Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. 1 Morgado in forests climate tropical Geml J´ozsef of in fungi communities role montane fungal the Tropical soil-borne reveals title: of Running gradients composition elevation taxonomic and Paleotropical functional and the Neotropical shaping of time. will over fungi sampling functionality forest and Comparative montane composition in tropical variation that in implies resulting elevation in factors change, along resulting environmental climate across turnover forests, to correlated or functional profiles sensitive tropical and and hosts functional be moist compositional temperature their likely of across pronounced by as similarity mechanisms The mainly well remarkable similar driven guilds. as low, by gradients functional driven fungi, is of be of regions proportions may among pools predictable proportions relatively shared species niche species local which functional of guild, in suggests biogeographic functional number differences regions Strong of the resulting taxa. irrespective Although and related often limitation closely orders, substrates. relatively dispersal taxonomic contrasting for reflects of found optimum by we level likely environmental addition, driven the structure and In at is constraints observed guilds. turnover physiological functional be Elevational shared factors within already suggests pH. environmental species can (northern soil of reflects preference Paleotropics and replacement habitat composition and and temperature that community groups Neo- particularly fungal functional the scales, among soil taxonomically in pantropical preferences that of environmental understanding gradients and found structure our elevational We regional the enhance along both to compared Borneo). soils at We setting and in ideal America, communities assemblies. an Central fungal community offer Argentina, soil and mountains diverse distributions factors, functionally biotic species and and underlie abiotic that in mechanisms gradients of steep their of Because Abstract 2020 5, May 6 5 4 3 2 1 Morgado Geml Jozsef in and communities forests functional fungal tropical the soil-borne shaping of in composition climate taxonomic of role the elevation reveals Paleotropical gradients and Neotropical of sampling Comparative auai idvriyCne,Driwg2 ..Bx91,20 ALie,TeNetherlands The Leiden, RA 2300 9517, Box P.O. 2, Darwinweg Center, Biodiversity Naturalis ueUniversity Duke CREAF-CSIC-UAB C´ordoba de Nacional Universidad Center Biodiversity Naturalis Arizona of University Eszterh´azy K´aroly Egyetem 1,5 3 ro Grau Oriol , ruOriol Grau , 1,2 1 ,A lzbt Arnold Elizabeth A. *, lzbt Arnold Elizabeth , 6,7 5 lcaIb´a˜nez Alicia , lcaIbanez Alicia , 2 ain Semenova-Nelsen Tatiana , 3 3 ain .Semenova-Nelsen A. Tatiana , Bal´azs Hegyi , 2 Bal´azs Hegyi , 1 8,9 Fran¸cois Lutzoni , 1 n aci Lutzoni Fancois and , 3 dad Nouhra Eduardo , 1 dad .Nouhra R. Eduardo , 10 6 4 Luis , 4 usN. Luis , Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. canadGyns21) onan rvd nqeopruiist etvroseooia hypotheses, ecological various test to 2001; opportunities (Lomolino studies unique rich- biogeographic provide species and Mountains in ecological 2010). changes to of Grytnes central documentation and been biota, have McCain mountain composition on community Humboldt mountains and von Antarctica, ness and outside Wallace (Spehn area (Lo- Darwin, species land endemism of terrestrial world’s high studies all the of of of areas one-third one-eighth as about well about harbor as representing hotspots Despite biodiversity 2001). as molino recognized are generally ecosystems Montane fungi time. soil over Panama, functionality metabarcoding, INTRODUCTION and DNA composition will ITS, in fungi Borneo, variation forest Argentina, in Words: montane resulting driven Key tropical change, gradients that climate elevation implies to along factors sensitive of environmental turnover be proportions correlated is functional likely predictable and regions and relatively temperature among be in compositional by shared may resulting pronounced mainly species proportions forests, The of niche tropical number functional guilds. moist species the suggests across functional Although regions mechanisms local across substrates. similar in profiles or by differences driven functional hosts guild, of resulting their similarity functional as and remarkable well of limitation low, as irrespective habitat dispersal fungi, often that reflects of orders, taxa. found likely pools related taxonomic we closely structure relatively of addition, for biogeographic level optimum among environmental In Strong the and preferences guilds. constraints at physiological environmental functional shared observed suggests contrasting within which be by species already driven of can particularly is replacement preference scales, turnover fungal and pantropical Elevational soil and groups that pH. regional functional found We soil both Borneo). at and and factors temperature America, environmental Central reflects Argentina, gradients elevational (northern composition our along Paleotropics community soils enhance and in to communities Neo- setting fungal the ideal soil the in diverse an compared functionally We offer assemblies. and community mountains taxonomically and factors, of distributions structure biotic species underlie and that abiotic mechanisms in of understanding gradients steep their of Because [email protected] Abstract [email protected], E-mail: 523-456; 36 (+36) Phone: Hungary Eger, H-3300 6, Eszterh´azy K´aroly Le´anyka Group, University, u. Research Microbiome Lend¨ulet Environmental MTA-EKE address: author, *Corresponding Hungary. Debrecen, 10 H-4002 t´er 1, Egyetem Debrecen, of University 9 8 Guiana French Kourou, 97310 7 6 C´ordoba,5 Argentina 5000 495, CC 1611, V´elezC´ordoba, Sarsfield Av. 4 U.S.A. AZ, Tucson, 3 Hungary. Eger, H-3300 6, u. 2 otrlSho fErhSineadDprmn o adcp rtcinadEvrnetlGeography, Environmental and Protection Landscape for Department and Science Earth of School Hungary. Doctoral Eger, H-3300 Eszterh´azy 6, Le´anyka K´aroly Centre, University, u. Agronomique, Knowledge Campus Innoregion Guyane), Univ. Antilles, Univ. Inra, CNRS, (AgroParisTech, EcoFoG Spain Vall`es, UMR Catalonia, del Cirad, Norway Cerdanyola Oslo, 08193, Oslo, Unit, of Ecology University Sciences, Global Natural CREAF, and Mathematics of Faculty Biosciences, of de Department Arizona, Nacional Universidad of FCEFyN, University CONICET, Biology, (IMBIV), Evolutionary Biolog´ıa Vegetal de and Multidisciplinario Ecology Instituto of Department and Sciences Plant of School Eszterh´azy Le´anyka K´aroly University, Group, Research Microbiome Lend¨ulet Environmental MTA-EKE eateto ilg,Dk nvriy uhm C 70 U.S.A. 27708 NC, Durham, University, Duke Biology, of Department 2 tal et 02 noel 05.Sneteeryscientific early the Since 2015). Antonelli 2012; . Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. ueadpeiiain eetn ffcso oleahcvralsadpatcmuiycmoiin Fungal composition. community plant and variables tempera- edaphic in soil changes on through effects communities reflecting fungal precipitation, structure fungal and would core ture gradients on Pantropical elevation factors of that forests. types; characterization edaphic hypothesized montane Neotropical first We forest and and among the elevational lowland climatic provide fungi of of major 4) of soils and influence in in groups mountains; possible mycobiomes communities functional tropical the fungal and in evaluate taxonomic soil composition 3) community in characterize Argentina. mountains; richness 1) northwestern Paleotropical of to in and patterns Yungas Sabah, were elevation in Andean work Range compare the Crocker this and 2) and Panama; of Kinabalu western aims Mt. of forest areas: The in unexplored. rainforests mountain fungi mostly the tropical of remain groups Borneo; three functional mountains in Malaysian diverse tropical gradients of richness elevation in and along groups composition soils fungal community compared of we range study, this wide In a of diversity beta and alpha (Merckx fungi AM (G´omez-Hern´andezand (Geml Mexico communities eastern fungal air in total and macrofungi of of food studies water, morphological clean include al. of These provision (2017). the Geml as by such diversity our services the in ecosystem of drivers to gap (Bakker contribute major This (Bagchi plants, as forests unknown. with tropical reported virtually in interactions been communities remain plant have of mountains fungi composition because tropical and concerning in particularly communities seems fungal knowledge of composition and Javis (Cordier (Bahram increasingly fungi fungi phyllosphere is root-associated groups: mountains importance functional in al. their fungal conducted specific and et been have of ecosystems, studies distribution Several of the health. al. functioning on human regions and the safety temperate in food in roles to respect key with plants with vascular recognized organisms on the focused living have to of gradients contributed elevation along have Grau richness Parris 2006; and species (e.g., of