Hysterangiales Revisited: Expanded Phylogeny Reveals New Genera and Two New Suborders

VOLUME 8 DECEMBER 2021 Fungal Systematics and Evolution PAGES 65–80

doi.org/10.3114/fuse.2021.08.06

Hysterangiales revisited: expanded phylogeny reveals new genera and two new suborders

N. Davoodian1*, T. Lebel1,2, M.A. Castellano3, K. Hosaka4

1Royal Botanic Gardens Victoria, South Yarra, VIC 3141, Australia 2Botanic Gardens and State Herbarium of South Australia, Adelaide, SA 5001, Australia; Royal Botanic Gardens Victoria, South Yarra, VIC 3141, Australia 3US Department of Agriculture Forest Service, Northern Research Station, Corvallis, OR 97331, USA 4National Museum of Nature and Science, Tsukuba, Ibaraki 305-0005, Japan (ORCID: 0000-0002-4469-8303)

*Corresponding author: [email protected]

Key words: Abstract: Hysterangiales (Phallomycetidae, Agaricomycetes, Basidiomycota) is a diverse, nearly cosmopolitan order atp6 of predominantly hypogeous, sequestrate, ectomycorrhizal fungi. Expanding on previously published phylogenies, we Phallogaster significantly increased sampling of Hysterangiales specimens, emphasizing representatives from Australia. Using protein- phylogenetics coding genes atp6 (adenosine triphosphate synthase subunit 6) and tef1 (translation elongation factor 1-α), we recovered tef1 26 provisional novel genera, and corroborated existing genera and families. Further, two new suborders (Phallogastrineae Trappea subord. nov. and Hysterangineae subord. nov.) and a new family (Phallogastraceae fam. nov.) are described, and three new combinations made to Phallogaster. Aspects of classification and biogeography are presented.

Citation: Davoodian N, Lebel T, Castellano MA, Hosaka K (2021). Hysterangiales revisited: expanded phylogeny reveals new genera and two new suborders. Fungal Systematics and Evolution 8: 65–80. doi: 10.3114/fuse.2021.08.06 Received: 22 January 2021; Accepted: 9July 2021; Effectively published online: 29 July 2021 Corresponding editor: P.W. Crous Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

INTRODUCTION of Hysterangiales in Australasia. Another impetus for the present study is that numerous novel Australasian Hysterangiales taxa Hysterangiales (Phallomycetidae, Agaricomycetes, Basidio- yet to be formally described have been known for over a decade mycota) was established in 2006 to accommodate a but generic boundaries were not strongly supported (Hosaka phylogenetically refined Hysterangiaceae and relatives 2005, Hosaka et al. 2006, 2008). We expanded on previous work (Hosaka et al. 2006). Hysterangiales comprises a wide array initiated by Hosaka et al. (2006, 2008) by sampling extensively of predominantly hypogeous, sequestrate, ectomycorrhizal from Australian specimens, which corroborated known genera fungi, but some taxa in the basal lineage (Phallogastraceae) (both formally described and provisional) and families, and are epigeous and non-mycorrhizal. Hysterangiales possess a elucidated additional provisional novel genera. range of unusual morphologies (Fig. 1), such as basidiomes with a powdery spore mass (and often with a sterile core) in the Mesophelliaceae (Trappe et al. 1996), the gelatinized or MATERIALS AND METHODS cartilaginous gleba with translucent columella in Gallaceaceae and Hysterangiaceae, or the utricle-encased spores of many Collections referred to in this study are housed at various Mesophelliaceae and Hysterangiaceae (Castellano & Beever fungaria including AD, ATH, BRI, OSC, and MEL. Collections were 1994; Fig. 2). Like many truffle-like fungi, Hysterangiales often initially sorted into morphotaxa based on macro- and micro- have distinct odours, which attract small animals for spore characters, and a broad selection of material encompassing dispersal via mycophagy (e.g. Fogel & Trappe 1978, Claridge as many genera as possible were then sampled for molecular & May 1994). The characteristics of being mostly hypogeous analyses. This included some material from non-Australasian and nearly universally dependent on other organisms for taxa that had not previously been sampled, to better support dispersal, combined with their worldwide (excluding Antarctica) previous taxonomic hypotheses. Thin sections of fungal tissue distribution and diversity, have led to the consideration of were mounted in KOH, Melzers’ reagent and occasionally Congo multiple biogeographic scenarios for Hysterangiales, ranging Red + KOH to examine peridium and glebal structure, and spore from ancient Pangaean vicariance to recent long-distance size, shape, presence of a utricle, and ornamentation. Imaging dispersal (Hosaka et al. 2008). was completed using either an Olympus BX-52 microscope with Hysterangiales taxa are remarkably abundant and highly DP-73 camera and measurement tools using Olympus cellSens diverse in Australia (a major impetus for the present study), standard (v. 1.16) or a Zeiss/Moticam 10 camera system. particularly the endemic familyMesophelliaceae . Ectomycorrhizal Adenosine triphosphate synthase subunit 6 (atp6; Myrtaceae and Nothofagaceae species are the primary symbionts mitochondrial) and translation elongation factor 1-α (tef1;

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

Fig. 1. Examples of a range of basidiome morphologies in Hysterangiales. A–C. Phallogastraceae A. "Gen. prov. 1" (AU, photo credit M. Castellano). B. Phallogaster (Trappea) phillipsii (photo credit N. Siegel). C. Phallogaster saccatus (photo credit D. Mitchell/B. Roody); D–F. Gallaceaceae. D. Austrogautieria sp. (NZ, photo credit K. Hosaka). E. Gallacea eburnea (photo credit N. Siegel). F. Gallacea scleroderma (photo credit N. Siegel); G–I. Hysterangiaceae. G. Aroramyces sp. (AU, photo credit M. Castellano). H. Hysterangium sp. (AU, photo credit Castellano). I. Hysterangium sp. (NZ, photo credit K. Hosaka); J–L. Mesophelliaceae. J. Chondrogaster sp. (photo credit M. Castellano). K. "Gen. prov. 3" (photo credit M. Castellano). L. Mesophellia sp. (photo credit D. Catcheside); M–O. Mesophelliaceae. M. Nothocastoreum cretaceum (photo credit D. Catcheside). N. Castoreum radicatum (photo credit M. Castellano). O. Gummiglobus pachythrix (photo credit M. Castellano). Scale bars = 10 mm.

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

Fig. 2. Three spore types observed in Hysterangiales. A. Mesophellia brevispora (photo credit M. Castellano): smooth, no utricle. B. "Genus prov. 7" (photo credit N. Davoodian): ornamented, utricle present. C. Austrogautieria (photo credit T. Lebel): ribbed, no utricle. Scale bars = 10 µm. nuclear) nucleotide sequence data for Hysterangiales available 1, nswapsval: 1, samplefreqval: 1 000, burninfracval: 0.5, on GenBank were downloaded. Further, new atp6 and tef1 vparam.stopval: 0.003, sump_burninfrac: 0.5, sumpnruns: 2, sequences were generated for 99 Hysterangiales specimens, sumt_burninfrac: 0.5, sumtnruns: 2, sumtntrees: 1. The average as well as one specimen of Phallales and one specimen of potential scale reduction factor reported for all parameters Cantharellales to use as outgroup (Table 1). For these newly was 1.038. The alignment and resultant consensus phylogram sequenced specimens, DNA was extracted from dried herbarium is deposited in TreeBASE under study number 27527. To material with the E.Z.N.A. Forensic DNA Kit (Omega Bio-tek). A corroborate our topology and support values, we conducted portion of the mitochondrial protein-coding gene atp6 was PCR additional separate analyses using different outgroup taxa amplified with ATP6-3 and ATP6-2 primers (Kretzer & Bruns (Geastrales), extended generations (ranging from 22 000 000 1999) using the following cycling protocol: 95 °C for 5 min; 8 to 59 000 000), different models of rate variation among sites, cycles of 94 °C for 35 s, 37 °C for 55 s, 72 °C for 60 s; 35 cycles and different partitions i.e. one partition, two partition (each of 72 °C for 60 s, 94 °C for 35 s, 45 °C for 55 s; 72 °C for 2 min. gene), or multi-partition (codon positions for each gene, as A portion of the nuclear protein-coding gene tef1 was PCR well as introns for tef1) analyses. We also conducted maximum amplified with EF1-983F and EF1-1567R, or EF1-1953R, primers likelihood (ML) analysis using RAxML software (Stamatakis et (Rehner & Buckley 2005) using the following cycling protocol: al. 2008) via the CIPRES REST API with the following options; 95 °C for 5 min; 10 cycles of 94 °C for 30 s, 65 °C for 60 s, 72 °C for dna_gtrcat: GTRGAMMA, select_analysis: fa, choose_bootstrap: 90 s; 30 cycles of 94 °C for 45 s, 55 °C for 60 s, 72 °C for 70 s; 72 °C x, bootstrap_value: 1 000. Taxon names at individual terminals for 5 min. Sanger sequencing was conducted in forward and in the phylogram are field-based, verbatim identifications, from reverse directions at the Australian Genome Research Facility various mycologists. (Melbourne, Victoria, Australia). Sequences are deposited in GenBank (Table 1). Each gene was aligned using MAFFT online (Katoh et al. RESULTS 2017), visually inspected, and manually adjusted as required. Phylogenetic analyses of each individual gene were conducted The BI analysis recovered Hysterangiales (Bayesian posterior using the methods outlined in the next paragraph. Dendrograms probability, bpp = 1) with the four constituent families from these analyses were then compared using PHYLO.IO Gallaceaceae, Hysterangiaceae, Mesophelliaceae, and (Robinson et al. 2016) to assess concordance. The topologies Phallogastraceae fam. nov. (all bpp = 1), containing numerous between each gene were found to be almost completely described and undescribed genera (Supplementary Fig. S1, Fig. concordant at the generic level and below, corroborating all 3 - Parts 1,2,3). In total, 26 provisional undescribed genera were genera and species (with the exception of specimen H381 inferred (25 Hysterangiales and 1 Phallales). The results of the which appeared in different positions). Also, many relationships additional BI analyses and ML analysis were not substantively between genera were corroborated between atp6 and tef1. different from our primary BI analysis, corroborating all taxa In the individual gene analyses, higher taxonomic levels (e.g. indicated in Fig. 3 and Table 1. Phallogastraceae is accommodated family, suborder) were either not recovered or recovered with by a newly established suborder, Phallogastrineae (described low support. below, Fig. 3 - Part 1), and the families Mesophelliaceae (Fig. 3 After the assessment of the individual gene phylogenies, - Part 1), Hysterangiaceae (Fig. 3 - Part 2) and Gallaceae (Fig. 3 - atp6 and tef1 were concatenated, resulting in an alignment Part 3) are accommodated in the new suborder Hysterangineae. of 1 464 sites long with 223 terminals. MrBayesS v. 3.2.7a Trappeaceae is not in Hysterangiales, since the type species of (Ronquist et al. 2012) was used via the CIPRES REST API (Miller Trappea, T. darkeri, is inferred to be in Phallales based on this and et al. 2015) for Bayesian inference (BI) analysis. An analysis using previous analyses (Hosaka et al. 2006, Sulzbacher et al. 2016; Fig. default priors and the following parameters was run for 20 000 3 - Part 3, Table 1). Neither Trappea pinyonensis nor T. phillipsii 000 generations; nstopts: 6, nucmodelopts: 4by4, rateopts: belong in Trappea given their position withinPhallogastraceae (Xu gamma, nrunsval: 2, nchainsval: 4, tempval: 0.200, swapfreqval: & Luo 2003, Hosaka et al. 2006; Fig. 3 - Part 1); the combination to

