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What are fungal species and how to delineate them?

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Abstract

This is the opening paper in the special issue of Fungal Diversity, which collates the data on defining species. Defining and recognizing species has long been a controversial issue. Since Darwin's proposed origin of species, over 30 species criteria have been brought forth and used to define species boundaries. In recent times, phylogenetic analyses based on multiple loci have been extensively used as a method to define species boundaries. However, only a few mycologists are aware that phylogenetic species criteria can mask discordances among fungal groups, leading to inaccurately defined species boundaries. In the current review, we discuss species recognition criteria, how and where these criteria can be applied along with their limitations and derived alternatives. In order to delimit fungal species, authors need to take into account not only the phylogenetic and phenotypic coherence, but also the timing of events that lead to fungal speciation and subsequent diversifications. Variations in the rate of phenotypic diversifications and convergent fungal evolution make it difficult to establish a universal species recognition criterion. The best practice can only be defined in the context of each fungal group. In this review, we provide a set of guidelines, encouraging an integrative taxonomic approach for species delimitation that can be used to define fungal species boundaries in the future. The other papers in this special issue deal with fungal speciation in Ascomycota, Dothideomycetes, Basidiomycota, basal fungi, lichen-forming fungi, plant pathogenic fungi, and yeasts.

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References

  • Achari SR, Kaur J, Dinh Q, Mann R, Sawbridge T, Summerell BA, Edwards J (2020) Phylogenetic relationship between Australian Fusarium oxysporum isolates and resolving the species complex using the multispecies coalescent model. BMC Genom 21:1–20

    Article  CAS  Google Scholar 

  • Agapow P-M, Bininda-Edmonds ORP, Crandall KA, Gittleman JL, Mace GM, Marshall JC, Purvis A (2004) The impact of species concept on biodiversity studies. Q Rev Biol 79:161–179

    Article  PubMed  Google Scholar 

  • Aldhebiani AY (2018) Species concept and speciation. Saudi J Biol Sci 25:437–440

    Article  PubMed  Google Scholar 

  • Aldrovandi MS, Johnson JE, O’Meara B, Petersen RH, Hughes KW (2015) The Xeromphalina campanella/kauffmanii complex: species delineation and biogeographical patterns of speciation. Mycologia 107:1270–1284

    Article  CAS  PubMed  Google Scholar 

  • Andersson L (1990) The driving force: species concepts and ecology. Taxon 39:375–382

    Article  Google Scholar 

  • Anderson JB, Kohn LM (1998) Genotyping, gene genealogies and genomics bring fungal population genetics above ground. Trends Ecol Evol 13:444–449

    Article  CAS  PubMed  Google Scholar 

  • Anderson JB, Kohn LM (1995) Clonality in soilborne, plant-pathogenic fungi. Annu Rev Phytopathol 33:369–391

    Article  CAS  PubMed  Google Scholar 

  • Anderson JB, Ullrich RC (1979) Biological species of Armillaria mellea in North America. Mycologia 71:402–414

    Article  Google Scholar 

  • Anderson JB, Korhonen K, Ullrich RC (1980) Relationships between European and North-American biological species of Armillaria. Exp Mycol 4:87–95

    Article  Google Scholar 

  • Araújo JPM, Evans HC, Kepler R, Hughes DP (2018) Zombie-ant fungi across continents: 15 new species and new combinations within Ophiocordyceps I. Myrmecophilous hirutelloid species. Stud Mycol 90:119–160

    Article  PubMed Central  PubMed  Google Scholar 

  • Ariyawansa HA, Hyde KD, Jayasiri SC, Buyck B, Chethana KWT, Dai DQ, Dai YC, Daranagama DA, Jayawardena RS, Lücking R, Ghobad-Nejhad M, Niskanen T, Thambugala KM, Voigt K, Zhao RL, Li GJ, Doilom M, Boonmee S, Yang ZL, Cai Q, Cui YY, Bahkali AH, Chen J, Cui BK, Chen JJ, Dayarathne MC, Dissanayake AJ, Ekanayaka AH, Hashimoto A, Hongsanan S, Jones EBG, Larsson E, Li WJ, Li QR, Liu JK, Luo ZL, Maharachchikumbura SSN, Mapook A, McKenzie EHC, Norphanphoun C, Konta S, Pang KL, Perera RH, Phookamsak R, Phukhamsakda C, Pinruan U, Randrianjohany E, Singtripop C, Tanaka K, Tian CM, Tibpromma S, Abdel-Wahab MA, Wanasinghe DN, Wijayawardene NN, Zhang JF, Zhang H, Abdel-Aziz FA, Wedin M, Westberg M, Ammirati JF, Bulgakov TS, Lima DX, Callaghan TM, Callac P, Chang CH, Coca LF, Dal-Forno M, Dollhofer V, Fliegerová K, Greiner K, Griffith GW, Ho HM, Hofstetter V, Jeewon R, Kang JC, Wen TC, Kirk PM, Kytovuori I, Lawrey JD, Xing J, Li H, Liu ZY, Liu XZ, Liimatainen K, Thorsten Lumbsch H, Matsumura M, Moncada B, Nuankaew S, Parnmen S, Santiago ALCMA, Sommai S, Song Y, de Souza CAF, de Souza-Motta CM, Su HY, Suetrong S, Wang Y, Fong WSYH, Zhou LW, Réblová M, Fournier J, Camporesi E, Luangsa-ard JJ, Tasanathai K, Khonsanit A, Thanakitpipattana D, Somrithipol S, Diederich P, Millanes AM, Common RS, Stadler M, Yan JY, Li XH, Lee HW, Nguyen TTT, Lee HB, Battistin E, Marsico O, Vizzini A, Vila J, Ercole E, Eberhardt U, Simonini G, Wen HA, Chen XH, Miettinen O, Spirin V, Hernawati H (2015) Fungal diversity notes 111–252 taxonomic and phylogenetic contributions to fungal taxa. Fungal Divers 75:27–274

    Article  Google Scholar 

  • Arnold ML (2004) Transfer and origin of adaptations through natural hybridization: were Anderson and Stebbins right? Plant Cell 16:562–570

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Aveskamp MM, de Gruyter J, Woudenberg JHC, Verkley GJM, Crous PW (2010) Highlights of the Didymellaceae: a polyphasic approach to characterise Phoma and related pleosporalean genera. Stud Mycol 65:1–60

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Avise JC (2000) Phylogeography: the history and formation of species. Harvard University Press, Cambridge, MA, p 447

    Book  Google Scholar 

  • Avise JC, Wollenberg K (1997) Phylogenetics and the origin of species. Proc Natl Acad Sci USA 94:7748

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Bakhshi M, Arzanlou M, Babai-Ahari A, Groenewald JZ, Braun U, Crous PW (2015) Application of the consolidated species concept to Cercospora spp. from Iran. Persoonia 34:65

    Article  CAS  PubMed  Google Scholar 

  • Banke S, Frisvad JC, Rosendahl S (1997) Taxonomy of Penicillium chrysogenum and related xerophilic species, based on isozyme analysis. Mycol Res 101:617–624

    Article  CAS  Google Scholar 

  • Basilico M, Chiericatti C, Aringoli E, Althaus R, Basilico J (2007) Influence of environmental factors on airborne fungi in houses of Santa Fe City, Argentina. Sci Tot Environ 376:143–150

    Article  CAS  Google Scholar 

  • Bastide P, Solís-Lemus C, Ané C, William Sparks K, Kriebel R (2018) Phylogenetic comparative methods on phylogenetic networks with reticulations. Syst Biol 67:800–820

    Article  PubMed  Google Scholar 

  • Bezerra JD, Oliveira RJ, Paiva LM, Silva GA, Groenewald JZ, Crous PW, Souza-Motta CM (2017) Bezerromycetales and Wiesneriomycetales ord. nov. (class Dothideomycetes), with two novel genera to accommodate endophytic fungi from Brazilian cactus. Mycol Prog 16:297–309

    Article  Google Scholar 

  • Bhunjun CS, Phukhamsakda C, Jayawardena RS, Jeewon R, Promputtha I, Hyde KD (2021) Investigating species boundaries in Colletotrichum. Fungal Divers 107:107–127

    Article  Google Scholar 

  • Bhunjun CS, Dong Y, Jayawardena RS, Jeewon R, Phukhamsakda C, Bundhun D, Hyde KD, Sheng J (2020) A polyphasic approach to delineate species in Bipolaris. Fungal Divers 102:225–256

    Article  Google Scholar 

  • Bhunjun CS, Jayawardena RS, Wei DP, Huanraluek N, Abeywickrama PD, Jeewon R, Monkai J, Hyde K (2019) Multigene phylogenetic characterisation of Colletotrichum artocarpicola sp. nov. from Artocarpus heterophyllus in Northern Thailand. Phytotaxa 418:273–286

    Article  Google Scholar 

  • Bobay LM, Ochman H (2017) The evolution of bacterial genome architecture. Front Genet 8:72

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Boekhout T, Aime MC, Begerow D, Gabaldón T, Heitman J, Kemler M, Khayhan K, Lachance A, Louis EJ, Sun S, Vu D, Yurkov A (2020) The evolving species concepts used for yeasts: from phenotypes and genomes to speciation networks. Fungal Divers. https://doi.org/10.1007/s13225-021-00475-9

  • Boerema GH, Loerakker WM, Hamers MEC (1996) Contributions towards a monograph of Phoma (Coelomycetes) - III. 2. Misapplications of the type species name and the generic synonyms of the section Plenodomus. Persoonia 16:141–190

    Google Scholar 

  • Brasier CM (1995) Episodic selection as a force in fungal microevolution, with special reference to clonal speciation and hybrid introgression. Can J Botany 73:S1213–S1221

    Article  Google Scholar 

  • Brasier CM, Kirk SA (2010) Rapid emergence of hybrids between the two subspecies of Ophiostoma novo-ulmi with a high level of pathogenic fitness. Plant Pathol 59:186–199

    Article  CAS  Google Scholar 

  • Brown EM, McTaggart LR, Dunn D, Pszczolko E, Tsui KG, Morris SK, Stephens D, Kus JV, Richardson SE (2018) Epidemiology and geographic distribution of Blastomycosis, Histoplasmosis, and Coccidioidomycosis, Ontario, Canada, 1990–2015. Emerg Infect Dis 24:1257–1266

