ibu ion
ac oss he globe is incomple e, hampe ing analysis and p edic ion
o di e si y- unc ioning ela ionships in changing en i onmen s.
The ela i ely ecen eme gence o high- h oughpu molecula
iden i ica ion o e s he ools o la ge-scale su eys o di e se
ungal communi ies (5–11). Pionee ing global s udies, inco po a -
ing hund eds o samples and ocusing on dominan phylo ypes (5,
7,9,12), ha e consis en ly epo ed a majo ole o p ecipi a ion in
soil ungal alpha di e si y (i.e., plo -scale di e si y) and he exis-
ence o a la i udinal di e si y g adien ( he pa e n o polewa d
declines o biodi e si y in insic o many la ge g oups o o gan-
isms). A ecen me as udy (13) in ol ing he e ogeneous da a om
a la ge numbe o soil samples (>3000) in e ed highe ungal di e -
si y in hype -a id and A c ic habi a s compa ed o much o he
opics, challenging he common unde s anding abou he global
dis ibu ion o ungal di e si y and o biodi e si y in gene al (5,7,
14–16).
While he biogeog aphic pa e ns seen in plan s and animals a e
explained by compe ing hypo heses in ol ing in e ac ions be ween
a ious ac o s o di e si y (16), he join e ec o key d i e s—
wa e and he mal ene gy supply—on he global dis ibu ion o
ungal alpha di e si y has no been assessed. The e is also a
no able lack o global da a p o iding join analysis o o he di e si y
aspec s, including be a di e si y (spa ial a iabili y o communi y
SCIENCE ADVANCES |RESEARCH ARTICLE
Mik yuko e al.,Sci. Ad . 9, eadj8016 (2023) 29 No embe 2023 1 o 15
Downloaded om h ps://www.science.o g a Jy askyla Uni e si y on No embe 30, 2023
composi ion), gamma di e si y ( egional species ichness), and
phylogene ic di e si y (inco po a ing he e olu iona y componen
o communi y di e si y). Conside ing hese di e si y aspec s o-
ge he is pi o al o comp ehending he spa ial o ganiza ion o bio-
di e si y and becomes an impe a i e in de elopmen o sus ainable
ecosys em managemen s a egies and e ec i e biodi e si y conse -
a ion schemes (15,17).
He e, we analyze global pa e ns o ungal axonomic and phy-
logene ic alpha, be a, and gamma di e si ies based on long- ead
DNA sequencing o ca. 4000 opsoil samples ( ig. S1), collec ed
du ing six global su eys ollowing compa able sampling and mo-
lecula analysis p o ocols: he Global Soil Mycobiome conso ium
(GSMc) (6), BIODESERT (18), MUSGONET and CLIMIFUN (19),
GlobalAM (20), and GlobalWe lands (21). The samples encompass
a wide a ie y o land co e ypes, including di e en ypes o
woody and he baceous plan communi ies, dese s, and ag icul u al
and u ban en i onmen s. To inc ease he disc imina o y powe o
he analysis compa ed o p e ious wo ks, we use he ull-leng h in-
e nal ansc ibed space (ITS) egion o he ibosomal RNA
( RNA) gene as a axonomic ma ke (22,23). The esul ing >14
million high-quali y sequences a e clus e ed in o 800,000 ope a-
ional axonomic uni s (OTUs; ig. S2) using a 98% simila i y
h eshold, a p ac ical comp omise o g ouping sequences oughly
co esponding o ungal species (6). Al hough he esul ing OTUs
may no always p ecisely ep esen dis inc species, hey se e as ax-
onomic p oxies whose dis ibu ion pa e ns a e likely simila o co -
esponding species.
While ungi exhibi immensely high unc ional di e si y, hey
a e, much like o he o ganisms, commonly ca ego ized by hei
mos p ominen oles in ecosys em unc ioning in o pa hogens,
symbion s, and decompose s o dead o ganic ma e . Le e aging
he expe -cu a ed FungalT ai s da abase (24), we ca ego ize
species in o unc ionally dis inc , minimally o e lapping g oups
( ig. S3 and able S1): a buscula myco hizal (AM) and ec omyco -
hizal (EcM) ungi, molds, nonmyco hizal Aga icomyce es (NMA;
mos ly ep esen ed by sap o ophic mac o ungi), and a g oup o
species ha a e nonsymbio ically bio ophic on a wide a ie y o o -
ganisms (he ea e “pa hogens”). We conside yeas s and nonyeas
unicellula ungi ha exhibi ma kedly di e en li es yles om he
p edominan mul icellula mycelial o ms sepa a ely (25,26). Gi en
hei impo ance om a public heal h s andpoin , we also dis in-
guish oppo unis ic human pa hogens (OHPs).
D awing on hese knowledge gaps and p e ious indings, we
add ess he ollowing ques ions. (i) Wha is he combined e ec
o p ecipi a ion and empe a u e on soil ungal alpha di e si y?
(ii) Do he global dis ibu ion and d i e s o alpha di e si y a y
1
Ins i u e o Ecology and Ea h Sciences, Uni e si y o Ta u, Ta u 50409, Es onia.
2
Depa men o Biology, Philipps-Uni e si y, Ma bu g 35032, Ge many.
3
Depa men o
Ecology, Swedish Uni e si y o Ag icul u al Sciences, Uppsala 75007, Sweden.
4
Depa men o Mycology and Plan Resis ance, School o Biology, V.N. Ka azin Kha ki
Na ional Uni e si y, Kha ki 61022, Uk aine.
5
Labo a o io de Biodi e sidad y Funcionamien o Ecosis emico, Ins i u o de Recu sos Na u ales y Ag obiología de Se illa
(IRNAS), Consejo Supe io de In es igaciones Cien í icas, Se illa 41012, Spain.
6
Ins i u o Mul idisciplina pa a el Es udio del Medio ‘Ramón Ma gale ’and Depa amen o
de Ecología, Uni e sidad de Alican e, Alican e 03690, Spain.
