available online at www.studiesinmycology.org
STUDIES
IN
MYCOLOGY 98: 100116 (2021).
Fusarium: more than a node or a foot-shaped basal cell
Studies in Mycology
P.W. Crous1,2, L. Lombard1*, M. Sandoval-Denis1,3*, K.A. Seifert4, H.-J. Schroers5, P. Chaverri6,7, J. Gene8, J. Guarro8, Y. Hirooka9,
K. Bensch1, G.H.J. Kema2, S.C. Lamprecht10, L. Cai11,12, A.Y. Rossman13, M. Stadler14, R.C. Summerbell15,16, J.W. Taylor17,
S. Ploch18, C.M. Visagie19, N. Yilmaz19, J.C. Frisvad20, A.M. Abdel-Azeem21, J. Abdollahzadeh22, A. Abdolrasouli23,24, A. Akulov25,
J.F. Alberts26, J.P.M. Araújo27, H.A. Ariyawansa28, M. Bakhshi29, M. Bendiksby30,31, A. Ben Hadj Amor1, J.D.P. Bezerra32, T. Boekhout1,
M.P.S. C^amara33, M. Carbia34, G. Cardinali35, R.F. Casta~neda-Ruiz36, A. Celis37, V. Chaturvedi38, J. Collemare1, D. Croll39,
U. Damm40, C.A. Decock41, R.P. de Vries1, C.N. Ezekiel42, X.L. Fan43, N.B. Fernandez44,45, E. Gaya46, C.D. Gonzalez47, D. Gramaje48,
J.Z. Groenewald1, M. Grube49, M. Guevara-Suarez50, V.K. Gupta51,52, V. Guarnaccia53, A. Haddaji54, F. Hagen1, D. Haelewaters55,56,
K. Hansen57, A. Hashimoto58, M. Hernandez-Restrepo1, J. Houbraken1, V. Hubka59, K.D. Hyde60, T. Iturriaga61, R. Jeewon62,
Z. Jurjevic64, _I. Karalti65, L. Korsten66, E.E. Kuramae3,67, I. Kusan68, R. Labuda69, D.P. Lawrence70, H.B. Lee71,
P.R. Johnston63,
72
C. Lechat , H.Y. Li73, Y.A. Litovka74,75, S.S.N. Maharachchikumbura76, Y. Marin-Felix14, B. Matio Kemkuignou14, N. Matocec68,
A.R. McTaggart77, P. Mlcoch78, L. Mugnai79, C. Nakashima80, R.H. Nilsson81, S.R. Noumeur82, I.N. Pavlov74,75, M.P. Peralta83,
A.J.L. Phillips84, J.I. Pitt85, G. Polizzi86, W. Quaedvlieg87, K.C. Rajeshkumar88, S. Restrepo89, A. Rhaiem90, J. Robert54, V. Robert1,
A.M. Rodrigues91, C. Salgado-Salazar92, R.A. Samson1, A.C.S. Santos93, R.G. Shivas94, C.M. Souza-Motta93, G.Y. Sun95,
W.J. Swart96, S. Szoke54, Y.P. Tan94,97, J.E. Taylor98, P.W.J. Taylor99, P.V. Tiago93, K.Z. Vaczy100, N. van de Wiele54,
N.A. van der Merwe19, G.J.M. Verkley1, W.A.S. Vieira33, A. Vizzini101, B.S. Weir63, N.N. Wijayawardene102, J.W. Xia103,
M.J. Ya~nez-Morales104, A. Yurkov105, J.C. Zamora106, R. Zare29, C.L. Zhang107, and M. Thines18,108,109
1
Westerdijk Fungal Biodiversity Institute, 3508 AD, Utrecht, the Netherlands; 2Wageningen University and Research Centre (WUR), Laboratory of Phytopathology,
Droevendaalsesteeg 1, 6708 PB, Wageningen, the Netherlands; 3Netherlands Institute of Ecology (NIOO-KNAW), Department of Microbial Ecology,
Droevendaalsesteeg 10, 6708 PB, Wageningen, the Netherlands; 4Department of Biology, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario, K1S 5B6,
Canada; 5Plant Protection Department, Agricultural Institute of Slovenia, Hacquetova ulica 17, 1000, Ljubljana, Slovenia; 6Department of Plant Science and
Landscape Architecture, University of Maryland, College Park, MD, USA; 7Escuela de Biología and Centro de Investigaciones en Productos Naturales, Universidad
de Costa Rica, San Pedro, Costa Rica; 8Unitat de Micologia, Facultat de Medicina i Ciencies de la Salut i Institut d’Investigacio Sanitaria Pere Virgili (IISPV),
Universitat Rovira i Virgili, 43201, Reus, Spain; 9Department of Clinical Plant Science, Faculty of Bioscience, Hosei University, 3-7-2 Kajino-cho, Koganei, Tokyo,
184-8584, Japan; 10ARC-Plant Health and Protection, Private Bag X5017, Stellenbosch, 7599, Western Cape, South Africa; 11State Key Laboratory of Mycology,
Institute of Microbiology, Chinese Academy of Sciences, Beijing, 100101, China; 12University of Chinese Academy of Sciences, Beijing, 100049, China;
13
Department of Botany & Plant Pathology, Oregon State University, Corvallis, OR, 97330, USA; 14Department of Microbial Drugs, Helmholtz Centre for Infection
Research GmbH (HZI), Inhoffenstrasse 7, 38124 Braunschweig, Germany; 15Sporometrics, Toronto, ON, Canada; 16Dalla Lana School of Public Health, University
of Toronto, Toronto, ON, Canada; 17Plant and Microbial Biology, 111 Koshland Hall, University of California, Berkeley, CA, 94720-3102, USA; 18Senckenberg
Biodiversity and Climate Research Center, Senckenberganlage 25, D-60325, Frankfurt am Main, Germany; 19Department of Biochemistry, Genetics and
Microbiology, Forestry and Agricultural Biotechnology Institute (FABI), Faculty of Natural and Agricultural Sciences, University of Pretoria, P. Bag X20, Hatfield,
0028, Pretoria, South Africa; 20Department of Biotechnology and Biomedicine, DTU-Bioengineering, Technical University of Denmark, 2800, Kongens Lyngby,
Denmark; 21Systematic Mycology Lab., Botany and Microbiology Department, Faculty of Science, Suez Canal University, Ismailia, 41522, Egypt; 22Department of
Plant Protection, Faculty of Agriculture, University of Kurdistan, P.O. Box 416, Sanandaj, Iran; 23Department of Medical Microbiology, King's College Hospital,
London, UK; 24Department of Infectious Diseases, Imperial College London, London, UK; 25Department of Mycology and Plant Resistance, V. N. Karazin Kharkiv
National University, Maidan Svobody 4, 61022, Kharkiv, Ukraine; 26Department of Food Science and Technology, Cape Peninsula University of Technology, P.O.
Box 1906, Bellville, 7535, South Africa; 27School of Forest Resources and Conservation, University of Florida, Gainesville, FL, USA; 28Department of Plant
Pathology and Microbiology, College of Bio-Resources and Agriculture, National Taiwan University, No.1, Sec.4, Roosevelt Road, Taipei, 106, Taiwan, ROC;
29
Iranian Research Institute of Plant Protection, Agricultural Research, Education and Extension Organization (AREEO), P.O. Box 19395-1454, Tehran, Iran;
30
Natural History Museum, University of Oslo, Norway; 31Department of Natural History, NTNU University Museum, Trondheim, Norway; 32Setor de Micologia/
Departamento de Bioci^encias e Tecnologia, Instituto de Patologia Tropical e Saúde Pública, Rua 235 - s/n – Setor Universitario - CEP: 74605-050, Universidade
Federal de Goias/Federal University of Goias, Goi^ania, Brazil; 33Departamento de Agronomia, Universidade Federal Rural de Pernambuco, Recife, 52171-900, PE,
Brazil; 34Departamento de Parasitología y Micología, Instituto de Higiene, Facultad de Medicina – Universidad de la República, Av. A. Navarro 3051, Montevideo,
Uruguay; 35Department of Pharmaceutical Science, University of Perugia, Via Borgo 20 Giugno, 74 Perugia, Italy; 36Instituto de Investigaciones Fundamentales en
Agricultura Tropical Alejandro de Humboldt (INIFAT), Academico Titular de la Academia de Ciencias de, Cuba; 37Grupo de Investigacion Celular y Molecular de
Microorganismos Patogenos (CeMoP), Departamento de Ciencias Biologicas, Universidad de Los Andes, Bogota, 111711, Colombia; 38Mycology Laboratory, New
York State Department of Health Wadsworth Center, Albany, NY, USA; 39Laboratory of Evolutionary Genetics, Institute of Biology, University of Neuchatel, CH-2000,
Neuchatel, Switzerland; 40Senckenberg Museum of Natural History Görlitz, PF 300 154, 02806, Görlitz, Germany; 41Mycotheque de l'Universite catholique de
Louvain (MUCL, BCCMTM), Earth and Life Institute – ELIM – Mycology, Universite catholique de Louvain, Croix du Sud 2 bte L7.05.06, B-1348, Louvain-laNeuve, Belgium; 42Department of Microbiology, Babcock University, Ilishan Remo, Ogun State, Nigeria; 43The Key Laboratory for Silviculture and Conservation of
Ministry of Education, Beijing Forestry University, Beijing, 100083, China; 44Laboratorio de Micología Clínica, Hospital de Clínicas, Universidad de Buenos Aires,
Buenos Aires, Argentina; 45Facultad de Farmacia y Bioquímica, Universidad de Buenos Aires, Buenos Aires, Argentina; 46Royal Botanic Gardens, Kew, Richmond,
Surrey, TW9 3DS, UK; 47Laboratorio de Salud de Bosques y Ecosistemas, Instituto de Conservacion, Biodiversidad y Territorio, Facultad de Ciencias Forestales y
Recursos Naturales, Universidad Austral de Chile, casilla 567, Valdivia, Chile; 48Institute of Grapevine and Wine Sciences (ICVV), Spanish National Research
Council (CSIC)-University of La Rioja-Government of La Rioja, Logro~no, 26007, Spain; 49Institut für Biologie, Karl-Franzens-Universit€at Graz, Holteigasse 6, 8010,
Graz, Austria; 50Applied genomics research group, Universidad de los Andes, Cr 1 # 18 a 12, Bogota, Colombia; 51Center for Safe and Improved Food, Scotland's
Rural College (SRUC), Kings Buildings, West Mains Road, Edinburgh, EH9 3JG, UK; 52Biorefining and Advanced Materials Research Center, Scotland's Rural
College (SRUC), Kings Buildings, West Mains Road, Edinburgh, EH9 3JG, UK; 53Department of Agricultural, Forestry and Food Sciences (DISAFA), University of
Torino, Largo P. Braccini 2, 10095, Grugliasco, TO, Italy; 54BioAware, Hannut, Belgium; 55Research Group Mycology, Department of Biology, Ghent University, 35
e Budejovice, Czech Republic;
K.L. Ledeganckstraat, 9000, Ghent, Belgium; 56Faculty of Science, University of South Bohemia, Branisovska 31, 370 05, Cesk
57
Department of Botany, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05, Stockholm, Sweden; 58Microbe Division/Japan Collection of
Microorganisms RIKEN BioResource Research Center, 3-1-1 Koyadai, Tsukuba, Ibaraki, 305-0074, Japan; 59Department of Botany, Charles University in Prague,
Peer review under responsibility of Westerdijk Fungal Biodiversity Institute.
© 2021 Westerdijk Fungal Biodiversity Institute. Production and hosting by ELSEVIER B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
1
CROUS
ET AL.
Prague, Czech Republic; 60Center of Excellence in Fungal Research, Mae Fah Luang University, Chaing Rai, 57100, Thailand; 61Cornell University, 334 Plant Science
Building, Ithaca, NY, 14850, USA; 62Department of Health Sciences, Faculty of Medicine and Health Sciences, University of Mauritius, Reduit, Mauritius; 63Manaaki
Whenua Landcare Research, Private Bag 92170, Auckland, 1142, New Zealand; 64EMSL Analytical, Inc., 200 Route 130 North, Cinnaminson, NJ, 08077, USA;
65
Department of Nutrition and Dietetics, Faculty of Health Sciences, Yeditepe University, Turkey; 66Department of Plant and Soil Sciences, University of Pretoria, P.
Bag X20 Hatfield, Pretoria, 0002, South Africa; 67Institute of Environmental Biology, Ecology and Biodiversity, Utrecht University, 3584 CH, Utrecht, the Netherlands;
68
Laboratory for Biological Diversity, RuCer Boskovic Institute, Bijenicka cesta 54, HR-10000, Zagreb, Croatia; 69University of Veterinary Medicine, Vienna (VetMed),
Institute of Food Safety, Food Technology and Veterinary Public Health, Veterinaerplatz 1, 1210 Vienna and BiMM – Bioactive Microbial Metabolites group, 3430
Tulln a.d. Donau, Austria; 70University of California, Davis, One Shields Ave., Davis, CA, 95616, USA; 71Department of Agricultural Biological Chemistry, College of
Agriculture & Life Sciences, Chonnam National University, Yongbong-Dong 300, Buk-Gu, Gwangju, 61186, South Korea; 72Ascofrance, 64 route de Chize, 79360,
Villiers-en-Bois, France; 73The Key Laboratory of Molecular Biology of Crop Pathogens and Insects of Ministry of Agriculture, The Key Laboratory of Biology of
Crop Pathogens and Insects of Zhejiang Province, Institute of Biotechnology, Zhejiang University, 866 Yuhangtang Road, Hangzhou, 310058, China; 74V.N.
Sukachev Institute of Forest SB RAS, Laboratory of Reforestation, Mycology and Plant Pathology, Krasnoyarsk, 660036, Russia; 75Reshetnev Siberian State
University of Science and Technology, Department of Chemical Technology of Wood and Biotechnology, Krasnoyarsk, 660037, Russia; 76School of Life Science and
Technology, University of Electronic Science and Technology of China, Chengdu, 611731, China; 77Queensland Alliance for Agriculture and Food Innovation, The
University of Queensland, Ecosciences Precinct, G.P.O. Box 267, Brisbane, 4001, Australia; 78Department of Botany, Faculty of Science, Palacký University
Olomouc, Slechtitelů
27, CZ-783 71, Olomouc, Czech Republic; 79Department of Agricultural, Food, Environmental and Forestry Science and Technology (DAGRI),
Plant Pathology and Entomology section, University of Florence, P.le delle Cascine 28, 50144, Firenze, Italy; 80Graduate school of Bioresources, Mie University,
Kurima-machiya 1577, Tsu, Mie, 514-8507, Japan; 81Gothenburg Global Biodiversity Center at the Department of Biological and Environmental Sciences, University
of Gothenburg, Box 461, 405 30, Gothenburg, Sweden; 82Department of Microbiology and Biochemistry, Faculty of Natural and Life Sciences, University of Batna
2, Batna, 05000, Algeria; 83Laboratorio de Micodiversidad y Micoprospeccion, PROIMI-CONICET, Av. Belgrano y Pje. Caseros, Argentina; 84Universidade de
Lisboa, Faculdade de Ci^encias, Biosystems and Integrative Sciences Institute (BioISI), Campo Grande, 1749-016, Lisbon, Portugal; 85Microbial Screening
Technologies, 28 Percival Rd, Smithfield, NSW, 2164, Australia; 86Dipartimento di Agricoltura, Alimentazione e Ambiente, sez. Patologia vegetale, University of
Catania, Via S. Sofia 100, 95123 Catania, Italy; 87Phytopathology, Van Zanten Breeding B.V., Lavendelweg 15, 1435 EW, Rijsenhout, the Netherlands; 88National
Fungal Culture Collection of India (NFCCI), Biodiversity and Palaeobiology (Fungi) Group, Agharkar Research Institute, Pune, Maharashtra, 411 004, India;
89
Laboratory of Mycology and Phytopathology – (LAMFU), Department of Chemical and Food Engineering, Universidad de los Andes, Cr 1 # 18 a 12, Bogota,
Colombia; 90Plant Pathology and Population Genetics, Laboratory of Microorganisms, National Gene Bank, Tunisia; 91Laboratory of Emerging Fungal Pathogens,
Department of Microbiology, Immunology, and Parasitology, Discipline of Cellular Biology, Federal University of S~ao Paulo (UNIFESP), S~ao Paulo, 04023062, Brazil;
92
USDA-ARS Mycology & Nematology Genetic Diversity & Biology Laboratory, Bldg. 010A, Rm. 212, BARC-West, 10300 Baltimore Ave, Beltsville, MD, 20705,
USA; 93Departamento de Micologia Prof. Chaves Batista, Universidade Federal de Pernambuco, Centro de Bioci^encias, Cidade Universitaria, Av. Prof. Moraes
Rego, s/n, Recife, PE, CEP: 50670-901, Brazil; 94Centre for Crop Health, University of Southern Queensland, Toowoomba, 4350, Queensland, Australia; 95College
of Plant Protection, Northwest A&F University, Yangling, Shaanxi, China; 96Faculty of Natural and Agricultural Sciences, Department of Plant Sciences, University of
the Free State, P.O. Box 339, Bloemfontein, 9300, South Africa; 97Queensland Plant Pathology Herbarium, Department of Agriculture and Fisheries, Dutton Park,
Queensland, 4102, Australia; 98Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh, EH3 5LR, United Kingdom; 99Faculty of Veterinary and Agricultural
Sciences, The University of Melbourne, Parkville, VIC, 3010, Australia; 100Food and Wine Research Institute, Eszterhazy Karoly University, 6 Leanyka Street, H3300, Eger, Hungary; 101Department of Life Sciences and Systems Biology, University of Torino and Institute for Sustainable Plant Protection (IPSP-SS Turin),
C.N.R, Viale P.A. Mattioli, 25, I-10125, Torino, Italy; 102Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource
and Food Engineering, Qujing Normal University, Qujing, Yunnan, 655011, China; 103Shandong Provincial Key Laboratory for Biology of Vegetable Diseases and
Insect Pests, College of Plant Protection, Shandong Agricultural University, Taian, 271018, China; 104Fitosanidad, Colegio de Postgraduados-Campus Montecillo,
Montecillo-Texcoco, 56230 Edo. de Mexico, Mexico; 105Leibniz Institute DSMZ-German Collection of Microorganisms and Cell Cultures GmbH, Inhoffenstrasse 7 B,
38124, Braunschweig, Germany; 106Museum of Evolution, Uppsala University, Norbyv€agen 16, SE-752 36, Uppsala, Sweden; 107Ministry of Agriculture Key
Laboratory of Molecular Biology of Crop Pathogens and Insects, Institute of Biotechnology, College of Agriculture and Biotechnology, Zhejiang University, No. 866
Yuhangtang Road, Hangzhou, 310058, China; 108Goethe-University Frankfurt am Main, Department of Biological Sciences, Institute of Ecology, Evolution and
Diversity, Max-von-Laue Str. 13, D-60438, Frankfurt am Main, Germany; 109LOEWE Centre for Translational Biodiversity Genomics, Georg-Voigt-Str. 14-16, D60325, Frankfurt am Main, Germany
*Correspondence: L. Lombard, l.lombard@wi.knaw.nl; M. Sandoval-Denis, m.sandoval@wi.knaw.nl
Abstract: Recent publications have argued that there are potentially serious consequences for researchers in recognising distinct genera in the terminal fusarioid clade
of the family Nectriaceae. Thus, an alternate hypothesis, namely a very broad concept of the genus Fusarium was proposed. In doing so, however, a significant body of
data that supports distinct genera in Nectriaceae based on morphology, biology, and phylogeny is disregarded. A DNA phylogeny based on 19 orthologous protein-coding
genes was presented to support a very broad concept of Fusarium at the F1 node in Nectriaceae. Here, we demonstrate that re-analyses of this dataset show that all 19
genes support the F3 node that represents Fusarium sensu stricto as defined by F. sambucinum (sexual morph synonym Gibberella pulicaris). The backbone of the
phylogeny is resolved by the concatenated alignment, but only six of the 19 genes fully support the F1 node, representing the broad circumscription of Fusarium.
Furthermore, a re-analysis of the concatenated dataset revealed alternate topologies in different phylogenetic algorithms, highlighting the deep divergence and unresolved placement of various Nectriaceae lineages proposed as members of Fusarium. Species of Fusarium s. str. are characterised by Gibberella sexual morphs,
asexual morphs with thin- or thick-walled macroconidia that have variously shaped apical and basal cells, and trichothecene mycotoxin production, which separates them
from other fusarioid genera. Here we show that the Wollenweber concept of Fusarium presently accounts for 20 segregate genera with clear-cut synapomorphic traits,
and that fusarioid macroconidia represent a character that has been gained or lost multiple times throughout Nectriaceae. Thus, the very broad circumscription of
Fusarium is blurry and without apparent synapomorphies, and does not include all genera with fusarium-like macroconidia, which are spread throughout Nectriaceae
(e.g., Cosmosporella, Macroconia, Microcera). In this study four new genera are introduced, along with 18 new species and 16 new combinations. These names convey
information about relationships, morphology, and ecological preference that would otherwise be lost in a broader definition of Fusarium. To assist users to correctly
identify fusarioid genera and species, we introduce a new online identification database, Fusarioid-ID, accessible at www.fusarium.org. The database comprises partial
sequences from multiple genes commonly used to identify fusarioid taxa (act1, CaM, his3, rpb1, rpb2, tef1, tub2, ITS, and LSU). In this paper, we also present a
nomenclator of names that have been introduced in Fusarium up to January 2021 as well as their current status, types, and diagnostic DNA barcode data. In this study,
researchers from 46 countries, representing taxonomists, plant pathologists, medical mycologists, quarantine officials, regulatory agencies, and students, strongly support
the application and use of a more precisely delimited Fusarium (= Gibberella) concept to accommodate taxa from the robust monophyletic node F3 on the basis of a welldefined and unique combination of morphological and biochemical features. This F3 node includes, among others, species of the F. fujikuroi, F. incarnatum-equiseti, F.
oxysporum, and F. sambucinum species complexes, but not species of Bisifusarium [F. dimerum species complex (SC)], Cyanonectria (F. buxicola SC), Geejayessia
(F. staphyleae SC), Neocosmospora (F. solani SC) or Rectifusarium (F. ventricosum SC). The present study represents the first step to generating a new online
monograph of Fusarium and allied fusarioid genera (www.fusarium.org).
Key words: Multi-gene phylogeny, Mycotoxins, Nectriaceae, Neocosmospora, Novel taxa, Pathogen, Taxonomy.
2
FUSARIUM
REDELIMITED
Taxonomic novelties: New genera: Luteonectria Sand.-Den., L. Lombard, Schroers & Rossman, Nothofusarium Crous, Sand.-Den. & L. Lombard, Scolecofusarium L.
Lombard, Sand.-Den. & Crous, Setofusarium (Nirenberg & Samuels) Crous & Sand.-Den.; New species: Fusarium echinatum Sand.-Den. & G.J. Marais, Fusarium
lyarnte J.L. Walsh, Sangal., L.W. Burgess, E.C.Y. Liew & Summerell, Fusarium palustre W.H. Elmer & Marra, Fusarium prieskaense G.J. Marais & Sand.-Den.,
Fusarium werrikimbe J.L. Walsh, L.W. Burgess, E.C.Y. Liew & B.A. Summerell, Fusicolla quarantenae J.D.P. Bezerra, Sand.-Den., Crous & Souza-Motta, Fusicolla
meniscoidea L. Lombard & Sand.-Den., Fusicolla sporellula Sand.-Den. & L. Lombard, Macroconia bulbipes Crous & Sand.-Den., Macroconia phlogioides Sand.Den. & Crous, Neocosmospora epipeda Quaedvl. & Sand.-Den., Neocosmospora merkxiana Quaedvl. & Sand.-Den., Neocosmospora neerlandica Crous & Sand.Den., Neocosmospora nelsonii Crous & Sand.-Den., Neocosmospora pseudopisi Sand.-Den. & L. Lombard, Nothofusarium devonianum L. Lombard, Crous &
Sand.-Den., Stylonectria corniculata Gr€afenhan, Crous & Sand.-Den., Stylonectria hetmanica Akulov, Crous & Sand.-Den.; New combinations: Apiognomonia
platani (Lev.) L. Lombard, Calloria tremelloides (Grev.) L. Lombard, Cosmosporella cavisperma (Corda) Sand.-Den., L. Lombard & Crous, Cylindrodendrum
orthosporum (Sacc. & P. Syd.) L. Lombard, Dialonectria volutella (Ellis & Everh.) L. Lombard & Sand.-Den., Fusarium armeniacum (G.A. Forbes et al.) L.W.
Burgess & Summerell, Hymenella aurea (Corda) L. Lombard, Hymenella spermogoniopsis (Jul. Müll.) L. Lombard & Sand.-Den., Luteonectria albida (Rossman)
Sand.-Den. & L. Lombard, Luteonectria nematophila (Nirenberg & Hagedorn) Sand.-Den. & L. Lombard, Neocosmospora floridana (T. Aoki et al.) L. Lombard &
Sand.-Den., Neocosmospora obliquiseptata (T. Aoki et al.) L. Lombard & Sand.-Den., Neocosmospora rekana (Lynn & Marinc.) L. Lombard & Sand.-Den.,
Neocosmospora tuaranensis (T. Aoki et al.) L. Lombard & Sand.-Den., Scolecofusarium ciliatum (Link) L. Lombard, Sand.-Den. & Crous, Setofusarium setosum
(Samuels & Nirenberg) Sand.-Den. & Crous.; Epitypes (basionyms): Fusarium buharicum Jacz. ex Babajan & Teterevn.-Babajan, Fusarium cavispermum Corda,
Fusarium flocciferum Corda, Fusarium graminearum Schwabe, Fusarium heterosporum Nees & T. Nees, Fusarium redolens Wollenw., Fusarium reticulatum Mont.,
Fusarium scirpi Lambotte & Fautrey, Fusarium stilboides Wollenw., Fusarium xylarioides Steyaert, Fusisporium culmorum Wm.G. Sm., Fusisporium incarnatum
Roberge ex Desm., Selenosporium equiseti Corda, Sphaeria sanguinea var. cicatricum Berk., Sporotrichum poae Peck.; Lectotypes (basionyms): Atractium
pallidum Bonord., Cephalosporium sacchari E.J. Butler, Fusarium aeruginosum Delacr., Fusarium agaricorum Sarrazin, Fusarium albidoviolaceum Dasz., Fusarium
aleyrodis Petch, Fusarium amentorum Lacroix, Fusarium annuum Leonian, Fusarium arcuatum Berk. & M.A. Curtis, Fusarium aridum O.A. Pratt, Fusarium
arthrosporioides Sherb., Fusarium asparagi Delacr., Fusarium batatas Wollenw., Fusarium biforme Sherb., Fusarium cactacearum Pasin. & Buzz.-Trav., Fusarium
cacti-maxonii Pasin. & Buzz.-Trav., Fusarium caudatum Wollenw., Fusarium cavispermum Corda, Fusarium cepae Hanzawa, Fusarium cesatii Rabenh., Fusarium
citriforme Jamal., Fusarium citrinum Wollenw., Fusarium citrulli Taubenh., Fusarium clavatum Sherb., Fusarium coccinellum Kalchbr., Fusarium cromyophthoron
Sideris, Fusarium cucurbitae Taubenh., Fusarium cuneiforme Sherb., Fusarium delacroixii Sacc., Fusarium dimerum var. nectrioides Wollenw., Fusarium epicoccum
McAlpine, Fusarium eucheliae Sartory, R. Sartory & J. Mey., Fusarium fissum Peyl, Fusarium flocciferum Corda, Fusarium gemmiperda Aderh., Fusarium
genevense Dasz., Fusarium graminearum Schwabe, Fusarium graminum Corda, Fusarium heterosporioides Fautrey, Fusarium heterosporum Nees & T. Nees,
Fusarium idahoanum O.A. Pratt, Fusarium juruanum Henn., Fusarium lanceolatum O.A. Pratt, Fusarium lateritium Nees, Fusarium loncheceras Sideris, Fusarium
malvacearum Taubenh., Fusarium martii f. phaseoli Burkh., Fusarium muentzii Delacr., Fusarium nigrum O.A. Pratt, Fusarium oxysporum var. asclerotium Sherb.,
Fusarium palczewskii Jacz., Fusarium polymorphum Matr., Fusarium poolense Taubenh., Fusarium prunorum McAlpine, Fusarium pusillum Wollenw., Fusarium
putrefaciens Osterw., Fusarium redolens Wollenw., Fusarium reticulatum Mont., Fusarium rhizochromatistes Sideris, Fusarium rhizophilum Corda, Fusarium
rhodellum McAlpine, Fusarium roesleri Thüm., Fusarium rostratum Appel & Wollenw., Fusarium rubiginosum Appel & Wollenw., Fusarium rubrum Parav., Fusarium
samoense Gehrm., Fusarium scirpi Lambotte & Fautrey, Fusarium secalis Jacz., Fusarium spinaciae Hungerf., Fusarium sporotrichioides Sherb., Fusarium stercoris
Fuckel, Fusarium stilboides Wollenw., Fusarium stillatum De Not. ex Sacc., Fusarium sublunatum Reinking, Fusarium succisae Schröt. ex Sacc., Fusarium
tabacivorum Delacr., Fusarium trichothecioides Wollenw., Fusarium tritici Liebman, Fusarium tuberivorum Wilcox & G.K. Link, Fusarium tumidum var. humi Reinking,
Fusarium ustilaginis Kellerm. & Swingle, Fusarium viticola Thüm., Fusarium willkommii Lindau, Fusarium xylarioides Steyaert, Fusarium zygopetali Delacr.,
Fusisporium andropogonis Cooke ex Thüm., Fusisporium anthophilum A. Braun, Fusisporium arundinis Corda, Fusisporium clypeaster Corda, Fusisporium
culmorum Wm.G. Sm., Fusisporium didymum Harting, Fusisporium elasticae Thüm., Fusisporium episphaericum Cooke & Ellis, Fusisporium flavidum Bonord.,
Fusisporium hordei Wm.G. Sm., Fusisporium incarnatum Roberge ex Desm., Fusisporium lolii Wm.G. Sm., Fusisporium pandani Corda, Gibberella
phyllostachydicola W. Yamam., Menispora penicillata Harz, Selenosporium equiseti Corda, Selenosporium hippocastani Corda, Selenosporium urticearum Corda.,
Sphaeria sanguinea var. cicatricum Berk.; Neotypes (basionyms): Atractium ciliatum Link, Fusarium longipes Wollenw. & Reinking, Fusisporium avenaceum Fr.,
Selenosporium sarcochroum Desm.
Published online xxx; https://doi.org/10.1016/j.simyco.2021.100116.
INTRODUCTION
The relevance and impact of Fusarium (Ascomycota, Hypocreales, Nectriaceae) to humankind is substantial. Over the past
100 years, it has attracted considerable attention from scientists
as the extent of species diversity and the impact on agriculture
and human health became clear. After an initial period of discovery and cataloguing by 19th century naturalists, its taxonomy
became the target of research from a broad range of scientists,
that resulted in the emergence of distinct “schools” that promoted
different taxonomic approaches to fusarium-like organisms. With
the advent of an objective and reproducible framework for
phylogenetic relationships inferred from molecular phylogenetics,
it might have been expected that controversies would melt away,
and a stable, universally accepted taxonomy of Fusarium and its
species would emerge, but this does not yet appear to be the
case (Fig. 1). However, all scientists working with Fusarium
desire a stable taxonomic system, and all agree that taxonomic
changes should be made with the aim of promoting stability.
Recently, Geiser et al. (2021), largely in response to papers
published by Gr€afenhan et al. (2011), Schroers et al. (2011),
Lombard et al. (2015), and Sandoval-Denis et al. (2019), proposed a cladistic solution to redelimit a generic concept for
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Fusarium. The generic treatment of Fusarium by Geiser et al.
(2013, 2021), produced an ill-delimited genus without clear
synapomorphies, as fusarium-like macroconidia are strongly
polyphyletic within Nectriaceae and also occur outside their very
broadly circumscribed Fusarium concept. We argue that a narrower concept of genera with a clear, unique combination of
features is needed for the majority of fusarioid species.
Dual nomenclature and consensus on the use
of the generic name Fusarium
In accordance with the single-name system for fungi, that was
adopted at the International Botanical Congress, Melbourne
(IBCM) in 2011, we are in full agreement with Geiser et al. (2013,
2021) and O'Donnell et al. (2020) that the name Fusarium applies to any genus with a delimitation that includes the conserved
lectotype of the type species, F. sambucinum (sexual morph
synonym Gibberella pulicaris), as stated by Rossman et al.
(2013). Unfortunately, a single joint paper explaining the choice
of this name supported by the entire Fusarium community was
planned but failed because of the insistence of a subset of authors to adopt a broad generic concept.
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Fig. 1. Timeline summarising important events in the taxonomy and nomenclature of Fusarium and related taxa.
Taxonomy and nomenclature are different concepts, although
they are frequently confused, leading to misinterpretations.
Support for dual nomenclature ended at the IBCM in August
2011. The significance of 1 January 2013 was to ensure the
formal nomenclatural validity of newly proposed dual names
(new species or new combinations) that were in press or part of
studies about to be submitted for publication. These dates have
no significance for names proposed in a single name system,
which can be done at any time. Despite these technicalities,
virtually all members of the Fusarium community accept that
Fusarium must be used over the sexually-typified name Gibberella in the single name system, a recommendation included in
the proposed list of Protected Names submitted to the Nomenclature Committee for Fungi, the body with the authority to
recommend its formal acceptance (Kirk et al. 2013). However,
statements in Geiser et al. (2013) seem to reflect a confusion
about how the nomenclatural decision affected taxonomic
concepts.
The name Fusarium has never been at risk during the
nomenclatural transition, and the community support for its use in
a single name system is unanimous. We fully agree with Geiser
et al. (2013, 2021) and Rossman et al. (2013) that Fusarium
equals Gibberella. Fusarium will always be applied to the clade
that includes the type species of Fusarium, F. sambucinum,
which is the same fungus that also typifies Gibberella. In this
study, we show that the clade defined as Fusarium s. str.
(O'Donnell et al. 2013, as Gibberella; Geiser et al. 2013, as
Clade B) combines monophyly, morphology of sexual and
asexual morphs, and biochemical data in a coherent way that
can logically be recognised at the generic rank. Expanding the
concept of Fusarium to node F1 sensu Geiser et al. (2013, 2021)
4
results in the combination of several distinct genera and does not
resolve the issue of fusarium-like macroconidia in genera outside
their broad circumscription of Fusarium.
Phylogenetic structure and distribution of
fusarioid asexual morphs in Nectriaceae
(Hypocreales)
Gr€afenhan et al. (2011) and Schroers et al. (2011) presented a
phylogenetic overview of selected Nectriaceae based on combined analyses of two different genes, namely the commonly
employed and phylogenetically informative RNA polymerase II
second largest subunit (rpb2) and exon regions of the larger
subunit of ATP citrate lyase (acl1). The two papers were the first
to apply a single name system to fusarioid fungi (i.e., genera with
fusarium-like macroconidia), and were written along with others
(see Rossman & Seifert 2011) to promote discussions that
eventually led to changes to the International Code of Nomenclature for algae, fungi, and plants (ICNafp) (Turland et al. 2018).
The main focus of the Gr€afenhan et al. (2011) paper was to
deal with extraneous elements that had long been included in
Fusarium. These fungi had distinct phenotypic characters, such
as thin, collapsing perithecial walls, slow growing agar colonies
lacking aerial mycelium, or sparsely septate macroconidia. Users
of the Gerlach & Nirenberg (1982) and Nelson et al. (1983)
identification manuals may be familiar with some of these species, then called Fusarium aquaeductuum, F. coccophilum and
F. merismoides. There was evidence in the first papers on the
molecular phylogeny of Fusarium that these species did not
belong to Fusarium (e.g., see O'Donnell 1993). It was not until
FUSARIUM
the study by Gr€afenhan et al. (2011) that other genera in the
family, such as members of the Cylindrocarpon generic complex
(Chaverri et al. 2011), Calonectria (Liu et al. 2020), Tubercularia
(Hirooka et al. 2012), and minor genera such as Mariannaea,
Pseudonectria, and Volutella (also see Lombard et al. 2015)
were adequately sampled to yield generic-level resolution. The
phylograms showed the division of fusarioid taxa into two large
groups, which Gr€afenhan et al. (2011) called the Terminal
Fusarium Clade (abbreviated TFC by Geiser et al. 2013) and the
ill-delineated Basal Fusarium Clade (BFC) that contained several
of the genera noted above. A single-genus recognition for the
BFC was not feasible because of the great morphological, genetic, and ecological divergence among the sampled species.
The BFC included seven genera, each with their monophyly
strongly supported and more or less ecologically coherent.
Species with fusarioid conidia were reclassified in the phylogenetically redefined but previously described genera Atractium,
Cosmospora, Dialonectria, Fusicolla, Macroconia, Microcera,
and Stylonectria (Gr€afenhan et al. 2011, Schroers et al. 2011).
Geiser et al. (2013) accepted these segregate genera in the BFC
as distinct from the TFC, while correctly pointing out the
weak support values obtained for the phylogenetic backbone
of the tree. One consequence of the widespread occurrence of
macroconidia in the taxon sampling (fusarioid genera, cylindrocarpon-like genera, and Calonectria) was the suggestion that
especially the fusarioid macroconidium is a plesiomorphic
character (that is, an ancestral character) and had been lost in
some lineages in Nectriaceae (Gr€afenhan et al. 2011).
The second paper by Schroers et al. (2011) recovered similar
phylogenies as Gr€afenhan et al. (2011), but focused on the TFC,
supplementing this with a five-gene analysis of a particular
subclade within the TFC intended to delimit phylogenetic genera
and a few species. This demonstrated the monophyly of the
treated genera and resulted in the acceptance of the previously
described Cyanonectria (Samuels et al. 2009), as well as the
description of the genus Geejayessia. Again, Geiser et al. (2013)
correctly criticised the weakness of the backbone of the tree,
especially in the BFC. About 75 % of the phylogenetic signal in
the analysis came from one gene, rpb2. Schroers et al. (2011)
did not discuss the taxonomic fate of Neocosmospora (the
Fusarium solani species complex, FSSC), which was represented by only two species in their analysis, but was excluded
from Fusarium s. str.
The call for more genetic markers and even genome analysis
by Geiser et al. (2013), to better resolve the phylogenetic
backbone of the TFC was justified, but the increased number of
markers should have been matched by increased taxon sampling of all known genera of Nectriaceae, as taxon sampling is
equally important for inferring robust and meaningful phylogenies
(Zwickl & Hillis 2002, Heath et al. 2008).
Lombard et al. (2015) greatly expanded both the number of
genetic markers and the taxon sampling in order to explore the
generic boundaries across the Nectriaceae, including all genera
known from culture and many genera for which no DNA data was
previously available. A 10-gene phylogeny was inferred including
all the markers previously used by Gr€afenhan et al. (2011),
Schroers et al. (2011), Geiser et al. (2013), and O'Donnell et al.
(2013), plus nrDNA sequences and other markers of known
phylogenetic utility, namely actin (act1), beta-tubulin (tub2),
calmodulin (CaM), histone (his3), and the translation elongation
factor 1-α (tef1). From this, a phylogeny of the TFC overall
congruent to that presented by Gr€afenhan et al. (2011) and
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REDELIMITED
Geiser et al. (2013) was obtained. Importantly, the monophyly of
Albonectria, Cyanonectria, Geejayessia, Fusarium, and Neocosmospora was reaffirmed and a few early diverging lineages
previously included in the TFC were segregated into new
fusarioid genera i.e., Bisifusarium (formerly the F. dimerum
species complex) and Rectifusarium (formerly the F. ventricosum
species complex) (Lombard et al. 2015).
After nearly a hundred years of quandary, a modern revision
was published for Neocosmospora (Sandoval-Denis et al. 2019),
In this study, many unnamed phylogenetic species were
morphologically characterised and given Latin binomials, while
old names were resurrected, epitypified, and linked to DNA
barcodes.
Two recent publications by O'Donnell et al. (2020) and Geiser
et al. (2021) argued for the broad Fusarium concept of Geiser
et al. (2013). Both papers present very similar phylogenetic
analyses, relying on 19 genes, including 12 newly sampled
markers, namely: cytochrome P450 reductase (cpr1), ATPdependent DNA helicase II (ku70), sphinganine palmitoyl
transferase subunit 2 (lcb2), DNA replication licensing factor
(mcm7), phosphoglycerate kinase (pgk1), topoisomerase (top1),
two subunits each of the DNA polymerase (dpa1 and dpe1), the
fatty acid synthase (fas1, fas2), alpha-tubulin (tub1), and tub2.
The previously employed marker his3 was not included, nor were
nrDNA markers. The results are in essence the same as those of
the previously published phylogenies, but with stronger support
for the backbone in the combined analyses (see Cummings &
Meyer 2005). Geiser et al. (2021) claimed that the F1 node
was supported by 12, and the F2 node by 14 of the individual
genes, but did not mention that all 19 genes supported the F3
node (Fusarium s. str. = the Gibberella clade).
In this study we re-investigated the Geiser et al. (2021)
dataset using several different high-resolution phylogenetic approaches, and we found that their evaluations of concordance
were based on an inadequate interpretation of Ultra-Fast bootstrap results (only values 95 % are to be deemed significant,
see Minh et al. 2013, Hoang et al. 2018). In addition to the topological incongruences among six genes (act1, CaM, DNA
polymerase epsilon subunit dpe1, ku70, pgk1, tef1, and tub2),
only six and 11 genes actually support the F1 and F2 nodes,
respectively, while all 19 genes support the F3 node. The low
internode certainty (IC) and IC All (ICA) values obtained for F1
(0.19 and 0.33, respectively) were misinterpreted by Geiser et al.
(2021) as IC values close to 0 indicate conflict between the
partitions (Salichos et al. 2014). The F3 node was well supported
with IC and ICA values at 1 (Geiser et al. 2021, Supplementary
Table. S1), which indicates the absence of conflict.
While the effort by O'Donnell et al. (2020) and Geiser et al.
(2021) to include a high diversity of DNA markers is
commendable, it is undermined by an imbalanced selection of
taxa for their analyses. Specifically, there is a marked overrepresentation of node F1 species, while sampling and taxon
selection across the Nectriaceae is almost absent. Excluding any
of the major genus-level clades, especially those relevant to the
recognition of Bisifusarium, Neocosmospora and Rectifusarium,
introduces taxon sampling biases in a way that reduce the reliability of phylogenetic inferences and support values with respect
to the backbone of the Nectriaceae. Furthermore, neither
O'Donnell et al. (2020) nor Geiser et al. (2021) give full
consideration to morphological and ecological evidence. In
principle, a genus should always be delimited as monophyletic,
supported by derived traits. In addition, its circumscription should
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depend on the systematic (phylogenetic and biological) structure
of the family it belongs to, in this case, the Nectriaceae.
Phylogenetics has rapidly advanced from a powerful adjunct
tool for understanding evolutionary relationships to the dominant
principle for classification, especially for delimitation of taxa at all
ranks. However, the resulting analyses and phylogenies are
compromised if they are not reconciled with other biological data.
The call for additional genomic data in the Fusarium clade (Geiser
et al. 2013, Aoki et al. 2019) may improve backbone node support
values, but the phylogenetic structure is unlikely to change; it is the
translation of that data into practicable taxonomy. The broad
Fusarium concept of Aoki et al. (2019), O'Donnell et al. (2020) and
Geiser et al. (2021) is phylogenetically possible, but it does not
offer a generic definition based on a combination of available
genetic, morphological, biochemical and ecological data. It is,
thus, impractical in that it is so broad that the genus would not have
any synapomorphies when compared to other genera of the
Nectriaceae outside their broad circumscription of Fusarium.
The arguments presented by Aoki et al. (2019), O'Donnell
et al. (2020) and Geiser et al. (2021) are centred around the
phylogenetic support of some nodes, which have never been a
key subject of the discussion, as the made observations
generally match the interpretations made by many authors. While
the very broad circumscription of Fusarium reflects as a monophyletic group in DNA phylogenetic analyses, the TFC is a
conglomerate of several monophyletic genera that has a common ancestor (node F1 in Geiser et al. 2013). Each of these
genera has a distinctive combination of morphological features.
An analogous situation was observed in the monophyletic sister
clade that was originally classified as Cylindrocarpon s. lat., but
that is currently viewed as composed of several monophyletic
genera i.e., Cinnamomeonectria, Corinectria, Cylindrodendrum,
Dactylonectria, Ilyonectria, Macronectria, Neonectria, Pleiocarpon, Rugonectria, Thelonectria and Tumenectria (Chaverri
et al. 2011, Gr€afenhan et al. 2011, Lombard et al. 2014,
Salgado-Salazar et al. 2016, Gonzalez & Chaverri 2017).
What is a genus?
Taxonomically, a genus is a group that is defined by a type
species, and that often includes additional species considered to
belong to the same group (Vellinga et al. 2015). The observations or category of data involved in delineating genera have
varied over time, and in many cases, the characters used to
delimit well accepted genera have proven to be homoplasious
and the genera polyphyletic (Crous et al. 2009). However, it is a
fundamental principle that taxonomic entities should reflect
evolutionary relationships.
This has led to inevitable splitting of well-known fungal taxa,
both genera and species, into smaller groups, but sometimes
also genera were merged with others based on the reappraisal or
discovery of derived characters (e.g., Voglmayr & Thines 2007).
This proceeds with each technological revolution providing ever
deeper insight into the biological/evolutionary relationships of
organisms, and has accelerated again since molecular phylogenetics came into widespread use. There is a prevailing notion
that nature made species, but that humans made all other
taxonomic ranks for their own convenience. However, it is
increasingly recognised that all taxonomic ranks, including the
species level, do not have solid boundaries but are more like a
steam cloud with fuzzy margins. At the genus level, these
6
boundaries are often even more obscure, but is a genus just an
arbitrary (but statistically well-supported) monophyletic convenience, a consensus accepted by a self-appointed committee?
Or is a genus a meaningful, definable unit resulting from
evolutionary processes, which can be recognised by patterns of
biological structure, biochemistry, behaviour, and adaptation to
specific niches? We believe that the latter should be the case.
While we recognise that generic delimitations will always depend
on a subjective choice, we believe that generic concepts should
always be guided in a phylogenetic context by morphological,
biochemical, or ecological characters that can both be used for
practical recognition and convey evolutionary information.
The generic concept for Fusarium proposed by Geiser et al.
(2013, 2021) is a rejection of this concept, as it merges lineages
with divergent characters that were accepted and applied not
only throughout the family Nectriaceae for the delimitation of
genera but also in other fungal families and orders. The very
broad genus Fusarium that it gives rise to does not have clearcut features, as the diversity of characters shared with the rest of
the Nectriaceae is so high that it could be extended almost
arbitrarily to the entire family. It would, in fact be as if the concept
of cryptic species was expanded to genera, that is, genera that
can only be recognised as a well-supported node on a phylogram, which is, in our view, in disagreement with fundamental
principles of practical classification. The node F1 selected by
Geiser et al. (2013, 2021) for defining Fusarium is devoid of
phenotypic support and includes several genera with distinct
evolutionary traits. Indeed, the Geiser et al. (2013, 2021) concept
of Fusarium is strictly phylogenetically defined and essentially
amounts to a list of the species bound within a selected clade.
Their morphological circumscription does not admit the existence
of synapomorphies (i.e., unique diagnostic characters possessed
by all included species), and it extends beyond their chosen node
to other groups in Nectriaceae. In this very wide definition of
Fusarium, phenotypic characters and ecological patterns that
correlate with well-supported monophyletic groups within the
larger, poorly supported TFC are disregarded as basis for
generic delineation.
Admittedly, phenotypic characters in the TFC are tricky to
interpret. The fusarioid macroconidium with or without a welldeveloped foot-shaped basal cell (i.e., basal conidial cell
showing an asymmetrical papillum, delimited from the rest of the
cell and forming a distinct notch) occurs in the majority but not all
of the species in the traditional generic concept, but is also a
feature present in a significant proportion of other members of
the Nectriaceae, or even of the unrelated genus Microdochium
(Amphisphaeriaceae). It is, therefore, not a unique feature for
generic delineation (Gr€afenhan et al. 2011).
Perithecial pigmentation has been used to delimit genera in
Nectriaceae. The orange/red perithecium is an ancestral character in the family and common also to members of the BFC and
early diverging lineages of the TFC, including all Neocosmospora species known to reproduce sexually, Setofusarium,
and some species of Cyanonectria and Geejayessia. These
structures are easily distinguished from the homogeneously
bluish/black perithecia of true Fusarium s. str. species in the
Gibberella clade sensu O'Donnell et al. (2013). Contrary to what
was suggested by Geiser et al. (2021), it is not Neocosmospora
which represents an interesting but morphologically aberrant
lineage, since neither its type nor the members of its modern
morphological circumscription (Nalim et al. 2011) exhibit aberrant
characteristics. It is the dark-coloured perithecia typical of
FUSARIUM
Fusarium s. str. (= Gibberella clade) that are aberrant and unusual within Nectriaceae.
The dark purple to black perithecium formerly used to characterise Fusarium s. str. (= Gibberella), represents a synapomorphic state. Ascomata with similar colours have evolved
independently in some, but not all, species of Geejayessia, while
heterogeneously coloured bluish black or bicoloured perithecia
can be observed in several species of Cyanonectria, which often
appears as a sister genus to Fusarium. However, Cyanonectria
and Geejayessia differ from Fusarium and Neocosmospora by
their typically well-developed stromata as well as their thinner
and smooth perithecial walls. Notably, pale yellowish perithecia
occur in several clades and are a derived character as well, and
one genus that we accept, Albonectria, was initially defined by
white perithecia (Rossman et al. 1999). Also, in terms of its
ascospores, Fusarium shows a derived state. With the exception
of Albonectria, which includes species with hyaline, ellipsoidal to
fusoid, 3-septate, smooth to finely striated ascospores, the
genera mentioned above present mostly pale yellow-brown ascospores. Ascospores of Fusarium s. str. are more often subhyaline, ellipsoidal to fusoid, 1–3-septate, and smooth-walled
when viewed with light microscopy. Ascospores of Neocosmospora are easily distinguished from those of Fusarium by
being ovoid to ellipsoidal, (0–)1-septate, pigmented, conspicuously striate or more rarely cerebriform or spinulose. It is worth
noting that most of the above-mentioned characters and differences are the same applied to define genera across the whole
Nectriaceae (Rossman et al. 1999, Lombard et al. 2015), where
they correlate well with phylogenetic inferences. Ascospores
showing similarly many septa as in Fusarium s. str. have independently evolved in Nectria diploa (now Microcera), as well as
in N. glabra, and N. decora (now Flammocladiella). The fact that
none of these species is a member of the TFC supports the
interpretation that multiseptate ascospores might be apomorphic
for Fusarium s. str., separating it clearly from other phylogenetically related genera.
Behaviour and other adaptations, determine how an organism
operates and survives in nature and are the ultimate determinants and products of natural selection. They may be
difficult to translate into nodes and other results of phylogenetic
analyses such as phylogenetic distance. Despite this, similarities
in adaptive traits are frequently used to calibrate phylogenetic
delimitations of genera. For example, all known species of
Microcera are pathogens of scale insects. It is easy to understand the hypothesis that the ancestor of this clade jumped to
these hosts, followed by subsequent radiation and speciation
(Thines 2019). This resulted in considerable micromorphological
diversity, while a core of adaptation resulting from the parasitic
life style remained conserved. Similarly, several of the genuslevel clades include mostly mycoparasitic species or pathogens
of plants. If we apply this kind of thinking to the well-supported
clades of the TFC, as noted by Schroers et al. (2011), species of
Cyanonectria and Geejayessia occur only on woody hosts
(mostly species of Buxus, Celtis and Staphylea) and would
typically not occur as soil-borne plant pathogens or pathogens of
grasses. They are also not known to produce trichothecene
mycotoxins. This is in stark contrast with the prevailing ecological
concept of Fusarium s. str. as a genus of primarily soil-borne
fungi, of which many are in a firm biological association with
grasses and herbs. Importantly, the vast majority of Fusarium s.
str. species produce trichothecene mycotoxins as a chemical
synapomorphy. Most of the strongly supported clades within the
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REDELIMITED
TFC can be supported by these kinds of morphological, chemical, and biological traits, allowing the possibility of non-arbitrary
recognition of biologically meaningful genera. One such clade is
Neocosmospora.
Arguments for and the practicality of
recognising Neocosmospora (the F. solani
species complex) as a genus
In the days of dual nomenclature, the distinction between the red
perithecia of Neocosmospora, as amended by Nalim et al.
(2011), and the typically purple or blackish perithecia of the
trichothecene-producing Gibberella species was generally
accepted by Fusarium taxonomists. The ecological distinctiveness of Neocosmospora as a group of soil fungi, often associated with roots and causing root rot and vascular wilt diseases,
was also generally acknowledged. In addition to the dissimilar
sexual characters mentioned above, the asexual morphs of this
group are also distinctive. The macroconidia are usually thickwalled, with blunt, rounded apical cells, and they usually have
inconspicuous foot-shaped basal cells. Microconidia are produced on very long, narrow phialides. Cultures of a vast majority
of species of this group can easily be recognised morphologically, even with a dissecting microscope.
The ecological similarities of the members of Neocosmospora
with F. oxysporum have to be acknowledged, as noted by Geiser
et al. (2013, 2021). However, these two groups of species are
morphologically distinct, even as asexual morphs. Fusarium
oxysporum produces macroconidia with acutely pointed apical
cells, and microconidia from phialides that are usually 5–10
times shorter than those of Neocosmospora species.
Geiser et al. (2013, 2021) have pointed out that microchromosomes or conditionally dispensable chromosomes occur
in Neocosmospora and members of their F3 clade, namely
F. oxysporum. Microchromosomes have been observed, however, also in phylogenetically distinct taxa such as Magnaporthe
oryzae (Yoshida et al. 2009, now Pyricularia oryzae), Mycosphaerella graminicola (Stukenbrock et al. 2010, now Zymoseptoria tritici), and Alternaria arborescens (Hu et al. 2012) and
might occur sporadically as a result of horizontal gene transfer.
They are thought to increase the ability of a pathogen to adapt to
the host's defence mechanisms. The ability to acquire conditionally dispensable chromosomes might thus be seen as a
general genetic tool allowing organisms to gain ecologically
advantageous genes. Similarly, they could present a general
driving force in co-evolutionary processes, but the per se
occurrence of conditionally dispensable chromosomes in two
taxa can hardly be used as a criterion for drawing conclusions on
or imply generic relatedness.