    Article  PubMed Central  PubMed  Google Scholar 

  • Brundrett M, Bougher N, Dell B, Grove T, Malajczuk N (1996) Working with mycorrhizas in forestry and agriculture, Vol. 32. Australian Centre for International Agricultural Research, Canberra, p. 374

  • Bustamante DE, Oliva M, Leiva S, Mendoza JE, Bobadilla L, Angulo G, Calderon MS (2019) Phylogeny and species delimitations in the entomopathogenic genus Beauveria (Hypocreales, Ascomycota), including the description of B. peruviensis sp. nov. MycoKeys 58:47–68

    Article  PubMed Central  PubMed  Google Scholar 

  • Cai L, Giraud T, Zhang N, Begerow D, Cai G, Shivas R (2011) The evolution of species concepts and species recognition criteria in plant pathogenic fungi. Fungal Divers 50:121–133

    Article  Google Scholar 

  • Cai L, Hyde KD, Taylor PWJ, Weir B, Waller JM, Abang MM, Zang JC, Yang YL, Phouliyong S, Prihastuti ZY, Shivas RG, McKenzie EHC, Johnston PR (2009) A polyphasic approach for studying Colletotrichum. Fungal Divers 39:183–204

    Google Scholar 

  • Campbell N, Reece J (2002) Biology. Benjamin Cummings, San Fransico

    Google Scholar 

  • Carbone I, Kohn LM (2001) A microbial population-species interface: nested cladistic and coalescent inference with multilocus data. Mol Ecol 10:947–964

    Article  CAS  PubMed  Google Scholar 

  • Carbone I, Anderson JB, Kohn LM (1999) Patterns of descent in clonal lineages and their multilocus fingerprints are resolved with combined gene genealogies. Evolution 53:11–21

    Article  CAS  PubMed  Google Scholar 

  • Carstens BC, Knowles LL (2007) Estimating species phylogeny from gene-tree probabilities despite incomplete lineage sorting: an example from Melanoplus grasshoppers. Syst Biol 56:400–411

    Article  PubMed  Google Scholar 

  • Caten CE (1981) Parasexual processes in fungi. In: Gull K, Oliver SG (eds) The fungal nucleus. Cambridge University Press, Cambridge, pp 191–214

    Google Scholar 

  • Chang Y, Wang S, Sekimoto S, Aerts AL, Choi C, Clum A, LaButti KM, Lindquist EA, Yee Ngan C, Ohm RA, Salamov AA (2015) Phylogenomic analyses indicate that early fungi evolved digesting cell walls of algal ancestors of land plants. Genome Biol Evol 7:1590–1601

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Chethana KWT, Niranjan M, Dong W, Samarakoon MC, Bao DF, Calabon MS, Chaiwan N, Chuankid B, Dayarathne MC, de Silva NI, Devadatha B, Dissanayake AJ, Goonasekara ID, Huanraluek N, Jayawardena RS, Karunarathna A, Luo ZL, Marasinghe DS, Ma XY, Norphanphoun C, Pem D, Perera RH, Rathnayaka AR, Raspé O, Samarakoon BC, Senwanna C, Sun YR, Tang X, Thiyagaraja V, Tennakoon DS, Zeng M, Zeng XY, Zhang JY, Zhang SN, Bulgakov TS, Camporesi E, Sarma VV, Wang Y, Bhat DJ, Hyde KD (2021) AJOM new records and collections of fungi: 101–150. Asian J Mycol 4:113–260

    Google Scholar 

  • Chethana KWT, Jayawardena RS, Hyde KD (2020) Hurdles in fungal taxonomy: Effectiveness of recent methods in discriminating taxa. Megataxa 1:114–122

    Google Scholar 

  • Chethana KWT, Zhou Y, Zhang W, Xing QK, Hyde KD, Yan JY, Li XH (2017) Coniella vitis sp. nov. is the common pathogen of white rot in Chinese vineyards. Plant Dis 101:2123–2136

    Article  CAS  PubMed  Google Scholar 

  • Chibucos MC, Soliman S, Gebremariam T, Lee H, Daugherty S, Orvis J, Shetty AC, Crabtree J, Hazen TH, Etienne KA, Kumari P (2016) An integrated genomic and transcriptomic survey of mucormycosis-causing fungi. Nat Commun 7:12218

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Chupp C (1954) A monograph of the fungus genus Cercospora. Plant Pathology Department, Cornell University, Ithaca, NY, p 200

    Google Scholar 

  • Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork PN (2006) Toward automatic reconstruction of a highly resolved tree of life. Science 311:1283–1287

    Article  CAS  PubMed  Google Scholar 

  • Clement M, Posada D, Crandall KA (2000) TCS: a computer program to estimate gene genealogies. Mol Ecol 9:1657–1659

    Article  CAS  PubMed  Google Scholar 

  • Corradi N, Lildhar L (2012) Meiotic genes in the arbuscular mycorrhizal fungi: What for? Commun Integr Biol 5:187–189

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Corsaro D, Walochnik J, Venditti D, Hauröder B, Michel R (2020) Solving an old enigma: Morellospora saccamoebae gen. nov., sp. nov. (Rozellomycota), a Sphaerita-like parasite of free-living amoebae. Parasitol Res 119:925–934

    Article  PubMed  Google Scholar 

  • Couch BC, Fudal I, Lebrun M-H, Tharreau D, Valent B, van Kim P, Nottéghem J-L, Kohn LM (2005) Origins of host-specific populations of the blast pathogen Magnaporthe oryzae in crop domestication with subsequent expansion of pandemic clones on rice and weeds of rice. Genetics 170:613–630

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Coyne JA (1994) Ernst Mayr and the origin of species. Evolution 48:19–30

    Article  PubMed  Google Scholar 

  • Coyne JA, Orr HA (2004) Speciation. Sinauer Associates, Sunderland, MA, p 545

    Google Scholar 

  • Crouch JA (2014) Colletotrichum caudatum sl is a species complex. IMA Fungus 5:17–30

    Article  PubMed Central  PubMed  Google Scholar 

  • Cronquist A (1978) Once again, what is a species? Biosystematics in agriculture. Beltsville Symp Agr Res 2:3–20

    Google Scholar 

  • Crous PW, Lombard L, Sandoval-Denis M, Hyde KD (2021) Fusarium: more than a node or a foot cell (In press)

  • Damm U, Woudenberg J, Cannon P, Crous P (2009) Colletotrichum species with curved conidia from herbaceous hosts. Fungal Divers 39:45–87

    Google Scholar 

  • Das K, Aminuzzaman FM (2017) Morphological and ecological characterization of xylotrophic fungi in mangrove forest regions of Bangladesh. J Adv Biol Biotechnol 28:1–5

    Article  Google Scholar 

  • Darwin C (1859) On the origin of species by means of natural selection or the preservation of favoured races in the struggle for life. Murray, London

    Book  Google Scholar 

  • Dawson CO, Gentles JC (1961) The perfect states of Keratinomyces ajelloi Vanbreuseghem, Trichophyton terrestre Durie & Frey and Microsporum nanum Fuentes. Sabouraudia 1:49–57

    Article  CAS  PubMed  Google Scholar 

  • Dayarathne MC, Boonmee S, Braun U, Crous PW, Daranagama DA, Dissanayake AJ, Ekanayaka H, Jayawardena R, Jones EB, Maharachchikumbura SS, Perera RH (2016) Taxonomic utility of old names in current fungal classification and nomenclature: Conflicts, confusion & clarifications. Mycosphere 7:1622–1648

    Article  Google Scholar 

  • De Candole A (1813) Théorie élémentaire de la botanique; ou, Exposition des prinicpes de la classification naturelle et de l’art de décrire et d’étudier les végétaux. London

  • De Queiroz K (2005) Ernst Mayr and the modern concept of species. Proc Natl Acad Sci 102:6600–6607

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • De Queiroz K (2007) Species concepts and species delimitation. Syst Biol 56:879–886

    Article  PubMed  Google Scholar 

  • De Vienne DM, Ollier S, Aguileta G (2012) Phylo-MCOA: a fast and efficient method to detect outlier genes and species in phylogenomics using multiple co-inertia analysis. Mol Biol Evol 29:1587–1598

    Article  CAS  PubMed  Google Scholar 

  • Delaux PM, Varala K, Edger PP, Coruzzi GM, Pires JC, Ané JM (2014) Comparative phylogenomics uncovers the impact of symbiotic associations on host genome evolution. PLoS Genet 10:e1004487

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Del-Prado R, Blanco O, Lumbsch HT, Divakar PK, Elix JA, Molina MC, Crespo A (2013) Molecular phylogeny and historical biogeography of the lichen-forming fungal genus Flavoparmelia (Ascomycota: Parmeliaceae). Taxon 62:928–939

    Article  Google Scholar 

  • Delsuc F, Brinkmann H, Philippe H (2005) Phylogenomics and the reconstruction of the tree of life. Nat Rev Genet 6:361–375

    Article  CAS  PubMed  Google Scholar 

  • Demos TC, Peterhans JCK, Agwanda B, Hickerson MJ (2014) Uncovering cryptic diversity and refugial persistence among small mammal lineages across the Eastern Afromontane biodiversity hotspot. Mol Phylogenet Evol 71:41–54

    Article  PubMed  Google Scholar 

  • Desjardin DE, Perry BA (2015) A new species of Scytinopogon from the island of Príncipe, Republic of São Tomé and Príncipe, West Africa. Mycosphere 6:434–441

    Article  Google Scholar 

  • Desjardins CA, Giamberardino C, Sykes SM, Yu C-H, Tenor JL, Chen Y, Yang T, Jones AM, Sun S, Haverkamp MR, Heitman J, Litvintseva AP, Perfect JR, Cuomo CA (2017) Population genomics and the evolution of virulence in the fungal pathogen Cryptococcus neoformans. Genome Res 27:1207–1219

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Dettman JR, Jacobson DJ, Taylor JW (2003) A multilocus genealogical approach to phylogenetic species recognition in the model eukaryote Neurospora. Evolution 57:2703–2720