7
Go henbu g Global Biodi e si y Cen e, Uni e si y o Go henbu g, Go henbu g 40530, Sweden.
8
Ins i u e o
Bo any, Uni e si y o he Punjab, Pakis an 54590, Pakis an.
9
Cen o de In es igaciones en Mic obiología y Bio ecnología-UR (CIMBIUR), Uni e sidad del Rosa io, Bogo á
111221, Colombia.
10
Ins i u e o Fo es y and Enginee ing, Es onian Uni e si y o Li e Sciences, Ta u 51006, Es onia.
11
G upo de BioMic o y Mic obiología Ambien al,
Escuela de Mic obiologia, Uni e sidad de An ioquia UdeA, Medellin 050010, Colombia.
12
Depa men o Ag icul u al, Food and Fo es Sciences, Uni e si y o Pale mo,
Pale mo 90128, I aly.
13
Depa men o Biomedical Sciences, Uni e si y o Caglia i, Caglia i 09124, I aly.
14
Depa men Biology, Ghen Uni e si y, Ghen 9000, Belgium.
15
Depa men o C op Science, Uni e si y o Dschang, Dschang, Came oon.
16
Depa men o Biological and En i onmen al Science, Uni e si y o Jy äskylä, Jy äskylä
40014, Finland.
17
Ins i u o Ju uá, Manaus 69083, B azil.
18
Depa men o Biological Sciences and Bio echnology, Bo swana In e na ional Uni e si y o Science and Tech-
nology, Palapye 10071, Bo swana.
19
Na u al His o y Museum o Zimbabwe, Bulawayo, Zimbabwe.
20
Cen o de In es igación e Inno ación pa a el Cambio Climá ico
(CiiCC), Uni e sidad San oTomás, Valdi ia, Chile.
21
Cen e o Mycology and Mic obiology, Uni e si y o Ta u, Ta u 50409, Es onia.
22
La ian S a e Fo es Resea ch Ins i u e
Sila a, Salaspils 2169, La ia.
23
Depa men o Bo any, Jawaha lal Neh u Rajkeeya Maha idyalaya, Pondiche y Uni e si y, Po Blai 744101, India.
24
College o Biological
Resou ce and Food Enginee ing, Qujing No mal Uni e si y, Qujing, Yunnan 655011, China.
25
Ins i u o Mul idisciplina io de Biología Vege al (CONICET), Uni e sidad Na-
cional de Có doba, Co doba 5000, A gen ina.
26
Na u al His o y Museum o Denma k, Copenhagen 1123, Denma k.
27
B i ish An a c ic Su ey, NERC, High C oss, Cam-
b idge CB3 0ET, UK.
28
Depa men o En i onmen al Science, Sain Ma y’s Uni e si y, Hali ax B3H 3C3, Canada.
29
Al ai S a e Uni e si y, Ba naul 656049, Russia.
30
Land and
Wa e , Commonweal h Scien i ic and Indus ial Resea ch O ganisa ion (CSIRO), Wembley 6014, Aus alia.
31
Depa men o Na u al Sciences, Manches e Me opoli an
Uni e si y, Manches e M1 5GD, UK.
32
Helmhol z Zen um München, Neuhe be g 85764, Ge many.
33
Biology Depa men , U ah Valley Uni e si y, O em, UT 84058, USA.
34
Plan , Soil and Mic obial Sciences, Michigan S a e Uni e si y, Eas Lansing, MI 48824-6254, USA.
35
Facul y o Na u al and En i onmen al Sciences, Ag icul u al Uni e si y
o Iceland, Reykja ík 112, Iceland.
36
Cen e o Mac oecology, E olu ion and Clima e, Uni e si y o Copenhagen, Copenhagen 1350, Denma k.
37
Vokė b anch, Ins i u e o
Ag icul u e, Li huanianResea ch Cen e o Ag icul u e andFo es y (LAMMC), Vilnius LT-02232, Li huania.
38
Depa men o Bo any, Facul y o Science, Uni e si yo Sou h
Bohemia, České Budějo ice 37005, Czech Republic.
39
Depa men o En i onmen al Sciences, Plan Ecology and Na u e Conse a ion, Wageningen Uni e si y and Re-
sea ch, Wageningen 6708, Ne he lands.
40
ELKH-EKKE Lendüle En i onmen al Mic obiome Resea ch G oup, Esz e házy Ká oly Ca holic Uni e si y, Ege 3300, Hunga y.
41
En i onmen al Science Cen e , Qa a Uni e si y, Doha, Qa a .
42
Ins i u o de Biología, Uni e sidad Nacional Au ónoma de México, Ciudad de México 04510, Mexico.
43
Na u al His o y Museum, Uni e si y o Ta u, Ta u 51003, Es onia.
44
Depa men o A c ic and Ma ine Biology, The A c ic Uni e si y o No way, T omsø 9019, No way.
45
Resea ch Uni T opical Mycology and Plan s-Soil Fungi In e ac ions, Uni e si y o Pa akou, Pa akou 00229, Benin.
46
Depa men o Sil icul u e and Ecology, Ins i u e o
Fo es y, Li huanian Resea ch Cen e o Ag icul u e and Fo es y (LAMMC), Gi ionys 53101, Li huania.
47
Cen e o Excellence in Fungal Resea ch, Mae Fah Luang Uni-
e si y, Chiang Rai 57100, Thailand.
48
Chai o Hyd obiology and Fishe y, Es onian Uni e si y o Li e Sciences, Ta u 51006, Es onia.
49
Depa men o Plan Biology, Facul y
o Li e Science, Uni e si y o Ilo in, Ilo in 240102, Nige ia.
50
Depa men o Fo es Sciences, Uni e si y o Helsinki, Helsinki 00014, Finland.
51
Go henbu g Cen e o Sus-
ainable De elopmen , Go henbu g 41133, Sweden.