Phallogaster is made here. It should be noted that true Protubera Notes: Phallogastraceae Locq. (1974) is invalid in accordance lies outside of Hysterangiales, since the type (P. maracuja) and with Art. 39:1 of the Shenzen code. This is the only clade in related species belong in Protophallaceae, Phallales (Trierveiler- Hysterangiales with an apparently non-mycorrhizal habit. Pereira et al. 2014). Thus, Protubera hautuensis, P. nothofagi, and Although the number of taxa is thus far very limited, the all other taxa appearing in Fig. 3 - Part 3) labelled as Protubera are phylogenetic relationships within the family/suborder strongly not true members of the genus (this includes P. canescens, which indicate reciprocal monophyly of the Northern Hemisphere is a synonym of Ileodictyon gracile in Phallaceae, Phallales). (“Phallogaster clade”) and Southern Hemisphere clades (“Gen.

prov. 1”) (Fig. 3 - Part 1). Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] Taxonomy In addition to a traditional morphology-based description for the corresponding suborder (Phallogastrineae subord. nov.), Order Hysterangiales K. Hosaka & Castellano we have also applied a phylogenetic taxon definition for the suborder (see Hibbett et al. 2018). More taxon sampling in the Phallogastrineae Castellano, T. Lebel, Davoodian & K. Hosaka, future may require further amendment of the morphological subord. nov. MycoBank MB 838485. definition of the suborder, whereas our phylogenetic taxon definition of Phallogastrinae will remain stable. Basidiomes sequestrate, small (0.2 cm wide) to larger (up to 5 cm long and 3 cm wide), subglobose to irregularly subovoid, with Phallogaster pinyonensis (States) K. Hosaka, Castellano, a tapered to irregularly shaped base, single or caespitose, with Davoodian & T. Lebel, comb. nov. MycoBank MB 838487. white rhizomorphs (rhizomorphs sometimes discolouring pinkish). Basionym: Trappea pinyonensis States, Mycotaxon 41: 128. Basidiome surface smooth to velvety to finely tomentose, white 1991. MycoBank MB 129849. to clay white (discolouring pinkish to salmon buff to reddish or faint lilac). Peridium 160–2 000 µm thick. Columella branched, Typus: USA, Arizona, Walnut Canyon National Monument, whitish to semi-translucent, gelatinized. Gleba varies from green, Coconino County, on sandy soil under Pinus edulis, 10 Oct. 1986, olivaceous green, greyish olive to pale olive; gelatinized, loculate, J. States AHF-530 (holotype ASC). locules small. Basidia clavate to narrowly clavate to irregularly cylindrical, thin-walled, hyaline, 6–8-spored. Sterigmata small Phallogaster phillipsii (Harkn.) K. Hosaka, Castellano, Davoodian to indistinct. Spores smooth, elongate-ellipsoid to oblong, thin- & T. Lebel, comb. nov. MycoBank MB 840085. walled, utricle absent, hyaline singly, olive buff to honey yellow or Basionym: Hysterangium phillipsii Harkn., Proceedings of the pale green in mass. Clamp connections present. California Academy of Sciences 1(8): 255. 1899. MycoBank MB 187085. Phylogenetic taxon definition of the suborder: The largest crown Synonym: Trappea phillipsii (Harkn.) Castellano, Mycotaxon 38: clade containing Phallogaster saccatus Morgan 1893, but not 7. 1990. MycoBank MB 127551. Hysterangium clathroides Vittad. 1831, Mesophellia arenaria Berk. 1857, and Gallacea scleroderma (Cooke) Lloyd 1905. This Typus: USA, California, Wire Bridge, Placer County, in soil, is a maximum crown-clade definition (Hibbett et al. 2018). associated with Quercus, Jan. 1899, C.L. Phillips (Harkness 234, holotype BPI) Type family: Phallogastraceae Castellano, T. Lebel, Davoodian & K. Hosaka non Phallogastraceae Locq., De Taxia Fungorum: I. Notes: Although the species was not included in the phylogenetic Syllabus 1A: 56. 1974. analysis in this study, it has clearly been demonstrated to be the sister taxon to Phallogaster (Trappea) pinyonensis by Hosaka et Phallogastraceae Castellano, T. Lebel, Davoodian & K. Hosaka, al. (2006). Its basidiome surfaces stain pinkish when bruised, fam. nov. MycoBank MB 838484. which is also a common characteristic with other species in the genus Phallogaster. Basidiomes sequestrate, small (0.2 cm wide) to larger (up to 5 cm long and 3 cm wide), subglobose to irregularly subovoid, with Excluded taxa a tapered to irregularly shaped base, single or caespitose, with white rhizomorphs (rhizomorphs sometimes discolouring pinkish). Trappea cinnamomea A-S. Xu & D.Q. Luo, Mycosystema 22: 192. Basidiome surface smooth to velvety to finely tomentose, white 2003. MycoBank MB 489004. to clay white (discolouring pinkish to salmon buff to reddish or faint lilac). Peridium 160–2 000 µm thick. Columella branched, Notes: Based on the original description of T. cinnamomea (Xu whitish to semi-translucent, gelatinized. Gleba varies from green, & Luo 2003), its basidiome surfaces have the staining reaction olivaceous green, greyish olive to pale olive; gelatinized, loculate, to pink, which is similar to Trappea phillipsii and T. pinyonensis. locules small. Basidia clavate to narrowly clavate to irregularly However, we so far have no access to the specimens and no cylindrical, thin-walled, hyaline, 6–8-spored. Sterigmata small DNA data from the species are currently available. Until more to indistinct. Spores smooth, elongate-ellipsoid to oblong, thin- definitive evidence becomes available, we keep this taxon in the walled, utricle absent, hyaline singly, olive buff to honey yellow or genus Trappea. pale green in mass. Clamp connections present. Odour sometimes fetid. Species apparently saprotrophic, occurring on decaying Hysterangineae Castellano, T. Lebel, Davoodian & K. Hosaka, wood or leaves, or partially buried in litter. subord. nov. MycoBank MB 838486.

Type genus: Phallogaster Morgan, J. Cincinnati Soc. Nat. Hist. 15: Basidiomes sequestrate, hypogeous or epigeous, and can be 171. 1893. small (0.1 cm wide) to larger (up to 10 cm wide), subglobose to

100/1 Trappea pinyonensis WNA AHF530 100/1 Phallogaster saccatus ENA T13202 Phallogaster 100/1 Protubera hautuensis NZ OSC59673 Phallogastraceae Protubera sp. AU T20068* gen. prov. 1 A 100/1 Hysterangium sp. AU:VIC AWC42 Aroramyces sp. AU:VIC H6547 fam. nov. 75/ 0.95 gen. prov. 2 Aroramyces sp. AU:NSW MEL2364203 Chondrogaster sp. AU:WA H170 100/1 Chondrogaster sp. AU:WA MEL2105021 Chondrogaster sp. AU:VIC MEL2137016 99/1 Chondrogaster sp. AU:NSW TL449 Chondrogaster sp. AU:NSW TL1487 Chondrogaster sp. AU:NSW TL1426 90/1 Chondrogaster sp. AU:NSW MEL2238556 58/95 99/1 Chondrogaster sp. AU:QLD H4362 Chondrogaster s.str. 100/1 Chondrogaster angustisporus BRZ OSC62041 HOLO