In the Nelson et al. (1983) manual and in one of the last
vestiges of the ultra-reductionist Snyder & Hansen (1941) system, F. solani was recognised as the only species of section
Martiella, even though the existence of several distinct mating
populations was known. The European system (exemplified by
Gerlach & Nirenberg 1982) accepted several more species,
derived from the classic Wollenweber & Reinking (1935) treatment. When molecular phylogenetic studies of this group began
in earnest, Neocosmospora included three major clades and
many species (O'Donnell 1993, 2000, O'Donnell et al. 2008a). To
date, 86 species are formally described in this group (Aoki et al.
2019, Sandoval-Denis et al. 2019, Guarnaccia et al. 2021), but
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ET AL.
additional novel phylogenetic lineages are recognised and await
formal description.
Thus, in Neocosmospora we have a group of species that can
easily be recognised morphologically by both sexual and asexual
morphs, exhibit generally consistent ecological behaviour, lack
trichothecene mycotoxins, and form a strongly supported
monophyletic group. This sounds like a biologically meaningful
calibration of a genus, but what about the practicality of doing
this? Presently, the data supporting the recognition of Neocosmospora (and equally, also Fusarium s. str., the F3 clade) is
stronger than the data supporting either of the nodes favoured for
designating a broader concept of Fusarium. If there are 100 plus
species in Neocosmospora, and hundreds of species in the
trichothecene-producing, Poaceae-loving Fusarium s. str. clade,
it will be useful for students, plant pathologists, clinical microbiologists, and other scientists to have different generic names for
each group. Those names will convey information about relationships and behaviour that are lost in a broader definition of
Fusarium with much greater diversity of ecological and
biochemical behaviours. Geiser et al. (2013) raised concerns
that grant evaluators, government regulators and medical practitioners who now believe they know what Fusarium means will
be confused by the segregation of these fusarioid fungi into
different genera, and that confusion could lead to unpredictable
consequences. However, in our experience these end users
continuously familiarise themselves with up-to-date, informative
taxonomic and nomenclatural concepts for socio-economically
important fungal groups, thus allowing them to predict the
possible real-world effects of reliably identified fungi with
increased precision. To them, the segregation of a heterogeneous concept of Fusarium into biologically and biochemically
predictive genera will be helpful.
With Neocosmospora accepted as a different genus, Albonectria, Cyanonectria, and Geejayessia, as defined by Schroers
et al. (2011), as well as Bisifusarium and Rectifusarium as
defined in Lombard et al. (2015) must also be accepted as
separate genera. As previously said, these are all monophyletic
groups, also characterised by distinctive ecological and
morphological traits.
The end consequence of our strategy is a series of phylogenetically well-supported genera, each with a recognisable
suite of morphological characters, and ecological, pathological,
and biochemical behaviour. Indeed, the results of such splitting
activities applied to what we called the Wollenweber concept of
Fusarium s. lat. accounts for 20 segregate genera. Most
importantly, both Fusarium and Neocosmospora will have
generic names to indicate their important but distinct significance. The extraneous species, with different ecology and
generally much lower economic or agricultural significance can
now justifiably be classified elsewhere, where they can be
appreciated for their own features without the need for the uncertainty inherent in a broad concept of the generic name
Fusarium.
The generic concept of Fusarium proposed by Geiser et al.
(2013, 2021) functions well as a phylogenetic concept only if
taxonomists turn their eyes away from all other kinds of data and
observations applied to the family Nectriaceae. It is a political
generic concept, meant to assuage the concerns of plant pathologists and other applied scientists, many of whom are
already upset by the proliferation of cryptic phylogenetic species.
Ironically, this late-blooming alleged pragmatism seems to betray
8
the cladistic ideals that many of its authors profess to adhere to
(Taylor 2014).
All authors agree on the use of the single name Fusarium,
have a common understanding of a phylogenetic structure of the
family Nectriaceae, and agree that removing Neocosmospora
from the main Fusarium core is the critical point of discussion.
Sequencing additional markers may lead to increased phylogenetic support, but it is a false comfort if the taxon sampling does
not include as many genera of Nectriaceae as possible.
Expanded representation of the TFC in the dataset will not solve
the controversy, and the resulting phylogenies will remain unbalanced. The segregation of Neocosmospora from Fusarium
certainly needs to be done efficiently by those who have the most
comprehensive expertise on the relevant species, which include
several of the co-authors of the Geiser et al. (2013, 2021) and
O'Donnell et al. (2020) papers as well as the present one.
Fusarium taxonomy has long been confused because of
the nine-species system of Snyder & Hansen (1940, 1941),
the misleading overlaps caused by convergent evolution and
character loss, the difficulty in characterising perithecia, the
phenomenon of cultural degeneration, and rigid opinions of
the taxonomists and plant pathologists who have worked on
them. To arrive at a stable taxonomy for Fusarium, the generic
concept needs to be fixed in a practical and evolutionary
reasonable manner so that future technologies and applications
will not disrupt it.
SECONDARY METABOLITES OF FUSARIOID
GENERA
The phylogenetic distribution of the fusarioid genera presented
here is further corroborated by their ability to produce genusspecific secondary metabolites. The commercial database Dictionary of Natural Products (DNP; http://dnp.chemnetbase.com),
was used to search for secondary metabolites produced by the
genera and species treated here. The database contained (as of
March 6, 2021) over 720 entries on metabolites from Fusarium s.
lat., even though some plant metabolites, discovered during
studies on the elicitation of phytoalexins by challenging plant
cells with a Fusarium strain, are included. The number of metabolites from Fusarium s. lat. is therefore estimated to be around
680, which is still behind Aspergillus s. lat. (over 3 000 entries)
and Penicillium s. lat. (over 2 700 entries). Hits that were
retrieved were confirmed by consulting the original literature. The
reported structures were corroborated, with a selection of these
compounds presented here (Figs 2–4).
It remains uncertain if the reported taxonomy is reliable, since
the producer strains may have been misidentified or determined
using one of many outdated taxonomic concepts. However,
several compound classes have been encountered multiple
times from the same species or species complex, and in some
instances, the strains were identified by experts and/or
sequenced later in phylogenetic studies (O'Donnell et al. 2018).
The situation is further complicated by the fact that certain
secondary metabolites have been given similar names, but
represent different molecules. The name solaniol has been given
to both a trichothecene (Fusarium s. str.) and a naphthoquinone
(Neocosmospora), and the fusariumins represent four different
secondary metabolites.
FUSARIUM
REDELIMITED
Fig. 2. Secondary metabolites from Fusarium spp. / Neocosmospora spp.
Typical metabolites of Fusarium s. str.
Fusarium sambucinum, the type species of the genus, has not
been studied in much detail, but among the 20 metabolites
known from this species, several metabolites are ranked in the
classes trichothecenes and enniatins. The trichothecenes represents a well-known and notoriously dangerous class of mycotoxins belonging to the scirpene terpenoid type. These
www.studiesinmycology.org
compounds are widely distributed within the genus Fusarium s.
str., including familiar plant pathogenic species such as,
F. culmorum, F. graminearum, F. sporotrichioides and
F. tricinctum (Bamburg et al. 1968, Tatsuno et al. 1968,
Yoshizawa & Morooka, 1973, Jimenez et al. 1997). The enniatins, known from 17 Fusarium s. str. species (Munkvold 2017,
O'Donnell et al. 2018), are cyclic depsipeptides that have strong
antibiotic activities (Plattner et al. 1948, German-Fattal 2001,
9
CROUS
ET AL.
Fig. 3. Some of the most important mycotoxins from Fusarium spp.
Bills & Gloer 2017). Similar to trichothecenes, they are only
known from Fusarium s. str. in the current taxonomic concept,
although Trichoderma and Beauveria, which belong to different
families of the Hypocreales, also produce trichothecenes or
enniatin-like beauvericins, respectively. However, trichothecenes
have not been reported from Neocosmospora or “F. solani”
except from two isolates misidentified as “F. solani” (Ueno et al.
1972, Sugimoto et al. 2002) (Supplementary Table S2)
Two other well-known classes of mycotoxins, the fumonisins
(Bezuidenhout et al. 1988) and zearalenone (Urry et al. 1966),
are also found frequently among species of Fusarium s. str.
Similarly, equisetin, also considered a “mycotoxin” and originally
found from a Fusarium sp. strain (NRRL 5537) in the FIESC
(Vesonder et al. 1979, Xia et al. 2019) is actually a strong
antibiotic. A more complex derivative known as fusarisetin A was
reported from an unidentified Fusarium sp. (Jang et al. 2011).
Some rather unique compounds only known from Fusarium s.
str., include wortmannin (Abbas & Mirocha, 1988) and oxysporizoline (Nenkep et al. 2016), which have interesting biological activities and may be species or even strain-specific.
Among the compounds that are not regarded as mycotoxins,
the antimicrobial sesquiterpenes of the fusarielin type (Sørensen
et al. 2013) and the antiparasitic and cytostatic cyclopeptides of
the apicidin type (Jiang et al. 2002, Von Bargen et al. 2013) have
been respectively isolated from Fusarium s. str. Additionally,
aurofusarin (Munkvold 2017, O'Donnell et al. 2018), chlamydosporol (Munkvold 2017, O'Donnell et al. 2018), fusapyrone
(Evidente et al. 1994), fusaric acid (Munkvold 2017, O'Donnell
et al. 2018), fusoxysporone (Abraham & Hannsen 1992),
fusaproliferin, moniliformin (Munkvold 2017, O'Donnell et al.
2018) and the terpestacins (Liu et al. 2013) are other examples of secondary metabolites found only in Fusarium s. str. Thus
far, only one report has indicated that a Neocosmospora species
can produce fusaric acid (Zhou et al. 2019). Both aurofusarin and
bikaverin produced by Fusarium s. str. and other bis-naphthoquinone and bis-naphthopyrone pigments protect fungi from
predation (Xu et al., 2019), while Neocosmospora species produce other naphthoquinones such as javanicin (Arnstein & Cook
1947, Kimura et al. 1981) as potential predator protectors. Some
unique compounds have been reported from marine strains of
10
certain Fusarium species, which include the mangicols, rare
sesterterpenes produced by a strain tentatively classified as
F. heterosporum (Renner et al. 2000).
Typical metabolites of Neocosmospora and
other fusarioid genera
Neocosmospora species and other fusarioid genera apparently
have a different secondary metabolism, or have not been
intensively studied in the past. A striking example are the
cyclosporins, which are immunosuppressive peptides. Originally,
these were obtained from Tolypocladium inflatum, but later also
found to be produced by species of Neocosmospora (Sawai et al.
1981, Nakajima et al. 1989). However, they have not been reported from Fusarium s. str. Other unique compounds only
known from Neocosmospora species, include dihydrofusarin
(Kurobane et al. 1980, Kyekyeku et al. 2017), the polyketides
neovasipyrones (Furumoto et al. 1995, Nakajima et al. 1995) and
vasinfectin A (Furumoto et al. 1997). The rare cyclopeptides of
the neosansalvamide type (Lee & Lee 2012) and the resorcylic
acid lactones of the monorden/monocillin type (Cutler et al. 1987,
Gao et al. 2013) are also known from Neocosmospora and other
fungi, but not from Fusarium s. str., even though the latter
compounds bear a high structural resemblance to zearalenone.
Several Neoscomospora species produce a range of naphthoquinones that are members of a widespread class of polyketides (Roos 1977).
The fusarioid genus Bisifusarium is known to produce the
PKS/NRPS hybrid siderophore, dimerumic acid (= dimerum acid)
(Diekmann 1970), and indole acetic acid (Reddy & Reddy 1992,
Kulkarni et al. 2011, 2013). The parnafungins, which are under
development as antimycotics, are only known from Microcera
larvarum (Parish et al. 2008). Additionally, Microcera larvarum is
also known to produce monocerin and fusarentins, which are not
known from any other fungi (Grove & Pople 1979), except a
Colletotrichum species (Tianpanich et al. 2011). The anticancer
agent balanol (azepinostatin) (Ohshima et al. 1994) is known to
be produced by two Fusicolla species, which might be applied as
a taxonomic marker for this genus, although it has also been
FUSARIUM
REDELIMITED
Fig. 4. Secondary metabolites from fusarioid Hypocreales.
found in species of the Ophiocordycipitaceae. Unfortunately,
there is no available information on secondary metabolites for the
other fusarioid genera treated here. However, secondary
www.studiesinmycology.org
metabolite studies of these missing genera will facilitate for the
discovery of novel molecules and help to elucidate the functional
biodiversity of these fungi.
11
CROUS
ET AL.
RECOMMENDED METHODS FOR THE
IDENTIFICATION AND CHARACTERISATION OF
FUSARIUM AND ALLIED GENERA
The following part of this study presents an overview of the
morphological and phylogenetic characters of Fusarium and
related genera as well as an account of recommended methods
for the identification and characterisation of these taxa. In
addition, novel genera and species are described and, in view of
the recent taxonomic data, a list of names that are applied to the
genus Fusarium s. lat. with their current scientific names is
presented.
Morphology
Current Fusarium taxonomy is dominated by molecular phylogenetic studies. Nonetheless, morphology is a fundamental
component of the generic and species concepts of fungi and
must not be overlooked. Key morphological features for generic
circumscription include characteristics of sexual morphs such as
perithecial colour, wall thickness and anatomy, surface structures
and the presence and nature of a basal stroma, ascospore
shape, septation, colour and surface ornamentation (Rossman
et al. 1999). Classification of taxa solely based on their
asexual morphs can be trickier than integrated systems using
sexual and asexual characters. However, the general shapes,
different types and combinations of conidiogenous structures and
conidia present in culture can be sufficient to allow a preliminary
identification (Fig. 5), especially if host data are also available
(Leslie & Summerell 2006). For species-level characterisation, a
number of morphological traits must be carefully studied,
particularly those of the asexual morph, while sexual morphs are
generally less suitable, especially as they are typically not produced in culture. Diagnostic characters for species identification
include colony characters such as colony morphology, pigmentation, and type of aerial mycelium. Also included are the dimensions and characteristics of aerial conidiophores and
conidiogenous cells (mono- vs polyphialides), presence/absence
and characteristics of sporodochia, the types of conidia produced, e.g., aerial microconidia, mesoconidia, and aerial and
sporodochial macroconidia. In examining conidia themselves,
consideration is given to the overall shape, septation and curvature of the macroconidia, as well as characteristics of their
apical and basal cells; with aerial microconidia, their dimensions,
shape, septation and spatial organisation (forming slimy heads,
chains or a combination of both) are noted. Finally, the presence
or absence of chlamydospores may be important.
Culture media and incubation
Vigorous growth, sporulation, and pigment production of fusarioid
fungi can be achieved on numerous agar formulations. The
morphology of fungal structures will vary dramatically depending
on the selection of media and growth conditions which may
compromise the identification process. In addition, it is also
common for fusaria to degenerate and lose viability in culture,
particularly when they are grown on nutrient-rich media (Nelson
et al. 1983, Nirenberg 1990, Summerell et al. 2003, Leslie &
Summerell 2006). Culture conditions and media have been
extensively summarised in the literature (Booth 1971, Nirenberg
12
1990, Nelson et al. 1994, Summerell et al. 2003, Leslie &
Summerell 2006). Consequently, we recommend the agar formulations listed in Table 1 to be employed for the isolation and
description of fusaria. A summary of the procedures and conditions suitable for work with fusarioid fungi is shown in Fig. 6.
An important condition that must be stressed is that the
identification must always be made on the basis of a monosporic
culture (a culture produced from a single sporulating conidium,
ascospore, or hyphal tip), as multiple species are commonly
found to co-occur in the same substrate tissue. A freshly isolated
fusarioid strain should be sub-cultured onto at least two different
culture media, a relatively rich one suitable for examination of
gross morphology, and a nutrient-poor one for micromorphological examination and for further culture propagation. The
standard culture setup for initial assessment of growth rates and
colony characters i.e., colony pigmentation, diffusible pigments,
and colour of sporodochia, is to use potato dextrose agar (PDA)
incubated for 1–2 wk. Fusarium and related genera will also
grow and sporulate well on malt extract agar (MEA, recipe in
Crous et al. 2019a), which can be a suitable alternative for initial
isolation and monosporic cultivation. However, MEA should not
be used to assess colony or morphological characters. Standard
incubation is commonly made in total darkness; however,
exposure to light will normally result in a faster and more intense
pigmentation. We have observed better colour formation using
in-house prepared media rather than commercial formulae.
While colony colour cannot be employed as a primary criterion
for species identification, it can provide useful means to grossly
distinguish related groups and to direct the identification process
towards determining genera or species complexes. The high
nutrient content of these agar media strongly affects sporulation,
commonly resulting in the development of atypical structures.
Therefore, we strongly discourage the use of PDA for micromorphological assessment or culture propagation of Fusarium
spp. (Nelson et al. 1994, Summerell et al. 2003). Oatmeal agar
(OA) is a suitable alternative for strain sub-culturing, allowing for
good sporulation with reduced strain degeneration; however, it is
not recommended for micromorphological studies.
Carnation leaf agar (CLA), synthetic nutrient-poor agar (SNA),
and water agar (WA) are the standard culture media for micromorphological analyses. Also, by reducing culture degeneration,
they allow for prolonged storage of actively growing cultures
(Nirenberg 1976, Nelson et al. 1983, Leslie & Summerell 2006).
Subcultures on CLA will normally produce abundant sporodochia
and macroconidia on the surface or around the carnation leaf
pieces with consistent morphological features. Incubation at
room temperature (20–25 °C) for 1–2 wk under a 12/12 h nearUV light (wavelength 320–400 nm)/dark or near-UV light/cool
fluorescent light cycles results in stronger sporulation and good
development of sporodochial pigmentation (Nirenberg 1990,
Seifert 1996, Summerell et al. 2003, Leslie & Summerell 2006).
The use of continuous near-UV light (also commonly termed
"blacklight" or UV-A light) is also suitable although it often results
in the formation of unusually long macroconidia (Nirenberg
1990), and it can suppress the development of useful morphological characters such as the globose microconidia of Fusarium
globosum. Nevertheless, incubation under near-UV light is
fundamental since isolates of some species such as Fusarium
poae and F. sacchari are known to lack macroconidia or to
produce them in only small quantities unless they are stimulated
by incubation under a near-UV light source (Leslie et al. 2005,
Leslie & Summerell 2006). Fusarium cultures also need
FUSARIUM
REDELIMITED
Fig. 5. Basic morphological features of fusarioid fungi. A. Macroconidial shapes. A1. Slender with no significant curvature. A2. Curved with parallel walls. A3. Unequally curved. A4.
Widest at the middle portion. A5. Widest at the apical third, wedge-shaped. A6. Widest at the basal portion. A7. Irregularly clavate and swollen. A8. Elongate, whip-like. A9. Distinctly
curved. B. Macroconidial apex. B1. Curved. B2. Long and tapered. B3. Pointed. B4. Blunt. B5. Hooked. B6. Elongated. C. Macroconidial base. C1. Obtuse, non foot-shaped. C2. Papillate,
non foot-shaped. C3. Poorly developed, foot-shaped. C4. Well-developed, foot-shaped. C5. Elongate, foot-shaped. D. Aerial phialides and microconidial organization. D1. Monophialide.
D2–D5. Polyphialides. D2. Simple polyphialide. D3–D4. Polyphialides with multiple conidiogenous loci. D5. Sympodially proliferating polyphialides. D6, D7. Microconidia forming false
heads. D8, D9. Microconidia in chains (D8. Dry chain. D9. Palisade). E. Sporodochial conidiophore and conidiogenous cells. F. Aerial conidiophore bearing mesoconidia. G. Mesoconidia.
H. Microconidial shapes. H1. Fusiform. H2. Oval. H3. Obovoid. H4. Reniform. H5. Allantoid. H6. Clavate. H7. Napiform. H8. Pyriform. H9. Limoniform.
www.studiesinmycology.org
13
CROUS
14
Table 1. Recommended agar media formulations for the isolation and cultivation of fusaria.
Components1
Preparation2
Incubation3
Application
Reference
Carnation leaf agar (CLA)
Sterilised carnation leaves
WA
Carnation leaves are cut into
approximately 5 × 5 mm pieces and
dried at 60 °C for 24 h; sterilise by
gamma radiation or autoclave; place
3–5 pieces on nearly solid 2 % WA
surface.
25 °C;
7–14 d under 12 h
near-UV-light/dark
cycle; 7–14 d under
24 h near-UV-light
Micromorphological
characterisation:
formation of
sporodochia;
sporodochial
macroconidia
Fisher et al. (1982),
Crous et al. (2019a)
Selective Fusarium Agar (SFA)
Glucose (Dextrose)
KH2PO4
NaNO3
MgSO4$7H2O
Yeast Extract
1 % FeSO4$7H2O (aquous)
Streptomycin
Neomycin
Dichloran
Agar
Water
20 g
1g
2g
0.5 g
1g
1 ml
(5 % w/v) 20 mL
(1 % w/v) 12 mL
(50 % w/v in ethanol) 13 mL
20 g
1 000 mL
Add all components, except antibiotics,
to water and autoclave; cool to
45–50 °C and add antibiotics. Dichloran
can be replaced by PCNB (0.75 g).
25 °C; 7–14 d in
dark
Selective isolation
of fusaria from soil
Tio et al. (1977),
Leslie & Summerell (2006)
Komada's Medium
D-Galactose
L-Asparagine
KH2PO4
KCl
MgSO4$7H2O
PCNB
Fe3Na EDTA
Streptomycin
Oxgall stock solution
Na2B4O7$10H2O (borax)
Agar
Water
20 g
2g
1g
0.5 g
0.5 g
0.75 g
0.01 g
(5 % w/v) 6 mL
0.5 g
0.5 g
15–20 g
1 000 mL
Add all components, except antibiotics,
oxgall and borax; to water and
autoclave; cool to 45–50 °C and add the
reamining components. Adjust pH to 3.8
± 0.2 prior to autoclaving.
25 °C; 7–14 d in
dark
Selective isolation
of fusaria from soil,
particularly those
belonging to the
Fusarium
oxysporum species
complex. Other
fusaria can be
inhibited by this
medium
Komada (1975),
Leslie & Summerell (2006)
Malachite Green Agar (MGA)
Peptone
KH2PO4
MgSO4$7H2O
Malachite green oxalate
Streptomycin
Penicillin
Agar
Water
15 g
1g
0.5 g
2.5 mg
(5 % w/v) 20 mL
(5 % w/v) 20 mL
20 g
1 000 mL
Add all components, except antibiotics,
to water and autoclave; cool to
45–50 °C and add antibiotics. Penicillin
can be also replaced by
chloramphenicol (5 % w/v) or neomycin
(1 % w/v).
25 °C; 7–14 d in
dark
Selective isolation
of fusaria from soil
and plant material,
with improved
inhibition of nonfusarioid
contaminants
Castella et al. (1997), Leslie &
Summerell (2006)
Oatmeal agar (OA)
Oatmeal extract
Agar
1 000 mL
15–20 g
Oatmeal flakes (30 g/L) are wrapped in
cloth and simmered in water for 2 h;
liquid is squeezed and filtered through
cloth.
25 °C; 7–14 d in
dark
Macromorphological
characterisation,
colony
characteristics
Crous et al. (2019a)
ET AL.
Agar media
Table 1. (Continued).
www.studiesinmycology.org
Agar media
Components1
Preparation2
Incubation3
Application
Reference
Potato dextrose agar (PDA)
Potato extract
Agar
Water
230 mL
15–20 g
770 mL
Potatoes (5 kg; peeled and sliced) are
minced; soak in water (300 mL/100 g
potato) overnight at 4 °C; filter through
cloth; adjust pH to 6.6.
25 °C; 7–14 d in
dark; 5–40 °C
(5 °C increments for
growth curves)
Inoculum
preparation, macromorphological
characterisation:
colony
characteristics;
growth curve
Crous et al. (2019a)
Peptone Pentachloronitrobenzene
(PCNB) agar (PPA)
Peptone
KH2PO4
MgSO4$7H2O
PCNB
Streptomycin
Penicillin
Agar
Water
15 g
1g
0.5 g
0.75 g
(5 % w/v) 20 mL
(5 % w/v) 20 mL
20 g
1 000 mL
Add all components, except antibiotics,
to water and autoclave; cool to
45–50 °C and add antibiotics. Penicillin
can be also replaced by
chloramphenicol (5 % w/v) or neomycin
(1 % w/v).
25 °C; 7–14 d in
dark
Selective isolation
of fusaria from soil
and plant material
Nash & Snyder (1962), Booth (1971),
Leslie & Summerell (2006)
Rose Bengal-Glycerine-Urea Medium
(RbGU)
Glycerol
Urea
L-Alaninw
PCNB
Rose Bengal
Streptomycin
Agar
Water
10 g
1g
0.5 g
1g
0.5 g
(5 % w/v) 20 mL
15 g
1 000 mL
Add all components, except antibiotics,
to water and autoclave; cool to
45–50 °C and add antibiotics.
25 °C; 7–14 d in
dark
Isolation of fusaria
from soil and plant
material
van Wyk et al. (1986), Leslie &
Summerell (2006)
Synthetic nutrient-poor agar (SNA)
KH2PO4
KNO3
MgSO4$7H2O
KCl
Glucose
Saccharose
Water
1g
1g
0.5 g
0.5 g
0.2 g
0.2 g
1 000 mL
Add all components to water and
autoclave.
25 °C; 7–14 d
under 12 h nearUV-light/dark cycle
Inoculum
preparation, micromorphological
characterisation:
aerial conidiophores
and micro- &
macroconidia;
chlamydospore
formation
Nirenberg (1976), Crous et al. (2019a)
Water agar (WA)
Agar
Water
15–20 g
1 000 mL
25 °C; 7–14 d in
dark
Inoculum
preparation, base
agar for CLA
Crous et al. (2019a)
1
FUSARIUM
Unless specified differently, antibiotic stock solutions are prepared in distilled water.
Water refers to distilled water; autoclave = 121 °C for 15 min.
3
Near-UV = near ultraviolet spectrum (wavelength 320–400 nm).
2
REDELIMITED
15
CROUS
ET AL.
Fig. 6. Flow diagram summarising recommended methods for the preservation, identification, and characterisation of fusarioid fungi.
adequate aeration to produce conidia reliably and to attain stable
growth rates, and hence we discourage the incubation of sealed
plates. Carnation leaf agar, SNA, and WA are also suitable for
the observation of conidiophore disposition and microconidial
arrangements such as the formation of false heads, chains or
both. These structures can easily be examined under a dissecting microscope or at low magnification under a compound
light microscope (Leslie & Summerell 2006). Examination of
micromorphological characters must be carried out using slide
preparations mounted in water. Lactic acid, lactophenol and
Shear's mounting media can cause considerable shrinking of the
structures and can alter the appearance of the cell surface;
hence we advise against the use of these mountants for examination of morphological characters in Fusarium and related
genera.
Additional culture media, incubation conditions, and protocols
are available for induction of sexual characters in Fusarium and
related genera (Klittich & Leslie 1988, Leslie & Summerell 2006,
Guo et al. 2018, Kim et al. 2019, Santos et al. 2019). Carrot agar
(CA) and half-strength CA are the most commonly used media.
16
The crossing procedures are often variations from the protocol of
Klittich & Leslie (1988), in which strains of opposite mating types
are paired in all possible combinations as male and female parents, together with crosses made against tester strains from
known mating populations (Leslie & Summerell 2006). The process can be shortened by reducing the number of combinations to
be crossed by first determining the MAT gene alleles carried by
each strain by means of specific mating type idiomorph PCR
primers (Kerenyi et al. 1999, 2004, Steenkamp et al. 2000).
Molecular studies
Several genes, primer combinations and PCR conditions have
been listed in the Fusarium literature (O'Donnell et al. 1998a, b,
2000a, b, 2007, 2010, 2013, Gr€afenhan et al. 2011, Lombard
et al. 2015, 2019a, b), including whole-genome sequencing to
mine for the desired genes (O'Donnell et al. 2020, Geiser et al.
2021). Here we detail those DNA markers that have shown the
best results in routine diagnosis (Table 2, Fig. 6).
FUSARIUM
REDELIMITED
Table 2. Recommended PCR primers for DNA amplification of Fusarium and related genera.
Gene/DNA region
1
2
Primer
Name
Abbreviation Name
Direction Sequence (5'/3')
Reference
28S large subunit of the nrDNA
LSU
LR0R
LR5
NL42
Forward
Reverse
Reverse
ACCCGCTGAACTTAAGC
ATCCTGAGGGAAACTTC
GGTCCGTGTTTCAAGACGG
Vilgalys & Sun (1994)
Vilgalys & Hester (1990)
Kurtzman & Robnett (1997)
ATP citrate lyase
acl1
230up
1220low
Forward
Reverse
AGCCCGATCAGCTCATCAAG
CCTGGCAGCAAGATCVAGGAAGT
Gr€afenhan et al. (2011)
Gr€afenhan et al. (2011)
Beta-tubulin
tub2
T1
TUB-2Fd2
TUB4RD
Forward
Forward
Reverse
AACATGCGTGAGATTGTAAGT
GTBCACCTYCARACCGGYCARTG
CCRGAYTGRCCRAARACRAAGTTGTC
O'Donnell & Cigelnik (1997)
Woudenberg et al. (2009)
Woudenberg et al. (2009)
Calmodulin
CaM
CAL-228f
CAL-CL12
CAL-CL2A2
CAL-2Rd
Forward
Forward
Reverse
Reverse
GAGTTCAAGGAGGCCTTCTCCC
GARTWCAAGGAGGCCTTCTC
TTTTTGCATCATGAGTTGGAC
TGRTCNGCCTCDCGGATCATCTC
Carbone & Kohn (1999)
O'Donnell et al. (2000b)
O'Donnell et al. (2000b)
Quaedvlieg et al. (2011)
Internal transcribed spacer
region of the nrDNA
ITS
ITS5
V9G2
ITS4
Forward
Forward
Reverse
GGAAGTAAAAGTCGTAACAAGG
TTACGTCCCTGCCCTTTGTA
TCCTCCGCTTATTGATATGC
White et al. (1990)
de Hoog & van den Ende (1998)
White et al. (1990)
RNA polymerase largest subunit
rpb1
Fa
F7
F81
F61
R8
R9
G2R1
Forward
Forward
Forward
Forward
Reverse
Reverse
Reverse
CAYAARGARTCYATGATGGGWC
CRACACAGAAGAGTTTGAAGG
TTCTTCCACGCCATGGCTGGTCG
CTGCTGGTGGTATCATTCACG
CAATGAGACCTTCTCGACCAGC
TCARGCCCATGCGAGAGTTGTC
GTCATYTGDGTDGCDGGYTCDCC
O'Donnell
O'Donnell
O'Donnell
O'Donnell
O'Donnell
O'Donnell
O'Donnell
RNA polymerase second
largest subunit
rpb2
RPB2-5f2
fRPB2-7cf
fRPB2-7cr
RPB2-11ar
Forward
Forward
Reverse
Reverse
GGGGWGAYCAGAAGAAGGC
ATGGGYAARCAAGCYATGGG
CCCATRGCTTGYTTRCCCAT
GCRTGGATCTTRTCRTCSACC
Reeb et al. (2004)
Liu et al. (1999)
Liu et al. (1999)
Liu et al. (1999)
Translation elongation factor 1-alpha
tef1
EF-1
EF-2
Forward
Reverse
ATGGGTAAGGARGACAAGAC
GGARGTACCAGTSATCATG
O'Donnell et al. (1998b)
O'Donnell et al. (1998b)
et
et
et
et
et
et
et
al.
al.
al.
al.
al.
al.
al.
(2010)
(2010)
(2010)
(2010)
(2010)
(2010)
(2010)
Used only for sequencing reactions.
Alternative primer, not used in this study.
Nuclear ribosomal DNA (nrDNA), including the internal transcribed spacer region cistron (ITS) and the 28S large subunit
nrDNA (LSU), are nearly useless for species recognition in
Fusarium and related genera. Nevertheless, given the ease
of amplification and the extensive data available for comparison in
public databases (Schoch et al. 2012), these markers are useful in
the discrimination between the multiple species complexes of
Fusarium, and for obtaining a confident genus-level identification
for Fusarium and related genera, allowing further DNA markers to
be incorporated in the analyses. The ITS region can still provide
valuable information at species level for related genera containing
species formerly included in Fusarium (Bisifusarium, Cosmosporella, Fusicolla, Macroconia, Microcera, and Stylonectria).
Many protein-coding genes have been explored for identification and taxonomic purposes in Fusarium and fusarioid fungi.
The two main genes used for identification are tef1 and rpb2.
Both offer high discriminatory power and are well represented in
public databases. Translation elongation factor 1-α is commonly
the first-choice identification marker as it has very good resolution power for most species in all the genera treated here, while
rpb2 allows for enhanced discrimination between closely related
species. For example, some species in the Fusarium fujikuroi
species complex (FFSC) and in Neocosmospora that are not
easily separated by using tef1 alone (O'Donnell 2000, Nalim
et al. 2011, Herron et al. 2015), can be resolved with rpb2. On
the other hand, PCR amplification and sequencing success are
often better for tef1 than for rpb2. When used for phylogenetic
analyses, sequence alignments of rpb2 sequences are much
www.studiesinmycology.org
more robust and less ambiguous than tef1 data, given the former
gene's advantageously low proportion of introns. An analogous
situation has been shown in Aspergillus (Samson et al. 2014)
and Penicillium (Visagie et al. 2014).
Additional genetic markers, often employed in association
with the previously mentioned genes in multigene phylogenetic
analyses include acl1, tub2, CaM, and rpb1. These markers have
variable resolution or applicability depending on the genus or
species complex. For example, use of CaM data may yield
conflicting clade resolutions in the FFSC (O'Donnell 2000, AlHatmi et al. 2019), while paralogous or xenologous gene
copies have been demonstrated for tub2 in the
F. chlamydosporum and F. incarnatum-equiseti species complexes (O'Donnell et al. 2009) as well as in Neocosmospora
(O'Donnell 2000, O'Donnell et al. 2008a).
The most widely used algorithm for fungal identification by
means of DNA markers is the Basic Local Alignment Search Tool
(BLAST), available at the NCBI's GenBank website. This is a
quick and useful method that can convey a great deal of information, but its results must be analysed with care given the
presence of a high proportion of misidentified strains and lowquality sequences that must be filtered out (Vilgalys 2003, Nilsson
et al. 2012). Sequences from type material are present in the
GenBank nucleotide database for most fusarioid species known
from culture, especially for rpb2 and tef1 barcodes, but the ex-type
status of these sequences is not always explicitly mentioned. In
many cases the names listed do not reflect the current
taxonomy, even for sequences derived from ex-type cultures.
17
CROUS
ET AL.
Some sequences used in past phylogenetic analyses
of O'Donnell et al. (2020) and Geiser et al. (2021) appear to
be linked to incorrect Fusarium names, likely due to errors in
the database used. For this reason, we recommend the use of
our curated database: Fusarioid-ID (https://www.fusarium.org). It
can also be used for sequence similarity-based analysis of
routine isolations and for identifications within several related
genera.
MALDI-TOF
A number of studies have thus far demonstrated the utility of
mass spectrometry (MS) for species determination of subgroups
of Fusarium, particularly members of the FFSC (Al-Hatmi et al.
2015, 2016, Wigmann et al. 2019). It is also useful for clinically relevant subgroups within several Fusarium species complexes (Marinach-Patrice et al. 2009, Triest et al. 2015, Sleiman
et al. 2016, Paziani et al. 2020) and clinically relevant Bisifusarium (Triest et al. 2015, Paziani et al. 2020) and Neocosmospora
species (Marinach-Patrice et al. 2009, Triest et al. 2015, Sleiman
et al. 2016, Paziani et al. 2020). These techniques show highly
accurate discriminative power, comparable to what has been
shown with bacteria and yeasts. Only a limited number of taxa
have thus far been evaluated, and a genus-wide evaluation of
applicability of MALDI-TOF to Fusarium and related taxa is
pending. The main limiting factor is, as usual, the current lack of
representation of these taxa in commercial spectrum databases,
a matter that can be resolved by constructing in-house, curated
reference databases of spectra. Online availability and comparison of MS spectra of Fusarium has been proposed by Triest
et al. (2015).
MATERIALS AND METHODS
Isolates and fungarium specimens
Fungal strains were obtained from the Westerdijk Fungal
Biodiversity Institute (WI) collection (CBS), the Belgian Coordinated Collections of Microorganisms (IHEM), the International
Mycological Institute (IMI), and the personal collection of Pedro
W. Crous (CPC) housed at WI. For the list of names applied to
the genus Fusarium and related fungarium specimens, the
following fungaria were approached for holotype specimens: B,
BM, BO, BP, BPI, BR, BRA, C, CBS, CO, DAOM, E, FH, H, HAL,
IMI, K(M), L, LEP, M, MASS, MPA, NY, PC, PAD, PARMA, PAV,
PH, PRM, ROVP, SIENA, STR, UPS, VPRI, W, and WIR.
DNA amplification and phylogeny
Total genomic DNA was extracted from isolates grown for 7 d on
PDA or MEA (recipes in Crous et al. 2019a; Table 1) incubated at
24 °C under a 12/12 h photoperiod using the Wizard® Genomic
DNA purification Kit (Promega Corporation, Madison, WI, USA),
following the manufacturer's instructions. Partial gene sequences
were determined for eight DNA markers, i.e., acl1, CaM, ITS,
LSU, rpb1, rpb2, tef1, and tub2 using PCR protocols described
elsewhere (O'Donnell et al. 1998b, 2007, 2010, Lombard et al.
2015). Primer pairs used for amplification and sequencing of
18
the respective gene regions are summarised in Table 2.
Consensus sequences for each marker were assembled in
Geneious R11 (Kearse et al. 2012) or SeqMan Pro v. 15.3.0
(DNASTAR, Madison, WI, USA). All sequences generated in this
study were deposited in GenBank (Table 3; also see Diagnostic
DNA Barcodes in list of Fusarium names). The multiple
sequence alignments and phylogenetic trees were deposited in
TreeBASE (study ID 28093).
Sequences of the individual markers, including introns, were
aligned using MAFFT v. 7.110 (Katoh et al. 2019) using default
parameters and manually corrected where necessary. Seven
multimarker datasets (Table 4) were assembled and analysed
using Maximum Likelihood (ML) and Bayesian Inference (BI).
For the ML analyses, concatenated phylogenies, where each
marker was treated as a separate partition, were determined
using IQ-TREE v. 2.1.2 (Nguyen et al. 2015, Minh et al. 2020b)
with ultrafast bootstrapping (UFBoot2; Hoang et al. 2018) for
estimation of branch support. The most suitable evolutionary
model for each partition was estimated using ModelFinder
(Kalyaanamoorthy et al. 2017; Minh et al. 2020b) as implemented in IQ-TREE. To assess whether the individual markers
were compatible, genealogical concordance factors (gCF) were
calculated using IQ-TREE (Minh et al. 2020a, b). Additional ML
analyses were performed using RAxML v. 8.2.12 (randomised
accelerated (sic) maximum likelihood for high performance
computing; Stamatakis 2014) with the system's default modelling
options. The robustness of the analysis was evaluated by
bootstrap support (BS) with the number of bootstrap replicates
automatically determined by the software. The BI analyses were
carried out through the CIPRES website (http://www.phylo.org)
using MrBayes v. 3.2.7a (Ronquist & Huelsenbeck 2003)
incorporating the best evolutionary models for each marker as
determined by MrModeltest v. 2.3 (Nylander 2004). Two parallel
Markov Chain Monte Carlo (MCMC) runs of four incrementally
heated chains (temp parameter = 0.2) were run starting from a
random tree topology. The MCMC analyses lasted for 5M generations, and convergence of the runs was checked by average
standard deviation of split frequencies below 0.01. Trees were
saved every 1 000 generations and the first 25 % of saved trees
were discarded as the “burn-in” phase. Posterior probabilities
(PP) were determined from the remaining trees. Proper mixing of
the MCMC runs was further confirmed by checking that all chains
converged (minimum and average Estimated Sampled Size
[ESS >200], Potential Scale Reduction Factor [PSRF = 1.0]) and
by plotting and analysing trace file results using Tracer v.1.7.1
(Rambaut et al. 2018).
The phylogenetic re-analysis of the dataset presented by
Geiser et al. (2021) was first made according to the original
exons-only alignment file and procedures as indicated in
Geiser et al. (2021) (Supplementary Table S1). Additionally, the
dataset was split into the 19 genes according to the original
partitioning file, and every gene was realigned using the
MAFFT webserver (v. 7, Katoh et al. 2019) applying the G-INSi algorithm. All other parameters were set to default. Six of the
19 genes exhibited a diverging alignment length. No subsequent changes were done to the alignments. The sequences
were merged using BioEdit (v. 7.2.5, Hall 1999), and the
phylogenetic trees were calculated using Minimum evolution
(ME) and ML algorithms, and BI. The ME tree was calculated
using FastTree 2 (Price et al. 2010) using standard settings
and 1 000 bootstraps (Felsenstein 1985). The ML analysis was
done using RAxML (v. 8.2.12, Stamatakis 2014) with the
Table 3. Details of strains included in the phylogenetic analyses.
Species name
Strain1
Substrate
GenBank accession number2
Country
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
www.studiesinmycology.org
Albonectria albosuccinea
NRRL 20459
Unidentified tree
Venezuela
—
—
JAADYS010000048.1*
JAADYS010000048.1*
JX171471
JX171585
JAADYS010002360.1*
—
A. rigidiuscula
CBS 133754
Bauhinia longicupsis
French Guiana
—
—
MW827602
MW827641
MW834177
MW833995
MW834269
—
Atractium crassum
CBS 180.31T = NRRL 20894
Water tap
Germany
—
—
KM231790
MH866623
MW834178
HQ897722
KM231919
—
At. stilbaster
DAOM 215627
Cut stump
Canada
—
—
—
HQ843769
—
HQ897748
—
—
Bisifusarium delphinoides
CBS 110140 = FRC E-0073 = NRRL 36160
Human eye
USA
—
—
MW827603
MW827642
JX171535
HM347219
EU926302
—
B. dimerum
CBS 108944ET = NRRL 36140
Human blood
Netherlands
—
—
JQ434586
JQ434514
—
HM347218
KR673912
—
T
B. nectrioides
CBS 176.31 = NRRL 20689
Humus
Honduras
—
—
EU926245
EU926245
JX171477
JX171591
EU926312
—
B. penzigii
CBS 116508 = ATCC 15621 = NRRL 20711
Human eye
Sri Lanka
—
—
EU926256
EU926256
JX171482
HM347217
EU926323
—
Corinectria fuckeliana
CBS 239.29 = IMI 039700
Picea sitchensis
Scotland
—
—
MW827604
MW827643
MW834179
MW833996
DQ789728
—
Co. tsugae
CBS 788.69T
Tsuga heterophylla
Canada
—
—
KM231763
KM231763
—
KM231763
MW834270
—
Cosmospora butyri
CBS 301.38T = MUCL 9950
Butter
Denmark
—
—
MW827605
MW827644
MW834180
HQ897729
—
—
Cs. coccinea
CBS 341.70
Inonotus nodulosus on Fagus sylvatica
Germany
—
—
MH859703
KM231692
MW834181
HQ897777
KM231947
—
Cs. khandalensis
CBS 356.65IT = ATCC 16091 = IMI 112790 = MUCL
Bambusa sp.
India
—
—
MH858608
NG_069711
—
MW833997
—
—
7974
Cs. lavitskiae
CBS 530.68T = ATCC 18666 = IMI 133984
Plant debris
Ukraine
—
—
KU563624
HQ231997
—
MW833998
MW834271
—
Cs. viridescens
CBS 102433
Tilia sp.
Czech Republic
—
—
KJ676148
KJ676185
MW834182
MW833999
KJ676343
—
Cosmosporella cavisperma
CBS 172.31ET = NRRL 13996
Pinus sylvestris
Norway
—
—
MW827606
MW827645
JX171465
MW834000
—
—
Dead terminal branches connected
Slovenia
—
—
NR_145049
MH875034
MW834183
MW834001
KM231939
—
France
—
—
FJ474076
MH874353
MW834184
MW834002
HM626647
—
Cyanonectria buxi
ET
CBS 125551
with alive Buxus sempervirens
var. elegantissima
C. cyanostoma
CBS 101734ET = CBS 115512 = GJS 98-127
Buxus sempervirens
Dialonectria episphaeria
CBS 125494
Old ascomycete ascomata
Canada
—
—
MH863609
MH875085
MW834185
HQ897756
KM231953
—
D. ullevolea
CBS 125493
Ascomycete on Fagus americana
USA
—
—
KM231821
KM231696
—
HQ897782
KM231952
—
Fusarium acutatum
CBS 402.97T = BBA 69580 = FRC O-1117 = NRRL
Unknown
India
—
MW402459
—
—
MW402653
MW402768
MW402125
MW402323
13309
Agapanthus sp.
Australia
—
KU900611
—
—
KU900620
KU900625
KU900630
KU900635
F. ananatum
CBS 118516T = CMW 18685 = MRC 8165
Ananas comosus fruit
South Africa
—
LT996175
—
—
LT996188
LT996137
LT996091
LT996112
F. andiyazi
CBS 119857T = NRRL 31727
Sorghum bicolor soil debris
South Africa
—
LT996176
—
—
LT996189
LT996138
LT996092
LT996113
F. anthophilum
CBS 737.97 = DAOM 225119 = FRC M-1355 = IMI
Hippeastrum sp.
Germany
—
LT996177
—
—
LT996190
LT996139
LT996093
LT996114
F. bactridioides
NRRL 20476
Cronartium conigenum
USA
—
AF158343
—
—
Not public
Not public
AF160290
U34434
Begonia elatior hybrid
Germany
—
AF158346
—
—
LT996191
LT996140
AF160293
U61543
Soil
New Caledonia
—
—
U61674
U61648
JX171506
JX171619
PVQB02000800*
—
375325 = NRRL 13602
F. begoniae
T
CBS 403.97 = BBA 67781 = DAOM 225116 = IMI
375315 = NRRL 25300
19
F. beomiforme
CBS 740.97 = BBA 65829 = DAOM 225123 = IMI
375328 = NRRL 25174
(continued on next page)
REDELIMITED
NRRL 54463T
FUSARIUM
F. agapanthi
CROUS
20
Table 3. (Continued).
F. brevicatenulatum
Strain1
Substrate
CBS 404.97T = BBA 69197 = DAOM 225122 = IMI
ET AL.
Species name
GenBank accession number2
Country
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
Striga asiatica
Madagascar
—
MW834108
—
—
—
MN534295
MN533995
MN534063
Hibiscus cannabinus
Iran
—
—
U34581
U34552
JX171449
JX171563
—
—
Nerine bowdenii
Germany
—
KF466327
—
—
KF466394
KF466404
KF466415
KF466437
Pinus radiata
USA
—
KM231393
—
—
JX171510
HM068354
KM231943
KM232080
375329 = NRRL 25446
F. buharicum
CBS 796.70 = ATCC 24135 = BBA 11122 = DSM
62165 = FRC R-4955 = IMI 141195 = NRRL 13371
F. bulbicola
CBS 220.76T = BBA 12293 = BBA 63628 = DAOM
225114 = IMI 202877 = IMI 375322 = NRRL 13618
F. circinatum
CBS 405.97T = BBA 69720 = DAOM 225113 = IMI
375321 = MRC 7541 = NRRL 25331
F. coicis
NRRL 66233T = RBG 5368
Coix gasteenii
Australia
—
LT996178
—
—
KP083269
KP083274
KP083251
LT996115
F. compactum
NRRL 13829
River sediments
Japan
—
—
—
—
JX171460
JX171574
—
—
T
Musa sapientum
Costa Rica
—
AF158335
—
—
LT996192
JF741086
AF160282
U61548
F. concentricum
CBS 450.97 = BBA 64354 = CBS 833.85 = DAOM
225146 = IMI 375352 = NRRL 25181
F. cugenangense
CBS 130308 = NRRL 25387 = ATCC 26225
Human toe nail
New Zealand
—
—
MW827607
MW827646
JX171512
JX171625
MH485011
—
F. curvatum
CBS 744.97 = IMI 375335 = NRRL 22902
Pseudotsuga menziesii
USA
—
AF158365
—
—
LT996203
LT575065
AF160312
U34424
F. denticulatum
CBS 735.97 = NRRL 25302
Ipomoea batatas
USA
—
AF158322
—
—
LT996195
LT996143
AF160269
U61550
F. dlaminii
CBS 119860T = BBA 69859 = FRC M-1637 = MRC
Soil debris in cornfield
South Africa
—
AF158330
—
—
KU171681
KU171701
AF160277
U34430
CBS 146496 = CPC 30814
Unidentified tree
South Africa
—
MW834109
—
—
MW834186
MW834003
MW834272
MW834300
CBS 146497T = CPC 30815
Unidentified tree
South Africa
—
MW834110
—
—
MW834187
MW834004
MW834273
MW834301
3032 = NRRL 13164
F. echinatum
F. equiseti
CBS 245.61 = NRRL 20697
Beta vulgaris
Chile
—
—
MH858038
MH869603
JX171481
JX171595
—
—
F. flocciferum
CBS 831.85 = BBA 64346 = NRRL 25473
Triticum aestivum
Germany
—
—
—
MW827647
JX171514
JX171627
—
—
F. fracticaudum
CBS 137234PT = CMW 25237
Pinus maximonoii
Colombia
—
LT996179
—
—
LT996196
LT996144
KJ541059
KJ541051
F fractiflexum
NRRL 28852T
Cymbidium sp.
Japan
—
AF158341
—
—
Not public
LT575064
AF160288
AF160315
F. fredkrugeri
T
CBS 144209 = CPC 33747
Melhania acuminata rhizosphere
South Africa
—
LT996181
—
—
LT996199
LT996147
LT996097
LT996117
F. fujikuroi
CBS 221.76T = BBA 12428 = BBA 63630 = IHEM
Oryza sativa
Taiwan
—
—
MW827608
MW827648
MW834188
MW834005
AF160279
—
Oryza sativa
China
—
AF158332
—
—
JX171456
JX171570
AF160279
U34415
Zea mays
South Africa
—
KF466329
—
—
KF466396
KF466406
KF466417
KF466439
3821 = IMI 196086 = IMI 202879 = NRRL
13620 = NRRL 13998 = NRRL 22174
NRRL 13566 = ATCC 38941 = DAOM 225143 = IMI
300793 = IMI 375349 = NRRL 5538 = NRRL A-26483
F. globosum
CBS 428.97T = DAOM 214966 = FRC M-8014 = IMI
375330 = MRC 6647 = NRRL 26131 = PREM 51878
F. graminearum
CBS 123657 = NRRL 31084
Zea mays
USA
—
—
DQ459823
DQ459823
JX171531
JX171644
AY452957
—
F. heterosporum
CBS 720.79 = NRRL 20693
Claviceps purpurea on Lolium perenne
Netherlands
—
—
MW827609
MW827649
JX171480
JX171594
JAAGWP010000622.1*
—
Vicia faba
Germany
—
AF158366
—
—
JX171469
JX171583
AF008479
U34435
F. inflexum
Table 3. (Continued).
Species name
Strain1
Substrate
GenBank accession number2
Country
www.studiesinmycology.org
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
T
CBS 716.74 = ATCC 32213 = BBA 63203 = DAOM
225130 = DSM 63203 = IMI 375336 = NRRL 20433
F. konzum
CBS 119849T = MRC 8427
Sorghastrum nuttans
USA
—
LT996182
—
—
LT996200
LT996148
LT996098
LT996118
F. lactis
CBS 411.97ET = BBA 68590 = DAOM 225145 = IMI
Ficus carica
USA
—
AF158325
—
—
LT996201
LT996149
AF160272
U61551
Ulmus sp.
USA
—
—
—
—
JX171457
JX171571
—
—
375351 = NRRL 25200
F. lateritium
NRRL 13622 = NRRL A-26433
F. longipes
NRRL 20723 = IMI 265540
Unknown
England
—
—
—
—
JX171483
JX171596
—
—
F. mangiferae
NRRL 25226 = BBA 69662 = DAOM 225155 = IMI
Mangifera indica
Israel
—
AF158334
—
—
JX171509
HM068353
AF160281
U61561
"F." melanochlorum
CBS 202.65 = ATCC 16069 = BBA 9831 = DSM
Fagus sylvatica
Austria
—
—
MH858541
MH870179
JX171537
JX171649
—
—
Mangifera indica
Mexico
—
GU737389
—
—
LR792579
LR792615
GU737416
GU737308
Pennisetum typhoides
Namibia
—
AF158319
—
—
HM347136
EF470117
AF160266
U34428
304063 = IMI 375361
62248 = NRRL 36353
F. mexicanum
F. napiforme
NRRL 47473
T
CBS 748.97 = BBA 69861 = DAOM 225147 = FRC
M-3563 = IMI 375353 = MRC 4144 = NRRL 13604
F. nurragi
CBS 392.96 = NRRL 36452
Soil
Australia
—
—
MW827610
MW827650
JX171538
JX171650
JAALXI010000436.1*
—
F. nygamai
CBS 749.97T = ATCC 58555 = BBA 69862 = DAOM
Sorghum bicolor
Australia
—
AF158326
—
—
LT996202
EF470114
AF160273
U34426
225148 = FRC M-1375 = IMI 375354 = NRRL 13448
F. parvisorum
CBS 137236T
Pinus patula
Colombia
—
LT996183
—
—
—
LT996150
KJ541060
KJ541055
F. phyllophilum
CBS 216.76T = BBA 11730 = BBA 63625 = DAOM
Dracaena deremensis
Italy
—
KF466333
—
—
KF466399
KF466410
KF466421
KF466443
F. poae
NRRL 13714 = FRC T-503 = MRC 2181
Overwintered wheat
Canada
—
—
—
—
JX171458
JX171572
—
—
225132 = IMI 202874 = IMI 375338 = NRRL 13617
F. prieskaense
F. phyllophilum
CPC 30825
Aloidendron dichotomum
South Africa
—
MW834111
—
—
MW834189
MW834006
MW834274
MW834302
CBS 146498T = CPC 30826
Prunus spinosa
South Africa
—
MW834112
—
—
MW834190
MW834007
MW834275
MW834303
CBS 146499 = CPC 30827
Prunus spinosa
South Africa
—
MW834113
—
—
MW834191
MW834008
MW834276
MW834304
CBS 217.76 = BBA 11341 = BBA 63624 = DAOM
Cattleya sp.
Germany
—
KF466333
U34558
U34529
JX171504
JX171617
AF160280
KF466443
Solanum sp.