52
Depa men o Biology, Sy acuse Uni e si y, Sy acuse 13244, USA.
53
Depa men o Plan Sciences, Quaid-i-Azam
Uni e si y, Islamabad 45320, Pakis an.
54
Depa men o Gene ics, Facul y o Na u al and Ag icul u al Sciences, Uni e si y o he F ee S a e, Bloem on ein 9300, Sou h
A ica.
55
Depa men o En i onmen , Facul y o Bioscience Enginee ing, Ghen Uni e si y, Ghen 9000, Belgium.
56
Mycology Wo king G oup, Goe he Uni e si y F ank u
am Main, F ank u am Main 60438, Ge many.
57
College o Biological Resou ce and Food Enginee ing, Qujing No mal Uni e si y, Qujing, China.
58
Ins i u e o Ag icul u al
and En i onmen al Sciences, Es onian Uni e si y o Li e Sciences, Ta u 51006, Es onia.
59
Cen e Fo Moun ain Fu u es, Kunming Ins i u e o Bo any, Chinese Academy o
Sciences, Kunming 650201, China.
60
F eie Uni e si ä Be lin, Ins i u ü Biologie, Be lin 14195, Ge many.
61
Ins i u e o Mic obiology, Czech Academy o Sciences, P ague,
Czech Republic.
62
Ins i u o Ciencias Ambien ales y E olu i as, Uni e sidad Aus al de Chile, Valdi ia, Chile.
63
Depa men o Zoology, College o Science, King Saud
Uni e si y, Riyadh 11451, Saudi A abia.
64
Depa men o Ecology and Plan Geog aphy, Moscow Lomonoso S a e Uni e si y, Moscow 119234, Russia.
65
Depa men
o Biology, College o Science, Uni ed A ab Emi a es Uni e si y (UAEU), Al Ain, UAE.
66
Depa men o Biological Sciences, Cali o nia S a e Poly echnic Uni e si y,
A ca a, CA 95521, USA.
67
Depa men o Food Science and Technology, Uni e si y o Bu undi, Bujumbu a Bu undi.
68
School o Biological Sciences and Ins i u e o Mi-
c obiology, Seoul Na ional Uni e si y, Seoul 08826, Ko ea.
69
Socie y o he P o ec ion o Unde g ound Ne wo ks (SPUN), Do e , DE 19901, USA.
70
Cen e o En i on-
men al Sciences, Hassel Uni e si y, Hassel 3500, Belgium.
71
Royal Bo anic Ga dens, Kew, Richmond TW9 3AE, UK.
*Co esponding au ho . Email: ladimi .mik yuko[email p o ec ed] (V.M.); [email p o ec ed] (L.T.)
†These au ho s con ibu ed equally o his wo k.
SCIENCE ADVANCES |RESEARCH ARTICLE
Mik yuko e al.,Sci. Ad . 9, eadj8016 (2023) 29 No embe 2023 2 o 15
Downloaded om h ps://www.science.o g a Jy askyla Uni e si y on No embe 30, 2023
among ungal ecological g oups, e lec ing hei niche a ibu es?
(iii) Wha mechanisms unde lie ungal species and phylogene ic di-
e si y ela ionships? (i ) Is clima e, h ough he e ec s on ungal
species anges (9,13), he p ima y d i e o ungal composi ional
u no e (i.e., be a di e si y)? ( ) Is he mal ene gy supply, he
main d i e o animal and plan di e si ica ion (27), also he
p ima y explana o y a iable o ungal gamma di e si y?
RESULTS AND DISCUSSION
Alpha di e si y
The la i udinal dis ibu ion o he o al numbe o ungal OTUs in
he samples (i.e., alpha di e si y, S
TOT
) exhibi s peaks in opical
and empe a e ecosys ems, while showing dep essions in dese ,
A c ic, and An a c ic ecosys ems (Fig. 1A and ig. S4). To iden i y
he ac o s explaining S
TOT
a iabili y, we used 135 publicly a ail-
able clima ic, ege a ion, and edaphic a iables ( able S2). We elim-
ina ed co ela ed a iables, p io i izing a iables al eady iden i ied
as ungal biodi e si y d i e s in global and con inen al-scale
s udies, and hen p eselec ed 35 in luen ial a iables o modeling
S
TOT
dis ibu ion wi hin each da ase using he ex eme g adien
boos ing (XGBoos ) machine lea ning echnique. Using a consen-
sus app oach ha accoun s o model pe o mance and ep esen a-
ion o en i onmen al condi ions in he aining da a, we
cons uc ed a high- esolu ion global map o p edic ed S
TOT
(S0
TOT
; Fig. 1B and igs. S5 o S9) wi h a 30–a c sec esolu ion equa -
ing o g id cells o <1 km
2
. The map highligh s he la ges ungal
di e si y “ho spo s”(a eas wi h S0
TOT
abo e he global 97.5 h pe -
cen ile) in he Eas A ican highlands, Gul o Guinea opical
o es s, Appalachian o es s, Cen al Ame ican d y o es s, and Hi-
malayan and New Guinean o es s ( ig. S10). The ho spo s pa ly
align wi h plan alpha di e si y peaks (14), p ima ily along he
equa o , excluding he we es pa s o equa o ial ain o es s wi h
S
TOT
dep essions. The S0
TOT
“coldspo s”(a eas wi h S0
TOT
below
he 2.5 h global pe cen ile) include he A acama, Saha a, and
A abian dese s, as well as he Cen al Asian d ylands, e lec ing
he p ima y limi ing e ec o wa e de ici on soil ungal alpha di-
e si y, simila o i s impac on soil es a e amoebae, ea hwo ms,
and o he soil-dwelling o ganisms (28–30).