Chondrogaster angustisporus BRZ OSC62042 Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 100/1 E-mail: [email protected] Chondrogaster sp. WNA T11067 * 100/1 Chondrogaster pachysporus AU OSC49298 100/1 Chondrogaster sp. CHN T17536 99/1 Chondrogaster sp. AU:QLD SEA10 * Andebbia pachythrix AU:NSW MEL2124354 100/1 Andebbia pachythrix AU:NSW MEL2238554 Andebbia Andebbia pachythrix AU:NSW OSC58809 100/1 Castoreum sp. AU:NSW MEL2238626 * 99/1 Castoreum sp. AU:NSW T18910 gen. prov. 26 100/1 37/ 0.67 Castoreum camphoratum AU:NSW MEL2063469 100/1 Castoreum camphoratum AU:NSW MEL2238635 gen. prov. 3 Castoreum camphoratum AU:NSW MEL2063470 99/1 Castoreum sp. AU:SA MEL2389702 32/ 0.67 Mesophellia clelandii AU:VIC MEL2042631 47/ 0.91 Mesophellia sp. AU:QLD SEA35B 44/ 0.89 Mesophellia sabulosa AU:VIC OSC55918 Malajczukia amicorum AU:VIC OSC59295 87/1 Malajczukia amicorum AU:TAS MEL2056858 Mesophellia glauca AU:TAS OSC56986 gen. prov. 4 99/1 95/1 Mesophellia brevispora AU:WA MEL2281255 Mesophellia sp. AU:QLD SEA37 Mesophellia sp. AU:SA MEL2389696 95/1 Mesophellia trabalis AU:VIC OSC59282 100/1 Malajczukia ingratissima AU:VIC OSC59296 64/0.9 Mesophellia clelandii AU:VIC OSC59292 Mesophellia arenaria AU:TAS OSC59306 Mesophellia s.str. 99/1 Nothocastoreum cretaceum AU:WA MEL2356006 100/1 Nothocastoreum cretaceum AU* OSC79832 Nothocastoreum Nothocastoreum cretaceum AU OSC79925 100/1 Chondrogaster sp. AU:QLD SEA6B * 100/1 Chondrogaster sp. AU: NSW TL1294 100/1 Chondrogaster sp. AU:VIC T19877 gen. prov. 5 Chondrogaster sp. AU:NSW TL1097 Mesophelliaceae Gummiglobus sp. AU:WA H7679 gen. prov. 6 22/0.54 Castoreum tasmanicum AU:VIC MEL2041148 85/1 Castoreum tasmanicum AU:VIC MEL2056682 100/0.99Castoreum radicatum AU:QLD MEL2354501 100/1 Castoreum sp. AU:QLD MEL2354440 99/1 Castoreum sp. AU:QLD SEA2 Castoreum s.str. Castoreum sp. AU:QLD SEA3 54/ 0.97 87/0.99 100/1 Mesophellia arenaria AU:TAS MEL2356530 Castoreum radicatum AU:NSW MEL2364205 100/1 Castoreum sp AU OSC122814 Gummiglobus agglutinosporus AU:VIC MEL2042096* Gummiglobus sp. AU:QLD SEA22 33/0.6 100/1 Gummiglobus agglutinosporus AU OSC58784 Gummiglobus joyceae AU:QLD MEL2314623 Gummiglobus Gummiglobus joyceae AU OSC59485 100/1 Gummiglobus agglutinosporus AU:VIC MEL2105039* 76/ 1 100/1 Gummivena potorooi AU:WA MEL2104390 Gummivena potorooi AU:WA MEL2314967 Gummivena Gummivena potorooi AU:WA MEL2105001 100/1 Chondrogaster sp. AU:QLD SEA9 * Aroramyces sp. AU:NSW TL1701 gen. prov. 7 Hysterangium inflatum AU H4035 91/1 Hysterangium sp. AU:NSW MEL2057692 91/1Hysterangium sp. PNG H5573 58/0.85 59/0.98Hysterangium sp. AU:QLD SEA27 Hysterangium sp. USA:CAL T36459 97/1Chondrogaster sp. AU:VIC H6586 94/1 Hysterangium sp. AU:ACT T16823 32/0.63Hysterangium sp. AU:VIC H6761 Chondrogaster sp. AU:SA TL2871 Hysterangium sp. AU:SA PSC4372 gen. prov. 8 100/1 Hysterangium sp. AU:WA T14749 Hysterangium sp. AU T6923 100/1 90/1 Chondrogaster sp. AU:NSW MEL2364057 100/1 Chondrogaster sp. AU:NSW MEL2364304 43/0.62 Chondrogaster sp. AU:SA TL2888 100/1 70/1 Hysterangium sp. AU:NT T15362 Hysterangium sp. AU:NT T15375 100/1 Hysterangium gardnerii AU T6950 100/1 Chondrogaster sp. AU:SA TL2889 Hysterangium sp. AU:TAS H381 gen. prov. 9 99/1 Hysterangium sp. AU H2022 100/1 Hysterangium sp. AU H4262 Hysterangium aggregatum AU:VIC MEL2049882 99/1 Chondrogaster sp. AU:QLD MEL2305115 gen. prov. 10 100/0.92 Hysterangium sp. AU:QLD T22535 Chondrogaster sp. AU:QLD SEA7B Hysterangium sp. AU:QLD SEA270 B 99/1

Hysterangiaceae 0.2

Fig. 3. Part 1. Phylogram from Bayesian analysis showing families and genera within Hysterangiales. Node A indicates Phallogastrineae subord. nov.; node B indicates Hysterangineae subord. nov. Maximum likelihood (ML) bootstrap values / Bayesian posterior probabilities (bpp) are shown at the nodes. The scale bar shows substitutions per site. Where bpp ≥ 0.95 and ML bootstrap ≥ 70%, branches are thickened. Type species indicated by a blue asterisk *. The asterisk in parentheses (*) denotes Ileodictyon, the currently accepted genus for Protubera canescens. Australian (AU) States and Territories indicated by: WA (Western Australia), NT (Northern Territory), QLD (Queensland), NSW (New South Wales), VIC (Victoria), TAS (Tasmania), SA (South Australia). United States (USA) states indicated by: ARIZ (Arizona), CAL (California), COL (Colorado), ID (Idaho), OR (Oregon). Other geographic areas indicated as follows: NZ (New Zealand), NewCal (New Caledonia), PNG (Papua New Guinea), BRZ (Brazil), MEX (Mexico), WNA (western North America), ENA (eastern North America), CHN (China), Guyana (Guyana), SSA (southern South America), NSA (northern South America), ASIA (Asia), SEASIA (southeast Asia), SING (Singapore), THAI (Thailand), IND (India), EUR (Europe), COSTA (Costa Rica), ZIMB (Zimbabwe).

Mesophelliaceae

100/1 99/1 Hysterangium sp. AU:TAS H6404 47/ 0.52 Hysterangium sp. AU H4749 98/1 Hysterangium sp. AU:WA T6889 90/1 Hysterangium sp. AU:WA T6965

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 100/1 Hysterangium sp. AU OSC122836 E-mail: [email protected] 72/ 0.95 100/1 Hysterangium sp. NZ OSC122721

100/1 Hysterangium sp. NZ T15545 91/1 Hysterangium sp. AU H5057 100/1 Hysterangium salmonaceum AU Beaton33 43/ 0.54 Hysterangium sp. AU:TAS MEL2078287 gen. prov. 11 55/ 0.97 Hysterangium sp. AU H6105 94/1 99/1 Hysterangium sp. PNG OSC122483 86/1 Hysterangium sp. NZ OSC59629 Hysterangium sp. AU H4123 Hysterangium sp. SSA Halling5741 100/1 95/ 0.99 Hysterangium sp. SSA T26347 35/ 0.68 Hysterangium spegazzinii SSA Singer3426 92/1 Hysterangium sp. SSA T26367 Hysterangium fragile ? Kers3971 Hysterangium epiroticum EUR T6116 93/ 0.99Hysterangium sp. WNA OSC122857 100/1 34/ 0.91 Hysterangium seperabile WNA OSC69030 67/ 0.89 Hysterangium youngii NZ OSC59645 gen. prov. 12 100/1 Hysterangium sp. WNA T17856 81/1 Hysterangium sp ASIA OSC122860 Hysterangium sp. ENA T591 100/1 Hysterangium sp. MEX T3296 99/1 Hysterangium sp. WNA T4794 gen. prov. 13 89/1 Hysterangium strobilus ENA T5285 91/1 Hysterangium rugisporum NZ OSC59662 B 43/ 0.98 Hysterangium rupticutis NZ OSC59667 97/ 0.98 Hysterangium sp. AU:SA H6973 gen. prov. 14 100/1 Aroramyces sp. AU:SA MEL2320943 96/1 100/1 Aroramyces sp. AU:SA MEL2320959 Hysterangium sp. AU:SA T16994 100/1 95/1 Hysterangium sp. USA:ARIZ AHF602 97/1 Hysterangium sp. MEX T19263 Hysterangiaceae Hysterangium pompholyx EUR Gross495 gen. prov. 15 Hysterangium calcareum EUR Gross97 99/1 Aroramyces gelatinosporus AU H4010 100/1 Aroramyces sp. NCAL OSC122590 95/1 * 39/ 0.86 Aroramyces sp. IND T9930 87/1 Aroramyces sp. NSA SM10030 65/1 Aroramyces radiatus ZIMB Verbeken99062 Aroramyces 79/ 0.99 99/1 Aroramyces sp. SEASIA T15013 35/ 0.91 Aroramyces sp. ASIA OSC122858 Hysterangium occidentale WNA OSC47048 gen. prov. 16 99/1 Hysterangium sp. GUYA SM10007

51/ 0.66 Hysterangium sp. GUYA SM10100 70/ 0.9 Hysterangium sp. GUYA SM10166 100/1 Hysterangium sp BRAZ UFRN2112 97/1 Hysterangium sp. BRAZ UFRN2115 48/ 0.78 gen. prov. 17 100/1 Hysterangium sp. THAI T13345