    PubMed  Google Scholar 

  • Dwidjoseputro D (1961) Studies on Monilia sitophila from Indonesia. Bull Torrey Bot Club 1:404–411

    Article  Google Scholar 

  • Dobzhansky T (1951) Experiments on sexual isolation in Drosophila: X. Reproductive isolation between Drosophila pseudoobscura and Drosophila persimilis under natural and under laboratory conditions. Proc Natl Acad Sci USA 37:792–796

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Dobzhansky T (1937) Genetics and the origin of species. Columbia University Press, New York

    Google Scholar 

  • Domingos FMCB, Bosque RJ, Cassimiro J, Colli GR, Rodrigues MT, Santos MG, Beheregaray LB (2014) Out of the deep: Cryptic speciation in a Neotropical gecko (Squamata, Phyllodactylidae) revealed by species delimitation methods. Mol Phylogenet Evol 80:113–124

    Article  PubMed  Google Scholar 

  • Egidi E, Hoog S, Isola D, Onofri S, Quaedvlieg W, de Vries ME, Verkley JGM, Stielow JB, Zucconi L, Selbmann L (2014) Phylogeny and taxonomy of meristematic rock-inhabiting black fungi in the Dothideomycetes based on multi-locus phylogenies. Fungal Divers 65:127–165

    Article  Google Scholar 

  • Ence DD, Carstens BC (2011) SpedeSTEM: a rapid and accurate method for species delimitation. Mol Ecol Resour 11:473–480

    Article  PubMed  Google Scholar 

  • Ereshefsky M (2007) Species, taxonomy, and systematics. Philosophy of biology. Elsevier, North-Holland, pp 403–427

    Chapter  Google Scholar 

  • Fisher RA (1930) The genetical theory of natural selection. Clarendon Press, Oxford

    Book  Google Scholar 

  • Fitzpatrick DA, Logue ME, Stajich JE, Butler G (2006) A fungal phylogeny based on 42 complete genomes derived from super tree and combined gene analysis. BMC Evol Biol 6:99

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Fournier E, Giraud T (2008) Sympatric genetic differentiation of a generalist pathogenic fungus, Botrytis cinerea, on two different host plants, grapevine and bramble. J Evol Biol 21:122–132

    Article  CAS  PubMed  Google Scholar 

  • Frenkel O, Peever TL, Chilvers MI, Özkilinc H, Can C, Abbo S, Shtienberg D, Sherman A (2010) Ecological genetic divergence of the fungal pathogen Didymella rabiei on sympatric wild and domesticated Cicer spp. (Chickpea). Appl Environ Microbiol 76:30–39

    Article  CAS  PubMed  Google Scholar 

  • Frisvad JC (2015) Taxonomy, chemo diversity and chemo consistency of Aspergillus, Penicillium, and Talaromyces species. Front Microbiol 5:773

    Article  PubMed Central  PubMed  Google Scholar 

  • Fuckel L (1872) Symbolae mycologicae. Beiträge zur Kenntniss der rheinischen Pilze. Erster Nachtrag Jahrb Nassau Ver Naturkd 25–26:287–346

    Google Scholar 

  • Fujita MK, Leache DA, Burbrink FT, McGuire JA, Moritz C (2012) Coalescent-based species delimitation in an integrative taxonomy. Trends Ecol Evol 27:480–488

    Article  PubMed  Google Scholar 

  • Fujisawa T, Barraclough TG (2013) Delimiting species using single-locus data and the Generalized Mixed Yule Coalescent approach: a revised method and evaluation on simulated data sets. Syst Biol 62:707–724

    Article  PubMed Central  PubMed  Google Scholar 

  • Giraud T, Refrégier G, Le Gac M, de Vienne DM, Hood ME (2008) Speciation in fungi. Fungal Genet Biol 45:791–802

    Article  CAS  PubMed  Google Scholar 

  • Gladieux P, Wilson BA, Perraudeau F, Montoya LA, Kowbel D, Hann-Soden C, Fischer M, Sylvain I, Jacobson DJ, Taylor JW (2015) Genomic sequencing reveals historical, demographic and selective factors associated with the diversification of the fire-associated fungus Neurospora discreta. Mol Ecol 24:5657–5675

    Article  PubMed  Google Scholar 

  • Gladieux P, Vercken E, Fontaine MC, Hood ME, Jonot O, Couloux A, Giraud T (2011) Maintenance of fungal pathogen species that are specialized to different hosts: allopatric divergence and introgression through secondary contact. Mol Biol Evol 28:459–471

    Article  CAS  PubMed  Google Scholar 

  • Gostinčar C (2020) Towards genomic criteria for delineating fungal species. J Fungi 6:246–264

    Article  CAS  Google Scholar 

  • Gostinčar C, Ohm RA, Kogej T, Sonjak S, Turk M, Zajc J, Zalar P, Grube M, Sun H, Han J, Sharma A (2014) Genome sequencing of four Aureobasidium pullulans varieties: biotechnological potential, stress tolerance, and description of new species. BMC Genom 15:1–29

    Article  CAS  Google Scholar 

  • Greig D, Louis EJ, Borts RH, Travisano M (2002) Hybrid speciation in experimental populations of yeast. Science 298:1773–1775

    Article  CAS  PubMed  Google Scholar 

  • Gregory TR (2009) Understanding natural selection: essential concepts and common misconceptions. Evolution 2:156–175

    Google Scholar 

  • Groenewald JZ, Nakashima C, Nishikawa J, Shin HD, Park JH, Jama AN, Groenewald M, Braun U, Crous PW (2013) Species concepts in Cercospora: spotting the weeds among the roses. Stud Mycol 75:115–170

    Article  CAS  PubMed  Google Scholar 

  • Gryganskyi A, Muszewska A (2014) Whole genome sequencing and the Zygomycota. Fungal Genom Biol 4:1000e116

    Google Scholar 

  • Gryganskyi AP, Golan J, Dolatabadi S, Mondo S, Robb S, Idnurm A, Muszewska A, Steczkiewicz K, Masonjones S, Liao HL, Gajdeczka MT (2018) Phylogenetic and phylogenomic definition of Rhizopus species. G3: Genes Genom Genet 8: 2007–2018

    Article  CAS  Google Scholar 

  • Guého E, Midgley G, Guillot J (1996) The genus Malassezia with description of four new species. Antonie Van Leeuwenhoek 69:337–355

    Article  PubMed  Google Scholar 

  • Gusfield D, Bansal V (2005) A fundamental decomposition theory for phylogenetic networks and incompatible characters. In: Annual international conference on research in computational molecular biology. Springer, Berlin

  • Haelewaters D, De Kesel A, Pfister DH (2018) Integrative taxonomy reveals hidden species within a common fungal parasite of ladybirds. Sci Rep 8:1–6

    Article  CAS  Google Scholar 

  • Hagen F, Khayhan K, Theelen B, Kolecka A, Polacheck I, Sionov E, Falk R, Parnmen S, Lumbsch HT, Boekhout T (2015) Recognition of seven species in the Cryptococcus gattii/Cryptococcus neoformans species complex. Fungal Genet Biol 78:16–48

    Article  CAS  PubMed  Google Scholar 

  • Hare MC (2001) Prospects for nuclear gene phylogeography. Trends Ecol Evol 16:700–706

    Article  Google Scholar 

  • Hardison RC (2003) Comparative genomics. PLoS Biol 17:e58

    Article  CAS  Google Scholar 

  • Haridas S, Albert R, Binder M, Bloem J, LaButti K, Salamov A, Andreopoulos B, Baker SE, Barry K, Bills G, Bluhm BH, Cannon C, Castanera R, Culley DE, Daum C, Ezra D, González JB, Henrissat B, Kuo A, Liang C, Lipzen A, Lutzoni F, Magnuson J, Mondo SJ, Nolan M, Ohm RA, Pangilinan J, Park H-J, Ramírez L, Alfaro M, Sun H, Tritt A, Yoshinaga Y, Zwiers L-H, Turgeon BG, Goodwin SB, Spatafora JW, Crous PW, Grigoriev IV (2020) 101 Dothideomycetes genomes: a test case for predicting lifestyles and emergence of pathogens. Stud Mycol 96:141–153

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Harrington T, Rizzo D (1999) Defining species in the fungi. Structure and dynamics of fungal populations. Springer, Dordrecht, pp 43–71

    Chapter  Google Scholar 

  • Harrington TC, Wingfield MJ (1998) The Ceratocystis species on conifers. Can J Bot 76:1446–1457

    Google Scholar 

  • Hausdorf B (2011) Progress toward a general species concept. Evolution 65:923–931

    Article  PubMed  Google Scholar 

  • Hawksworth DL, Lücking R (2017) Fungal diversity revisited: 2.2 to 3.8 million species. In: Heitman J, Howlett B, Crous P, Stukenbrock E, James T, Gow N (eds) The Fungal Kingdom, vol 5. ASM Press, Washington, pp 79–95

    Chapter  Google Scholar 

  • Hawlitschek O, Nagy ZT, Berger J, Glaw F (2013) Reliable DNA barcoding performance proved for species and island populations of Cormoran squamate reptiles. PLoS ONE 8:e73368

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Hedin M, Carlson D, Coyle F (2015) Sky island diversification meets the multispecies coalescent divergence in the spruce-fir moss spider (Microhexura montivaga, Araneae, Mygalomorphae) on the highest peaks of southern Appalachia. Mol Ecol 24:3467–3484

    Article  PubMed  Google Scholar 

  • Hennig W (1965) Phylogenetic systematics. Annu Rev Entomol 10:97–116

    Article  Google Scholar 

  • Heled J, Drummond AJ (2010) Bayesian inference of species trees from multilocus data. Mol Biol Evol 27:570–580

    Article  CAS  PubMed  Google Scholar 

  • Hey J (2006) On the failure of modern species concepts. Trends Ecol Evol 21:447–450

    Article  PubMed  Google Scholar 

  • Hibbett DS, Fukumasa-Nakai Y, Tsuneda A, Donoghue MJ (1995) Phylogenetic diversity in Shiitake inferred from nuclear ribosomal DNA sequences. Mycologia 87:618–638

    Article  CAS  Google Scholar 

  • Hilário S, Santos L, Alves A (2021) Diaporthe amygdali, a species complex or a complex species? Fungal Biol 125:505