While S0
TOT
displays a nea -mono onic inc ease along he g adi-
en s o mean annual p ecipi a ion (MAP) and empe a u e (MAT)
(Fig. 1C), co obo a ing p e ious wo ks [(5,7,12,30), bu see (13)],
condi ional e ec analysis shows ha S0
TOT
esponse o bo h p edic-
o s a ies subs an ially ac oss clima e ypes. Speci ically, i is posi-
i ely ela ed o annual empe a u e in cold humid o empe a e
humid egions, and nega i ely so in ho o a id egions (Fig. 1D).
The S0
TOT
esponse o annual p ecipi a ion is posi i e in ho cli-
ma es wi h up o ca. 2000 mm o annual ain all, bu nega i e in
hype -humid o cold a eas. Co espondingly, while opical and
sub opical ain o es s gene ally ha e high S0
TOT
(Fig. 1E, ig. S11,
and able S3), ela i ely lowe alues a e p edic ed o he cen al
we es pa s o he Amazon and Congo basins and o he Indo-
Malay swamp o es s, whe e hyd ic soils p edomina e. Ac oss em-
pe a e b oadlea and mixed o es s, S0
TOT
peaks in he Balkans, Fa
Eas , and Appalachians, bu has he lowes alues in Wes e n and
Cen al Eu opean humid lowlands. In und a and bo eal o es
biomes, S0
TOT
peaks in he Fa Eas and declines in suba c ic and
A c ic ecosys ems.
Acco ding o he model p edic ions, land co e ype also cons i-
u es a majo in luence o ungal alpha di e si y, wi h deciduous
o es s suppo ing he highes S0
TOT
in all biomes ( able S3). A a
b oade scale, his e ec is exempli ied by S0
TOT
g adien s om
s eppes o ansi ional o es -g assland ege a ion o o es ed
biomes. The di e ence in S0
TOT
be ween deciduous o es s and
ei he c oplands o u ban a eas is pa icula ly p onounced in he
opical d y o es biome. Consequen ly, ag icul u al expansion
and u baniza ion, which co e o e 30% o he biome a ea, lead
o no able and ex ensi e declines in S0
TOT
, as exempli ied in he
Deccan pla eau, Cen al Indochina, and Cen al Ame ica. This e-
lec s conce ns abou he loss o opical biodi e si y in o he bio ic
g oups (15). In addi ion, while less p onounced, ye ex ensi e S0
TOT
declines due o ag icul u al land ans o ma ion a e also p edic ed
in he Ame icas and Eu ope ac oss Medi e anean and empe a e
o es biomes, as well as empe a e g asslands.
The p esen analysis o he d i e s o ungal di e si y dis ibu-
ion is based on he mos ex ensi e global sample collec ed using
a uni ied p o ocol and co e ing a la ge olume in he mul idimen-
sional space o en i onmen al p ope ies. Howe e , ce ain habi a s,
cha ac e ized by unique combina ions o en i onmen al cha ac e -
is ics, ha e lowe p edic ion p ecision and dese e u u e esea ch
ocus ( igs. S8 and S9). The la ge numbe o species o ming he
basis o ou analyses suppo s he expec a ion o global ungal di e -
si y (31), while i s exceeding he numbe o cu en ly named species
unde sco es he necessi y o addi ional e o s in desc ibing myco-
bio a om a ious habi a s (32).
Alpha di e si y o ungal ecological g oups
We examine and align he wo ldwide dis ibu ions o key ungal
g oups d i ing impo an soil p ocesses on Ea h h ough pa ho-
genesis, symbiosis, and decomposi ion. We ind ha edaphic pa-
ame e s, ecosys em p oduc i i y, and wa e a ailabili y a e
gene ally mo e impo an o sap o ophic g oups, while empe a-
u e and ege a ion composi ion a e undamen al o myco hizal
ungi (Fig. 2A and able S4). Concep ually mi o ing p e ious ind-
ings (13,33,34), he la i udinal dis ibu ions o he p edic ed ich-
ness o AM and EcM ungi (S0
AM
and S0
ECM
, espec i ely) a e almos
opposi e ( =−0.54; ig. S12, A and B, and Supplemen a y Tex )
p ima ily di ided by empe a u e niche (i.e., wi h op imal MAT
>20°C o S0
AM
and −15°C o +5°C o S0
ECM
; see ig. S13) and
abundance o po en ial hos plan s. By o e laying he maps o
S0
AM
and S0
ECM
(Fig. 2B), we ou line wo dis inc a eas wi h high
S0
AM
(gene ally in opical o es s and sub opical g asslands) and
S0
ECM
(in empe a e and bo eal o es s) wi h ansi ional ege a ion
such as o es -g asslands, o es -s eppes, and sa annas, inhabi ed by
species- ich communi ies o bo h g oups.
While he di e si y ho spo s o molds and p edominan ly sap o-
ophic NMA a e in e ed, espec i ely, o empe a e (e.g., in he
Japanese A chipelago, Tasmania, and he Appalachians) and op-
ical (e.g., he Gul o Guinea, Oceanian, and Cen al Ame ica
o es s) ain o es s, S0
MOLD
and S0
NMA
a e s ongly co ela ed ( =
0.70) and espond e y simila ly o p ima y d i e s ( igs. S12 and
S13). Molds, howe e , exhibi a s onge demand o soil ca bon
and ni ogen con en and p e e ence o mo e acid soils han
NMA. Unlike o he sap o ophic g oups, yeas di e si y is less e-
sponsi e o MAP and nu ien a ailabili y. We p edic ela i ely
high S0
YEAST
o a id alkaline soils, in ag eemen wi h he ex em-
ophilic habi o many yeas species and suppo ing he idea o an
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unde s udied yeas bio a om ex eme habi a s (25,35). Con e sely,
nonyeas unicellula ungi exhibi ed na owe op imum anges o
clima ic and edaphic pa ame e s, and had p edic ed ho spo s in
empe a e mesic o es s bu coldspo s in a id egions and opical
hyd ic soils.