46/ 0.96 80/ 0.95 Hysterangium sp. SING T17501 54/ 0.66 100/1 Hysterangium sp. AU:QLD SEA29 Hysterangium sp. AU:QLD SEA30 100/1 Hysterangium sp. NZ PDD82853 Hysterangium affine AU:WA T6884 gen. prov. 18 Hysterangium sp. MEX T3328 gen. prov. 19 Hysterangium sp. MEX Garcia3779 49/ 0.7 100/1 100/1 100/1Hysterangium crassum OSC110447 41/ 0.7 Hysterangium cistophilum EUR T1088 100/1 Hysterangium clathroides EUR Zemere 100/1 Hysterangium sp. ASIA OSC122859 Hysterangium s.str. Hysterangium setchellii WNA OSC58071 * 91/1 Hysterangium album ENA T15139 100/1 Hysterangium sp. WNA T8997 Hysterangium membranaceum EUR T12836 83/1 Hysterangium coriaceum EUR Kers4984 80/ 0.96 Hysterangium coriaceum ENA OSC55265 87/ 0.99 Hysterangium sp. WNA T22832 61/ 0.84 90/ 0.99 Hysterangium coriaceum WNA OSC64939 gen. prov. 20 100/1 Hysterangium aureum WNA OSC56988 Hysterangium crassirhachis WNA OSC58056 Hysterangium sp. COSTA T27921

Gallaceaceae 0.2

Fig. 3. (Continued) - Part 2. irregularly subovoid without tapered base, single or gregarious, elastic, glutinous or hard and brittle, 1–4-layered, sometimes often covered with adhering sand and soil or encased in debris incorporating mycorrhizae. Columella branched, whitish to and rootlets, usually with conspicuous white to off-white semi-translucent and gelatinized or a soft to corky or rubbery rhizomorphs at base or along sides of basidiomata. Basidiome central core or lacking such structures. Gleba varies from green, surface smooth to velvety to finely tomentose, often white to olivaceous green, greyish olive, brown to ochraceous or pinkish; off-white in colour and often discolouring pinkish, reddish, gelatinous to cartilaginous or with a powdery spore mass at yellowish or brownish, but purple to violet in some species. maturity, often with labyrinthine to elongated to circular locules. Peridium adherent to often readily separable from gleba, Basidia clavate to narrowly clavate to irregularly cylindrical,

76/1

100/1 Mesophelliaceae 99/1

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

96/1 Hysterangiaceae

90/1 89/1

85/1 Austrogautieria sp. AU Beaton66 B 100/1 Austrogautieria clelandii AU:VIC OSC80012 100/1 Austrogautieria clelandii AU:VIC OSC62178 gen. prov. 24 86/ 0.99 Austrogautieria sp. NZ OSC122637 100/1 Hysterangium sp. AU :TAS H1199 92/1 Hysterangium sp. AU:VIC T19862 Hallingea Hallingea purpurea SSA Garido418A 55/ 0.98 Gallacea sp. NZ Beever718 44/ 0.88 Gallacea sp. AU OSC122813* 94/1 Mesophellia sp. AU:TAS HO130165 96/1 100/1 Hysterangium sp. AU:TAS HO130161 99/1 Austrogautieria sp. AU:QLD H4096 Gallacea sp. AU:VIC MEL2053228 gen. prov. 21 100/1 99/1 65/0.97 Chondrogaster sp. AU:VIC MEL2059688 64/ 0.92 Gallacea subalpina AU:NSW MEL2236389 Gallacea subalpina AU:NSW OSC80855 96/1 Chondrogaster sp. AU:QLD H4263 91/1 87/1 Hysterangium sp. NZ Beever536 Gallacea sp. NZ OSC122728 100/1 Hysterangium sp. NZ KHNZ04117 Gallaceaceae 88/1 Gallacea sp. NZ REB2364 gen. prov. 22 100/1 Gallacea sp. NZ KHNZ04123 100/1 Gallacea eburnea NZ OSC59601 Protubera nothofagi NZ OSC59699 100/1 Hysterangium sp. AU:TAS H5277 Hysterangium sp. AU Beaton 80/1 95/1 Hysterangium sp. AU:QLD SEA28 42/ 0.68 Hysterangium sp. AU:QLD T13462 gen. prov. 23 80/ 0.97 Austrogautieria sp. AU:NSW T24466 99/1 Gallacea dingleya NZ OSC59606 100/1 Gallacea sp. NCALT25038 100/1 Austrogautieria chlorospora AU:SA MEL2320844 94/1 Austrogautieria chlorospora AU:SA OSC46596 99/1 Austrogautieria manjimupana AU OSC59545 94/1 Austrogautieria manjimupana AU OSC55900 Austrogautieria s.str. 41/ 0.56 98/1 Austrogautieria costata AU:SA MEL2320846 Austrogautieria sp. AU OSC80139 73/ 0.96 Austrogautieria sp. AU OSC80140 Gallacea scleroderma NZ OSC59621 Gallacea s.str. 98/1 Aroramyces sp. AU:WA MEL2236309* gen. prov. 25 Trappea darkeri USA:OR OSC65085 Trappea s.str. 100/1 Protubera canescens AU:WA MEL2063471 Protubera canescens AU:VIC MEL2105035* Ileodictyon (*)

0.2 Fig. 3. (Continued) - Part 3. thin-walled, hyaline, mostly 4-spored. Spores ellipsoid to Notes: The Hysterangineae is exclusively composed of oblong to fusoid, smooth to minutely verrucose or with long sequestrate taxa which are ectomycorrhizal with various spines beneath a utricle, or sometimes with longitudinal ridges, vascular plants insofar as inferred from available studies. Most thin- to thick-walled, often covered with a wrinkled to inflated species in Hysterangineae have rather specific ectomycorrhizal or ephemeral utricle, hyaline, pale green, or brown in KOH, hosts, usually restricted to a single plant family. For example, inamyloid, sometimes weakly dextrinoid. Odour faint, fetid, or species distributed in the Southern Hemisphere are associated sometimes sweet. Clamp connections present or absent. All with Myrtaceae (mostly with Eucalyptus spp. and/or Kunzea/ species are presumably ectomycorrhizal with various vascular Leptospermum spp.) or Nothofagaceae (Castellano & Beever plants including Dipterocarpaceae, Fagaceae, Myrtaceae, 1994, Hosaka et al. 2008). Nothofagaceae, and Pinaceae. This suborder is much more speciose than Phallogastrineae Phylogenetic taxon definition of the suborder: The largest and many more new species, especially from Australia, remain crown clade containing Hysterangium clathroides Vittad. 1831, to be described for all three families, i.e., Hysterangiaceae, but not Phallogaster saccatus Morgan 1893. This is a maximum Mesophelliaceae and Gallaceaceae. We believe that the crown-clade definition (Hibbett et al. 2018). family-level phylogeny is robust (Fig. 3, Supplementary Fig. S1), but more taxon sampling in the future may require further Type family: Hysterangiaceae E. Fisch., in Engler & Prantl, Nat. amendment of the morphological definition of the suborder. Pflanzenfam., Teil. I (Leipzig): 304. 1899. [1900] Therefore, as is the case for Phallogastrineae, we have also applied a phylogenetic taxon definition approach as summarized by Hibbett et al. (2018).

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Phallogastrineae ( Hysterangiales Phallogastrineae ( Hysterangiales Phallogastrineae ( Hysterangiales Phallogastrineae ( Hysterangiales Hysterangineae ( Phallales ( Phallales ( Phallales ( Phallales ) ( Geastrales ) ( Geastrales ) ( Geastrales ) ( Geastrales ) ( Geastrales ) ( Geastrales ( Cantharellales Suborder (Order) Suborder Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Phallogastraceae Phallogastraceae Phallogastraceae Phallogastraceae Gallaceaceae Clathraceae Clathraceae Trappeaceae Trappeaceae Geastraceae Geastraceae Geastraceae Geastraceae Geastraceae Geastraceae Cantharellaceae Family MW264846 DQ219144 MW302308 MW302290 MW302306 MW302309 MW302289 MW302288 DQ219145 DQ219127 DQ219126 DQ219132 DQ219217 DQ219220 DQ219218 DQ219133 DQ219221 DQ219284 DQ219283 DQ219292 MW302302 DQ219234 DQ219230 DQ219229 DQ219228 DQ219227 DQ219226 MW302303 TEF1 MW048612 DQ218829 MW449882 MW048615 MW048613 MW417403 MW048610 MW449866 n/a DQ218817 DQ218816 DQ218819 DQ218882 DQ218883 DQ218801 DQ218820 DQ218884 DQ218932 DQ218931 DQ218938 MW438321 DQ218896 DQ218770 DQ218889 DQ218888 DQ218769 DQ218887 MT828543 ATP6 VIC AUS NZ QLD VIC QLD VIC TAS SSA VIC VIC NZ ENA AUS NZ AUS WNA VIC WA USA - OR USA WA — USA - ID USA VIC NSW USA - CO USA USA - OR USA SA Geography MEL2053228 OSC122813 Beever718 H4263 MEL2059688 H4096 T19862 H1199 Garido418A OSC80012 OSC62178 OSC122637 T13202 T20068 OSC59673 Beaton66 AHF530 MEL2105035 MEL2063471 OSC65085 MEL2236309 OSC58979 OSC41996 MEL2096557 MEL2087743 OSC29328 OSC60730 TL2893 Collector or Collector Herbarium ID ] ] sp. sp. sp. sp. sp. sp. sp. purpurea clelandii clelandii sp. saccatus saccatus sp. hautuensis sp. pinyonensis canescens [ gracile canescens canescens [ gracile canescens darkeri sp. fuscogleba recolligens pectinatum fornicatum floriforme fimbriatum sp. Species verbatim Species verbatim determination ] ] [ Ileodictyon [ Ileodictyon Gallacea Gallacea Gallacea Chondrogaster Chondrogaster Austrogautieria indet. Hysterangium Hallingea Austrogautieria Austrogautieria Austrogautieria Phallogaster Protubera Protubera Austrogautieria Trappea Protubera Protubera Trappea Aroramyces Radiigera Geastrum Geastrum Geastrum Geastrum Geastrum Cantharellus Cantharellus Genus verbatim Genus verbatim determination ) . bold