research studies animals evolutionary most and mechanisms. scale, and underlying global their ecological a and On for biodiversity of point increasing patterns with focal spatial structure factors community of a abiotic in understanding changes for been Resulting requirements have species. physiological other elevation their tropical with occupy (Stevens to dynamics of Organisms interaction elevations. constraints according vegetation high their gradients geometric characteristic to and elevation mid- to most at along due the clouds habitats area persistent perhaps cloud different largely surface forests, by radiation, characterized habitat cloud are solar as habitats, example, montane well including For as productivity, 1995). content, Rosenzweig biological nutrient 1992; interplay determine and factors type to environmental soil Related or precipitation cover, 2010). gradient and elevation Grytnes an and temperature along (McCain rainfall with peak with in mid-elevation increase variation to moderate tends little of a changes precipitation show relationships exhibit latitudes, organisms, typically complex high mountains living to and tropical due mid- for whereas In general water elevation, 2008). in of (Barry predictable 0.6 topography importance less and ca. much crucial climate of are regional the gradients decrease elevation Despite average along 2008). an increasing precipitation (Barry with with in elevation predictably predictable, in less most or the increase more is m change temperature communities these, biological Among shape elevation. as that such factors composition abiotic factors community Numerous abiotic Guo and 2001; of patterns, (Lomolino gradients distributional systems diversity, by montane regarding characterized in questions are fundamental they (Guo to as moisture answers change, available climate and to temperature relevant those as such 02 n rswtracmctsi h euinAds(Shearer Andes Peruvian the in ascomycetes freshwater and 2012) 02 Coince 2012; tal. et 00,absua yoria A)fni(Gai fungi (AM) mycorrhizal arbuscular 2010), tal et 05 Rinc´on2015; tal et 09.Tefwsuista aeeaie ug ntoia onan eervee recently reviewed were mountains tropical in fungi examined have that studies few The 2019). . tal et 07 Grytnes 2007; . 04,bypyeascae ug (Davey fungi bryophyte-associated 2014), . tal. et tal. et tal. et 05 omnadAnl 08 Sch¨on 2018; Arnold and Bowman 2015; 05 n C ug (Geml fungi ECM and 2015) 92 Wood 1992; tal. et tal. et tal. et 02 Nouhra 2012; 08 Liew 2008; 04 n Mfni(el21)i h uua-oiinYungas, Tucuman-Bolivian the in 2017) (Geml fungi AM and 2014) tal. et tal. et tal. et 03 Perrigo 2013; 03.Hwvr nms raimlgop esillack still we groups organismal most in However, 2013). 93 o 01 Cardel´us 2001; Nor 1993; tal et tal. et 3 tal. et 00.Fnirpeetoeo h ags groups largest the of one represent Fungi 2010). . 02 Coince 2012; tal. et 02,adetmcrhzl(C)adother and (ECM) ectomycorrhizal and 2012), tal. et tal. et 07 nMlyinBre.Consequently, Borneo. Malaysian in 2017) tal. et 2020). tal. et tal et 04 n eas ug,truhtheir through fungi, because and 2014) 03,wo-naiigfni(Meier fungi wood-inhabiting 2013), tal. et 05,evrnetlDAstudies DNA environmental 2015), 08.Hwvr pce richness species However, 2018). . tal. et 04 Miyamoto 2014; 06 Ghalambor 2006; tal et ° e 100 per C 2014; . tal. et et et Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. er,addy(10 mprya)frss h ao oettpssmldi hssuywr classified were study this greater in per is sampled mm there types (1500-3000 forest but moist in major year), drier, occur The per somewhat to mm forests. be known (>3000 year) to are wet per of tends species mm mosaic 2300 side (<1500 a dry Pacific least in and at The resulting year), alone, year. rainfall, in trees is per variation to that precipitation regional hotspot respect mm biodiversity With 3000 a world. than represent the America in (Condit South richest Panama and the For as of Central metals. of number labelled in heavy were one small countries in forests relatively neighboring rich boulder a respectively. and excluded and granite forests, of Panama nutrients were and montane up in montane, upper samples made poor dipterocarp, and forest is montane the are rock lower it regions, that lowland, ultramafic zone: other soils elevation the paper, serpentine with the surviving this comparisons forests for of For montane typical capable Range. and not species Crocker forests plant is (1200–2700 dipterocarp the (3000–4095 vegetation only be zone forests of the these, summit can montane parts because the Of Kinabalu other masl), of 1992). forests Kitayama Mt. in 1200 1990; boulder more of occur (< granite Beaman includes vegetation forests and and and masl), The (Beaman dipterocarp world, (2700–3000 masl) 2004). the lowland forests of Anderson rock zones: Himalayas biotas ultramafic and discrete the masl), species-rich between (Beaman four most mountain of species the into tallest height the of plant average divided is one an vascular Kinabalu has with 5000 Mt. (Malaysia) Kinabalu masl, than Sabah Mount 4095 in Guinea. of range elevation New mountain an and highest With the masl. is 1800 Range ca. Crocker the Borneo, In masl), (400–700 (Brown forest masl) piedmont (1500–3000 types: forest purposes. elevational cloud major 2000). montane three Prado and into 1976; al. masl), 1995; classified (Cabrera (700–1500 Manzano domain are forest and biogeographic region montane Lavilla Amazonian this 1989; the in Mares of forests well and limit The relatively Ojeda southern been have (e.g., the Yungas endemics constitute the Yungas Brown of in fauna Roug´es rich 1997; and and flora and eastern diverse The the Blake rains. very on orographic are developed of forests and result a montane studied as humid Andes subtropical the and of tropical slopes comprise Yungas the Argentina, Geml In by published subset originally small Argentina a Geml here. for northwestern by analyzed except from published Panama, Borneo dataset and from Borneo Yungas fungi novel: the ECM are representing these data of Two the datasets. of three from data analyzed We flora in Methods similarities and consequent Materials and to connection substantially land pools 3). with to histories, (Hypothesis species due biogeographic regions fauna fungal distinct Neotropical and expected two to Nouhra we the due between 2010, Finally, regions overlap biogeographic 2b). Grytnes and more three of host- (Hypothesis and the richness higher among McCain decomposition different species to for 2005, be available due higher (Rahbek elevations energy mid-elevations expected lower more to we at and plant-associated low wood saprotrophs addition, intimate- for at extent generalist In those richness composition and some 2a). substrate than community decomposers to (Hypothesis type wood and and types vegetation pathogens, fungi, richness forest plant by root-associated in elevational affected pathogens, differences among plant less greater fungi ECM, be in as to resulting example, such plants For decomposers, plants, 2). with with (Hypothesis associated zones associated elevation not ly among fungi functionality strate- inferred expected life in distinct we differences would to in structure host corresponding community resulting optima of of and environmental patterns different gies abundance geo- elevational reflecting of groups, and that irrespective functional hypothesized diversity gradients among also elevation by differ We sampled 1). fungi, the (Hypothesis along ECM structure region community of graphic fungal mean case of (MAT), drivers the temperature strongest annual the in mean and, by pH, influenced (Tedersoo strongly plants soil is (MAP), scale precipitation global a annual on distribution and richness 05,hr osdrda oln,lwmnaeaduprmnaefrss epciey o comparative for respectively, forests, montane upper and montane low lowland, as considered here 2005), tal et tal et 04 Vˇetrovsk´y 2014; . 01.TeCrbensd fteitmsadtecnrlmutisrciemore receive mountains central the and isthmus the of side Caribbean The 2011). . tal. et 01.Tgte ihajcn,saoal r idotfrss the forests, piedmont dry seasonally adjacent, with Together 2001). tal. et 09.Teeoe eepce A,MP n olp obe to pH soil and MAP, MAT, expected we Therefore, 2019). 4 tal. et 21) hs eecmie with combined were These (2017). tal et 21) hc sre- is which (2014), . tal. et 01 Brown 2001; tal. et 2018) et Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. ilISsqecs n eeueteegid shpteia ucinlgop.Scnl,frgnr that genera for Secondly, groups. functional hypothetical guilds as functional guilds putative these assigned use we here First, and steps. sequences, two 3 ITS in of tial groups total (Nguyen a functional representing FunGuild to that OTUs, using OTUs OTUs fungal fungal included 620 that assigned 20 only OTUs We contained we artifactual dataset analyses minimize final sequences. subsequent To The high-quality in sequence. samples. 