    Article  PubMed  Google Scholar 

  • Hirayama K, Tanaka K (2011) Taxonomic revision of Lophiostoma and Lophiotrema based on re-evaluation of morphological characters and molecular analyses. Mycoscience 52:401–412

    Article  Google Scholar 

  • Hsuan H, Salleh B, Zakaria L (2011) Molecular identification of Fusarium species in Gibberella fujikuroi species complex from Rice, Sugarcane and Maize from Peninsular Malaysia. Int J Mol Sci 12:6722–6732

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Huelsenbeck JP (1995) Performance of phylogenetic methods in simulation. Syst Biol 44:17–48

    Article  Google Scholar 

  • Hudler GW, Jensen-Tracy S, Banik MT (1998) Rhytisma americanum sp. nov.: a previously undescribed species of Rhytisma on maples (Acer spp.). Mycotaxon 68:405–416

    Google Scholar 

  • Hudson RR, Coyne JA (2002) Mathematical consequences of the genealogical species concept. Evolution 56:1557–1565

    PubMed  Google Scholar 

  • Huson DH, Bryant D (2006) Application of phylogenetic networks in evolutionary studies. Mol Biol Evol 23:254–267

    Article  CAS  PubMed  Google Scholar 

  • Hyde KD, Jeewon R, Chen YJ, Bhunjun CS, Calabon MS, Jiang HB, Lin CG, Norphanphoun C, Sysouphanthong P, Pem D, Tibpromma S (2020a) The numbers of fungi: is the descriptive curve flattening? Fungal Divers 103:219–271

    Article  Google Scholar 

  • Hyde KD, Dong Y, Phookamsak R, Jeewon R, Bhat DJ, Jones EBG, Liu NG, Abeywickrama PD, Mapook A, Wei D, Perera RH, Manawasinghe IS, Pem D, Bundhun D, Karunarathna A, Ekanayaka AH, Bao DF, Li J, Samarakoon MC, Chaiwan N, Lin CG, Phutthacharoen K, Zhang SN, Senanayake IC, Goonasekara ID, Thambugala KM, Phukhamsakda C, Tennakoon DS, Jiang HB, Yang J, Zeng M, Huanraluek N, Liu JK, Wijesinghe SN, Tian Q, Tibpromma S, Brahmanage RS, Boonmee S, Huang SK, Thiyagaraja V, Lu YZ, Jayawardena RS, Dong W, Yang EF, Singh SK, Singh SM, Rana S, Lad SS, Anand G, Devadatha B, Niranjan M, Sarma VV, Liimatainen K, Aguirre-Hudson B, Niskanen T, Overall A, Alvarenga RLM, Gibertoni TB, Pfliegler WP, Horváth E, Imre A, Alves AL, Santos ACS, Tiago AV, Bulgakov TS, Wanasinghe DN, Bahkali AH, Doilom M, Elgorban AM, Maharachchikumbura SSN, Rajeshkumar KC, Haelewaters D, Mortimer PE, Zhao Q, Lumyong S, Xu J, Sheng J (2020b) Fungal diversity notes 1151–1276: taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Divers 100:5–277

    Article  Google Scholar 

  • Hyde KD, de Silva NI, Jeewon R, Bhat DJ, Phookamsak R, Doilom M, Boonmee S, Jayawardena RS, Maharachchikumbura SSN, Senanayake IC, Manawasinghe IS, Liu NG, Abeywickrama PD, Chaiwan N, Karunarathna A, Pem D, Lin CG, Sysouphanthong P, Luo ZL, Wei DP, Wanasinghe DN, Norphanphoun C, Tennakoon DS, Samarakoon MC, Jayasiri SC, Jiang HB, Zeng XY, Li JF, Wijesinghe SN, Devadatha B, Goonasekara ID, Brahmanage RS, Yang EF, Aluthmuhandiram JVS, Dayarathne MC, Marasinghe DS, Li WJ, Dissanayake LS, Dong W, Huanraluek N, Lumyong S, Liu JK, Karunarathna SC, Jones EBG, Al-Sadi AM, Xu JC, Harishchandra D, Sarma VV (2020c) AJOM new records and collections of fungi 1–100. Asian J Mycol 3:22–294

    Article  Google Scholar 

  • Hyde KD, Xu JC, Rapior S, Jeewon R, Lumyong S, Niego AGT, Abeywickrama PD, Aluthmuhandiram JVS, Brahamanage RS, Brooks S, Chaiyasen A, Chethana KWT, Chomnunti P, Chepkirui C, Chuankid B, de Silva NI, Doilom M, Faulds C, Gentekaki E, Gopalan V, Kakumyan P, Harishchandra D, Hemachandran H, Hongsanan S, Karunarathna A, Karunarathna SC, Khan S, Kumla J, Jayawardena RS, Liu JK, Liu N, Luangharn T, Macabeo APG, Marasinghe DS, Meeks D, Mortimer PE, Mueller P, Nadir S, Nataraja KN, Nontachaiyapoom S, O’Brien M, Penkhrue W, Phukhamsakda C, Ramanan US, Rathnayaka AR, Sadaba RB, Sandargo B, Samarakoon BC, Tennakoon DS, Siva R, Sriprom W, Suryanarayanan TS, Sujarit K, Suwannarach N, Suwunwong T, Thongbai B, Thongklang N, Wei DP, Wijesinghe SN, Winiski J, Yan J, Yasanthika E, Stadler M (2019) The amazing potential of fungi: 50 ways we can exploit fungi industrially. Fungal Divers 97:1–136

    Article  Google Scholar 

  • Isaac NJB, Mallet J, Mace GM (2004) Taxonomic inflation: its influence on macroecology and conservation. Trends Ecol Evol 19:464–469

    Article  PubMed  Google Scholar 

  • Jayawardena RS, Camporesi E, Elgorban AM, Bahkali AH, Yan J, Hyde KD (2017) A new species of Colletotrichum from Sonchus sp. in Italy. Phytotaxa 314:55–63

    Article  Google Scholar 

  • Jayawardena RS, Hyde KD, Damm U, Cai L, Liu M, Li XH, Zhang W, Zhao WS, Yan JY (2016) Notes on currently accepted species of Colletotrichum. Mycosphere 7:1192–1260

    Article  Google Scholar 

  • Jeewon R, Hyde K (2016) Establishing species boundaries and new taxa among fungi: recommendations to resolve taxonomic ambiguities. Mycosphere 7:1669–1677

    Article  Google Scholar 

  • Jeewon R, Liew ECY, Hyde KD (2004) Phylogenetic evaluation of species nomenclature of Pestalotiopsis in relation to host association. Fungal Divers 17:39–55

    Google Scholar 

  • Johnston PR, Park D, Ho WWH, Alexander BJR (2017) Genetic validation of historical plant pathology records – a case study based on the fungal genus Phoma from the ICMP culture collection. Plant Pathol 66:1424–1431

    Article  Google Scholar 

  • Justo A, Minnis AM, Ghignone S, Menolli N, Capelari M, Rodríguez O, Malysheva E, Contu M, Vizzini A (2011) Species recognition in Pluteus and Volvopluteus (Pluteaceae, Agaricales): morphology, geography and phylogeny. Mycol Progress 10:453–479

    Article  Google Scholar 

  • Kamel L, Keller-Pearson M, Roux C, Ané JM (2017) Biology and evolution of arbuscular mycorrhizal symbiosis in the light of genomics. New Phytol 213:531–536

    Article  CAS  PubMed  Google Scholar 

  • Kanz B, von Brackel W, Cezanne R, Eichler M, Hohmann ML, Teuber D, Printzen C (2015) DNA barcodes for the distinction of reindeer lichens: a case study using Cladonia rangiferina and C. stygia. Herzogia 28:445–464

    Article  Google Scholar 

  • Karimi K, Ahari AB, Arzanlou M, Amini J, Pertot I, Rota-Stabelli O (2017) Application of the consolidated species concept to identify the causal agent of strawberry anthracnose in Iran and initial molecular dating of the Colletotrichum acutatum species complex. Eur J Plant Pathol 147:375–387

    Article  Google Scholar 

  • Kasuga T, White TJ, Koenig G, McEwen J, Restrepo A, Castañeda E, Da Silva LC, Heins-Vaccari EM, De Freitas RS, Zancopé-Oliveira RM, Qin Z (2003) Phylogeography of the fungal pathogen Histoplasma capsulatum. Mol Ecol 12:3383–3401

    Article  CAS  PubMed  Google Scholar 

  • Kekkonen M, Mutanen M, Kaila L, Nieminen M, Hebert PD (2015) Delineating species with DNA barcodes: a case of taxon dependent method performance in moths. PLoS ONE 10:e0122481

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Kerrigan RW, Imbernon M, Callac P, Billette C, Olivier JM (1994) The heterothallic life cycle of Agaricus bisporus var. burnettii and the inheritance of its tetrasporic trait. Exp Mycol 18:193–210

    Article  Google Scholar 

  • Kobmoo N, Mongkolsamrit S, Arnamnart N, Luangsa-ard JJ, Giraud T (2019) Population genomics revealed cryptic species within host-specific zombie-ant fungi (Ophiocordyceps unilateralis). Mol Phylogenet Evol 140:106580

    Article  PubMed  Google Scholar 

  • Kodsueb R, Lumyong S, McKenzie EHC, Bahkali AH, Hyde KD (2016) Relationships between terrestrial and freshwater lignicolous fungi. Fungal Ecol 19:155–168

    Article  Google Scholar 

  • Kohn LM (2005) Mechanisms of fungal speciation. Annu Rev Phytopathol 43:279–308

    Article  CAS  PubMed  Google Scholar 

  • Korhonen K (1978a) Intersterility groups of Heterobasidion annosum. Commun Inst Fenn 94:1–25

    Google Scholar 

  • Korhonen K (1978b) Interfertility and clonal size in the Armillariella mellea complex. Karstenia 18:31–42

    Article  Google Scholar 

  • Kruse J, Piątek M, Lutz M, Thines M (2018) Broad host range species in specialised pathogen groups should be treated with suspicion – a case study on Entyloma infecting Ranunculus. Persoonia 41:175–201

