Published May 4, 2026 | Version v1
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Revisiting infrageneric concepts in Geastrum (Geastraceae, Geastrales) with three new species

  • 1. Ludong University, Yantai, China
  • 2. Northeast Forestry University, Harbin, China
  • 3. National and Local Joint Engineering Research Center for Breeding & Cultivation of Features Edible Fungi, Fuzhou, China|Fujian Academy of Agricultural Sciences, Fuzhou, China

Description

Geastrum is a typical saprotrophic genus with nearly 130 species distributed worldwide, yet its infrageneric classification has traditionally relied on morphological characteristics whose phylogenetic significance remains uncertain. Here, we revisit infrageneric concepts in Geastrum using Chinese material, integrating detailed morphological observations with multi-locus phylogenetic analyses. Eight specimens collected from temperate forests in Northeast and East China yielded three new species: G. capillitiononramosum sp. nov., G. villopannosum sp. nov., and G. kunyuense sp. nov. These species share a sessile endoperidial body lacking a collar and apophysis, a distinctly delimited fibrillose peristome, and globose to subglobose basidiospores with columnar ornamentation but differ mainly in the structure of the mycelial layer and the branching and ornamentation of capillitial hyphae. Phylogenetic analyses based on a concatenated ITS–nrLSU–rpb1 dataset recovered 12 well-supported clades and confirmed the distinct phylogenetic positions of the three new species. The resulting topology further demonstrates that several traditional infrageneric groupings defined by key morphological traits, such as the hygroscopic behavior of the exoperidium and debris-encrusted mycelial layers, are only partially congruent with molecular relationships, reflecting repeated character evolution across distant lineages. Nevertheless, specific combinations of peristome structure, mycelial layer features, and capillitial morphology remain informative at the species level. With the descriptions of three new species and an updated key to 43 Chinese taxa, this study refines current knowledge of Geastrum diversity in China and provides new evidence for reassessing infrageneric classification within the genus.

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References

  • Accioly T, Sousa JO, Moreau PA, Lécuru C, Silva BDB, Roy M, Gardes M, Baseia IG, Martín MP (2019) Hidden fungal diversity from the Neotropics: Geastrum hirsutum, G. schweinitzii (Basidiomycota, Geastrales) and their allies. PLoS ONE 14(2): e0211388. https://doi.org/10.1371/journal.pone.0211388
  • Bandala VM, Montoya L (2000) A taxonomic revision of some American Crepidotus. Mycologia 92: 341–353. https://doi.org/10.1080/00275514.2000.12061165
  • Baseia IG, Milanez AI (2002) Geastrum setiferum (Gasteromycetes): A new species with a setose endoperidium. Mycotaxon 84: 135–139. https://doi.org/10.5962/p.418292
  • Bottomley AM (1948) Gasteromycetes of South Afica. Bothalia 4(3): 473–642. https://doi.org/10.4102/abc.v4i3.1859
  • Cabral TS, Sousa JO, Silva BDB, Martín MP, Clement CR, Baseia IG (2017) A remarkable new species of Geastrum with an elongated branched stipe. Mycoscience 58(5): 344–350. https://doi.org/10.1016/j.myc.2017.03.004
  • Caffot ML, Amarilla LD, Crespo EM, Domínguez LS (2016) Geastrum minutisporum sp. nov. from Polylepis australis woodlands in central Argentina. Mycoscience 57(3): 157–163. https://doi.org/10.1016/j.myc.2015.08.004
  • Coker WC (1924) The geasters of the United States and Canada. Journal of Elisha Mitchell Scientific Society 39(3/4): 170–224. http://www.jstor.org/stable/24330732