204 work, two reference 962 molecular least fungal at the a in during to occurred generated length Hypothesis been alignment Species have pairwise to may bp assignments 150 with < sequences (K˜oljalgor thresholds taxonomic fungal similarity identified to dynamic database on sequence containing sequences based ITS groups 2016), fungal assigned USEARCH, UNITE+INSD We 22, in curated removed. the August similarity were against were all (released sequence searches sequences sequences similarity from 97% identical pairwise chimeric sequences on at sample, putative to based quality-filtered (OTUs) groups each and truncated The units For singletons counts. were taxonomic discarded. global their operational sequences were and into preserving were all 1 grouped while Sequences > settings: were types Zealand). error following samples sequence New expected the unique (BioMatters, with into with 5.6.1 Galaxy sequences collapsed Pro 2010) in and Geneious (Edgar trimmed removed bp in v.8.0 were were 200 limit USEARCH ends probability tags) with error poor-quality (identification filtered 0.02 and adapters a removed and on were samples based sequences to Primer according (https://main.g2.bx.psu.edu/root). sorted were Sequences sites. sampling the of Bioinformatics Geml coordinates in database geographic WorldClim described the the as on from out based obtained (www.worldclim.org) carried were data were Climate Borneo Panam´a (IDIAP). de and Investigaci´on Argentina Agropecuaria Geml from and samples (2014) an soil with of (Naturalis) Center analyses Biodiversity (Ihrmark Chemical Naturalis fITS7 at primers 250 sequenced 318 with was (ca. NucleoSpin Ion library PCR region amplicon the via ITS2 The amplified The (MIDs). with was DNA-tags pooled. sample repeat were (White sample, DNA each extracts ITS4 each the ribosomal from and For and nuclear soil out the protocol. carried of dry manufacturer’s were bp) the to of extractions provide according DNA ml D¨uren, to independent Germany) 0.5 two zones. Co., dataset elevation & combined from different Gmbh the the extracted (Macherey-Nagel of use was characteristic fungi did and i.e., DNA We forests, regions Genomic kept lowland paper. Paleotropical and samples was and this montane soil Neotropical of sample of mycobiome the location, each focus pantropical among remote ‘core’ of the shared putative more g are the the of that 20 characterization of site analyses first ca. because each the Panama, Borneo, in for and 30-35 In collected region Argentina at later. were immediately weeks in dimensions dried 2 collected were above ca. samples sampling lyophilization the the each until of at to frozen taken cores respect were soil With deep, 10 cm site. Panama, 20 ca. In 4 and repeatedly. (ca. diameter genet S1-3). in same (Figs. cm regions 2 the three cores, 10 the soil (ca. to 40 site and corresponding Borneo, coordinates, maps and geographic on Argentina elevation, del and type, In Forest S1 Bocas regions. Table in elevation respective in 2016, the entire shown June, in are the in forests in represent Borneo; localities of and Sabah, sites types Argentina; various of sampling of of Parks Tucum´an The range provinces Range and Panama. Crocker Salta of and Jujuy, Panam´a Chiriqu´ı, provinces Kinabalu Col´on in and Toro, in 2013, 2012, and September, 2011 in May, collected were montane samples upper Soil and masl), work (800–1500 molecular forest and montane Sampling lower masl), (Condit (0–800 masl) forests (1500–3000 moist forests and wet lowland as × TM ) olcrstkna ie iewr old eutn nacmoiesi apefreach for sample soil composite a in resulting pooled, were site given a at taken cores Soil m). 5 hpada o orn esnlGnm ahn Lf ehoois ulod T U.S.A.). CT, Guilford, Technologies, (Life Machine Genome Personal Torrent Ion an and Chip × 5m.Crswr olce a rmec te omnmz h rbblt fsampling of probability the minimize to other each from m 2 ca. collected were Cores m). 25 tal. et tal. et 21) epciey h ape rmPnm eeaaye yteIsiuode Instituto the by analysed were Panama from samples The respectively. (2017), 90.TeIS rmrwslble ihsml-pcfi utpe Identification Multiplex sample-specific with labelled was primer ITS4 The 1990). tal et 05.W eonz h iiain ffntoa neec ae npar- on based inference functional of limitations the recognize We 2015). . ° tal et .Bcueo hs ieecs ehnldtedt eaaeyfreach for separately data the handled we differences, these of Because C. 2011). . 5 tal et 03.W xlddOU ih<8%similarity 80% < with OTUs excluded We 2013). . tal. et ® olkit soil 2012) tal. et Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. niiulOU ihseiceeainlfrs ye tapnrpclsaewt the with of scale associations pantropical quantify a to at 1997) types the Legendre forest (Dufrˆene elevational for and the analysis specific as with species well OTUs indicator as the individual out region with carried geographic standardized we each variables, Finally, in For environmental including test model. abiotic parameters, Mantel the final partial and of used with the selection we forward group in comparison, a variables cross-regional functional performed environmental each we significant and variables, only of environmental fungi estimate sources among all To (PerMANOVA) as correlations variance for variables region. for of (continuous) out any analysis environmental multivariate carried in and permutational was types composition, (categorical) community type forest fungal forest elevational in variation of across importance the dispersions relative via group the variances of equality homogeneity the the with multivariate ordinations of GNMDS tests the on Statistical elevation of we- isolines geographic ( composition plotted coefficient We each community ordinations. correlation fungal in in Pearson and plotted fungi the measure. variables with all distance environmental calculated Bray-Curtis between for re with significance as run statistical well per and as values iterations mentioned 999 groups variables to functional environmental subjected for containing the separately matrix with run region secondary were a Ordinations the and above. used table we OTU types, Hellinger-transformed forest and Panama. elevational content, and across eleva- Argentina nitrogen nen between in composition total content relationships community matter, P examine organic compare total forest To pH, to as among soil R, well MAP, them as in MAT, richness compared ratio, also as proportional (C/N) and analyses, such carbon/nitrogen of basis variables regression values environmental per-sample HSD quadratic Tukey’s calculated and a used with We tion on We functio- ANOVA 2013). above. groups and via Team as functional classes types Core types taxonomic of forest Development of abundance elevational richness (R proportional major OTU R and three as in the well implemented among as test, compared richness were fungal groups total nal region, geographic biogeography core estimate of each putative to effects In reads possible of composition. 2000 the community characterization the distinguish to on and for rarefied the evaluate variables dataset reads For to environmental a 241 and Panama). and used regions (14 for we geographic region among fungi, 2000 that overlap lowland species and in and size montane Borneo, pantropical library rarefying for of smallest by communities 812 the analyses to 24 statistical subsequent sequences Yungas, for fungal Argentinian table high-quality OTU of the number normalized the we region, biogeographic each For versions, first the are paper analyses this Statistical respectively. in KDPZ01000000, deposited described and been versions KDPY01000000, have (Ma- The KDPX01000000, regions (Panama). KDPY00000000 i.e. Argentina, Yungas), KDPZ00000000 study groups, (Argentinian from and functional quality- three Borneo), to KDPX00000000 samples the The laysian assigned accessions the to 65%). were under in corresponding OTUs (ca. DDBJ/EMBL/GenBank OTUs 2896 projects OTUs at and fungal Study 356 5078, 4498 Locus 5917, 13 and Targeted which for respectively. of 7829, utilize respectively, made 8720, Panama, can were contained and generally fungi assignments Borneo, dataset that separated functional rarefied saprotrophs we total, generalist and Similarly, respective- from In filtered fungi. pathogens, wood sugars. plant saprotrophic of simple or purely carbohydrates only animal as from complex as grow degrading distinction designated can of ecological were that capable Fungi pathogens the available database. with plant acknowledge basis UNITE or to case the animal by ly, in as case Hypothesis a well on Species as OTU matching saprotrophs each the to for function ecological information assigned ecological we Sebacinales), (e.g., the guilds in functional genera multiple from species comprise indicspecies tal et 05 ornGnrlzdNnercMliiesoa cln GMS riain nthe on ordinations (GNMDS) Scaling Multidimensional Nonmetric Generalized run to 2015) . akg nR(eC´aceres (De R in package metaMDS envfit ucin hc sssvrlrno trst n tbeslto.Dt were Data solution. stable a find to starts random several uses which function, ucin n etr fvralswt ttsial infiatcreain were correlations significant statistically with variables of vectors and function, tal et 2012). . betadisper 6 Amanita vegan ucinidctdn infiatdffrnein difference significant no indicated function scale odffrnit h ffcso pta distance spatial of effects the differentiate to , Entoloma ucini ,o