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Kubatko LS, Carstens BC, Knowles LL (2009) STEM: species tree estimation using maximum likelihood for gene trees under coalescence. Bioinformatics 25:971–973

    Article  CAS  PubMed  Google Scholar 

  • Kuhnert E, Surup F, Sir EB, Lambert C, Hyde KD, Hladki AI, Romero AI, Stadler M (2015) Lenormandins A—G, new azaphilones from Hypoxylon lenormandii and Hypoxylon jaklitschii sp. nov., recognised by chemotaxonomic data. Fungal Divers 71:165–184

    Article  Google Scholar 

  • Kurtzman CP, Fell JW, Boekhout T, Robert V (2011) Methods for isolation, phenotypic characterization and maintenance of yeasts. In: Kurtzman C, Fel JW, Boekhout T (eds) The yeasts: a taxonomic study. Elsevier, Amsterdam, pp 7–110

    Google Scholar 

  • Kwon-Chung KJ, Bennett JE, Wickes BL, Meyer W, Cuomo CA, Wollenburg KR, Bicanic TA, Castañeda E, Chang YC, Chen J, Cogliati M (2017) The case for adopting the “species complex” nomenclature for the etiologic agents of cryptococcosis. Msphere 2:e00357-e416

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Kytövuori I (1988) The Tricholoma caligatum group in Europe and North Africa. Karstenia 28:65–77

    Article  Google Scholar 

  • Laurence M, Summerell B, Burgess L, Liew ECY (2014) Genealogical concordance phylogenetic species recognition in the Fusarium oxysporum species complex. Fungal Biol 118:374–384

    Article  PubMed  Google Scholar 

  • Leavitt SD, Grewe F, Widhelm T, Muggia L, Wray B, Lumbsch HT (2016) Resolving evolutionary relationships in lichen-forming fungi using diverse phylogenomic datasets and analytical approaches. Sci Rep 6:22262

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Leavitt S, Fernández-Mendoza F, Pérez-Ortega S, Sohrabi M, Divakar P, Lumbsch T, Clair LS (2013) DNA barcode identification of lichen-forming fungal species in the Rhizoplaca melanophthalma species-complex (Lecanorales, Lecanoraceae), including five new species. MycoKeys 7:1–22

    Article  Google Scholar 

  • Leavitt S, Johnson L, Clai L (2011) Species delimitation and evolution in morphologically and chemically diverse communities of the lichen-forming genus Xanthoparmelia (Parmeliaceae, Ascomycota) in Western North America. Am J Bot 98:175–188

    Article  PubMed  Google Scholar 

  • Levetin E, Horner W, Scott J (2015) Taxonomy of allergenic fungi. J Allergy Clin Immunol Pract 4:375–385

    Article  PubMed  Google Scholar 

  • Li WJ, Bhat JD, Hyde KD, Wang Y (2016) Towards a natural classification of Dothideomycetes 4: The genera Bryopelta, Bryorella, Bryosphaeria, Lophiosphaerella and Maireella (Dothideomycetes incertae sedis). Phytotaxa 176:28–41

    Google Scholar 

  • Lin X, Stur E, Ekrem T (2015) Exploring genetic divergence in a species-rich insect genus using 2790 DNA barcodes. PLoS ONE 10:e0138993

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Linder CR, Rieseberg LH (2004) Reconstructing patterns of reticulate evolution in plants. Am J Bot 91:1700–1708

    Article  PubMed Central  PubMed  Google Scholar 

  • Linnaeus C (1753) Species Plantarum. In: Facsimile (Ed) (1957) London, Ray Society, British Museum

  • Litvintseva AP, Xu J, Mitchell TG (2011) Population structure and ecology of Cryptococcus. In: Heitman J, Kozel TR, Kwon-Chung KJ, Perfect JR, Casadevall A (eds) Cryptococcus: from human pathogen to model yeast. ASM Press, Washington DC, pp 97–111

    Google Scholar 

  • Liu F, Wang M, Damm U, Crous PW, Cai L (2016) Species boundaries in plant pathogenic fungi: a Colletotrichum case study. BMC Evol Biol 16:81

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Looney BP, Adamčík S, Matheny PB (2020) Coalescent-based delimitation and species-tree estimations reveal Appalachian origin and Neogene diversification in Russula subsection Roseinae. Mol Phylogenet Evol 147:106787

    Article  PubMed  Google Scholar 

  • Lorch JM, Palmer JM, Vanderwolf KJ, Schmidt KZ, Verant ML, Weller TJ, Blehert DS (2018) Malassezia vespertilionis sp. nov.: a new cold-tolerant species of yeast isolated from bats. Persoonia 41:56–70

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Lücking R, Aime MC, Robbertse B, Miller AN, Ariyawansa HA, Aoki T, Cardinali G, Crous PW, Druzhinina IS, Geiser DM, Hawksworth DL (2020) Unambiguous identification of fungi: where do we stand and how accurate and precise is fungal DNA barcoding? IMA Fungus 11:1–32

    Article  PubMed Central  PubMed  Google Scholar 

  • Lücking R, Dal-Forno M, Sikaroodi M, Gillevet PM, Bungartz F, Moncada B, Yánez-Ayabaca A, Chaves JL, Coca LF, Lawrey JD (2014) A single macrolichen constitutes hundreds of unrecognized species. Proc Natl Acad Sci USA 111:11091–11096

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Lutsak T, Fernández-Mendoza F, Kirika P, Wondafrash M, Printzen C (2020) Coalescence-based species delimitation using genome-wide data reveals hidden diversity in a cosmopolitan group of lichens. Org Divers Evol 14:1–30

    Google Scholar 

  • Maddison WP (1997) Gene trees in species trees. Syst Biol 46:523–536

    Article  Google Scholar 

  • Magain N, Miadlikowska J, Mueller O, Gajdeczka M, Truong C, Salamov AA, Dubchak I, Grigoriev IV, Goffinet B, Sérusiaux E, Lutzoni F (2017) Conserved genomic collinearity as a source of broadly applicable, fast evolving, markers to resolve species complexes: a case study using the lichen-forming genus Peltigera section Polydactylon. Mol Phylogenet Evol 117:10–29

    Article  PubMed  Google Scholar 

  • Maharachchikumbura SSN, Guo LD, Chukeatirote E, Bahkali AH, Hyde KD (2011) Pestalotiopsis - morphology, phylogeny, biochemistry and diversity. Fungal Divers 50:167–187

    Article  Google Scholar 

  • Mallet J (1995) A species definition for the Modern Synthesis. Trends Ecol Evol 10:294–299

    Article  CAS  PubMed  Google Scholar 

  • Manawasinghe IS, Dissanayake AJ, Li X, Liu M, Wanasinghe DN, Xu J, Zhao W, Zhang W, Zhou Y, Hyde KD, Brooks S (2019) High genetic diversity and species complexity of Diaporthe associated with grapevine dieback in China. Front Microbiol 10:1936

    Article  PubMed Central  PubMed  Google Scholar 

  • Mark K, Saag L, Leavitt SD, Will-Wolf S, Nelsen MP, Tõrra T, Saag A, Randlane T, Lumbsch HT (2016) Evaluation of traditionally circumscribed species in the lichen-forming genus Usnea, section Usnea (Parmeliaceae, Ascomycota) using a six-locus dataset. Org Divers Evol 16:497–524

    Article  Google Scholar 

  • Matute DR, Sepúlveda VE (2019) Fungal species boundaries in the genomics era. Fungal Genet Biol 131:103249

    Article  PubMed Central  PubMed  Google Scholar 

  • Mayden RL (1997) A hierarchy of species concepts: the denouement in the saga of the species problem. In: Claridge MF, Dawah HA, Wilson MR (eds) Species: the units of diversity. Chapman & Hall, New York, pp 381–423

    Google Scholar 

  • Mayr E (1982) The growth of biological thought. Harvard University Press, Cambridge, MA

    Google Scholar 

  • Mayr E (1963) Animal species and evolution. Harvard University Press, Harvard, MA

    Book  Google Scholar 

  • Mayr E (1942) Systematics and the origin of species. Columbia University Press, New York

    Google Scholar 

  • Mayr E, Provine WB (1998) The evolutionary synthesis. Harvard University Press, Harvard, MA

    Google Scholar 

  • McCarthy CG, Fitzpatrick DA (2017) Phylogenomic reconstruction of the oomycete phylogeny derived from 37 genomes. Msphere 2:e00095-e117

    Article  PubMed Central  PubMed  Google Scholar 

  • Meier R, Willmann R (2000) The Hennigian species concept. In: Wheeler QD, Meier R (eds) Species concepts and phylogenetic theory: a debate. Columbia University Press, New York, pp 30–43

    Google Scholar 

  • Meier R, Shiyang K, Vaidya G, Ng PK (2006) DNA barcoding and taxonomy in Diptera: a tale of high intraspecific variability and low identification success. Syst Biol 55:715–728

    Article  PubMed  Google Scholar 

  • Menardo F, Praz CR, Wyder S, Ben-David R, Bourras S, Matsumae H, McNally KE, Parlange F, Riba A, Roffler S, Schaefer LK (2016) Hybridization of powdery mildew strains gives rise to pathogens on novel agricultural crop species. Nat Genet 48:201–205

    Article  CAS  PubMed  Google Scholar 

  • Menkis A, Urbina H, James TY, Rosling A (2014) Archaeorhizomyces borealis sp. nov. and a sequence-based classification of related soil fungal species. Fungal Biol 118:943–955

    Article  CAS  PubMed  Google Scholar 

  • Michener CD (1970) Diverse approaches to systematics. Evol Biol 4:1–38

    Google Scholar 

  • Milgroom MG (2017) Population biology of plant pathogens: genetics, ecology, and evolution. American Phytopathological Society, St. Paul, Minnesota

    Book  Google Scholar 

  • Millanes AM, Truong C, Westberg M, Diederich P, Wedin M (2014) Host switching promotes diversity in host-specialized mycoparasitic fungi: uncoupled evolution in the Biatoropsis-Usnea system. Evolution 68:1576–1593

    Article  CAS  PubMed  Google Scholar 

  • Mishler BD, Theriot E (2000) The phylogenetic species concept (sensu Mishler and Theriot): Monophyly, apomorphy, and phylogenetic species concepts. Species Concepts and Phylogenetic Theory: A Debate. Columbia University Press, New York, pp 44–54