Compa ed o he o he ungal g oups, he e ec o en i onmen-
al a iables (including land co e ypes) on S
PATH
is weake , esul -
ing in a lowe S0
PATH
a iabili y ac oss he globe ( ig. S12 and able
S5). P ima ily explained by diu nal empe a u e ampli ude (DTA),
S0
PATH
is highe han a e age a DTA o ca. 7° o 13°C (i.e., op imal
o mos plan s and ec o he ms and ungal pa hogens’p ima y
hos s) while sha ply declining ou side his ange (i.e., in ex eme
hype -a id, alpine, and A c ic habi a s). These indings, summa iz-
ing he a ie y o ela ionships be ween pa hogen di e si y and hos
communi y p ope ies (36–39), sugges ha a bio ic communi y
Fig. 1. Global dis ibu ion o soil ungal alpha di e si y. (A) La i udinal dis ibu ions o o al ungal alpha di e si y (S
TOT
). (B) Consensus map o p edic ed o al ungal
alpha di e si y (S0
TOT
), wi h inse s showing S0
TOT
la i udinal dis ibu ions in No he n (g een) and Sou he n (blue) Hemisphe es; a eas co e ed wi h pe ennial ice a e
colo ed in g ay. (C) S0
TOT
(median and in e qua ile ange) dis ibu ion in he global g adien s o MAP and MAT; g een and g ay deno e he p edic ions wi hin and
beyond aining da a co e age, espec i ely. (D) Condi ional e ec s o MAP and MAT on S0
TOT
. (E) S0
TOT
ac oss e es ial biomes, wi h do s showing median S0
TOT
o
818 e es ial eco egions (47). Biomes sha ing he same le e indica e no signi ican di e ence a he 5% le el o signi icance a e P alue adjus men wi h he
Holm-Bon e oni me hod.
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exceeding a ce ain low h eshold o di e si y o biomass in a iably
shel e s a ela i ely di e se communi y o ungal pa hogens.
Ou alpha di e si y maps p o ide he basis o conse ing ungal
ecological g oups and, hus, he ecosys em unc ions pe o med by
hem and he o ganisms elian on hem. Fu he mo e, he p edic -
ed unc ional p o iles o soil mycobio a, buil by summa izing he
g oups’di e si y, subs an ially a y ac oss he globe (Fig. 2C). The
unique unc ional p o iles o e es ial biomes and eco egions ( igs.
S14 and S15) p o ide dis inc ecosys em se ices and unc ions in
global nu ien cycling, i idly exempli ied by he con ibu ion o
myco hizal ungi in global ca bon alloca ion (4).
A eas wi h high S0
OHP
(Fig. 2D) co e he egions wi h he g ea -
es human popula ion densi y (Fig 2E). These esul s, while sup-
po ing he link be ween he equency o OHP in ec ions and
social ac o s epo ed in epidemiological s udies (40), may also
poin o an unde explo ed pa hogenici y po en ial o ungal bio a
in less popula ed a eas. In ligh o he ele a ed OHP po en ial in
molds, we expec a la ge numbe o ye undisco e ed OHP
species wi hin S0
MOLD
ho spo s, whe e espec i e epidemiological
su eillance is needed o mi iga e public heal h isks.
Phylogene ic alpha di e si y
Inco po a ing species ela edness in o biodi e si y es ima es
h ough phylogene ic di e si y indices iden i ies he mechanisms
o communi y assembly (41), and hei po en ial unc ional alue,
and co espondingly unde lies conse a ion p io i iza ion (15,42).
In ungal biogeog aphy, phylogene ic di e si y is a ely conside ed
alongside axonomic di e si y [bu see (33)]. In ou da ase , he
simples measu e o phylogene ic alpha di e si y— o al phyloge-
ne ic b anch leng hs in a communi y (S
PD
)—is igh ly posi i ely
linked wi h species ichness ( = 0.71 be ween S
PD
and S
TOT-
GSMc
). Acco dingly, S0
PD
b oadly co ela es wi h S0
TOT
( ig. S17).
Meanwhile, phylogene ic dispe sion (SES
PD
), ep esen ing s an-
da dized di e ence be ween ac ual S
PD
and S
PD
o a andomly as-
sembled communi y o he same ichness, illus a es ha
app oxima ely 93% o he sampled communi ies a e phylogene i-
cally clus e ed (SES
PD
< 0). This highligh s ha en i onmen al il-
e ing o closely ela ed, and hus unc ionally simila , species is he
main global mechanism shaping local mycobio a. Phylogene ically
o e dispe sed communi ies (SES
PD
> 0), assembled wi h a g ea e
con ibu ion o compe i i e displacemen among unc ionally
simila species, a e ound in he equa o ial opics. Dec easing pole-
wa d (Fig. 3A) and wi h al i ude in he opical highlands (Fig. 3B),
SES
PD
is bes explained by MAT ( able S4). While SES
PD
is indepen-
den o S
TOT
( = 0.01), i is a ec ed by he ichness o ce ain ungal
g oups in a communi y (posi i ely by S
AM
and S
UCELL
and nega i e-
ly by S
ECM
and S
MOLD
; able S6).
Fig. 2. Alpha di e si yo soil ungal ecological g oups. (A) Impo ance o key p edic o s and he shape o hei ela ionships wi h he local ichness o ungal ecological
g oups. (B) Model-p edic ed global dis ibu ion o EcM and AM ungal local ichness. Colo legend deno es low, medium, and high EcM o AM ungal p edic ed ichness;
a eas co e ed wi h pe ennial ice a e colo ed in g ay. (C) Func ional p o iles o soil mycobio a in di e en biomes based on p edic ed alpha di e si y o ungal ecological
g oups. (D) Global dis ibu ion o p edic ed alpha di e si y o OHP ungi and (E) o e lap be ween a eas wi h high p edic ed ichness o OHP ungi and high human
popula ion densi y (D).