omnt structure. community on R, in function adonis , ucinin function Ramaria vegan , multipatt Sebacina vegan akg (Oksa- package R ordisurf oaccount To . n related and ucinin function function. r ) Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. mn lvtoa oettpsi l he ein Fg ,Fg 5.Ro-soitdfni h only the to compared fungi, fungi Root-associated plant-associated the 2a. in regions, among Hypothesis types meaningfully two S5). with vegetation differ consistent in among not are richness Fig. differences did guilds in Greater non-plant-associated generally 3, saprotrophs significantly types. generalist differed forest (Fig. and and elevational pathogens, mycoparasites pathogens, regions significantly of animal plant differed three richness fungi, richness and while all (ECM proportional guild, plants in as plant-associated well with types at in as remaining associated types forest richness except in all forest OTU elevational types, other groups, groups observed forest among the functional which functional elevational than for three three OTUs all decomposers), found all fungal We in wood in more observed significantly similar 3). harbored generally were forests (Fig. was also montane richness lower types fungal where Total Panama, forest region. elevational We one in among least groups 1). than richness 2a. (Table functional plant-associated Panama Hypothesis in in the to in types in Differences contrast forest 31.5% 26.5% elevational in and among and groups, turnover 13.0% regions, 11.5% functional community groups, between all between non-plant-associated greater of functional and In for in and all Borneo, terms support fungi vectors. almost find in in all not as in 32.6% in did groups displayed composition and variation orders community functional 15.3% the in taxonomic between largest of variation of Argentina, 20% four of richness source the with ca. significant edaphic 2, of explaining a and ordinations was elevation Fig. plots type with in GNMDS GNMDS forest shown correlations The 2. are similar Hypothesis richness showed above. general OTU groups detailed our functional with as to consistent variables corresponding regions, datasets geographic the all of in types forest ( elevational significant exhibited in groups functional composition. important All groups pH, community MAT, especially fungal functional to of soil fungal being respect of of proportions MAP with drivers patterns particularly significant and as Elevational 1, MAP, regions explaining extent Hypothesis two with lesser OM consistent a least and are varied to at results variables and P These in abiotic with fungal 1). structure regions other (Table total regions, community of three Argentina the contribution fungal all geographic The in in in and variation variation model for. groups greatest combined accounted functional the the was explain among in parameters to contributors among significant correlation tended remained when pH and and MAT composition community analyses, (all PerMANOVA significant the not was In proximity controlled. were spatial variables whereas tests sampling abiotic matrix), Mantel all when (control Partial in 0.1) for regions. accounted structure was community the proximity to among spatial robustly importance linked (all were their (Argentina: regions pH variables in areas three soil environmental differed and all abiotic MAT variables in that variables, edaphic indicated composition abiotic and community to climatic fungal respect three other With with all 1). strongly in (Fig. correlated elevation respectively were to Panama, (Fig. according and Kinabalu communities Borneo, Mt. values: Argentina, fungal of correlations of zone correlated (Pearson’s structuring summit were regions strong the N, revealed extent in lesser ordinations trend a not GNMDS of to was change and trend with (OM), the again material Panama zone general, organic in S4). summit soil in vegetated although manner, elevation sparsely regions, related with the a all representing positively zone masl, In in forest generally 3500 Kinabalu. elevation montane above lower MAP Mt. with again the of regions. increased decrease in pH three highest to Borneo, was all tended In it significant. pH in regions. where Argentina, elevation Soil sampled in S4). three with except the (Fig. elevation correlation in with negative correlated degree strong not varying was showed a MAT to elevation expected, with As correlated variables environmental Abiotic drivers Environmental Results r 0.636, = p .0;Borneo: 0.001; < r 2 p 0.8633, = .5 ieecsi omnt opsto mn h three the among composition community in differences 0.05) < r 0.444, = 7 r 2 .46 and 0.9406, = p .0;Panama 0.001; < r 2 .61 all 0.9661; = r 0.576, = p .0) whereas 0.001), = p .0) when 0.001), < p .0)in 0.001) = p > Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. obndi ata atltss eutn nsgicn orlto fevrnetlfcoswt fungal with factors environmental of correlation significant in resulting tests, Mantel Neotropical partial two in the community ( fungal combined among tests between with Mantel overlap correlated separate species strongly little were in shared location i.e., structure geographic more and 3, variables between environmental with Hypothesis abiotic shared Both with regions, were consistent biogeographic 192 the are and three between findings shared Borneo, the These were regions. and of 376 Yungas 4). but pools the three region, all between species (Fig. geographic among Panama shared one shared were and were to OTUs Borneo 188 restricted 211 were regions, only OTUs Neotropical these, of two Of majority OTUs. The 7612 included regions. dataset combined forests rarefied, montane The and lowland lowland in Similarly, richness of S7). detected lower mycobiome and Argentina. core only (Fig. Pantropical higher were respectively) in (i.e., forests, types significant observed montane forest were not upper patterns elevational was in contrasting the where than difference in increased among Panama, rich the Agaricomycetes richness and more Borneo of though their always in richness of lowlands, (i.e., was in richness Argentina the subphylum differences in Mortierellomycotina in particular, significant The patterns than In contrasting forests forests respectively). showed montane montane elevation, S7). lower they with in but decreased peaked (Fig. than Panama, Saccharomycetes and montane regions and and upper Archaeorhizomycetes three Borneo in of respectively, all in Richness lower, in and forests. higher lowland types was in forest consistently elevational Sordariomycetes and significantly the Leotiomycetes differed among at Sordariomycetes richness Xylariales and and in Saccharomycetes, Agaricales Leotiomycetes, and Archaeorhizomycetes, Helo- elevations, Overall, decomposers, higher wood at 2). Among forests (Fig. in forests. elevations richness montane lower highest in Chaetothyriales, showed saprotrophic richness for consistently higher vectors Eurotiales, tiales contrast, indicated Agaricales, In Sebacinales included regions. and two elevations least signifi- Helotiales, lower at with in at each values Sordariales, forests correlation and Pearson’s in cant (Panama). Sporidiobolales, richer Saccharomycetales, forests Kickxellales, were montane Hypocreales, that upper pathogenic Geastrales, fungi or putatively (Borneo) saprotrophic whereas forests of forests, montane Orders lower lowland in towards rich Rhi- mostly more correlations Polyporales, directed were significant Ophiostomatales, were Helotiales with Hypocreales, pathogens Xylariales) in plant and Diaporthales, richness of zophydiales, Cantharellales, highest groups Botryosphaeriales, taxonomic with in most Thelephorales 2). of (e.g., the and richness (Fig. towards the Sordariales forests directed representing montane were of Vectors upper fungi exception in root-associated the Helotiales Borneo, and and with In ECM forests, of sites, lowland Archaeorhi- orders Thelephorales). forest Agaricales, taxonomic and montane (e.g., most forests lower Sebacinales, for significantly montane richness Russulales, correlated upper of that Hysteriales, in vectors sites richness Helotiales, Neotropical highest the Eurotiales, exhibited in zomycetales, axes fungi ordination root-associated and the ECM with of orders taxonomic elevation All among groups decomposers taxonomic Borneo, wood of in of patterns S6). forest abundance Elevational montane (Fig. proportional lower regions the the three in in all peak differences saprotrophs abundance in significant pronounced of zones of a minimum of lack abundance exceptions the the notable and richness fungi, the and proportional with fungi to ECM abundance, in respect mycoparasitic diverse proportional with contrast, of most observed in were were In extent they also some patterns but in to Similar forests, example, montane and 2b. For Borneo. upper in S5). the Hypothesis forests in montane Fig. richness lesser lower with 3, in the a peaked montane consistent (Fig. fungi lower patterns to ECM largely and/or mixed forests, showed and, lowland Neotropical forests, fungi in decomposers, root-associated montane higher non-ECM wood and be upper