Ghana
—
AF158324
—
—
LT996204
LT996151
AF160271
U34427
Hordeum vulgare
Australia
—
—
DQ459871
DQ459871
JX171524
JX171637
AF212468
—
Pennisetum typhoides
Nigeria
—
AF158316
—
—
LT996205
LT996152
AF160263
U34421
Ficus carica
USA
—
KF466335
—
—
KF466401
KF466412
KF466423
KF466445
Pseudotsuga menziesii
Canada
—
—
U34565
U34536
JX171503
JX171616
MT409452
—
225133 = IMI 202873 = IMI 375339 = NRRL 22944
F. pseudocircinatum
CBS 449.97T = ATCC 24379 = BBA 69636 = CBS
126.73 = IMI 105384 = NRRL 22946
F. pseudograminearum
F. pseudonygamai
CBS 109956T = NRRL 28062
T
CBS 417.97 = BBA 69552 = FRC M-1166 = IMI
F. ramigenum
CBS 418.98T = BBA 68592 = DAOM 225137 = IMI
375343 = NRRL 25208
CBS 743.97 = DAOM 225128 = IMI 375334 = NRRL
22901
21
(continued on next page)
REDELIMITED
F. redolens
FUSARIUM
375342 = NRRL 13592
CROUS
22
Table 3. (Continued).
F. sacchari
Strain1
Substrate
CBS 223.76ET = BBA 63340 = DAOM 225138 = IMI
ET AL.
Species name
GenBank accession number2
Country
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
Saccharum officinarum
India
—
AF158331
—
—
JX171466
JX171580
AF160278
U34414
Solanum tuberosum
England
—
—
—
—
JX171493
JX171606
MW834277
—
202881 = NRRL 13999
F. sambucinum
CBS 146.95 = BBA 64226 = NRRL 22187 = NRRL
20727
F. sarcochroum
CBS 745.79 = BBA 63714 = NRRL 20472
Viscum album
Switzerland
—
—
MW827611
MW827651
JX171472
JX171586
MW834278
—
F. scirpi
NRRL 13402
Soil
Australia
—
—
GQ505681
GQ505681
JX171452
JX171566
GQ505592
—
F. sororula
CBS 137242T = CMW 40578
Pinus patula
Colombia
—
LT996184
—
—
LT996206
LT996153
KJ541067
KJ541057
Fusarium sp.
CBS 102163 = GJS 84-426
Bamboo
Venezuela
—
—
KM231812
KM231681
MW834193
MW834009
KM231940
—
F. sterilihyposum
NRRL 25623
Mango
South Africa
—
AF158353
—
—
MW402713
MN193897
AF160300
AF160316
F. stilboides
NRRL 20429 = ATCC 15662
Coffea sp.
Nyasaland
—
—
—
—
JX171468
JX171582
—
—
Zea mays
USA
—
AF158342
—
—
JX171486
JX171599
AF160289
U34417
Soil
Costa Rica
—
—
HQ897830
KM231680
JX171451
JX171565
—
—
Succisa pratensis
Germany
—
AF158344
—
—
LT996207
LT996154
AF160291
U34419
Striga hermonthica
Sudan
—
LT996185
—
—
LT996208
LT996155
KU711697
KU603909
F. subglutinans
CBS 747.97
ET
= BBA 62451 = DAOM 225141 = FRC
M-36 = MRC 8554 = NRRL 22016 = NRRL 22114
F. sublunatum
CBS 189.34T = BBA 62431 = DSM 62431 = NRRL
20840 = NRRL 13384
F. succisae
CBS 219.76ET = BBA 12287 = BBA 63627 = DAOM
225142 = IMI 202876 = IMI 375347 = NRRL 13613
F. sudanense
CBS 454.97T = BBA 65862 = NRRL 25451 = NRRL
26793
F. temperatum
NRRL 25622 = NRRL 26616
Zea mays
South Africa
—
AF158354
—
—
Not public
Not public
AF160301
AF160317
F. terricola
CBS 483.94T = FRC M-1650
Soil
Australia
—
KU603951
—
—
LT996209
LT996156
KU711698
KU603908
F. thapsinum
CBS 733.97 = DAOM 225109 = IMI 375317 = MRC
Sorghum bicolor
South Africa
—
LT996186
—
—
JX171487
JX171600
AF160270
U34418
6002 = NRRL 22045
F. tjaetaba
CBS 144400T = NRRL 66243 = RBG 5361
Sorghum interjectum
Australia
—
LT996187
—
—
MW834192
KP083275
KP083263
GU737296
F. torreyae
CBS 133858T = NRRL 54151
Torreya sp.
USA
—
—
HM068344
MW827652
JX171548
JX171660
HM068337
—
ET
F. tricinctum
CBS 393.93
Winter wheat culm base
Germany
—
—
HM068317
HM068317
JX171516
JX171629
AB674263
—
F. tupiense
NRRL 53984
Mangifera indica
Brazil
—
GU737377
—
—
LR792583
LR792619
GU737404
GU737296
F. udum
CBS 178.32 = BBA 1813 = DAOM 225111 = IMI
Lactarius pubescens
Germany
—
AF158328
—
—
LT996220
LT996172
AF160275
U34433
= BBA 64485 = NRRL 25481
375319 = NRRL 22949
F. venenatum
NRRL 22196 = BBA 65031
Zea mays
Germany
—
—
—
—
JX171494
JX171607
—
—
F. verticillioides
CBS 734.97 = BBA 62264 = IMI 375318 = NRRL
Zea mays
Germany
—
AF158315
—
—
LT996221
EF470122
AF160262
U34413
F. xylarioides
CBS 258.52ET = NRRL 25486
22172
Fusicolla acetilerea
Coffea sp.
Ivory Coast
—
—
—
—
JX171517
HM068355
AY707136
AY707118
Polluted soil
Japan
—
—
HQ897790
U88108
—
HQ897701
—
—
Table 3. (Continued).
Species name
Strain1
Substrate
GenBank accession number2
Country
www.studiesinmycology.org
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
T
BBA 63789 = IMI 181488 = NRRL
20827
Fu. aquaeductuum
BBA 63789 = IMI 181488 = NRRL 20827
Polluted soil
Japan
HQ897839
—
HQ897790
U88108
—
HQ897701
—
—
CBS 734.79 = BBA 63669 = NRRL 20686
Drinking water
Germany
—
—
MW827612
MW827653
JX171476
HQ897742
MW847905
—
CBS 268.53
Rubber tubing
Netherlands
—
—
MH857190
MH868728
—
—
—
—
CBS 837.85ET =BBA 64559 = NRRL 20865 = NRRL
Plug in water tap
Germany
—
—
KM231823
KM231699
—
—
—
KM232094
BBA 64317ET
Triticum aestivum
Germany
HQ897917
—
MH855265
MH866717
—
HQ897781
—
—
T
Avicennia marina
India
—
—
MK152510
MK152511
—
MK157022
—
MK376462
Cassia fistula
Thailand
—
—
MT215497
MT215549
—
—
—
—
Diatrypella sp., on Betula sp.
England
HQ897901
—
—
AF228352
—
HQ897765
—
—
37595
Fu. betae
Fu. bharatavarshae
NFCCI 4423
Fu. cassiae-fistulae
MFLUCC 19-0318T
Fu. epistroma
BBA 62201
ET
= ATCC 24369 = IMI 85601 = NRRL
20439 = NRRL 20461
Fu. gigantispora
Fu. matuoi
HKAS 101990
Bruguiera sp.
Thailand
—
—
MN047106
MN017870
—
—
—
—
MFLU 161206T
Avicennia marina
Thailand
—
—
MN047105
MN017876
—
—
—
—
CBS 581.78 = ATCC 18694 = MAFF 238445 = NRRL
Albizzia julibrissin
Japan
HQ897858
—
KM231822
KM231698
MW834194
HQ897720
KM231954
KM232093
Melogramma campylosporum on Carpinus sp.
England
—
—
KX897140
KY092489
—
HQ897720
—
MW834305
20427
Fu. melogrammae
CBS 141092T
Fu. meniscoidea
CBS 110189 = FRC E-0086
Soil
Australia
MW834043
—
MW827613
MW827654
—
MW834010
MW834279
MW834306
Fu. merismoides
CBS 186.34 = BBA 1867a = NRRL 20895
Acer sp.
Germany
—
—
MH855482
MH866963
—
—
—
—
Bone of wild boar
Belgium
—
—
MF628022
MF628021
—
MW834011
MW834280
MW834307
Melocactus zehntneri
Brazil
—
—
MW553789
MW553788
—
MW556626
MW556625
MW556624
T
Fu. ossicola
CBS 140161
Fu. quarantenae
URM 8367T = CBS 141541
T
Fu. septimanifiniscientiae
CBS 144935
Soil
Netherlands
—
—
MK069422
MK069418
—
—
MK077808
MK069408
Fu. siamensis
MFLUCC 17-2577T
Cassia fistula
Thailand
—
—
MT215498
MT215550
—
—
—
—
Fu. sporellula
CBS 110191 = FRC E-0139
Soil
South Africa
MW834044
—
MW827614
MW827655
—
MW834012
MW834281
MW834308
Fu. violacea
CBS 634.76T = BBA 62461 = NRRL 20896
Quadraspidiotus perniciosus
Iran
—
—
KM231824
U88112
MW834195
HQ897696
KM231956
KM232095
Geejayessia atrofusca
CBS 125482 = DAOM 238117
Staphylea trifolia
Canada
—
—
MH863592
MH875066
MW834196
HQ897775
MW834282
—
NRRL 22316
Staphylea trifolia
USA
—
—
AF178423
AF178392
JX171496
EU329502
AF178361
—
CBS 125502T
Celtis occidentalis
Canada
HM626625
—
HM626657
HM626669
MW834197
MW834013
HM626638
KM232074
G. cicatricum
CBS 125550
Dead twig connected with alive Buxus sempervirens
Slovenia
—
—
HM626654
HM626666
MW834198
HQ897697
HM626642
—
Dead twig
Slovenia
HQ728171
—
HQ728145
MH875038
—
HQ728153
HM626644
—
var. elegantissima
CBS 125552
T
CBS 132815
Protea sp.
South Africa
—
—
NR_152887
NG_070049
MW834199
MW834014
JX231119
—
I. destructans
CBS 264.65
Cyclamen persicum
Sweden
—
—
MH858563
KM515927
—
MW834015
JF735695
—
Luteonectria albida
CBS 102683 = GJS 99-73 = GJS 8522A
Tree bark
Costa Rica
—
—
MW827615
MH874402
MW834200
MW834016
MW834283
—
23
(continued on next page)
REDELIMITED
Ilyonectria capensis
FUSARIUM
G. celtidicola
CROUS
24
Table 3. (Continued).
Strain1
Substrate
GenBank accession number2
Country
acl1
T
L. nematophila
Macroconia bulbipes
ET AL.
Species name
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
NRRL 22152 = NRRL 13950
Woody stem bark
Jamaica
—
—
JABFEP010000142.1*
JABFEP010000142.1*
JX171492
JX171605
JABFEP010002685.1*
—
NRRL 54600
Unknown
Germany
—
—
JABFFA010000104.1*
JABFFA010000104.1*
JX171552
JX171664
JABFFA010003988.1*
—
CBS 146678 = CPC 37137
Erica sp. associated with Dimerosporiopsis engleriana
South Africa
MW834045
MW834114
MW827616
MW827656
MW834201
MW834017
—
MW834309
CBS 146679T = CPC 37138
Erica sp. associated with Dimerosporiopsis engleriana
South Africa
MW834046
MW834115
MW827617
MW827657
MW834202
MW834018
—
MW834310
Stylodothis sp. on unidentified tree
China
—
—
EF121864
EF121870
—
—
—
—
Rotten stem of bamboo associated with other fungi
China
—
—
EF121853
EF121869
—
—
—
—
T
Ma. cupularis
HMAS 173240
Ma. gigas
HMAS 173239T
Ma. leptosphaeriae
CBS 100001
Leptosphaeria on dead stem of Urtica dioica
Netherlands
HQ897891
MW834116
HQ897810
HQ897755
MW834203
HQ728164
KM231959
KM232097
Ma. papillionacearum
CBS 125495
Ascomycete on Fabaceae
USA
HQ897912
MW834117
HQ897826
MH875086
MW834204
HQ897776
—
KM232096
Ma. phlogioides
Ma. sphaeriae
Mariannaea elegans
CBS 125496
Quercus sp., branch in stream
USA
HQ897868
MW834118
MW827618
MW827658
MW834205
HQ897732
MW834284
MW834311
CBS 146500 = CPC 35388
Encephalartos sp. leaf
South Africa
MW834047
MW834119
MW827619
MW827659
MW834206
MW834019
—
MW834312
CBS 146501T = CPC 35389
Encephalartos sp. leaf
South Africa
MW834048
MW834120
MW827620
MW827660
MW834207
MW834020
—
MW834313
CBS 717.74
Pyrenomycete on Coronilla emerus
France
MW834049
MW834121
MW827621
MW827661
—
KM232390
—
KM232099
CBS 112770
Cucurbitaria laburni on Laburnum anagyroides
Austria
KM231061
KM231413
MW827622
MW827662
MW834208
MW834021
—
KM232098
DAOM 226709
Betula sp.
Canada
—
—
—
HQ843768
—
HQ897747
—
—
T
M. samuelsii
CBS 125515 = DAOM 235814
Soil
Guatemala
—
—
NR_137767
NG_060269
—
HQ897752
—
—
Microcera coccophila
CBS 310.34 = NRRL 13962
Scale insect
Italy
—
—
MH855540
KM231703
JX171462
JX171576
—
—
Mi. diploa
CBS 735.79 = BBA 61173 = NRRL 36545
Quadraspidiotus perniciosus
Iran
—
—
MW827623
MW827663
JX171463
JX171577
—
—
Mi. larvarum
CBS 738.79 = BBA 62239 = DSM 62239 = MUCL
Quadraspidiotus perniciosus
Iran
—
—
KM231825
KM231701
JX171473
JX171587
KM231957
—
Quadraspidiotus perniciosus on Prunus domestica
Iran
HQ897903
KM231409
MH861019
MH872790
—
HQ897767
—
MW834314
19033 = NRRL 20473
Mi. rubra
CBS 638.76IT = BBA 62460 = NRRL 20475; NRRL
22111; NRRL 22170
Microcera sp.
NRRL 26790
Parmelia rudecta
USA
—
—
—
—
JX171523
JX171636
—
—
Nectria cinnabarina
CBS 125165ET
Aesculus sp.
France
KM231074
—
HM484548
HM484562
—
KM232402
HM484527
—
"Nt." flavoviridis
CBS 124353 = BBA 65542 = NRRL 22093
Decorticated wood
USA
—
—
HQ897791
MW827664
MW834209
HQ897702
—
—
Neocosmospora acutispora
CBS 145461T = NRRL 22574 = BBA 62213
Coffea arabica
Guatemala
MW834050
MW834122
LR583700
LR583908
MW834210
LR583814
LR583593
—
CBS 146509 = CPC 37127
Citrus sinensis
South Africa
MW218004
MW218051
MW173041
MW173032
MW218097
MW446574
MW248740
—
CBS 146510T = CPC 37128
Citrus sinensis
South Africa
MW218005
MW218052
MW173042
MW173033
MW218098
MW446575
MW248741
—
Euwallacea fornicatus
India
—
—
EU329669
EU329669
MW834211
EU329503
FJ240350
—
N. addoensis
N. ambrosia
CBS 571.94
ET
= NRRL 22346 = BBA 65390 = MAFF
246287
N. ampla
N. bataticola
NRRL 20438 = IMI 296597
Xyleborus fornicatus
India
—
—
AF178397
AF178366
JX171470
JX171584
NIZV01000014.1*
—
CBS 202.32T = BBA 4170
Coffea sp.
German East Africa
MW834051
MW834123
LR583701
LR583909
MW834212
LR583815
LR583594
—
CBS 144397 = NRRL 22400 = BBA 64683
Ipomoea batatas
USA
MW218006
MW218053
AF178407
AF178376
MW218099
EU329509
AF178343
—
CBS 144398T = NRRL 22402 = BBA 64954 = FRC S-
Ipomoea batatas
USA
MW218007
MW218054
AF178408
AF178377
MW218100
FJ240381
AF178344
—
0567
Table 3. (Continued).
Species name
www.studiesinmycology.org
N. borneensis
Strain1
Substrate
CBS 145462ET = NRRL 22579 = BBA 65095 = GJS
Bark or recently dead tree
GenBank accession number2
Country
Indonesia
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
MW834052
MW834124
AF178415
AF178384
MW834213
EU329515
AF178352
—
85-197
CBS 144.25NT
Soil
Honduras
MW218008
MW218055
LR583704
LR583912
MW218101
LR583818
LR583597
—
CBS 392.66 = NRRL 25325 = BBA 69595
Bertholletia excelsa
Unknown
MW218009
MW218056
LR583705
LR583913
MW218102
LR583819
LR583598
—
N. brevicona
CBS 204.31ET = NRRL 22659 = BBA 2123
Gladiolus sp.
Indonesia
MW218010
MW218057
LR583707
LR583915
MW218103
LR583821
LR583600
—
N. brevis
CBS 130326 = NRRL 28009 = CDC B-5543
Human eye
USA
MW834053
MW834125
DQ094351
DQ236393
MW834214
EF470136
DQ246869
—
CBS 143228 = NRRL 54992 = UTHSC 09-1008
Stegostoma fasciatum
USA
MW218011
MW218058
KC808255
KC808255
MW218104
KC808354
KC808213
—
CBS 143229T = NRRL 54993 = UTHSC 09-1009
Stegostoma fasciatum
USA
MW218012
MW218059
KC808256
KC808256
MW218105
KC808355
KC808214
—
N. bostrycoides
N. catenata
N. citricola
CBS 146512 = CPC 37130
Citrus sinensis
South Africa
MW218014
MW218061
MW173047
MW173035
MW218107
MW446580
MW248746
—
CBS 146513T = CPC 37131
Citrus sinensis
South Africa
MW218015
MW218062
MW173048
MW173036
MW218108
MW446581
MW248747
—
T
N. crassa
CBS 144386 = MUCL 11420
Unknown
France
MW218016
MW218063
LR583709
LR583917
MW218109
LR583823
LR583604
—
N. cryptoseptata
CBS 145463T = NRRL 22412 = BBA 65024
Bark
French Guiana
MW834054
MW834126
AF178414
AF178383
MW834215
EU329510
AF178351
—
CBS 410.62 = NRRL 22658 = CECT 2864
Cucurbita viciifolia
Netherlands
MW834055
MW834127
LR583710
LR583918
MW834216
LR583824
DQ247640
—
CBS 616.66T = NRRL 22399 = BBA 64411
Cucurbita viciifolia
Netherlands
MW834056
MW834128
LR583711
LR583919
MW834217
LR583825
DQ247592
—
N. cucurbitae
CBS 518.82T
Human foot
Netherlands
MW218017
MW218064
AB190389
LR583920
MW218110
LR583826
LR583605
—
CBS 637.82
Human foot
Netherlands
MW218018
MW218065
LR583712
LR583921
MW218111
LR583827
LR583606
—
N. diminuta
CBS 144390T = MUCL 18798
Coelocaryon preusii
Unknown
MW834057
MW834129
LR583713
LR583922
MW834218
LR583828
LR583607
—
N. elegans
CBS 144395 = NRRL 22163 = MAFF 238540 = ATCC
Xanthoxylum piperitum
Japan
MW218019
MW218066
AF178394
AF178363
MW218112
EU329496
AF178328
—
Xanthoxylum piperitum
Japan
MW218020
MW218067
AF178401
AF178370
MW218113
FJ240380
AF178336
—
N. cyanescens
18690
CBS 144396ET = NRRL 22277 = MAFF
238541 = ATCC 42366
CBS 146523T = CPC 38310
Bouvardia sp. imported from Uganda
Netherlands
MW834058
MW834130
MW827624
MW827665
MW834219
MW834022
MW834285
—
CBS 146524 = CPC 38311
Bouvardia sp. imported from Uganda
Netherlands
MW834059
MW834131
MW827625
MW827666
MW834220
MW834023
MW834286
—
N. euwallaceae
CBS 135854T = NRRL 54722
Euwallacea sp.
Israel
—
—
JQ038014
JQ038014
JQ038021
JQ038028
JQ038007
—
N. falciformis
CBS 475.67T = IMI 268681
Human mycetoma
Puerto Rico
MW218021
MW218068
MG189935
MG189915
MW218114
LT960558
LT906669
—
CBS 121450
Declined grape vine
Syria
MW218022
MW218069
JX435211
JX435211
MW218115
JX435261
JX435161
—
NRRL 43529 = CDC 2006743575
Human cornea
USA
—
—
EF453117
EF453117
JX171541
JX171653
EF452965
—
N. epipeda
Switzerland
MW834060
MW834132
DQ094446
DQ236488
MW834221
EU329581
DQ246979
—
Citrus sinensis
Italy
MW834061
MW834133
LT746276
LT746276
MW834222
LT746341
LR583602
—
N. floridana
NRRL 62628T = MAFF 246849
Euwallacea interjectus
USA
—
—
KC691563
KC691563
KC691593
KC691624, KC691653
KC691535
—
N. gamsii
CBS 143207T = NRRL 32323 = UTHSC 99-205
Human bronchoalveolar lavage fluid
USA
MW834062
MW834134
DQ094420
DQ236462
MW834223
EU329622
DQ247103
—
CBS 143211 = NRRL 32794 = FRC S-1152
Humidifier coolant
USA
MW834063
MW834135
DQ094563
DQ236605
MW834224
EU329576
DQ246951
—
CBS 146502T = VG16 = CPC 37120
Citrus sinensis
South Africa
MW218023
MW218070
MW173063
MW173038
MW218116
MW446611
MW248762
—
Dying tree
Sri Lanka
MW834064
MW834136
KM231797
KM231664
—
LT960561
DQ247510
N. gamtoosensis
N. haematococca
ET
CBS 119600
= FRC S-1832
—
(continued on next page)
25
REDELIMITED
Human subcutaneous nodule
CPC 28194
FUSARIUM
CBS 109028T = NRRL 32437
N. ferruginea
CROUS
26
Table 3. (Continued).
Strain1
Substrate
GenBank accession number2
Country
acl1
N. hypothenemi
N. illudens
ET AL.
Species name
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
CBS 145464T = NRRL 52782 = ARSEF 5878
Hypothenemus hampei
Benin
MW218024
—
LR583715
LR583923
MW218117
JF741176
JF740850
—
CBS 145466 = NRRL 52783 = ARSEF 5879
Hypothenemus hampei
Uganda
MW218025
MW218071
MW827626
MW827667
MW218118
MW834024
MW834287
—
CBS 147303 = NRRL 22090 = BBA 67606 = GJS 82-
Beilschmiedia tawa
New Zealand
MW834065
MW834137
AF178393
AF178362
JX171488
JX171601
AF178326
—
CBS 353.87 = NRRL 22657
Gerbera sp.
Netherlands
MW218026
MW218072
LR583717
LR583925
MW218119
LR583831
DQ247639
—
CBS 833.97
Rosa sp.
Netherlands
MW218027
MW218073
LR583719
LR583927
MW218120
LR583833
LR583611
—
= FRC S-1837 = GJS 02-114
Branch of unidentified tree
Sri Lanka
MW834066
MW834138
LR583720
LR583928
MW834225
LR583834
LR583612
—
CBS 125722PT = FRC S-1836 = GJS 02-114
Branch of unidentified tree
Sri Lanka
MW834067
MW834139
JF433039
JF433039
MW834226
LR583835
DQ247515
—
98
N. ipomoeae
N. keleraja
N. keratoplastica
CBS 125720
CBS 490.63
PT
T
CBS 144389 = MUCL 18301
T
Human
Japan
MW218028
MW218074
LR583721
LR583929
MW218121
LT960562
LT906670
—
Greenhouse humic soil
Belgium
MW218029
MW218075
LR583722
LR583930
MW218122
LR583836
LR583613
—
N. kuroshio
CBS 142642
Euwallacea sp.
USA
MW834068
MW834140
LR583723
LR583931
MW834227
LR583837
KX262216
—
N. kurunegalensis
CBS 119599T = GJS 02-94
Recently cut tree
Sri Lanka
MW834069
MW834141
JF433036
JF433036
MW834228
LR583838
DQ247511
—
N. lerouxii
CBS 146514T = CPC 37132
Citrus sinensis
South Africa
MW218030
MW218076
MW173069
MW173039
MW218123
MW446617
MW248768
—
N. lichenicola
CBS 509.63 = MUCL 8050 = IMUR 410
Air
Brazil
MW834070
MW834142
LR583728
LR583936
MW834229
LR583843
LR583618
—
CBS 623.92ET
Human
Germany
MW834071
MW834143
LR583730
LR583938
—
LR583845
LR583620
—
CBS 117481T = NRRL 22389 = BBA 67587 = GJS 91-
Liriodendron tulipifera
USA
MW218031
MW218077
AF178404
AF178373
MW218124
EU329506
AF178340
—
—
N. liriodendri
148
N. longissima
CBS 126407T = GJS 85-72
Tree bark
New Zealand
MW834072
MW834144
LR583731
LR583939
MW834230
LR583846
LR583621
N. macrospora
CBS 142424T = CPC 28191
Citrus sinensis
Italy
MW218032
MW218078
LT746266
LT746281
MW218125
LT746331
LT746218
—
CPC 28193
Citrus sinensis
Italy
MW218033
MW218079
LT746268
LT746283
MW218126
LT746333
LT746220
—
Dead branch on live tree
Sri Lanka
MW834073
MW834145
JF433045
JF433045
MW834231
LT960563
DQ247513
—
—
N. mahasenii
CBS 119594T
ET
N. martii
CBS 115659
Solanum tuberosum
Germany
MW834074
MW834146
JX435206
JX435206
MW834232
JX435256
JX435156
N. merkxiana
CBS 146525T
Chrysanthemum sp. imported from Uganda
Netherlands
MW834075
MW834147
MW827627
MW827668
MW834233
MW834025
MW834288
—
CBS 146526
Chrysanthemum sp. imported from Uganda
Netherlands
MW834076
MW834148
MW827628
MW827669
MW834234
MW834026
MW834289
—
—
= FRC S-0679 = MRC 2198
CBS 135789T
Human pleural effusion
Greece
MW218034
MW218080
LR583738
LR583946
MW218127
LR583849
LR583627
CBS 143219 = NRRL 46708 = FMR 8634
Human foot
Spain
MW218035
MW218081
LR583744
LR583948
MW218128
LR583851
LR583629
—
N. mori
CBS 145467T = NRRL 22230 = MAFF 238539
Morus alba
Japan
MW834077
MW834149
DQ094305
DQ236347
MW834235
EU329499
AF178358
—
CBS 145468 = NRRL 22157 = MAFF 238538
Morus alba
Japan
MW834078
MW834150
DQ094306
DQ236348
MW834236
EU329493
AF178359
—
N. neerlandica
CBS 232.34T
Pisum sativum
Netherlands
MW834079
MW834151
MW827629
MW827670
MW834237
MW847903
MW847906
—
T
N. metavorans
N. nelsonii
CBS 309.75
Pisum sativum
Unknown
MW834080
MW834152
MW827630
MW827671
MW834238
MW847904
MW847907
—
N. nirenbergiana
CBS 145469T = NRRL 22387 = BBA 65023 = GJS 87-
Bark
French Guiana
MW834081
MW834153
AF178403
AF178372
—
EU329505
AF178339
—
N. noneumartii
CBS 115658T = FRC S-0661
Solanum tuberosum
Israel
MW218036
MW218082
LR583745
LR583949
MW218129
MW446618
LR583630
—
N. obliquiseptata
NRRL 62611 = MAFF 246845
Euwallacea sp.
Australia
—
—
KC691576
KC691576
KC691606
KC691637, KC691666
KC691548
—
127
Table 3. (Continued).
Species name
Strain1
Substrate
GenBank accession number2
Country
www.studiesinmycology.org
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
N. oblonga
CBS 130325T = NRRL 28008 = CDC B-4701
Human eye
USA
MW834082
MW834154
LR583746
LR583950
MW834239
LR583853
LR583631
—
N. oligoseptata
CBS 143241T = NRRL 62579 = FRC
Euwallacea validus
USA
MW834083
MW834155
KC691566
KC691566
KC691596
LR583854
KC691538
—
Solanum tuberosum
Argentina
MW834084
MW834156
LR583747
LR583951
MW834240
LR583855
DQ247549
—
S-2581 = MAFF 246283
N. paraeumartii
CBS 487.76T = NRRL 13997 = BBA 62215
T
N. parceramosa
CBS 115695
Soil
South Africa
MW218037
MW218083
JX435199
JX435199
—
JX435249
JX435149
—
N. perseae
CBS 144142T = CPC 26829
Persea americana
Italy
MW218038
MW218084
LT991940
LT991947
MW218130
LT991909
LT991902
—
N. petroliphila
N. phaseoli
N. piperis
CBS 203.32 = NRRL 13952
Pelargonium sp.
South Africa
MW218039
MW218085
DQ094320
DQ236362
MW218131
LR583857
DQ246835
—
CBS 224.34 = NRRL 28579
Human toenail
Cuba
MW218040
MW218086
DQ094383
DQ236425
MW218132
LR583858
DQ246910
—
CBS 265.50
Phaseolus sp.
USA
MW834085
MW834157
LR583750
LR583954
—
KJ511278
FJ919464
—
NRRL 22276 = ATCC 38466
Phaseolus vulgaris
USA
—
—
EU329668
EU329668
JX171495
JX171608
AY220186
—
CBS 145470T = NRRL 22570 = GJS
Piper nigrum
Brazil
MW834086
MW834158
AF178422
AF178391
MW834241
EU329513
AF178360
—
89-14 = CML 1888
N. pisi
CBS 123669ET = NRRL 45880 = ATCC MYA-4622
Progeny of parentals from Pisum sativum and soil
USA
MW834087
MW834159
LR583753
LR583957
MW834242
LR583862
LR583636
—
CBS 142372
Trifolium subterraneum
Germany
MW834088
MW834160
LR583755
LR583959
MW834243
LR583864
KY556454
—
—
N. plagianthi
NRRL 22632 = GJS 83-146
Hoheria glabrata
New Zealand
—
—
AF178417
AF178386
JX171501
JX171614
AF178354
N. protoensiformis
CBS 145471T = NRRL 22178 = GJS 90-168
Dicot tree
Venezuela
MW834089
MW834161
AF178399
AF178368
MW834244
EU329498
AF178334
—
N. pseudensiformis
CBS 130.78 = NRRL 22575 = NRRL 22653
Cocos nucifera
Indonesia
MW834090
MW834162
LR583759
LR583963
MW834245
LR583868
DQ247635
—
N. pseudopisi
CBS 266.50
Pisum sativum
Unknown
MW834091
MW834163
MW827631
MW827672
MW834246
MW834027
MW834290
—
N. pseudoradicicola
CBS 145472T = NRRL 25137 = ARSEF 2313
Diseased cocoa pods
Papua New Guinea
MW218041
MW218087
JF740899
JF740899
MW218133
JF741084
JF740757
—
T
N. quercicola
CBS 141.90 = NRRL 22652
Quercus cerris
Italy
MW834092
MW834164
LR583760
LR583964
MW834247
LR583869
DQ247634
—
N. rectiphora
CBS 125726 = FRC S-1842
Dead tree
Sri Lanka
MW834093
MW834165
JF433043
JF433043
MW834248
MW834028
JF433026
—
N. regularis
N. rekana
CBS 125727T = GJS 02-89 = FRC S-1831
Dead tree
Sri Lanka
MW834094
MW834166
JF433034
JF433034
MW834249
LR583871
DQ247509
—
CBS 190.35
Phaseolus sp.
USA
MW834095
MW834167
LR583762
LR583966
MW834250
LR583872
LR583642
—
CBS 230.34T
Pisum sativum
Netherlands
MW834096
MW834168
LR583763
LR583967
—
MW834029
LR583643
—
CMW 52862T
Euwallacea perbrevis
Indonesia
—
—
MN249094
—
—
MN249137, MN249108
MN249151
—
T
CBS 145473 = NRRL 22395 = BBA 65682
Bark
Venezuela
—
MW834169
AF178405
LR583968
MW834251
EU329507
AF178341
—
CBS 114067T = GJS 89-70
Bark
Guyana
MW834097
MW834170
LR583764
LR583969
MW834252
LR583874
LR583644
—
CBS 119601 = GJS 98-135
Populus nigra
France
MW834098
MW834171
LR583765
LR583970
MW834253
LR583875
LR583645
—
CBS 123846T = GJS 04-147
Liriodendron tulipifera
USA
MW834099
MW834172
LR583766
LR583971
MW834254
LR583876
LR583646
—
Solanum tuberosum
Slovenia
MW218042
MW218088
KT313633
KT313633
MW218134
KT313623
KT313611
—
N. silvicola
N. solani
CBS 140079
ET
= NRRL 66304 = GJS
09-1466 = FRC S-2364
CBS 145474T = NRRL 28541 = UTHSC 98-1305
Human synovial fluid
USA
MW218045
MW218091
EU329674
EU329674
MW218137
EU329542
DQ246882
—
N. stercicola
CBS 142481T = DSM 106211
Compost yard debris
Germany
MW834100
MW834173
LR583779
LR583984
MW834255
LR583887
LR583658
—
(continued on next page)
27
REDELIMITED
N. spathulata
FUSARIUM
N. robusta
N. samuelsii
CROUS
28
Table 3. (Continued).
Strain1
Substrate
GenBank accession number2
Country
acl1
N. suttoniana
N. tonkinensis
N. tuaranensis
ET AL.
Species name
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
CBS 144388 = MUCL 18299
Greenhouse humic soil
Belgium
MW834101
MW834174
LR583780
LR583985
MW834256
LR583888
LR583659
—
CBS 143214T = NRRL 32858
Human wound
USA
MW218046
MW218092
DQ094617
DQ236659
MW218138
EU329630
DQ247163
—
CBS 143224 = NRRL 54972
Equine eye
USA
MW218047
MW218093
MG189940
MG189925
MW218139
KC808336
KC808197
—
CBS 115.40T
Musa sapientum
Vietnam
MW218048
MW218094
MG189941
MG189926
MW218140
LT960564
LT906672
—
CBS 118931
Solanum lycopersicum
UK
MW218049
MW218095
LR583784
LR583989
MW218141
LR583891
LR583662
—
Hevea brasiliensis damaged by unknown ambrosia
Malaysia
—
—
KC691570
KC691570
KC691600
KC691631, KC691660
KC691542
—
South Africa
—
—
AF178412
AF178381
JX171497
JX171610
AF178348
—
NRRL 22231T = ATCC 16563 = MAFF 246842
beetle
N. vasinfecta
CBS 325.54 = ATCC 16238 = IFO 7591 = IMI
Soil
251386 = NRRL 22436
CBS 446.93 = IMI 316967 = NHL 2919
Soil
Japan
MW834102
MW834175
LR583791
LR583996
MW834257
LR583898
LR583670
—
CBS 533.65 = IMI 302625
Unknown
India
MW834103
MW834176
LR583792
LR583997
MW834258
LR583899
LR583671
—
Neonectria coccinea
CBS 125484
Fagus sylvatica
Germany
—
—
HQ897832
MH875068
MW834259
HQ897785
—
—
Ne. ditissima
CBS 125486
Fagus americana
Canada
—
—
HQ897824
MH877864
—
HQ897774
—
—
Nothofusarium devonianum
CBS 147304T = NRRL 22134
Ruscus aculeatus
United Kingdom
—
—
MW827632
MW827673
JX171490
JX171603
MW834291
—
Pseudofusicolla belgica
CBS 147300 = IHEM 5322
Recycled water from air-conditioning humidifier
Belgium
—
—
KJ125590
KJ126478
—
KP835473
KJ126182
—
CBS 147301T = IHEM 2413
Recycled water, spray humidifier in air-conditioned
Belgium
—
—
KJ125588
KJ126476
—
KP835474
KJ126180
—
CBS 147302 = IHEM 2440
Humidifier water from air-conditioning
Belgium
—
—
KJ125589
KJ126477
—
KP835475
KJ126181
—
IHEM 2105
Recycled humidifier water from airconditioning
Belgium
—
—
KP835478
KP835480
—
KP835476
KP835484
—
building
Rectifusarium robinianum
CBS 430.91T = NRRL 25729
Robinia pseudoacacia
Germany
—
—
KM231794
NG_058096
JX171520
JX171633
KM231923
—
R. ventricosum
CBS 748.79T = BBA 62452 = NRRL 20846 = NRRL
Wheat field soil
Germany
—
—
HQ897816
KM231658
JX171484
JX171597
KM231924
—
Rugonectria castaneicola
CBS 128360
Bark
China
—
—
MH864901
MH876352
MW834260
MW834030
MW834292
—
22113
Ru. neobalansae
CBS 125120 = GJS 85-219
Dead tree
Indonesia
—
—
KM231750
HM364322
—
MW834031
KM231874
—
Ru. rugulosa
CBS 126565 = GJS 09-1245
Dead wood
Venezuela
—
—
KM231749
MH877897
MW834261
MW834032
KM231873
—
Setofusarium setosum
Scolecofusarium ciliatum
CBS 574.94 = BBA 65063
Unknown
French Guiana
—
—
MW827633
MW827674
MW834262
MW834033
MW834293
—
CBS 635.92ET = GJS 88-12 = NRRL 36526
Tree bark
French Guiana
—
—
MW827634
MW827675
JX171539
JX171651
MW834294
—
CBS 155.86 = NRRL 22284
Hordeum vulgare mouldy grain, associated with scale
Denmark
—
—
MW827635
MW827676
MW834263
MW834034
MW834295
—
Germany
—
—
MW827636
MW827677
MW834264
MW834035
MW834296
—
insects
CBS 191.65NT = ATCC 16068 = ATCC 24137 = BBA
Fagus sylvatica
9661 = DSM 62172 = IMI 112499 = NRRL 20431
CBS 144385 = IHEM 2989
Fagus sylvatica
Belgium
—
—
KJ125591
KJ126479
MW834265
KP835472
MW834297
—
CBS 125489
Unidentified ascomycete on Betula sp.
Canada
HQ897875
—
HQ897805
KM231689
—
HQ897739
KM231944
—
St. carpini
DAOM 235819
Melanconis spodiaea on Carpinus betulus
Austria
HQ897909
—
HQ897823
—
—
HQ897773
—
—
St. corniculata
CBS 125491T
Unidentified ascomycete on Carpinus sp.
Germany
HQ897915
—
HQ897829
KM231691
—
HQ897779
KM231946
—
Diaporthe sp. on Frangula alnus
Ukraine
MW834104
—
MW827637
—
—
MW834036
—
—
Stylonectria applanata
St. hetmanica
T
CBS 147305 = CPC 38725
Table 3. (Continued).
Species name
Strain1
Substrate
GenBank accession number2
Country
www.studiesinmycology.org
acl1
St. norvegica
CaM
ITS
LSU
rpb1
rpb2
tef1
tub2
CBS 147306 = CPC 38848
Dothiorella sarmentorum on Acer platanoides
Ukraine
MW834105
—
MW827638
—
—
MW834037
—
—
CBS 139239T
Dead sporodochia of fusarium state on pyrenomycete
Norway
MW834106
—
KR605485
—
—
MW834038
—
—
Norway
MW834107
—
MW827639
—
—
MW834039
—
—
Picea abies
Germany
HQ897919
—
HQ897831
—
—
HQ897783
—
—
(presumably Amphiporthe sp.)
CBS 139242
On sporodochia of fusarium-like on unidentified
pyrenomycete
St. purtonii
DAOM 235818
T
St. qilianshanensis
HMAS 255803
Unknown ascomycete on Picea asperata
China
MT087289
—
—
—
—
MT087288
—
—
St. wegelianiana
CBS 125490
Hapalycystis bicaudata on Ulmus glabra
Austria
HQ897890
—
KM231817
KM231690
—
HQ897754
KM231945
—
Thelonectria discophora
CBS 125487
Aesculus hippocastanum
Germany
—
—
HQ897789
MW827678
MW834266
HQ897700
MW834298
—
T. olida
CBS 215.67NT = ATCC 16548 = DSM 62520 = IMI
Asparagus officinalis
Germany
—
—
MW827640
MW827679
MW834267
MW834040
MW834299
—
116873
Tumenectria laetidisca
CBS 100284
Bamboo
Japan
—
—
KJ022017
KJ022066
—
MW834041
KJ022400
—
CBS 101909ET
Bamboo
Jamaica
—
—
KJ022018
KJ022067
MW834268
MW834042
KJ022401
—
1
ARSEF: Collection of entomopathogenic fungal cultures, US Department of Agriculture (USDA), Agricultural Research Service (ARS), Ithaca, NY, USA; ATCC: American Type Culture Collection, Manassas, VA, USA; BBA: Biologische Bundesanstalt
für Land- und Forstwirtschaft, Institut für Mikrobiologie, Berlin, Germany; CBS: Westerdijk Fungal Biodiverity Institute (WI), Utrecht, The Netherlands; CDC: Centers for Disease Control and Prevention, Atlanta, GA, USA; CECT: Spanish Type Culture
Collection, Universidad de Valencia, Burjassot, Spain; CML: Coleç~ao Micologica de Lavras, Universidade Federal de Lavras, Minas Gerais, Brazil; CMW: Culture collection at the FABI, University of Pretoria, South Africa; CPC: Collection of P.W. Crous,
held at WI; DAOM: Canadian National Mycological Herbarium and Culture Collection, AAFC, Ottawa, Ontario, Canada; DSM: DSMZ-Deutsche Sammlung von Mikroorganismen und Zellkulturen GmbH, Braunschweig, Germany; FMR: Facultat de
Medicina i Ciencies de la Salut, Reus, Spain; FRC: Fusarium Research Center, Pennsylvannia State University, PA, USA; GJS: Collection of G.J. Samuels, USDA-ARS, USA; HKAS: Herbarium of Cryptogams, Kunming Institute of Botany, Kunming,
China; HMAS: Herbarium Mycologicum Academiae Sinicae, Chinese Academy of Sciences, Beijing, China; IFO: Institute for Fermentation, Osaka, Yodogawa-ku, Osaka, Japan; IHEM: Biomedical Fungi and Yeasts Collection, Scientific Institute of
Public Health, Belgium; IMI: CABI Bioscience, Egham, UK; IMUR: Institute of Mycology, University of Recife, Recife, Brazil; MAFF: Ministry of Agriculture, Forestry and Fisheries, Tsukuba, Ibaraki, Japan; MFLU: Mae Fah Luang University herbarium,
Chiang Rai, Thailand; MRC: Microbial Culture Collection, South African Medical Research Council, Tygerberg, South Africa; MUCL: Mycotheque de lUniversite Catholique de Louvain, Louvain-la-Neuve, Belgium; NHL: National Institute of Hygienic
Sciences, Tokyo, Japan; NRRL: Agricultural Research Service Culture Collection, National Center for Agricultural Utilization Research, USDA, Peoria, IL, USA; RBG: Royal Botanic Gardens Trust, Sydney, New South Wales, Australia; URM: Micoteca
do Departmento de Micologia, Universidade Federal de Pernambuco, Recife, Brazil; UTHSC: Fungus Testing Laboratory, Department of Pathology, University of Texas Health Science Center, San Antonio, USA. ET: Ex-epitype; IT: Ex-isotype; NT: Exneotype; PT: Ex-paratype; T: Ex-type.
2
acl1 = ATP citrate lyase; CaM = Calmodulin; ITS = Internal transcribed spacer region of the nrDNA; LSU = 28S large subunit of the nrDNA; rpb1 = RNA polymerase largest subunit; rpb2 = RNA polymerase second largest subunit; tef1 = translation
elongation factor 1-alpha; tub2 = Beta-tubulin. Sequences generated in this study are shown in bold; Not public = sequences not available at GenBank, obtained from K. O’Donnell's alignment datasets; * = Whole genome sequence contig accession
numbers.
FUSARIUM
REDELIMITED
29
CROUS
ET AL.
Table 4. Summary of phylogenetic information generated in this study.
Analysis
Nuclear
region
Length + gap
PI
Var.
BI unique
site patterns
Model (AIC)
Model (BIC)
ML -InL (IQ)
Generic delimitation
ITS
LSU
rpb1
rpb2
tef1
Combined
626
435
1 371
1 761
699
4 892
249
90
705
834
448
2 326
310
109
755
892
489
2 555
378
118
823
989
551
2 859
GTR+I+G
GTR+I+G
GTR+I+G
GTR+I+G
GTR+I+G
n/d
TIMe+I+G4
TIM2+F+I+G4
TIM3e+I+G4
GTR+F+I+G4
TIM2e+I+G4
n/d
-3099.276
-15223.682
-27263.487
-8493.378
-40875.16
-94954.982
Ex-type strains
rpb1
rpb2
tef1
Combined
1 724
1 789
859
4 372
980
788
463
2 231
550
916
301
1 767
1 358
1 056
700
3 114
GTR+I+G
GTR+I+G
GTR+I+G
n/d
TIM3e+R4
TIM2e+R6
GTR+F+I+G4
n/d
-37377.092
-44286.314
-25546.628
-113450.62
Fusarium fujikuroi
species complex
CaM
rpb1
rpb2
tef1
tub2
Combined
545
1 534
1 541
676
488
4 794
76
201
241
137
76
731
131
340
362
243
150
1 226
150
344
365
305
182
1 346
SYM+G
SYM+G
GTR+I+G
GTR+I+G
SYM+G
n/d
G4TNe+G4
TIM2e+G4
TNe+G4
TNe+I+G4
TNe+G4
n/d
-4032.663
-5669.761
-7415.729
-2062.906
-1930.688
-22043.423
Fusicolla
acl1
ITS
LSU
rpb2
tef1
tub2
Combined
908
518
476
1 702
476
484
4 564
153
54
34
258
109
83
691
346
111
69
447
216
162
1 351
241
128
72
359
202
159
1 161
GTR+G
GTR+I+G
K80+I
SYM+I+G
SYM+I
GTR+G
n/d
TNe+I
TIM2e+I+G4
K80+R2
TIM2e+G4
TIM2+F+G4
K80+G4
n/d
-3238.214
-1704.698
-1229.69
-5692.247
-2051.471
-1780.157
-16092.82
Macroconia
acl1
CaM
ITS
LSU
rpb1
rpb2
tub2
Combined
801
551
540
694
814
778
519
4 697
207
150
36
21
116
160
101
791
332
223
64
37
182
618
168
1 624
205
159
94
3
96
151
142
850
SYM+I
K80+I
GTR+I
GTR+I
SYM+G
SYM+I
SYM+G
n/d
K80+I
K80+I
TNe+G4
TNe+I
TNe+G4
TNe+G4
TNe+G4
n/d
-1241.031
-2092.487
-2259.518
-3097.338
-2620.526
-1784.381
-1205.535
-14388.257
Neocosmospora
acl1
CaM
ITS
LSU
rpb1
rpb2
tef1
Combined
630
586
476
482
1 492
1 613
688
5 967
173
171
119
36
390
449
230
1 568
271
231
357
63
506
564
323
2 315
297
280
211
76
636
621
370
2 491
K80+I+G
HKY+I+G
GTR+I+G
GTR+I+G
GTR+I+G
GTR+I+G
GTR+I+G
n/d
TIM3e+I+G4
TIM2e+R3
TNe+G4
TIM3e+I+G4
TIM2e+R3
TIM2e+I+G4
K80+G4
n/d
-13572.514
-5595.928
-4164.678
-10056.777
-2888.743
-1496.116
-4087.046
-46528.083
Stylonectria
acl1
ITS
rpb2
Combined
897
544
1 631
3 072
254
21
183
458
426
39
442
907
416
47
299
762
GTR+G
HKY+I
GTR+G
n/d
K80+I
TNe+G4
TNe+G4
n/d
-1022.317
-5181.494
-4061.543
-10441.718
PI = parsimony informative characters; Var. = variable characters; BI = Bayesian inference; Model (AIC) = evolutionary model selected by MrModeltest; Model
(BIC) = evolutionary model selected by ModelFinder in IQ-TREE; ML -InL (R) = best tree score determined using RAxML; ML -InL(IQ) = best tree score determined in IQTREE. F = Empirical base frequencies; G = Rate of discrete Gamma categories; GTR = General time reversible model; HKY = Unequal transition/transversion rates and
unequal base frequencies; I = Proportion of invariable sites; K80 = Unequal transition/transversion rates and equal base frequencies; R = FreeRate model;
SYM = Symmetric model; TIM2 = Transition model, AC = AT, CG = GT and unequal base frequencies; TIM2e = TIM2 with equal base frequencies; TIM3e = Transition
model, AC = CG, AT = GT with equal base frequencies; TNe = Unequal transition/transversion rates with unequal purine/pyrimidine rates and equal base frequencies;
TPM2 = AC = AT, AG = CT, CG = GT and equal base frequencies.
generalized time-reversible (GTR) model and applying the
partitioning option, which estimates the Gamma-shape
parameter and the proportion of invariable sites for every gene
separately. Again 1 000 bootstraps were calculated to estimate
branch support. Bayesian inference was conducted using
MrBayes v. 3.2.7 (Ronquist & Huelsenbeck 2003) with the
partitioned dataset. The Gamma-shape parameter and proportion of invariable sites were estimated independently for
each partition. MrBayes was run for 5 M generations with
every 500th tree sampled and a burn-in of 30 % of the sampled
trees to ensure sampling from the stationary phase. All other
parameters were set to default.
30
Morphology
Morphological characterisation followed standard procedures
as described by Leslie & Summerell (2006) using PDA, SNA
(Nirenberg 1976), and CLA (Fisher et al. 1982). Colony
morphology and pigmentation were evaluated on PDA after 7
to 14 d at 25 °C in darkness. Colour notation was based on
the colour charts of Rayner (1970). Fungarium specimens were
rehydrated in 3 % aqueous KOH for a few minutes and then
rinsed by replacing the KOH solution with sterile distilled water
or 100 % lactic acid (Samuels 1976a, b, Samuels et al. 1990).
Unless otherwise mentioned, micromorphological characters
FUSARIUM
were examined using water as mounting medium on a Zeiss
Axioskop 2 plus or a Nikon Eclipse 80i, both equipped with
Differential Interference Contrast (DIC) optics and a Nikon
AZ100 dissecting microscope all fitted with Nikon DS-Ri2 highdefinition colour digital cameras to photo-document fungal
structures. Measurements were taken using the Nikon software
NIS-elements D v. 4.50. The dimensions of at least 30
randomly selected elements were recorded for every fungal
structure. Average, standard deviation, and maximum–
minimum values were determined for elements using five or
more individual measurements. To facilitate the comparison of
relevant micro- and macroconidial features, composite photo
plates were assembled from separate photo micrographs using
Adobe Photoshop CC.
RESULTS
DNA phylogeny
The results of DNA evolutionary model selection, alignment
length, and composition as well as tree statistics for all the
multimarker datasets included in this study are summarised in
Table 4.
Re-analysis of the dataset of Geiser et al. (2021): A reanalysis of the dataset of Geiser et al. (2021) revealed no major
differences in the ML analysis. However, in ME analysis
(Supplementary Fig. S3), we found that the backbone architecture is less solid than previously thought and a large monophyletic clade containing Neocosmospora, Albonectria, and
several other genera formed as sister group to Fusarium s. str.
with strong support.
Generic delimitation of fusarioid taxa in Nectriaceae: The analyses included nectriaceous taxa historically ascribed to Fusarium s. lat., including several recently segregated fusarioid genera
(Gr€afenhan et al. 2011, Schroers et al. 2011, Lombard et al. 2015),
cylindrocarpon-like taxa (Chaverri et al. 2011), and the closely
related – although morphologically distinct – phylogenetic relatives Cosmospora and Mariannaea. Analyses using ML and BI of
the individual genes and combined datasets resulted in phylogenies with congruent topologies. Therefore, only IQ-TREE-ML
topologies are presented with RAxML-BS, UFboot2-BS, BI-PP
and gCF support values superimposed (Fig. 7).
The combined alignment of ITS, LSU, rpb1, rpb2 and tef1
comprised 100 strains representing 92 species, including the
outgroup Nectria cinnabarina (CBS 125165). Phylogenetic analyses resolved 27 monophyletic genera, of which 19 contain taxa
with fusarioid asexual morphs and nectria- or cosmospora-like
sexual morphs. Of these, 15 clades represent currently
described genera, namely Albonectria, Atractium, Bisifusarium,
Cosmosporella, Cyanonectria, Dialonectria, Fusarium, Fusicolla,
Geejayessia, Macroconia, Microcera, Neocosmospora, Pseudofusicolla, Rectifusarium, and Stylonectria. The fusarioid
genera Cosmosporella and Dialonectria, both of which have
cosmospora-like sexual morphs, clustered as sister clades to
Cosmospora; the latter, however, differ by having acremoniumlike asexual morphs. The remaining four clades with fusarioid
morphology represent undescribed taxa, formally described
here as the new genera Luteonectria, Nothofusarium, Scolecofusarium, and Setofusarium. A strongly supported clade
comprising six cylindrocarpon-like genera (Corinectria, Ilyonectria, Neonectria, Rugonectria, Thelonectria, and Tumenectria)
www.studiesinmycology.org
REDELIMITED
and the genus Mariannaea resolved as successive sister groups
to the F1 node.
Twenty-four out of the 27 genera included in the analysis
resolved as fully supported clades, including all but one
(Nothofusarium with RAxML-BS = 99 % / UFboot-BS = 92 % /
PP = 1) of the fusarioid genera (Fig. 7). The two remaining
clades (Cosmospora and Neonectria), however, received high
statistical support (RAxML-BS = 99 % / UFboot-BS = 100 % /
PP = 1 and RAxML-BS = 92 % / UFboot-BS = 95 % / PP = 1,
respectively). Similarly, the combined phylogeny resolved most
of the internal nodes with high to full bootstrap and Bayesian PP
support including the nodes F1, F2, and F3 sensu Geiser et al.
(2013, 2021) and O'Donnell et al. (2013, 2020). Nevertheless,
only F3 was resolved with confidence by all the individual marker
phylogenies (Supplementary Fig. S4). Node F2 was resolved
with high statistical support in the ITS, rpb1, and tef1 phylogenies, but unsupported in the LSU and rpb2 trees, while node F1
resolved without bootstrap and PP support in the ITS, rpb1, rpb2,
and tef1 phylogenies and was not recovered in the LSU tree.
To illustrate shared and differential morphological characters
among the different genera recognised here, a tree was constructed based on the phylogeny presented in Fig. 7, and the main
morphological features were plotted for each clade/genus (Fig. 8).
In addition to the genera recognised above, the recently
described aquatic fusarioid genus Varicosporella (Lechat &
Fournier 2015) is not included in the phylogenetic analyses due
to lack of available sequences; however, is accepted here based
on its distinct morphology. Non-molecular character variation
supports the phylogenetic relationship of fusarioid taxa in Nectriaceae. The 20 fusarioid genera in Nectriaceae are characterised by phialidic asexual morphs with variously septate, falcate
conidia with diverse degrees of foot-shaped basal cell development, formed on aerial or sporodochial conidiophores, with or
without additional production of microconidia. Characteristic
macroconidial foot-shaped basal cells are found most of the time,
but not always (e.g., Fusarium caeruleum) in clade F1, i.e.,
Albonectria, Bisifusarium, Cyanonectria, Fusarium, Geejayessia,
Luteonectria, Neocosmospora, Nothofusarium, Rectifusarium,
and Setofusarium, but are also present in distantly related genera
such as Cosmosporella, Dialonectria, Macroconia, and Microcera. Setofusarium is clearly recognisable by the formation of
thick-walled, slightly rugose setae on its sporodochia.