    Google Scholar 

  • Moon CD, Miles CO, Jarlfors U, Schardl CL (2002) The evolutionary origins of three new Neotyphodium endophyte species from grasses indigenous to the Southern Hemisphere. Mycologia 94:694–711

    Article  CAS  PubMed  Google Scholar 

  • Morin E, Miyauchi S, San Clemente H, Chen ECH, Pelin A, de la Providencia I, Ndikumana S, Beaudet D, Hainaut M, Drula E, Kuo A, Tang N, Roy S, Viala J, Henrissat B, Grigoriev IV, Corradi N, Roux C, Martin FM (2019) Comparative genomics of Rhizophagus irregularis, R. cerebriforme, R. diaphanus and Gigaspora rosea highlights specific genetic features in Glomeromycotina. New Phytol 222:1584–1598

    Article  CAS  PubMed  Google Scholar 

  • Moritz C, Cicero C (2004) DNA barcoding: promise and pitfalls. PLoS Biol 2:1529–1531

    Article  CAS  Google Scholar 

  • Muggia L, Kopun T, Grube M (2017) Effects of growth media on the diversity of culturable fungi from lichens. Molecules 22:824

    Article  PubMed Central  CAS  Google Scholar 

  • Myers EA, Rodriguez-Robles JA, Denardo DF, Staub RE, Stropoli A, Ruane S, Burbrink FT (2013) Multilocus phylogeographic assessment of the California Mountain Kingsnake (Lampropeltis zonata) suggests alternative patterns of diversification for the California Floristic Province. Mol Ecol 22:5418–5429

    Article  CAS  PubMed  Google Scholar 

  • Naranjo-Ortiz MA, Gabaldón T (2019) Fungal evolution: diversity, taxonomy and phylogeny of the Fungi. Biol Rev Camb Philos Soc 94:2101–2137

    Article  PubMed Central  PubMed  Google Scholar 

  • Nguyen HDT, Sultana T, Kesanakurti P, Hambleton S (2019) Genome sequencing and comparison of five Tilletia species to identify candidate genes for the detection of regulated species infecting wheat. IMA Fungus 10:1–7

    Article  Google Scholar 

  • Nguyen HD, Jančič S, Meijer M, Tanney JB, Zalar P, Gunde-Cimerman N, Seifert KA (2015) Application of the phylogenetic species concept to Wallemia sebi from house dust and indoor air revealed by multi-locus genealogical concordance. PLoS ONE 10:e0120894

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Niemiller MR, Fitzpatrick BM, Miller BT (2008) Recent divergence with gene flow in Tennessee cave salamanders (Plethodontidae: Gyrinophilus) inferred from gene genealogies. Mol Ecol 17:2258–2275

    Article  CAS  PubMed  Google Scholar 

  • Nilsson RH, Hyde KD, Pawlowska J, Ryberg M, Tedersoo L, Aas AB, Alias SA, Alves A, Anderson CL, Antonelli A, Arnold AE, Bahnmann B, Bahram M, Bengtsson-Palme J, Berlin A, Branco S, Chomnunti P, Dissanayake A, Drenkhan R, Friberg H, Frøslev TG, Halwachs B, Hartmann M, Henricot B, Jayawardena R, Jumpponen A, Kauserud H, Koskela S, Kulik T, Liimatainen K, Lindahl BD, Lindner D, Liu JK, Maharachchikumbura S, Manamgoda D, Martinsson S, Neves MA, Niskanen T, Nylinder S, Pereira OL, Pinho DB, Porter TM, Queloz V, Riit T, Sánchez-García M, de Sousa F, Stefańczyk E, Tadych M, Takamatsu S, Tian Q, Udayanga D, Unterseher M, Wang Z, Wikee S, Yan J, Larsson E, Larsson KH, Kõljalg U, Abarenkovet K (2014) Improving ITS sequence data for identification of plant pathogenic fungi. Fungal Divers 67:11–19

    Article  Google Scholar 

  • Nilsson RH, Tedersoo L, Abarenkov K, Ryberg M, Kristiansson E, Hartmann M, Schoch CL, Nylander JA, Bergsten J, Porter TM, Jumpponen A (2012) Five simple guidelines for establishing basic authenticity and reliability of newly generated fungal ITS sequences. MycoKeys 4:37–63

    Article  Google Scholar 

  • Nirenberg HI, O’Donnell K (1998) New Fusarium species and combinations within the Gibberella fujikuroi species complex. Mycologia 90:434–458

    Article  Google Scholar 

  • Nixon KC, Wheeler QD (1990) An amplification of the phylogenetic species concept. Cladistics 6:211–223

    Article  Google Scholar 

  • Noble R, Grogan H, Elliott T (1995) Variation in morphology, growth, and fructification of isolates in the Agaricus subfloccosus complex. Mycol Res 99:1453–1461

    Article  Google Scholar 

  • Nobrega MA, Pennacchio LA (2004) Comparative genomic analysis as a tool for biological discovery. J Physiol 554:31–39

    Article  CAS  PubMed  Google Scholar 

  • Norphanphoun C, Hongsanan S, Gentekaki E, Chen YJ, Kuo C-H, Hyde KD (2020) Differentiation of species complexes in Phyllosticta enables better species resolution. Mycosphere 11:2542–2628

    Article  Google Scholar 

  • Nosil P (2012) Ecological speciation. Oxford University Press, Oxford

    Book  Google Scholar 

  • Nosil P, Crespi BJ, Sandoval CP (2002) Host-plant adaptation drives the parallel evolution of reproductive isolation. Nature 417:440–443

    Article  CAS  PubMed  Google Scholar 

  • O’Meara BC (2010) New heuristic methods for joint species delimitation and species tree inference. Syst Biol 59:59–73

    Article  PubMed  Google Scholar 

  • Padial JM, Miralles A, De la Riva I, Vences M (2010) The integrative future of taxonomy. Front Zool 7:1–14

    Article  Google Scholar 

  • Papagianni M (2014) Characterization of fungal morphology using digital image analysis techniques. J Microb Biochem Technol 6:189–194

    Article  Google Scholar 

  • Park HJ, Jin G, Nakhleh L (2010) Bootstrap-based support of HGT inferred by maximum parsimony. BMC Evol Biol 10:131

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Parmasto E (1985) The species concept in Hymenochaetaceae (Fungi, Hymenomycetes). Proc Plant Sci 94:369

    Article  Google Scholar 

  • Parnmen S, Rangsiruji A, Mongkolsuk P, Boonpragob K, Nutakki A, Lumbsch HT (2012) Using phylogenetic and coalescent methods to understand the species diversity in the Cladia aggregata complex (Ascomycota, Lecanorales). PLoS ONE 7:e52245

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Petersen RH (1995) Contributions of mating studies to systematics of Agaricales. Can J Bot 73:S831–S842

    Article  Google Scholar 

  • Petersen RH, Hughes KW (1999) Species and speciation in mushrooms: development of a species concept poses difficulties. Bioscience 49:440–452

    Article  Google Scholar 

  • Petersen RH, Hughes KW (1998) Mating systems in Omphalotus (Paxillaceae, Agaricales). Plant Syst Evol 211:217–229

    Article  Google Scholar 

  • Petersen RH, Hughes KW (1993) Intercontinental interbreeding collections of Pleurotus pulmonarius, with notes on P. ostreatus and other species. Sydowia 45:139–152

    Google Scholar 

  • Phillips MJ, Delsuc F, Penny D (2004) Genome-scale phylogeny and the detection of systematic biases. Mol Biol Evol 21:1455–1458

    Article  CAS  PubMed  Google Scholar 

  • Phukhamsakda C, McKenzie EH, Phillips AJ, Jones EG, Bhat DJ, Stadler M, Bhunjun CS, Wanasinghe DN, Thongbai B, Camporesi E, Ertz D, Jayawardena RS, Perera RH, Ekanayake AH, Tibpromma S, Doilom M, Xu J, Hyde KD (2020) Microfungi associated with Clematis (Ranunculaceae) with an integrated approach to delimiting species boundaries. Fungal Divers 102:1–203

    Article  Google Scholar 

  • Plazzi F, Ferrucci RR, Passamonti M (2010) Phylogenetic representativeness: a new method for evaluating taxon sampling in evolutionary studies. BMC Bioinf 11:209

    Article  CAS  Google Scholar 

  • Pons J, Barraclough TG, Gomez-Zurita J, Cardoso A, Duran DP, Hazell S, Kamoun S, Sumlin WD, Vogler AP (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Syst Biol 55:595–609

    Article  PubMed  Google Scholar 

  • Postaire B, Magalon H, Bourmaud CAF, Bruggemann JH (2016) Molecular species delimitation methods and population genetics data reveal extensive lineage diversity and cryptic species in Aglaopheniidae (Hydrozoa). Mol Phylogenet Evol 105:36–49

    Article  PubMed  Google Scholar 

  • Price TD (2007) Speciation in Birds. Woodbury, NY: Roberts and Company

  • Puillandre N, Lambert A, Brouillet S, Achaz G (2012) ABGD, Automatic Barcode Gap Discovery for primary species delimitation. Mol Ecol 21:1864–1877

    Article  CAS  PubMed  Google Scholar 

  • Quaedvlieg W, Binder M, Groenewald JZ, Summerell BA, Carnegie AJ, Burgess TI, Crous PW (2014) Introducing the consolidated species concept to resolve species in the Teratosphaeriaceae. Persoonia 33:1–40

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Ray J (1686) Historia plantarum: species hactenus editas aliasque insuper multas noviter inventas & descriptas complectens. In qua agitur primo de plantis in genere, earumque partibus, accidentibus & differentiis; deinde genera omnia tum summa tum subalterna ad species usque infi mas, notis suis certis & characteristicis definita, methodo naturae vestigiis insistente disponuntur, vol. 1, Typis Mariae Clark, prostant apud Henricum Faithorne, London, England

  • Regan CT (1925) Organic evolution 1. Nature 116:398–401

    Article  Google Scholar 

  • Regan CT (1926) The pediculate fishes of the suborder Ceratioidea. Dana Oceanogr Rep 2:1–45