mycoparasites, to and in tended pathogens than saprotrophs, plant generalist forests and of pathogens richness animal proportional extent, and richness Observed r .5 and 0.658 = r .9,rsetvl;all respectively; 0.497, = 8 p .0) n hnte were they when and 0.001), < Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. olp lopasa motn oei hpn eogon uglcmuiis(Porter communities fungal seasonality. belowground the shaping pronounced al. in in less role et moisture important far available an have plays of which also amounts pH Panama, Soil greater and environmental and Borneo stronger variability of a sub- limited represent forests mainly the may tropical The (Brown than stress, wet functional Panama. drought assembly elevations community temporary the and lower resulting for Borneo of community at the filter in of majority and particularly insignificant terms moisture, the seasonality, in or in in pronounced differences marginal example, 1-2, variation has For remained (Figs. explained Yungas region-dependent. it composition of more tropical while community contributor be Argentina, fungal to strongest in of seems the groups driver MAP was influential of MAP most role alone. composition, the the area is while habitat communities MAT 1), biological than that Table of mountains composition indicate in as richness results well fungal Our as of factors drivers the edaphic S1- showed influential three and groups (Figs. more the climatic other Panama far that elevation, of and no likely are increasing any Borneo with is were in richness in It there in forests trend. particularly decreased patterns: montane opposite areas, groups richness larger upper functional observed much increasing some and cover Although the with lowland forests 3). for area among lowland explanation richness though habitat richness even satisfying fungal decreasing regions, species total a the with in offer fungi, positively difference to for correlated statistical However, seem is not 1998). often (Gotelli does area elevation groups habitat cover and taxonomic generally ones many habitats elevation elevation relative in lower higher influences Moreover, than elevation strongly conditions. different areas edaphic in which smaller found and temperature, types mesoclimatic forest in biotic distinct the Consequently, have and differences processes. zones abiotic chemical by through soil indirectly mediated and moisture, is or soil richness humidity, directly and either composition community driven fungal factors on also elevation of forest effect montane the The lower of the communities types, forest between drivers two intermediate Environmental other as fungal the and of regarded as taxa. climatic characteristic composition although, be several Furthermore, contrasting forests, may possesses vegetation. montane to forests in upper according montane differences and filtering lower lowland extent, environmental the smaller resulting in a the communities to and and (Cardel´us elevation conditions, mountains by edaphic American driven Central fungal be in in richness peak to richness mid-elevation fungal plant a the in of vascular where differences lack exception, latitudinal Prada with The of an concordant lack was Borneo. is Panama the and (Vˇetrovsk´yrichness 2019). to Argentina scale similar al. in global is et zones a richness elevational on fungal montane three in upper the pattern than among elevational richness species strong vascular more fungal to soil harbor Contrary typically in Brown Paleotropics. strongly forests differences 1999; the is montane Kitayama and lower groups, and functional Neo- the (Aiba all the and forests of lowland both that the in presented as data types where well sequence plants, forest as deep community, elevational The similar fungal to total from rare. according the are data structured scales of compare global composition to and that local were opportunities show at and here types 72 known, forest and tropical little diverse remains across 24, guilds forests 40, tropical in these, biodiversity S2). Fungal Of (Table respectively forests, comparison. montane pantropical upper the and Discussion montane, in lower included lowland, the were for and indicators consistent ( Paleotropics model, significant and combined with Neo- the OTUs in fungal contributions the 360 significant Of of pH with 5.8% 7.4%, all and variation, explained 1. 7.3% the MAP Hypothesis explained variables, of with abiotic type 5.7% among MAT forest while and and respectively, 6.9%, region composition, geographic community that in variation indicated ( PerMANOVA for accounted 0.001). was < location when structure community 00 Lauber 2000; tal. et 07.Hwvr nalfntoa rusadi l ein,cmoiinlsrcueappears structure compositional regions, all in and groups functional all in However, 2017). tal. et 08 Rousk 2008; p tal. et .5 niao au o oettp,16ocre ohi the in both occurred 136 type, forest a for value indicator 0.05) < tal. et 00 Geml 2010; 01 canadGyns21) eddntfidsubstantial find not did we 2010), Grytnes and McCain 2001; 9 r 0.553, = tal et 04 Tedersoo 2014; . p .0) and 0.001), < tal et ieversa vice tal. et 04 Glassman 2014; . tal et 97 Coughlan 1987; 01.Local 2001). . ( r tal. et 0.279, = 2006, et p Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. oet n eln nuprmnaefrss hc srflce eeadi icse ndti yGeml by detail in discussed is montane lower and and here lowland reflected in is high which are highest forests, trees reported montane ECM that upper of al Neotropics in (Mueller density wet decline forests and the and richness montane forests of soils, upper areas non-ultramafic in other in fungi in Borneo, ECM studies of of sporocarp-based density richness with highest the agreement between the in fungi harbor are ECM to of tend (Malizia patterns Panama forests tree elevational montane on the upper Malizia studies with acuminata in 1999; Neotropics, previous i.e., Kitayama differences the trees, from and important In ECM (Aiba inferred in Palaeotropics. types as results and (Tedersoo forest Palaeotropics, also Neo- sampled scales This and the regional with density 2012). Neo- of and and positively . richness the composition global host correlates and both mirrors on richness richness richness composition in fungal community both community fungi, ECM gradients plants that ECM and elevation suggest host For richness here along presented ECM plants. species data of the with for Similarly, density 2014). associated type and fungi vegetation richness of of taxonomic groups importance functional the of suggest various in results observed Our been have (Cardel´us also animals function composition and a and as Guo plants composition richness of in in groups differences functional envi- differences strong different in functional elevational observed by Even we favored Similar zones. richness, elevation. elevational be in the to of differences among seem significant differences strategies no life functional with different by groups with indicated fungi as that conditions, show ronmental here presented pH. results sparsely soil The communities in low fungal sharply in soil drops result in accumulation not OM dynamics does whereas Elevational and and forests, Kinabalu accumulation summit montane Mt. OM the the of greater near in zone with the habitats However, summit associated pH MAT. vegetated granitic are in of vegetated temperatures decrease that sparsely lower simultaneous from habitat, the a effects forest in their the OM disentangle within in and to that decrease suggest difficult and fungi is pH soil it in between and increase elevation, observed interactions with competitive OM in and availability increase nutrient altering (Rousk by bacteria e.g., indirect, mainly Nevarez 1991; al. ne h ucinlgid fpatptoesadsporpsmypoet eedpye,riigan raising endophytes, be identified to fungi prove the may of saprotrophs some that and and note pathogens specificity We habitat plant well. suggests of as groups, guilds commu- guilds functional competitive strong functional all functional of richness, the in within result in observed under saprotrophs species a differences elevation, of of as of of turnover richness possibly function Regardless elevational fungi, proportional a effect”. as lowest ECM Gadgil structure of the “the nity richness i.e. regions, patterns above, proportional three richness mentioned highest but all interactions elevations, the In lower with at predictable. coincided higher less predictably always was were fungi, saprotrophs pathogenic (G´omez-Hern´andez plant of extent scales some global to to landscape-level and from repor- studies al as of temperature et range with wide positively ECM- correlates 2016). a in generally by Kennedy fungi as ted and saprotrophic rhizosphere, Fernandez and pathogenic the 1971; plant to Gadgil of Richness C and contribute fixed (Gadgil likely of decomposition forests addition inhibit montane continuous that in the fungi fungi through (Clemmensen ECM forests OM of boreal soil in abundance dominated of patterns and to accumulation diversity above tends the greater richness The to The fungal study. ECM 2018). this (Bahram where Arnold in elevation mountains, and included increasing temperate with in not observed monotonically were those decrease which from 1999), differ Kitayama mountains tropical and (Aiba soils ultramafic 21) thg lvtos C ot uhas such hosts ECM elevations, high At (2017). . 