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The map o model-p edic ed SES
PD
(SES0
PD
) shows s ong la i-
udinal and ele a ional g adien s o phylogene ic dispe sion, mi -
o ing plan and animal biogeog aphical pa e ns (15,43). On he
basis o he known pa e ns in EcM and AM ungal e olu iona y
biogeog aphy (33,34,44,45), he links be ween SES
PD
and he ich-
ness o ecological g oups highligh he impac o apid specia ion in
speci ic gene a o EcM ungi and he selec i e exclusion o AM ungi
in colde clima es on he global dis ibu ion o he phylogene ic
s uc u e o he whole soil ungal communi y. Resea ch o he
mechanisms in ol ing unicellula ungi and molds, axa wi h a
la gely unexplo ed e olu iona y biogeog aphy, ep esen s a p om-
ising a enue o u u e in es iga ions.
Be a di e si y
Be a di e si y, measu ing he composi ional he e ogenei y o bio a
among loca ions, is a c ucial pa ame e o he de elopmen o s a -
egies o egional- and con inen al-scale biodi e si y main enance
(17). We examined global pa e ns o soil ungal be a di e si y
among GSMc samples based on Simpson’s pai wise dissimila i y
in he composi ion o OTUs [D
TAX
( axonomic dissimila i y)]
and phylogene ic lineages [D
PD
(phylogene ic dissimila i y)]. As
he indices a e s ongly co ela ed (Man el’s = 0.86), hei e ealed
d i e s and dis ibu ion maps a e la gely simila ( ig. S19). Speci i-
cally, while ou sampling cap u es high composi ional dissimila i y
a he landscape scale, i also e eals he g ea e impo ance o he
en i onmen a he han geog aphical dis ance o ungal composi-
ional a iabili y a mac oscales, suppo ing he esul s o he
species dis ibu ion analyses (Fig. 4A) (5,9,13,46). The s onges
en i onmen al p edic o s o ungal spa ial composi ional u no e
include MAT, soil pH, and N con en (Fig. 4A and ig. S19).
We cons uc ed maps o he model-p edic ed D
TAX
and D
PD
(D0
TAX
and D0
PD
, espec i ely) (Fig. 4B and ig. S20), which
b oadly illus a e a s a is ically signi ican di e gence o all e es i-
al biomes in mycobio a composi ion (Fig. 4C, ig. S18, and able S7),
wi h opical mois o es s ha bo ing he mos unique communi ies.
Mang o es, opical coni e ous o es s, and looded and moun ain
g asslands [i.e., small biomes associa ed wi h speci ic e ains “em-
bedded”in la ge con inuous biomes (47)] sha e he la ges pa o
Fig. 3. Global pa e ns in soil ungal phylogene ic s uc u e. (A) Global dis ibu ion o p edic ed phylogene ic dispe sion (SES0
PD
) o soil ungi, wi h accompanying
la i udinal dis ibu ions o SES0
PD
in he No he n (g een) and Sou he n (blue) Hemisphe es. (B) Associa ion be ween SES
PD
and ele a ion in opical biomes. Fo cla i y,
he yaxis is clipped, and wo da a poin s displaying low SES
PD
alues om he Neo opical biome a e excluded.
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hei ungal bio a wi h su ounding biomes. Howe e , gi en he
speci ici y o such ecosys ems, we conside ha he uniqueness o
hei mycobio a could be unde es ima ed in ou s udy (and o he
ongoing wo ks) due o he lack o a iables accu a ely desc ibing
such habi a s (as emphasized in moun ain biodi e si y s udies) (48).
By a e aging D0
TAX
wi hin e es ial eco egions (Fig. 4D) and a
150-km adius a ea, we p edic ha he mos p onounced spa ial
u no e o soil ungi wi hin sub opical and opical egions
a ises om g ea e di e ences in al i ude and, consequen ly, en i-
onmen al he e ogenei y ( igs. S21 and S22). When accoun ing o
he opog aphy e ec , we ound ha he D0
TAX
- o-D0
PD
a io de-
clines wi h dis ance om he equa o (β < −0.01, P< 0.001;
Fig. 4E). In e ms o opical conse a ism heo y, his implies a
nonlinea inc ease o opical lineage loss wi h dis ance om he
en i onmen al op imum.
Gamma di e si y
Fungi a e es ima ed o comp ise 2 million o 3 million species (31),
wi h he global dis ibu ion o egional species ichness emaining
one o he leas s udied aspec s o ungal di e si y. On he basis o
ou global su eys and museum collec ions’sequences deposi ed in
he UNITE (including GenBank) da abase (49), we e alua e gamma
di e si y as OTU ichness in a supe eco egion [as de ined by (46)],
accoun ing o a ia ions in sampling and sequencing e o ( able
S8). The o e all ungal gamma di e si y (G
TOT
) is linked o egional
a e age S0
TOT
(Fig. 5A) and, co espondingly, esponds o he index
Fig. 4. Global pa e ns o soil ungal be a di e si y. (A) Composi ional u no e o ungal communi ies o e he anges o he mos in luen ial a iables: geog aphical
dis ance (0.7 o 19,948 km), MAT (−11.8° o 29.3°C), soil pH (4.0 o 8.7), soil N con en (0.4 o 16.7 g kg
−1
), abo eg ound biomass (0 o 1828 Mg C ha
−1
), o ganic C s ocks (9.4
o 142.5 Mg ha
−1
), and win e leng h (0 o 365 days). (B) P edic ed composi ional dissimila i y o ungal communi ies, whe e colo g adien s ep esen species compo-
si ion g adien s de i ed om en i onmen al p edic o s and combined in o h ee GDM componen s; he eby, colo simila i y among loca ions in he map is p opo ional
o composi ional simila i y o ungal communi ies. (C) G aph o composi ional pai wise ela edness o e es ial biomes, wi h node colo indica i e o he biome’s po-
si ion in GDM componen s and edge wid h p opo ional o mycobio a composi ional simila i y be ween he biomes. (D) P edic ed composi ional dissimila i y among soil
ungal communi ies wi hin eco egions (median o D0
TAX
). (E) La i udinal dis ibu ion o eco egional D0
TAX
- o-D0
PD
a io in a eas wi h a small, medium, and la ge al i ude
di e ence (ele a ion in e qua ile ange).