With the exception of Atractium, Bisifusarium, Nothofusarium,
and Pseudofusicolla, most fusarioid genera have sexual morphs,
usually seen as nectria-like or cosmospora-like perithecial
ascomata. The ascomata show various colour reactions or no
reaction in KOH; the colour reaction correlates with the phylogenetic distribution. Apart from Albonectria, with white to pale
yellow perithecia, Luteonectria, with white to buff coloured perithecia and Fusarium, with dark blue-violet to black perithecia,
Fusicolla, with yellow-orange perithecia and Varicosporella, with
yellow perithecia, the rest of fusarioid genera all present orange
to red perithecial ascomata. Going beyond this prototypical
group, perithecia of Cyanonectria species are often unequally
red to dark blue, while those of Geejayessia can be bright red or
black. Anatomically, two types of perithecial walls can be
distinguished among the known fusarioid genera, based on wall
thickness: thin-walled perithecia, in which a single region can be
identified, and thick-walled perithecia, on which distinctive inner
and outer regions can be recognised (but see Schroers et al.
2011 for differing interpretations). The former is seen in Cosmosporella, Cyanonectria, Dialonectria, Fusicolla, Geejayessia,
31
CROUS
32
ET AL.
FUSARIUM
Luteonectria, Macroconia, Microcera, Scolecofusarium, and
Varicosporella; and the latter is found in Albonectria, Fusarium,
Neocosmospora, Rectifusarium, Setofusarium and Stylonectria.
With the exception of Rectifusarium and Stylonectria, the perithecial surface of the thick-walled genera is typically warted;
nevertheless, those of Setofusarium often present additional
scaly protrusions, while smooth perithecia can be rarely found in
Neocosmospora (i.e., N. vasinfecta). Additionally, both Cyanonectria and Geejayessia most commonly have smooth perithecial walls. The remaining genera, that is Cosmosporella,
Dialonectria, Fusicolla, Luteonectria, Macroconia, Microcera,
Rectifusarium, Scolecofusarium, Stylonectria, and Varicosporella, all form smooth-walled perithecia.
Significant variation also exists among fusarioid genera
regarding ascospore characteristics. Most genera consistently
form 1-septate ascospores. These are seen in Cosmosporella,
Cyanonectria, Dialonectria, Fusicolla, Geejayessia, Macroconia,
Microcera, Rectifusarium, Scolecofusarium, Setofusarium, Stylonectria, and Varicosporella. Except for Cyanonectria, in which
the ascospores remain hyaline and smooth; Setofusarium, in
which the ascospores surface is finely striated, and Varicosporella, in which the ascospore surface is ribbed, ascospores of the
above-mentioned genera are often pale yellow to pale brown and
smooth at first, becoming finely spinulose or tuberculate. The
genus Neocosmospora forms (0–)1-septate, yellow-brown ascospores, which are often markedly striate, or more rarely cerebriform (i.e., N. vasinfecta) or spiny (i.e., N. spinulosa). Albonectria
and Luteonectria form characteristic 3-septate, pale yellow-brown,
faintly striate ascospores, while Fusarium produces 1–3-septate,
hyaline to pale yellow-brown and smooth ascospores.
Based on the morphological variation observed in these taxa,
an identification scheme is presented for fusarioid genera of the
Nectriaceae (Fig. 9).
Ex-type strain phylogeny: The analyses included partial rpb1,
rpb2 and tef1 sequences of only the ex-, epi- and neotype strains
as indicated in the nomenclator list of all the names that have been
introduced in Fusarium. The analyses used both ML inferences
and BI of the individual genes and combined datasets, and they
resulted in phylogenies with congruent topologies. Therefore, the
RAxML topology is presented with RAxML-BS, UFboot2-BS, BIPP and gCF support values superimposed (Fig. 10).
The combined alignment comprised 325 strains from 309
species of 14 fusarioid genera including Atractium stilbaster
(CBS 410.67) as the outgroup. A total of 14 fusarioid genera
were resolved of which six (Cosmosporella, Microcera, Nothofusarium, Rectifusarium, Scolecofusarium, and Setofusarium)
were represented by single lineages, mostly due to a lack of
living isolates directly linked to type material available for other
species recognised within these genera at present. The genera
Fusarium (224 strains; 220 accepted species) and Neocosmospora (83 strains; 71 accepted species) both represented
the largest sampling of living isolates directly linked to type
material available. The remaining five genera were represented
by two or more strains and include Bisifusarium (five species and
REDELIMITED
strains), Cyanonectria (two species and strains), Fusicolla (three
species and strains), Geejayessia (two species and strains), and
Luteonectria (two species and strains).
In order to describe novel species found for the genera
treated in this study, additional phylogenies were constructed for
the Fusarium fujikuroi species complex (FFSC), Fusicolla,
Macroconia, Neocosmospora, and Stylonectria.
Fusarium fujikuroi SC phylogeny: The analyses included
partial sequences of five genes (CaM, rpb1, rpb2, tef1 and tub2)
from 52 strains representing 46 species of the FFSC, and two
outgroup taxa (F. curvatum CBS 744.97 and F. inflexum CBS
716.74) (Fig. 11). The analysis of the combined dataset fully
supported five main clades corresponding to the African,
American and Asian clades sensu O'Donnell et al. (2000b), plus
the African B-clade (Sandoval-Denis et al. 2018b, Yilmaz et al.
2021) and a fifth, monotypic clade, which formed the sister
clade to the joint American and African B clades and which is
here termed African C. The latter clade included two strains
showing a clear genealogical and morphological separation from
their closest phylogenetic relatives; both came from an unknown
tree species in South Africa. This clade is here described as the
novel species F. echinatum. Another fully supported novel
monophyletic group was found within the main African clade,
related to but distinct from F. brevicatenulatum and
F. pseudonygamai. This novel group, represented by isolates of
South African origin isolated from Prunus spinosa and from the
South African indigenous species Aloidendron dichotomum, is
here recognised as the novel species F. prieskaense.
Fusicolla phylogeny: The alignment consisted of partial acl1,
ITS, LSU, rpb2, tef1, and tub2 sequences from 20 type or
reference strains, representing 17 species of Fusicolla (Fu.) plus
one outgroup taxon (Macroconia leptosphaeriae CBS 100001).
The analysis confidently resolved 11 ingroup taxa (Fig. 12),
including three novel monotypic lineages, represented by strains
URM 8367, CBS 110189, and CBS 110191, described here as
the new species Fu. quarantenae, Fu. meniscoidea and Fu.
sporellula. Due to a partial lack of sequence data, six species
could not be clearly resolved. Fusicolla cassiae-fistulae and Fu.
siamensis did not receive statistical support in the combined
analysis but are well-resolved using nrDNA sequence data (data
not shown). Fusicolla acetilerea and Fu. bharatavarshae, while
well-delimited in the individual ITS, LSU and rpb2 analyses (data
not shown), were ill-supported in the 6-marker combined analysis. Similarly, Fu. epistroma and Fu. ossicola were not differentiated in either the multimarker analysis, or in the individual
rpb2 analysis. The lack of sequences available to allow comparison with Fu. epistroma, for which only LSU and rpb2 sequences are available, prevented further analysis, as did a
similar problem with Fu. bharatavarshae, for which only nrDNA
and rpb2 are available.
Macroconia phylogeny: The analysis consisted of partial acl1,
CaM, ITS, LSU, rpb1, rpb2, and tub2 sequences from 12 strains
representing seven lineages of Macroconia (Ma.) plus one outgroup taxon (Microcera rubra CBS 638.76) (Fig. 13). Four out of
Fig. 7. Maximum-Likelihood (IQ-TREE-ML) consensus tree inferred from the combined ITS, LSU, rpb1, rpb2 and tef1 multiple sequence alignment of members of Nectriaceae.
Numbers at the branches indicate support values (RAxML-BS / UFboot2-BS / BI-PP / gCF) above 70 % / 0.95 with thickened branches indicating full support (RAxML-BS /
UFboot2-BS / gCF = 100 %; BI-PP = 1). The scale bar indicates expected changes per site. The tree is rooted to Nectria cinnabarina (CBS 125165). Arrows “F1”, “F2” and “F3”
indicate the three alternative Fusarium hypotheses sensu Geiser et al. (2013). Ex-epitype, ex-isotype, ex-neotype and ex-type strains are indicated with ET, IT, NT, and T,
respectively.
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33
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34
ET AL.
FUSARIUM
REDELIMITED
Fig. 8. (Continued).
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35
CROUS
ET AL.
Fig. 8. (Continued).
36
FUSARIUM
REDELIMITED
Fig. 8. Morphological features and phylogenetic affinities of fusarioid genera of Nectriaceae and close relatives. The tree was delineated based on the phylogeny presented in
Fig. 7 and does not indicate phylogenetic distances. Fully supported branches are indicated in bold. The genus Fusarium is indicated in blue. Arrows “F1”, “F2” and “F3”
indicate the three alternative Fusarium hypotheses sensu Geiser et al. (2013). Fusarium. A, B. Ascomata. C–E. Ascospores. F, G. Conidiogenous cells. H–J. Macroconidia.
(B. Adapted from Schroers et al. 2011). Cyanonectria. A, B. Ascomata. C–E. Ascospores. F. Conidiogenous cells. G. Macroconidia. Neocosmospora. A, B. Ascomata. C–E.
Ascospores. F, G. Conidiogenous cells. H, I. Macroconidia. [A. Adapted from Sandoval-Denis & Crous (2018). G. Adapted from Sandoval-Denis et al. (2019)]. Albonectria. A,
B. Ascomata. C–E. Ascospores. F, G. Conidiophores and conidiogenous cells. H. Macroconidia. Setofusarium. A, B. Ascomata. C–E. Ascospores. F–H. Setae formed on
sporodochia. I. Conidiophore. J. Conidia. Geejayessia. A, B. Ascomata. C–E. Ascospores. F, G. Conidiophores and conidiogenous cells. H, I. Macroconidia. [A. Adapted from
Schroers et al. (2011)]. Nothofusarium. A–D. Conidiophores and conidiogenous cells. E. Conidia. Luteonectria. A, B. Ascomata. C–D. Ascospores. F, G. Conidiophores and
conidiogenous cells. H. Conidia. Rectifusarium. A–D. Conidiophores and conidiogenous cells. E, F. Conidia. Bisifusarium. A–D. Conidiophores and conidiogenous cells.
E, F. Conidia. Mariannaea. A, B. Conidiophores. C, D. Conidiogenous cells. E. Conidia. Tumenectria. A, B. Ascomata. C. Ascospores. D, E. Conidiophores and conidiogenous
cells. F. Conidia. [A–C. Adapted from Salgado-Salazar et al. (2016)]. Rugonectria. A, B. Ascomata. C–E. Ascospores. F, G. Conidiophores and conidiogenous cells.
H. Conidia. Thelonectria. A, B. Ascomata. C, D. Ascospores. E, F. Conidiophores and conidiogenous cells. G. Conidia. Corinectria. A, B. Ascomata. C–E. Ascospores.
F, G. Conidiophores and conidiogenous cells. H. Conidia. (H. Picture by C. Gonzalez). Neonectria. A, B. Ascomata. C, D. Ascospores. E, F. Conidiophores and conidiogenous
cells. G, H. Conidia. [A. Adapted from Chaverri et al. (2011)]. Ilyonectria. A, B. Ascomata. C, D. Ascospores. E, F. Conidiophores and conidiogenous cells. G, H. Conidia.
Atractium. A, B. Conidiophores. C, D. Conidiogenous cells. E, F. Conidia. Fusicolla. A, B. Ascomata. C. Ascospores. D, E. Conidiogenous cells. F, G. Conidia. (A–C. Pictures
by C. Lechat). Scolecofusarium. A. Ascomata. B, C. Ascospores. D, E. Conidiophores and conidiogenous cells. F. Conidia. Microcera. A. Ascomata. B. Ascospores.
C, D. Conidiogenous cells. E, F. Conidia. (A, B. Pictures by N. Aplin, Fungi of Great Britain and Ireland). Macroconia. A, B. Ascomata. C–E. Ascospores. F, G. Conidiophores
and conidiogenous cells. H, I. Conidia. (B. Picture by P. Mlcoch). Pseudofusicolla. A, B. Conidiophores and conidiogenous cells. C, D. Conidia. [A–D. Adapted from Triest et al.
(2016)]. Cosmospora. A, B. Ascomata. C, D. Ascospores. E, F. Conidiophores and conidiogenous cells. G. Conidia. Dialonectria. A, B. Ascomata. C–E. Ascospores.
F, G. Conidiophores and conidiogenous cells. H. Conidia. (A. Picture by P. Mlcoch). Cosmosporella. A, B. Ascomata. C–E. Ascospores. F, G. Conidiophores and conidiogenous cells. H, I. Conidia. (A–E. Pictures by P. Mlcoch). Stylonectria. A, B. Ascomata. C–E. Ascospores. F–I. Conidiophores and conidiogenous cells. J. Conidia.
(A–C, E. Pictures by B. Wergen).
the five Macroconia spp. previously known from culture, Ma.
gigas, Ma. leptosphaeriae, Ma. papilionacearum, and Ma.
sphaeriae, resolved as highly to fully-supported lineages. The
poorly resolved position of the ex-type isolate of Ma. cupularis
(HMAS 173240) should be interpreted in light of the fact that only
nrDNA sequences were available for analysis. However, separate ITS and LSU comparisons demonstrated it as distinct (data
not shown). Two distinct and highly supported novel lineages of
South African origin were determined and are described here as
the novel species, Ma. bulbipes and Ma. phlogioides.
Neocosmospora phylogeny: The alignment consisted of
partial acl1, CaM, ITS, LSU, rpb1, rpb2, and tef1 sequences of
107 ex-type and reference strains, including two outgroup taxa
(Geejayessia atrofusca NRRL 22316 and G. cicatricum CBS
125552). The analysis resolved 76 terminal clades, of which 71
correspond to known species of Neocosmospora (Fig. 14).
www.studiesinmycology.org
Seventy of these clades resolved with high support from two or
more independent algorithms (RAxML, IQ-TREE-ML, and BI).
The position of the ex-type of N. crassa (CBS 144386) is poorly
resolved and only partially supported by BI. Similarly, except for
the types of N. ambrosia (CBS 571.94), N. obliquiseptata (NRRL
62611), N. rekana (CMW 52862), and the reference strain of
N. pseudensiformis (CBS 130.78), the position of most members
of the well-delimited Ambrosia-clade of Neocosmospora were
only partially supported by the individual analyses (only BI in
N. kuroshio, N. oligoseptata, and N. tuaranensis, and only IQTREE-ML-BS for N. euwallaceae and N. floridana). All these
lineages were represented by single isolates in these analyses.
Of the five unnamed phylogenetic clades, one corresponded to a
species previously known from phylogenetic analyses (FSSC 41,
Cardoso 2015), for which a Latin binomial is lacking; this species
is here formally described as N. merkxiana. The four additional
37
CROUS
ET AL.
novel lineages discovered here are proposed as the novel
species N. neerlandica, N. nelsonii, N. pseudopisi, and
N. epipeda.
Stylonectria phylogeny: The alignment consisted of partial
acl1, ITS and rpb2 sequences of 11 strains, including the outgroup (Nectria cinnabarina CBS 125165). The analyses (Fig. 15)
identified eight species-level clades, of which six represented
previously known species of the genus: St. applanata, St. carpini,
St. norvegica, St. purtonii, St. qilianshanensis, and St. wegeliniana. One strain, CBS 125491, isolated from an unknown
ascomycetous host, corresponded to a previously known unnamed and fully supported monophyletic lineage, which is
formally described here as St. corniculata. In addition, a fully
supported clade formed by two strains, CBS 147305 from Diaporthe sp. and CBS 147306 from Dothiorella sarmentorum, is
here recognised as the novel species St. hetmanica.
Taxonomy
Albonectria Rossman & Samuels, Stud. Mycol. 42: 105. 1999.
Figs 8, 16.
Type species: Albonectria rigidiuscula (Berk. & Broome) Rossman & Samuels, Stud. Mycol. 42: 105. 1999.
(See F. colorans in List section for synonyms)
Ascomata perithecial, solitary or gregarious, superficial on a
sparse to well-developed, pseudoparenchymatous stroma,
globose to subglobose to ellipsoidal or ovoid to obovoid, not
collapsing or laterally pinched when dry, off-white to pale yellow to
pale ochraceous, not changing in KOH, strongly tuberculate and
thick-walled, with or without a small, pointed papilla, lacking hairs
or appendages. Ascomatal wall of three regions: outer region of
thick-walled, textura angularis to textura globulosa; middle region
of elongate thick-walled cells; inner region with thin-walled, hyaline elongated cells. Asci narrowly to broadly clavate or ellipsoidal, 4–8-spored, ascospores obliquely uniseriate or biseriate.
Ascospores ellipsoidal to long-ellipsoidal or fusoid to long-fusoid,
3- to multiseptate, hyaline to yellow-brown, smooth to striate, not
to slightly constricted at the septum. Conidiophores mononematous (aerial conidiophores) or grouped on sporodochia;
aerial conidiophores unbranched or irregularly branched, bearing
terminal or lateral phialides, often reduced to single phialides;
conidiogenouhs cells monophialidic, cylindrical to subcylindrical,
smooth- and thin-walled, with periclinal thickening inconspicuous
or absent, producing arial micro- and macroconidia. Microconidia
hyaline, thin-walled, 0- or 1-septate, ovoid to obovoid, with or
without a flattened basal papilla, borne in dry chains or small
slimy heads. Macroconidia falcate, multiseptate, thick-walled,
with a blunt to hooked apical cell and well-developed foot-shaped
basal cell or distinctly beaked at both ends. Sporodochia cream to
yellow; sporodochial conidiophores verticillately branched and
densely packed, consisting of short, smooth- and thin-walled
stipes bearing apical whorls of 2–4 monophialides; sporodochial
conidiogenous cells monophialidic, cylindrical to subulate,
smooth- and thin-walled, with reduced or flared collarette.
Sporodochial macroconidia formed in off-white or creamy slimy
masses, falcate, 5–9-septate, thick-walled, gently curved to
straight, with a blunt to hooked apical cell and distinct welldeveloped foot-shaped basal cell. Chlamydospores absent.
[Description adapted from Rossman et al. (1999), Booth (1971)
and Lombard et al. (2015)].
38
Diagnostic features: Off-white to pale yellow to pale ochraceous
perithecia producing narrowly or broadly clavate to ellipsoidal
asci containing (long) ellipsoidal to fusoid, 3- to multiseptate
ascospores; fusarioid asexual morph characterised by monophialides producing distinctly long, robust, slightly curved to
straight multiseptate macroconidia and dry chains or small slimy
heads of ovoid microconidia. Chlamydospores absent.
Atractium Link, Mag. Ges. Naturf. Freunde Berlin 3: 10. 1809
(Fries, Syst. Mycol. 1: XLI. 1821, nom. sanct.). Figs 8, 17.
Type species: Atractium stilbaster Link, Mag. Ges. Naturf.
Freunde Berlin 3: 10. 1809.
Ascomata unknown. Conidiophores aggregated into sporodochia
or synnemata, non-stromatic; synnemata determinate, pale
brown, composed of a stipe of parallel hyphae and a divergent
capitulum of conidiophores giving rise to a slimy conidial mass;
conidiophore branching once or twice monochasial, 2-level
verticillate, monoverticillate or irregularly biverticillate. Conidiogenous cells monophialidic, hyaline, subulate with conspicuous periclinal thickening, producing micro- and macroconidia.
Microconidia hyaline, thin-walled, 0- or 1-septate, ellipsoidal,
allantoid, broadly lunate to reniform, straight or slightly curved,
tapering towards both apices with rounded base. Macroconidia
3–5-septate, falcate, gently curved, with a rounded to blunt
apical cell, and obtuse, non foot-shaped basal cell, forming
yellow to orange masses.
[Description adapted from Gr€afenhan et al. (2011)].
Diagnostic features: Synnematous asexual morph characterised
by fusarioidal macroconidia lacking foot-shaped basal cells.
Bisifusarium L. Lombard et al., Stud. Mycol. 80: 223. 2015. Figs
8, 18.
Type species: Bisifusarium dimerum (Penz.) L. Lombard &
Crous, Stud. Mycol. 80: 225. 2015.
(See F. dimerum in List section for synonyms)
Ascomata unknown. Conidiophores mononematous (aerial conidiophores) or grouped on sporodochia; aerial conidiophores
simple, unbranched or irregularly branched, mostly reduced to
terminal or single lateral conidiogenous cells. Conidiogenous
cells often formed as (i) lateral phialidic pegs arising from superficial or submerged intercalary hyphal cells or, (ii) cylindrical
and slightly tapering towards apex or ampulliform, smooth- and
thin-walled monophialides, rarely polyphialides, with inconspicuous or absent periclinal thickening, solitary or aggregated to
represent a poorly developed pionnotal sporodochial-like structure, producing micro- and macroconidia. Microconidia hyaline,
thin-walled, 0- or 1-septate, ellipsoidal, allantoid, broadly lunate
to reniform, straight or curved, tapering towards both ends.
Macroconidia falcate, (0–)1–2(–3)-septate, thick-walled, curved
to lunate, with a blunt to hooked apical cell and obtuse to poorly
developed, foot-shaped basal cell, typically formed on sporodochia. Sporodochia pale yellow to orange; sporodochial conidiophores verticillately branched and densely packed,
consisting of short, smooth- and thin-walled stipes bearing an
apical whorl of 2–3 monophialides; sporodochial conidiogenous
cells monophialidic, cylindrical to subulate, smooth- and thinwalled, with reduced or flared collarette. Chlamydospores, if
present, globose to subglobose to ellipsoidal, solitary or in
chains, sometimes aggregated in sclerotia.
FUSARIUM
REDELIMITED
Fig. 9. Characters for morphological identification of fusarioid genera in Nectriaceae. The rings show, from inside to outside: conidial morphology; ascospore morphology,
septation and surface; colour reaction of ascomata in 3 % KOH/lactic acid (nr = no reaction); ascomata wall thickness; and general colour, appearance and wall surface of
ascomata.
[Description adapted from Schroers et al. (2009) and Lombard
et al. (2015)].
Diagnostic features: Fusarioid asexual morph characterised by
lateral phialidic pegs arising from superficial or submerged
intercalary hyphal cells or solitarily formed monophialides producing microconidia; distinctly short (< 30 μm long), curved to
lunate, (0–)1–2(–3)-septate macroconidia typically formed on
sporodochia on plant tissue such as carnation leaf pieces.
Corinectria C. Gonzalez & P. Chaverri, Mycol. Progr. 16: 1021.
2017. Fig. 8.
Type species: Corinectria fuckeliana (C. Booth) C. Gonzalez & P.
Chaverri, Mycol. Progr. 16: 1023. 2017.
www.studiesinmycology.org
Basionym: Nectria fuckeliana C. Booth, Mycol. Pap. 73: 56. 1959.
Synonym: Neonectria fuckeliana (C. Booth) Castl. & Rossman,
Canad. J. Bot. 84: 1428. 2006.
Ascomata perithecial, gregarious, seated on an erumpent
stroma, superficial, globose to subglobose, orange, red to dark
red darkening around ostiolar region, turning black in KOH,
pigment dissolving in lactic acid, not collapsing when dry, slightly
papillate to papillate, smooth-walled, lacking hairs or appendages. Ascomatal wall of 2–3 regions: outer region of thick-walled, pigmented cells forming a textura epidermoidea; middle and
inner regions of globose to elongate, hyaline, thin-walled cells,
becoming thinner toward the centrum. Asci cylindrical, 8-spored,
with an apical ring, uniseriate. Ascospores ellipsoidal to fusoid, 139
CROUS
40
ET AL.
FUSARIUM
REDELIMITED
Fig. 10. (Continued).
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41
CROUS
42
ET AL.
FUSARIUM
REDELIMITED
Fig. 10. Maximum-Likelihood (IQ-TREE-ML) consensus tree inferred from the combined rpb1, rpb2 and tef1 sequence alignment of the living type strains as indicated in the
nomenclator list. Numbers at the branches indicate support values (RAxML-BS / UFboot2-BS / BI-PP) above 70 % / 0.95 with thickened branches indicating full support
(RAxML-BS / UFboot2-BS = 100 %; BI-PP = 1). The scale bar indicates expected changes per site. The tree is rooted to Atractium stilbaster (CBS 410.67). Names indicated in
bold are in current use. Subdivision of the Fusarium clade (blue block) represent the recognised species complexes.
septate, hyaline, smooth. Conidiophores mononematous, hyaline, septate, unbranched or sparsely branched, terminating in
1–2 phialides or reduced to lateral phialides. Conidiogenous
cells monophialidic, cylindrical, tapering towards the apex, with
inconspicuous periclinal thickening and collarettes. Sporodochia
not formed. Microconidia ellipsoidal to obovoid, hyaline, aseptate, sometimes forming false heads on phialides. Macroconidia
cylindrical, mostly straight, (3–)5–7-septate, with rounded ends.
Chlamydospores unknown.
[Description adapted from Gonzalez & Chaverri (2017)].
Diagnostic features: Orange to dark red, smooth-walled perithecia with papilla producing cylindrical asci bearing ellipsoidal to
fusoid, 1-septate ascospores and cylindrocarpon-like asexual
morph characterised by (3–)5–7-septate macroconidia.
Cosmospora Rabenh., Hedwigia 2: 59. 1862. Fig. 8.
Synonyms: Crysogluten Briosi & Farneti, Atti Ist. Bot. Univ. Lab.
Crittog. Pavia 8: 117. 1904.
Botryocrea Petr., Sydowia 3: 140. 1949.
Type species: Cosmospora coccinea Rabenh., Hedwigia 2: 59.
1862 [non Nectria coccinea (Pers.) Fr. 1849].
Synonyms: Nectria cosmariospora Ces. & De Not., Comment.
Soc. Crittog. Ital. 1: 195. 1863.
Dialonectria cosmariospora (Ces. & De Not.) Cooke, Grevillea
12: 110. 1884.
Cucurbitaria cosmariospora (Ces. & De Not.) Kuntze, Revis.
Gen. Pl. 3: 461. 1898.
www.studiesinmycology.org
Dialonectria cosmariospora (Ces. & De Not.) Z. Moravec, Ceska
Mykol. 8: 92. 1954, an isonym, Art. 6.3, Note 2.
Verticillium olivaceum W. Gams, Cephalosporium-artige Schimmelpilze: 129. 1971.
Ascomata perithecial, solitary or gregarious, with inconspicuous
or absent stroma, obpyriform with an acute or papillate apex,
orange red or bright red, turning dark red in KOH, smooth walled.
Asci narrowly clavate to cylindrical, with an apical ring, 8spored. Ascospores initially hyaline, becoming yellow brown to
reddish brown, 1-septate, becoming tuberculate when mature.
Conidiophores acremonium-like, consisting of lateral phialides on
somatic hyphae, or with one or two levels of monochasial
branching, or verticillate, hyaline. Conidiogenous cells monophialidic, cylindrical to subulate to subclavate, hyaline. Microconidia ellipsoidal, oblong or clavate or slightly allantoid,
aseptate, hyaline, forming slimy heads. Macroconidia absent or
rare, subcylindrical, curved, slightly narrowing towards each end,
apical cell often slightly hooked with a more or less pointed apex,
basal cell obtuse to poorly developed, foot-shaped, 3–5-septate,
hyaline.
[Description adapted from Rossman et al. (1999) and Gr€afenhan
et al. (2011)].
Diagnostic features: Orange-red to bright red perithecia with an
acute or papillate apex producing cylindrical to narrowly clavate
asci, yellow brown to reddish brown, 1-septate, tuberculate ascospores and acremonium-like asexual morph.
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CROUS
ET AL.
Cosmosporella S.K. Huang et al., Cryptog. Mycol. 39: 179.
2018. Figs 8, 19.
Type species: Cosmosporella olivacea S.K. Huang et al., Cryptog. Mycol. 39: 181. 2018.
Ascomata perithecial, solitary to gregarious, superficial, on
immersed to erumpent stroma, ovoid, globose to obpyriform,
collapsing laterally when dry, orange red, red to pale yellow, not
reacting in KOH, with a central ostiole, with hyaline periphyses.
Ascomatal wall membranous, composed of orange to hyaline
cells of textura angularis, with septate paraphyses. Asci
cylindrical to slightly clavate, apically rounded, with evanescent
wall, pedicel combined with paraphyses, 8-spored, unitunicate.
Ascospores hyaline to pale brown, ellipsoidal to ovoid, 0- or
1-septate. Conidiophores acremonium-like, mononematous,
hyaline, septate, consisting of lateral phialides on somatic
hyphae, or with one or two levels of monochasial branching, or
irregularly branched. Conidiogenous cells monophialidic, cylindrical, producing micro- and macroconidia. Microconidia ellipsoidal to obovoid, 0- or 1-septate, hyaline, forming a false head
on phialides. Macroconidia falcate, almost straight to curved,
1–3-septate, with a blunt to hooked apical cell and poorly to welldeveloped foot-shaped basal cell. Chlamydospores unknown.
[Description adapted from Huang et al. (2018)].
Diagnostic features: Pale yellow to orange-red perithecia lacking
a papilla producing cylindrical to narrowly clavate asci, pale
brown, 1-septate ascospores and fusarioid asexual morph
characterised by overly long, 1–3-septate macroconidia.
Cosmosporella cavisperma (Corda) Sand.-Den., L. Lombard &
Crous, comb. nov. MycoBank MB 838659.
Basionym: Fusarium cavispermum Corda, Icon. Fung. 1: 3. 1837.
Synonyms: Fusarium aquaeductuum var. cavispermum (Corda)
Raillo, Fungi of the Genus Fusarium: 280. 1950.
Fusarium oxydendri Ellis & Everh., Bull. Torrey Bot. Club 24: 477.
1897.
Fusarium cavispermum var. minus Wollenw., Fusaria Autogr.
Delin. 3: 848. 1930.
Lectotypus: Czech Republic, near Carlsstein, on pine resin,
A.K.J. Corda, Icon. Fung. 1: tab. I, fig. 58 (MBT 10001322 hic
designatus). Epitype of Fusarium cavispermum (CBS 172.31,
MBT 10000645 hic designatus, a metabolically inactive culture).
Norway, from Pinus sylvestris, 1929, H.W. Wollenweber, culture
ex-epitype CBS 172.31 = NRRL 13996.
Notes: The genus Cosmosporella was erected by Huang et al.
(2018) to accommodate Cm. olivacea and the superfluous taxon
Cm. obscura, shown to cluster within a subset of taxa pertaining to
Cosmospora s. lat. (Rossman et al. 1999), former members of the
Nectria episphaeria group sensu Booth (1959) and Nectria subgenus Dialonectria (Samuels et al. 1991) characterised by cosmospora-like sexual morphs and fusarioid asexual morphs. More
recently, this monophyletic clade had been ascribed to the Fusarium
cavispermum species complex (O'Donnell et al. 2013) and, separated from any of the polyphyletic taxa formerly classified in
Fusarium section Eupionnotes (O'Donnell 1993, Schroers et al.
2009, Gr€afenhan et al. 2011). “Fusarium” melanochlorum, its purposed sexual morph “Nectria” flavoviridis (Gerlach & Nirenberg
1982), and “Fusarium” cavispermum have also been resolved as
members of this clade (Gr€afenhan et al. 2011, O'Donnell et al. 2013,
Huang et al. 2018, and Fig. 7 in this paper). Here, the new combination Cm. cavisperma is proposed, lectotypified, and an epitype
44
is designated to stabilise the application of the name based on
material studied by Wollenweber [number 849 in Wollenweber
(1916–1935) and Gerlach & Nirenberg (1982)]. The suggested
conspecificity of “F”. melanochlorum and “N”. flavoviridis, however,
is questioned given the large phylogenetic distance between the
currently available strains. Fresh isolations and a thorough phylogenetic revision of the entire group including additional Cosmospora
s. lat. taxa having fusarioid asexual morphs are necessary.
Cyanonectria Samuels & Chaverri, Mycol. Progr. 8: 56. 2009.
Figs 8, 20.
Type species: Cyanonectria cyanostoma (Sacc. & Flageolet)
Samuels & Chaverri, Mycol. Progr. 8: 56. 2009.
Basionym: Nectria cyanostoma Sacc. & Flageolet, Rendiconti
Congr. Bot. Palermo 1902: 53. 1902.
Synonym: Fusarium cyanostomum (Sacc. & Flageolet) O'Donnell & Geiser, Phytopathology 103: 404. 2013.
Ascomata perithecial, gregarious or caespitose, with a reduced
or well-developed prosenchymatous stroma, smooth- and thinwalled, ampulliform to obpyriform to pyriform, apex dark bluish
purple to bluish black and body less intensely dark bluish or red
or reddish brown, turning darker in KOH, pigment dissolving in
lactic acid to become red to yellow, non-papillate, lacking hairs or
appendages. Ascomatal wall consisting of a single region,
comprising several layers of morphologically similar cells. Asci
cylindrical to narrowly clavate, with rounded to flattened thickened apex, with or without refractive ring, 8-spored, ascospores
overlapping uniseriate or biseriate above and uniseriate below.
Ascospores ellipsoidal, 1-septate, not or slightly constricted at
septum, pale yellow-brown, smooth-walled or finely verrucose.
Conidiophores mononematous (aerial conidiophores) or grouped
on sporodochia; aerial conidiophores unbranched or rarely
branched, bearing terminal or lateral phialides, often reduced to
single phialides. Conidiogenous cells monophialidic, cylindrical
to subcylindrical, smooth- and thin-walled, with periclinal thickening inconspicuous or absent. Sporodochia white to bluish;
sporodochial conidiophores verticillately branched and densely
packed, consisting of short, smooth- and thin-walled stipes
bearing apical whorls of 2–3 monophialides; sporodochial
conidiogenous cells monophialidic, cylindrical to subulate,
smooth- and thin-walled, with reduced or flared collarette.
Macroconidia formed in off-white or creamy or greyish blue slimy
masses, falcate, straight to gently curved, with inequilateral
fusoid or hooked apical cell and well-developed, foot-shaped
basal cell. Microconidia unknown. Chlamydospores absent or
rarely formed from cells of the macroconidia, subglobose.
[Description adapted from Samuels et al. (2009) and Schroers
et al. (2011)].
Diagnostic features: Bicoloured or dark bluish purple to bluish
black perithecia producing cylindrical to narrowly clavate asci
containing ellipsoidal, 1-septate ascospores and fusarioid
asexual morph characterised by monophialides producing long,
narrow, almost straight macroconidia, lacking microconidia and
hyphal-borne chlamydospores.
Dialonectria (Sacc.) Cooke, Grevillea 12: 77, 109. 1884. Figs 8, 21.
Basionym: Nectria subgen. Dialonectria Sacc., Syll. Fung. 2:
490. 1883.
Type species: Dialonectria episphaeria (Tode) Cooke (as
“episphaerica”), Grevillea 12: 82. 1884.
FUSARIUM
REDELIMITED
Fig. 11. Maximum-Likelihood (IQ-TREE-ML) consensus tree inferred from the combined CaM, rpb1, rpb2, tef1, and tub2 sequence alignment of members of the Fusarium
fujikuroi species complex. Numbers at the branches indicate support values (RAxML-BS / UFboot2-BS / BI-PP) above 70 % / 0.95 with thickened branches indicating full
support (RAxML-BS / UFboot2-BS = 100 %; BI-PP = 1). Novel taxa are indicated in bold. The scale bar indicates expected changes per site. The tree is rooted to Fusarium
curvatum CBS 744.97 and Fusarium inflexum CBS 716.74. Ex-epitype, ex-neotype and ex-type strains are indicated with ET, NT, and T, respectively.
Basionym: Sphaeria episphaeria Tode, Fung. mecklenb. sel. 2:
21. 1791.
Ascomata perithecial, solitary or gregarious, with inconspicuous
or absent stroma, obpyriform with an acute or round papilla,
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orange red to carmine red, turning dark red in KOH, smoothwalled. Asci narrowly clavate to cylindrical, with an apical ring, 8spored, uniseriate. Ascospores initially hyaline, becoming pale
brown, 1-septate, becoming tuberculate when mature. Conidiophores mononematous, initially as lateral phialides on
45
CROUS
ET AL.
Fig. 12. Maximum-Likelihood (IQ-TREE-ML) consensus tree inferred from the combined acl1, ITS, LSU, rpb2, tef1, and tub2 sequence alignment of members of the genus
Fusicolla. Numbers at the branches indicate support values (RAxML-BS / UFboot2-BS / BI-PP) above 70 % / 0.95 with thickened branches indicating full support (RAxML-BS /
UFboot2-BS = 100 %; BI-PP = 1). Novel taxa are indicated in bold. The scale bar indicates expected changes per site. The tree is rooted to Macroconia leptosphaeriae CBS
100001. Ex-epitype and ex-type strains are indicated with ET, and T, respectively.
somatic hyphae, sometimes verticillate, hyaline. Conidiogenous
cells monophialidic, subulate to subclavate, hyaline. Microconidia
ellipsoidal to clavate, aseptate, hyaline, abundant. Macroconidia
if present subcylindrical, moderately curved, slightly narrowing
towards each end, apical cell often slightly hooked with a more or
less pointed tip, basal cell obtuse to poorly developed, footshaped, predominantly 3–5-septate, hyaline. Chlamydospores
unknown.
[Description adapted from Rossman et al. (1999) and Gr€afenhan
et al. (2011)].
Diagnostic features: Orange-red to carmine-red perithecia with
an acute or round papilla producing cylindrical to narrowly
clavate asci, pale brown, 1-septate, tuberculate ascospores and
asexual morph that rarely produces macroconidia.
Fusarium Link, Mag. Ges. Naturf. Freunde Berlin 3: 10. 1809.
Figs 8, 22.
Synonyms: Fusisporium Link, Mag. Ges. Naturf. Freunde Berlin
3: 19. 1809.
Selenosporium Corda, Icon. Fung. 1: 7. 1837.
Gibberella Sacc., Michelia 1: 43. 1877.
Lisea Sacc., Michelia 1: 43. 1877.
Sporotrichella P. Karst., Meddel. Soc. Fauna Fl. Fenn. 14: 96.
1887.
Gibberella subgen. Lisiella Cooke & Massee, Grevillea 16: 5.
1887.
46
Lisiella (Cooke & Massee) Sacc., Syll. Fung. 9: 945. 1891.
Septorella Allesch., Hedwigia 36: 241. 1897.
Ustilaginoidella Essed, Ann. Bot. 25: 351. 1911.
Rachisia Linder, Deutsche Essigind. 17: 467. 1913.
Stagonostroma Died., Krypt.-Fl. Mark Brandenb. 9: 561. 1914.
Fusidomus Grove, J. Bot. 67: 201. 1929.
Pseudofusarium Matsush., Microfungi Solomon Isl. Papua-New
Guinea: 46. 1971.
Type species: Fusarium sambucinum Fuckel, Fungi Rhen. Exs.,
Fasc. 3, no. 211. 1863, nom. cons.
(See List section for synonyms)
Ascomata perithecial, mostly gregarious, non-stromatic or on a thin
stroma erumpent through the epidermis, superficial, subglobose to
globose to broadly pyriform, not collapsing or laterally pinched
when dry, bluish purple to black, turning dark purple in KOH,
pigment dissolving in lactic acid, non-papillate, slightly rugose to
tuberculate, lacking hairs or appendages. Ascomatal wall of two
regions: outer region of thick-walled, pigmented cells forming a
textura angularis or textura globulosa; inner region of elongate,
hyaline, thin-walled cells, becoming thinner towards the centrum.
Asci clavate, apex simple, 8-spored often with an apical ring,
biseriate to pluriseriate. Ascospores ellipsoidal to cylindrical, 1–3septate, not or slightly constricted at the septa, pale tan, smoothwalled. Conidiophores mononematous (aerial conidiophores) or
grouped on sporodochia; aerial conidiophores, if consistenly
FUSARIUM
REDELIMITED
Fig. 13. Maximum-Likelihood (ML) consensus tree inferred from the combined acl1, CaM, ITS, LSU, rpb1, rpb2, and tub2 sequence alignment of members of the genus
Macroconia. Numbers at the branches indicate support values (RAxML-BS / UFboot2-BS / BI-PP / gCF) above 70 % / 0.95 with thickened branches indicating full support
(RAxML-BS / UFboot2-BS = 100 %; BI-PP = 1). Novel taxa are indicated in bold. The scale bar indicates expected changes per site. The tree is rooted to Microcera rubra CBS
638.76. Ex-type and ex-isotype strains are indicated with T, and IT, respectively.
formed, unbranched, sympodial or irregularly branched, bearing
terminal or lateral phialides, often reduced to single phialides.
Conidiogenous cells mono- or polyphialidic, subulate to subcylindrical, smooth- and thin-walled, sometimes proliferating percurrently, with periclinal thickening inconspicuous or absent. Aerial
conidia hyaline, smooth- and thin-walled, of three types: microconidia ellipsoidal to fusoid to ovoid to obovoid to reniform to
allantoid to clavate to napiform to pyriform to limoniform, 0–5septate, borne in false heads or chains on the phialides; mesoconidia (occurring in some species or species complexes) falcate,
slender with no significant curvature to curved with parallel walls,
1–5-septate, tapering towards both ends, with a pointed to blunt
apical cell and obtuse to flattened basal cell; macroconidia, typically formed on sporodochia, falcate, slightly to strongly curved
dorsiventrally, 1-septate to multiseptate, with a curved, long and
tapering, pointed, blunt, hooked or elongated apical cell and
obtuse, poorly developed, well-developed, to elongate, foot-shaped basal cell. Sporodochia cream to pale tan to orange to saffron
to blue; sporodochial conidiophores verticillately branched and
densely packed, consisting of short, smooth- and thin-walled stipes
bearing an apical whorl of 2–4 monophialides; sporodochial conidiogenous cells subulate to subcylindrical, smooth- and thin-walled, with reduced or flared collarette; sporodochial (macro)conidia
falcate, smooth- and thin-walled, distinctly curved to curved with
parallel walls to unequally curved, tapering towards both ends, with
pointed, blunt, papillate, hooked, or elongate apical cell and obtuse,
poorly developed, well-developed, to elongate, foot-shaped basal
cell. Chlamydospores globose to subglobose to ovoid to obovoid,
hyaline to subhyaline, smooth-walled to slightly verrucose, terminal
or intercalary, solitary or in pairs or forming chains or aggregating to
form microsclerotia.
[Description adapted from Rossman et al. (1999) and Lombard
et al. (2015)].
www.studiesinmycology.org
Diagnostic features: Dark blue to black perithecia producing
clavate asci bearing ellipsoidal to cylindrical 1- to multiseptate
ascospores and asexual morphs producing micro- and macroconidia, and sometimes mesoconidia on aerial conidiophores
with mono- and/or polyphialides or only macroconidia in sporodochia. Chlamydospores form in hyphae, rarely in
macroconidia.
Fusarium echinatum Sand.-Den. & G.J. Marais, sp. nov.
MycoBank MB 838660. Fig. 23.
Etymology: From the Latin echinatus, prickly, referring to the
spiny appearance of its multiloculate, often swollen and rather
deformed conidiogenous cells.
Typus: South Africa, unidentified tree species, 2010, A. Lubben
(holotype CBS H-24658, culture ex-type CBS 146497 = CPC
30815 = CAMS 000733).
Conidiophores on aerial mycelium 10–120 μm tall, unbranched
or irregularly laterally branched, bearing lateral and terminal
single phialides; aerial conidiogenous cells polyphialidic, subulate, subcylindrical or more commonly irregularly shaped, curved,
swollen and distorted due to abundant conidiogenous loci,
smooth- and thin-walled, 6.5–36.5 × 2–3.5 μm, polyphialides
with 2–3 or more commonly 10–18 conidiogenous openings,
with inconspicuous to absent periclinal thickening and collarettes.
Aerial microconidia forming small false heads on tips of phialides, hyaline, smooth, and thin-walled, commonly ovoid to
ellipsoidal, 0- or 1-septate, 4–11(–19) × 2–3.5(–4.5) μm (av.
7.5 × 2.7 μm), and more rarely napiform, smooth and thin-walled,
0-septate, (5–)5.5–7 × (3.5–)4.5–5.5 μm (av. 6.4 × 4.5 μm).
Sporodochial conidiophores 28.5–60(–68.5) μm tall, irregularly
branched, bearing terminal solitary monophialides or whorls
of up to three monophialides. Sporodochial conidiogenous
47
CROUS
48
ET AL.
FUSARIUM
REDELIMITED
Fig. 14. Maximum-Likelihood (IQ-TREE-ML) consensus tree inferred from the combined acl1, CaM, ITS, LSU, rpb1, rpb2, and tef1 sequence alignment of members of the
genus Neocosmospora. Numbers at the branches indicate support values (RAxML-BS / UFboot2-BS / I-PP) above 70 % / 0.95 with thickened branches indicating full support
(RAxML-BS / UFboot2-BS = 100 %; BI-PP = 1). Novel taxa are indicated in bold. The scale bar indicates expected changes per site. The tree is rooted to Geejayessia
atrofusca NRRL 22316 and G. cicatricum CBS 125552. Ex-epitype, ex-neotype, ex-paratype and ex-type strains are indicated with ET, NT, PT, and T, respectively.
www.studiesinmycology.org
49
CROUS
ET AL.
cells monophialidic, subulate to subcylindrical, smooth- and thinwalled, (8.5–)11.5–16(–17.5) × (1.5–)2.5–3.5 μm. Sporodochial macroconidia moderately curved to wedge-shaped,
slender, tapering towards the basal part, apical cell of equal size
than the adjacent cell, blunt to slightly hooked; basal cell poorly
to well-developed, foot-shaped, (1–)2–3(–4)-septate, hyaline,
thin- and smooth-walled: 1-septate conidia: (16.5–)
19.5–32.5(–36) × 2.5–3.5 μm (av. 26.1 × 2.9 μm); 2-septate
conidia: (19.5–)25–36(–37.5) × 2.5–3.5 μm (av. 30.5 × 3.1 μm);
3-septate
conidia:
(20.5–)28.5–36(–40)
×
(2.5–)
3–3.5(–4.5) μm (av. 32.5 × 3.2 μm); 4-septate conidia: (27–)
30.5–39(–40.5) × 3–4 μm (av. 35.4 × 3.6 μm); overall: (19.5–)
28.6–36.5(–40.5) × (2.5–)3–3.5(–4.5) μm (av. 32.4 × 3.2 μm).
Chlamydospores not observed.
Culture characteristics: Colonies on PDA reaching
31–63 mm diam at 25 °C after 7 d. Surface white, pale luteus to
sulphur yellow, flat, woolly to cottony with radial patches of white
aerial mycelium, margin regular and filiform. Reverse white,
sulphur yellow to pure yellow at centre. On OA pale luteus to
sulphur yellow, flat, membranous at first, quickly becoming
velvety to dusty, margin regular. Reverse sulphur yellow.
Additional material examined: South Africa, unidentified tree species, 2010, A.
Lubben, culture CBS 146496 = CPC 30814 = CAMS 000730.
Notes: Yilmaz et al. (2021) recently revised the FFSC, including
formal descriptions for several species, while fixing the typification of relevant plant pathogenic and toxigenic species. Species
in this complex have been traditionally organised according to
their biogeographic patterns, which roughly match their phylogenetic distribution. Apart from the monophyletic American and
Asian clades, the complex contains a non-monophyletic African
clade, which is currently known to cluster into two distinct clades:
the speciose core African clade and the African “B” clade
encompassing F. dlaminii and F. fredkrugeri (O'Donnell et al.
2000b, Herron et al. 2015, Sandoval-Denis et al. 2018b, Yilmaz
et al. 2021). The novel South African species F. echinatum,
however, formed a fully-supported single lineage that did not
belong to any of the currently known biogeographically defined
clades (Fig. 11). The most noticeable morphological feature that
distinguishes F. echinatum is the presence of well-developed
polyphialides bearing multiple conidiogenous openings that are
often concentrated in large numbers and that cause a deformation of the apical region. Somewhat similar, conspicuous polyphialides can be found in Fusarium chlamydosporum and
F. concolor (syn. F. polyphialidicum); however, these species are
not directly related, in that they belong to two different species
complexes, the F. chlamydosporum and F. concolor species
complexes, respectively (Fig. 10). The polyphialides formed by
these two species do not show as many conidiogenous loci as do
those of F. echinatum.
Fusarium prieskaense G.J. Marais & Sand.-Den., sp. nov.
MycoBank MB 838661. Fig. 24.
Etymology: Referring to Prieska, a town in Northern Cape
Province, South Africa, where the type was collected.
Typus: South Africa, Northern Cape Province, Prieska,
on Prunus spinosa, 2010, F.J.J. van der Walt & G.J. Marais
(holotype CBS H-24660, culture ex-type CBS 146498 = CPC
30826 = CAMS 001176).
50
Conidiophores on aerial mycelium 12.5–43.5 μm tall, unbranched or rarely irregularly or sympodially branched and
proliferating, bearing terminal single phialides or whorls of 2–3
phialides, commonly reduced to solitary conidiogenous cells
borne laterally on hyphae; aerial conidiogenous cells mono- and
polyphialides, subulate to subcylindrical, smooth- and thin-walled, 8–29.5 × 2–5 μm, polyphialides often with 2–3 conidiogenous openings, periclinal thickening and collarettes often
inconspicuous or absent. Aerial microconidia forming small
false heads and short chains on phialide tips, hyaline, obovoid to
short clavate, smooth and thin-walled, 0-septate, (4.5–)
6–9(–13) × 2–3(–4) μm (av. 7.4 × 2.6 ìm). Sporodochial conidiophores 24.5–39(–45) μm tall, irregularly branched, bearing
terminal solitary or whorls of 2–3 phialides. Sporodochial conidiogenous
cells
monophialidic,
doliiform,
subulate
to subcylindrical, smooth- and thin-walled, (8.5–)
10–14(–15) × 2–4.5 μm. Sporodochial conidia straight to
moderately curved and slender, tapering towards the basal part,
apical cell more or less equally sized as the adjacent cell, blunt to
slightly hooked; basal cell well-developed, foot-shaped, rarely
papillate, (1–)3–4-septate, hyaline, thin- and smooth-walled: 1septate conidia: 23.5 × 3.5 μm; 3-septate conidia: (33.5–)
44.5–58(–68.5) × (3–)3.5–4.5(–5) μm (av. 51.1 × 4 μm); 4septate conidia: (52.5–)55.5–67.5(–71) × 3.5–4.5 μm (av.
61.3 × 4.1 μm); overall: (23–)44–59(–71) × 3–4(–5) μm (av.
51.3 × 4 μm). Chlamydospores not observed.
Culture characteristics: Colonies on PDA reaching
42–68 mm diam at 25 °C after 7 d. Surface pale luteous, luteous
to pale sienna, flat, velvety to felty, sometimes with small white
patches of aerial mycelium, margin filiform and regular. Reverse
sulphur yellow to amber, pale orange at centre. On OA, sienna to
pale umber, flat, membranous to dusty, margin entire and regular; reverse sienna to pale umber.
Additional material examined: South Africa, Northern Cape Province, Prieska,
on Prunus spinosa, 2010, F.J.J. van der Walt & G.J. Marais, culture CBS
146499 = CPC 30827 = CAMS 001177; on Aloidendron dichotomum, 2010, F.J.J.
van der Walt & G.J. Marais, culture CPC 30825 = CAMS 001175.
Notes: Fusarium prieskaense is nested within the core African
clade of the FFSC (Fig. 11). Similar to most members of this
clade, this species is characterised by forming mostly monophialides and occasional to frequent polyphialides, sometimes
proliferating and producing aerial conidia typically organised in a
combination of false heads and short to long chains. Fusarium
prieskaense is morphologically and phylogenetically related to
Fusarium brevicatenulatum and F. pseudonygamai from which it
can be differentiated by its pale luteus to yellow colony pigmentation on PDA, versus the orange to dark blue or violet pigments
produced by the two latter species (Leslie & Summerell 2006).
Additionally, sporodochia and macroconidia are commonly and
abundantly produced by F. prieskaense, whereas these structures
are relatively rare in the two aforementioned species. Moreover,
the obovoid to short clavate microconidia of F. prieskaense also
distinguishes this species from F. brevicatenulatum, which is
characterised by long oval to obovoid microconidia (Nirenberg
et al. 1998).
Fusicolla Bonord., Handb. Allg. Mykol.: 150. 1851. Figs 8, 25.
Type species: Fusicolla betae (Desm.) Bonord., Handb. Allg.
Mykol.: 150. 1851.
FUSARIUM
REDELIMITED
Fig. 15. Maximum-Likelihood (IQ-TREE-ML) consensus tree inferred from the combined acl1, ITS, and rpb2 sequence alignment of members of the genus Stylonectria.
Numbers at the branches indicate support values (RAxML-BS / UFboot2-BS / BI-PP) above 70 % / 0.95 with thickened branches indicating full support (RAxML-BS / UFboot2BS = 100 %; BI-PP = 1). Novel taxa are indicated in bold. The scale bar indicates expected changes per site. The tree is rooted to Macroconia leptosphaeriae CBS 100001. Exepitype and ex-type strains are indicated with ET and T, respectively.
Fig. 16. Albonectria spp. A–C. Ascomata on natural substrate. D. Surface view of perithecial wall in 2 % KOH. E–K. Asci and ascospores (J, K. Surface view). L–P. Conidiophores
and conidiogenous cells. Q, R. Microconidia. S. Macroconidia. A, C–F, H–J. Albonectria rigidiuscula (BPI 553050). B, G, K. Albonectria rigidiuscula (BPI 1104484). L, M, P–S.
Albonectria rigidiuscula (CBS 122570). N, O. Albonectria rigidiuscula (CBS 133.25). Scale bars: A–C = 100 μm; all others = 10 μm (G applies to H–K).
www.studiesinmycology.org
51
CROUS
ET AL.
Fig. 17. Atractium spp. A, B. Synnemata. C–G. Conidiophores and conidiogenous cells. H. Microconidia. I. Macroconidia. A–D, H. Atractium stilbaster (CBS 410.67). E–G, I.
Atractium crassum (CBS 180.31). Scale bars: A = 100 μm; all others = 10 μm.
Fig. 18. Bisifusarium spp. A–D, F–J. Conidiophores and conidiogenous cells. K, L. Microconidia. E, M. Macroconidia. A–E. Bisifusarium dimerum (CBS 108944). F–M.
Bisifusarium delphinoides (CBS 120718). Scale bars: H, J = 5 μm; all others = 10 μm.