    Google Scholar 

  • Rehner SA, Buckley E (2005) A Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: evidence for cryptic diversification and links to Cordyceps teleomorphs. Mycologia 97:84–98

    CAS  PubMed  Google Scholar 

  • Rhodes J, Desjardins CA, Sykes SM, Beale MA, Vanhove M, Sakthikumar S, Chen Y, Gujja S, Saif S, Chowdhary A, Lawson DJ, Ponzio V, Colombo AL, Meyer W, Engelthaler DM, Hagen F, Illnait-Zaragozi MT, Alanio A, Vreulink J-M, Heitman J, Perfect JR, Litvintseva AP, Bicanic T, Harrison TS, Fisher MC, Cuomo CA (2017) Tracing genetic exchange and biogeography of Cryptococcus neoformans at the global population level. Genetics 207:327–346

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Riley R, Charron P, Idnurm A, Farinelli L, Dalpé Y, Martin FM, Corradi N (2014) Extreme diversification of the mating type-high-mobility group (MATA-HMG) gene family in a plant-associated arbuscular mycorrhizal fungus. New Phytol 201:254–268

    Article  CAS  PubMed  Google Scholar 

  • Rokas A, Williams BL, King N, Carroll SB (2003) Genome-scale approaches to resolving incongruence in molecular phylogenies. Nature 425:798–804

    Article  CAS  PubMed  Google Scholar 

  • Rosen DE (1979) Fishes from the uplands and intermontane basins of Guatemala: Revisionary studies and comparative geography. Bull Amer Mus Nat Hist 162:267–376

    Google Scholar 

  • Rosendahl S (2008) Communities, populations and individuals of arbuscular mycorrhizal fungi. New Phytol 178:253–266

    Article  PubMed  Google Scholar 

  • Saccardo PA (1883) Sylloge Fungorum 2: i, 1–815, ii-lxix, 1–77. P.A. Saccardo, Padua

  • Samarakoon MC, Thongbai B, Hyde KD, Brönstrup M, Beutling U, Lambert C, Miller AN, Liu JK, Promputtha I, Stadler M (2020) Elucidation of the life cycle of the endophytic genus Muscodor and its transfer to Induratia in Induratiaceae fam. nov., based on a polyphasic taxonomic approach. Fungal Divers 101:177–210

    Article  Google Scholar 

  • Sanders KL, Malhotra A, Thorpe RS (2006) Combining molecular, morphological and ecological data to infer species boundaries in a cryptic tropical pit viper. Biol J Linn Soc 87:343–364

    Article  Google Scholar 

  • Satler JD, Carstens BC, Hedin M (2013) Multilocus species delimitation in a complex of morphologically conserved trapdoor spiders (Mygalomorphae, Antrodiaetidae, Aliatypus). Syst Biol 62:805–823

  • Schardl CL (2001) Epichloë festucae and related mutualistic symbionts of grasses. Fungal Genet Biol 33:69–82

    Article  CAS  PubMed  Google Scholar 

  • Schardl CL, Craven KD (2003) Interspecific hybridization in plant-associated fungi and Oomycetes: a review. Mol Ecol 12:2861–2873

    Article  CAS  PubMed  Google Scholar 

  • Schardl CL, Leuchtmann A, Tsai H-F, Collett MA, Watt DM, Scott DB (1994) Origin of a fungal symbiont of perennial ryegrass by interspecific hybridization of a mutualist with the ryegrass choke pathogen, Epichloë typhina. Genetics 136:1307–1317

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Schubert K, Groenewald JZ, Braun U, Dijksterhuis J, Starink M, Hill CF, Zalar P, De Hoog GS, Crous PW (2007) Biodiversity in the Cladosporium herbarum complex (Davidiellaceae, Capnodiales), with standardisation of methods for Cladosporium taxonomy and diagnostics. Stud Mycol 58:105–156

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Seifert KA, Rossman AY (2010) How to describe a new fungal species. IMA Fungus 1:109–111

    Article  PubMed Central  PubMed  Google Scholar 

  • Senwanna C, Wanasinghe DN, Bulgakov TS, Wang Y, Bhat DJ, Tang AMC et al (2019) Towards a natural classification of Dothidotthia and Thyrostroma in Dothidotthiaceae (Pleosporineae, Pleosporales). Mycosphere 10:701–738

    Article  Google Scholar 

  • Sepúlveda VE, Márquez R, Turissini DA, Goldman WE, Matute DR (2017) Genome sequences reveal cryptic speciation in the human pathogen Histoplasma capsulatum. Mbio 8:e01339-e1417

    Article  PubMed Central  PubMed  Google Scholar 

  • Seto K, Degawa Y (2018) Collimyces mutans gen. et sp. nov. (Rhizophydiales, Collimycetaceae fam. nov.), a new chytrid parasite of Microglena (Volvocales, clade Monadinia). Protist 169:507–520

    Article  PubMed  Google Scholar 

  • Seto K, Matsuzawa T, Kuno H, Kagami M (2020) Morphology, ultrastructure, and molecular phylogeny of Aphelidium collabens sp. nov. (Aphelida), a parasitoid of a green alga Coccomyxa sp. Protist 171:125728

    Article  CAS  PubMed  Google Scholar 

  • Shen XX, Zhou X, Kominek J, Kurtzman CP, Hittinger CT, Rokas A (2016) Reconstructing the backbone of the Saccharomycotina yeast phylogeny using genome-scale data. G3 6:3927–3939

    Article  PubMed Central  PubMed  Google Scholar 

  • Simpson GG (1951) The species concept. Evolution 5:285–298

    Article  Google Scholar 

  • Singh G, Dal Grande F, Divakar PK, Otte J, Leavitt SD, Szczepanska K, Crespo A, Rico VJ, Aptroot A, da Silva Caceres ME, Lumbsch HT (2015) Coalescent-based species delimitation approach uncovers high cryptic diversity in the cosmopolitan lichen-forming fungal genus Protoparmelia (Lecanorales, Ascomycota). PLoS ONE 10:0124625

    Google Scholar 

  • Sobel JM, Chen GF, Watt LR, Schemske DW (2010) The biology of speciation. Evolution 64:295–315

    Article  PubMed  Google Scholar 

  • Sokal RR, Crovello TJ (1970) The biological species concept: a critical evaluation. Am Nat 104:127–153

    Article  Google Scholar 

  • Spatafora JW, Chang Y, Benny GL, Lazarus K, Smith ME, Berbee ML, Bonito G, Corradi N, Grigoriev I, Gryganskyi A, James TY (2016) A phylum-level phylogenetic classification of zygomycete fungi based on genome-scale data. Mycologia 108:1028–1046

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Stace CA (1989) Plant Taxonomy and Biosystematics, 2nd edn. Edward Arnold, a division of Hoddoer and Stoughton, London

    Google Scholar 

  • Steele PR, Pires JC (2011) Biodiversity assessment: State-of-the-art techniques in phylogenomics and species identification. Am J Bot 98:415–425

    Article  PubMed  Google Scholar 

  • Steenwyk JL, Shen X-X, Lind AL, Goldman GH, Rokas A (2019) A robust phylogenomic time tree for biotechnologically and medically important fungi in the genera Aspergillus and Penicillium. Mbio 10:e00925-e1019

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Stenlid J, Karlsson J-O (1991) Partial intersterility in Heterobasidion annosum. Mycol Res 95:1153–1159

    Article  Google Scholar 

  • Stewart JE, Timmer LW, Lawrence CB, Pryor BM, Peever TL (2014) Discord between morphological and phylogenetic species boundaries: incomplete lineage sorting and recombination results in fuzzy species boundaries in an asexual fungal pathogen. BMC Evol Biol 14:38

    Article  PubMed Central  PubMed  Google Scholar 

  • Stielow JB, Lévesque CA, Seifert KA, Meyer W, Iriny L, Smits D et al (2015) One fungus which genes? Development and assessment of universal primers for potential secondary fungal DNA barcodes. Persoonia 35:242–263

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Stockdale PM (1961) Nannizzia incurvata gen. nov., sp. nov., a perfect state of Microsporum gypseum (Bodin) Guiart et Grigorakis. Sabouraudia 1:41–48

    Article  Google Scholar 

  • Stukenbrock EH (2013) Evolution, selection and isolation: a genomic view of speciation in fungal plant pathogens. New Phytol 199:895–907

    Article  PubMed  Google Scholar 

  • Stukenbrock EH (2016) The role of hybridization in the evolution and emergence of new fungal plant pathogens. Phytopathology 106:104–112

    Article  CAS  PubMed  Google Scholar 

  • Stukenbrock EH, McDonald BA (2009) Population genetics of fungal and oomycete effectors involved in gene-for-gene interactions. Mol Plant Microbe Interact 22:371–380

    Article  CAS  PubMed  Google Scholar 

  • Stukenbrock EH, Quaedvlieg W, Javan-Nikhah M, Zala M, Crous PW, McDonald BA (2012a) Zymoseptoria ardabiliae and Z. pseudotritici, two progenitor species of the septoria tritici leaf blotch fungus Z. tritici (synonym: Mycosphaerella graminicola). Mycologia 104:1397–1407

    Article  PubMed  Google Scholar 

  • Stukenbrock EH, Christiansen FB, Hansen TT, Dutheil JY, Schierup MH (2012b) Fusion of two divergent fungal individuals led to the recent emergence of a unique widespread pathogen species. Proc Natl Acad Sci USA 109:10954–10959

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Stukenbrock EH, Banke S, Javan-Nikkhah M, McDonald BA (2007) Origin and domestication of the fungal wheat pathogen Mycosphaerella graminicola via sympatric speciation. Mol Biol Evol 24:398–411

    Article  CAS  PubMed  Google Scholar 

  • Sukumaran J, Knowles LL (2017) Multispecies coalescent delimits structure, not species. Proc Natl Acad Sci 114:1607–1612

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Sullivan DJ, Westerneng TJ, Haynes KA, Bennett DE, Coleman DC (1995) Candida dubliniensis sp. nov.: phenotypic and molecular characterization of a novel species associated with oral candidosis in HIV-infected individuals. Microbiology 141:1507–1521

    Article  CAS  PubMed  Google Scholar 

  • Summerell BA, Laurence MH, Liew EC, Leslie JF (2010) Biogeography and phylogeography of Fusarium: a review. Fungal Divers 44:3–13