21 Hosaka Gen. prov. Hallingea Gen. prov. 1 Hosaka Gen. prov. & Hosaka 24 Castellano Gen. prov. Phallogaster Ileodictyon Trappea Trappea 25 Lebel & Davoodian Gen. prov. Geastrum ( Radiigera Geastrum

Geastrum Cantharellaceae Tabulation of specimens and Clade sequences names Fig. used from are 3. in1. Tabulation Generic thisstudy. names withoutdetermination) verbatim (genus a species name determination) (species verbatim Table (Western WA by: indicated Territories and States Australian (AUS) based micromorphologically. are were examined on collectionsfield identification; with species determination) namesverbatim (species in (USA) states bold. indicated are States United areas Extra-Australian SA (South Australia). (Tasmania), QLD (Queensland), VIC NSW (New (Victoria), TAS South Wales), NT (Northern Territory), Australia), BRZ Guinea), New (Papua PNG Caledonia), (New NEWCAL Zealand), (New NZ follows: as indicated areas geographic Other (Oregon). OR (Idaho), ID (Colorado), CO (California), CA (Arizona), AZ by: indicated (Brazil), MEX (Mexico), WNA (western North America), ENA (eastern North America), CHN (China), (Guyana), GUYANA SSA (southern South America), NSA (northern South America), ASIA (Asia), SEASIA (southeast Asia), SING THAI (Singapore), (Thailand), IND (India), (Costa Rica), ZIMB EUR (Zimbabwe). (Europe), COSTA Novel DNA sequences for this generated study are indicated by GenBank accessions in Clade

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae Suborder (Order) Suborder Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Gallaceaceae Family DQ219196 DQ219205 DQ219204 DQ219158 MW302282 DQ219139 DQ219129 DQ219128 MW302296 DQ219125 MW302295 DQ219131 DQ219130 DQ219174 MW302298 MW302297 MW302294 DQ219137 DQ219141 MW302299 DQ219219 MW302292 MW302291 DQ219138 MW302293 DQ219140 DQ219143 MW302305 MW302304 DQ219142 MW302307 TEF1 DQ218865 DQ218874 DQ218873 DQ218781 MW417406 AY574787 DQ218762 DQ218818 MW438327 DQ218761 MW048618 DQ218764 DQ218763 DQ218790 MW048617 MW438329 MW048616 DQ218824 DQ218826 MW438326 AY574786 MW438330 MW417402 DQ218766 MW449865 DQ218825 DQ218828 MW048611 MW438331 DQ218827 MW048614 ATP6 AUS SSA SSA SSA TAS NZ AUS AUS SA AUS SA AUS AUS AUS QLD QLD TAS NZ NEWCAL NSW NZ NZ NZ NZ NZ NZ NZ TAS TAS NSW NSW Geography OSC122836 T26367 T26347 Halling5741 H6404 OSC59621 OSC59545 OSC55900 MEL2320846 OSC46596 MEL2320844 OSC80140 OSC80139 KandGBeaton T13462 SEA28 H5277 OSC59606 T25038 T24466 OSC59699 KHNZ04117 Beever536 OSC59601 KHNZ04123 REB2364 OSC122728 HO130165 HO130161 OSC80855 MEL2236389 Collector or Collector Herbarium ID sp. sp. sp. sp. sp. scleroderma manjimupana manjimupana costata chlorospora chlorospora sp. sp. sp. sp. sp. sp. dingleyae sp. sp. nothofagi sp. sp. eburnea sp. sp. sp. sp. sp. subalpina subalpina Species verbatim Species verbatim determination Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Gallacea Austrogautieria Austrogautieria Austrogautieria Austrogautieria Austrogautieria Austrogautieria Austrogautieria Hysterangium Hysterangium Hysterangium Hysterangium Gallacea Gallacea Austrogauteria Protubera Hysterangium Hysterangium Gallacea Gallacea Gallacea Gallacea Mesophellia Hysterangium Gallacea Gallacea Genus verbatim Genus verbatim determination (Continued). (Continued).

11 Hosaka Gen. prov. 11 Hosaka Gen. prov. 11 Hosaka Gen. prov. Gallacea Gallacea

Austrogautieria

23 Hosaka Gen. prov.

22 Castellano Gen. prov.

Table 1. Table Clade

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Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae Suborder (Order) Suborder Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Family DQ219185 MW302285 MW302284 n/a DQ219164 MW302287 MW302286 DQ219168 DQ219177 DQ219172 DQ219169 DQ219166 DQ219156 DQ219155 DQ219170 DQ219188 DQ219191 DQ219182 n/a DQ219165 DQ219189 MW302283 DQ219184 DQ219197 DQ219198 DQ219173 DQ219195 DQ219193 DQ219192 DQ219176 DQ219161 TEF1 DQ218854 MW218373 MW218372 DQ218838 DQ218784 MW218375 MW218374 DQ218788 DQ218846 DQ218842 DQ218789 DQ218786 DQ218780 DQ218779 DQ218840 DQ218857 DQ218860 DQ218851 DQ218787 DQ218785 DQ218858 MW438322 DQ218853 DQ218866 DQ218867 DQ218843 DQ218864 DQ218862 DQ218861 DQ218845 DQ218837 ATP6 USA - AZ USA VIC SA NZ NZ SA SA ENA WNA MEX NZ WNA — EUR ENA WNA ASIA WNA SSA AUS WA WA NZ NZ PNG TAS AUS AUS AUS AUS NZ Geography AHF602 T16994 H6973 OSC59667 OSC59662 MEL2320959 MEL2320943 T5285 T4794 T3296 OSC59645 OSC69030 Kers3971 T6116 T591 T17856 OSC122860 OSC122857 Singer3426 Beaton33 T6889 T6965 OSC59629 OSC122721 OSC122483 MEL2078287 H6105 H5057 H4749 H4123 T15545 Collector or Collector Herbarium ID sp. sp. sp. rupticutis rugisporum sp. sp. strobilus sp. sp. youngii separabile fragile epiroticum sp. sp. sp. sp. spegazzinii salmonaceum ” “ rhodocarpum “irregular” sp. sp. sp. sp. sp. sp. sp. sp. sp. Species verbatim Species verbatim determination Hysterangium Hysterogaster Hysterogaster Hysterangium Hysterangium Aroramyces Aroramyces Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Genus verbatim Genus verbatim determination (Continued). (Continued). 15 Castellano Gen. prov.

14 Lebel Gen. prov.

13 Castellano Gen. prov.

12 Castellano Gen. prov.

Table 1. Table Clade

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae Suborder (Order) Suborder Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Family DQ219160 DQ219154 DQ219151 DQ219150 DQ219147 DQ219207 DQ219190 DQ219178 DQ219183 n/a DQ219199 LT635646 LT635645 DQ219171 DQ219203 DQ219202 DQ219201 DQ219200 MW302301 MW302300 DQ219162 DQ219121 DQ219120 DQ219124 DQ219122 DQ219123 DQ219119 DQ219118 DQ219163 DQ219149 DQ219180 TEF1 DQ218782 AY574827 DQ218832 DQ218777 DQ218774 DQ218876 DQ218859 DQ218847 DQ218852 DQ218831 DQ218868 LT635648 LT635647 DQ218841 DQ218872 DQ218871 DQ218870 DQ218869 MW438323 MW438328 AY574825 DQ218812 DQ218811 DQ218815 DQ218813 DQ218814 DQ218810 DQ218809 DQ218783 DQ218776 DQ218849 ATP6 EUR — EUR EUR ENA WNA ASIA MEX MEX AUS AUS NZ BRZ BRZ SING THAI GUYANA GUYANA GUYANA QLD QLD WNA IND SEASIA NSA ASIA NEWCAL ZIMB AUS EUR EUR MEX Geography T12836 OSC110447 Zemere T1088 T15139 T8997 OSC122859 Garcia3779 T3328 T6884 PDD82853 UFRN2115 UFRN2112 T17501 T13345 SM10166 SM10100 SM10007 SEA30 SEA29 OSC47048 T9930 T15013 SM10030 OSC122858 OSC122590 Verbeken99062 H4010 Gross495 Gross97 T19263 Collector or Collector Herbarium ID membranaceum crassum clathroides cistophilum album sp. sp. sp. sp. affine sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. occidentale sp. sp. sp. sp. sp. radiatus gelatinosporus pompholyx calcareum sp. Species verbatim Species verbatim determination Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Aroramyces Aroramyces Aroramyces Aroramyces Aroramyces Aroramyces Aroramyces Hysterangium Hysterangium Hysterangium Genus verbatim Genus verbatim determination (Continued). (Continued).

Hysterangium 19 Castellano Gen. prov.