07 el21) eas ayfna pce aearltvl iep piu eg,Wheeler (e.g., optimum pH wide relatively a have species fungal many Because 2019). Geml 2017, tal. et 04 el21) norsuy ihes(ohpootoa n boue fwo decomposers, wood of absolute) and proportional (both richness study, our In 2019). Geml 2014, . 2013). n several and tal. et tal. et tal et 09,i slkl htteosre orlto fp ihcmuiycmoiinis composition community with pH of correlation observed the that likely is it 2009), lu acuminata Alnus 00 n te olboa nalrgos eosre infiatdces np and pH in decrease significant observed we regions, all In biota. soil other and 2010) 02 aple21;Wicaksono 2016; Kappelle 2012; . Quercus p.i h onanrnesann otenCsaRc n northern and Rica Costa southern spanning range mountain the in spp. tal. et stepicplEMhs nteYna nAgnia and Argentina, in Yungas the in host ECM principal the as 03,ada euto eaieitrcin ihsaprotrophic with interactions negative of result a as and 2013), tal et 10 etsemmrecurvum Leptospermum 06 can20;MCi n rte 2010; Grytnes and McCain 2009; McCain 2006; . tal et tal. et tal. et 07 Nouhra 2017; . 02 Nouhra 2012; 06 G´omez-Hern´andez2006; edt edmnn nyin only dominant be to tend tal. et tal. et 02 u e Bowman see but 2012; 08.Teefindings These 2018). tal. et tal. et 02 Tedersoo 2012, 02.In 2012). Alnus tal. et tal. et tal et et Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. ein sicmlt,i spsil htmr hrdtx ilb dnie hog h cuuainof accumulation Peay low- 2011; in the Geml species through many (e.g., these identified as patterns in fungi, phylogeographic be in mycota pronounced will widespread and hyperdiverse al is taxa limitation et the endemism shared dispersal regional of show Nonetheless, more mid-latitudes sampling future. that to the our possible in Because data is field Neotropics. it more the endemism incomplete, of of is level high parts regions relatively a different indicate within regions geographic even among differences compositional observed The of irrespective often orders, patterns closely pantropical taxonomic relatively Emerging of for optimum level environmental the and at constraints taxa. observed physiological related be shared (Semenova suggests already ecosystems guild, can tundra functional preference arctic decom- in habitat ascomycetes wood that of and fact order apparent saprotrophs, diverse similar is pathogens, most a plant al. the forests In be et fungi, cloud elevations. to montane root-associated low appear as in at Helotiales such Helotiales, soil posers. groups, particularly in boreal functional lowland prevalence Leotiomycetes, in several tropical their of Unlike in in with richness saprotrophs. fungi agreement higher endophytic and as in the plant host well 2007), of manner, higher Lutzoni as class with and dominant pathogens agreement the (Arnold plant in are forests include is Sordariomycetes which and forests, groups temperate fungi, Xyla- functional and these and various for ex- Sordariales in For diversity apparent Hypocreales, niches. functional substrate is particularly environmental elevations across Sordariomycetes, to lower groups of respect at richness with taxonomic riales, conservatism higher of phylogenetic consistently distribution of the level ample, the certain regarding a follow patterns suggesting broadly guilds, consistent will several endophytes overlap observed sequences, phylogenetic We ITS and of ecological terms here. the in described given taxa patterns that pathogenic the suggests and work saprotrophic, Our endophytic, study. of future for direction interesting ug ln lvtoa rdet navreyo as .. yaetn irba rcse eg,decompo- (e.g., processes microbial and affecting Neo- by the e.g., of ways, in distribution of gradients the variety elevation shaping a factor along in driving fungi gradients the of elevational be along to structure appears community fungi temperature, comparing particularly Neotropical study Climate, in first Paleotropics. OTUs the of is This 2-4% and 4-12% reach and and Conclusions Neotropical in rainforests, OTUs respectively. lowland of forests, Paleotropical 1-5% function). montane and in unknown upper 1-2% currently represent 1-2% with fungi fungi and mycoparasites including root-associated and community 4-8% non-ECM representing whole 2-4%, and the pathogens ECM saprotrophs for addition, animal calculated forests: In mechanisms 6-15%, (as lowland OTUs decomposers similar in the in wood resulting by of 11-17%, guilds 1-2% forests, driven pathogens functional Paleotropical plant be the and remar- 35-40%, may is Neo- of ca. proportions regions throughout representations three gradients niche predictable elevation the functional relatively along among that groups filtering hypothesize environmental functional We the of of S5). formation). richness cloud there (Fig. and proportional conditions kable humidity of environmental relative similarity characteristic higher unique the accumulation, the a OM Likewise, to represents increased likely specialized fidelity pH, forest and species and montane specificity of temperature upper lower high proportion The (i.e. with analyses. higher taxa species a indicator in the with by rich of habitat shown particularly distributions as are habitat, the forests their shaping montane to in upper factors in (Vˇetrovsk´y type. communities abiotic scale Fungal forest of global fungi elevational a importance among certain on the even a fungi with that for common agreement suggests specialists most in habitat This of is are zone. drivers observation the many elevational important of This distributions, an two-thirds as geographic Almost for least widespread history). indicators at biogeographic with were still (i.e. OTUs are geography variables pantropical as environmental fungi putatively tropical species, of shared composition of community number low the Despite 02 Branco 2012; . 05,adteaoetedcnrsta ayHltae aatrv nrltvl odrciae.The climates. colder relatively in thrive taxa Helotiales many that confirms trend above the and 2015), tal et 05,rsligi ieecsi h opsto frgoa pce pools. species regional of composition the in differences in resulting 2015), . tal. et 11 2019). Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. akrM,BunrI swo F Ashwood I, Brunner MR, Bakker emnJ,Baa S(90 iest n itiuinpten nteflr fMutKnbl.I:Baas In: Kinabalu. Mount of flora (eds.) the R in Geesink patterns distribution P, and Kalkman Diversity P, (1990) RS Beaman JH, Beaman (2004) C Winteraceae. Anderson JH, Beaman (2008) RG Barry forests. European of northern services of forests Hyrcanian ectomycorrhizal the of (2014) in patterns gradient OT altitudinal local Lewis an and RP, along Regional Iran. structure Freckleton (2012) community L CE, and Tedersoo diversity Addis S, fungal composition. L, Zarre K˜oljalg and U, P˜olme Narayan S, diversity M, SJ, plant Bahram rainforest Gurr drive S, herbivores biodi- insect Gripenberg leaves and RE, tropical Pathogens are Gallery endophytes: R, fungal foliar Bagchi of range host and Diversity hotspots? (2007) versity F life. Lutzoni mountain AE, of of Arnold origins matrix Multiple altitude-substrate Biodiversity: an (2015) Borneo. in A diversity Kinabalu, Antonelli Mount species on and communities composition tree Structure, forest (1999) rain K Kitayama S, Aiba and REFERENCES KDPY01000000, KDPX01000000, DDBJ/EMBL/GenBank; metagenome; Soil KDPZ01000000 by 2020; support J; acknowledges Geml Geml [dataset] J. interest. T´amogatott sequencing. of authors Kutat´ocsoportok Irod´aja. Torrent the conflict The Universiti Ion no of Eurlings declare 96049) the and Marcel Scientific (no. conducting for Parks Lend¨ulet The programme and for for MTA-EKE Sabah fieldwork, permits. the (Naturalis) during Institute and support Duijm export Panamanian logistical Panama Elza providing and the in for and access Malaysia (INDICASAT-AIP) Agentina, in issuing (UMS) Services in Sabah for Technology Fundaci´onMalaysia ProYungas Malaysia High the in and in for Parks Panama, sampled Research in grateful Sabah reserves Environment of provincial and are Ministry and Center authors Panamanian parks, the Biodiversity S1), provincial Sabah Table reserves, in and Lutzoni. national (listed Panama F. parks, and to was national Malaysia grant Argentina, all work GoLife NSF of Molecular above members Lutzoni. the staff and F. Geml to J. and 1541548) to thank expedition, grant (DEB authors Range Initiative grant Kinabalu-Crocker The Research the Naturalis C´ordoba, GoLife a of de (NSF) by part Nacional Foundation financed Universidad as Science Parks (SECYT Sabah Nouhra National E. and the by Naturalis provided 124/13), was 162/12; fieldwork future 26/11; the near for the fungi. support in many Financial possible for be habitats will suitable the it studies, model Acknowledgements and metabarcoding accumu- niches from the climatic functionality. characterization taxa With the in efforts. habitat fungal determine conservation also for and to and but points monitoring assessments composition, data biodiversity for in spatial in possibilities only fungi lating offers communities not incorporating fungi future ones advocate we many current guilds, and from by functional considerably broad exhibited differ the groups specificity may taxonomic within Habitat several site them of given among preferences a differences are habitat at functional fun- contrasting forests affect possible the Montane undoubtedly Given the will dynamics. warming ecosystems. and interaction and these change in species climate communities to altering ecosystems gal by terrestrial vulnerable and most factors, the among edaphic and vegetation, sition), e Phytologist New oa oai adn,Kew. Gardens, Botanic Royal Ecology onanWahradClimate and Weather Mountain , 193, , 88, rnir nFrssadGoa Change Global and Forests in Frontiers 465–473. 