  • Sun S, Xu J (2007) Genetic analyses of a hybrid cross between serotypes A and D strains of the human pathogenic fungus Cryptococcus neoformans. Genetics 177:1475–1486

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Tang AMC, Jeewon R, Hyde KD (2007) Phylogenetic relationships of Nemania plumbea sp. nov. and related taxa based on ribosomal ITS and RPB2 sequences. Fungal Biol 111:392–402

    CAS  Google Scholar 

  • Taylor HR, Harris WE (2012) An emergent science on the brink of irrelevance: a review of the past 8 years of DNA barcoding. Mol Ecol Resour 12:377–388

    Article  CAS  PubMed  Google Scholar 

  • Taylor JW, Turner E, Townsend JP, Dettman JR, Jacobson D (2006) Eukaryotic microbes, species recognition and the geographic limits of species: examples from the kingdom fungi. Philos Trans R Soc B 361:1947–1963

    Article  Google Scholar 

  • Taylor J, Jacobson D, Kroken S, Kasuga T, Geiser D, Hibbett D, Fisher M (2000) Phylogenetic species recognition and species concepts in fungi. Fungal Genet Biol 31:21–32

    Article  CAS  PubMed  Google Scholar 

  • Treindl AD, Leuchtmann A (2019) Assortative mating in sympatric ascomycete fungi revealed by experimental fertilizations. Fungal Biol 123:676–686

    Article  PubMed  Google Scholar 

  • Trejo D, Guzmán G, Lara L, Zulueta R, Palenzuela J, Sánchez-Castro I, Silva GA, Sieverding E, Oehl F (2015) Morphology and phylogeny of Acaulospora foveata (Glomeromycetes) from Mexico. Sydowia 67:119–126

    Google Scholar 

  • Trudell SA, Xu J, Saar I, Justo A, Cifuentes J (2017) North American matsutake: names clarified and a new species described. Mycologia 109:379–390

    Article  PubMed  Google Scholar 

  • Tsai H-F, Liu J-S, Staben C, Christensen MJ, Latch GC, Siegel MR, Schardl CL (1994) Evolutionary diversification of fungal endophytes of tall fescue grass by hybridization with Epichloë species. Proc Natl Acad Sci USA 91:2542–2546

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Udayanga D, Castlebury LA, Rossman AY, Chukeatirote E, Hyde KD (2014a) Insights into the genus Diaporthe: phylogenetic species delimitation in the D. eres species complex. Fungal Divers 67:203–229

    Article  Google Scholar 

  • Udayanga D, Castlebury LA, Rossman AY, Hyde KD (2014b) Species limits in Diaporthe: molecular re-assessment of D. citri, D. cytosporella, D. cystosporella, D. foeniculina and D. rudis. Persoocnia 32:83–101

    Article  CAS  Google Scholar 

  • Udayanga D, Liu X, Crous PW, McKenzie EHC, Chukeatirote E, Hyde KD (2012) A multi-locus phylogenetic evaluation of Diaporthe (Phomopsis). Fungal Divers 56:157–171

    Article  Google Scholar 

  • Via S (2001) Sympatric speciation in animals: the ugly duckling grows up. Trends Ecol Evol 16:381–390

    Article  CAS  PubMed  Google Scholar 

  • Van Valen L (1976) Ecological species, multispecies, and Oaks. Taxon 25:233–239

    Article  Google Scholar 

  • Verkley GJ, Dukik K, Renfurm R, Göker M, Stielow JB (2014) Novel genera and species of coniothyrium-like fungi in Montagnulaceae (Ascomycota). Persoonia 32:25–51

  • Videira SIR, Groenewald JZ, Nakashima C, Braun U, Barreto RW, de Wit PJG, Crous PW (2017) Mycosphaerellaceae – Chaos or clarity? Stud Mycol 87:257–421

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Vijaykrishna D, Mukerji R, Smith GJD (2015) RNA virus reassortment: an evolutionary mechanism for host jumps and immune evasion. PLoS Pathog 11:e1004902

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Vilgalys R (1991) Speciation and species concepts in the Collybia dryophila complex. Mycologia 83:758–773

    Article  Google Scholar 

  • Vilgalys R, Sun BL (1994) Ancient and recent patterns of geographic speciation in the oyster mushroom Pleurotus revealed by phylogenetic analysis of ribosomal DNA sequences. Proc Natl Acad Sci USA 91:4599–4603

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Villamil J, Avila LJ, Morando M, Sites JW Jr, Leaché AD, Maneyro R, Camargo A (2019) Coalescent-based species delimitation in the sand lizards of the Liolaemus wiegmannii complex (Squamata: Liolaemidae). Mol Phyl Evol 138:89–101

    Article  Google Scholar 

  • Wallen RM, Perlin MH (2018) An overview of the function and maintenance of sexual reproduction in Dikaryotic fungi. Front Microbiol 9:503

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Wanasinghe DN, Hyde KD, Jeewon R, Crous PW, Wijayawardene NN, Jones EBG et al (2017) Phylogenetic revision of Camarosporium (Pleosporineae, Dothideomycetes) and allied genera. Stud Mycol 87:207–256

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Wang H, Xu Z, Gao L, Hao B (2009) A fungal phylogeny based on 82 complete genomes using the composition vector method. BMC Evol Biol 9:195

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Whiston E, Taylor JW (2016) Comparative phylogenomics of pathogenic and nonpathogenic species. G3 6:235–244

    Article  Google Scholar 

  • White BP, Pilgrim EM, Boykin LM, Stein ED, Mazor RD (2014) Comparison of four species-delimitation methods applied to a DNA barcode data set of insect larvae for use in routine bioassessment. Freshw Sci 33:338–348

    Article  Google Scholar 

  • Willis KJ (2018) State of the world’s fungi 2018. Report. Royal Botanic Gardens, Kew

    Google Scholar 

  • Wu G, Li YC, Zhu XT, Zhao K, Han LH, Cui YY, Li F, Xu JP, Yang ZL (2016) One hundred noteworthy Boletes from China. Fungal Divers 81:25–188

    Article  Google Scholar 

  • Wulandari NF, To-Anun C, Hyde KD, Duong LM, De Gruyter J, Meffert JP, Groenewald JZ, Crous PW (2009) Phyllosticta citriasiana sp. nov., the cause of Citrus tan spot of Citrus maxima in Asia. Fungal Divers 34:23–39

    Google Scholar 

  • Xu J (2014) Genomics and fungal speciation: What can we expect in the clinical laboratory? Curr Clin Microbiol Rep 1:19–26

    Article  Google Scholar 

  • Xu J (2020) Fungal species concepts in the genomics era. Genome 63:459–468

    Article  PubMed  Google Scholar 

  • Xu B, Yang Z (2016) Challenges in species tree estimation under the multispecies coalescent model. Genetics 204:1353–1368

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Yang Z, Rannala B (2010) Bayesian species delimitation using multilocus sequence data. Proc Natl Acad Sci 107:9264–9269

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Yang ZL (2011) Molecular techniques revolutionize knowledge of basidiomycete evolution. Fungal Divers 50:47

    Article  Google Scholar 

  • Yang Q, Fan X-L, Guarnaccia V, Tian C-M (2018) High diversity of Diaporthe species associated with dieback diseases in China, with twelve new species described. MycoKeys 39:97–149

    Article  Google Scholar 

  • Yeung CK, Tsai PW, Chesser RT, Lin RC, Yao CT, Tian XH, Li SH (2011) Testing founder effect speciation: divergence population genetics of the spoonbills Platalea regia and Pl. minor (Threskiornithidae, Aves). Mol Biol Evol 28:473–482

    Article  CAS  PubMed  Google Scholar 

  • Yu Y, Than C, Degnan JH, Nakhleh L (2011) Coalescent histories on phylogenetic networks and detection of hybridization despite incomplete lineage sorting. Syst Biol 60:138–149

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Yu G, Rao D, Matsui M, Yang J (2017) Coalescent-based delimitation outperforms distance-based methods for delineating less divergent species: the case of Kurixalus odontotarsus species group. Sci Rep 7:16124

    Article  PubMed Central  CAS  PubMed  Google Scholar 

  • Zachos FE (2016) Species concepts in biology: historical development, theoretical foundations and practical relevance. Springer Nature, Dodrecht

    Book  Google Scholar 

  • Zang M (1987) Some new and noteworthy higher fungi from eastern Himalayas. Acta Bot Yunnanica 9:81–88

    Google Scholar 

  • Zeng NK, Cai Q, Yang ZL (2012) Corneroboletus, a new genus to accommodate the southeastern Asian Boletus indecorus. Mycologia 104:1420–1432

    Article  CAS  PubMed  Google Scholar 

  • Zhang W, Groenewald JZ, Lombard L, Schumacher RK, Phillips AJ, Crous PW (2021) Evaluating species in Botryosphaeriales. Persoonia 46:63–115

    Google Scholar 

  • Zhang J, Kapli P, Pavlidis P, Stamatakis A (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29:2869–2876

    Article  CAS  PubMed Central  PubMed  Google Scholar 

  • Zhang Y, Crous PW, Schoch CL, Bahkali AH, Guo LD, Hyde KD (2011) A molecular, morphological and ecological re-appraisal of Venturiales - a new order of Dothideomycetes. Fungal Divers 51:249–277

    Article  PubMed Central  PubMed  Google Scholar 

  • Zhang Y, Wang HK, Fournier J, Crous PW, Jeewon R, Pointing SB, Hyde KD (2009) Towards a phylogenetic clarification of Lophiostoma / Massarina and morphologically similar genera in the Pleosporales. Fungal Divers 38:225–251

    Google Scholar 

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Acknowledgements

Authors would like to thank the Thailand Research Fund entitled “Impact of climate change on fungal diversity and biogeography in the Greater Mekong Sub region” (grant number RDG6130001). Kevin D Hyde thanks Chiang Mai University for the award of Visiting Professor.

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Chethana, K.W.T., Manawasinghe, I.S., Hurdeal, V.G. et al. What are fungal species and how to delineate them?. Fungal Diversity 109, 1–25 (2021). https://doi.org/10.1007/s13225-021-00483-9

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