  • Crous PW, Luangsa-ard JJ, Wingfield MJ, Carnegie AJ, Hernández-Restrepo M, Lombard L, Roux J, Barreto RW, Baseia IG, Cano-Lira JF, Martín MP, Morozova OV, Stchigel AM, Summerell BA, Brandrud TE, Dima B, García D, Giraldo A, Guarro J, Gusmão LFP, Khamsuntorn P, Noordeloos ME, Nuankaew S, Pinruan U, Rodríguez-Andrade E, Souza-Motta CM, Thangavel R, Van Iperen AL, Abreu VP, Accioly T, Alves JL, Andrade JP, Bahram M, Baral H-O, Barbier E, Barnes CW, Bendiksen E, Bernard E, Bezerra JDP, Bezerra JL, Bizio E, Blair JE, Bulyonkova TM, Cabral TS, Caiafa MV, Cantillo T, Colmán AA, Conceição LB, Cruz S, Cunha AOB, Darveaux BA, Da Silva AL, Da Silva GA, Da Silva GM, Da Silva RMF, De Oliveira RJV, Oliveira RL, De Souza JT, Dueñas M, Evans HC, Epifani F, Felipe MTC, Fernández-López J, Ferreira BW, Figueiredo CN, Filippova NV, Flores JA, Gené J, Ghorbani G, Gibertoni TB, Glushakova AM, Healy R, Huhndorf SM, Iturrieta-González I, Javan-Nikkhah M, Juciano RF, Jurjević Ž, Kachalkin AV, Keochanpheng K, Krisai-Greilhuber I, Li Y-C, Lima AA, Machado AR, Madrid H, Magalhães OMC, Marbach PAS, Melanda GCS, Miller AN, Mongkolsamrit S, Nascimento RP, Oliveira TGL, Ordoñez ME, Orzes R, Palma MA, Pearce CJ, Pereira OL, Perrone G, Peterson SW, Pham THG, Piontelli E, Pordel A, Quijada L, Raja HA, Rosas De Paz E, Ryvarden L, Saitta A, Salcedo SS, Sandoval-Denis M, Santos TAB, Seifert KA, Silva BDB, Smith ME, Soares AM, Sommai S, Sousa JO, Suetrong S, Susca A, Tedersoo L, Telleria MT, Thanakitpipattana D, Valenzuela-Lopez N, Visagie CM, Zapata M, Groenewald JZ (2018) Fungal Planet description sheets: 785–867. Persoonia-Molecular Phylogeny and Evolution of Fungi 41: 238–417. https://doi.org/10.3767/persoonia.2018.41.12
  • Crous PW, Osieck ER, Shivas RG, Tan YP, Bishop-Hurley SL, Esteve-Raventós F, Larsson E, Luangsa-ard JJ, Pancorbo F, Balashov S, Baseia IG, Boekhout T, Chandranayaka S, Cowan DA, Cruz RHSF, Czachura P, De La Peña-Lastra S, Dovana F, Drury B, Fell J, Flakus A, Fotedar R, Jurjević Ž, Kolecka A, Mack J, Maggs-Kölling G, Mahadevakumar S, Mateos A, Mongkolsamrit S, Noisripoom W, Plaza M, Overy DP, Pitek M, Sandoval-Denis M, Vauras J, Wingfield MJ, Abell SE, Ahmadpour A, Akulov A, Alavi F, Alavi Z, Altés A, Alvarado P, Anand G, Ashtekar N, Assyov B, Banc-Prandi G, Barbosa KD, Barreto GG, Bellanger JM, Bezerra JL, Bhat DJ, Bilański P, Bose T, Bozok F, Chaves J, Costa-Rezende DH, Danteswari C, Darmostuk V, Delgado G, Denman S, Eichmeier A, Etayo J, Eyssartier G, Faulwetter S, Ganga KGG, Ghosta Y, Goh J, Góis JS, Gramaje D, Granit L, Groenewald M, Gulden G, Gusmão LFP, Hammerbacher A, Heidarian Z, Hywel-Jones N, Jankowiak R, Kaliyaperumal M, Kaygusuz O, Kezo K, Khonsanit A, Kumar S, Kuo CH, Laessøe T, Latha KPD, Loizides M, Luo SM, Maciá-Vicente JG, Manimohan P, Marbach PAS, Marinho P, Marney TS, Marques G, Martín MP, Miller AN, Mondello F, Moreno G, Mufeeda KT, Mun HY, Nau T, Nkomo T, Okrasińska A, Oliveira JPAF, Oliveira RL, Ortiz DA, Pawłowska J, Pérez-De-Gregorio M'A, Podile AR, Portugal A, Privitera N, Rajeshkumar KC, Rauf I, Rian B, Rigueiro-Rodríguez A, Rivas-Torres GF, Rodriguez-Flakus P, Romero-Gordillo M, Saar I, Saba M, Santos CD, Sarma PVSRN, Siquier JL, Sleiman S, Spetik M, Sridhar KR, Stryjak-Bogacka M, Szczepańska K, Taşikn H, Tennakoon DS, Thanakitpipattana D, Trovão J, Türkekul A, Van Iperen AL, Van 'T Hof P, Vasquez G, Visagie CM, Wingfield BD, Wong PTW, Yang WX, Yarar M, Yarden O, Yilmaz N, Zhang N, Zhu YN, Groenewald JZ (2023) Fungal Planet description sheets: 1478–1549. Persoonia-Molecular Phylogeny and Evolution of Fungi 50: 158–310. https://doi.org/10.3767/persoonia.2023.50.05
  • Cunningham GH (1944) The Gasteromycetes of Australia and New Zealand. John Mclndee Printer, Dunedin, 236 pp.