541–549. h ln iest fMalesia of diversity plant The h lnso on iaau .Dctldnfmle anlaeeto Magnoliaceae families Dicotyledon 5. Kinabalu. Mount of plants The tal et 21)Blwrudboiest eae oiieyt ecosystem to positively relates biodiversity Belowground (2019) . abig nvriyPes abig,UK. Cambridge, Press, University Cambridge . 12 ln Ecology Plant uwrAaei ulses Dordrecht, Publishers, Academic Kulwar . Nature , 2, , , 6 140, 524, 139–157. 300–301. Nature , 506, 85–88. Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. aglR,Gdi D(91 yoriaadlte decomposition. litter and Mycorrhiza control (1971) interactions PD fungal Gadgil interguild RI, do Gadgil effect’: soils? ‘Gadgil forest the in Revisiting cycling (2016) carbon PG Kennedy CW, BLAST. 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Data may be preliminary. abrC,Srcln S rdodM,Fee 20)Teiflec fsi rpriso h structure the on properties soil of types. influence land-use The across (2008) communities N fungal Fierer and MA, bacterial Bradford of MS, Strickland CL, Lauber fungi. Borneo. of Kinabalu, K cation Abarenkov Mount RH, on vegetation Nilsson K˜oljalg the U, of study ed., transect M, altitudinal 102 Kappelle An In: (1992) Talamanca. K de Kitayama Cordillera the of Forests Cloud ecosystems Montane Rican The ectomycor- gradient. (2016) of elevation distributions M m) the Kappelle (300 in short partitioning a altitudinal Strong along (2015) fungi AFS rhizal Taylor S, Woodward SG, Javis communities. natural and artificial 677. of 454-sequencing by K evaluation Cruz-Martinez B¨odeker ITM, K, Ihrmark Z Sun analyses DA, simulation Kelt matters: Borneo. Q, effect Kinabalu, Guo mid-domain Mount The from (2008) data C distribution Rahbek bryophytes range-size TS, of of Romdal patterns JH, richness Beaman Himalaya. species JA, Central elevational Grytnes Nepal, of in comparison groups A plant (2007) HUB other Birks with JA, Grytnes O, analy- Grau metacommunity and gradient com- (1998) fungal NJ macromycete Gotelli for of options Patterns gradient: (2012) altitudinal DJ an ses. along Lodge assemblage R, communitymunity Guevara drives nutrients G, soil scales. and Williams-Linera G´omez-Hern´andez spatial pH by fine M, filtering at Environmental fungi (2017) Janzen’s in TD tropics? assembly Bruns the IJ, in Wang higher SI, passes Glassman mountain Are Borneo. (2006) Wang in G. Revisited. 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Mount of plants The , 20, ora fBiogeography of Journal , 241–248. 21, , ea:CmuiyEooyPackage Ecology Community Vegan: 15 185, 53–73. 55–68. SEJournal ISME , Mycologia 42, lblEooyadBiogeography and Ecology Global 1083–1090 , , 104, , eiilu glabrum Penicillium 37, 8, nentoa ora fFood of Journal International 45–52. 1739–1746. nylpdao ieSciences Life of Encyclopedia 1071–1078. eso 2.3-2. Version . lblEcology Global Biodiversity tanLCP strain , Labora- . 19, Nature Notho- Royal 346– , Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. otn lu oet:acs td fteAda le ( Contracting alder (2017) Andean J Geml the S, of Pacheco study N, Raes case N, a Pastor forests: ER, Nouhra cloud J, montane of Gutierrez eds. RNA Aguirre TJ, species ribosomal CY, White toxigenic fungal Wicaksono JJ, of Sninsky some sequencing DH, direct of Gelfand Applications and growth MA, and Amplification Innis Methods (1990) the In: JW Taylor on phylogenetics. S, for pH Lee genes T, of Bruns Influence TJ, White (1991) JI Pitt BF, Aspergillus Hurdman KA, Wheeler patterns. Thu climate-driven AM, Vasco-Palacios LV, Kopeck´y M, Ruiz P, R, Vˇetrovsk´y Kohout Wijensundera NS, T, Yorou U, A, Koljalg S, Suija QP, rule Polme latitudinal M, Bahram Rapoport’s L, of Tedersoo extension an range: elevational in altitude. gradient to elevational The (1992) GC biodiversity. of Stevens Mountain patterns (2012) Distributional C Korner (2015) 301–302. K, JP Peru. 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Press, University Cambridge . lu acuminata Alnus , , 17, 24, , 3009–3024 13, 659–662. e0208493. Science , 16, ln clg n Diversity and Ecology Plant 4122–4136. n soitdfnii the in fungi associated and ) ora fVgtto Science Vegetation of Journal C rtcl:AGieto Guide A Protocols: PCR , ora fBiogeography of Journal 346, , 12, 1256688. 141–150. Biodiversity Molecular . Ecology , 4, , Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. i.4 hrdadecuieOU costetresmldgorpi ein ae nterrfidPan- rarefied the on based regions geographic sampled text. three the in the detail across in OTUs dataset. described exclusive tropical are and types Shared forest Forest 4. elevational forest. Fig. tests, montane the HSD upper Tukey’s across ( = and groups ANOVA UM differences using functional forest, compared significant of montane were denoting Means richness regions. letters and geographic richness with sampled three fungal the total in phosphorus of types soil Comparison = P 3. content, nitrogen Fig. = soil light MAP = with N temperature, content, indicated annual matter mean are organic = soil sites content. MAT = ordi- forest OM Abbreviations: with montane respectively. precipitation, annual correlation upper symbols, mean black significant and and montane with grey, lower functional orders dark displayed Lowland, grey, taxonomic elevation fungal displayed. and with are of data, variables axes Hellinger-transformed environmental plots nation on of ordination based Vectors types (GNMDS) isolines. forest as elevational scaling organic sampled respectively. multidimensional soil the content. symbols, in = phosphorus non-metric black groups soil OM Generalized and = precipitation, 2. grey, P annual content, Fig. dark mean nitrogen grey, = soil light MAP = with N temperature, envi- content, annual indicated of matter montane mean Vectors are lower S1. = Lowland, sites Table MAT displayed. in forest Abbreviations: are given montane axes are ordination elevation upper sites with with sampling and correlation data, the significant Hellinger-transformed of with on descriptions variables based and ronmental regions Localities three in isolines. communities the fungal as each of of displayed types for plots forest elevation) ordination elevational (GNMDS) sampled (without Climatic scaling the multidimensional model bold. non-metric in composite Generalized 1. are final (*). Fig. results asterisk the by Significant in indicated matrix. significant are community remained group fungal fungal that type the variables forest on elevational multiva- edaphic based by permutational and variance, explained with of independently composition analysis community calculated variables riate fungal manuscript environmental in the continuous of (%) and revisions variation (categorical) the of to Proportion contributed Figures authors 1. version. for all Table maps submitted and the first paper prepared the the Hegyi of in B. draft J. resulted scripts. statistical first R that sites. the the contributed wrote and sampling Morgado Geml bioinformatics L.N. selected J. the and S1-3. and Geml completed Grau J. Geml O. research fieldwork. which J. the the to labwork. analyses, performed designed the Ib´a˜nez Morgado performed Arnold A. L.N. Semenova-Nelsen Arnold, A.E. and T.A. A.E. and and Lutzoni, F. Lutzoni, at Nouhra, F. E.R. DDBJ/EMBL/GenBank Geml, Nouhra, in E.R. Geml, available J. openly are study contributions: KD- Author this and of KDPY01000000, findings KDPX01000000, PZ01000000. the numbers support i.e. reference versions, https://www.ncbi.nlm.nih.gov/genbank/, that first data DD- the The are at paper deposited this been Availability: in have Data described respectively. regions versions KDPZ01000000, study and The KDPY01000000, (Malaysian (Panama). three KDPX01000000, KDPY00000000 KDPZ00000000 the Yungas), and (Argentinian to Borneo), KDPX00000000 corresponding accessions under projects BJ/EMBL/GenBank Study Locus Targeted Accessibility: Data U.K. London, (1993) Kew, JH Beaman RS, Beaman JJ, Wood Yungas. Biotropica , 49, 141–152. h lnso on iaau .Orchids. 2. Kinabalu. Mount of plants The p .5.Abeitos L=lwadfrs,L lower = LM forest, lowland = LL Abbreviations: 0.05). < 17 oa oai Gardens, Botanic Royal Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. Borneo - All fungi - Borneo Panama - All fungi All -Panama Yungas - All fungi All - Yungas 18 Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary.