(See F. betae in List section for synonyms)
Ascomata perithecial, solitary, rarely gregarious, with erumpent
stroma, fully or partially immersed in a slimy, pale orange sheet of
hyphae over the substrate, globose to pyriform with a short acute or
disk-like papilla, not collapsing when dry, yellow, pale buff to orange,
not changing colour in KOH, smooth-walled, rarely tuberculate,
generally lacking hairs or with short, thick-walled hyphae-like
structures. Asci cylindrical to narrowly clavate, with an apical ring, 8spored. Ascospores broadly ellipsoidal, 1-septate, slightly constricted at the septum, verrucose, hyaline to pale brown. Conidiophores initially as lateral phialides on somatic hyphae,
sometimes monochasial, verticillate or penicillate, hyaline. Conidiogenous cells monophialidic, cylindrical to subulate, hyaline.
Microconidia absent or sparse, ellipsoidal to allantoid, aseptate,
hyaline. Macroconidia falcate, straight to curved, narrowing towards
the ends, apical cell often hooked with a pointed tip, basal cell poorly
52
developed, foot-shaped, 1–3-septate or 3–5-septate or up to 10septate, hyaline. Chlamydospores absent to abundant, globose,
single, in pairs or chains, sometimes formed in macroconidia.
[Description adapted from Gerlach & Nirenberg (1982) and
Gr€afenhan et al. (2011)].
Diagnostic features: Yellow to orange, mostly smooth-walled
perithecia with a short acute or disk-like papilla producing cylindrical to narrowly clavate asci bearing broadly ellipsoidal, 1septate, verrucose ascospores and fusarioid asexual conidia.
Fusicolla quarantenae J.D.P. Bezerra, Sand.-Den., Crous &
Souza-Motta, sp. nov. MycoBank MB 838692. Fig. 26.
Etymology: The epithet refers to the quarantine period during the
2020–2021 coronavirus pandemic, which killed thousands of people on five continents, and during which this species was described.
FUSARIUM
REDELIMITED
Fig. 19. Cosmosporella spp. A–D. Ascomata on natural substrate. E. Surface view of perithecial wall. F. Asci. G–J. Ascospores. K, L. Conidiophores. M. Microconidia. N, O.
Macroconidia. A–J. “Cosmospora” flavoviridis (photos P. Mlcoch). K–N. “Cosmospora” flavoviridis (CBS 124353). O. Cosmosporella cavisperma (CBS 172.31). Scale bars:
A–D = 300 μm; E = 50 μm; G–J = 5 μm; all others = 10 μm.
Fig. 20. Cyanonectria spp. A–C. Ascomata on natural substrate. D. Longitudinal section through perithecium in Shears. E. Surface view of perithecial wall in 2 % KOH. F, G.
Asci. H–K. Ascospores (K. Surface view). L–O. Conidiogenous cells. P. Macroconidia. A–C, E–J. Cyanonectria buxi (CBS H-20380). D, K. Cyanonectria buxi (CBS H-20379).
L–N. Cyanonectria buxi (CBS 130.97). O, P. Cyanonectria buxi (CBS 125551). [A, D, L. adapted from Schroers et al. (2011).] Scale bars: A–D = 100 μm; H–K = 5 μm (H
applies to I and J); all others = 10 μm.
Typus: Brazil, Pernambuco state, Itaíba municipality, Curral
Velho Farm, 9º08.895 S 37º12.069 W, on cladodes of Melocactus
zehntneri, Sep. 2013, J.D.P. Bezerra (holotype URM 94407,
culture ex-type URM 8367 = CBS 141541).
Conidiophores arising laterally from somatic hyphae,
simple, straight, hyaline, thin- and smooth-walled, septate,
25–116 × 1.5–2.5 μm, or reduced to solitary conidiogenous
cells. Conidiogenous cells monophialidic, arising laterally from
hyphae, cylindrical to subulate, straight, hyaline, thin- and
smooth-walled, 1–22 × 0.5–2 μm, or as short lateral pegs.
Macroconidia falcate, more or less straight, slightly narrowing
towards the ends, apical cell often hooked with a more or
less pointed tip, basal cell poorly developed, foot-shaped,
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hyaline, thin- and smooth-walled, 3-septate, (21–)
27–35(–38.5) × 2–2.5(–3) μm (av. 29.5 × 2.5 μm, n = 30).
Microconidia, chlamydospores and sexual morph not observed.
Culture characteristics: Colonies on PDA reaching 15 mm diam
after at 25 °C after 7 d. Surface yellow to apricot in centre, peach
to brick in middle, and salmon at margin, flat, aerial mycelium
absent, slimy, with entire margin; reverse yellow to brick.
Notes: Fusicolla quarantenae, an endophyte of Melocactus
zehntneri, is morphologically reminiscent of Fu. betae, Fu. epistroma, and Fu. septimanifiniscientiae, all of which produce mainly
3-septate macroconidia. Fusicolla betae and Fu. epistroma differ
by having larger conidia (50–60 μm and 19–45 μm long,
53
CROUS
ET AL.
Fig. 21. Dialonectria spp. A–D. Ascomata on natural substrate. E. Surface view of perithecial wall in 2 % KOH. F–H. Asci. I–M. Ascospores (L, M. Surface view). N, O.
Conidiophores and conidiogenous cells. P. Macroconidia. A, B. Dialonectria episphaeria (photos P. Mlcoch). C, D, F, M. Dialonectria episphaeria (CBS H-19716). E, G, K.
Dialonectria sanguinea (CBS H-2127). H–J, L. Dialonectria episphaeria (CBS H-2662). N–P. Dialonectria episphaeria (CBS 125494). Scale bars: A–D = 100 μm; I, L, M = 5 μm
(I applies to J and K); all others = 10 μm.
Fig. 22. Fusarium spp. A–D. Ascomata on natural substrate. E. Surface view of perithecial wall in 2 % KOH. F–H. Asci. I–M. Ascospores (M. Surface view). N–P. Conidiophores and conidiogenous cells. Q–T. Microconidia. U–A2. Macroconidia. A. Fusarium graminearum (photo P. Cannon). B, C, F. Fusarium sambucinum [adapted from
Wergen (2018)]. D. Fusarium sp. (HPC 2244). E. Fusarium cf. tricinctum (CBS H-12819). G, I. Fusarium lateritium (photo P. Cannon). H, K. Fusarium equiseti (CBS H-12817). J.
Fusarium sambucinum (BPI 632307). L, M. Fusarium sambucinum (CBS H-12818). N. Fusarium avenaceum (CPC 30660). O, Q. Fusarium fredkrugerii (CBS 144209). P, W.
Fusarium prieskaense (CBS 146498). R. Fusarium madaense (CBS 146669). S. Fusarium globosum (CBS 428.97). T. Fusarium echinatum (CBS 146497). U. Fusarium
avenaceum (CBS 408.86). V. Fusarium caeruleum (CBS 146590). X. Fusarium longicaudatum [CBS 123.73, adapted from Xia et al. (2019)]. Y. Fusarium transvaalense [CBS
144211, adapted from Sandoval-Denis et al. (2018b)]. Z. Fusarium gamsii (CBS 143610). A1. Fusarium oxysporum [CBS 144134, adapted from Lombard et al. (2019b)]. A2.
Fusarium convolutans [CBS 144207, adapted from Sandoval-Denis et al. (2018b)]. Scale bars: A–D = 100 μm; I–M, Q–T = 5 μm; all others = 10 μm.
54
FUSARIUM
REDELIMITED
Fig. 23. Fusarium echinatum (CBS 146497). A–D. Aerial conidiophores. E–G. Conidiogenous cells on aerial conidiophores. H, I. Microconidia. J. Sporodochia formed on the
surface of carnation leaves. K. Sporodochial conidiophores and conidiogenous cells. L. Macroconidia. Scale bars: A = 20 μm; J = 100 μm; all others = 10 μm.
respectively, Gerlach & Nirenberg 1982). The absence of chlamydospores in Fu. quarantenae further differentiates this species
from Fu. epistroma and Fu. septimanifiniscientiae (Gerlach &
Nirenberg 1982, Crous et al. 2018).
Fusicolla meniscoidea L. Lombard & Sand.-Den., sp. nov.
MycoBank MB 838662. Fig. 27.
Etymology: From Greek meniskos, crescent, in reference to the
shape of its conidia.
Typus: Australia, from soil, unknown collection date (before
1978), unknown collector (holotype CBS H-24662, culture extype CBS 110189 = FRC E-0086).1
Conidiophores arising laterally or terminally from somatic hyphae
50–70 μm long, simple or sparingly branched laterally, straight,
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hyaline, smooth- and thin-walled, bearing terminal and lateral
conidiogenous cells, or more commonly reduced to single conidiogenous cells borne laterally on the substrate and aerial hyphae.
Conidiogenous cells monophialidic, subcylindrical, cylindrical to
slightly subulate, 10.5–35 × 2–3.5 μm, smooth- and thin-walled,
without noticeable periclinal thickening, a minute apical collarette
can be present. Macroconidia falcate, tapering gently towards both
ends, apical cell often hooked with a blunt to pointy apex, basal cell
obtuse to poorly developed, foot-shaped, 0–2(–3)-septate, predominantly 1-septate, hyaline, smooth- and thin-walled; 0-septate
(8–)9–13(–15) × 2–3.5 μm (av. 11.1 × 2.9 μm); 1-septate, (9–)
11.5–15(–17.5) × 2.5–3.5 μm (av. 13.1 × 2.9 μm); 2-septate,
13–17.5(–18) × 2.5–4 μm (av. 15.4 × 3 μm); 3-septate,
20–24.5(–25.5) × 3–3.5 μm (av. 22.6 × 3.3 μm). Microconidia,
chlamydospores and sexual morph not observed.
55
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ET AL.
Fig. 24. Fusarium prieskaense (CBS 146498). A–D. Aerial conidiophores. E–G. Conidiogenous cells on aerial conidiophores. H. Microconidia. I. Sporodochia formed on the
surface of carnation leaves. J–L. Sporodochial conidiophores and conidiogenous cells. M. Macroconidia. Scale bars: A, B = 20 μm; I = 100 μm; all others = 10 μm.
Culture characteristics: Colonies on PDA reaching
21–30 mm diam at 25 °C after 7 d. Surface white to pale luteus
at periphery, centre salmon to pale orange, flat to slightly radially
folded, membranous to slimy, margin entire to slightly undulate;
reverse luteous to pale salmon at centre. On OA, pale luteous to
pale salmon, flat, membranous, margin entire; reverse pale
luteous.
Notes: Fusicolla meniscoidea is here introduced based on an
isolate originally misidentified as Bisifusarium dimerum.
Despite the great genetic differences and phylogenetic distance, the two taxa share similar morphological traits, particularly regarding macroscopic aspects of colonial growth, and the
shape and size of conidiophores and conidia. However, unlike
in B. dimerum, conidia of Fu. meniscoidea present a much
more pronounced curvature involving both conidial planes
(somewhat parallel walls), while foot-shaped basal cells are
56
less evident or absent. Fusicolla aqueductuum, Fu. betae, Fu.
quarantenae, and Fu. violacea are all morphologically related to
Fu. meniscoidea by showing similar conidial septation ranges
and lacking chlamydospores. Conidial size in Fu. meniscoidea
is, however, markedly reduced and often closer to the lower
limits of the conidial size of all the aforementioned species.
Another species also described here, Fusicolla sporellula, lacks
chlamydospores but has similar, although smaller, conidia with
a reduced range of septa (0- or 1-septate). It furthermore differs
from Fu. meniscoidea by its shorter and doliiform conidiogenous cells.
Fusicolla sporellula Sand.-Den. & L. Lombard, sp. nov.
MycoBank MB 838663. Fig. 28.
Etymology: From Latin, very small spores, in reference to its very
small conidia.
FUSARIUM
REDELIMITED
Fig. 25. Fusicolla spp. A. Slimy macroscopic growth on natural substrate. B–E. Ascomata on natural substrate. F. Ostiolar hairs. G. Asci. H. Ascospores. I–K. Conidiophores
and conidiogenous cells. L–N. Macroconidia. A. Fusicolla merismoides (photo J. Cunningham). B. Fusicolla melogrammae [CLL 16006, adapted from Crous et al. (2016)].
C–H. Fusicolla ossicola (photos N. Aplin and P. Cannon). I. Fusicolla merismoides (photo P. Cannon). J, K, M. Fusicolla aquaeductuum (CBS 734.79). L. Fusicolla violacea
(CPC 38810). N. Fusicolla matuoi (CBS 581.78). Scale bars: B–E = 100 μm; F, H. 5 μm; all others = 10 μm.
Fig. 26. Fusicolla quarantenae (URM 8367). A. Host. B–G. Conidiophores, conidiogenous cells and conidia. H. Macroconidia. Scale bars = 10 μm.
Typus: South Africa, Transkei, from soil, unknown collection
date (before 1983), unknown collector (holotype CBS H-24663,
culture ex-type CBS 110191 = FRC E-0139).
Conidiophores arising laterally from substrate and aerial
hyphae 14–35 μm long, simple or laterally and verticillately
branched, straight, hyaline, smooth- and thin-walled, or
reduced to single conidiogenous cells. Conidiogenous cells
monophialidic, doliiform, short lageniform to subulate
7.5–20 × 2.5–4 μm, smooth- and thin-walled, with or without
inconspicuous periclinal thickening, collarettes absent; or
reduced to short phialidic pegs emerging laterally from hyphae,
1–5 × 1–2.5 μm, smooth- and thin-walled, with inconspicuous
periclinal thickening and an often conspicuously flared collarette. Macroconidia lunate to falcate, moderately to strongly
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dorsiventrally curved, slightly narrowing towards both ends,
apical cell blunt, more or less hooked, basal cell obtuse
to poorly developed, foot-shaped, hyaline, thin- and smoothwalled, 0- or 1-septate, predominantly 1-septate, 0-septate:
(11–)12–14( –15) × 2–3( –3.5) μm (av. 13.2 × 2.7 μm),
1-septate: (11.5–)13–16.5( –20) × 2.5–3.5 μm (av.
14.6 × 2.8 μm). Microconidia, chlamydospores, and sexual
morph not observed.
Culture characteristics: Colonies on PDA reaching 24–31 mm
diam at 25 °C after 7 d. Surface white, luteous to orange, flat to
slightly radially folded, membranous to slimy, margin entire;
reverse pale luteous to saffron, peach at centre. On OA, pale
luteous to peach, flat, membranous with filiform to undulate
margins; reverse pale peach to saffron.
57
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ET AL.
Fig. 27. Fusicolla meniscoidea (CBS 110189). A–D. Conidiophores. E–H. Conidiogenous cells. I. Macroconidia. Scale bars: A–D, G–I = 10 μm; E, F = 5 μm.
Fig. 28. Fusicolla sporellula (CBS 110191). A–C. Conidiophores. D–F. Conidiogenous cells. G. Macroconidia. Scale bars = 10 μm.
Notes: Fusicolla sporellula presents the smallest conidia
described to date for any species in this genus. This taxon is
phylogenetically and morphologically related to Fu. meniscoidea,
from which it can be differentiated by its smaller and less septate
conidia, and by the characteristic doliiform shape of its conidiogenous cells.
Geejayessia Schroers et al., Stud. Mycol. 68: 124. 2011. Figs
8, 29.
Type species: Geejayessia cicatricum (Berk.) Schroers, Stud.
Mycol. 68: 124. 2011.
(See F. cicatricum in List section for synonyms)
Ascomata perithecial, caespitose, with erumpent, byssoid or
densely prosenchymatous stroma, superficial, broadly ampulliform with short ostiolar neck to broadly ellipsoidal, not collapsing
when dry, pale orange, brownish to reddish orange, bright,
reddish black or black, changing colour in KOH if not black and
58
becoming purple in lactic acid, mostly smooth-walled, lacking
hairs or appendages. Ascomatal wall consists of a single region,
comprising several layers of morphologically similar cells. Asci
cylindrical to clavate, with a broadly rounded or flattened apex,
with or without a minute refractive ring, 8-spored, mostly overlapping, uniseriate or biseriate above and uniseriate below. Ascospores broadly ellipsoidal to ellipsoidal, 1-septate, slightly
constricted at the septum, verrucose, hyaline to pale brown.
Conidiophores mononematous (aerial conidiophores) or grouped
on sporodochia. Aerial conidiophores unbranched, sympodial or
irregularly branched, bearing terminal or lateral phialides, often
reduced to single phialides. Conidiogenous cells monophialidic,
subcylindrical to cylindrical, smooth- and thin-walled, with periclinal thickening inconspicuous or absent. Aerial conidia hyaline,
smooth- and thin-walled, of two types: microconidia, present in
some species, ellipsoidal to fusoid, 0- or 1-septate, with rounded
ends, straight to slightly curved; macroconidia typically formed on
sporodochia, falcate, straight to gently curved dorsiventrally,
FUSARIUM
REDELIMITED
Fig. 29. Geejayessia spp. A–E. Ascomata on natural substrate. F. Surface view of perithecial wall in 2 % KOH. G–I. Asci. J–M. Ascospores. N, O. Conidiophores and
conidiogenous cells. P, Q. Macroconidia. A, C. Geejayessia cicatricum [CBS H-20375, adapted from Schroers et al. (2011)]. C. Geejayessia cicatricum (CBS H-20374). D, H, K,
M. Geejayessia atrofusca (CBS H-20381). E–G, I, J, L. Geejayessia desmazieri (CBS H-20372). N, O, Q. Geejayessia atrofusca (CBS 502.94). P. Geejayessia cicatricum (CBS
125549). Scale bars: A = 500 μm; B, D, E = 200 μm; C = 100 μm; J–M = 5 μm; all others = 10 μm.
3–8-septate, with a blunt apical cell and well-developed footshaped basal cell. Sporodochia cream to pale yellow; sporodochial conidiophores verticillately branched and densely packed,
consisting of short, smooth- and thin-walled stipes bearing an
apical whorl of 2–3 monophialides; sporodochial conidiogenous
cells monophialidic, cylindrical to subcylindrical, smooth- and
thin-walled, with reduced or flared collarette. Chlamydospores
unknown.
[Description adapted from Schroers et al. (2011) and Lombard
et al. (2015)].
Diagnostic features: Pale orange, brownish to reddish orange,
bright red, reddish black to black, mostly smooth-walled perithecia with short ostiolar neck producing clavate to cylindrical
asci bearing ellipsoidal, 1-septate, verrucose ascospores and
asexual morphs producing only macroconidia on sporodochia or
micro- and macroconidia on elongate subulate to subcylindrical
aerial conidiophores with monophialides. Chlamydospores
absent.
aggregated, unbranched or sparsely branched, bearing up to
three phialides; complex conidiophores solitary or aggregated in
small sporodochia, repeatedly and irregularly branched. Conidiogenous cells monophialidic, cylindrical, tapering towards the
apex. Microconidia 0- or 1-septate, ovoid to fusoid to ellipsoidal,
with a minutely or clearly laterally displaced hilum, formed in
heads on solitary conidiophores or as masses on sporodochia.
Macroconidia straight, cylindrical, 1–3(–4)-septate, with both
ends obtusely rounded, base sometimes with a visible, centrally
located to laterally displaced hilum, forming flat domes of slimy
masses. Chlamydospores globose to subglobose, thick-walled,
intercalary or solitary, initially hyaline, becoming brown with age.
[Description adapted from Chaverri et al. (2011)].
Diagnostic features: Red, mostly smooth-walled perithecia with
conical papilla or flattened apex producing cylindrical asci
bearing ellipsoidal, 1-septate ascospores and cylindrocarpon-like
asexual morph characterised by 1–3(–4)-septate macroconidia
with centrally located to laterally displaced hilum.
Ilyonectria P. Chaverri & C. Salgado, Stud. Mycol. 68: 69. 2011.
Fig. 8.
Luteonectria Sand.-Den., L. Lombard, Schroers & Rossman,
gen. nov. MycoBank MB 838664. Figs 8, 30.
Type species: Ilyonectria destructans (Zinssm.) Rossman et al.,
Stud. Mycol. 80: 217. 2015.
(See F. aderholdii in List section for synonyms)
Etymology: Name refers to the luteous coloured, nectria-like
ascomata characteristic of these fungi.
Ascomata perithecial, solitary or gregarious, non-stromatic, superficial, globose to subglobose or ovoid to obpyriform, red,
turning purple to dark purple in KOH, pigment dissolving in lactic
acid, not collapsing when dry, with broadly conical papilla or
flattened apex, smooth to slightly rugulose, lacking hairs or appendages. Ascomatal wall of two regions: outer region of thickwalled, pigmented cells forming a textura globosa; inner region of
compressed, flattened cells, becoming thinner towards the
centrum. Asci narrowly clavate to cylindrical, 8-spored, apex
subtruncate, with inconspicuous apical ring, uniseriate. Ascospores ellipsoidal, 1-septate, hyaline, smooth. Conidiophores
simple or complex or sporodochial; simple conidiophores arising
laterally or terminally from aerial mycelium, solitary or loosely
www.studiesinmycology.org
Type species: Luteonectria albida (Rossman) Sand.-Den. & L.
Lombard
Ascomata perithecial, gregarious on a well-developed stroma
composed of pseudoparenchymatous cells, covered with loose,
white hyphae, smooth and thin-walled, globose to pyriform, offwhite to pale luteous, becoming ochraceous when dry, with a
broadly rounded and papillate apical region, not changing colour
in KOH or lactic acid, short setae-like hairs sometimes emerging
from perithecial wall. Asci clavate with simple apex, 8-spored,
ascospores overlapping irregularly uniseriate to biseriate. Ascospores fusiform with rounded ends, 3-septate, slightly constricted at septum, hyaline, becoming pale yellow-brown,
smooth-walled to finely striate. Conidiophores mononematous,
59
CROUS
ET AL.
Fig. 30. Luteonectria albida. A–C. Ascomata on natural substrate. D. Surface view of perithecial wall in lactic acid. E. Detail of ascomata hair. F. Asci. G–J. Ascospores (J.
Surface view). K, L. Conidiophores and conidiogenous cells. M. Macroconidia. A, C. BPI 550103. B. BPI 1108874. D–J. BPI 1108875. K–M. CBS 102683. Scale bars: A,
B = 100 μm; C = 50 μm; all others = 10 μm.
septate and irregularly branched, bearing terminal phialides.
Conidiogenous cells monophialidic, cylindrical to subcylindrical,
smooth- and thin-walled, with periclinal thickening inconspicuous
to absent. Macroconidia fusoid and multiseptate, 1–7-septate,
curved, hyaline, with a wide, blunt apical cell and a poorly- to
well-developed, foot-shaped basal cell. Micro- and mesoconidia
unknown. Chlamydospores unknown.
[Description adapted from Rossman (1983) and Schroers et al.
(2011)].
1999, G.J. Samuels et al., BPI 746587, culture CBS 102683. Jamaica, Newcastle, Chesterville Youth Developmental Camp, on undetermined host, 8 Jan.
1971, A.Y. Rossman, BPI 550103. Venezuela, Los Venados, El Avila, along Trail
1–2 km above Los Venados, El Avila, on undetermined substrate, 24 Jul. 1972,
K.P. Dumont et al., BPI 1108875.
Diagnostic features: Off-white to pale luteous perithecia that do
not change colour on KOH or lactic acid, formed on welldeveloped stroma producing clavate asci containing fusiform, 3septate, finely striate ascospores and fusarioid asexual morph
characterised by monophialides producing robust multiseptate
conidia from aerial conidiophores, lacking micro- and mesoconidia, and chlamydospores.
Typus: Germany, Berlin, from soil with roots of Hedera helix,
unknown date and collector (holotype BBA 72279 in B, culture
ex-type BBA 72279 = NRRL 54600).
Luteonectria albida (Rossman) Sand.-Den. & L. Lombard,
comb. nov. MycoBank MB 838665.
Basionym: Nectria albida Rossman, Mycol. Pap. 150: 79. 1983.
Synonyms: Albonectria albida (Rossman) Guu & Y.M. Ju, Bot.
Stud. (Taipei) 48: 189. 2007.
Fusarium albidum (Rossman) O'Donnell & Geiser, Phytopathology 103: 404. 2013.
Typus: Jamaica, Hanover Parish, Dolphin Head Mt. near
Askenish, on bark of woody stem of unknown host, 22 Jan. 1971,
R.P. Korf et al. (holotype CUP-MJ 942, culture ex-type ATCC
44543 = CTR 71-110 = BBA 67603 = NRRL 13950 = NRRL
22152).
Description and illustration: Rossman (1983), Guu et al. (2007),
Schroers et al. (2011).
Additional material examined: Costa Rica, Limon, Central Distrito Valle, Valle del
Estrella, Selva Biologia Hitoi Caneri, 100 –150 m alt, on bark of living tree, 7 Jul.
60
Luteonectria nematophila (Nirenberg & Hagedorn) Sand.-Den.
& L. Lombard, comb. nov. MycoBank MB 838666.
Basionym: Fusarium nematophilum Nirenberg & Hagedorn,
Nachrichtenbl. Deutsch. Pflanzenschutzdienstes 60: 214. 2008.
Description and illustration: Nirenberg & Hagedorn (2008).
Macroconia (Wollenw.) Gr€afenhan et al., Stud. Mycol. 68: 101.
2011. Figs 8, 31.
Basionym: Nectria sect. Macroconia Wollenw., Angew. Bot. 8:
179. 1926.
Type species: Macroconia leptosphaeriae (Niessl) Gr€afenhan &
Schroers, Stud. Mycol. 68: 102. 2011.
Synonyms: Nectria leptosphaeriae Niessl, in Krieger, Fungi
Saxon. Exs.: no. 165. 1886.
Cucurbitaria leptosphaeriae (Niessl) Kuntze, Revis. Gen. Pl. 3:
461. 1898.
Hypomyces leptosphaeriae (Niessl) Wollenw., Fusaria Autogr.
Delin. 1: 57. 1916.
Lasionectria leptosphaeriae (Niessl) Petch, Trans. Brit. Mycol.
Soc. 21: 268. 1938.
Cosmospora leptosphaeriae (Niessl) Rossman & Samuels, Stud.
Mycol. 42: 122. 1999.
Ascomata perithecial, solitary, with stroma inconspicuous or
absent, subglobose with or without a small apical papilla, orange
to carmine red, turning dark red to violet in KOH, sometimes with
FUSARIUM
hyphal hairs arising from the outer wall. Asci cylindrical to
narrowly clavate, with a simple apex, 8-spored, uniseriate or
partially biseriate. Ascospores yellowish, 1-septate, smooth,
sometimes becoming striate when mature. Conidiophores initially
as lateral phialides on somatic hyphae, later monochasial to
verticillate, hyaline. Conidiogenous cells monophialidic, cylindrical to subulate, hyaline. Microconidia rare or absent, ellipsoidal to
allantoid, hyaline. Macroconidia subcylindrical to curved, apical
cell conical or hooked, basal cell poorly- to well-developed, footshaped, 3–7(–14)-septate, hyaline. Chlamydospores absent to
rare, globose, single, in pairs or chains in hyphae.
[Description adapted from Gr€afenhan et al. (2011)].
Diagnostic features: Orange-red to carmine-red perithecia with or
without a small papilla producing cylindrical to narrowly clavate
asci bearing 1-septate ascospores that sometimes become
striate when mature, and asexual morphs characterised by
verticillate conidiophores producing large, multiseptate fusarioid
macroconidia.
Macroconia bulbipes Crous & Sand.-Den., sp. nov. MycoBank
MB 838667. Fig. 32.
Etymology: Named after the shape of the basal cell, which is
commonly swollen, bulbous.
Typus: South Africa, Western Cape Province, Swellendam,
Bontebok National Park, from Erica sp. associated with Dimerosporiopsis engleriana, 24 Sep. 2018, A.R. Wood (holotype
CBS H-24664, culture ex-type CBS 146679 = CPC 37138).
Conidiophores commonly aggregated into sporodochia, more
rarely simple (aerial). Aerial conidiophores borne laterally on hyphae and commonly reduced to single conidiogenous cells, hyaline,
thin- and smooth-walled, 23.5–39.6 μm long; conidiogenous cells
monophialidic, subcylindrical to cylindrical, hyaline, (23–)
24–25(–26.5) × 3–4 μm, without discernible periclinal thickening or
collarettes. Sporodochia abundantly formed on carnation leaves
and on the agar surface, pink to pink-brown coloured. Sporodochia
REDELIMITED
light orange-peach, turning dark brick coloured in old cultures;
sporodochial conidiophores irregularly or verticillately branched,
40–55.5 μm long, irregularly branched, bearing lateral and terminal
solitary monophialides. Sporodochial conidiogenous cells monophialidic, cylindrical to subcylindrical to subulate, (8–)
14.5–26.5(–30.5) × 3.5–5.5 μm with inconspicuous periclinal
thickening, flared collarettes absent. Microconidia absent. Macroconidia straight to moderately dorsiventrally curved, tapering toward
the apex, apical cell conical or hooked, and slightly extended, basal
cell well-developed, foot shaped, commonly irregularly swollen
at bottom, (2–)3–5(–6)-septate, predominantly 4-septate,
hyaline, thick- and smooth-walled: 2-septate conidia:
43–45.5 × 5–5.5 μm (av. 44.2 × 5.1 μm); 3-septate conidia: (38.5–)
41–53(–55) × 5–6 μm (av. 48.1 × 5.4 μm); 4-septate conidia:
(45.5–)50–62(–67.5) × 5–6(–7) μm (av. 56.1 × 5.8 μm); 5-septate
conidia: (58–)61–77(–80.5) × 5–6.5 μm (av. 68.9 × 5.8 μm); 6septate conidia: (70–)71–74 × 5.5–6.5(–7) μm (av. 72.1 × 6.4 μm);
overall: (38.5–)48–68(–80.5) × 5–6(–7) μm (av. 58 × 5.7 μm).
Chlamydospores commonly formed in the substrate mycelium and
conidia, spherical to subspherical, 8.5–11(–12.5) μm diam, hyaline
and smooth-walled. Sexual morph not observed.
Culture characteristics: Colonies on PDA reaching
21–24 mm diam at 25 °C after 7 d. Surface salmon to buff, flat,
membranous to velvety, with scant aerial mycelium and pionnotal, margin white and regular; reverse pale salmon with radial
white to pale yellow patches. On OA, salmon to buff, flat,
membranous and pionnotal, with regular margin; reverse pale
pink to salmon.
Additional material examined: South Africa, Western Cape Province,
Swellendam, Bontebok National Park, from Erica sp. associated with Dimerosporiopsis engleriana, 24 Sep. 2018, A.R. Wood, culture CBS 146678 = CPC
37137.
Notes: Macroconia bulbipes resolved as the closest phylogenetic
relative to Ma. gigas and Ma. papilionacearum (Fig. 13). The
former is, however, clearly distinguished morphologically by its
Fig. 31. Macroconia spp. A–D. Ascomata on natural substrate. E. Surface view of perithecial wall in 2 % KOH. F. Asci. G–J. Ascospores (J. Surface view). K–M. Conidiophores and conidiogenous cells. N. Microconidia. O, P. Macroconidia. A. Macroconia cupularis [HMAS 97514, adapted from Luo & Zhuang (2008)]. B, C. Macroconia
leptosphaeriae (photo P. Mlcoch). D. Macroconia gigas [HMAS 99592, adapted from Luo & Zhuang (2008)]. E–J. Macroconia leptosphaeriae (CBS H-15051). K, L. Macroconia
phlogioides (CBS 125496). M, N. Macroconia leptosphaeriae (CBS 10001). O. Macroconia phlogioides (CBS 146500). P. Macroconia bulbipes (CBS 146679). Scale bars:
A–D = 100 μm; G–J = 5 μm; all others = 10 μm.
www.studiesinmycology.org
61
CROUS
ET AL.
Fig. 32. Macroconia bulbipes (CBS 146679). A–D. Conidiophores and conidiogenous cells. E, F. Sporodochia formed on the agar surface. G, H. Detail of macroconidia basal
cells. I. Macroconidia. Scale bars: E, F = 100 μm; G, H = 5 μm; all others = 10 μm.
smaller and less septate conidia (rarely up to 80.5 μm long and
up to 6-septate vs longer than 100 μm and more than 10-septate
in the latter two species). On the contrary, the asexual morph of
Ma. bulbipes is closer to that of Ma. leptosphaeriae and Ma.
sphaeriae (recognised as two distinct species in Gr€afenhan et al.
2011). The conidia of Ma. bulbipes, however, differ by having
commonly swollen basal cells.
Macroconia phlogioides Sand.-Den. & Crous, sp. nov.
MycoBank MB 838668. Fig. 33.
Etymology: From Greek floga. Referring to the flame-like
macroscopic semblance of the sporodochia.
Typus: South Africa, Limpopo Province, Tzaneen, on leaf of
Encephalartos sp., 2019, P.W. Crous (holotype CBS H-24665,
culture ex-type CBS 146501 = CPC 35389).
Conidiophores simple (aerial) or aggregated into sporodochia.
Aerial conidiophores often borne laterally on hyphae and
reduced to single conidiogenous cells, rarely 1-septate, hyaline,
thin- and smooth-walled, 13–17 × 26–32 μm; conidiogenous
cells monophialidic, subcylindrical to cylindrical, hyaline, (13–)
16–24(–27.5) × (3.5–)4–5 μm conidiogenous opening rather
wide, with inconspicuous periclinal thickening and no discernible
apical collarettes. Sporodochia orange-pink to pink-brown coloured, often acquiring a flame-like, somewhat pointy macroscopic appearance and later merging into pionnotal crusts;
sporodochial conidiophores irregularly or verticillately branched,
37.5–46 μm long, often bearing groups of 2–3 conidiogenous
cells; sporodochial conidiogenous cells monophialidic, subcylindrical to subulate, (10–)18.5–26(–30) × (2.5–)3.5–5 μm
with inconspicuous periclinal thickening, collarettes absent.
Microconidia absent. Macroconidia robust, often with a nearly
62
straight central portion and markedly curved and tapering towards both ends, apical cell conical to hooked, basal cell welldeveloped, foot-shaped, (1–)9–15(–19)-septate, predominantly
11-septate, hyaline, thick- and smooth-walled: 9-septate conidia:
(106.5–)119.5–140(–143.5) × 7.5–8.5(–9) μm (av. 129 × 8 μm);
10-septate conidia: (116–)120–144.5(–164) × (7–)7.5–9 μm
(av. 132 × 8 μm); 11-septate conidia: (122–)127–140
(–153.5) × 7.5–9(–9.5) μm (av. 134 × 8.5 μm); 12-septate
conidia: (119–)127.5–146.5(–153) × 7.5–9.5(–10) μm (av.
137 × 8.5 μm); 13-septate conidia: (128–)132–155
(–172) × (7–)8–9(–10) μm (av. 143.5 × 8.5 μm); 14-septate
conidia: (133.5–)136–157(–168) × 8–9.5 μm (av.
146.5 × 9 μm); 15-septate conidia: 147–163.5(–173.5) ×
8.5–9.5(–10) μm (av. 155 × 9 μm); overall: (86–)
123.5–150(–175) × (7–)8–9(–10) μm (av. 137 × 8.5 μm).
Chlamydospores and sexual morph not observed.
Culture characteristics: Colonies on PDA reaching
17–25 mm diam at 25 °C after 7 d. Surface salmon, buff to rosy
buff, flat to slightly raised at centre, glabrous or with central
patches of white, dense aerial mycelium; membranous to dusty
with regular margin; reverse pale luteous to sulphur yellow, with
salmon patches. On OA, salmon, flat, membranous, inconspicuously radially folded with regular margin; reverse pale pink to
luteous with more intense salmon-coloured patches.
Additional material examined: South Africa, Limpopo Province, Tzaneen, on leaf
of Encephalartos sp., 2019, P.W. Crous, culture CBS 146500 = CPC 35388. USA,
Arizona, Huachuca Mountains, Miller Canyon, on branch of Quercus sp. in
stream, 1 Oct. 2008, T. Gr€afenhan, culture CBS 125496.
Notes: Macroconia phlogioides is morphologically related to Ma.
papilionacearum and Ma. gigas. These three species are characterised by producing robust and large (often above 100 μm
FUSARIUM
REDELIMITED
Fig. 33. Macroconia phlogioides (CBS 146501). A–C. Conidiophores. D, E. Conidiogenous cells. F, G. Sporodochia formed on the agar surface. H. Macroconidia. Scale bars:
B, C = 20 μm; F, G = 50 μm; all others = 10 μm.
long) macroconidia. Unlike the above-mentioned species, however, conidia of Ma. phlogioides tend to present a higher number
of septa (up to 19 vs up to 12 and 14, for Ma. papilionacearum
and Ma. gigas, respectively), with rounder and less tapered
apical cells, contrasting with the elongated conidial apices of Ma.
gigas. Conidia of Ma. phlogioides also differ by having a more
pronounced and continuous curvature compared to Ma. gigas
and Ma. papilionacearum. These three species are clearly
different phylogenetically, clustering in distant monophyletic lineages of the genus (Fig. 13).
Mariannaea G. Arnaud ex Samson, Stud. Mycol. 6: 74. 1974.
Fig. 8.
Type species: Mariannaea elegans (Corda) Samson, Stud.
Mycol. 6: 75. 1974.
Basionym: Penicillium elegans Corda, Icon. Fung. 2: 17. 1838.
Synonyms: Hormodendron elegans (Corda) Bonorden, Handb.
Allg. Mykol.: 76. 1851.
Spicaria elegans (Corda) Harz., Bull. Soc. Imp. Naturalistes
Moscou 44: 238. 1871.
Paecilomyces elegans (Corda) Mason & Hughes, Mycol. Pap.
45: 27. 1951.
Ascomata perithecial, solitary, non-stromatic or on inconspicuous
stroma, superficial, globose with flat apex, not collapsing or
laterally pinched when dry, pale yellow, orange or brown, not
reacting in KOH, smooth-walled to slightly rugose, lacking hairs
or appendages. Asci cylindrical to narrowly clavate, 8-spored
sometimes with inconspicuous apical ring, uniseriate to apically
biseriate. Ascospores 1-septate, hyaline, smooth-walled to
spinulose. Conidiophores verticillate to penicillate, hyaline, with
phialides arising directly from the stipe or forming whorls of
metulae on lower parts of stipe; stipe hyaline, becoming yellow-
www.studiesinmycology.org
brown at the base. Conidiogenous cells monophialidic, ampulliform, hyaline, usually with obvious periclinal thickening and
inconspicuous collarettes. Conidia limoniform, aseptate, hyaline,
in chains that collapse to form slimy heads. Chlamydospores
globose to ellipsoidal, hyaline, formed in intercalary chains.
[Description adapted from Samson (1974), Gr€afenhan et al.
(2011) and Lombard et al. (2015)].
Diagnostic features: Pale yellow, orange to brown perithecia with
flattened apex producing cylindrical to narrowly clavate asci bearing
1-septate ascospores and asexual morphs characterised by verticillate to penicillate conidiophores producing small, aseptate,
limoniform conidia in chains that collapse into slimy heads.
Microcera Desm., Ann. Sci. Nat. Bot., ser. 3, 10: 359. 1848. Figs
8, 34.
Synonym: Pseudomicrocera Petch, Trans. Brit. Mycol. Soc. 7:
164. 1921.
Type species: Microcera coccophila Desm., Ann. Sci. Nat. Bot.,
ser. 3, 10: 359. 1848.
(See F. coccophilum in List section for synonyms)
Ascomata perithecial, solitary or gregarious, with stroma and/or
byssus covering host, globose, with a blunt papilla, orange to
dark red, turning dark red or violet in KOH, finely roughened. Asci
cylindrical to narrowly clavate, with an apical ring, 8-spored.
Ascospores hyaline to pale yellow-brown, 1(–3)-septate,
smooth, sometimes becoming tuberculate when mature. Conidiophores as lateral phialides on somatic hyphae, becoming
monochasial, verticillate to penicillate, hyaline, forming discrete
sporodochia or synnemata on the host. Conidiogenous cells
monophialidic, cylindrical to subulate to subclavate, hyaline.
Macroconidia pale, orange, pink or bright red in mass, subcylindrical, moderately or conspicuously curved, apical cell often
63
CROUS
ET AL.
slightly or conspicuously hooked, basal cell papillate to welldeveloped, foot-shaped, (0–)3–5(–12)-septate, hyaline.
[Description adapted from Gr€afenhan et al. (2011)].
Diagnostic features: Orange to dark red perithecia with a blunt
papilla producing cylindrical to narrowly clavate asci bearing
yellow-brown, 1(–3)-septate ascospores; asexual morphs characterised by verticillate to penicillate conidiophores producing
small macroconidia; species typically associated with scale
insects.
Neocosmospora E.F. Sm., Bull. U.S.D.A. 17: 45. 1899. Figs
8, 35.
Type species: Neocosmospora vasinfecta E.F. Sm., Bull.
U.S.D.A. 17: 45. 1899.
(See F. neocosmosporiellum in List section for synonyms)
Ascomata perithecial, solitary or gregarious, non-stromatic or
with reduced basal stroma, superficial, globose to pyriform, not
collapsing when dry, orange-brown to bright red, darkening or
becoming purple in KOH, papillate or with short ostiolar neck,
commonly tuberculate, rarely smooth-walled, lacking hairs or
appendages. Ascomatal wall of two regions: outer region of thickwalled, pigmented cells forming a textura angularis; inner region
of elongate, hyaline, thin-walled cells, becoming thinner towards
the centrum. Asci saccate, clavate to cylindrical, unitunicate,
apex simple, rounded or flattened, 8-spored, uniseriate to
irregularly biseriate. Ascospores globose to ellipsoidal, with or
without slightly truncate ends, typically 1-septate, hyaline when
young becoming yellow golden-brown at maturity, thick-walled,
longitudinally striate; ascospores in some species 0-septate,
cerebriform or spinulose. Conidiophores mononematous (aerial)
or grouped on sporodochia, or somewhat erect, loosely
branched sporodochial pustules. Aerial conidiophores simple,
sparsely to highly branched; aerial conidiogenous cells monophialidic, elongate subulate to subcylindrical. Aerial conidia
hyaline, smooth- and thick-walled, of two types: microconidia
subglobose, ellipsoidal to somewhat clavate, 0–2(–4)-septate,
borne in false heads on phialides; macroconidia falcate, slightly
to strongly curved dorsiventrally, 1-septate to multiseptate, with
blunt to hooked to slightly pointed apical cell and papillate to welldeveloped foot-shaped basal cell. Sporodochia cream, pale
luteous, light green, olivaceous, bluish, hazel to greyish sepia;
sporodochial conidiophores verticillately or sympodially branched
or sparingly branched and densely packed, consisting of short,
smooth- and thin-walled stipes bearing apical whorl of 2–4
monophialides; sporodochial conidiogenous cells monophialidic,
doliiform, short subcylindrical to subulate, smooth- and thinwalled, periclinal thickening and collarettes inconspicuous or
absent. Sporodochial macroconidia falcate, smooth- and thickwalled, straight or curved with parallel walls to unequally curved,
in some species clavate and asymmetrical, tapering towards
both ends, with a pointed to blunt to hooked apical cell and
papillate to well-developed foot-shaped basal cell. Chlamydospores globose to subglobose to ovoid to obovoid, hyaline to
pale golden brown, smooth-walled to slightly verrucose, terminal
or intercalary, solitary or in pairs or forming chains or aggregating
in some species to form buff, olive aeruginous or bluish
microsclerotia.
[Description adapted from Rossman et al. (1999) and SandovalDenis et al. (2019)].
Diagnostic features: Orange-brown to frequently bright, blood red
warted perithecia with papillate or short ostiolar neck producing
saccate, clavate to cylindrical asci bearing globose to ellipsoidal, 0or 1-septate, longitudinally striate, cerebriform or spinulose ascospores and asexual morphs producing micro- and macroconidia
on elongate subulate to subcylindrical aerial conidiophores with
monophialides or only macroconidia in sporodochia. Chlamydospores formed in hyphae, rarely observed in macroconidia.
Neocosmospora epipeda Quaedvl. & Sand.-Den., sp. nov.
MycoBank MB 838669. Fig. 36.
Fig. 34. Microcera spp. A–C. Ascomata on natural substrate. D. Surface view of perithecial wall in 2 % KOH. E, F. Asci. G–K. Ascospores (J, K. Surface view). L–N.
Conidiophores and conidiogenous cells. O–Q. Macroconidia. A. Microcera auranticola (photo N. Aplin). B, O. Microcera coccophila [adapted from Gr€afenhan et al. (2011)]. C.
Microcera larvarum [adapted from Gr€afenhan et al. (2011)]. D, F–J. Microcera coccophila (K(M) 165807). E, K. Microcera larvarum (photo P. Cannon). L, M, Q. Microcera rubra
(CBS 638.76). N, P. Microcera larvarum (CBS 169.30). Scale bars: A, B = 100 μm; G–K = 5 μm; all others = 10 μm.
64
FUSARIUM
REDELIMITED
Fig. 35. Neocosmospora spp. A–E. Ascomata on culture. F. Surface view of perithecial wall in 2 % KOH. G–I. Asci. J–Q. Ascospores (K, M, O, Q. Surface view). R–U. Aerial
conidiophores. V. Sporodochial conidiophores. W, X. Microconidia. Y–A5. Macroconidia. A, I, N, O. Neocosmospora vasinfecta (CBS 446.93). B. Neocosmospora sp. (CPC
34617). C, S, W, A1. Neocosmospora elegans (CBS 144396). D. Neocosmospora vasinfecta (CBS 863.70). E. Neocosmospora bataticola (CBS 144398). F, L, M. Neocosmospora ipomoeae (CBS 833.97). G. Neocosmospora robiniae (CBS 119601). H, J, K. Neocosmospora diminuta (CBS 144390). O, Q. Neocosmospora spinulosa (CBS H5443). R, V, A3. Neocosmospora solani (CBS 140079). T. Neocosmospora bataticola (CBS 144398). U. Neocosmospora suttoniana (CBS 143214). X. Neocosmospora
tonkinensis (CBS 115.40). Y. Neocosmospora longissima (CBS 126407). Z. Neocosmospora mori (CBS 145467). A2. Neocosmospora pseudoradicicola (CBS 145472). A4.
Neocosmospora keratoplastica (CBS 490.63). A5. Neocosmospora oligoseptata (CBS 143241). [A, C, S, T, W, Y, Z, A1, A2. Adapted from adapted from Sandoval-Denis et al.
(2019). R, V, A3, Adapted from Crous et al. (2019a). U, X, A4. Adapted from Sandoval-Denis & Crous (2018)]. Scale bars: A, B = 200 μm; C–E 100 μm; R–T = 20 μm; J–Q, W,
X = 5 μm; all others = 10 μm.
Etymology: From the Greek επ ίπεδα, flat; referring to the microconidia of this species commonly being flattened on one side.
Typus: Netherlands, from Bouvardia sp. imported from Uganda,
2019, W. Quaedvlieg (holotype CBS H-24666, culture ex-type
CBS 146523 = CPC 38310).
Conidiophores borne on the agar substrate and aerial mycelium,
78–230 μm tall, unbranched or more commonly sympodially
branched at various levels, bearing terminal single phialides;
www.studiesinmycology.org
aerial conidiogenous cells monophialidic, subulate, subcylindrical
to acicular, smooth- and thin-walled, 27.5–62 × 2–3.5 μm, short
apical collarettes and periclinal thickening inconspicuous or absent. Aerial conidia microconidial, arranged in false heads on
phialide tips, hyaline, broadly ellipsoidal, ellipsoidal to short
clavate, commonly asymmetrical with a somewhat flattened side,
smooth- and thin-walled, aseptate, (4.5–)6–10(–13.5) × (2–)
3–5 μm (av. 8 × 3.5 μm). Sporodochia pale luteous to orange,
formed abundantly on the surface of carnation leaves;
65
CROUS
ET AL.
sporodochial conidiophores laterally and irregularly branched
bearing apical groups of 2–3 monophialides; sporodochial
conidiogenous cells monophialidic, subulate to subcylindrical,
11–19.5 × 3–4.5 μm, smooth and thin-walled, with short, nonflared collarettes and inconspicuous or absent periclinal thickening. Sporodochial conidia falcate, almost straight to slightly
curved dorsoventrally, broadest near the half portion or the
upper third, tapering towards both ends, with a blunt to
somewhat pointy and slightly curved apical cell and an often welldeveloped foot-shaped basal cell, (3–)4–7(–8)-septate, predominantly 5-septate, hyaline, smooth- and thick-walled; 3septate conidia: 42.5 × 4.4 μm; 4-septate conidia: (41.5–)
44–58(–60) × 4–5 μm (av. 51.1 × 4.4 μm); 5-septate conidia:
(53.5–)59–69.5(–76) × 4–6 μm (av. 64.3 × 5 μm); 6-septate
conidia: 68–75.5(–79.5) × 4.5–6 μm (av. 71.7 × 5.3 μm);
7-septate conidia: (68–)69–74.5(–77) × 5–6 μm (av.
71.7 × 5.5 μm); 8-septate conidia: 74–75.5 × 5–6 μm (av.
74.7 × 5.3 μm); overall: (42.5–)59–73.5(–79.5) × (4–)5–6 μm
(av. 66.3 × 5.1 μm). Chlamydospores and sexual morph not
observed.
Culture characteristics: Colonies on PDA reaching
38–53 mm diam at 25 °C after 7 d. Surface white to
sulphur yellow with scarce pale ochreous to pale rust patches,
flat to slightly raised with abundant white aerial mycelium, cottony
to woolly, margin filiform; reverse pale luteous to sulphur yellow,
pale apricot to pale rust at centre. On OA, pale luteous, flat,
membranous with entire margin; reverse pale luteous.
Additional material examined: Netherlands, from Bouvardia sp. imported from
Uganda, 2019, W. Quaedvlieg, culture CBS 146524 = CPC 38311.
Notes: The name N. epipeda is coined here for a novel phylogenetic lineage discovered on a Bouvardia sp. imported from
Uganda. The new species clusters as the closest phylogenetic
relative of N. catenata (Fig. 14), an opportunistic animal-pathogenic species characterised by abundant production of catenate
to clustered, pigmented chlamydospores, and by the absence
(as far as known) of macroconidia (O'Donnell et al. 2016,
Sandoval-Denis & Crous 2018). These characters form the most
notable differences with respect to N. epipeda. Additionally,
N. epipeda can be differentiated from N. catenata by its less
Fig. 36. Neocosmospora epipeda (CBS 146524). A–C. Aerial conidiophores and conidiogenous cells. D. Microconidia. E, F. Sporodochia formed on the surface of carnation
leaves. G. Sporodochial conidiophores and conidiogenous cells. H. Macroconidia. Scale bars: A–C = 20 μm; E, F = 200 μm; D, G, H = 10 μm.
66
FUSARIUM
septate and shorter microconidia (aseptate and up to 13.5 μm vs
up to 1-septate and 11 μm in N. catenata). Other species producing macroconidia of similar size and shape to those of
N. epipeda include N. quercicola, N. robusta, and N. silvicola;
however, the three latter species are genetically distant in that
they belong to monophyletic lineages of clade 3 (N. quercicola
and N. silvicola) and clade 1 (N. robusta) of Neocosmospora
sensu O'Donnell et al. (2008a). Neocosmospora epipeda can be
distinguished morphologically from N. robusta by the production
of microconidia with absence of aerial macroconidia in the former
species. Morphological differentiation of the novel species from
N. quercicola and N. silvicola is difficult because of overlapping
features; nevertheless, subtle differences exist in the size and
morphology of the microconidia (aseptate in N. epipeda vs up to
1-septate in both N. quercicola and N. silvicola, being also
reniform and longer in the latter species) and sporodochial colour
(pale luteous to orange in N. epipeda vs greenish to citrine in
N. quercicola and N. silvicola, respectively).
Neocosmospora merkxiana Quaedvl. & Sand.-Den., sp. nov.
MycoBank MB 838670. Fig. 37.
Etymology: Named after Trix Merkx, senior technician at the
Westerdijk Fungal Biodiversity Institute, in recognition of her
career as the foremost link in strain handling between the
research groups and the culture collection.
Typus: Netherlands, from Chrysanthemum sp. imported from
Uganda, unknown date, W. Quaedvlieg (holotype CBS H24669, culture ex-type CBS 146525 = CPC 38701).
Conidiophores borne on the agar substrate and aerial mycelium,
99–205 μm tall, unbranched or rarely laterally branched,
bearing terminal single phialides; aerial conidiogenous cells
monophialidic, subulate to subcylindrical, smooth- and thin-walled, 41.5–77 × 2.5–4.5 μm, with short and flared apical collarettes and inconspicuous periclinal thickening. Aerial conidia of
two types: microconidia oval to broadly ellipsoidal, straight to
slightly curved and asymmetrical, smooth- and thin-walled,
0(–1)-aseptate, (8.5–)9–15.5(–18.5) × 3–5.5 μm (av. 12.4 × 4.3
ìm), arranged in false heads on phialide tips; macroconidia
falcate to navicular, smooth- and thin-walled, almost straight to
slightly dorsiventrally curved, ventral face almost straight, with a
blunt apical cell, basal cell obtuse to poorly-developed, footshaped, 1–3-septate, predominantly 1-septate, 1-septate conidia: (17.5–)20.5–27(–30.5) × (4.5–)5–6.5(–7.5) μm (av.
23.8
×
5.8
μm);
2-septate
conidia:
(25.5–)
27–30(–32) × 5.5–7 μm (av. 28.4 × 6 μm); 3-septate conidia:
(27–)28.5–33.5(–35.5) × 5–7.5 μm (av. 31.1 × 6.3 μm); overall: (17.5–)22–31(–35.5) × (4.5–)5–6.5(–7.5) μm (av.
26.4 × 6 μm), arranged in false heads at the tip of monophialides
and produced intermixed with microconidia. Sporodochia
pale luteous, formed on aerial and substrate mycelium, uncommon on carnation leaves. Sporodochial conidiophores
laterally and irregularly branched bearing apical groups of
2–3 monophialides; sporodochial conidiogenous cells monophialidic, doliiform, subulate to subcylindrical, (13.5–)
15–21.5(–27) × 2.5–5.5 μm, smooth and thin-walled, lacking
apical collarettes and with inconspicuous periclinal thickening.
Sporodochial macroconidia falcate, straight to slightly dorsiventrally curved, broadest at the half portion and tapering towards
both ends, apical cell blunt and slightly curved, basal cell poorlyto well-developed, foot-shaped, (1–)3–5-septate, predominantly
4-septate, hyaline, smooth- and thick-walled; 1-septate conidia:
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REDELIMITED
(23.5–)24.5–28.5 × 5–6.5 μm (av. 25.8 × 5.6 μm); 2-septate
conidia: 27–29 × 5.5–6.5 μm (av. 28 × 6 μm); 3-septate conidia:
(29–)35–45 × (4.5–)5–6 μm (av. 40.1 × 5.3 μm); 4-septate
conidia: (41–)44.5–49.5(–51.5) × 4.5–6.5 μm (av. 47 × 5.6 μm);
5-septate conidia: (42–)45.5–51.5(–52.5) × 5–6 μm
(av. 48.5 × 5.6 μm); overall: (24.5–)39–51.5(–52.5)
× 4.5–6(–6.5) μm (av. 45.2 × 5.6 μm). Chlamydospores obovoidal, subspherical to spherical, hyaline to pale yellow brown,
smooth-walled to slightly roughened, thick-walled, 5–13.5 μm,
single or in chains, terminal, intercalary or produced on short
lateral stipes.