  • Da Silva Alfredo D, De Oliveira Sousa J, Jacinto De Souza E, Nunes Conrado LM, Goulart Baseia I (2016) Novelties of gasteroid fungi, earthstars and puffballs, from the Brazilian Atlantic rainforest. Anales del Jardín Botánico de Madrid 73(2): 1–10. https://doi.org/10.3989/ajbm.2422
  • De Toni GB (1887) Revisio monographica generis Geasteris Mich. e tribu Gasteromycetum. Revue De Mycologie 9(34): 61–77.
  • Dissing H, Lange M (1962) Gasteromycetes of Congo. Bulletin du Jardin Botanique de l'État à Bruxelles 32: 325–416. https://doi.org/10.2307/3667249
  • Dramani R, Hegbe ADMT, Tabe A, Badou AS, Furneaux B, Ryberg M, Yorou NS (2020) How are basidiospore size measurements affected by drying? Current Research in Environmental & Applied Mycology 10: 63–70. https://doi.org/10.5943/cream/10/1/7
  • Dring DM (1964) Gasteromycetes of West Tropical Africa. Commonwealth Mycological Institute, Kew, 98–109.
  • Edler D, Klein J, Antonelli A, Silvestro D (2021) raxmlGUI 2.0: A graphical interface and toolkit for phylogenetic analyses using RAxML. Methods in Ecology and Evolution 12: 373–377. https://doi.org/10.1111/2041-210X.13512
  • Fries EM (1829) Systema Mycologicum (in Latin). 3. Ernesti Mauritii, Greifswald, 16 pp.
  • Gube M, Piepenbring M (2009) Preliminary annotated checklist of Gasteromycetes in Panama. Nova Hedwigia 89: 519–543. https://doi.org/10.1127/0029–5035/2009/0089–0519
  • Hall TA (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98. https://doi.org/10.1007/978-1-4757-0905-6_31
  • Han BX, Bau T (2016) New records of Geastrum from China. Mycosystema 35(11): 1416–1424. https://doi.org/10.3969/j.issn.1672–6472.2002.04.008
  • He MQ, Zhao RL, Hyde KD, Begerow D, Kemler M, Yurkov A, McKenzie EHC, Raspé O, Kakishima M, Sánchez-Ramírez S, Vellinga EC, Halling R, Papp V, Zmitrovich IV, Buyck B, Ertz D, Wijayawardene NN, Cui BK, Schoutteten N, Liu XZ, Li TH, Yao YJ, Zhu XY, Liu AQ, Li GJ, Zhang MZ, Ling ZL, Cao B, Antonín V, Boekhout T, Da Silva BDB, De Crop E, Decock C, Dima B, Dutta AK, Fell JW, Geml J, Ghobad-Nejhad M, Giachini AJ, Gibertoni TB, Gorjón SP, Haelewaters D, He SH, Hodkinson BP, Horak E, Hoshino T, Justo A, Lim YW, Menolli N, Mešić A, Moncalvo JM, Mueller GM, Nagy LG, Nilsson RH, Noordeloos M, Nuytinck J, Orihara T, Ratchadawan C, Rajchenberg M, Silva-Filho AGS, Sulzbacher MA, Tkalčec Z, Valenzuela R, Verbeken A, Vizzini A, Wartchow F, Wei TZ, Weiß M, Zhao CL, Kirk PM (2019) Notes, outline and divergence times of Basidiomycota. Fungal Diversity 99: 105–367. https://doi.org/10.1007/s13225–019–00435–4