18 Castellano & Lebel 18 Castellano Gen. prov.

17 Lebel & Castellano Gen. prov. Gen. prov. 16 Lebel & Castellano Gen. prov.

Aroramyces

Table 1. Table Clade

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae Suborder (Order) Suborder Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Hysterangiaceae Family MW302241 MW302230 MW302229 DQ219117 MW302228 MW302227 DQ219136 MW302222 DQ219135 MW302216 MW302226 MW302223 MW302225 MW302224 MW302221 MW302220 MW302219 MW302218 MW302217 MW302215 MW302212 MW302214 MW302213 DQ219153 DQ219152 n/a n/a DQ219148 DQ219206 DQ219181 DQ219167 TEF1 MW417408 MW417409 MW449869 DQ218808 MW449880 MW449879 DQ218823 MW449876 DQ218822 MW417407 MW417405 MW438324 MW449868 MW449874 MW417413 MW417414 MW417415 MW417412 MW449881 MW438325 MW417411 MW417417 MW417410 DQ218778 AY574826 DQ218834 DQ218833 DQ218775 DQ218875 DQ218850 DQ218839 ATP6 SA NSW NSW NSW NSW NSW AUS BRZ BRZ WA NSW WNA QLD CHN NSW NSW NSW VIC QLD WA VIC NSW VIC WNA WNA ENA EUR WNA COSTA WNA WNA Geography MEL2389702 T18910 MEL2238626 OSC58809 MEL2238554 MEL2124354 OSC49298 OSC62042 OSC62041 MEL2105021 MEL2238556 T11067 SEA10 T17536 TL1426 TL1487 TL449 MEL2137016 H4362 H170 AWC42 MEL2364203 H6547 OSC58056 OSC64939 OSC55265 Kers4984 OSC56988 T27921 T22832 OSC58071 Collector or Collector Herbarium ID sp. sp. sp. pachythrix pachythrix pachythrix pachysporus angustisporus angustisporus angustisporus sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. crassirhachis coriaceum coriaceum coriaceum aureum sp. sp. setchellii Species verbatim Species verbatim determination Castoreum Castoreum Castoreum Andebbia Andebbia Andebbia Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Hysterangium Aroramyces Aroramyces Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Genus verbatim Genus verbatim determination (Continued). (Continued). 3 Castellano Gen. prov.

26 Castellano Gen. prov.

Andebbia

Chondrogaster

2 Lebel Gen. prov.

20 Castellano Gen. prov.

Table 1. Table Clade

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae Suborder (Order) Suborder Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Family MW302248 MW302243 MW302250 MW302244 DQ219134 MW302247 MW302246 MW302242 MW302276 MW302275 MW302274 MW302273 DQ219216 DQ219215 MW302240 DQ219210 DQ219214 DQ219213 DQ219212 DQ219211 MW302234 MW302236 MW302239 MW302238 MW302237 DQ219209 DQ219208 MW302235 MW302233 MW302232 MW302231 TEF1 MW246683 MW246695 MW246690 MW246672 DQ218821 MW246668 MW246674 MW449873 MW246685 MW246684 MW246677 MW246665 n/a DQ218881 MW246680 DQ218877 DQ218880 DQ218879 DQ218878 DQ218795 MW449878 MW449872 MW438333 MW449870 MW449875 DQ218793 DQ218792 MW449867 MW449877 MW417416 MW417404 ATP6 VIC VIC NSW QLD AUS QLD QLD WA NSW VIC NSW QLD AUS AUS WA TAS VIC VIC TAS VIC VIC WA QLD SA QLD VIC VIC TAS NSW NSW NSW Geography MEL2056682 MEL2041148 MEL2364205 MEL2354501 OSC122814 SEA3 SEA2 H7679 TL1097 T19877 TL1294 SEA6B OSC79925 OSC79832 MEL2356006 OSC59306 OSC59282 OSC55918 OSC56986 OSC59292 MEL2042631 MEL2281255 SEA35B MEL2389696 SEA37 OSC59296 OSC59295 MEL2056858 MEL2063470 MEL2238635 MEL2063469 Collector or Collector Herbarium ID tasmanicum tasmanicum radicatum radicatum sp. sp. sp. sp. sp. sp. sp. sp. cretaceum cretaceum cretaceum arenaria trabalis sabulosa glauca clelandii clelandii brevispora sp. sp. sp. ingratissima amicorum amicorum camphoratum camphoratum camphoratum camphoratum camphoratum camphoratum Species verbatim Species verbatim determination Castoreum Castoreum Castoreum Castoreum Castoreum Castoreum Castoreum Gummiglobus Chondrogaster Chondrogaster Chondrogaster Chondrogaster Nothocastoreum Nothocastoreum Nothocastoreum Mesophellia Mesophelllia Mesophellia Mesophellia Mesophellia Mesophellia Mesophellia Mesophellia Mesophellia Mesophellia Malajczukia Malajczukia Malajczukia Castoreum Castoreum Castoreum Genus verbatim Genus verbatim determination (Continued). (Continued).

Castoreum 6 Lebel & Davoodian Gen. prov.

5 Lebel Gen. prov.

Nothocastoreum Mesophellia

4 Castellano & Lebel 4 Castellano Gen. prov.

Table 1. Table Clade

) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) ) )

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected]

( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae Suborder (Order) Suborder Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Family DQ219179 MW302272 MW302271 DQ219186 MW302266 MW302264 MW302265 MW302263 MW302261 MW302259 MW302258 MW302270 MW302269 MW302268 MW302267 MW302262 MW302260 MW264847 MT947438 MW302257 MW302256 MW302255 n/a MW302253 MW302254 MW302251 n/a MW302252 MW302249 MW302245 TEF1 DQ218848 MW246676 MW246675 DQ218855 MW246698 MW246669 MW246681 MW246666 MW246679 MW246694 MW246686 MW246687 MW246689 MW246697 MW246671 MW246692 MW246682 MT947436 MT947437 MW246673 MW246696 MW246678 DQ218772 MW438332 MW246670 MW246693 DQ218830 MW246691 MW246688 MW246667 ATP6 NSW NT NT AUS WA PNG QLD VIC SA USA - CA USA ACT SA SA NSW NSW SA VIC QLD NSW WA WA WA AUS QLD VIC VIC AUS QLD TAS QLD Geography MEL2057692 T15375 T15362 T6923 T14749 H5573 SEA27 H6761 PSC4372 T36459 T16823 TL2889 TL2888 MEL2364304 MEL2364057 TL2871 H6586 SEA9 TL1701 MEL2105001 MEL2314967 MEL2104390 OSC59485 MEL2314623 MEL2105039 MEL2042096 OSC58784 SEA22 MEL2356530 MEL2354440 Collector or Collector Herbarium ID sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. sp. potorooi potorooi potorooi joyceae joyceae agglutinosporus agglutinosporus agglutinosporus sp. arenaria sp. Species verbatim Species verbatim determination Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Chondrogaster Aroramyces Gummivena Gummivena Gummivena Gummiglobus Gummiglobus Gummiglobus Gummiglobus Gummiglobus Gummiglobus Mesophellia Fusicastoreum Genus verbatim Genus verbatim determination (Continued). (Continued).

8 Hosaka Gen. prov.

7 Davoodian, Lebel & 7 Davoodian, Gen. prov. Castellano

Gummivena

Gummiglobus

Table 1. Table Clade

) ) ) ) ) ) ) ) ) ) DISCUSSION

In addition to revealing new taxa, our analysis highlights prominent biogeographic patterns in Hysterangiales. Trans-Tasman distributions are numerous throughout Hysterangiales; almost every New Zealand lineage inferred in our phylogram is sister to an Australian lineage, at various taxonomic scales. Gondwanan disjunctions are also represented, for example "Gen. prov. 17" (Fig. 3 - Part 2) contains an Australian and

Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] paleotropical clade sister to a northeastern South American clade. Some human-mediated introduction events are apparent as well e.g. ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae ( Hysterangiales Hysterangineae Suborder (Order) Suborder Chondrogaster angustisporus (OSC62041 - holotype) is from planted Eucalyptus in Brazil (Giachini et al. 2000). Hosaka et al. (2008) postulated that the Hysterangiales originated in Australia or Eastern Gondwana and expanded in range over time, possibly through long-distance dispersal. Sheedy et al. (2016) inferred that Hysterangiales was the first group of Agaricomycetes to become sequestrate in Australia (ca. 83 million Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Mesophelliaceae Family years ago). The rate of sequestration of Agaricomycetes was inferred to have increased in Australia after separation from Antarctica, the timing of which overlaps with the radiation of potential mycorrhizal plant associates as well as the emergence of specialized mycophagous DQ219175 MW302279 MW302278 DQ219146 DQ219187 MW302280 MW302277 MW302281 DQ219159 DQ219157 TEF1 marsupials. Although periods of aridification were evidently not the sole driver of sequestration, they likely had a major influence on the diversity of sequestrate fungi in Australia, including Hysterangiales (Sheedy et al. 2016). Our analysis indicates phylogenetic signals for some prominent DQ218844 MW449863 MW449862 DQ218773 DQ218856 MW449864 MW449861 MW449871 DQ218836 DQ218835 ATP6 macroscopic features. In the Mesophelliaceae, the clade unifying Gummivena, Gummiglobus, and Castoreum is comprised of species that have gummy-like mycelium and/or outer glebal tissue (Fig. 1N, O). Species in the clade containing Chondrogaster, Andebbia, Mesophellia, VIC QLD QLD AUS AUS QLD QLD TAS AUS AUS Geography Nothocastoreum, and related provisional genera have a powdery spore mass at maturity and also incorporate roots and soil in a multi-layered mycelial crust surrounding the basidiome (in Nothocastoreum this falls off easily and remains in the soil) (Fig. 1J–M). In provisional genera 8 through 10, the basidiomes are typically encased in a mycelial soil crust, and the MEL2049882 SEA270 T22535 H4262 H2022 SEA7B MEL2305115 H381 H4035 T6950 Collector or Collector Herbarium ID glebal tissue is gelatinized. Macroscopic characters of Hysterangiaceae overlap somewhat with various clades in the Mesophelliaceae and Gallaceaceae, including having a gelatinized gleba and varying degrees of rhizomorph development across the surface of basidiomes (Fig. 1G– I). In Gallaceaceae, most lineages have a firm, cartilaginous peridium surrounding a gelatinized gleba (Fig. 1D–F), and spores lacking an utricle (Fig. 2A). The condition of having ridged spores (Fig. 2C) is not aggregatum sp. sp. sp. sp. sp. sp. sp. inflatum gardnerii Species verbatim Species verbatim determination monophyletic in Hysterangiales; this character was previously thought to be diagnostic for the genus Austrogautieria (Stewart & Trappe 1985) but now appears in two well-supported, distinct clades in Gallaceaceae (Austrogautieria and "Gen. prov. 24"; Fig. 3 - Part 3). Numerous provisional genera and species have been revealed in this study, which will be described by the respective authors of this paper and other collaborators in upcoming publications. We believe our current approach of recognising numerous provisional genera is a more accurate Hysterangium Hysterangium Hysterangium Hysterangium Hysterangium Chondrogaster Chondrogaster Hysterangium Hysterangium Hysterangium Genus verbatim Genus verbatim determination and conservative estimate of diversity than to lump them into fewer, larger genera. In addition to strong phylogenetic signals, each provisional genus is readily distinguishable by macroscopic and microscopic characters, but species level differentiation within genera requires more sampling in many of these taxa. Moreover, our phylogenetic analyses show that some existing genera, such as Castoreum and Gummiglobus in Mesophelliaceae, are closely related to each other within their respective families, but our provisional genera are more distantly related to these existing genera (Fig. 3, Supplementary Fig. S1). More sampling

(Continued). (Continued). throughout Australasia and South America will likely uncover additional clades and aid in further refining generic and specific boundaries.