gnmds2 gnmds2 gnmds2 gnmds2 −0.5 0.0 0.5 −0.6−0.4−0.2 0.0 0.2 0.4 0.6 −1.0−0.5 0.0 0.5 1.0 −2−1 0 1 2 −1.51.00.5 0.0 0.5 −1.00.5 0.0 −1.51.00.5 0.0 Archaeorhizomycetales −1.00.5 0.0 Mucorales Sebacinales Trechisporales Leotiomycetes i.s. Leotiomycetes OM N Chaetosphaeriales Russulales Helotiales OMNP Dothideales Rhizophydiales Helotiales Agaricales OM Helotiales Thelephorales OM Pezizomycotina i.s. Pezizomycotina N N Trichosporonales Plant pathogens Plant ECM and root fungi Wood decomposers Wood Chaetothyriales Yungas Yungas Saprotrophs Malasseziales Agaricales Eurotiales gnmds1 gnmds1 gnmds1 gnmds1 Hypocreales Orbiliales Polyporales MAP P Kickxellales MAT pH 0.5 1.0 Hypocreales MAT Agaricales Pleosporales Cantharellales Xylariales 1.0 1.5 2.0 Sporidiobolales Pezizomycotina i.s. Pezizomycotina 0.5 1.0 Agaricales MAP lMAP 0.5 1.0 Onygenales pH Geastrales pH MAT MAP Botryosphaeriales MAT pH Xylariales

gnmds2 gnmds2 gnmds2 gnmds2 −1.5−1.0−0.5 0.0 0.5 1.0 −1.5−1.0−0.5 0.0 0.5 1.0 1.5 −1.0−0.5 0.0 0.5 1.0 −1.5−1.0−0.5 0.0 0.5 1.0 1.5 Ophiostomatales Botryosphaeriales −1.00.5 0.0 −21 0 −21 0 −1.00.5 0.0 Xylariales Xylariales Diaporthales MAT pH MAT Polyporales Thelephorales Agaricales Pezizomycotina i.s. Pezizomycotina Hypocreales MAT Microascales Sporidiobolales Sordariales MAT Sordariomycetidae i.s. Sordariomycetidae pH Sordariales Chaetosphaeriales Polyporales pH Xylariales Cantharellales Cantharellales pH Hymenochaetales Boletales Eurotiales Saccharomycetales Helotiales Hypocreales Russulales Agaricales Ostropales Wood decomposers Wood Geastrales Gomphales Plant pathogens Plant Sebacinales Rhytismatales ECM and root fungi ECM androot 19 Agaricales Borneo Chaetosphaeriales Saprotrophs gnmds1 gnmds1 gnmds1 gnmds1 Trichosporonales OM Ostropales Chaetothyriales Tremellales Chaetothyriales Archaeorhizomycetales Dothideales C/N Dothideomycetidae i.s. Dothideomycetidae Zoopagales C/N Mortierellales 0.5 1.0 0.5 1.0 1 2 Ostropales Helotiales C/N Helotiales C/N Helotiales Pezizales Venturiales Sebacinales Entorrhizales Leucosporidiales 1 2 Cystofilobasidiales Leotiomycetes i.s. Leotiomycetes 1.5

gnmds2 gnmds2 gnmds2 gnmds2 −0.4−0.2 0.0 0.2 0.4 −2−1 0 1 2 −1.0−0.5 0.0 0.5 1.0 −1.0−0.5 0.0 0.5 1.0 −1.00.5 0.0 −42 0 Agaricales Telephorales −1.00.5 0.0 −1.00.5 0.0 Sebacinales OM Russulales P Helotiales Hysteriales OM Cantharellales Helotiales P Helotiales Eurotiales Helotiales Tremellales OM Hysteriales OM Dothideales P Trichosphaeriales Sebacinales ECM and root fungi ECM androot Plant pathogens Plant Wood decomposers Wood P Panama Saprotrophs Mortierellales gnmds1 gnmds1 gnmds1 gnmds1 Pezizales N MAP Pleosporales pH Sordariales Kickxellales Chaetosphaeriales Hypocreales Xylariales 0.5 1.0 0.5 1.0 Venturiales Hypocreales Diaporthales Tubeufiales Saccharomycetales Rhizophydiales pH Xylariales pH 0.5 1.0 2 4 MAT Polyporales MAT MAT MAT 1.5 1.5 Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary.

Richness (OTUs) Richness (OTUs) Richness (OTUs) Richness (OTUs)

2 4 6 8 10 14 0 5 10 15 20 25 10 15 20 25 300 400 500 600 a LL LM UM LL LM UM Yungas LL LM UM LL LM UM a a a b b

4 6 8 10 12 14 0 20 40 60 80 5 10 15 20 300 500 700 Animal pathogens ab Mycoparasites LL LM UM LL LM UM LL LM UM LL LM UM ECM fungi Borneo All fungi a b

0 2 4 6 8 10 0 10 20 30 40 50 4 6 8 10 12 14 250 300 350 400 450 LL LM UM LL LM UM LL LM UM LL LM UM a Panama a a a b a b b b a a a 20

15 25 35 45 150 200 250 0 5 10 20 30 40 50 60 70 80 90 a Yungas LL LM UM LL LM UM LL LM UM LL LM UM a a a a ab b b b

20 40 60 80 100 100 150 200 250 5 10 15 20 25 30 30 50 70 90 Wood decomposers Wood Plant pathogens a a Saprotrophs LL LM UM LL LM UM LL LM UM LL LM UM Borneo Root fungi ab a b b

10 15 20 25 30 100 120 140 160 5 10 15 20 25 30 35 20 30 40 50 60 ab ab LL LM UM LL LM UM LL LM UM LL LM UM a a Panama a a b b a b a b Posted on Authorea 4 Mar 2020 | CC BY 4.0 | https://doi.org/10.22541/au.158333914.43615523 | This a preprint and has not been peer reviewed. Data may be preliminary. communities-in-tropical-forests climate-in-shaping-the-functional-and-taxonomic-composition-of-soil-borne-fungal- sampling-of-neotropical-and-paleotropical-elevation-gradients-reveals-the-role-of- Table1.doc file Hosted vial at available 7612 OTUs Borneo https://authorea.com/users/301763/articles/431622-comparative- 2603 188 Yungas Yungas 192 211 2639 21 Panama 376 1403