  • He MQ, Cao B, Liu F, Boekhout T, Denchev TT, Schoutteten N, Denchev CM, Kemler M, Gorjón SP, Begerow D, Valenzuela R, Davoodian N, Niskanen T, Vizzini A, Redhead SA, Ramírez-Cruz V, Papp V, Dudka VA, Dutta AK, García-Sandoval R, Liu X-Z, Kijpornyongpan T, Savchenko A, Tedersoo L, Theelen B, Trierveiler-Pereira L, Wu F, Zamora JC, Zeng X-Y, Zhou L-W, Liu S-L, Ghobad-Nejhad M, Giachini AJ, Li G-J, Kakishima M, Olariaga I, Haelewaters D, Sulistyo B, Sugiyama J, Svantesson S, Yurkov A, Alvarado P, Antonín V, Da Silva AF, Druzhinina I, Gibertoni TB, Guzmán-Dávalos L, Justo A, Karunarathna SC, Galappaththi MCA, Toome-Heller M, Hosoya T, Liimatainen K, Márquez R, Mešić A, Moncalvo J-M, Nagy LG, Varga T, Orihara T, Raymundo T, Salcedo I, Silva-Filho AGS, Tkalčec Z, Wartchow F, Zhao CL, Bau T, Cabarroi-Hernández M, Cortés-Pérez A, Decock C, De Lange R, Weiss M, Menolli N, Nilsson RH, Fan Y-G, Verbeken A, Gafforov Y, Meiras-Ottoni A, Mendes-Alvarenga RL, Zeng N-K, Wu Q, Hyde KD, Kirk PM, Zhao R-L (2024) Phylogenomics, divergence times and notes of orders in Basidiomycota. Fungal Diversity 126: 127–406. https://doi.org/10.1007/s13225–024–00535–w
  • Hemmes DE, Desjardin DE (2011) Earthstars (Geastrum, Myriostoma) of the Hawaiian Islands including two new species, Geastrum litchiforme and Geastrum reticulatum. Pacific Science 65: 477–496. https://doi.org/10.2984/65.4.477
  • Hopple Jr JS, Vilgalys R (1999) Phylogenetic relationships in the mushroom genus Coprinus and dark-spored allies based on sequence data from the nuclear gene coding for the large ribosomal subunit RNA: Divergent domains, outgroups, and monophyly. Molecular Phylogenetics and Evolution 13(1): 1–19. https://doi.org/10.1006/mpev.1999.0634
  • Jeppson M, Nilsson RH, Larsson E (2013) European earthstars in Geastraceae (Geastrales, Phallomycetidae) – a systematic approach using morphology and molecular sequence data. Systematics and Biodiversity 11: 437–465. https://doi.org/10.1080/14772000.2013.857367
  • Karun NC, Sridhar KR (2014) Geasters in the Western Ghats and west coast of India. Acta Mycologica 49(2): 207–219. https://doi.org/10.5586/am.2014.023
  • Kasuya T, Hosaka K, Uno K, Kakishima M (2012) Phylogenetic placement of Geastrum melanocephalum and polyphyly of Geastrum triplex. Mycoscience 53: 411–426. https://doi.org/10.1007/S10267–012–0186–Z
  • Katoh K, Misawa K, Kuma K, Miyata T (2002) MAFFT: A novel method for rapid multiple sequence alignment based on fast fourier transform. Nucleic Acids Research 30(14): 3059–3066. https://doi.org/10.1093/nar/gkf436
  • Kirk P, Cannon P, Stalpers J, Minter DW (2008) Dictionary of the Fungi (10th ed.). Centre for Agriculture and Bioscience International, Wallingford, 771 pp.
  • Kotlaba F, Zehnálek P (2018) Sixteen gasteromycetes collected in Cuba 50 years ago. Czech Mycology 70(2): 185–209. https://doi.org/10.33585/CMY.70206
  • Kuhar F, Castiglia V, Papinutti L (2013) Geastrum species of the La Rioja province, Argentina. Mycotaxon 122(1): 145–156. https://doi.org/10.5248/122.145
  • Leite AG, Assis HK, Silva BDB, Sotão HMP, Baseia IG (2011) Geastrum species from the Amazon Forest, Brazil. Mycotaxon 118(1): 383–392. https://doi.org/10.5248/118.383
  • Liu B (1984) The Gasteromycetes of China. Beihefte zur Nova Hedwigia 76: 1–235.