Culture characteristics: Colonies on PDA reaching
45–56 mm diam at 25 °C after 7 d. Surface pale luteus to sulphur
yellow, becoming buff to honey, flat with abundant aerial mycelium,
cottony to woolly with entire to filiform margin; reverse luteous to
buff, pale scarlet to bay at centre. On OA pale luteous to peach
with sparse white cushions of aerial mycelium, flat, velvety to
cottony; reverse pale luteous, peach to pale scarlet.
Additional material examined: Netherlands, from Chrysanthemum sp. imported
from Uganda, unknown date, W. Quaedvlieg, culture CBS 146526 = CPC 38702.
Notes: Neocosmospora merkxiana represents the phylogenetic
species formerly known as “FSSC 41”, one of the few previously
known clades lacking a Latin binomial, originally reported as an
agent of collar rot on Passiflora edulis f. flavicarpa in Brazil
(Cardoso 2015, Sandoval-Denis et al. 2019). Here, this species
is reported causing collar and stem rot symptoms in Chrysanthemum imported from Uganda.
In the phylogenetic analysis (Fig. 14), N. merkxiana
resolved as the most basal taxon within a lineage containing
the morphologically similar species N. ipomoeae, N. martii,
and N. noneumartii, all characterised by producing both aerial
microconidia and macroconidia, in addition to relatively long
sporodochial conidia. Differing from the aforementioned species, N. merkxiana can be differentiated by its fewer septate
and shorter aerial and sporodochial macroconidia formed on
pale luteous sporodochia, and its pale luteous colonies on
PDA, thus contrasting with the greenish sporodochial colouration observed in both N. ipomoeae and N. noneumartii, and
the red pigmentation on PDA typical of N. martii. Sexual
morphs were not observed in the isolates studied here; however, this lineage was reported as heterothallic, and fertile
perithecial ascomata have been induced in vitro (Cardoso
characterised
by
ascomata
measuring
2015),
230–355 × 175–290 μm, 57.5–75 × 5 μm asci producing 1septate, 10–12.5 × 5 μm ascospores.
Neocosmospora neerlandica Crous & Sand.-Den., sp. nov.
MycoBank MB 838671. Fig. 38.
Etymology: Named after the country where the type was isolated,
the Netherlands.
Typus: Netherlands, Zeeland Province, Zuid-Beveland, near
Wolphaartsdijk, from Pisum sativum, unknown date, J.C. Went
(holotype CBS H-24667, culture ex-type CBS 232.34).
Conidiophores borne on agar substrate and aerial mycelium up
to 290 μm tall, unbranched or irregularly laterally branched,
bearing terminal single monophialides, commonly proliferating
percurrently; aerial conidiogenous cells monophialidic, subulate
to subcylindrical, commonly extended percurrently, smooth- and
thin-walled, 21–87 × 1.5–3.5 μm, with short and flared apical
67
CROUS
ET AL.
Fig. 37. Neocosmospora merkxiana (CBS 146525). A–E. Aerial conidiophores and conidiogenous cells. F. Sporodochium on aerial mycelium. G, H. Chlamydospores. I, J.
Sporodochial conidiophores and conidiogenous cells. K. Microconidia. L. Aerial macroconidia. M. Sporodochial macroconidia. Scale bars: A, E = 100 μm; C = 20 μm; all
others = 10 μm.
collarettes and rather evident periclinal thickening. Aerial conidia
of two types: microconidia oval to broadly ellipsoidal, smoothand thin-walled, 0- or 1-septate, (5.5–)8–14(–30) × (2–)
3–4.5(–5.5) μm (av. 11 × 3.8 μm), arranged in false heads
on phialide tips; macroconidia fusiform to falcate, smooth- and
thick-walled, straight to slightly curved, with a blunt apical
cell, basal cell often flattened to obtuse, (1–)2–3-septate, predominantly 3-septate, 1-septate conidia: 22.5–26 × 4.5–6 μm
(av. 24.4 × 5.1 μm); 2-septate conidia: (22.5–)
68
23.5–32 × 3.5–5 μm (av. 27 × 4.3 μm); 3-septate conidia: (24–)
25–32.5(–38.5) × (3.5–)4.5–5.5(–6) μm (av. 28.7 × 4.8 μm);
overall: (22.5–)24–31.5(–38.5) × (3.5–)4.5–6 μm (av.
27.7 × 4.8 μm), arranged in false heads at the tip of monophialides and produced intermixed with microconidia. Chlamydospores subspherical to spherical, pale golden brown,
smooth- and thick-walled, 6–8 μm, single or in pairs, terminal or
more often formed intercalary on hyphae. Sexual morph and
sporodochia unknown.
FUSARIUM
REDELIMITED
Fig. 38. Neocosmospora neerlandica (CBS 232.34). A–C. Conidiophores. D. Microconidia. E, F. Chlamydospores. G. Macroconidia. Scale bars: F = 5 μm; all others = 10 μm.
Culture characteristics: Colonies on PDA reaching
42–51 mm diam at 25 °C after 7 d. Surface white to pale luteous,
flat with abundant dense aerial mycelium, velvety to cottony,
margin regular and filiform; reverse pale luteous to sulphur yellow. On OA white to pale luteous, flat to slightly raised, velvety to
cottony, margin regular and filiform; reverse pale luteous.
Notes: The type of N. neerlandica was originally deposited as
N. pisi, an important root pathogen of Pisum sativum. Besides
sharing the same host association, both species are genetically
related, but cluster in distinct phylogenetic lineages and have a
different morphology. Although N. pisi produces typical wedgeshaped, larger macroconidia (up to 46 um long) on abundant
sic et al. 2018b), N. neerlandica is characterised
sporodochia (Si
by short falcate macroconidia (up to 38.5 um long) produced on
aerial conidiophores, while sporodochia are not formed. The latter
features relate N. neerlandica to N. diminuta, a phylogenetically
distant species that produces the shortest falcate conidia known in
Neocosmospora (Sandoval-Denis et al. 2019). Nevertheless,
N. diminuta is a homothallic species that conspicuously produces
sexual structures, while a sexual morph is not known for
N. neerlandica. Additionally, macroconidia of N. neerlandica differ
from those of N. diminuta by having less curved apices and poorly
developed or non foot-shaped basal cells.
Neocosmospora nelsonii Crous & Sand.-Den., sp. nov.
MycoBank MB 838672. Fig. 39.
Etymology: In honour of Paul E. Nelson, prominent Fusarium
researcher and collector of the ex-type strain of this species.
Typus: Unknown country, from Pisum sativum, unknown date,
P.E. Nelson (holotype CBS H-12719, culture ex-type CBS
309.75).
Conidiophores borne on agar substrate and aerial mycelium,
59–330 μm tall, often simple and reduced to solitary phialides
borne laterally from hyphae, or laterally irregularly and sympodially branching one or two times, bearing terminal single phialides; aerial conidiogenous cells monophialidic, subulate to
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subcylindrical, smooth- and thin-walled, 21–57.5 × 2–5 μm,
flared apical collarettes and periclinal thickening present. Aerial
microconidia arranged in false heads on phialide tips, hyaline,
broadly ellipsoidal, obovate to broadly clavate, smooth- and thinwalled, 0(–1)-septate, (5–)7–13(–17) × 2.5–5 μm (av.
10.1 × 3.7 μm). Sporodochia (from holotype specimen) pale
citrine to olivaceous; sporodochial conidiophores copiously
branched, laterally, verticillate and irregularly, bearing apical
groups of 2–3 monophialides and lateral solitary phialides;
sporodochial conidiogenous cells monophialidic, doliiform, subulate to subcylindrical, 6–21.5 × 3–4.5 μm, smooth and thinwalled, with short, conspicuously flared collarettes and conspicuous periclinal thickening, profusely proliferating percurrently. Sporodochial macroconidia falcate, gently and regularly
curved dorsoventrally or with an almost straight ventral line,
broadest at the middle portion, apical cell blunt and slightly
hooked, basal cell papillate to well-developed, foot-shaped,
1–3(–4)-septate, predominantly 3-septate, hyaline, smooth- and
thick-walled; 1-septate conidia: (17.5–)19–26(–29.5) × 4–5 μm
(av. 22.4 × 4.4 μm); 2-septate conidia: (26–)
27–34 × 3.5–5.5 μm (av. 30 × 4.7 μm); 3-septate conidia:
(25.5–)30.5–38(–42) × 4–5.5 μm (av. 34.3 × 4.8 μm); 4-septate
conidia: 38.5–43.5 × 4.5–5.5 μm (av. 40.7 × 5.0 μm); overall:
(17.5–)27–38(–43.5) × (3.5–)4–5.5 μm (av. 32.5 × 4.7 μm).
Chlamydospores subspherical to spherical, pale golden brown,
smooth- and thick-walled, 4–11.5 μm, formed singly and terminally on hyphae. Sexual morph not observed.
Culture characteristics: Colonies on PDA reaching
35–49 mm diam at 25 °C after 7 d. Surface pale luteous, pale
saffron to sulphur yellow, flat with abundant dense and short
aerial mycelium, velvety to woolly, margin filiform; reverse
sulphur yellow. On OA pale luteous, flat, membranous to dusty
with filiform margin; reverse pale luteous.
Notes: The ex-type of N. nelsonii, originally determined as “F.”
solani, currently presents a very simple microconidial
morphology with a rather acremonioid touch given its slender,
generally simple conidiophores and mostly aseptate
69
CROUS
ET AL.
Fig. 39. Neocosmospora nelsonii (CBS 309.75). A–D. Conidiophores and conidiogenous cells. E, F. Chlamydospores. G. Microconidia. H. Sporodochium. I, J. Sporodochial
conidiophores and conidiogenous cells. K. Macroconidia. Scale bars: E, F = 5 μm; all others = 10 μm.
microconidia. Hence, there are no clear phenotypic characters to
differentiate the species. Failed attempts to induce formation of
sporodochia indicate that the ex-type strain may have lost the
ability to produce macroconidia in vitro. The holotype material, is,
however, a dried subculture from the type strain dated from 1982.
It still contains a large amount of well-preserved sporodochia and
sporodochial conidia, which we describe here. These macroconidia are comparable in size to those observed in closely
related species such as N. brevis, N. pisi, and N. neerlandica.
However, macroconidia in N. brevis and N. neerlandica are
produced only in the aerial mycelium, while N. nelsonii produces
only a single type of aerial conidia (microconidia), which also
differ from those observed in the aforementioned species by their
reduced size. In addition, sporodochial conidia in N. nelsonii are
shorter and stout, with shorter and rounder apices compared to
those of N. pisi.
Neocosmospora pseudopisi Sand.-Den. & L. Lombard, sp.
nov. MycoBank MB 838673. Fig. 40.
Etymology: Named after its morphological, phylogenetic and host
affinity with Neocosmospora pisi.
Typus: Unknown country, from Pisum sativum, unknown date
and collector (holotype CBS H-24668, culture ex-type CBS
266.50).
Conidiophores borne on agar substrate and aerial
mycelium, erect and prostrate, up to 340 μm tall, unbranched or
irregularly laterally branched, bearing terminal single phialides,
rarely proliferating percurrently; aerial conidiogenous cells
monophialidic, rarely extended percurrently, subulate to subcylindrical, smooth- and thin-walled, 24.5–74 × 2–4 μm, with
cup-shaped, elongated, and flared apical collarettes and
70
conspicuous periclinal thickening. Aerial microconidia arranged
in false heads on phialide tips, hyaline, broadly ellipsoidal
to clavate, often lightly curved and asymmetrical, smooth- and
thin-walled, 0(–1)-septate, (4.5–)6.5–11(–17.5) × (2–)
3–4(–5) μm (av. 8.6 × 3.2 μm). Sporodochia pale luteous to pale
sienna coloured, rarely formed on the surface of carnation
leaves, agar surface or on aerial mycelium; sporodochial conidiophores unbranched or laterally and irregularly branched
bearing single monophialides or groups of groups of up to three
monophialides; sporodochial conidiogenous cells monophialidic,
subulate to subcylindrical, 10–25 × 2–5 μm, smooth and thinwalled, collarettes and periclinal thickening present. Sporodochial macroconidia falcate, gently tapering towards both ends,
slightly curved dorsoventrally to almost straight, apical cell blunt
to inconspicuously papillate, basal cell obtuse to poorly-developed, foot-shaped, 1–4(–5)-septate, predominantly 4-septate,
hyaline, smooth- and thick-walled; 1-septate conidia:
21.5–26(–27.5) × 4–5 μm (av. 24.7 × 4.3 μm); 2-septate conidia: 28–30 × 4.5–5 μm; 3-septate conidia: (28.5–)
34–46.5(–50) × 4–5.5 μm (av. 40.1 × 4.7 μm); 4-septate conidia: (36–)42.5–54(–56) × 4–5.5 μm (av. 48 × 4.9 μm); 5septate conidia: 50.5 × 5 μm; overall: (21.5–)
34.5–51.5(–56) × 4–5.5 μm (av. 42.9 × 4.8 μm). Chlamydospores subspherical to spherical, hyaline to pale yellow, smoothwalled, thick-walled, 5.5–10.5 μm, single or in pairs, terminal or
intercalary. Sexual morph not observed.
Culture characteristics: Colonies on PDA reaching
35–48 mm diam at 25 °C after 7 d. Surface pale luteous to pale
sulphur yellow, flat with abundant short aerial mycelium, velvety
to dusty, margin regular entire to filiform; reverse pale luteous
to sulphur yellow. On OA pale luteous to pale sulphur yellow,
FUSARIUM
REDELIMITED
Fig. 40. Neocosmospora pseudopisi (CBS 266.50). A–C. Conidiophores and conidiogenous cells. D. Microconidia. E. Sporodochia formed on aerial hyphae. F. Macroconidia
and chlamydospores. G. Macroconidia. Scale bars: C = 20 μm; E = 100 μm; all others = 10 μm.
flat, velvety to dusty, margin entire to filiform; reverse pale
luteous.
Notes: The type of N. pseudopisi was determined as pathogenic
to Pisum sativum and deposited in WI by W.C. Snyder. It is
phylogenetically and morphologically related to N. pisi, a major
sic et al. 2018b). However, both
pathogen of Pisum sativum (Si
species resolved as very closely related lineages in the sevenmarker phylogeny (Fig. 14), as well as on the individual CaM, ITS,
rpb1, and rpb2 phylogenies (data not shown). Morphologically,
N. pseudopisi can be differentiated from N. pisi by its longer
sporodochial conidia (up to 56 μm long vs up to 46 μm long in
sic et al. 2018b). Based on the features of its macroN. pisi, Si
conidia, N. pseudopisi resembles N. crassa and
N. pseudotonkinensis; the two latter species, though, are
phylogenetically well-separated. Neocosmospora pseudopisi,
however, differs from N. crassa and N. pseudotonkinensis by the
absence of aerial macroconidia in the former species, while unlike N. crassa, the sporodochial conidia of N. pseudopisi are often
wider on its apical third (vs wider at its basal part in N. crassa).
centrum. Asci cylindrical, 8-spored, without an apical ring, uniseriate. Ascospores ellipsoidal to fusoid, 1-septate, hyaline,
smooth or finely spinulose. Sporodochia not formed. Conidiophores mononematous, hyaline, septate, unbranched or
irregularly branched, terminating in 1–3 phialides or reduced to
lateral phialides. Conidiogenous cells monophialidic, cylindrical,
tapering towards the apex, with inconspicuous periclinal thickening and collarettes. Microconidia abundant, ellipsoidal to
obovoid, hyaline, aseptate, sometimes forming false heads on
phialides. Macroconidia cylindrical, mostly straight, 3–7(–9)septate, with rounded ends. Chlamydospores globose to subglobose, hyaline to subhyaline, smooth-walled to slightly verrucose, terminal or intercalary, solitary or in pairs or forming chains.
[Description adapted from Chaverri et al. (2011)].
Diagnostic features: Red, mostly smooth-walled perithecia lacking
papilla producing cylindrical asci bearing ellipsoidal to fusoid, 1septate ascospores and Cylindrocarpon asexual morph.
Nothofusarium Crous, Sand.-Den. & L. Lombard, gen. nov.
MycoBank MB 838674. Fig. 8.
Neonectria Wollenw., Ann. Mycol. 15: 52. 1917, nom. cons.
prop. Fig. 8.
Synonym: Cylindrocarpon Wollenw., Phytopathology 3: 225.
1913.
(see Chaverri et al. 2011 for additional synonyms)
Etymology: From the Greek prefix notho-, false, illegitimate; and
Fusarium, in reference to the genetic affinity and morphological
resemblance to the genus Fusarium s. str.
Type species: Neonectria ramulariae Wollenw., Ann. Mycol. 15:
52. 1917.
Ascomata
unknown.
Conidiophores
mononematous
(aerial conidiophores) or grouped on sporodochia. Aerial conidiophores simple, unbranched or irregularly branched, sometimes reduced to single lateral phialides or phialidic pegs on the
hyphae; conidiogenous cells monophialidic, cylindrical, tapering
towards apex, smooth- and thin-walled, with periclinal thickening
inconspicuous or absent, solitary. Microconidia not formed. Aerial
macroconidia falcate, 1–5(–6)-septate, thick-walled, curved to
lunate, with a blunt apical cell and often obtuse, poorly- to welldeveloped foot-shaped basal cell. Sporodochia white, pale
luteous to pale citrine. Sporodochial conidiophores irregularly
Ascomata perithecial, gregarious, seated on an erumpent
stroma, superficial, subglobose to broadly obpyriform, red,
turning dark red in KOH, pigment dissolving in lactic acid, not
collapsing when dry, with blunt to acute apex, rarely papillate,
smooth to slightly rugulose, lacking hairs or appendages.
Ascomatal wall of two regions: outer region of thick-walled,
pigmented cells forming a textura epidermoidea; inner region of
elongate, hyaline, thin-walled cells, becoming thinner toward the
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Type species: Nothofusarium devonianum L. Lombard, Crous &
Sand.-Den.
71
CROUS
ET AL.
Fig. 41. Nothofusarium devonianum (CBS 147304). A–F. Aerial conidiophores and conidiogenous cells. G–I. Sporodochia formed on the surface of carnation leaves. J–O.
Sporodochial conidiophores and conidiogenous cells. P, Q. Chlamydospores. R. Macroconidia. Scale bars: B, D = 20 μm; G, H = 200 μm; O, P = 5 μm; all others = 10 μm.
and verticillately branched, consisting of short, smooth- and thinto thick-walled stipes bearing apical whorls of mono- and polyphialides. Sporodochial conidiogenous cells monophialidic and
polyphialidic, doliiform, subulate to subcylindrical, smooth- and
thin-walled, with reduced apical collarette. Sporodochial macroconidia similar to aerial macroconidia. Chlamydospores subglobose to ellipsoidal, solitary or most commonly in chains.
Diagnostic features: Fusarioid asexual morph characterised by
aerial monophialides and sporodochial mono- and polyphialides
producing slightly curved and slender, mostly 3-septate
macroconidia.
Nothofusarium devonianum L. Lombard, Crous & Sand.-Den.,
sp. nov. MycoBank MB 838675. Fig. 41.
Etymology: The epithet refers to Devon, the English county
where the type specimen was collected.
Typus: UK, England, Devon, Totnes, Berry Pomeroy, Loventor
Manor, on dead cladodes of Ruscus aculeatus, 17 Jul. 1983,
B.C. Sutton & A.V. Sutton (holotype CBS H-24670, culture extype CBS 147304 = IMI 279297 = NRRL 22134).
Conidiophores borne on substrate mycelium, prostrate or erect
and quickly collapsing to the agar surface, 70–240 μm tall, unbranched or less commonly irregularly laterally branched,
bearing terminal single phialides; aerial conidiogenous cells
monophialidic, subulate to cylindrical, smooth- and thin-walled,
9–34 μm long, 2–5 μm at the widest part, or reduced to short
phialidic pegs, 3–6 × 2–3.5 μm, formed laterally on aerial
72
hyphae, apical collarettes short or lacking, periclinal thickening
absent. Aerial macroconidia borne on tips of conidiogenous cells
on aerial conidiophores, almost straight or slightly curved,
falcate, 1–5(–6)-septate, predominantly 3-septate, hyaline,
smooth- and thick-walled, with a blunt apical cell and obtuse,
sometimes papillate to poorly-developed, foot-shaped basal cell,
1-septate conidia: (15.5–)19–28(–32) × 2.5–4 μm (av.
23.5 × 4.3 μm); 2-septate conidia: (25.5–)27–31 × 2.5–4 μm
(av. 28.8 × 3.2 μm); 3-septate conidia: (13–)
41–57(–63.5) × 3–4(–4.5) μm (av. 49 × 3.6 μm); 4-septate
conidia: (48.5–)50–60(–61.5) × 3–4.5 μm (av. 55.1 × 3.8 μm);
5-septate conidia: (47–)50–64(–71) × 3.5–4.5 μm (av.
56.9 × 3.9 μm); 6-septate conidia: (54–)55–71.5 × 3.5–4 μm
(av. 62.3 × 3.8 μm); overall: (13–)35.5–59(–71.5) × 2.5–4.5 μm
(av. 47.2 × 3.6 μm). Sporodochia pale luteous to pale citrine
coloured, small, formed abundantly on the agar surface and less
regularly on the surface of carnation leaves; sporodochial conidiophores irregularly verticillately branched bearing solitary
lateral and terminal phialides or apical groups of 2–3 phialides;
sporodochial conidiogenous cells mono- and polyphialidic,
doliiform, subulate to subcylindrical, 3–25.5 × 2.5–5 μm, smooth
and thin-walled, commonly proliferating sympodially, collarettes
and periclinal thickening absent or inconspicuous. Sporodochial
conidia undifferentiable from aerial conidia. Chlamydospores
subglobose to ellipsoidal, solitary or most commonly in chains.
Sexual morph unknown.
Culture characteristics: Colonies on PDA reaching
23–27 mm diam at 25 °C after 7 d. Surface straw-coloured, pale
FUSARIUM
REDELIMITED
Fig. 42. Pseudofusicolla belgica. A, B. Conidiophores. C–G. Conidiogenous cells. H. Microconidia. I. Chlamydospores. J. Macroconidia. A, B, D–J. IHEM 2413. C. IHEM 5322.
Scale bars: A = 20 μm; F, G = 5 μm; all others = 10 μm.
luteous to pale ochreous, flat, dusty to velvety; reverse white to
pale luteous without diffusible pigments. On OA, grey-white to
pale luteous, flat, membranous to dusty, with irregular velvety
peripheral patches cottony; reverse pale luteous.
Notes: The type of No. devonianum was erroneously assigned to
Trichofusarium rusci (Sutton, 1986) and recombined in Fusarium
(Fusarium rusci, Geiser et al. 2013). Nevertheless, the
morphology exhibited by this strain does not match in respect
with the original description of the supposed basionym nor its
purported synonym Pycnofusarium rusci, as confirmed also by
examination of authentic material of T. rusci (BPI 453152A and
IMI 291476). The latter taxon is characterised by a setose
sporodochial asexual morph with small, fusoid, aseptate conidia,
more reminiscent of the genus Alfaria (Stachybotryaceae, Crous
et al. 2014).
Pseudofusicolla D. Triest, Mycobiology 44: 127. 2016. Figs 8, 42.
Type species: Pseudofusicolla belgica D. Triest, Mycobiology 44:
127. 2016.
Ascomata unknown. Conidiophores initially as lateral phialides
on somatic hyphae, sometimes monochasial, verticillate or
penicillate, hyaline. Conidiogenous cells monophialidic, cylindrical to subulate, hyaline, producing micro- and macroconidia.
Microconidia strongly falcate, 0- or 1-septate, hyaline. Macroconidia strongly falcate, narrowing towards the ends, apical cell
hooked with a pointed tip, basal cell papillate to poorly-developed, foot-shaped, 0–3-septate, hyaline. Chlamydospores
globose, in terminal pairs or intercalary chains.
[Description adapted from Triest et al. (2016)].
Basionym: Fusarium ventricosum Appel & Wollenw., Phytopathology 3: 32. 1913.
(See F. ventricosum in List section for synonyms)
Ascomata perithecial, mostly gregarious, non-stromatic or on a
thin stroma erumpent through the epidermis, superficial, subglobose to globose, laterally pinched when dry, dark red, with
short ostiolar neck, smooth-walled, lacking hairs and appendages. >Ascomatal wall of two regions: outer region of
thick-walled, pigmented cells forming a textura angularis or
textura globulosa; inner region of elongate, hyaline, thin-walled
cells, becoming thinner towards the centrum. Asci clavate,
apex rounded with distinct pore, 8-spored often with an apical
ring, uniseriate to biseriate. Ascospores ellipsoidal, 1-septate,
constricted at the septum, pale tan, verrucose. Sporodochia
not formed. Conidiophores simple, mononematous, straight to
flexuous, hyaline, septate, unbranched or rarely branched,
terminating in single phialides. Conidiogenous cells monophialidic, cylindrical, tapering towards the apex, with periclinal
thickening and flared collarettes, usually producing macroconidia. Microconidia rarely formed, ellipsoidal to fusoid, 0- or
1-septate, hyaline. Macroconidia falcate, straight to slightly
curved dorsiventrally, 3-septate, with blunt to slightly pointed
apical cell and poorly-developed foot-shaped basal cell.
Chlamydospores globose to subglobose to ovoid, hyaline to
subhyaline, verrucose, terminal or intercalary, solitary or in
pairs or forming chains or developing directly from
macroconidia.
[Description adapted from Booth (1971), Gerlach & Nirenberg
(1982) and Lombard et al. (2015)].
Rectifusarium L. Lombard et al., Stud. Mycol. 80: 229. 2015.
Figs 8, 43.
Diagnostic features: Dark red, smooth-walled perithecia with
short ostiolar neck producing clavate asci bearing ellipsoidal, 1septate ascospores and asexual morphs producing micro- and
macroconidia on elongate cylindrical aerial conidiophores with
monophialides, and not forming sporodochia. Chlamydospores
formed in hyphae and macroconidia.
Type species: Rectifusarium ventricosum (Appel & Wollenw.) L.
Lombard & Crous, Stud. Mycol. 80: 229. 2015.
Rugonectria P. Chaverri & Samuels, Stud. Mycol. 68: 73. 2011.
Fig. 8.
Diagnostic features: Fusarioid asexual morph that produces
strongly curved, 0- or 1-septate microconidia, and 0–3-septate
macroconidia.
www.studiesinmycology.org
73
CROUS
ET AL.
Fig. 43. Rectifusarium spp. A–F. Conidiophores and conidiogenous cells. G. Microconidia. H. Macroconidia. A–D, H. Rectifusarium robinianum (CBS 430.91). E–G. Rectifusarium ventricosum (CBS 748.79). Scale bars = 10 μm.
Type species: Rugonectria rugulosa (Pat. & Gaillard) Samuels
et al., Stud. Mycol. 68: 73. 2011.
Basionym: Nectria rugulosa Pat. & Gaillard, Bull. Soc. Mycol.
France 4: 115. 1889.
Synonyms: Cucurbitaria rugulosa (Pat. & Gaillard) Kuntze,
Revis. Gen. Pl. 3: 461. 1898.
Neonectria rugulosa (Pat. & Gaillard) Mantiri & Samuels, Canad.
J. Bot. 79: 339. 2001.
Cylindrocarpon rugulosum Brayford & Samuels, Sydowia 46:
148. 1994.
Ascomata perithecial, solitary or gregarious, stromatic, superficial or partly immersed in stroma, subglobose to globose, orange
to red, turning dark red in KOH, pigment dissolving in lactic acid,
non-papillate, rugose to tuberculate, lacking hairs or appendages. Ascomatal wall of two regions: outer region of thick-walled,
pigmented cells forming a textura angularis; inner region of
elongate, hyaline, thin-walled cells, becoming thinner towards the
centrum. Asci clavate, apex simple, 8-spored. Ascospores
ellipsoidal to oblong, 1-septate, not to slightly constricted at the
septum, pale yellow, striate. Sporodochia not formed. Conidiophores simple, mononematous, straight to flexuous, hyaline,
septate, unbranched or rarely to irregularly branched, terminating
in single phialides. Conidiogenous cells monophialidic, cylindrical, tapering towards the apex, with periclinal thickening and
flared collarettes, producing micro- and macroconidia. Microconidia ovoid to cylindrical, 0- or 1-septate, hyaline. Macroconidia fusoid, curved, (3–)5–7(–9)-septate, tapering to both
ends, basal cell obtuse with inconspicuous hilum. Chlamydospores not observed.
[Description adapted from Samuels et al. (1990), Samuels &
Brayford (1994) and Chaverri et al. (2011)].
Diagnostic features: Orange to red, rugose to tuberculate,
partially immersed perithecia producing clavate asci bearing
fusoid, 1-septate yellowish, striate ascospores and cylindrocarpon-like asexual morph characterised by curved, multiseptate macroconidia with inconspicuous hilum.
Scolecofusarium L. Lombard, Sand.-Den. & Crous, gen. nov.
MycoBank MB 838676. Figs 8, 44.
Etymology: From Greek skṓlex, worm, in reference to the wormlike appearance of the macroconidia.
Type species: Scolecofusarium ciliatum (Link) L. Lombard,
Sand.-Den. & Crous
Ascomata perithecial, solitary or gregarious, partially immersed
on a stroma, smooth- and thin-walled, globose to broadly pyriform, red, with a broad, discoid apical region, turning darker in
74
KOH, pigment dissolving in lactic acid to become yellow, lacking
hairs and warts. Ascomatal wall of a single region composed of
unevenly thickened cells of textura epidermoidea. Asci cylindrical, apex with an obscure refractive ring, 8-spored, ascospores
uniseriate. Ascospores ellipsoidal to fusiform-ellipsoidal, 1septate, not constricted at septum, yellow-brown, finely spinulose. Conidiophores mononematous (aerial) or grouped on
sporodochia. Aerial conidiophores unbranched to loosely irregularly branched, bearing terminal phialides; conidiogenous cells
monophialidic, subcylindrical, smooth- and thin-walled, with
evident periclinal thickening and a non-flared collarette, producing only macroconidia. Sporodochia pink, orange to salmon
coloured; sporodochial conidiophores irregularly and verticillately
branched, consisting of short, often swollen, smooth- and thinwalled stipes bearing single terminal monophialides or apical
whorls of 2–3 monophialides; sporodochial conidiogenous cells
monophialidic, cylindrical to subcylindrical, smooth- and thinwalled, with evident periclinal thickening. Macroconidia formed in
pink to salmon slimy masses, subcylindrical, (0–)3–7(–10)septate, straight or slightly curved, with blunt apical cell and
obtuse to poorly developed, foot-shaped basal cell. Microconidia
unknown. Chlamydospores unknown.
[Description adapted from Samuels et al. (1991) & Gerlach &
Nirenberg (1982)].
Diagnostic features: Red perithecia producing cylindrical
asci containing ellipsoidal, 1-septate, finely spinulose ascospores
and fusarioid asexual morph characterised by monophialides
producing slender and delicate, almost cylindrical macroconidia
from aerial conidiophores and pink to salmon coloured sporodochia, lacking microconidia as well as chlamydospores.
Scolecofusarium ciliatum (Link) L. Lombard, Sand.-Den. &
Crous, comb. nov. MycoBank MB 838677.
Basionym: Atractium ciliatum Link, Mag. Ges. Naturf. Freunde
Berlin 7: 32. 1816.
Synonyms: Fusarium ciliatum (Link) Link, in Willdenow, Sp. Pl.,
Ed. 4, 6: 105. 1825.
Microcera ciliata (Link) Wollenw., Fusaria Autogr. Delin. 1: 435.
1916.
Calonectria ciliata (Link) W.C. Snyder & H.N. Hansen, Amer. J.
Bot. 32: 664. 1945.
Sphaeria agnina Desm., Ann. Sci. Nat., Bot. ser. 3, 6: 72. 1846.
Calonectria agnina (Desm.) Sacc., Michelia 1: 311. 1878.
Dialonectria agnina (Desm.) Cooke, Grevillea 12: 111. 1884.
Fusarium peltigerae Westend., Herb. Crypt. Belg. 9: no. 414.
1849.
Fusarium parasiticum Westend., Bull. Seances Cl. Sci. Acad.
Roy. Sci. Belgique, ser. 2, 11: 652. 1861.
FUSARIUM
REDELIMITED
Fig. 44. Scolecofusarium ciliatum. A, B. Ascomata on natural substrate. C, D. Asci. E–G. Ascospores (G. Surface view). H. Pionnote on agar surface. I. Sporodochium. J–L.
Conidiophores and conidiogenous cells. M. Macroconidia. A–H, J–L. CBS 146674. I. CBS 146676. M. CBS 144385. Scale bars: A, B = 100 μm; E–G = 5 μm; H = 1 mm;
I = 20 μm; all others = 10 μm.
Nectria massariae Pass., in Rabenhorst, Fungi Eur. Exs. no.
1827. 1874.
Microcera massariae Sacc., Michelia 1: 263. 1878.
Calonectria massariae (Pass.) Sacc., Michelia 1: 312. 1878.
Fusisporium filisporum Cooke, Grevillea 8: 8. 1879.
Fusarium filisporum (Cooke) Sacc., Syll. Fung. 4: 708. 1886.
Fusarium scolecoides Sacc. & Ellis, Atti Reale Ist. Veneto Sci.
Lett. Arti, ser. 6, 3: 728. 1885.
Fusarium elongatum Cooke, Grevillea 19: 4. 1890.
Calonectria dearnessii Ellis & Everh., Proc. Acad. Nat. Sci.
Philadelphia 42: 245. 1891.
Typus: Germany, on branch canker of Fagus sylvatica, 1961, W.
Gerlach (neotype of Atractium ciliatum CBS H-12687 hic designatus, MBT 10000646, culture ex-neotype CBS
191.65 = ATCC 16068 = ATCC 24137 = BBA 9661 = DSM
62172 = IMI 112499 = NRRL 20431).
Additional descriptions and illustrations: Wollenweber & Reinking
(1935), Doidge (1938), Gerlach & Nirenberg (1982).
Additional material examined: Belgium, Mons, Pommeroeul, on leaf of Fagus
sylvatica, 1984, unknown collector, culture CBS 144385 = IHEM 2989. Denmark,
on Hordeum vulgare mouldy grain, associated with scale insects, 1986, U.
Thrane, culture CBS 155.86 = NRRL 22284. Netherlands, Noord-Brabant
Province, Boxmeer, on Quercus sp., Mar. 2016, S. Helleman, cultures CBS
146672 = CPC 30654; CBS 146673 = CPC 30655; CBS 146674 = CPC 30656;
CBS 146675 = CPC 30657; CBS 146676 = CPC 30658; CBS 146677 = CPC
30659.
Notes: No existent holotype material was located for At. ciliatum.
Therefore, a neotype is designated here. The neotype specimen
originates from a representative isolate studied by Gerlach &
Nirenberg (1982).
Setofusarium (Nirenberg & Samuels) Crous & Sand.-Den., gen.
et stat. nov. MycoBank MB 838678. Figs 8, 45.
Basionym: Fusarium sect. Setofusarium Nirenberg & Samuels,
Canad. J. Bot. 67: 3376. 1989.
Etymology: The name refers to the presence of setose sporodochia and to its resemblance to the genus Fusarium.
www.studiesinmycology.org
Type species: Fusarium setosum Nirenberg & Samuels, Canad.
J. Bot. 67: 3372. 1989.
Ascomata perithecial, solitary or gregarious on a well-developed
immersed stroma composed of pseudoparenchymatous to hyphal
cells, scaly to warty and thick-walled, pyriform, dark red with an
often darker red-coloured, flattened and non-papillate apical region, turning darker in KOH, pigment dissolving in lactic acid to
become yellow, lacking hairs. >Ascomatal wall of two regions:
outer region of thick-walled, pigmented cells of textura angularis to
textura globulosa at warts cells; inner region of elongate, hyaline,
thin-walled cells, becoming thinner towards the centrum. Asci
cylindrical to clavate, with rounded to flattened simple apex, 8spored, ascospores overlapping uniseriate to biseriate.
Ascospores ellipsoidal, 1-septate, not constricted at septum, pale
yellow-brown, smooth-walled to finely striate. Conidiophores
mononematous (aerial) or grouped on sporodochia. Aerial conidiophores unbranched or rarely branched, bearing terminal
phialides; conidiogenous cells monophialidic, cylindrical to subcylindrical, smooth- and thin-walled, with periclinal thickening
inconspicuous to evident, producing only macroconidia. Sporodochia grey; setae arising between and around sporodochia, stiff,
erect, thick-walled with acute tip, at first hyaline later becoming
pale golden brown; sporodochial conidiophores irregularly and
verticillately branched and densely packed, consisting of short,
often swollen, smooth- and thin-walled stipes bearing apical whorl
of 2–3 monophialides or single, terminal monophialides; sporodochial conidiogenous cells monophialidic, cylindrical to subcylindrical, smooth- and thin-walled, with inconspicuous to evident
periclinal thickening. Macroconidia formed in off-white or grey slimy
masses, cylindrical, (0–)3–5(–7)-septate, gently curved, with a
blunt apical cell and an obtuse to poorly developed foot-shaped
basal cell. Microconidia unknown. Chlamydospores unknown.
[Description adapted from Samuels & Nirenberg (1989)].
Diagnostic features: Dark red perithecia producing cylindrical to
clavate asci containing ellipsoidal, 1-septate, finely striate ascospores and fusarioid asexual morph characterised by monophialides producing robust, almost cylindrical macroconidia from
75
CROUS
ET AL.
Fig. 45. Setofusarium setosum. A–C. Ascomata on natural substrate. D. Surface view of perithecial wall in lactic acid. E. Ascus. F–H. Ascospores (H. Surface view). I, J.
Setose sporodochia. K–M. Setae. N–P. Detail of setae (N. Base. O. Middle portion wall. P. Surface view of apical wall). Q. Conidiophore. R. Macroconidia. A–H. BPI 882043.
I–R. CBS 635.92. Scale bars: A–C, I, Q = 100 μm; J–L = 20 μm; H, P = 5 μm; all others = 10 μm.
aerial conidiophores and setose sporodochia, lacking microconidia as well as chlamydospores.
Setofusarium setosum (Nirenberg & Samuels) Sand.-Den. &
Crous, comb. nov. MycoBank MB 838679.
Basionym: Fusarium setosum Nirenberg & Samuels, Canad. J.
Bot. 67: 3372. 1989.
Synonym: Nectria setofusarii Samuels & Nirenberg, Canad. J.
Bot. 67: 3372. 1989.
Typus: French Guiana, piste de Saint-Elie: km 16 on road between Sinnamary and St. Elie, ORSTOM research area
“ECEREX”, on bark of living liana, Mar. 1986, G.J. Samuels,
holotype NY00927992. Epitype of F. setosum (CBS H-24723
hic designatus, MBT 10000647): French Guiana, Vic. Cayenne,
15 km from Remise, trail to Vidal-old farm, secondary forest, from
bark, 25 Feb. 1988, A.Y. Rossman, culture ex-epitype CBS
635.92 = G.J.S. 88-12.
Description and illustrations: Samuels & Nirenberg (1989).
Additional material examined: French Guiana, unknown host and collection date,
A.Y. Rossman, culture CBS 574.94; from wood from unknown host, Feb. 1988,
A.Y. Rossman, IMI 324476. Ghana, Western Region: Wiawso District, Bia
National Park, trail from camp 1, disturbed forest, on living liana, J.G. Samuels &
H.C. Evans, BPI 882043.
Notes: The monotypic, former Fusarium section Setofusarium is
here elevated to generic rank to accommodate “Fusarium
setosum”, a genetically and morphologically divergent taxon
76
easily differentiated from any known fusarioid taxa by the production of setose sporodochia (Samuels & Nirenberg 1989). No
living ex-type culture could be located for this taxon. Isolate CBS
635.92 (as G.J.S. 88-12) is an authentic strain of Fusarium
setosum (Samuels & Nirenberg 1989). Therefore, a dried culture
from this strain is designated as epitype here.
Stylonectria Höhn., Sitzungsber. Kaiserl. Akad. Wiss. Wien,
Math.-Naturwiss. Cl., Abt. 1, 124: 52. 1915. Figs 8, 46.
Type species: Stylonectria applanata Höhn., Sitzungsber. Kaiserl.
Akad. Wiss. Wien, Math.-Naturwiss. Kl., Abt. 1, 124: 52. 1915.
Ascomata perithecial, gregarious in groups of up to 20, on a thin,
white to yellow hyphal or subiculum-like stroma, superficial,
subglobose, pyriform to subcylindrical, pale yellow, orange-red,
orange-brown, or pale to dark red, becoming dark red to purple in
KOH, with a rounded or broad, circular, flat disc on a venter-like
neck, smooth to slightly rugulose, lacking hairs or appendages.
Ascomatal wall consisting of two layers; inner layer of hyaline,
thin-walled, compressed, elongated cells and outer layer of
distinct, isodiametric to oblong, angular or globose, thick-walled
cells. Asci cylindrical to clavate, 8-spored, with simple apex or
apical ring. Ascospores cylindrical to allantoid to ellipsoidal, 1septate, hyaline or yellow to pale brown, smooth or tuberculate.
Conidiophores initially formed as unbranched phialides on somatic hyphae, sometimes loosely branched, sometimes forming
small sporodochia. Conidiogenous cells monophialidic,
FUSARIUM
REDELIMITED
Fig. 46. Stylonectria spp. A–D. Ascomata on natural substrate. E. Ascomata on culture. F. Surface view of perithecial wall in lactic acid. G, H. Asci. I–K. Ascospores. L–M.
Conidiophores and conidiogenous cells. O, P. Microconidia. Q, R. Macroconidia. A, G. Stylonectria qilianshanensis [HMAS 255803, adapted from Zeng et al. (2020)]. B.
Stylonectria norvegica [CLL14047, adapted from Lechat et al. (2015)]. C. Stylonectria purtonii (photo P. Mlcoch). D, I, K. Stylonectria wegeliana (photo B. Bergen). E, F, Q.
Stylonectria hetmanica (CBS 147306). H, J, O. Stylonectria sp. (HPC 2668). L, M. Stylonectria corniculata (CBS 125491). N, P, R. Stylonectria applanata (CBS 125489). Scale
bars: A–E = 100 μm; I–K = 5 μm; all others = 10 μm.
cylindrical to subcylindrical, with a distinct collarette. Microconidia
sparse, allantoid to lunulate, slightly or strongly curved, aseptate,
in slimy heads. Macroconidia orange in mass, subcylindrical or
moderately to strongly curved, falcate, 0- or 1-septate, apex
narrower than base, apical cell blunt or hooked, basal cell not or
scarcely foot-shaped.
[Description adapted from Höhnel (1915) and Gr€afenhan et al.
(2011)].
Diagnostic features: Pale yellow to dark red, mostly smoothwalled perithecia with rounded or broad, circular, flat disc on a
venter-like neck, producing cylindrical to clavate asci bearing
cylindrical to allantoid to ellipsoidal, 1-septate hyaline or yellow to
pale brown ascospores and fusarioid asexual morph characterised by 0- or 1-septate macroconidia with blunt or hooked
apical cell, lacking a foot-shaped basal cell.
Stylonectria corniculata Gr€afenhan, Crous & Sand.-Den., sp.
nov. MycoBank MB 838680. Fig. 47.
Etymology: From Latin corniculum, little horn. Referring to the
shape of the conidiophores.
Typus: Germany, Brandenburg, Stolpe, near Gellmersdorfer
Forst, from unidentified ascomycete on Carpinus sp., 1 Mar.
2007, T. Gr€afenhan, holotype CBS H-24671, culture ex-type
CBS 125491.
Conidiophores often as single phialides borne laterally
on substrate and aerial hyphae, or irregularly branched and
crowded with phialides produced laterally and terminally, hyaline,
thin- and smooth-walled, 24–89 μm long. Conidiogenous
cells monophialidic, short doliiform, subcylindrical to subulate,
6–28.5 × 2–3.5 μm, often with a conspicuous flared collarette,
periclinal thickening absent, producing micro- and macroconidia.
Microconidia cylindrical to allantoid, hyaline, thin- and smoothwalled, 0(–1)-septate, (4.5–)6–13.5(–21) × (1.5–)2–3 μm (av.
9.7 × 2.1 μm). Macroconidia falcate, almost straight or gently
www.studiesinmycology.org
dorsiventrally curved, tapering toward the basal portion, (0–)1septate, with a blunt apical cell and obtuse basal cell, (20–)
28–47(–56) × 2–3.5 μm (av. 37.6 × 2.5 μm). Chlamydospores
and sexual morph not observed.
Culture characteristics: Colonies on PDA reaching
16–20 mm diam at 25 °C after 7 d. Surface at first white and
membranous, becoming slimy, saffron to orange, to bright orange at the centre, flat, aerial mycelium absent, moisty at the
centre, velvety at the margin, margin regular, filiform to undulate;
reverse white, pale saffron to orange at centre. On OA, white to
pale orange, flat, membranous to slimy, with regular and undulate margin; reverse pale luteous to pale saffron.
Notes: The species is here described based on its morphology
in vitro, where only the asexual morph was obtained. This prevents further comparisons with known species of this genus. The
only known collection, CBS 125491, has been shown to represent the most basal lineage in Stylonectria in previous phylogenetic studies (Gr€afenhan et al. 2011, Lechat et al. 2015), which
was confirmed here (Fig. 15). Although with neither a clear host
association – an important character for species recognition in
Stylonectria – nor any known sexual morphology, St. corniculata
shows a distinctive morphology when it comes to its asexual
morph, especially regarding the branching pattern and the shape
of its mature conidiophores, which can be very elaborate and
largely resemble antlers (Fig. 47).
Stylonectria hetmanica Akulov, Crous & Sand.-Den., sp. nov.
MycoBank MB 838681. Fig. 48.
Etymology: The epithet refers to the Cossack Hetmanate
(Ukrainian Hetmanscyna), the name of the former Cossack state
territories where the type was collected.
Typus: Ukraine, Sumy, Okhtyrka, vicinities of Klymentove
village, Hetmanskyi National Nature Park, on the ascomata of
Diaporthe sp., associated with Phomopsis asexual morph, on
77
CROUS
ET AL.
Fig. 47. Stylonectria corniculata (CBS 125491). A–E. Conidiophores and conidiogenous cells. F. Microconidia. G. Macroconidia. Scale bars = 10 μm.
dead branches of Frangula alnus still attached to the tree, 13
Oct. 2019, Ya. Mieshkov, CWU (Myc) AS 7177, holotype CBS
H-24672, culture ex-type CBS 147305 = CPC 38725.
Ascomata perithecial, gregarious or solitary, broadly pyriform,
220–310 μm wide, with a distinctive flat and discoid papilla,
130–225 μm wide, dark red, becoming darker in 3 % KOH and
light yellow in lactic acid. Ascomatal wall smooth, 30–45 um
thick, composed of two regions: outer region 25–40 μm thick,
of irregularly shaped cells of textura intricata to textura epidermoidea; inner region 5–10 μm thick of thin-walled, flattened
cells of textura prismatica to textura angularis.
Asci subcylindrical, 45–72 × 4–8 μm, 8-spored, apices rounded
and simple, uniseriate or irregularly biseriate. Ascospores ellipsoidal, 1-septate, often constricted at septum, (7.5–)
8.5–11(–12.5) × 3–4.5(–5.5) μm, smooth to finely spinulose,
thick-walled, hyaline at first, becoming pale golden brown at
maturity. Conidiophores often as single phialides or short phialidic pegs borne laterally on the substrate and aerial hyphae,
rarely irregularly to verticillately branched. Conidiogenous cells
monophialidic, short doliiform, subcylindrical to subulate,
4–21(–27.5) × 2–3.5 μm, often with a conspicuous flared
collarette, periclinal thickening absent, producing micro- and
macroconidia. Microconidia allantoid, hyaline, smooth- and thinwalled, 0(–1)-septate, (9–)10.5–13.5(–15) × 2–3 μm (av.
12 × 2.4 μm). Macroconidia subcylindrical to falcate, almost
straight or moderately dorsiventrally curved, tapering towards
both ends, 0–1(–2)-septate, apical cell blunt to slightly hooked,
basal cell obtuse to poorly-developed, foot-shaped (11.5–)
16.5–28(–34) × 2–3 μm (av. 22.2 × 2.5 μm). Chlamydospores
not observed.
Culture characteristics: Colonies on PDA reaching
2.5–3 mm diam at 25 °C after 7 d. Surface straw-coloured to
78
luteous, pale orange at centre, flat or radially folded, membranous to slimy, margin filiform to undulate; reverse pale luteous to
pale orange. On OA orange to pale apricot, flat, membranous to
slimy, margin filiform with abundant submerged mycelium;
reverse pale orange.
Additional material examined: Ukraine, Sumy, Okhtyrka, in the vicinities of the
village Klymentove, Hetmanskyi National Nature Park, on the conidiomata of
Dothiorella sarmentorum, on recently dead branches of Acer platanoides still
attached to the tree, 13 Oct. 2019, A. Akulov, CWU (Myc) AS 7278, culture CBS
147306 = CPC 38848.
Notes: The morphological description of St. hetmanica is
based on its growth on OA, where both studied strains
showed optimal growth and sporulation. Contrary to most
fusarioid genera, St. hetmanica grows very poorly and fails
to sporulate on SNA and WA. Stylonectria hetmanica is
morphologically comparable and genetically close to St.
purtonii, St. norvegica, and St. wegeliana. Nevertheless,
ascospores of St. hetmanica are smaller than those of St.
purtonii and St. wegeliana. Additionally, macroconidia of St.
hetmanica, while similar in size to those of St. purtonii, are
less septate (0- or 1-septate, rarely 2-septate in St. hetmanica, and up to 3-septate in St. purtonii). The sexual
morph of the recently described St. norvegica is very similar
to that of St. hetmanica, although both species are genetically less closely related. The latter species can be distinguished by the production of shorter macroconidia.
Thelonectria P. Chaverri & C. Salgado, Stud. Mycol. 68: 76.
2011. Fig. 8.
Type species: Thelonectria discophora (Mont.) P. Chaverri & C.
Salgado, Stud. Mycol. 68: 76. 2011.
Basionym: Sphaeria discophora Mont., Ann. Sci. Nat. Bot., ser. 2,
3: 353. 1835.
FUSARIUM
REDELIMITED
Fig. 48. Stylonectria hetmanica (CBS 147305). A–C. Ascomata (A. On natural substrate. B, C. In culture). D. Surface view of perithecial wall in lactic acid. E–G. Ascospores
(G. Surface view). H–K. Conidiophores and conidiogenous cells. L. Macroconidia. Scale bars: A–C = 100 μm; E–I = 5 μm; all others = 10 μm.
Synonyms: Nectria discophora (Mont.) Mont., Fl. Chil. 7: 454.
1850.
Cucurbitaria discophora (Mont.) Kuntze, Revis. Gen. Pl. 3: 461.
1898.
Neonectria discophora (Mont.) Mantiri & Samuels, Canad. J. Bot.
79: 339. 2001.
Nectria tasmanica Berk., in Hooker, Bot. Antarct. Voy. III, Fl.
Tasman. 2: 279. 1860.
Cucurbitaria tasmanica (Berk.) Kuntze, Revis. Gen. Pl. 3: 462.
1898.
Nectria umbilicata Henn., Hedwigia 41: 3. 1902.
Creonectria discostiolata Chardon, Bol. Soc. Venez. Ci. Nat. 5:
341. 1939.
Ascomata perithecial, solitary to gregarious, non-stromatic or
sometimes seated on an immersed inconspicuous stroma, superficial, globose to subglobose or pyriform to elongated, orange
to red, with prominent areolate papilla or darkly pigmented apex,
smooth to slightly rugulose, lacking hairs or appendages.
Ascomatal wall of two regions: outer region of thick-walled,
pigmented cells forming a textura epidermoidea; inner region of
elongate, hyaline, thin-walled cells, becoming thinner towards the
centrum. Asci cylindrical to narrowly clavate, 8-spored, with an
apical ring, uniseriate. Ascospores ellipsoidal to fusoid, 1septate, hyaline, smooth or finely spinulose or striate. Sporodochia not formed. Conidiophores mononematous, hyaline,
septate, irregularly branched, terminating in 1–3 phialides or
reduced to lateral phialides. Conidiogenous cells monophialidic,
cylindrical or slightly swollen, tapering towards the apex, with
periclinal thickening and flared collarettes, producing usually
macroconidia. Microconidia rarely formed, globose to ovoid,
hyaline, aseptate, with displaced inconspicuous hilum. Macroconidia subcylindrical to slightly fusoid, curved, broadest at upper
third, (3–)5–7(–9)-septate, with rounded ends or flattened at the
basal cell. Chlamydospores unknown.
www.studiesinmycology.org
[Description adapted from Chaverri et al. 2011)].
Diagnostic features: Orange to red, mostly smooth-walled perithecia with prominent darkened papilla producing cylindrical to
narrowly clavate asci bearing ellipsoidal to fusoid, 1-septate
ascospores and cylindrocarpon-like asexual morph.
Tumenectria C. Salgado & Rossman, Fungal Diversity 80: 451.
2016. Fig. 8.
Type species: Tumenectria laetidisca (Rossman) C. Salgado &
Rossman, Fungal Diversity 80: 451. 2016.
Basionym: Nectria laetidisca Rossman, Mycol. Pap. 150: 36.
1983.
Synonym: Cylindrocarpon bambusicola Matsush., Matsush.
Mycol. Mem. 5: 9. 1987.
Ascomata perithecial, mostly solitary to gregarious, non-stromatic, superficial, broadly pyriform, not collapsing when dry,
orange to sienna, turning blood red in KOH, pigment dissolving in lactic acid, broadly rounded to flattened papilla,
smooth-walled, lacking hairs and appendages. Ascomatal wall
of two regions: outer region of thick-walled, pigmented cells
forming a textura angularis; inner region of elongate, hyaline,
thin-walled cells, becoming thinner towards the centrum. Asci
narrowly clavate, apex simple, 8-spored, lacking an apical
ring, irregularly multiseriate. Ascospores fusoid, 3-septate,
hyaline, smooth or finely spinulose. Sporodochia not formed.
Conidiophores simple, mononematous, straight to flexuous,
hyaline, septate, unbranched or rarely branched, terminating
in a single phialide or reduced to lateral phialides. Conidiogenous cells monophialidic, cylindrical or slightly swollen,
tapering towards the apex, with periclinal thickening and flared
collarettes. Microconidia not formed. Macroconidia cylindrical
to slightly fusoid, straight to slightly curved, 3–6-septate, with
rounded ends. Chlamydospores unknown.