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o d ough (Fig. 5B). The s ong posi i e e ec o spa ial a iabili y
o he mal ene gy supply (i.e., o po en ial e apo anspi a ion) on
G
TOT
ma ches he classical posi i e ela ionships be ween en i on-
men al he e ogenei y and egional plan and animal biodi e si y
(50). Gi en ha empe a u e is he main d i e o composi ional
u no e and SES
PD
, he e ealed e ec o ene gy supply a iabili y
on G
TOT
also links egional ichness wi h be a di e si y and phylo-
gene ic dispe sion. The signi icance o he annual ange o soil em-
pe a u e o G
TOT
sugges s ha he ela ionships be ween
physiological ole ance o empe a u e luc ua ions and dis ibu ion
anges o plan and ec o he mic species (51) also apply o ungi.
Meanwhile, MacA hu and Wilson’s (52) classic heo y o island
biogeog aphy does no hold ue o ou da ase , as G
TOT
is no
ela ed o island a ea o dis ance om mainland ( able S9).
In he G
TOT
map, much like he si ua ion in o he kingdoms (48,
53,54), mos o he G
TOT
peaks in he opics a e loca ed no only in
moun ainous egions such as he eas e n A ican highlands, New
Guinea, and eas e n Himalayas bu also in coas al lowlands in
Cen al A ica and Aus alia (Fig. 5C). No ably, he eas e n slopes
o he opical Andes, a egion known o i s high plan and e e-
b a e ichness (48,53,54), a e no among he iden i ied a eas wi h
high ungal G
TOT
. In he empe a e zones, G
TOT
peaks in no heas -
e n Uni ed S a es, Ana olian o es s, he Japanese A chipelago, and
Ussu i o es s. Be ingia exhibi s he highes gamma di e si y among
A c ic supe eco egions, possibly because o i s ela i ely long
glacie - ee his o y (55). The lowes G
TOT
alues a e obse ed in
dese a eas.
While egional ichness o ungal ecological g oups gene ally co-
incides wi h alpha di e si y, peaking in he same geog aphical
egions and la gely d i en by simila a iables ( igs. S24 and S25,
ables S10 and S11, and Supplemen a y Tex ), gamma di e si y o
pa hogens is also egula ed by he egional a iabili y o clima ic
p edic o s and ele a ion, which highligh s he impo ance o en i-
onmen al he e ogenei y o his wide- anging ecological g oup.
Syn hesis
Toge he , he esul s o ou mul idimensional analyses indica e ha ,
alongside o he kingdoms, soil ungi exhibi peaks in alpha, be a,
and gamma di e si ies in opical (mos ly moun ainous) egions
while dec easing owa d he poles and ab up ly d opping in
hype -a id egions. While wa e de ici in d ylands is an e iden
limi ing ac o o alpha di e si y, ou models inco po a ing
wa e -ene gy balance also explain he declines in local di e si y in
opical hype -humid soils—one o se e al un esol ed issues in
ungal biogeog aphy. In addi ion, phylogene ic and species u n-
o e a e linked o gamma di e si y by spa ial a iabili y in
Fig. 5. Gamma di e si y o soil ungal communi ies. (A) Rela ionships o o al ungal egional ichness (G
TOT
) wi h median S
TOT
and dis ance om he equa o . Sepa a e
eg ession lines a e shown o he No he n (g een) and Sou he n (blue) Hemisphe es. Dese s and d ylands ( ed do s) a e excluded om he eg ession analysis. (B)
Ma ginal e ec s o he signi ican p edic o s on G
TOT
. Po en ial e apo anspi a ion (PET) spa ial a iabili y is he in e qua ile ange o Tho n hwai e’s PET, and he Ellen-
be g quo ien , he a io o mean empe a u e o he wa mes mon h o annual p ecipi a ion, is an index o d ough . (C) Map o G
TOT
a he eco egion scale (46).
SCIENCE ADVANCES |RESEARCH ARTICLE
Mik yuko e al.,Sci. Ad . 9, eadj8016 (2023) 29 No embe 2023 8 o 15
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AUA g an om esea ch council o UAE Uni e si y: G00003654 ( o S.M.), Ghen Uni e si y:
Bijzonde Onde zoeks onds ( o A.V.), Ghen Uni e si y: Bijzonde Onde zoeks onds (BOF-PDO-
2017-001201) ( o E.D.C.), Ghen Uni e si y: The Facul y Commi ee Scien i ic Resea ch, FCWO
( o E.D.C. and A.V.), The King Leopold III Fund o Na u e Explo a ion and Conse a ion ( o A.V.
and E.D.C.), The Resea ch Founda ion—Flande s (FWO) ( o E.D.C. and A.V.), The High-Le el
Talen Rec ui men Plan o Yunnan P o inces: “Young Talen s”P og am ( o D.-Q.D.), The High-
Le el Talen Rec ui men Plan o Yunnan P o inces: “High-End Fo eign Expe s" P og am ( o N.
N.W.), IRIS schola ship o p og essi e and ambi ious women ( o L.H.), Es onian Uni e si yo Li e
Sciences: P190250PKKH ( o K .Pa.), Hunga ian Academy o Sciences: Lendüle P og amme
(96049) ( o J.G.), Eö ös Lo ánd Resea ch Ne wo k ( o J.G.), Bo swana In e na ional Uni e si y o
Science and Technology ( o C.N.), and Highe Educa ion Commision (HEC, Islamabad, Pakis an):
Indigenous and In e na ional esea ch suppo ini ia i e p og am (IRSIP) schola ship ( o M.S.).