  • Lloyd CG (1902) The Geastrae. Bulletin of the Lloyd Library of Botany 2: 1–43.
  • Mao XL (1998) Economic Fungi of China. Science Press, Beijing, 762 pp.
  • Na Q, Hu YP, Liu ZW, Zeng H, Qi LL, Ding H, Cheng XH, Ge YP (2021) The first reported occurrence of Leucoinocybe (Porotheleaceae, Agaricales) in China: Leucoinocybe lishuiensis sp. nov. from Zhejiang Province. Nova Hedwig 113: 453–469. https://doi.org/10.1127/nova_hedwigia/2021/0661
  • Na Q, Hu YP, Zeng H, Song ZZ, Ding H, Cheng XH, Ge YP (2022) Updated taxonomy on Gerronema (Porotheleaceae, Agaricales) with three new taxa and one new record from China. MycoKeys 89: 87–120. https://doi.org/10.3897/mycokeys.89.79864
  • Na Q, Zeng H, Hu YP, Ding H, Ke BR, Zeng ZH, Liu CJ, Cheng XH, Ge YP (2024) Morphological and phylogenetic analyses reveal five new species of Porotheleaceae (Agaricales, Basidiomycota) from China. MycoKeys 105: 49–95. https://doi.org/10.3897/mycokeys.105.118826
  • Nylander JAA (2004) MrModeltest v2. Program distributed by the author Evolutionary Biology Centre, Uppsala University.
  • Pegler DN, Læssøe T, Spooner BM (1995) British Puffballs, Earthstars and Stinkhorns. An Account of the British Gasteroid Fungi. Royal Botanic Gardens, England, 255 pp.
  • Persoon CH (1794) Neuer Versuch einer systematischen Einteilung der Schwämme. Neues Magazin für die Botanik 1: 63–128.
  • Ponce de León P (1968) A revision of the family Geastraceae. Fieldiana Botany 31: 301–349. https://doi.org/10.5962/bhl.title.2389
  • Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior summarization in bayesian phylogenetics using Tracer 1.7. Systematic biology 67(5): 901–904. https://doi.org/10.1093/sysbio/syy032
  • Rebriev YA, Bulakh EM, Biketova AY (2023) Geastrum juliae (Geastrales, Agaricomycetes), a new species from the Russian Far East. Botanica Serbica 47(1): 41–46. https://doi.org/10.2298/BOTSERB2301041R
  • Ridgway R (1912) Color Standards and Color Nomenclature. Robert Ridgway, Washington D.C, 110 pp. https://doi.org/10.5962/bhl.title.144788
  • Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19(12): 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
  • Soto MK, Wright JE (2000) Taxonomía del género Geastrum (Basidiomycetes, Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la Sociedad Argentina de Botánica 34(3–4): 185–201.
  • Sousa JO, Baracho GS, Baseia IG (2015) Geastrum laevisporum: A new earthstar fungus with uncommon smooth spores. Mycosphere 6(4): 501–507. https://doi.org/10.5943/mycosphere/6/4/12
  • Sousa JO, Suz LM, García MA, Alfredo DS, Conrado LM, Marinho P, Ainsworth AM, Baseia IG, Martín MP (2017) More than one fungus in the pepper pot: Integrative taxonomy unmasks hidden species within Myriostoma coliforme (Geastraceae, Basidiomycota). PLoS ONE 12(6): e0177873. https://doi.org/10.1371/journal.pone.0177873
  • Staněk VJ (1956) Hvězdovka Šmardova-Geastrum Šmardae sp. n. Česká Mykologie 10(1): 18–23.
  • Staněk VJ (1958) Geastraceae. In: Pilát A (Ed.) Flora ČSR B1: Gasteromycetes. Českolovenská Akademie Vĕd, Prague, 392–526. [777–795.]
  • Sunhede S (1989) Geastraceae (Basidiomycotina), Mophology, Ecology and Systematies with Special Emphasison the North European Species. Flora-Oslo-Norway, 534 pp.
  • Tai FL (1979) Sylloge Fungorum Sinicorum. Science Press, Beijing, 1527 pp.
  • Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011) MEGA5: Molecular Evolutionary Genetics Analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28: 2731–2739. https://doi.org/10.1093/molbev/msr121
  • Teng SC (1963) Fungi of China. Science Press, Beijing, 808 pp.