79
CROUS
ET AL.
[Description adapted from Rossman (1983) and Salgado-Salazar
et al. (2016)].
Diagnostic features: Orange to sienna, smooth-walled perithecia
with broadly rounded to flattened papilla producing narrowly
clavate asci bearing fusoid, 3-septate phragmo-ascospores and
cylindrocarpon-like asexual morph.
FUSARIUM AND ALLIED GENERA: LIST OF
ACCEPTED NAMES
The following nomenclator lists names that have been introduced
in Fusarium up to January 2021, as well as their current status
(with accepted names indicated in bold and underlined for easier
recognition). Where type specimens have been located,
these details, as well as any ex-type cultures and diagnostic
DNA barcode data are provided, along with notes regarding
potential synonymy. This list will be updated and republished at
regular intervals, and will form the basis for a monograph of
Fusarium and allied genera that will be freely available on www.
Fusarium.org.
aberrans Fusarium J.W. Xia et al., Persoonia 43: 192. 2019.
Holotypus: CBS H-24050.
Ex-type culture: CBS 131385.
Type locality: Australia, Northern Territory, Roper River area.
Type substrate: Stem of Oryza australiensis.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170378; tef1: MN170445.
acaciae Fusarium Berl. & Voglino, Syll. Fung., Addit. I–IV: 201.
1886, nom. illegit., Art. 53.1.
(See Fusarium acaciae Cooke & Harkn.)
acaciae Fusarium Cooke & Harkn., Grevillea 12: 96. 1884.
Synonyms: ?Fusarium acaciae Berl. & Voglino, Syll. Fung.,
Addit. I–IV: 201. 1886, nom. illegit., Art. 53.1.
?Fusarium acaciae Sacc., Syll. Fung. 9: 958. 1891, nom. illegit.,
Art. 53.1.
(See Fusarium lateritium)
Holotypus: ?BPI 451718.
Type locality: USA, California.
Type substrate: Stem of Acacia sp.
Note: Synonym fide Wollenweber & Reinking (1935).
acaciae Fusarium Sacc., Syll. Fung. 9: 958. 1891, nom. illegit.,
Art. 53.1.
(See Fusarium acaciae Cooke & Harkn.)
acaciae-mearnsii Fusarium O'Donnell et al., Fungal Genet.
Biol. 41: 619. 2004.
Holotypus: BPI 843477.
Ex-type culture: CBS 110254 = MRC 5120 = NRRL 25754.
Type locality: South Africa, KwaZulu-Natal, Pietermaritzburg.
Type substrate: Acacia mearnsii.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: JAAWUD010000100; rpb2:
JAAWUD010000080; tef1: AF212448.
acicola Fusarium Bres., in Strasser, Verh. Zool.-Bot. Ges. Wien
60: 328. 1910.
Holotypus: Not located.
Type locality: Austria, Sonntagberg.
Type substrate: Rotting needles of Pinus sp.
80
Descriptions and illustrations: See Strasser (1910).
Note: Status unclear. Not Fusarium fide Wollenweber & Reinking
(1935).
acremoniopsis Fusarium Vincens, Bull. Soc. Mycol. France 31:
26. 1915.
(See Fusarium larvarum)
Holotypus: ?PC.
Type locality: Brazil, Para, Belem.
Type substrate: Agrotis sp. (cutworm).
Descriptions and illustrations: See Vincens (1915).
Note: Synonym fide Wollenweber & Reinking (1935).
acridiorum Fusarium (Trab.) Brongn. & Delacr., Bull. Seances
Soc. Natl. Agric. France 51: 631. 1891.
Trichothecium acridiorum (Trab.) Madelin, Trans. Brit. Mycol.
Soc. 49: 284. 1966.
Basionym: Botrytis acridiorum Trab., Compt. Rend. Hebd.
Seances Acad. Sci. 112: 1383. 1891.
Synonym: Lachnidium acridiorum Giard, Compt. Rend. Hebd.
Seances Acad. Sci. 112: 1520. 1891.
Holotypus: Not located.
Type locality: Algeria.
Type substrate: Acrididae (locust).
Description and illustrations: See Madelin (1966).
acuminatum Fusarium Ellis & Everh., Proc. Acad. Nat. Sci.
Philadelphia 47: 441. 1895.
Synonyms: Microcera acuminata (Ellis & Everh.) Höhn., in
Weese, Sitzungsber. Akad. Wiss. Wien, Math.-Naturwiss. Kl.,
Abt. 1. 128: 729. 1919.
Fusarium scirpi var. acuminatum (Ellis & Everh.) Wollenw.,
Fusaria Autogr. Delin. 3: 930. 1930.
Fusarium scirpi subsp. acuminatum (Ellis & Everh.) Raillo, Fungi
of the Genus Fusarium: 177. 1950.
Fusarium gibbosum var. acuminatum (Ellis & Everh.) Bilaĭ,
Mikrobiol. Zhurn. 49: 6. 1987.
?Selenosporium hippocastani Corda, Icon. Fung. 2: 7. 1838.
Fusarium hippocastani (Corda) Sacc., Syll. Fung. 4: 703. 1886.
Fusarium erubescens Appel & Oven, Landwirtsch. Jahrb. 1905,
nom. illegit., Art. 53.1.
Fusarium caudatum Wollenw., J. Agric. Res. 2: 262. 1914.
Fusarium scirpi var. caudatum (Wollenw.) Wollenw., Fusaria
Autogr. Delin. 3: 934 & 935. 1930.
Fusarium equiseti var. caudatum (Wollenw.) Joffe, Mycopathol.
Mycol. Appl. 53(1–4): 220. 1974.
Fusarium arcuosporum Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 186. 1915.
Fusarium ferruginosum Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 190. 1915.
Fusarium sanguineum Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 193. 1915.
Fusarium lanceolatum O.A. Pratt, J. Agric. Res. 13: 83. 1918.
Fusarium pseudoeffusum Murashk., Arb. Landwirtsch. Akad.
Omsk. 3: 19. 1924.
Fusarium moronei Curzi, Revista Biol. (Lisbon)10: 141. 1928.
Fusarium russianum Manns, Bull. N. Dakota Agric. Exp. Sta.
259: 34. 1932.
Gibberella acuminata Wollenw., Fusarien: 68. 1935.
Spermospora oryza M. Rao, Sci. & Cult. 32: 94. 1966.
Gibberella acuminata C. Booth, The Genus Fusarium: 161.
1971, nom. illegit., Art. 53.1.
Holotypus: NY00928689.
FUSARIUM
Type locality: USA, New York, Geneva.
Type substrate: Solanum tuberosum.
Descriptions and illustrations: See Sherbakoff (1915), Booth
(1971), Gerlach & Nirenberg (1982), Burgess & Summerell
(2000) and Leslie & Summerell (2006).
Notes: Fusarium acuminatum is an established name in the
Fusarium literature, but it lacks living type material to confirm its
taxonomic position. Although an older epithet, based on Selenosporium hippocastani, could be used, we refrain from
providing a new combination for this well-known species due to a
lack of DNA-based evidence to support this combination.
Moreover, Holubova-Jechova et al. (1994) could not locate any
holotype material for S. hippocastani, abstaining from introducing
a neotype, which they argued would cause nomenclatural
instability, a view we fully support.
acutatum Fusarium Nirenberg & O'Donnell, Persoonia 46: 144.
2021.
Synonym: Fusarium acutatum Nirenberg & O'Donnell, Mycologia
90: 435. 1998, nom. inval., Art. 40.1.
Holotypus: B 70 0001695.
Ex-type culture: BBA 69580 = CBS 402.97 = FRC 0-1117 = IMI
376110 = NRRL 13309.
Type locality: India.
Type substrate: Unknown.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998)
and Yilmaz et al. (2021).
Diagnostic DNA barcodes: rpb1: MT010947; rpb2: KT154005;
tef1: KR071754.
acutisporum Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 1. 2020.
Neocosmospora acutispora Sand.-Den. & Crous, Persoonia
43: 108. 2019.
Holotypus: CBS H-23969.
Ex-type culture: BBA 62213 = CBS 145461 = NRRL 22574.
Type locality: Guatemala.
Type substrate: Coffea arabica
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834210; rpb2: LR583814;
tef1: LR583593.
aderholdii Fusarium Osterw., Bericht Schweiz. Versuchsanst.
Obst-, Wein- und Gartenbau 1913/14: 519. 1915.
Ilyonectria destructans (Zinssm.) Rossman et al., Stud. Mycol.
80: 217. 2015.
Basionym: Ramularia destructans Zinssm., Phytopathology 8:
570. 1918.
Synonyms: Cylindrocarpon destructans (Zinssm.) Scholten,
Netherlands J. Plant Pathol. 70 suppl. (2): 9. 1964.
Fusarium polymorphum Marchal, Bull. Soc. Roy. Bot. Belgique
34: 145. 1895, nom. illegit., Art. 53.1.
Cylindrocarpon radicicola Wollenw., Fusaria Autogr. Delin. 2:
651. 1924.
Nectria radicicola Gerlach & L. Nilsson, Phytopathol. Z. 48: 255.
1963.
Neonectria radicicola (Gerlach & L. Nilsson) Mantiri & Samuels,
Canad. J. Bot. 79: 339. 2001.
Ilyonectria radicicola (Gerlach & L. Nilsson) P. Chaverri & C.G.
Salgado, Stud. Mycol. 68: 71. 2011.
?Fusarium rhizogenum Aderh., Centralbl. Bacteriol. Parasitenk.,
2. Abth., 6: 623. 1900, nom. illegit., Art. 53.1.
www.studiesinmycology.org
REDELIMITED
?Septocylindrium radicicola Aderh., Centralbl. Bakteriol. Parasitenk., 2. Abth., 6: 623, 1900, nom. illegit., Art. 53.1.
?Septocylindrium aderholdii Sacc & P. Syd., Syll. Fung. 16: 1048.
1902.
Holotypus: Not located.
Type locality: Germany.
Type substrate: Unknown.
Notes: Synonymy fide Wollenweber & Reinking (1935). Although
older epithets are available for Ilyonectria destructans, we refrain
from providing a new combination for this well-known species
due to a lack of DNA-based evidence to support this
combination.
adesmiae Fusarium Henn., Hedwigia 36: 246. 1897.
Synonym: Ramularia adesmiae (Henn.) Wollenw., Fusaria
Autogr. Delin. 1: 466. 1916.
Holotypus: In B fide Hein (1988).
Type locality: Chile, Bío-Bío Province.
Type substrate: Adesmia sp.
Note: Status unclear, not Ramularia fide Braun (1998).
aduncisporum Fusarium Weimer & Harter, J. Agric. Res. 32: 312.
1926.
(See Fusarium solani)
Lectotypus: BPI 451321, designated in Sandoval-Denis et al.
(2019).
Lectotype locality: USA, California, Ventura.
Lectotype substrate: Stems of Melilotus alba.
Note: Synonym fide Wollenweber & Reinking (1935).
aecidii-tussilaginis Fusarium Allesch., Ber. Bot. Vereines Landshut 12: 131. 1892.
(See Fusarium avenaceum)
Holotypus: In M.
Type locality: Germany, Oberammergau.
Type substrate: Aecidium tussilaginis.
Note: Synonym fide Wollenweber & Reinking (1935).
aeruginosum Fusarium Delacr., Bull. Soc. Mycol. France 7: 110.
1891.
(See Fusarium caeruleum)
Lectotypus (hic designatus, MBT 10000648): France, Paris, from
Solanum tuberosum, April 1891, G. Delacroix, Bull. Soc. Mycol.
France 7: pl. VIII, fig. h.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
aethiopicum Fusarium O'Donnell et al., Fungal Genet. Biol. 45:
1521. 2008.
Holotypus: BPI 878409.
Ex-type culture: CBS 122858 = NRRL 46726.
Type locality: Ethiopia, Bure district, west Gojjam zone of
Amhara region.
Type substrate: Triticum aestivum.
Descriptions and illustrations: See O'Donnell et al. (2008b).
Diagnostic DNA barcodes: rpb1: MW233298; rpb2: MW233470;
tef1: FJ240298.
affine Fusarium Fautrey & Lambotte, Rev. Mycol. (Toulouse) 18:
68. 1896.
Syntypes: ILL00221136 (Roumeguere, Fungi Sel. Gall. Exs. no.
6927) & ILL00221137 (Roumeguere, Fungi Sel. Gall. Exs. no. 6928).
Type locality: France.
Type substrate: Solanum tuberosum.
81
CROUS
ET AL.
Notes: Booth (1971) examined the exsiccatae (Fung. Sel. Gall.
Exs., No. 6927 & 6928) of F. affine and found that one part (no.
6927) is F. solani and the other part (no. 6928) represented
another fungus that was interpreted as Hymenula affinis by
Wollenweber (1916–1935). Booth (1971) indicated that F. affine
might be a possible synonym of F. tabacinum, which is now
regarded as Plectosphaerella cucumerina (Palm et al. 1995,
Giraldo & Crous 2019). However, both Gams & Gerlach (1968)
and Palm et al. (1995) considered F. affine as a misapplied
synonym of P. cucumerina. Sherbakoff (1915) also treated the
fungus as F. affine, which was later reinterpreted as Septomyxa
affine by Wollenweber (1916–1935). Therefore, the current
status of F. affine is uncertain and requires further investigation.
agapanthi Fusarium O'Donnell et al., Mycologia 108: 987.
2016.
Holotypus: VPRI 41777.
Ex-type culture: NRRL 54463 = VPRI 41777.
Type locality: Australia, Victoria, Melbourne, Royal Botanic
Gardens.
Type substrate: Agapanthus praecox.
Descriptions and illustrations: See Edwards et al. (2016).
Diagnostic DNA barcodes: rpb1: KU900620; rpb2: KU900625;
tef1: KU900630.
agaricorum Fusarium Sarrazin, Rev. Mycol. (Toulouse) 9: 170.
1887.
Lectotypus (hic designatus, MBT 10000649): France, on the cap
of Psalliota campestris (syn. Agaricus campestris), 1887, F.
Sarrazin, ILL00218415 (Roumeguere, Fungi Sel. Gall. Exs. no.
4298).
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
ailanthinum Fusarium Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 6: 350. 1899.
(See Fusarium lateritium)
Holotypus: In LPS (Fungi Argent. n.v.c. #864).
Type locality: Argentina.
Type substrate: Trunk and branches of Ailanthus glandulosa.
Note: Synonym fide Wollenweber & Reinking (1935).
alabamense Fusarium Sacc., Syll. Fung. 4: 722. 1886, nom.
illegit., Art. 52.1.
Synonym: Fusarium erubescens Berk. & M.A. Curtis, Grevillea 3:
98. 1875.
Holotypus: ?K(M).
Type locality: USA, Alabama, Beaumont.
Type substrate: Bark.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
albedinis Fusarium Kill. & Maire ex Malençon, Compt. Rend.
Hebd. Seances Acad. Sci. 198: 1261. 1934, nom. inval., Art.
6.10.
Synonym: Cylindrophora albedinis Kill. & Maire, Bull. Soc. Hist.
Nat. Afrique N. 21: 97. 1930, nom. inval., Art. 36.1.
(See Fusarium oxysporum)
Authentic material: Not located.
Original locality: Indicated as ‘oasis in Sahara’.
Original substrate: Dead trunk and leaf of Phoenix dactylifera.
Note: Synonym fide Booth (1971).
albertii Fusarium Roum., Fungi Sel. Gall. Exs., Cent. 19: no.
1867. 1881, nom. inval., Art. 38.1(a).
82
(See Fusarium lateritium)
Authentic material: BR5020140140720.
Original locality: France.
Original substrate: Petiole of Ziziphus volubilis.
Note: Synonym fide Wollenweber & Reinking (1935).
albidoviolaceum Fusarium Dasz. (as ‘albido-violaceum’), Bull.
Soc. Bot. Geneve, ser. 2, 4: 293. 1912.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000650): Switzerland,
Geneva, from soil, 1912, W. Daszewska, Bull. Soc. Bot. Geneve,
2 ser. 4: 292, fig. 15.
Notes: Wollenweber (1916–1935; Fusaria Autogr. Delin. 1: 361)
indicated that the living ex-type culture was lodged in the laboratory of W.C. Scholten in Amsterdam, which in turn has been
accessioned into the CBS. However, no record or culture can be
located in the CBS collection. Therefore, an illustration accompanying the original protologue is designated as lectotype here.
albidum Fusarium (Rossman) O'Donnell & Geiser, Phytopathology 103: 404. 2013.
Luteonectria albida (Rossman) Sand.-Den. & L. Lombard,
Stud. Mycol. 98 (no. 100116): 60. 2021.
Basionym: Nectria albida Rossman, Mycol. Pap. 150: 79. 1983.
Synonym: Albonectria albida (Rossman) Guu & Y.M. Ju, Bot.
Stud. (Taipei) 48: 189. 2007.
Holotypus: CUP-MJ 942.
Ex-type culture: ATCC 44543 = BBA 67603 = CTR 71110 = NRRL 13950 = NRRL 22152.
Type locality: Jamaica, Hanover Parish, Dolphin Head Mountain,
near Askenish.
Type substrate: Erumpent through thin bark of woody stem.
Diagnostic DNA barcode: rpb1: JX171492; rpb2: HQ897738;
tef1: MW834283.
albiziae Fusarium Woron., Vestn. Tiflissk. Bot Sada 48: 34. 1920.
(See Fusarium merismoides)
Syntypes: BPI 451733, BPI 451734 & CUP-017160.
Type locality: Georgia, Batumi, Adjara.
Type substrate: Albizia julibrissin.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification requires further investigation of the syntypes.
albocarneum Fusarium (Cooke & Harkn.) Sacc., Syll. Fung. 4:
720. 1886.
Basionym: Fusidium albocarneum Cooke & Harkn., Grevillea 9:
129. 1881.
Syntype: BPI 408577.
Type locality: USA, California, San Francisco, San Francisco
Odd Fellows Cemetery.
Type substrate: Eucalyptus sp.
Notes: The generic name Cylindrocarpon (= Neonectria;
Rossman et al. 2013) was conserved over Fusidium, making the
latter generic name a nom. rej. (Art. 14.1, 14.6 & 14.7). Therefore, Fusidium albocarneum should be transferred to Neonectria
after further investigation. Lectotypification requires further
investigation of the syntype.
albosuccineum Fusarium (Pat.) O'Donnell & Geiser, Phytopathology 103: 404. 2013.
Albonectria albosuccinea (Pat.) Rossman & Samuels, Stud.
Mycol. 42: 107. 1999.
Basionym: Calonectria albosuccinea Pat., Bull. Soc. Mycol.
France 8: 132. 1892.
FUSARIUM
REDELIMITED
Synonyms: Nectria albosuccinea (Pat.) Rossman, Mycotaxon 8:
487. 1979.
Calonectria ecuadorica Petrak, Sydowia 4: 463. 1950.
Holotypus: In FH fide Rossman (1983).
Type locality: Ecuador, Puente Chimbo.
Type substrate: Bark.
Holotypus: EMC, Palacios-Prü, 3 April 1998.
Type locality: Venezuela, Merida State, south of Sierra La
Culata, Valle de San Javier, Los Pinos.
Type substrate: Beetle immersed in kerosene.
Descriptions and illustrations: See Marcano et al. (2001).
Note: No living type material could be located.
album Fusarium Sacc., Michelia 1: 82. 1877.
Neonectria punicea (J.C. Schmidt) Castl. & Rossman, Canad.
J. Bot. 84: 1425. 2006.
Basionym: Sphaeria punicea J.C. Schmidt, in Schmidt & Kunze,
Mykol. Hefte 1: 61. 1817.
Synonyms: Nectria punicea (J.C. Schmidt) Fr., Summa Veg.
Scand. 2: 387. 1849.
Cucurbitaria punicea (J.C. Schmidt) Kuntze, Revis. Gen. Pl. 3:
461. 1898.
Cylindrocarpon album (Sacc.) Wollenw., Fusaria Autogr. Delin. 1:
no. 473. 1916.
Nectria punicea f. ilicicola Rehm, Ascomyceten: no. 337. 1876.
Nectria punicea var. ilicis C. Booth, Mycol. Pap. 73: 54. 1959.
Cylindrocarpon album var. majus Wollenw., Z. Parasitenk.
(Berlin) 1: 154. 1928.
Fusarium album var. abietinum Beeli, Bull. Soc. Roy. Bot. Belgique 62: 131. 1930.
Holotypus: Not located.
Type locality: Italy.
Type substrate: Bark of Pinus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
allescheri Fusarium Sacc. & P. Syd., Syll. Fung. 14: 1128. 1899.
Replaced synonym: Fusarium glandicola Allesch., Ber. Bot.
Vereines Landshut 12: 130. 1892, nom. illegit., Art. 53.1; non
Cooke & W.R. Gerard 1878.
(See Fusarium melanochlorum)
Holotypus: In M.
Type locality: Germany, München.
Type substrate: Quercus pedunculata.
Note: Synonym fide Wollenweber & Reinking (1935).
aleurinum Fusarium Ellis & Everh., Bull. Torrey Bot. Club 24:
476. 1897.
(See Fusarium avenaceum)
Syntypes: In BPI, BRU, CLEMS, CUP, F, FLAS, ILL, ILLS, ISC,
MICH, MSC, MU, NCU, NEB, OSC, PH, PUL, UC, WIS & WSP.
Type locality: USA, West Virginia, Fayette County Nuttallburg,
south of Edmond.
Type substrate: Wheat flour that had been on the ground for four
months.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification requires further investigation of the syntypes.
aleyrodis Fusarium Petch, Trans. Brit. Mycol. Soc. 7: 164. 1921.
Lectotypus (hic designatus, MBT 10000651): USA, Florida,
Sutherland, from Aleyrodes citri, 13 Sep. 1907, F. Wills, in Petch
1921, Trans. Brit. Mycol. Soc. 7, pl. V, fig. 12.
Notes: Wollenweber & Reinking (1935) considered this species
as a synonym of F. scirpi. However, based on the descriptions
and illustrations provided by Fawcett (1908) and Petch (1920),
this species belongs to the genus Microcera, which is also in
agreement with its aetiology. Therefore, a new combination will
presumably be required after further investigation.
algeriense Fusarium Laraba & O'Donnell, Mycologia 109: 944.
2017.
Holotypus: BPI 910347.
Ex-type culture: CBS 142638 = NRRL 66647.
Type locality: Algeria, Guelma Province, Djeballah Khemissi.
Type substrate: Triticum durum.
Descriptions and illustrations: See Laraba et al. (2017).
Diagnostic DNA barcodes: rpb1: MF120488; rpb2: MF120499;
tef1: MF120510.
alkanophilum Fusarium Palacios-Prü & V. Marcano, Rev. Ecol.
Latinoamer. 8: 5. 2001.
www.studiesinmycology.org
allescherianum Fusarium Henn., Verh. Bot. Vereins Prov.
Brandenburg 40: 175. 1899.
Synonyms: Gloeosporium allescherianum (Henn.) Wollenw.,
Fusaria Autogr. Delin. 1: 495. 1916.
?Fusarium personatum Cooke, in Harkness, Grevillea 7: 12.
1878.
Holotypus: In B fide Hein (1988).
Type locality: Germany.
Type substrate: Leaves of Ocotea foetens.
Notes: Status unclear. The taxonomic status of Gloeosporium
allescherianum is questionable. Furthermore, there is no DNAbased evidence linking F. allescherianum to F. personatum
although Wollenweber & Reinking (1935) considered them both
synonyms under G. allescherianum.
allii-sativi Fusarium Allesch., Ber. Bot. Vereines Landshut 12:
131. 1892.
(See Fusarium solani)
Holotypus: In M.
Type locality: Germany, Unterammergau.
Type substrate: Allium sativum.
alluviale Fusarium Wollenw. & Reinking, Phytopathology 15: 167.
1925.
(See Fusarium solani)
Holotypus: Not located.
Type locality: Honduras.
Type substrate: Alluvial soil.
aloes Fusarium Kalchbr. & Cooke, Grevillea 9: 23. 1880.
(See Fusarium scirpi)
Holotypus: ?K(M).
Type locality: South Africa, Eastern Cape Province, Somerset
East.
Type substrate: Aloe arborescens.
Note: Synonym fide Wollenweber & Reinking (1935).
ambrosium Fusarium (Gadd & Loos) Agnihothr. & Nirenberg,
Stud. Mycol. 32: 98. 1990.
Neocosmospora ambrosia (Gadd & Loos) L. Lombard &
Crous, Stud. Mycol. 80: 227. 2015.
Basionym: Monacrosporium ambrosium Gadd & Loos, Trans.
Brit. Mycol. Soc. 31: 17. 1947.
Synonyms: Dactylella ambrosia (Gadd & Loos) K.Q. Zhang et al.,
Mycosystema 7: 112. 1995.
Fusarium bugnicourtii Brayford, Trans. Brit. Mycol. Soc. 89: 350.
1987.
83
CROUS
ET AL.
Lectotypus: Trans. Brit. Mycol. Soc. 31: 16, Text-fig. 5. 1947,
designated by Aoki et al. (2018).
Lectotype locality: Sri Lanka.
Lectotype substrate: Gallery of Euwallacea fornicatus infesting
Camellia sinensis.
Epitypus: BPI 910524, designated by Aoki et al. (2018).
Ex-epitype culture: BBA 65390 = CBS 571.94 = NRRL
22346 = MAFF 246287.
Epitype locality: India, Upasi Tea Institute.
Epitype substrate: Gallery of Euwallacea fornicatus infesting
Camellia sinensis.
Diagnostic DNA barcodes: rpb1: KC691587; rpb2: EU329503;
tef1: FJ240350
amenti Fusarium Rostr., Bot. Tidsskr. 14: 240. 1885.
(See Fusarium avenaceum)
Holotypus: F-604398 in UPS.
Type locality: Denmark, Fyn, Holmdrup.
Type substrate: Salix cinerea.
Note: Synonym fide Wollenweber & Reinking (1935).
amentorum Fusarium Lacroix, Fl. Maine-et-Loire 2 (Suppl.): [1].
1854.
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000652): France, St.
Romain-sur-Vienne, from Salix cinerea, date unknown, J.B.H.J.
Desmazieres, BR5020140143752.
Note: Synonym fide Wollenweber & Reinking (1935).
amethysteum Fusarium P. Crouan & H. Crouan, Fl. Finistere: 14.
1867.
Holotypus: Not located.
Type locality: France.
Type substrate: Dead stem of Urtica sp.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
ampelodesmi Fusarium Fautrey & Roum., in Roumeguere, Rev.
Mycol. (Toulouse) 13: 82. 1891.
(See Fusarium reticulatum)
Syntype: ILL00219841 (Roumeguere, Fungi Sel. Gall. Exs. no.
5687).
Type locality: France, Jardin de Noidan.
Type substrate: Ampelodesmos tenax
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification requires further investigation of the syntype.
amplum Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 1. 2020.
Neocosmospora ampla Sand.-Denis & Crous, Persoonia 43:
110. 2019.
Holotypus: CBS H-23970.
Ex-type culture: BBA 4170 = CBS 202.32.
Type locality: German East Africa.
Type substrate: Coffea sp.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834212; rpb2: LR583815;
tef1: LR583594.
ananatum Fusarium A. Jacobs et al., Fung. Biol. 114: 522.
2010.
Holotypus: PREM 58713.
Ex-type culture: CBS 118516 = CMW 18685 = FCC 2986 = MRC
8165.
Type locality: South Africa, KwaZulu-Natal Province, Hluhluwe.
84
Type substrate: Ananas comosus.
Descriptions and illustrations: See Jacobs et al. (2010).
Diagnostic DNA barcodes: rpb1: MT010937; rpb2: LT996137;
tef1: MT010996.
andinum Fusarium Syd., Ann. Mycol. 37: 437. 1939.
Holotypus: S-F 45569.
Type locality: Ecuador, Tungurahua.
Type substrate: Chusquea serrulata.
Descriptions and illustrations: See Sydow & Sydow (1939).
andiyazi Fusarium Marasas et al., Mycologia 93: 1205. 2001.
Holotypus: BPI 748223.
Ex-type culture: CBS 119857 = IMI 386078 = KSU 4804 = MRC
6122.
Type locality: South Africa, KwaZulu-Natal Province, Greytown.
Type substrate: Soil debris of Sorghum bicolor.
Descriptions and illustrations: See Marasas et al. (2001) and
Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: LT996189; rpb2: LT996138;
tef1: LT996092.
andropogonis Fusarium Cooke ex Sacc., Syll. Fung. 10: 726.
1892.
Synonyms: Fusisporium andropogonis Cooke ex Thüm., Mycoth.
Univ. 7: no. 676. 1877, nom. inval., Art. 38.1(a).
Ramularia andropogonis (Cooke ex Sacc.) Wollenw., Fusaria
Autogr. Delin. 1: 469. 1916.
Lectotypus (hic designatus, MBT 10000653): USA, New Jersey,
Newfield, from dead stem of Andropogon virginicus, Oct. 1874,
J.B. Ellis, BR5020081431482 (Thümen, Mycoth. Univ. 7: no.
676).
Notes: Status unclear, not Ramularia fide Braun (1998). Synonym fide Wollenweber & Reinking (1935).
anguioides Fusarium Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 169. 1915.
Typus: ?CUP-007479.
Type locality: USA, New York, Castile.
Type substrate: Solanum tuberosum.
Descriptions and illustrations: See Sherbakoff (1915), Gerlach &
Nirenberg (1982) and Nelson et al. (1995).
Notes: Nelson et al. (1995) designated BPI 72044 as neotype of
F. anguioides, erroneously stating that no materials were available for epi- and lectotypification. However, Sherbakoff (1915)
did provide an illustration with the original protologue of
F. anguioides and placed material in CUP, as CUP-007479.
Furthermore, the neotype (BPI 72044) of Nelson et al. (1995)
originated from China and was isolated from soil in a bamboo
grove. An isolate from the original locality (USA) and host (Solanum tuberosum) needs to be selected. Lectotypification
pending study of material lodged in CUP.
angustum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 203. 1915.
(See Fusarium oxysporum)
Typus: ?CUP-007435.
Type locality: USA, New York, Ithaca.
Type substrate: Solanum tuberosum.
Descriptions and illustrations: See Sherbakoff (1915).
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in CUP.
anisophilum Fusarium Picado, J. Dept. Agric. Porto Rico 16: 391.
1932.
FUSARIUM
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Costa Rica.
Type substrate: Living stem of Coffea sp.
Note: Synonym fide Wollenweber & Reinking (1935).
annulatum Fusarium Bugnic., Rev. Gen. Bot. 59: 17. 1952.
Holotypus: IMI 202878.
Ex-type culture: BBA 63629 = CBS 258.54 = IMI 202878 = MUCL
8059 = NRRL 13619.
Type locality: New Caledonia.
Type substrate: Grain of Oryza sativa.
Descriptions and illustrations: See Bugnicourt (1952), Yilmaz
et al. (2021).
Diagnostic DNA barcodes: rpb1: MT010944; rpb2: MT010983;
tef1: MT010994.
annuum Fusarium Leonian, Bull. New Mex. Coll. Agric. Mech.
Arts 121: 9. 1919.
Lectotypus (hic designatus, MBT 10000654): USA, New Mexico,
from Capsicum annuum, 1919, L.H. Leonian, In Bull. New Mex.
Coll. Agric. Mech. Arts 121: 32, fig. 7.
Notes: No type specimen could be located. Wollenweber &
Reinking (1935) mentioned this species but did not study or
treat it any further. A new collection is required for epitypification
from the type locality and substrate.
anomalum Fusarium Berk. & M.A. Curtis, in Berkeley, Grevillea
3: 99. 1875.
Holotypus: ?K(M).
Type locality: USA, the New England region.
Type substrate: Gleditsia sp.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
anthophilum Fusarium (A. Braun) Wollenw., Ann. Mycol. 15:
14. 1917.
Basionym: Fusisporium anthophilum A. Braun, in Rabenhorst,
Fungi Eur. Exs.: no. 1964. 1875.
Synonyms: Fusarium moniliforme var. anthophilum (A. Braun)
Wollenw., Fusaria Autogr. Delin. 3: 975. 1930.
Fusarium tricinctum var. anthophilum (A. Braun) Bilaĭ, Fusarii
(Biologija I sistematika): 251. 1955.
Fusarium sporotrichiella var. anthophilum (A. Braun) Bilaĭ, Mikrobiol. Zhurn. 49: 7. 1987.
Fusarium sanguineum var. pallidius Sherb., Mem. Cornell Univ.
Agric. Exp. Sta. 6: 196. 1915.
Fusarium wollenweberi Raillo, Fungi of the Genus Fusarium:
189. 1950, nom. inval., Art. 41.1.
Lectotypus: Rabenhorst, Fungi Eur. Exs. no. 1964 in B, designated by Yilmaz et al. (2021).
Lectotype locality: Germany, Berchtesgaden.
Lectotype substrate: Succisa pratensis.
Epitypus: CBS 222.76 (preserved as metabolically inactive culture), designated by Yilmaz et al. (2021).
Ex-epitype culture: BBA 63270 = CBS 222.76 = IMI 196084 = IMI
202880 = NRRL 22943 = NRRL 25216.
Epitype locality: Germany, Berlin.
Epitype substrate: Euphorbia pulcherrima.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Nirenberg (1976), Gerlach & Nirenberg (1982), Nelson
et al. (1983) and Leslie & Summerell (2006).
www.studiesinmycology.org
REDELIMITED
Diagnostic DNA barcodes: rpb1: MW402641; rpb2: MW402811;
tef1: MW402114.
apii Fusarium P.E. Nelson & Sherb., Techn. Bull. Michigan Agric.
Exp. Sta. 155: 42. 1937.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: USA.
Type substrate: Apium graveolens var. dulce.
apiogenum Fusarium Sacc., Syll. Fung. 4: 717. 1886.
(See Fusarium lactis)
Holotypus: Not located.
Type locality: Germany.
Type substrate: Rotten fruit.
aquaeductuum Fusarium (Radlk. & Rabenh.) Lagerh. & Rabenh.,
Centralbl. Bakteriol. Parasitenk. Abth.9: 655. 1891.
Fusicolla aquaeductuum (Radlk. & Rabenh.) Gr€afenhan et al.,
Stud. Mycol. 68: 100. 2011.
Basionym: Selenosporium aquaeductuum Radlk. & Rabenh.,
Kunst- und Gewerbeblatt des Polytechnischen Vereins des
Königreichs Bayern 41(1): 10. 1863.
Synonyms: Fusisporium moschatum Kitasato, Centralbl. Bakteriol. Parasitenk., 1. Abth. 5: 368. 1889.
Fusarium moschatum (Kitasato) Sacc., Syll. Fung. 10: 729.
1892.
Fusarium magnusianum Allesch., Fungi Bav. no. 400. 1895.
Fusarium aquaeductuum var. pusillum Wollenw., Ann. Mycol. 15:
53. 1917.
Fusarium aquaeductuum var. volutum Wollenw., Ann. Mycol. 15:
53. 1917.
Fusarium aquaeductuum var. elongatum Wollenw., Fusaria
Autogr. Delin. 3: 847. 1930.
Fusarium aquaeductuum var. majus Wollenw., Fusaria Autogr.
Delin. 3: 845. 1930.
Fusarium bicellulare Kirschst., Hedwigia 80: 136. 1941.
Lectotypus: B 700014034, designated in Gr€afenhan et al. (2011).
Lectotype locality: Germany, Bayern, München, Gasteigberg.
Lectotype substrate: Water in water fountain.
Epitypus: BBA 64559, designated in Gr€afenhan et al. (2011).
Ex-epitype culture: BBA 64559 = CBS 837.85 = NRRL
20865 = NRRL 37595.
Epitype locality: Germany.
Epitype substrate: Water from plugged water tap in BBA.
Descriptions and illustrations: See Gerlach & Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: KM232250; rpb2: HQ897744;
tef1: KM231955.
arachnoideum Fusarium (Corda) Sacc., Syll. Fung. 4: 721. 1886.
Basionym: Fusisporium arachnoideum Corda, Icon. Fung. 1: 11.
1837.
(See Fusarium merismoides)
Typus: In PRM.
Type locality: Czech Republic, Prague.
Type substrate: Soil.
Note: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in PRM.
arcuatisporum Fusarium M.M. Wang et al., Persoonia 43: 78.
2019.
Holotypus: HAMS 248034.
Ex-type culture: CGMCC 3.19493 = LC 12147.
Type locality: China, Hubei.
85
CROUS
ET AL.
Type substrate: Pollen of Brassica campestris.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289799; rpb2: MK289739;
tef1: MK289584.
(1993) paper, rather than the intended basionym alone,
rendering the combination invalid (Art. 41.5, Ex. 15). Here we
validate the new combination with the correct citation of the
basionym.
arcuatum Fusarium Berk. & M.A. Curtis, Grevillea 3: 99. 1875.
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000655): USA, South Carolina, Malus pumila (syn. Pyrus malus), date unknown, M.A.
Curtis, PH00005557.
Note: Synonym fide Wollenweber & Reinking (1935).
arthrosporioides Fusarium Sherb., Mem. Cornell Univ. Agric.
Exp. Sta. 6: 175. 1915.
Typus: ?CUP-007467.
Type locality: USA, New York, Castile.
Type substrate: Solanum tuberosum.
Descriptions and illustrations: See Sherbakoff (1915), Booth
(1971) and Gerlach & Nirenberg (1982).
Notes: Synonym fide Wollenweber & Reinking (1935). Lectoand epitypification pending study of material lodged in CUP.
arcuosporum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 186. 1915.
(See Fusarium acuminatum)
Typus: ?CUP-007477.
Type locality: USA, New York, Castile.
Type substrate: Solanum tuberosum.
Descriptions and illustrations: See Sherbakoff (1915).
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in CUP.
argillaceum Fusarium (Fr.) Sacc., Syll. Fung. 4: 718. 1886.
Basionym: Fusisporium argillaceum Fr., Syst. Mycol. 3: 446.
1832.
Synonyms: Fusarium solani var. argillaceum (Fr.) Bilaĭ, Mikrobiol.
Zhurn. 49: 7. 1987.
Nectria solani Reinke & Berthold, Untersuch. Bot. Lab. Univ.
Göttingen 1: 39. 1879.
Dialonectria solani (Reinke & Berthold) Cooke, Grevillea 12: 111.
1884.
Cucurbitaria solani (Reinke & Berthold) Kuntze, Revis. Gen. Pl.
3: 461. 1898.
Holotypus: Not located.
Type locality: Unknown.
Type substrate: Periderm of Cucumis sp.
Notes: Status unclear. Requires recollection from type locality
and substrate.
aridum Fusarium O.A. Pratt, J. Agric. Res. 13: 89. 1918.
(See Fusarium sambucinum)
Lectotypus (hic designatus, MBT 10000656): USA, Idaho, from
soil, 1918, O.A. Pratt, in J. Agric. Res.13: 87, fig. 2Q.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
armeniacum Fusarium (G.A. Forbes et al.) L.W. Burgess &
Summerell, comb. nov. MycoBank MB 837636.
Basionym: Fusarium acuminatum subsp. armeniacum G.A.
Forbes et al., Mycologia 85: 120. 1993.
Holotypus: DAR 67507.
Ex-type culture: ATCC 90020 = CBS 485.94 = FRC R-9335 = IMI
352099 = MRC 6230 = NRRL 26908 = NRRL 25141 = NRRL
29133.
Type locality: Australia, New South Wales, Edgeroi.
Type substrate: Triticum aestivum.
Descriptions and illustrations: See Burgess et al. (1993), Burgess
& Summerell (2000) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: KT597715; rpb2: GQ915485;
tef1: GQ915501.
Notes: When proposing F. armeniacum, Burgess & Summerell
(2000) cited the basionym as F. acuminatum subsp. armeniacum with reference to the entire pagination of Burgess et al.'s
86
arundinis Fusarium (Corda) Sacc., Syll. Fung. 4: 724. 1886.
Trichoderma viride Pers., Neues Mag. Bot. 1: 92. 1794, nom.
sanct. [Fr., Syst. Mycol. 3: 215. 1829].
Synonyms: Pyrenium lignorum Tode, Fung. Mecklenb. Sel. 1: 33,
tab. 3, fig. 29. 1790.
Trichoderma lignorum (Tode) Harz, Bull. Soc. Imp. Naturalistes
Moscou 44: 116. 1871.
Trichoderma viride Schumach., Enum. Pl. 2: 235. 1803, nom.
illegit., Art. 53.1.
Fusisporium arundinis Corda, Icon. Fung. 1: 11. 1837.
Trichoderma glaucum E.V. Abbott, Iowa State Coll. J. Sci. 1: 27.
1927.
Lectotypus (hic designatus, MBT 10000657): Czech Republic,
Prague, rotten leaves of reeds, 1837, A.C.J. Corda, Icon. Fung.
1, tab. II, fig. 163.
Notes: Synonyms fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
arvense Fusarium Speg., Anales Soc. Ci. Argent. 10: 60. 1880.
(See Fusarium merismoides)
Holotypus: In LPS (Fungi Argent. pug. 2, #153).
Type locality: Argentina.
Type substrate: Dried fruits of Solanum elaeagnifolium.
Note: Synonym fide Wollenweber & Reinking (1935).
asclepiadeum Fusarium Fautrey, Rev. Mycol. (Toulouse) 18: 68.
1896.
(See Fusarium lateritium)
Syntype: ILL00221138 (Fungi Sel. Gall. Exs. #6929).
Type locality: France, Montagne de Bard.
Type substrate: Vincetoxicum officinale (syn. V. hirundinaria).
Note: Synonym fide Wollenweber & Reinking (1935).
asclerotium Fusarium (Sherb.) Wollenw., Fusaria Autogr. Delin.
1: 364. 1916.
Basionym: Fusarium oxysporum var. asclerotium Sherb., Mem.
Cornell Univ. Agric. Exp. Sta. 6: 222. 1915.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000658): USA, New
York, Atlanta, rotten tuber of Solanum tuberosum, 1915, C.D.
Sherbakoff, in Mem. Cornell Univ. Agric. Exp. Sta. 6: 221, fig. 35
B–J.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
asiaticum Fusarium O'Donnell et al., Fungal Genet. Biol. 41:
619. 2004.
Holotypus: BPI 843478.
FUSARIUM
REDELIMITED
Ex-type culture: CBS 110257 = FRC R-5469 = NRRL 13818.
Type locality: Japan.
Type substrate: Hordeum vulgare.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: JX171459; rpb2: JX171573;
tef1: AF212451.
Typus: No. 156060 in PRM.
Isotypus: IMI 133948 (slide).
Type locality: France.
Type substrate: Dead branch.
Note: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in PRM.
asparagi Fusarium Briard, Rev. Mycol. (Toulouse) 12: 142. 1890.
(See Fusarium incarnatum)
Holotypus: ?PC.
Type locality: France, Aube, Troyes.
Type substrate: Asparagus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
aureum Fusarium Corda, Icon. Fung. 1: 4. 1837.
Hymenella aurea (Corda) L. Lombard, comb. nov. MycoBank
MB 837637.
Basionym: Fusarium aureum Corda, Icon. Fung. 1: 4. 1837.
Synonym: Hymenula aurea (Corda) Wollenw., Fusarien: 319.
1935.
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, Prague.
Type substrate: Rotten vegetables.
Notes: Wollenweber & Reinking (1935) provided a new combination for F. aureum in the genus Hymenula. However, the generic
name Hymenella (1822) predates the generic name Hymenula
(1828) and therefore we provide a new combination in the latter
genus. Lectotypification pending study of material lodged in PRM.
asparagi Fusarium Delacr., Bull. Soc. Mycol. France 6: 99. 1890,
nom. illegit., Art. 53.1., non Fusarium asparagi Briard 1890.
Replacing synonym: Fusarium delacroixii Sacc., Syll. Fung. 10:
725. 1892.
(See Fusarium sambucinum)
Notes: Synonym fide Wollenweber & Reinking (1935). See
F. delacroixii for lectotypification.
asperifoliorum Fusarium (Westend.) Sacc., Syll. Fung. 4: 703.
1886.
Basionym: Selenosporium asperifoliorum Westend., Bull. Acad.
Roy. Sci. Belgique, ser. 2, 11: 652. 1861.
Holotypus: BR5020140146784.
Type locality: Belgium, Oudenaarde.
Type substrate: Borago officinalis.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
aspidioti Fusarium Sawada, Bot. Mag. (Tokyo) 28: 312. 1914.
(See Fusarium larvarum)
Holotypus: TNS-F-218710.
Type locality: Japan, Shizuoka.
Type substrate: Quadraspidiotus perniciosus (= Aspidiotus perniciosus) (San Jose scale).
Note: Synonym fide Wollenweber & Reinking (1935).
atrovinosum Fusarium L. Lombard & Crous, Fungal Syst. Evol.
4: 190. 2019.
Holotypus: CBS H-24015.
Ex-type culture: CBS 445.67 = BBA 10357 = DSM 62169 = IMI
096270 = NRRL 26852 = NRRL 26913.
Type locality: Australia.
Type substrate: Triticum aestivum.
Descriptions and illustrations: See Lombard et al. (2019a).
Diagnostic DNA barcodes: rpb1: MN120713; rpb2: MW928822;
tef1: MN120752.
atrovirens Fusarium (Berk.) Mussat, Syll. Fung. 15: 144. 1901,
nom. inval., Arts. 35.1, 36.1(a), (c).
Fusariella atrovirens (Berk.) Sacc., Atti Reale Ist. Veneto Sci.
Lett. Arti, ser. 6, 2: 463. 1884.
Basionym: Fusisporium atrovirens Berk., in Smith, Engl. Fl. 5 (2):
351. 1836.
Holotypus: ?K(M).
Type locality: UK, Northamptonshire, Kings Cliffe.
Type substrate: Allium sp.
Note: Synonym fide Wollenweber & Reinking (1935).
aurantiacum Fusarium Corda, in Sturm, Deutschl. Fl., 3 Abt.
(Pilze Deutschl.) 2: 19. 1829.
(See Fusarium oxysporum)
www.studiesinmycology.org
austroafricanum Fusarium A. Jacobs et al., Mycologia 110:
1197. 2018.
Holotypus: PREM 62137.
Ex-type culture: NRRL 66741 = PPRI 10408.
Type locality: South Africa, Eastern Cape Province,
Humansdorp.
Type substrate: Endophyte of Pennisetum clandestinum.
Descriptions and illustrations: See Jacobs-Venter et al. (2018).
Diagnostic DNA barcodes: rpb1: MH742537; rpb2: MH742616;
tef1: MH742687.
austroamericanum Fusarium T. Aoki et al., Fungal Genet. Biol.
41: 617. 2004.
Holotypus: BPI 843473.
Ex-type culture: CBS 110244 = NRRL 2903.
Type locality: Brazil.
Type substrate: Polypore.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: JAAMOD010000230; rpb2:
JAAMOD010000315; tef1: JAAMOD010000079.
avenaceum Fusarium (Fr.) Sacc., Syll. Fung. 4: 713. 1886.
Basionym: Fusisporium avenaceum Fr., Syst. Mycol. 2: 238.
1822, nom. sanct. [Fr., l.c.].
Synonyms: Sarcopodium avenaceum (Fr.) Fr., Summa Veg.
Scand. 2: 472. 1849.
Fusarium herbarum var. avenaceum (Fr.) Wollenw., Fusaria
Autogr. Delin. 3: 899. 1930.
Fusarium roseum var. avenaceum (Fr.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 32: 663. 1945.
Fusisporium pyrinum Fr., Syst. Mycol. 3: 445. 1832, nom. sanct.
[Fr., l.c.].
Fusarium pyrinum (Fr.) Sacc., Syll. Fung. 4: 720. 1886, nom.
illegit., Art. 53.1.
Fusarium tenue Corda, Icon. Fung. 1: 3. 1837.
Selenosporium tubercularioides Corda, Icon. Fung. 1: 7. 1837.
Fusarium tubercularioides (Corda) Sacc., Syll. Fung. 4: 697. 1886.
Fusarium herbarum var. tubercularioides (Corda) Wollenw.,
Fusaria Autogr. Delin. 3: 892. 1930.
Selenosporium herbarum Corda, Icon. Fung. 3: 34. 1839.
Fusarium herbarum (Corda) Fr., Summa Veg. Scand. 2: 472.
1849.
87
CROUS
ET AL.
Fusarium graminum var. herbarum (Corda) Wollenw., Fusaria
Autogr. Delin. 3: 941. 1930.
Fusarium avenaceum var. herbarum (Corda) Bilaĭ, Fusarii
(Biologija i sistematika): 95. 1955.
Fusarium tritici Liebman bis, Tidsskr. Landoekon., n.s., 2: 515.
1840.
Fusisporium zeae Westend., Bull. Acad. Roy. Sci. Belgique 18:
414. 1851.
Fusarium zeae (Westend.) Sacc., Syll. Fung. 4: 713. 1886.
Fusarium amentorum Lacroix, Fl. Maine-et-Loire 2 (Suppl.): [1].
1854.
Gloeosporium amentorum (Lacroix) Lind, Ann. Mycol. 3: 431. 1905.
Calogloeum amentorum (Lacroix) Nannf., Svensk Bot. Tidskr.
25: 25. 1931.
Platycarpium amentorum (Lacroix) Petr., Sydowia 7: 296. 1953.
Fusamen amentorum (Lacroix) Arx, Verh. Kon. Akad.
Wetensch., Afd. Natuurk. 51: 57. 1957.
Fusisporium incarcerans Berk., Intellectual Observ. 2: 11. 1863.
Fusarium incarcerans (Berk.) Sacc., Syll. Fung. 4: 713. 1886.
Fusarium stercoris Fuckel, Fungi Rhen. Exs., Suppl. Fasc. 5: no.
1921, 1867.
Menispora penicillata Harz, Bull. Soc. Imp. Naturalistes Moscou
44: 127. 1871.
Fusarium penicillatum (Harz) Sacc., Syll. Fung. 4: 710. 1886.
Fusisporium schiedermayeri Thüm., Fungi Austr. Exs. Cent. 1:
no. 78. 1871.
Fusarium schiedermayeri (Thüm.) Sacc., Syll. Fung. 4: 712. 1886.
Fusarium arcuatum Berk. & M.A. Curtis, Grevillea 3: 99. 1875.
Fusarium viticola Thüm., Pilze Weinst.: 52. 1878.
Fusarium herbarum var. viticola (Thüm.) Wollenw., Fusaria
Autogr. Delin. 3: 898. 1930.
Fusarium gaudefroyanum Sacc., Michelia 2: 132. 1880.
Fusisporium cucurbitariae Pat., Rev. Mycol. (Toulouse) 3: 10. 1881.
Fusarium cucurbitariae (Pat.) Sacc., Syll. Fung. 4: 708. 1886, nom.
illegit., Art. 53.1, non Fusarium cucurbitariae Peyronel 1918.
Fusarium amenti Rostr., Bot. Tidsskr. 14: 240. 1885.
Fusarium urenidicola Jul. Müll., Ber. Deutsch. Bot. Ges. 3: 395.
1885.
Fusarium diffusum Carmich., Grevillea 16: 81. 1888.
Fusarium iridis Oudem., Ned. Kruidk. Arch., ser. 2, 5: 515. 1889.
Fusarium ustilaginis Kellerm. & Swingle, Rep. (Annual) Kansas
Agric. Exp. Sta. 2: 285. 1890.
Fusarium ruberrimum Delacr., Bull. Soc. Mycol. France 6: 139.
1890.
Fusarium peckii Sacc., Syll. Fung. 10: 727. 1892, nom. illegit.,
Art. 53.1 [pro. p. fide Wollenweber & Reinking (1935)].
Fusarium aecidii-tussilaginis Allesch., Ber. Bot. Vereines Landshut 12: 131. 1892.
Fusarium subviolaceum Roum. & Fautrey, Rev. Mycol. (Toulouse) 14: 106. 1892.
Fusarium granulosum Ellis & Everh., Proc. Acad. Nat. Sci.
Philadelphia 45: 466. 1894 [1893].
Fusarium jungiae Pat., Bull. Soc. Mycol. France 11: 234. 1895.
Fusarium schnablianum Allesch., Hedwigia 34: 289. 1895.
Fusarium seemenianum Henn., Allg. Bot. Z. Syst. 2: 83. 1896.
Fusarium aleurinum Ellis & Everh., Bull. Torrey Bot. Club 24:
476. 1897.
Fusarium pseudonectria Speg., Anales Mus. Nac. Hist. Nat.
Buenos Aires 6: 351. 1899.
Fusarium limosum Rostr., Bot. Tidsskr. 22: 263. 1899.
Fusarium gracile McAlpine, Proc. Linn. Soc. New South Wales
28: 554. 1903.
88
Fusarium putrefaciens Osterw., Mitth. Thurgauischen Naturf.
Ges. 16: 123. 1904.
Fusarium paspali Henn., Bot. Jahrb. Syst. 38: 129. 1905.
Fusarium sorghi Henn., Ann. Mus. Congo Belge, Bot., Ser. 5, 2:
105. 1907.
Fusarium speiseri Lindau, Rabenh. Krypt.-Fl., ed. 2, 1(9): 580.
1909.
Fusarium palczewskii Jacz., Bull. Soc. Mycol. France 28: 345. 1912.
Fusarium pseudoheterosporum Jacz., Bull. Soc. Mycol. France
28: 347. 1912.
Fusarium metachroum Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 141. 1913.
Fusarium subulatum Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 131. 1913.
Fusarium biforme Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6:
166. 1915.
Fusarium lucidum Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6:
157. 1915.
Fusarium metachroum var. minus Sherb., Mem. Cornell Univ.
Agric. Exp. Sta. 6: 145. 1915.
Fusarium subulatum var. brevius Sherb., Mem. Cornell Univ.
Agric. Exp. Sta. 6: 149. 1915.
Fusarium truncatum Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 155. 1915.
Fusarium avenaceum var. pallens Wollenw., Fusaria. Autogr.
Delin. 2: 575. 1924.
Fusarium venerorum Dounin & Goldmacher. Actes du premier
Congres Internat. des Sylvicult.: 284–298. 1927.
Fusarium herbarum var. volutum Wollenw., Fusaria Autogr.
Delin. 3: 893. 1930.
Fusarium avenaceum var. volutum (Wollenw.) Wollenw. &
Reinking, Fusarien: 56. 1935.
Fusarium avenaceum subsp. volutum (Wollenw.) Raillo, Fungi of
the Genus Fusarium: 188. 1950.
Fusarium avenaceum var. fabae T.F. Yu, Phytopathology 34:
392. 1944.
Fusarium avenaceum f. fabae (T.F. Yu) W. Yamam., Sci. Rep.
Hyogo Univ. Agric., Ser. Agr. Biol. 2: 60. 1955.