10 Hosaka Gen. prov. 9 Davoodian Gen. prov.

Table 1. Table Clade

ACKNOWLEDGEMENTS Kretzer AM, Bruns TD (1999). Use of atp6 in fungal phylogenetics: an example from the Boletales. Molecular Phylogenetics and Evolution A special thanks to Dr S. Abell (James Cook University and Australian 13: 483–492. Tropical Herbarium) for access to Hysterangiales collections and early Miller MA, Schwartz T, Pickett BE,et al. (2015). A RESTful API for access collaboration on this diverse lineage. This project was completed with to phylogenetic tools via the CIPRES science gateway. Evolutionary funding and support from the Australian Biological Resources Study (grant Bioinformatics 11: EBO–S21501. RF217-63: Truffle-like fungi [Basidiomycetes]: taxonomy, systematics and Rehner SA, Buckley E (2005). A Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: evidence for cryptic diversification a revised classification for all Australian taxa), BioPlatforms Australia, Editor-in-Chief Prof. dr P.W. Crous, Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] the Friends of the Botanic Gardens Victoria, and the Royal Botanic and links to Cordyceps teleomorphs. Mycologia 97: 84–98. Gardens Victoria. Dr M. Castellano was also supported in part by the Robinson O, Dylus D, Dessimoz C (2016). Phylo.io: interactive viewing ABRS grant RF217. Dr M. Smith at the University of Florida is thanked and comparison of large phylogenetic trees on the web. Molecular for helpful comments to this manuscript. Much appreciation also to the Biology and Evolution 33: 2163–2166. curation staff at the National Herbarium of Victoria (MEL), Queensland Ronquist F, Teslenko M, Van Der Mark P, et al. (2012). MrBayes 3.2: State Herbarium (BRI), and State Herbarium of South Australia (AD) efficient Bayesian phylogenetic inference and model choice across for assistance with loans and processing of material. Assistance with a large model space. Systematic Biology 61: 539–542. processing material and sequencing was provided by technical staff at Sheedy EM, Martin AF, Ryberg M,et al. (2016). Dating the emergence of MEL: R. Jordan, Dr M. Hirst and A. Boxshall. This study was financed in truffle-like fungi in Australia, by using an augmented meta-analysis. part by JSPS KAKENHI Grants 24680085 and 20K06805 to Dr K. Hosaka. Australian Systematic Botany 29: 284–302. Stamatakis A, Hoover P, Rougemont J (2008). A rapid bootstrap Conflict of interest: The authors declare that there is no conflict algorithm for the RAxML web servers. Systematic Biology 57: 758– of interest. 771. Stewart EL, Trappe JM (1985). The new genus Austrogautieria (Basidiomycotina), segregate from Gautieria. Mycologia 77: 674– REFERENCES 687. Sulzbacher MA, Grebenc T, Cabral TS, et al. (2016). Restingomyces, a Castellano MA, Beever RE (1994). Truffle-like Basidiomycotina of New new sequestrate genus from the Brazilian Atlantic rainforest that Zealand: Gallacea, Hysterangium, Phallobata, and Protubera. New is phylogenetically related to early-diverging taxa in Trappeaceae Zealand Journal of Botany 32: 305–328. (Phallales). Mycologia 108: 954–966. Claridge AW, May TW (1994). Mycophagy among Australian mammals. Trappe JM, Castellano MA, Malajczuk N (1996). Australasia truffle- Australian Journal of Ecology 19: 251–275. like fungi. VII. Mesophellia (Basidiomycotina, Mesophelliaceae). Fogel R, Trappe JM (1978). Fungus consumption (mycophagy) by small Australian Systematic Botany 9: 773–802. animals. Northwest Science 52: 1–31. Trierveiler-Pereira L, Meijer AA, Hosaka K, et al. (2014). Updates on Giachini A, Castellano MA, Trappe JM (2000). Ectomycorrhizal fungi in Protubera (Protophallaceae, Phallales) and additional notes on P. Eucalyptus and Pinus plantations in southern Brazil. Mycologia 92: maracuja. 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66/0.99 Cantharellus sp A:SA TL2893 53/0.99 Geastrum fornicatum AU:NSW MEL2087743 66/1 Geastrum floriforme USA:COL OSC29328 Geastrum fimbriatum USA:OR OSC60730 Outgroup 100/1 100/1 Geastrum recolligens USA:ID OSC41996 Geastrum Radiigera fuscogleba OSC58979 Geastrum pectinatum AU:VIC MEL2096557 100/1 Trappea pinyonensis WNA AHF530 100/1 Phallogaster saccatus ENA T13202 Phallogaster 100/1 Protubera hautuensis NZ OSC59673 Protubera sp. AU* T20068 gen. prov. 1 Phallogastraceae 100/1 Hysterangium sp. AU:VIC AWC42 Aroramyces sp. AU:VIC H6547 75/0.95 gen. prov. 2 A Aroramyces sp. AU:NSW MEL2364203 Chondrogaster sp. AU:WA H170 fam. nov. 100/1 Chondrogaster sp. AU:WA MEL2105021 Chondrogaster sp. AU:VIC MEL2137016 99/1 Chondrogaster sp. AU:NSW TL449 Chondrogaster sp. AU:NSW TL1487

90/1 Chondrogaster sp. AU:NSW TL1426 Chondrogaster sp. AU:NSW MEL2238556 99/1 56/0.95 Chondrogaster sp. AU:QLD H4362 Chondrogaster s.str. 100/1 Chondrogaster angustisporus BRZ OSC62041 HOLO Chondrogaster angustisporus BRZ OSC62042 100/1 Chondrogaster sp. WNA T11067 * 100/1 Chondrogaster pachysporus AU OSC49298 100/1 Chondrogaster sp. CHN T17536 99/1 Chondrogaster sp. AU:QLD SEA10 * 100/1 100/0.69 Andebbia pachythrix AU:NSW MEL2124354 Andebbia pachythrix AU:NSW MEL2238554 Andebbia pachythrix AU:NSW OSC58809 Andebbia 100/1 Castoreum sp. AU:NSW MEL2238626 * 99/1 Castoreum sp. AU:NSW T18910 100/1 gen. prov. 26 37/0.67 Castoreum camphoratum AU:NSW MEL2063469 100/1 Castoreum camphoratum AU:NSW MEL2238635 Castoreum camphoratum AU:NSW MEL2063470 gen. prov. 3 Castoreum sp. AU:SA MEL2389702 99/1 32/0.67 Mesophellia clelandii AU:VIC MEL2042631 47/0.91 Mesophellia sp. AU:QLD SEA35B 44/0.89 Mesophellia sabulosa AU:VIC OSC55918

87/1 Malajczukia amicorum AU:VIC OSC59295 Malajczukia amicorum AU:TAS MEL2056858 95/1 Mesophellia glauca AU:TAS OSC56986 gen. prov. 4 99/1 Mesophellia brevispora AU:WA MEL2281255 Mesophellia sp. AU:QLD SEA37