  • Trierveiler-Pereira L, Bezerra KM, Bezerra JL, Baseia IG (2009) First records of Geastraceae and Nidulariaceae (Basidiomycota, Fungi) from Bahia, Northeastern Brazil. Revista Brasileira de Biociências 7(3): 316–319. https://api.semanticscholar.org/CorpusID:83507428
  • Wang X, Bau T (2023) Seven new species of the genus Geastrum (Geastrales, Geastraceae) in China. Journal of Fungi 9(2): 251–276. https://doi.org/10.3390/jof9020251
  • White T, Bruns T, Lee S, Taylor J, Innis M, Gelfand D, Sninsky J (1990) Amplification and Direct Sequencing of Fungal Ribosomal RNA Genes for Phylogenetics. Academic Press, New York, 315–322. https://doi.org/10.1016/B978–0–12–372180–8.50042–1
  • Wu YQ (2023) Taxonomic study on the fungi resources of the genus Geastrum and Volvariella in Shanxi Province. Master thesis, Shanxi Agricultural University, Shanxi, 88 pp.
  • Wu YQ, Liu S, Wang T, Meng J, Chang M, Wang S (2024) Geastrum sanglinense, a new species from the Manghe Rhesus Monkey National Nature Reserve, China. Mycoscience 65(1): 12–18. https://doi.org/10.47371/mycosci.2023.11.001
  • Yang X, Su JQ, Zhou HM, Zhao CL (2024) Morphological characteristics and phylogenetic analyses revealed Geastrum yunnanense sp. nov. (Geastrales, Basidiomycota) from Southwest China. Phytotaxa 665: 179–192. https://doi.org/10.11646/phytotaxa.665.3.1
  • Yang X, Zhu YG, Duan SJ, Wu XX, Zhao CL (2025) Morphology and multigene phylogeny revealed four new species of Geastrum (Geastrales, Basidiomycota) from China. MycoKeys 113: 73–100. https://doi.org/10.3897/mycokeys.113.139672
  • Zamora JC, Calonge FD, Martín MP (2013) New sources of taxonomic information for earthstars (Geastrum, Geastraceae, Basidiomycota): Phenoloxidases and rhizomorph crystals. Phytotaxa 132: 1–20. https://doi.org/10.11646/phytotaxa.132.1.1
  • Zamora JC, Calonge FD, Hosaka K, Martín MP (2014a) Systematics of the genus Geastrum (Fungi: Basidiomycota) revisited. Taxon 63(3): 477–497. https://doi.org/10.12705/633.36
  • Zamora JC (2014b) Proposal to conserve the name Geastrum (Geastrales, Basidiomycota) with a conserved type. Taxon 63(3): 679–680. https://doi.org/10.12705/633.22
  • Zamora JC, Calonge FD, Martín MP (2014c) Combining morphological and phylogenetic analyses to unravel systematics in Geastrum sect. Schmidelia. Mycologia 106(6): 1199–1211. https://doi.org/10.3852/14–072
  • Zamora JC, Calonge FD, Martín MP (2015) Integrative taxonomy reveals an unexpected diversity in Geastrum section Geastrum (Geastrales, Basidiomycota). Persoonia 34(1): 130–165. https://doi.org/10.3767/003158515X687443
  • Zamora JC, Dios MM, Moreno G (2017) Clarifying the identity of Geastrum campestre var. famatinum (Geastrales, Basidiomycota). Phytotaxa 328(2): 159–166. https://doi.org/10.11646/phytotaxa.328.2.6
  • Zhang ZQ, Li CH, Li L, Shen HW, He J, Su XJ, Luo ZL (2023) Geastrum suae sp. nov. (Geastraceae, Basidiomycota) a new species from Yunnan Province, China. Biodiversity Data Journal 11: e99027. https://doi.org/10.3897/BDJ.11.e99027
  • Zhou H, Li JQ, Hou CL (2022) Two new species of the genus Geastrum from the Yanshan Montains in China. Mycosystema 41(1): 1–16. https://doi.org/10.3897/mycokeys.88.81437
  • Zhou TX, Chen YH, Zhao LZ, Fu H, Yang B (2007) Flora Fungorum Sinicorum (Vol. 36). Geastraceae, Nidulariaceae. Science Press, Beijing, 167 pp.