Au ho con ibu ions: Concep ualiza ion: L.T., V.M., O.D., and U.K. Me hodology: V.M., O.D., A.
Z., Mo.Ba., O.P., L.T., and R.P. In es iga ion: V.M., O.D., and L.T. Fo mal analysis: V.M., O.D., L.T., A.Z.,
Mo.Ba., O.P., and N.H.-D. Resou ces: L.T. and U.K. Funding acquisi ion: L.T., R.D., U.K., Ka.Po., M.D.-
B., F.T.M., S .An., K.-A.B., M.T., M.O., M.Z., Ü.M., M.E., M.M., Mo.Ba., Al.An., Ke.A ., Je.An., N.S., K.R., D.
K., I.H., K.T.I., M.P., N.Y., C.M., R.G., T.R., T.H., G.B., R.G.-O., J.D., K.K.N., E.M.B., Ju.Al., V.F., V.O., E.N., Pe.
Mo., S.M., A.V., E.D.C., D.-Q.D., N.N.W., L.H., K .Pa., J.G., C.N., and M.S. P ojec adminis a ion: L.T.
Supe ision: L.T. Visualiza ion: O.D. and V.M., N.H.-D., L.T., and A.Z. W i ing—o iginal d a : O.D.,
V.M., and L.T. W i ing— e iew and edi ing: All au ho s. Compe ing in e es s: The au ho s
decla e ha hey ha e no compe ing in e es s. Da a and ma e ials a ailabili y: All da a
needed o e alua e he conclusions in he pape a e p esen in he pape and/o he
Supplemen a y Ma e ials. Addi ional da a used in his a icle and he gene a ed g idded maps
a e a ailable h ough Zenodo DOI:10.5281/zenodo.8013448. The code used o da a analysis
and isualiza ion o he main esul s o he s udy is a ailable a Gi Hub: h ps://gi hub.com/
Mycology-Mic obiology-Cen e /Global_ ungal_di e si y (DOI: 10.5281/zenodo.8407259).
Addi ional de ails on he GSMc a e a ailable a h ps://GSMc- ungi.gi hub.io/ (DOI:10.5281/
zenodo.10003080). Fungal DNA sequences and OTU occu ences can be accessed om he
Plu oF biodi e si y da a managemen pla o m. The aw sequence da a gene a ed in his s udy
a e deposi ed in he Eu opean Nucleo ide A chi e (ENA) a EMBL-EBI unde accession numbe
PRJEB63550 (S udy ID ERP148695, un accessions ERR11608487–ERR11609485 and
ERR11609494–ERR11613474). All soil samples collec ed a e p ese ed in he Uni e si y o Ta u
Na u al His o y Museum’s collec ion o DNA and en i onmen al samples.
Submi ed 17 July 2023
Accep ed 30 Oc obe 2023
Published 29 No embe 2023
10.1126/sciad .adj8016
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Connec ing he mul iple dimensions o global soil ungal di e si y
Vladimi Mik yuko , Olesya Dulya, Alexande Zizka, Mohammad Bah am, Nilou a Hagh-Dous , S en Anslan, Oleh
P ylu skyi, Manuel Delgado-Baque izo, Fe nando T. Maes e, Hen ik Nilsson, Jaan Pä n, Maa ja Öpik, Ma i Moo a,
Ma in Zobel, Mikk Espenbe g, Ülo Mande , Abdul Nasi Khalid, Ad iana Co ales, Ah o Agan, Aída-M. Vasco-Palacios,
Alessand o Sai a, And ea Rinaldi, Annemieke Ve beken, Bobby Sulis yo, Bo is Tamgnoue, B endan Fu neaux, Camila
Dua e Ri e , Caspe Nyamukondiwa, Ca hy Sha p, Césa Ma ín, Daniyal Goha , Da a Kla ina, Dipon Sha mah, Dong-
Qin Dai, Edua do Nouh a, Elisabe h Mach eld Bie sma, Elisabe h Rähn, E in Came on, Eske De C op, E eli O sing,
E geny Da ydo , Felipe Albo noz, F ancis B ea ley, F anz Buegge , Geo ey Zahn, G ego y Boni o, Inga Hiiesalu, Isabel
Ba io, Jacob Heilmann-Clausen, Jelena Ankuda, Ji i Doležal, John Kupagme, Jose Maciá-Vicen e, Joseph Djeugap Fo o,
Józse Geml, Juha Ala alo, Julie a Al a ez-Manja ez, Kad i Põldmaa, Kad i Runnel, Kale Adamson, Ka i-Anne B å hen,
Ka in P i sch, Kassim Tchan Issi ou, K#s u is A molai is, Ke in Hyde, Ke in K. Newsham, K is el Panksep, Adebola Azeez
La ee , Linda Hansson, Louis Lami , Malka Saba, Ma ia Tuomi, Ma ieka G yzenhou , Ma ijn Bau e s, Meike Piepenb ing,
Nalin N. Wijayawa dene, Nou ou Yo ou, Ola i Ku ina, Pe e Mo ime , Pe e Meidl, Pe Kohou , Rasmus Puusepp, Rein
D enkhan, Robe o Ga ibay-O ijel, Robe o Godoy, Saad Alkah ani, Saleh Rahimlou, Se gey Dudo , Se gei Põlme,
Soumya Ghosh, Sunil Mund a, Talaa Ahmed, Ta quin Ne he way, Te y Henkel, Tomas Roslin, Vincen N ezi yayo,
Vladimi Fedoso , Vladimi Onipchenko, Wee agalle A achchillage E andi Yasan hika, Young Lim, Michael Van Nuland,
Nadejda Soudzilo skaia, Alexand e An onelli, U mas Kõljalg, Kessy Aba enko , and Leho Tede soo
Sci. Ad . 9 (48), eadj8016. DOI: 10.1126/sciad .adj8016
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