95/1 Mesophellia sp. AU:SA MEL2389696 Mesophellia trabalis AU:VIC OSC59282 100/1 Malajczukia ingratissima AU:VIC OSC59296 64/0.9 Mesophellia clelandii AU:VIC OSC59292 Mesophellia arenaria AU:TAS OSC59306 Mesophellia s.str. 99/1 Nothocastoreum cretaceum AU:WA MEL2356006 100/1 Nothocastoreum cretaceum* AU OSC79832 Nothocastoreum cretaceum AU OSC79925 Nothocastoreum 100/1 Chondrogaster sp. AU:QLD SEA6B * 100/1 Chondrogaster sp. AU: NSW TL1294 Mesophelliaceae 100/1 Chondrogaster sp. AU:VIC T19877 gen. prov. 5 Chondrogaster sp. AU:NSW TL1097 Gummiglobus sp. AU:WA H7679 22/0.54 gen. prov. 6 85/1 Castoreum tasmanicum AU:VIC MEL2041148 100/0.99 Castoreum tasmanicum AU:VIC MEL2056682 Castoreum radicatum AU:QLD MEL2354501 100/1 Castoreum sp. AU:QLD MEL2354440 1 Castoreum sp. AU:QLD SEA2 99/1 87/0.99 Castoreum s.str. Castoreum sp. AU:QLD SEA3 54/0.97 100/1 Mesophellia arenaria AU:TAS MEL2356530 100/1Castoreum radicatum AU:NSW MEL2364205 Castoreum sp. AU OSC122814 Gummiglobus agglutinosporus AU:VIC MEL2042096* Gummiglobus sp. AU:QLD SEA22 33/0.6 100/1 Gummiglobus agglutinosporus AU OSC58784 64/0.81 Gummiglobus joyceae AU:QLD MEL2314623 Gummiglobus Gummiglobus joyceae AU OSC59485 Gummiglobus agglutinosporus AU:VIC MEL2105039* 76/1 100/1 100/1 Gummivena potorooi AU:WA MEL2104390 Gummivena potorooi AU:WA MEL2314967 Gummivena Gummivena potorooi AU:WA MEL2105001 100/1 Chondrogaster sp. AU:QLD SEA9* Aroramyces sp. AU:NSW TL1701 gen. prov. 7 91/1 Hysterangium inflatum AU H4035 Hysterangium sp. AU:NSW MEL2057692 Hysterangium sp. PNG H5573 100/1 91/1 58/0.85 59/0.98 Hysterangium sp. AU:QLD SEA27 59/1Hysterangium sp. USA:CAL T36459 Chondrogaster sp. AU:VIC H6586 97/1 94/1 Hysterangium sp. AU:ACT T16823 32/0.63Hysterangium sp. AU:VIC H6761 Chondrogaster sp. AU:SA TL2871 Hysterangium sp. AU:SA PSC4372 gen. prov. 8 100/1 Hysterangium sp. AU:WA T14749 Hysterangium sp. AU T6923 90/1 100/1 Chondrogaster sp. AU:NSW MEL2364057 100/1 Chondrogaster sp. AU:NSW MEL2364304 43/0.62 Chondrogaster sp. AU:SA TL2888 100/1 70/1 Hysterangium sp. AU:NT T15362 Hysterangium sp. AU:NT T15375 100/1 Hysterangium gardnerii AU T6950 Chondrogaster sp. AU:SA TL2889 Hysterangium sp. AU:TAS H381 gen. prov. 9 99/1 Hysterangium sp. AU H2022 100/1 Hysterangium sp. AU H4262 99/1 Hysterangium aggregatum AU:VIC MEL2049882 Chondrogaster sp. AU:QLD MEL2305115 100/0.92 gen. prov. 10 1 Hysterangium sp. AU:QLD T22535 Chondrogaster sp. AU:QLD SEA7B Hysterangium sp. AU:QLD SEA270 99/1 100/1 Hysterangium sp. AU:TAS H6404 47/0.52 98/1 Hysterangium sp. AU H4749 90/1 Hysterangium sp. AU:WA T6889 Hysterangium sp. AU:WA T6965 72/0.95 100/1 Hysterangium sp. AU OSC122836 100/1 Hysterangium sp. NZ OSC122721 Hysterangium sp. NZ T15545 91/1 Hysterangium sp. AU H5057 100/1 Hysterangium salmonaceum AU:VIC Beaton33 43/0.54 Hysterangium sp. AU:TAS MEL2078287 55/0.97 gen. prov. 11 Hysterangium sp. AU H6105 94/1 99/1 86/1 Hysterangium sp. PNG OSC122483 Hysterangium sp. NZ OSC59629 Hysterangium sp. AU H4123 Hysterangium sp. SSA Halling5741 100/1 95/0.99 35/0.68 Hysterangium sp. SSA T26347 Hysterangium spegazzinii SSA Singer3426 92/1 Hysterangium sp. SSA T26367 Hysterangium fragile Kers3971 Hysterangium epiroticum EUR T6116 93/0.99 100/1 Hysterangium sp. WNA OSC122857 34/0.91 Hysterangium seperabile WNA OSC69030 67/0.89 Hysterangium youngii NZ OSC59645 gen. prov. 12 100/1 Hysterangium sp. WNA T17856 81/1 Hysterangium sp. ASIA OSC122860 Hysterangium sp. ENA T591 100/1 99/1 Hysterangium sp. MEX T3296 Hysterangium sp. WNA T4794 Hysterangium strobilus ENA T5285 gen. prov. 13 91/1 Hysterangium rugisporum NZ OSC59662 43/0.98 Hysterangium rupticutis NZ OSC59667 97//0.98 Hysterangium sp. AU:SA H6973 100/1 gen. prov. 14 96/1 Aroramyces sp. AU:SA MEL2320943 100/1 Aroramyces sp. AU:SA MEL2320959 Hysterangium sp. AU:SA T16994 95/1 100/1 Hysterangium sp. USA:ARIZ AHF602 97/1 Hysterangium sp. MEX T19263 Hysterangiaceae Hysterangium pompholyx EUR Gross495 gen. prov. 15 Hysterangium calcareum EUR Gross97 99/1 100/1 Aroramyces gelatinosporus AU H4010 95/1 Aroramyces sp. NCAL OSC122590 87/1 Aroramyces sp. IND T9930 * 29/0.86 65/1 Aroramyces sp. NSA SM10030 Aroramyces radiatus ZIMB Verbeken99062 Aroramyces s.str. 90/1 99/1 Aroramyces sp. SEASIA T15013 35/0.91 Aroramyces sp. ASIA OSC122858 79/0.99 Hysterangium occidentale WNA OSC47048 99/1 Hysterangium sp. GUYA SM10007 gen. prov. 16 89/1 70/0.9 Hysterangium sp. GUYA SM10100 51/0.66 Hysterangium sp. GUYA SM10166 100/1 Hysterangium sp. BRAZ UFRN2112 97/1 Hysterangium sp. BRAZ UFRN2115 48/0.78 80/0.95 100/1 Hysterangium sp. THAI T13345 gen. prov. 17 46/0.96 Hysterangium sp. SING T17501 B 100/1 54/0.66 Hysterangium sp. AU:QLD SEA29 Hysterangium sp. AU:QLD SEA30 100/1 Hysterangium sp. NZ PDD82853 Hysterangium affine AU:WA T6884 gen. prov. 18 Hysterangium sp. MEX T3328 Hysterangium sp. MEX Garcia3779 gen. prov. 19 100/1 100/1 49/0.7 100/1 Hysterangium crassum OSC110447 41/0.7 Hysterangium cistophilum EUR T1088 100/1 Hysterangium clathroides EUR Zemere 100/1 Hysterangium sp. ASIA OSC122859 Hysterangium setchellii WNA OSC58071 * Hysterangium s.str. 91/1 Hysterangium album ENA T15139 100/1 Hysterangium sp. WNA T8997 Hysterangium membranaceum EUR T12836 83/1 Hysterangium coriaceum EUR Kers4984 80/0.96 87/0.99 Hysterangium coriaceum ENA OSC55265 Hysterangium sp. WNA T22832 61/0.84 Hysterangium coriaceum WNA OSC64939 90/0.99 gen. prov. 20 100/1 Hysterangium aureum WNA OSC56988 Hysterangium crassirhachis WNA OSC58056 Hysterangium sp. COSTA T27921 85/1 100/1 Austrogautieria sp. AU:VIC Beaton66 100/1 Austrogautieria clelandii AU:VIC OSC80012 Austrogautieria clelandii AU:VIC OSC62178 gen. prov. 24 0.99 Austrogautieria sp. NZ OSC122637 100/1 Hysterangium sp. AU:TAS H1199 92/1 65/0.97 Hysterangium sp. AU:VIC T19862 Hallingea Hallingea purpurea SSA Garido418A 55/0.98 Gallacea sp. NZ* Beever718 44/0.88 94/1 Gallacea sp. AU OSC122813 Mesophellia sp. AU:TAS HO130165 96/1 100/1 Hysterangium sp. AU:TAS HO130161 99/1 Austrogautieria sp. AU:QLD H4096 100/1 Gallacea sp. AU:VIC MEL2053228 gen. prov. 21 Chondrogaster sp. AU:VIC MEL2059688 99/1 Gallacea subalpina AU:NSW MEL2236389 64/0.92 Gallacea subalpina AU:NSW OSC80855 96/1 Chondrogaster sp. AU:QLD H4263 87/1 Hysterangium sp. NZ Beever536 52/0.55 Gallacea sp. NZ OSC122728 100/1 Gallaceaceae Hysterangium sp. NZ KHNZ04117 88/1 Gallacea sp. NZ REB2364 100/1 Gallacea sp. NZ KHNZ04123 gen. prov. 22 91/1 100/1 Gallacea eburnea NZ OSC59601 Protubera nothofagi NZ OSC59699 100/1 Hysterangium sp. AU:TAS H5277 Hysterangium sp. AU:VIC Beaton 80/1 95/1 Hysterangium sp. AU:QLD SEA28 42/0.68 80/0.97 Hysterangium sp. AU:QLD T13462 gen. prov. 23 99/1 Austrogautieria sp. AU:NSW T24466 Gallacea dingleya NZ OSC59606 100/1 Gallacea sp. NCAL T25038 100/1 94/1 Austrogautieria chlorospora AU:SA MEL2320844 Austrogautieria chlorospora AU:SA OSC46596 99/1 94/1 Austrogautieria manjimupana AU OSC59545 Austrogautieria manjimupana AU OSC55900 98/1 Austrogautieria s.str. 41/0.56 Austrogautieria costata AU:SA MEL2320846 Austrogautieria sp. AU OSC80139 73/0.96 Austrogautieria sp. AU OSC80140 Gallacea scleroderma NZ OSC59621 Gallacea s.str. 98/1 Aroramyces sp. AU:WA MEL2236309 Trappea darkeri USA:OR OSC65085 * gen. prov. 25 100/1 Protubera canescens AU:WA MEL2063471 Trappea Protubera canescens AU:VIC MEL2105035 *Ileodictyon (*)

0.2

Supplementary Fig. S1. Overview Phylogram from Bayesian analysis showing relationship between Hysterangiales, some Phallales and outgroups (OG). Node A indicates Phallogastrineae subord. nov.; node B indicates Hysterangineae subord. nov. Maximum likelihood (ML) bootstrap values / Bayesian posterior probabilities (bpp) are shown at the nodes. Where bpp ≥ 0.95 and ML bootstrap ≥ 70 %, branches are thickened. Type species indicated by a blue asterisk *. The asterisk in parentheses (*) denotes the currently accepted genus for Protubera canescens, Ileodictyon. Countries and States indicated with same acronyms as Fig. 3.