Gibberella avenacea R.J. Cook, Phytopathology 57: 735. 1967.
Fusarium avenaceum f. fabalis X.Y. Ruan et al., Acta Phytopathol. Sin. 12: 32. 1982, nom. inval., Art. 39.1.
Fusarium avenaceum f. fabarum X.Y. Ruan et al., Acta Phytopathol. Sin. 12: 32. 1982, nom. inval., Art. 39.1.
Neotypus (hic designatus, MBT 10000659): Denmark, Hordeum
vulgare, 3 Feb. 1986, U. Thrane, CBS 408.86 (preserved as
metabolically inactive culture).
Ex-neotype culture: CBS 408.86 = FRC R-8510 = IMI
309354 = NRRL 26850 = NRRL 26911.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982), Nelson
et al. (1983) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: MG282372; rpb2: MG282401;
tef1: MW928836.
Notes: No type material could be located for this species. Therefore, to provide taxonomic stability to this important cereal-associated Fusarium species, CBS 408.86 is designated here as exneotype of Fusisporium avenaceum (= Fusarium avenaceum).
awaxy Fusarium Petters-Vandresen et al., Persoonia 43: 363.
2019.
Holotypus: UPCB93138-H.
Ex-type culture: CMRP 4013 = LGMF1930.
FUSARIUM
Type locality: Brazil, Parana, Guarapuava.
Type substrate: Rotten stalks of Zea mays.
Descriptions and illustrations: See Crous et al. (2019b).
Diagnostic DNA barcodes: rpb2: MK766941; tef1: MG839004.
aywerte Fusarium (Sangal. & L.W. Burgess) Benyon & L.W.
Burgess, Mycol. Res. 104: 1171. 2000.
Basionym: Fusarium avenaceum subsp. aywerte Sangal. & L.W.
Burgess, Mycol. Res. 99: 287. 1995.
Holotypus: DAR 69501.
Ex-type culture: F10108 = NRRL 25410.
Type locality: Australia, Northern Territory, Deep Well.
Type substrate: Soil.
Descriptions and illustrations: See Sangalang et al. (1995),
Benyon et al. (2000) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: JX171513; rpb2: JX171626;
tef1: JABCQV010000336.
azedarachinum Fusarium (Thüm.) Sacc., Syll. Fung. 4: 704. 1886.
Basionym: Fusisporium azedarachinum Thüm., Mycoth. Univ.
14: no. 1379. 1879.
(See Fusarium lateritium)
Syntypes: In BPI, CUP, ILL, NEB, NY, NYS PH & PUL (Mycotheca Universalis no. 1379).
Type locality: USA, South Carolina, Aiken.
Type substrate: Melia azedarach.
Note: Synonym fide Wollenweber & Reinking (1935).
azukiicola Fusarium T. Aoki et al. (as ‘azukicola’), Mycologia 104:
1075. 2012.
Neocosmospora phaseoli (Burkh.) L. Lombard & Crous, Stud.
Mycol. 80: 227. 2015.
Basionym: Fusarium martii f. phaseoli Burkh., Mem. Cornell Univ.
Agric. Exp. Sta. 26: 1007. 1919.
Synonyms: Fusarium solani f. phaseoli (Burkh.) W.C. Snyder &
H.N. Hansen, Amer. J. Bot. 28: 740. 1941.
Fusarium phaseoli (Burkh.) T. Aoki & O'Donnell, Mycologia 95:
671. 2003.
?Fusarium epimyces Cooke, Grevillea 17: 15. 1888.
?Fusarium pestis Sorauer, Atlas Pfl.-Krankh. 4: 19, pl. XXV. 1890.
?Fusarium martii var. viride Sherb., Mem. Cornell Univ. Agric.
Exp. Sta. 6: 247. 1915.
Fusarium solani var. martii Appel & Wollenw. f. 3 Snyder, Centralbl. Bakteriol. Parasitenk., 2. Abth. 91: 179. 1934.
Fusarium solani f. sp. glycines K. Roy, Pl. Dis. 81: 264. 1997.
Fusarium tucumaniae T. Aoki et al., Mycologia 95: 664. 2003.
Neocosmospora tucumaniae (T. Aoki et al.) L. Lombard & Crous,
Stud. Mycol. 80: 228. 2015.
Fusarium virguliforme O'Donnell & T. Aoki, Mycologia 95: 667.
2003.
Neocosmospora virguliformis (O'Donnell & T. Aoki) L. Lombard &
Crous, Stud. Mycol. 80: 228. 2015.
Fusarium brasiliense T. Aoki & O'Donnell, Mycoscience 46: 166.
2005.
Fusarium cuneirostrum O'Donnell & T. Aoki, Mycoscience 46:
170. 2005.
Fusarium crassistipitatum Scandiani et al., Mycoscience 53: 171.
2011.
Holotypus: BPI 881712.
Ex-type culture: MAFF 242371 = NRRL 54364.
Type locality: Japan, Hokkaido, Tokachi, Urahoro.
Type substrate: Roots of Vigna angularis.
Descriptions and illustrations: See Aoki et al. (2012b).
www.studiesinmycology.org
REDELIMITED
Diagnostic DNA barcodes: rpb1: KJ511276; rpb2: KJ511287;
tef1: JQ670137.
babinda Fusarium Summerell et al., Mycol. Res. 99: 1345. 1995.
Holotypus: DAR 70287.
Ex-type culture: BBA 69872 = F11217 = NRRL 25807.
Type locality: Australia, Queensland, Mount Lewis.
Type substrate: Plant material in soil.
Descriptions and illustrations: See Summerell et al. (1995) and
Leslie & Summerell (2006).
Diagnostic DNA barcode: rpb2: MN534245; tef1: AF160305.
Note: The Fusarium babinda species complex encompassed
strains incorrectly assigned to this taxon, based on reference
strains of F. babinda, plus one unnamed Fusarium species
(O'Donnell et al. 2013, Jacobs-Venter et al. 2019, Geiser et al.
2021). However, DNA sequences from diverse gene regions
and phylogenetic analyses made by several authors place the
ex-type of F. babinda (NRRL 25807) within the Fusarium fujikuroi
species complex, as confirmed here (Fig. 8) (O'Donnell et al.
2000b, Lima et al. 2012, Herron et al. 2015, Crous et al.
2019b). Hence, the species in FBSC need to be reassessed
and the species complex renamed accordingly.
baccharidicola Fusarium Henn., Hedwigia 48: 20. 1908.
(See Fusarium coccophilum)
Syntype: Puttemans no. 1274 in B (syntype fide Hein (1988).
Type locality: Brazil, S~ao Paulo, Pirutuba.
Type substrate: Baccharis dracunculifolia in association with
cochineal (Dactylopius coccus)
Note: Synonym fide Wollenweber & Reinking (1935).
bacilligerum Fusarium (Berk. & Broome) Sacc., Syll. Fung. 4:
711. 1886.
Pseudocercospora bacilligera (Berk. & Broome) Y.L. Guo &
X.J. Liu, Mycosystema 2: 229. 1989.
Basionym: Fusisporium bacilligerum Berk. & Broome, Ann. Mag.
Nat. Hist., ser. 2, 7: 178. 1851.
Synonyms: Cercospora bacilligera (Berk. & Broome) Wollenw.,
Fusaria Autogr. Delin. 1: 450. 1916.
Fusisporium erubescens Durieu & Mont., Exploration scientifique
de l'Algerie 1–9: 351.1848.
Fusarium erubescens (Durieu & Mont.) Sacc., Syll. Fung. 4: 719.
1886, nom. illegit., Art. 53.1.
Holotypus: ?K(M).
Type locality: UK, Wiltshire, Spye Park.
Type substrate: Leaves of Rhamnus alaternus.
Note: Synonyms fide Wollenweber & Reinking (1935).
bactridioides Fusarium Wollenw., Science, N.Y. 79: 572. 1934.
Lectotypus: NY00936830, designated in Seifert & Gr€afenhan
(2012).
Ex-type culture: BBA 4748 = BBA 63602 = CBS 100057 = CBS
177.35 = DAOM 225115 = IMI 375323 = NRRL 22201.
Type locality: USA, Arizona, Chiricahua Mountains.
Type substrate: Parasitic on Cronartium conigenum growing on a
mummified cone of Pinus leiophylla.
Descriptions and illustrations: See Wollenweber (1934), Gerlach
& Nirenberg (1982) and Seifert & Gr€afenhan (2012).
Diagnostic DNA barcodes: rpb1: MT010939; rpb2: MT010963;
tef1: KC514053.
bagnisianum Fusarium Thüm., Nuovo Giorn. Bot. Ital. 8: 252. 1876.
Ascochyta caricis Fuckel, Fungi Rhen. Suppl. Fasc. 2: no.
1697. 1866.
89
CROUS
ET AL.
Synonyms: Phyllosticta caricis (Fuckel) Sacc., Syll. Fung. 3: 61.
1884.
Ascochyta caricis Lambotte & Fautrey, Rev. Mycol. (Toulouse)
19: 141. 1897, nom. illegit., Art. 53.1.
Syntypes: In BPI, ILL, NEB, NY, PUL & S.
Type locality: Italy, Rome, Insugherata.
Type substrate: Spartium junceum.
Note: Synonym fide Wollenweber & Reinking (1935).
bambusae Fusarium (Teng) Z.Q. Zeng & W.Y. Zhuang,
Mycosystema 36: 279. 2017.
Basionym: Lisea australis var. bambusae Teng, Sinensia 4: 278.
1934.
Synonym: Gibberella bambusae (Teng) W.Y. Zhuang & X.M.
Zhang, Nova Hedwigia 76: 195. 2003.
Holotypus: BPI 631179.
Type locality: China, Anhui, Huang-shan.
Type substrate: Bambusoideae culm.
Descriptions and illustrations: See Zhang & Zhuang (2003) and
Zeng & Zhuang (2017a).
bambusicola Fusarium Hara, Bot. Mag. (Tokyo) 27: 255. 1913.
Holotypus: Not located.
Type locality: Japan, Tokyo.
Type substrate: Phyllostachys reticulata.
Note: Type material (specimen(s) and/or living ex-type culture)
not located.
baptisiae Fusarium Henn., Notizbl. Bot. Gart. Berlin 2: 383. 1899.
(See Fusarium dimerum)
Holotypus: In B fide Hein (1988).
Type locality: Germany, Berlin, Botanical Garden.
Type substrate: Baptisia tinctoria.
Note: Synonym fide Wollenweber & Reinking (1935).
barbatum Fusarium Ellis & Everh., J. Mycol. 4: 45. 1888.
Raffaelea barbata (Ellis & Everh.) D. Hawksw. (as ‘barbatum'’),
Bull. Brit. Mus. (Nat. Hist.), Bot. 6: 272. 1979.
Holotypus: NY00928690.
Type locality: USA, New Jersey, Newfield.
Type substrate: Usnea barbata.
bartholomaei Fusarium Peck, Bull. Torrey Bot. Club 36: 157.
1909.
Septogloeum bartholomaei (Peck) Wollenw., Fusaria Autogr.
Delin. 2: 638. 1924.
Synonym: Trichofusarium bartholomaei (Peck) Sacc., Syll. Fung.
22: 1473. 1913.
Holotypus: NYS-F-000437.
Type locality: USA, Kansas, Stockton.
Type substrate: Sorghastrum nutans.
Note: Synonym fide Wollenweber & Reinking (1935).
batatas Fusarium Wollenw. (as ‘batatae’), J. Agric. Res. 2: 268.
1914.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000660): USA, Washington,
Ipomoea batatas, 1914, L.L. Harter & E.C. Field, in Wollenweber,
J. Agric. Res. 2: 268, pl. XVI, figs A–E.
Notes: Synonym fide Wollenweber & Reinking (1935). As
no holotype specimen could be located, an illustration accompanying the original protologue is designated here as lectotype.
bataticola Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 1. 2020.
90
Neocosmospora bataticola Sand.-Den. & Crous, Persoonia 43:
112. 2019.
Synonym: ?Fusarium solani f. batatas T.T. McClure, Phytopathology 41: 75. 1951, nom. inval., Art. 39.1.
Holotypus: CBS H-23971.
Ex-type culture: BBA 64954 = CBS 144398 = FRC S0567 = NRRL 22402.
Type locality: USA, North Carolina.
Type substrate: Ipomoea batatas.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218100; rpb2: FJ240381;
tef1: AF178344.
begoniae Fusarium Nirenberg & O'Donnell, Mycologia 90: 437.
1998.
Holotypus: B 70 0001694.
Ex-type culture: BBA 67781 = CBS 403.97 = IMI 375315 = NRRL
25300.
Type locality: Germany.
Type substrate: Begonia elatior.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998)
and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: JAAOAG010000375; rpb2:
MN193886; tef1: AF160293.
beomiforme Fusarium P.E. Nelson et al., Mycologia 79: 886.
1987.
Holotypus: DAOM 196987.
Ex-type culture: CBS 100160 = DAR 58880 = FRC M-1425 = IMI
316127 = MRC 4593 = NRRL 13606.
Type locality: Australia, Queensland, Rockhampton.
Type substrate: Soil.
Descriptions and illustrations: See Nelson et al. (1987) and Leslie
& Summerell (2006).
Diagnostic DNA barcodes: rpb1: MF120485; rpb2: MF120496;
tef1: MF120507.
berenice Fusarium (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
721. 1886.
Ascocalyx berenice (Berk. & M.A. Curtis) Baschien, IMA
Fungus 5: 93. 2014.
Basionym: Fusisporium berenice Berk. & M.A. Curtis, in Berkeley, Grevillea 3: 147. 1875.
Synonyms: Bothrodiscus berenice (Berk. & M.A. Curtis) J.W.
Groves, Canad. J. Bot. 46: 1273. 1968.
Holotypus: ?K(M).
Type locality: USA, Massachusetts, Boston, Murray.
Type substrate: Peziza sp.
berkeleyi Fusarium (Mont.) Berk. & Broome, North Amer. Fung.:
108. 1875.
Basionym: Gloeosporium berkeleyi Mont., Ann. Sci. Nat., Bot.,
ser. 3, 12: 296. 1849.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: USA, Alabama.
Type substrate: Leaves of Hibiscus syriacus.
Note: Synonym fide Wollenweber & Reinking (1935).
betae Fusarium (Desm.) Sacc., Michelia 2: 132. 1880.
Fusicolla betae (Desm.) Bonord., Handb. Mykol.: 150. 1851.
Basionym: Fusisporium betae Desm., Ann. Sci. Nat., Bot., Ser. 1,
19: 436. 1830.
FUSARIUM
Synonyms: Pionnotes betae (Desm.) Sacc., Syll. Fung. 4: 726.
1886.
Pionnotes rhizophila var. betae (Desm.) De Wild. & Durieu,
Prodr. Fl. Belg. 2: 367. 1898.
Lectotypus: K(M) 167520, designated in Gr€afenhan et al. 2011.
Lectotype locality: France.
Lectotype substrate: Tuber of Beta vulgaris.
Epitypus: BBA 64317, designated in Gr€afenhan et al. 2011.
Ex-epitype culture: BBA 64317.
Epitype locality: Germany, Schleswig-Holstein, Kiel.
Epitype substrate: Triticum aestivum.
Descriptions and illustrations: See Gr€afenhan et al. (2011).
Diagnostic DNA barcodes: rpb2: HQ897781.
beticola Fusarium A.B. Frank, Kampfbuch gegen die Sch€adlinge
unserer Feldfrüchte: 137. 1897.
(See Fusarium oxysporum)
Holotypus: ?NY.
Type locality: Germany.
Type substrate: Beta sp.
Note: Synonym fide Wollenweber & Reinking (1935).
biasolettianum Fusarium Corda, Icon. Fung. 2: 3. 1838.
(See Fusarium merismoides)
Typus: PRM 155487.
Type locality: Czech Republic, near Prague.
Type substrate: Young stalks of Betula sp.
Notes: Synonym fide Wollenweber & Reinking (1935). Synonymy under Fusicolla merismoides still questionable (See
Gr€afenhan et al. 2011). Lectotypification pending study of material lodged in PRM.
bicellulare Fusarium Kirschst., Hedwigia 80: 136. 1941.
(See Fusarium aquaeductuum)
Holotypus: B 70 0100184.
Type locality: Germany.
Type substrate: Parasitic on Cryptosporella hypodermia with
Nectria episphaeria.
Note: Synonym fide Wollenweber & Reinking (1935).
biforme Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6:
166. 1915.
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000661): USA, Wisconsin,
rotten tubers of Solanum tuberosum, date unknown, C.D.
Sherbakoff, in Mem. Cornell Univ. Agric. Exp. Sta. 6: 166, fig. 17
(1915).
Notes: Synonym fide Wollenweber & Reinking (1935). As no
holotype specimen could be located, an illustration accompanying the original protologue is designated here as lectotype.
bipunctatum Fusarium Preuss, Linnaea 25: 741. 1852.
(See Fusarium tortuosum)
Holotypus: ?B.
Type locality: Germany.
Type substrate: Branches of unknown tree.
Note: Synonym fide Wollenweber & Reinking (1935).
biseptatum Fusarium Sawada, Special Publ. Coll. Agric. Natl.
Taiwan Univ. 8: 228. 1959, nom. inval., Art. 39.1.
Authentic material: Not located.
Original locality: Taiwan.
Original substrate: Leaves of Stephania cepharantha.
www.studiesinmycology.org
REDELIMITED
biseptatum Fusarium Schroers et al., Mycologia 101: 59. 2009.
(non Fusarium biseptatum Sawada 1959).
Bisifusarium biseptatum (Schroers et al.) L. Lombard & Crous,
Stud. Mycol. 80: 224. 2015.
Holotypus: CBS H-20126.
Ex-type culture: CBS 110311 = FRC E-0228 = NRRL 36184.
Type locality: South Africa, Transkei.
Type substrate: Soil.
Descriptions and illustrations: See Schroers et al. (2009).
Diagnostic DNA barcode: tef1: EU926319.
blackmannii Fusarium W. Br. & A.S. Horne (as ‘blackmanni'’),
Ann. Bot. (London) 38: 379. 1924.
(See Fusarium lateritium)
Notes: Name withdrawn by original author (W. Brown),
See Brown (1928). Synonym fide Wollenweber & Reinking
(1935).
blasticola Fusarium Rostr. (as ‘blasticolum’), Gartn.-Tidende
1895: 122. 1895.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: Germany.
Type substrate: Pinus sylvestris.
Note: Synonym fide Wollenweber & Reinking (1935).
bomiense Fusarium (Z.Q. Zeng & W.Y. Zhuang) O'Donnell et al.,
Index Fungorum 440: 1. 2020.
Neocosmospora bomiensis Z.Q. Zeng & W.Y. Zhuang, Phytotaxa 319: 177. 2017.
Holotypus: HMAS 254519.
Ex-type culture: HMAS 248885.
Type locality: China, Tibet Autonomous Region, Bom^e County.
Type substrate: Twigs.
Descriptions and illustrations: See Zeng & Zhuang (2017b).
Diagnostic DNA barcode: tef1: KY829449.
bonordenii Fusarium Sacc., Syll. Fung. 4: 699. 1886.
Replaced synonym: Selenosporium aurantiacum Bonord., Abh.
Naturf. Ges. Halle 8: 97. 1864, nom. illegit., Art. 53.1, non
Fusarium aurantiacum Corda 1829.
(See Fusarium dimerum)
Holotypus: Not preserved fide Holubova-Jechova et al. (1994).
Type locality: Germany.
Type substrate: Branches of unknown tree.
Note: Synonym fide Wollenweber & Reinking (1935).
boothii Fusarium O'Donnell et al., Fungal Genet. Biol. 41: 618.
2004.
Holotypus: BPI 843475.
Ex-type culture: CBS 316.73 = IMI 160243 = NRRL 26916.
Type locality: South Africa.
Type substrate: Zea mays.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: KM361641; rpb2: KM361659;
tef1: GQ915503.
borneense Fusarium (Petr.) O'Donnell et al., Index Fungorum
440: 1. 2020.
Neocosmospora borneensis (Petr.) Sand.-Den. & Crous,
Persoonia 43: 115. 2019.
Basionym: Nectria borneensis Petr., Sydowia 8: 20. 1954.
Holotypus: K(M) 252860.
Epitypus: CBS H-23972, designated in Sandoval-Denis et al. (2019).
91
CROUS
ET AL.
Ex-epitype culture: BBA 65095 = CBS 145462 = G.J.S. 85197 = NRRL 22579.
Epitype locality: Indonesia, North Sulawesi, Bogani Nani Wartabone National Park.
Epitype substrate: Bark of a recently dead unidentified tree.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834213; rpb2: FJ240381;
tef1: AF178344.
bostrycoides Fusarium Wollenw. & Reinking, Phytopathology 15:
166. 1925.
Neocosmospora bostrycoides (Wollenw. & Reinking) Sand.Den. et al., Persoonia 43: 115. 2019.
Neotypus: CBS H-23973, designated in Sandoval-Denis et al.
(2019).
Ex-neotype culture: CBS 144.25.
Neotype locality: Honduras, Tela.
Neotype substrate: Soil.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218101; rpb2: LR583818;
tef1: LR583597.
brachiariae Fusarium M.M. Costa et al., Mycol. Progr. 20: 67.
2021.
Holotypus: UB 24188.
Ex-type culture: CML 3032.
Type locality: Brazil, Mato Grosso do Sul, Campo Grande.
Type substrate: Seed of Brachiaria decumbens.
Descriptions and illustrations: See Costa et al. (2021).
Diagnostic DNA barcodes: rpb2: MT901314; tef1: MT901348.
brachygibbosum Fusarium Padwick, Mycol. Pap. 12: 11. 1945.
Holotypus: IMI 268019.
Ex-type culture: BBA 64691 = NRRL 20954.
Type locality: India, Telangana, Hyderabad, Parbhani.
Type substrate: Sorghum vulgare.
Descriptions and illustrations: See Padwick (1945).
Diagnostic DNA barcodes: rpb1: MW233246; rpb2: MW233418;
tef1: MW233075.
brasilicum Fusarium T. Aoki et al., Fungal Genet. Biol. 41: 620.
2004.
Holotypus: BPI 843480.
Ex-type culture: CBS 119180 = NRRL 31281.
Type locality: Brazil.
Type substrate: Avena sativa.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: JABCJS010000032; rpb2:
JABCJS010000357; tef1: AY452964.
brasiliense Fusarium T. Aoki & O'Donnell, Mycoscience 46: 166.
2005.
(See Fusarium azukiicola)
Holotypus: BPI 843352.
Ex-type culture: MAFF 239050 = NRRL 31757.
Type locality: Brazil, Districto Federal, Brasilia.
Type substrate: Glycines max.
Descriptions and illustrations: See Aoki et al. (2005).
Diagnostic DNA barcodes: rpb1: MAEC01003448; rpb2:
EU329565; tef1: MAEC01004196.
brassicae Fusarium Lib. ex Cooke, Grevillea 8: 83. 1880.
(See Fusarium candidum Ehrenb.)
Holotypus: In B, Libert s.n. fide Index Fungorum.
92
Type locality: France.
Type substrate: Stem of Brassica oleracea.
Note: Synonym fide Wollenweber & Reinking (1935).
brassicae Fusarium (Thüm.) Sacc., Syll. Fung. 4: 701. 1886,
nom. illegit., Art. 53.1.
Basionym: Selenosporium brassicae Thüm., Hedwigia 19: 191.
1880.
(See Fusarium avenaceum)
Holotypus: Not located.
Type locality: Belgium.
Type substrate: Stem of Brassica oleracea.
Note: Synonym fide Wollenweber & Reinking (1935).
breve Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 1. 2020.
Neocosmospora brevis Sand.-Den. & Crous, Persoonia 43:
119. 2019.
Holotypus: CBS H-23975.
Ex-type culture: CBS 144387 = MUCL 16108.
Type locality: Belgium, Heverlee.
Type substrate: Soil-water polluted with diethylene glycerol and
ethylene glycerol.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834214; rpb2: LR583822;
tef1: LR583601.
brevicatenulatum Fusarium Nirenberg et al., Mycologia 90:
460. 1998.
Holotypus: Specimen in B fide Nirenberg et al. (1998).
Ex-type culture: BBA 69197 = CBS 404.97 = DAOM 225122= IMI
375329 = NRRL 25446.
Type locality: Madagascar.
Type substrate: Striga asiatica.
Descriptions and illustrations: See Nirenberg et al. (1998) and
Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: MT010948; rpb2: MT010979;
tef1: MT011005.
brevicaudatum Fusarium J.W. Xia et al., Persoonia 43: 195.
2019.
Holotypus: CBS H-24051.
Ex-type culture: NRRL 43638 = UTHSC R-3500.
Type locality: USA, Florida.
Type substrate: Trichechus sp. (manatee).
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb1: KC808322; rpb2: GQ505843;
tef1: GQ505665.
breviconum Fusarium (Wollenw.) O'Donnell et al., Index Fungorum 440: 1. 2020.
Neocosmospora brevicona (Wollenw.) Sand.-Den. & Crous,
Persoonia 43: 117. 2019.
Basionym: Hypomyces haematococcus var. breviconus Wollenw., Fusaria Autogr. Delin. 3: no. 828 (1930).
Synonyms: Fusarium solani var. minus Wollenw., Fusarien: 134.
1935.
Nectria haematococca var. brevicona (Wollenw.) Gerlach,
Fusarium: Diseases, Biology, and Taxonomy (Philadelphia): 422.
1981.
Lectotypus: Fig. 828 in Wollenweber (1930), designated in
Sandoval-Denis et al. (2019).
Epitypus: CBS H-23974 designated in Sandoval-Denis et al.
(2019).
FUSARIUM
Ex-epitype culture: BBA 2123 = CBS 204.31 = NRRL 22659.
Epitype locality: Indonesia, West Java, Bogor.
Epitype substrate: Gladiolus sp.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218103; rpb2: LR583821;
tef1: LR583600.
briosianum Fusarium Ferraris, Fl. Ital. Crypt. Hyphales, Fasc. 13:
857. 1914.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Italy, Pavia.
Type substrate: Branches of Styphnolobium japonicum (syn.
Sophora japonica).
Note: Synonym fide Wollenweber & Reinking (1935).
bubalinum Fusarium J.W. Xia et al., Persoonia 43: 195. 2019
Holotypus: CBS H-24052.
Ex-type culture: CBS 161.25 = NRRL 26857 = NRRL 26918.
Type locality: Australia.
Type substrate: Unknown.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170381; tef1: MN170448.
bufonicola Fusarium (Speg.) Sacc. & Trotter, Syll. Fung. 22:
1486. 1913.
(See Fusarium graminearum)
Basionym: Selenosporium bufonicola Speg., Anales Mus. Nac.
Buenos Aires, ser. 3, 13: 458. 1910.
Holotypus: In LPS (Myc. Argent. ser. 5, no. 1166) fide Farr
(1973).
Type locality: Argentina, Buenos Aires.
Type substrate: Decaying body of Amphibia (toad).
Note: Synonym fide Wollenweber & Reinking (1935).
bugnicourtii Fusarium Brayford, Trans. Brit. Mycol. Soc. 89: 350.
1987.
(See Fusarium ambrosium)
Synonym: Fusarium tumidum var. coeruleum Bugnic., Encycl.
Mycol. 11: 83. 1939.
Holotypus: IMI 296597.
Ex-type culture: IMI 296597 = NRRL 20438 = MAFF 246291.
Type locality: India, Chinchona.
Type substrate: Euwallacea fornicatus on Camellia sinensis.
Descriptions and illustrations: See Brayford (1987).
Diagnostic DNA barcodes: rpb1: JX171470; rpb2: JX171584;
tef1: AF178332.
buharicum Fusarium Jacz. ex Babajan & Teterevn.-Babajan,
Mater. Mikol. Fitopatol. 8: 216. 1929.
Holotypus: LEP 127667.
Epitypus (hic designatus, MBT 10000662): Uzbekistan, Tashkent, on Gossypium herbaceum, 1928, A.I. Raillo, CBS 178.35
(preserved as metabolically inactive culture).
Ex-epitype culture: CBS 178.35 = DSM 62166 = NRRL 25488.
Descriptions and illustrations: See Gerlach & Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: KX302920; rpb2: KX302928;
tef1: KX302912.
Notes: Gerlach & Nirenberg (1982) designated CBS 178.35 as
neotype of F. buharicum as they were unable to locate
the type specimen. However, A. Jaczweski did place a
specimen in LEP. Therefore, the neotype designation is superseded here (Art. 9.13) and CBS 178.35 is retained as
epitype for this species.
www.studiesinmycology.org
REDELIMITED
bulbicola Fusarium Nirenberg & O'Donnell, Mycologia 90: 452.
1998.
Replaced synonym: Fusarium sacchari var. elongatum Nirenberg, Mitt. Biol. Bundesanst. Land- Forstw. Berlin-Dahlem 169:
59. 1976, non Fusarium elongatum Reinking 1934.
Holotypus: IMI 202877.
Ex-type culture: BBA 63628 = CBS 220.76 = DAOM
225114 = IMI 375322 = NRRL 13618.
Type locality: Germany.
Type substrate: Haemanthus sp.
Descriptions and illustrations: See Nirenberg (1976), Nirenberg &
O'Donnell (1998) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: KF466394; rpb2: KF466404;
tef1: AF160294.
bulbigenum Fusarium Cooke & Massee, Grevillea 16: 49. 1887.
(See Fusarium oxysporum)
Holotypus: ?K(M).
Type locality: UK.
Type substrate: Narcissus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
bullatum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6:
198. 1915.
(See Fusarium equiseti)
Typus: CUP-007455.
Type locality: USA, Iowa.
Type substrate: Rotten tuber of Solanum tuberosum. Lectotypification pending study of material lodged in CUP.
burgessii Fusarium M.H. Laurence et al., Fungal Diversity 49:
109. 2011.
Holotypus: CBS 125537 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 125537 = NRRL 66654 = RBG 5315.
Type locality: Australia, Queensland, Idalia National Park.
Type substrate: Soil.
Descriptions and illustrations: See Laurence et al. (2011).
Diagnostic DNA barcodes: rpb1: MT409440; rpb2: HQ646393;
tef1: HQ667148.
butleri Fusarium Wollenw., Phytopathology 3: 38. 1913, nom.
illegit., Art. 52.1.
(See Fusarium udum)
Authentic material: ?B.
Original locality: India.
Original substrate: Cajanus cajan.
Note: Synonym fide Wollenweber & Reinking (1935).
butleri Fusarium Kr.P. Singh & Edward, Allahabad Farmer 49: 94.
1979, nom. illegit., Art. 53.1, non Fusarium butleri Wollenw. 1913.
Synonym: Gibberella butleri Kr.P. Singh & Edward, Allahabad
Farmer 49: 92. 1979.
Authentic material: Not located.
Original locality: India.
Original substrate: Cajanus cajan.
Notes: Status unclear. No further records available for this taxon.
buxi Fusarium Spreng., Syst. Veg., ed. 16, 4: 565. 1827.
Pseudonectria buxi (DC.) Seifert et al., Stud. Mycol. 68: 107.
2011.
Basionym: Tubercularia buxi DC., Fl. Franç., ed. 3, 5/6: 110.
1815.
Synonyms: Fusisporium buxi (DC.) Fr., Syst. Mycol. 3: 447.
1832, nom. sanct. [Fr., l.c.]
93
CROUS
ET AL.
Psilonia buxi (DC.) Fr., Syst. Mycol. 3: 447. 1832, nom. inval.,
Art. 36.1(c).
Chaetostroma buxi (DC.) Corda, Icon. Fung. 2: 31. 1838.
Volutella buxi (DC.) Berk. & Broome, Ann. Mag. Nat. Hist., ser. 2,
5: 465. 1850.
Chaetodochium buxi (DC.) Höhn., Mitt. Bot. Inst. T. H. Wien 9:
45. 1932.
Nectria rousseliana Mont., Ann. Sci. Nat., Bot., ser. 3, 16: 44.
1851.
Stigmatea rousseliana (Mont.) Fuckel, Jahrb. Nassauischen
Vereins Naturk. 23–24: 97. 1870.
Nectriella rousseliana (Mont.) Sacc., Syll. Fung. 2: 452. 1883.
Lasionectria rousseliana (Mont.) Cooke (as ‘rousselliana’), Grevillea 12: 111. 1884.
Pseudonectria rousseliana (Mont.) Wollenw., Z. Parasitenk.
(Berlin) 3: 489. 1931.
Notarisiella rousseliana (Mont.) Clem. & Shear, The genera of
Fungi: 280. 1931.
Nectria rousseliana var. viridis Berk. & Broome, Ann. Mag. Nat.
Hist., ser. 3, 3: 376. 1859.
Volutella buxi f. rusci Sacc., Michelia 2: 644. 1882.
Holotypus: ?PC.
Type locality: ?Germany/France.
Type substrate: Leaf of Buxus sp.
buxicola Fusarium Sacc., Syll. Fung. 2: 518. 1883.
Cyanonectria buxi (Fuckel) Schroers et al., Stud. Mycol. 68:
120. 2011.
Basionym: Gibbera buxi Fuckel, Jahrb. Nassauischen Vereins
Naturk. 27–28: 32. 1874.
Synonyms: Lisea buxi (Fuckel) Sacc., Syll. Fung. 2: 518. 1883.
Gibberella buxi (Fuckel) Cooke, Grevillea 12: 112. 1884.
Fusarium subcorticale Oudem., Ned. Kruidk. Arch., ser. 3, 3:
135. 1898.
Fusarium dimorphum J.V. Almeida & Sousa da C^amara, Revista
Agron. (Lisbon) 1: 306. 1903.
Fusarium buxicola var. chlamydosporum Batikyan (as ‘chlamydosporeae’), Biol. Zhurn. Armenii 22: 90. 1969.
Fusarium lateritium var. buxi C. Booth, The Genus Fusarium:
113. 1971.
Lectotypus: G 00111019, selected in Schroers et al. (2011).
Epitypus: CBS H-20379, designated in Schroers et al. (2011).
Ex-epitype culture: CBS 125551.
Epitype locality: Slovenia, Arboretum Volcji Potok.
Epitype substrate: Decaying twig of Buxus sempervirens var.
elegantissima.
Descriptions and illustrations: See Schroers et al. (2011).
Diagnostic DNA barcodes: rpb2: HM626689; tef1: HM626648.
Holotypus: UB 24189.
Ex-type culture: CML 3657.
Type locality: Brazil, S~ao Paulo, Guaíra.
Type substrate: Brachiaria brizantha.
Descriptions and illustrations: See Costa et al. (2021).
Diagnostic DNA barcodes: rpb2: MT901316; tef1: MT901350.
caatingaense Fusarium A.C.S. Santos et al., Mycologia 111:
248. 2019.
Holotypus: URM 91192.
Ex-type culture: MUM 1859 = URM 6779.
Type locality: Brazil, Pernambuco, Ibimirim.
Type substrate: Dactylopius opuntiae.
Descriptions and illustrations: See Santos et al. (2019).
Diagnostic DNA barcodes: rpb2: LS398495; tef1: LS398466.
cactacearum Fusarium Pasin. & Buzz.-Trav., Nuovo Giorn. Bot.
Ital. 42: 120. 1935.
Lectotypus (hic designatus, MBT 10000663): Italy, Milan, Thelocactus nidulans, 1935, L. Pasinetti & A. Buzzati-Traverso, in
Nuovo Giorn. Bot. Ital. 42: Pl. I, fig. 1.
Descriptions and illustrations: See Pasinetti & Buzzati-Traverso
(1935).
Notes: Based on illustrations by Pasinetti & Buzzati-Traverso
(1935), this species could be a synonym of Neocosmospora
solani but requires further investigation. No holotype specimen
could be located and therefore an illustration is designated as
lectotype.
cacti-maxonii Fusarium Pasin. & Buzz.-Trav., Nuovo Giorn.
Bot. Ital. 42: 120. 1935.
Lectotypus (hic designatus, MBT 10000664): Italy, Milan, Cactus
maxonii, 1935, L. Pasinetti & A. Buzzati-Traverso, in Nuovo
Giorn. Bot. Ital. 42: Pl. I, fig. 4.
Descriptions and illustrations: See Pasinetti & Buzzati-Traverso
(1935).
Notes: Based on illustrations by Pasinetti & Buzzati-Traverso
(1935), this species could be a synonym of Fusarium oxysporum but requires further investigation. No holotype specimen
could be located and therefore an illustration is designated as
lectotype.
byssinum Fusarium McAlpine, Proc. Linn. Soc. New South
Wales 22: 698. 1897.
Holotypus: VPRI 2556.
Type locality: Australia, New South Wales, Murwillumbah.
Type substrate: Desmodium sp.
Notes: Status unclear. This species was considered a member of
Diymopsis by Saccardo (1899); Hymenula by Wollenweber &
Reinking (1935); and Aschersonia by Walker (1962), who
examined the type specimen and found that the fungus occurred
in association with a scale insect on Desmodium. It is likely that
this species belongs in Microcera, which are usually parasites of
scale insects.
caeruleum Fusarium Lib. ex Sacc. (as ‘cæruleum’), Syll. Fung. 4:
705. 1886.
Synonyms: Fusarium solani var. caeruleum (Lib. ex Sacc.) Bilaĭ,
Fusarii (Biologija i sistematika): 287. 1955, nom. inval., Art. 41.5.
Fusarium solani var. caeruleum (Lib. ex Sacc.) C. Booth, The
Genus Fusarium: 51. 1971.
?Fusarium violaceum Fuckel, Fungi Rhen. Exs., Fasc. 3: no.
209. 1863.
Fusarium aeruginosum Delacr., Bull. Soc. Mycol. France 7: 110.
1891.
Selenosporium caeruleum Lib., 1834. (in herb.; nom. inval., Art.
38.1a).
Fusarium caeruleum var. cellulosae Sartory et al., Papier 38: 43.
1935.
?Hypomyces asclepiadis Zerova, Zhurn. Inst. Bot. Vseukraïns'k.
Akad. Nauk 11: 103. 1937.
Holotypus: BR5020140171069.
Type locality: Belgium.
Type substrate: Solanum tuberosum.
Notes: Status doubtful. See Sandoval-Denis et al. (2019).
caapi Fusarium M.M. Costa et al., Mycol. Progr. 20: 67. 2021.
calcareum Fusarium (Thüm.) Sacc., Syll. Fung. 4: 712. 1886.
94
FUSARIUM
(See Fusarium oxysporum)
Basionym: Fusisporium calcareum Thüm., Inst. Coimbra 28: 262.
1881.
Holotypus: S-F45605.
Type locality: Portugal, Coimbra.
Type substrate: Lagenaria vulgaris.
Note: Synonym fide Wollenweber & Reinking (1935).
?calidariorum Fusarium Sacc., Ann. Mycol. 4: 274. 1906.
Colletotrichum anthurii Delacr., Bull. Soc. Mycol. France 13:
110. 1897.
Synonyms: Fusoma calidariorum Sacc., Ann. Mycol. 4: 274.
1906.
Fusoma calidariorum var. acanthi Lindegg, Riv. Patol. Veg. 25:
233. 1935.
Holotypus: In PAD.
Type locality: Italy, Padua, botanical garden.
Type substrate: Anthurium scherzerianum.
Notes: Synonym fide Wollenweber & Reinking (1935). No record
could be located for the transfer of this epithet to the genus
Fusarium. In Saccardo (1906) on p. 274, no new combination is
provided and only the new name Fusoma calidariorum was
introduced. Similarly, Lindegg (1935) introduced a new variety as
Fusoma calidariorum var. acanthi, not in the genus Fusarium.
Although Wollenweber & Reinking (1935) did treat this as
Fusoma, Booth (1971) incorrectly treated the variety acanthi in
the genus Fusarium.
callistephi Fusarium L. Lombard & Crous, Persoonia 43: 15.
2018 [2019].
Holotypus: CBS H-23608.
Ex-type culture: CBS 187.53 = NRRL 36330.
Type locality: Netherlands, Oostenbrink.
Type substrate: Callistephus chinensis.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb2: MH484875; tef1: MH484966.
callosporum Fusarium Pat., Bull. Soc. Mycol. France 9: 164.
1893.
(See Fusarium coccophilum)
Holotypus: Not located.
Type locality: Ecuador, Quito.
Type substrate: Parasitic on Septobasidium pedicellatum.
Note: Synonym fide Wollenweber & Reinking (1935).
camerunense Fusarium Henn., Bot. Jahrb. Syst. 22: 81. 1895.
Gloeosporium camerunense (Henn.) Wollenw., Fusaria
Autogr. Delin. 1: 499. 1916.
Holotypus: In B fide Hein (1988).
Type locality: Cameroon, Itoki.
Type substrate: Bark of unknown tree.
Note: Synonym fide Wollenweber & Reinking (1935).
camptoceras Fusarium Wollenw. & Reinking, Phytopathology
15: 158. 1925.
Neotypus: CBS H-24077, designated in Xia et al. (2019).
Ex-neotype culture: ATCC 16065 = ATCC 24364 = BBA
9810 = CBS 193.65 = DSM 62167 = IMI 112500 = NRRL
20716 = NRRL 36344.
Neotype locality: Costa Rica.
Neotype substrate: Cushion gall of Theobroma cacao.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982), Marasas
et al. (1998) and Leslie & Summerell (2006).
www.studiesinmycology.org
REDELIMITED
Diagnostic DNA barcodes: rpb1: MW928800; rpb2: MN170383;
tef1: MN170450.
campylopodii Fusarium Weir, Mycologia 60: 374. 1968, nom.
inval., Art. 38.1(a).
Authentic material: Not located.
Original locality: USA, Washington.
Original substrate: Arceuthobium sp.
Note: The name is mentioned but neither a diagnosis nor a
description was provided.
candidulum Fusarium Sacc., Ann. Mycol. 6: 567. 1908.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: Mexico.
Type substrate: Myrtillocactus geometrizans.
Note: Synonym fide Wollenweber & Reinking (1935).
candidum Fusarium Ehrenb., Sylv. Mycol. Berol.: 24. 1818.
Neonectria candida (Ehrenb.) Rossman et al., Stud. Mycol. 80:
217. 2015.
Synonyms: Ramularia candida (Ehrenb.) Wollenw., Phytopathology 1: 220. 1913.
Cylindrocarpon ehrenbergii Wollenw., Fusaria Autogr. Delin. 1:
461. 1916.
Sclerotium castaneum Lib., in herb. 1832, nom. nud.
Fusarium castaneum Lindau (as “(Lib.) Lindau”), Rabenh. Krypt.Fl. 1(9): 556. 1909.
?Fusidium candidum Willk., Die mikroskopischen Feinde des
Waldes 1: 103. 1866, nom. illegit., Art. 53.1.
?Fusarium candidum Sacc. & D. Sacc., Syll. Fung. 18: 674.
1906, nom. illegit., Art. 53.1.
?Fusarium candidum Dasz., Bull. Soc. Bot. Geneve, 2 ser. 4:
293. 1913, nom. illegit., Art. 53.1.
Fusarium brassicae Lib. ex Cooke, Grevillea 8: 83. 1880.
Selenosporium brassicae Thüm., Hedwigia 19: 191. 1880.
Fusarium brassicae (Thüm.) Sacc., Syll. Fung. 4: 701. 1886,
nom. illegit., Art. 53.1.
Fusarium obtusiusculum Sacc., Michelia 2: 297. 1881.
Fusarium rhizogenum Pound & Clem., Bot. Surv. Nebraska 3:
12. 1894.
Fusarium oxysporum var. obtusiusculum (Sacc.) Cif., Ann. Bot.
(Rome) 16: 221. 1924.
Cylindrocarpon obtusiusculum (Sacc.) U. Braun, Cryptog. Bot. 4:
113. 1993.
Fusarium eichleri Bres., Ann. Mycol. 1: 130. 1903.
Neonectria ramulariae Wollenw., Ann. Mycol. 15: 52. 1917.
Nectria ramulariae (Wollenw.) E. Müll., Beitr. Kryptogamenfl.
Schweiz 11: 634. 1962.
Cylindrocarpon magnusianum Wollenw., Z. Parasitenk. (Berlin)
1: 172. 1928.
Holotypus: Not located.
Type locality: Germany, Berlin.
Type substrate: Unknown.
candidum Fusarium (Link) Sacc., Syll. Fung. 4: 720. 1886, nom.
illegit., Art. 53.1.
Neonectria ditissima (Tul. & C. Tul.) Samuels & Rossman, CBS
Biodiversity Ser. 4: 134. 2006.
Basionym: Nectria ditissima Tul. & C. Tul., Select. Fung. Carpol.
3: 73. 1865.
Synonyms: Cucurbitaria ditissima (Tul. & C. Tul.) Kuntze, Revis.
Gen. Pl. 3: 461. 1898.
95
CROUS
ET AL.
Fusidium candidum Link, Mag. Neuesten Entdeck. Gesammten
Naturk. Ges. Naturf. Freunde Berlin 3: 8. 1809, nom. sanct. [Fr.,
Syst. Mycol. 3: 481. 1832].
Cylindrocarpon candidum (Link) Wollenw., Fusaria Autogr. Delin.
1: 476. 1916.
?Fusisporium cylindricum Mont., Ann. Sci. Nat., Bot., ser. 2, 17:
120. 1842.
?Fusarium cylindricum (Mont.) Sacc., Syll. Fung. 4: 720. 1886.
Fusarium fissum Peyl, Lotos 8: 30. 1858.
?Fusarium heteronemum Berk. & Broome (as ‘heteronema'’),
Ann. Mag. Nat. Hist., Ser. 3, 15: 402. 1865.
?Cylindrocarpon heteronema (Berk. & Broome) Wollenw. (as
‘heteronemum’), Fusaria Autogr. Delin. 1: 460. 1916.
?Ramularia heteronema (Berk. & Broome) Wollenw. (as ‘heteronemum'’), Fusaria Autogr. Delin. 1: 460. 1916.
Fusarium ulmi P. Crouan & H. Crouan, Fl. Finistere: 14. 1867.
Fusarium fragrans P. Crouan & H. Crouan, Fl. Finistere: 14.
1867.
Fusarium decipiens Cooke & Massee, in Cooke, Handb. Austral.
Fungi: 388. 1892, nom. inval., Art. 39.1.
Fusarium mali Allesch., Ber. Bot. Vereines Landshut 12: 130.
1892.
Fusarium sarcochroum f. mali (Allesch.) Ferraris, 1910.
Cylindrocarpon mali (Allesch.) Wollenw., Phytopathology 18:
225. 1928.
Sporotrichum amenti P. Karst., Hedwigia 31: 296. 1892.
Fusarium fractum Sacc. & Cavara, Nuovo Giorn. Bot. Ital., n.s. 7:
308. 1900.
Cylindrocarpon fractum (Sacc. & Cavara) Wollenw., Fusaria
Autogr. Delin. 2: 655. 1924.
Nectria galligena Bres., in Strasser, Verh. K. K. Zool.-Bot. Ges.
Wien 51: 413. 1901.
Dialonectria galligena (Bres.) Petch ex E.W. Mason & Grainger,
Cat. Yorkshire Fung.: 32. 1937.
Neonectria galligena (Bres.) Rossman & Samuels, Stud. Mycol.
42: 159. 1999.
Fusarium prunorum McAlpine, Fungus Diseases of stone-fruit
trees in Australia: 91. 1902.
Fusarium willkommii Lindau, Rabenh. Krypt.-Fl. ed. 2, 1(9): 551.
1909.
Cylindrocarpon willkommii (Lindau) Wollenw., Z. Parasitenk.
(Berlin) 1: 150. 1928.
Fusarium luteum Parav., Ann. Mycol. 16: 302. 1918, nom. illegit.,
Art. 53.1.
Nectria ditissima var. arctica Wollenw., Angew. Bot. 8: 189. 1926.
Cylindrocarpon candidum var. medium Wollenw., Z. Parasitenk.
(Berlin) 1: 158. 1928.
Cylindrocarpon candidum var. majus Wollenw., Z. Parasitenk.
(Berlin) 1: 158. 1928.
Cylindrocarpon candidum var. minus Wollenw., Z. Parasitenk.
(Berlin) 1: 155. 1928.
Cylindrocarpon mali var. flavum Wollenw., Z. Parasitenk. (Berlin)
1: 150. 1928.
Cylindrocarpon willkommii var. pluriseptatum Wollenw., Z. Parasitenk. (Berlin) 1: 152. 1928.
Cylindrocarpon willkommii var. minus Wollenw., Z. Parasitenk.
(Berlin) 1: 152. 1928.
Holotypus: Not located.
Type locality: Unknown.
Type substrate: Branch.
Notes: Synonyms fide Wollenweber & Reinking (1935). Several
names that include Fusidium candidum (1809), Fusisporium
96
cylindricum (1842) and Fusarium fissum (1858) should take
preference for this taxon. However, the epithet “candidum” is
already occupied in the genus Neonectria and cannot be used.
Furthermore, the link between Fusisporium cylindricum and
Fusarium fissum with Neonectria ditissima still needs to be
established. Therefore, we choose to retain the name Neonectria
ditissima for this taxon.
candidum Fusarium Sacc. & D. Sacc., Syll. Fung. 18: 674. 1906,
nom. illegit., Art. 53.1, non Fusarium candidum Ehrenb. 1818.
Basionym: Fusidium candidum Willk., Die mikroskopischen
Feinde des Waldes 1: 103. 1866.
Replacing synonym: Fusarium willkommii Lindau, Rabenh.
Krypt.-Fl. ed. 2, 1(9): 551. 1910.
(See Fusarium willkommii)
capitatum Fusarium Schwein., Trans. Amer. Philos. Soc., n.s., 4:
302. 1832.
Synonym: Pionnotes capitata (Schwein.) Fr., Summa Veg.
Scand. 2: 481. 1849.
Holotypus: PH00081394.
Type locality: USA, Pennsylvania.
Type substrate: Tsuga canadensis.
Notes: The type material of Fusarium capitatum, type species of
the genus Pionnotes, was re-examined by Seifert (2013). It
represents not a hyphomycete but a basidiomycete identical to
Dacrymyces chrysospermus. Therefore, the generic name
Pionnotes is a synonym of Dacrymyces rather than Fusarium.
Further evaluations are necessary in future phylogenetic revisions of the Dacrymycetales.
caricis Fusarium Oudem., Verslagen Meded. Afd. Natuurk. Kon.
Akad. Wetensch., ser. 3, 7: 325. 1890.
(See Fusarium graminearum)
Holotypus: ?L.
Type locality: Netherlands, Zuid-Holland Province, Den Haag.
Type substrate: Leaves of Carex sp.
Note: Synonym fide Wollenweber & Reinking (1935).
caries Fusarium Nees, Nova Acta Phys.-Med. Acad. Caes.
Leop.-Carol. Nat. Cur. 19, Suppl. 1: 478. 1843.
Holotypus: ?B, L or STR.
Type locality: China.
Type substrate: Meoschium lodiculare.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
carminascens Fusarium L. Lombard et al., Persoonia 43: 19.
2018 [2019].
Holotypus: CBS H-23609.
Ex-type culture: CBS 144738 = CPC 25800.
Type locality: South Africa, KwaZulu-Natal Province.
Type substrate: Zea mays.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928801; rpb2: MH484937;
tef1: MH485028.
carneolum Fusarium P. Karst., Meddeland. Soc. Fauna Fl. Fenn.
16: 35. 1888.
Vermicularia herbarum Westend., Herb. Crypt. Belg. no. 393.
1849.
Holotypus: ?H.
Type locality: Finland, Tammela.
Type substrate: Iris pseudacorus.
Note: Synonym fide Wollenweber & Reinking (1935).
FUSARIUM
carneoroseum Fusarium Cooke, Grevillea 19: 4. 1890.
(See Fusarium lateritium)
Holotypus: In K(M), Colenso 538 fide Index Fungorum.
Type locality: New Zealand.
Type substrate: Bark.
Note: Synonym fide Wollenweber & Reinking (1935).
carneum Fusarium (Mont.) Sacc., Syll. Fung. 4: 724. 1886.
Basionym: Fusisporium carneum Mont., Ann. Sci. Nat., Bot., ser.
2, 17: 120. 1842.
Holotypus: ?PC.
Type locality: Cuba.
Type substrate: Leaf of monocotyledon.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
carniforme Fusarium Ellis & Tracy, nom. inval., Art. 38.1(a).
Synonym: Ramularia carniformis Sherb., Phytopathology 18:
149. 1928.
Authentic material: NY0093683.
Original locality: USA, Mississippi, Starkville.
Original substrate: Tripsacum dactyloides.
Notes: Status unclear. Braun (1998) considered this species
doubtful as conidia appeared microdochium-like.
carpineum Fusarium Davis, Trans. Wisconsin Acad. Sci. 18:
106. 1915.
Holotypus: BPI 442722.
Type locality: USA, Wisconsin, Wyalusing.
Type substrate: Carpinus caroliniana.
Notes: This species was not treated by any of Wollenweber &
Reinking (1935), Booth (1971), or Gerlach & Nirenberg (1982).
A literature search could not find any additional information
pertaining to this species.
carpini Fusarium Schulzer & Sacc., Hedwigia 23: 128. 1884.
(See Fusarium expansum)
Holotypus: Not located.
Type locality: Croatia, Vinkovci.
Type substrate: Carpinus betulus.
Note: Synonym fide Wollenweber & Reinking (1935).
cassiae Fusarium R.H. Perera et al., Mycosphere 11: 2138.
2020.
Holotypus: MFLU 18-2751.
Ex-type culture: MFLUCC 18-0573.
Type locality: Thailand, Phayao Province.
Type substrate: Pods of Cassia fistula.
Descriptions and illustrations: See Perera et al. (2020).
Diagnostic DNA barcodes: rpb2: MT212197; tef1: MT212205.
castagnei Fusarium Mont., Ann. Sci. Nat., Bot., ser. 3, 12: 296.
1849.
Myxosporium castagnei (Mont.) Wollenw., Fusaria. Autogr.
Delin. 1: 489. 1916.
Holotypus: ?PC.
Type locality: France, Marseille.
Type substrate: Psoralea bituminosa.
Note: Synonym fide Wollenweber & Reinking (1935).
castaneicola Fusarium W. Yamam., Trans. Mycol. Soc. Japan 3:
114. 1962, nom. inval., Art. 39.1 & 40.1.
Rugonectria castaneicola (W. Yamam. & Oyasu) Hirooka & P.
Chaverri, Stud. Mycol. 68: 73. 2011.
www.studiesinmycology.org
REDELIMITED
Basionym: Nectria castaneicola W. Yamam. & Oyasu, Sci. Rep.
Hyogo Univ. Agric. 3: 15. 1957.
Synonyms: Neonectria castaneicola (W. Yamam. & Oyasu) Tak.
Kobay. & Hirooka, J. Gen. Pl. Pathol. 71: 126. 2005, nom. inval.,
Art. 41.5.
Cylindrocarpon castaneicola Tak. Kobay. & Hirooka, J. Gen. Pl.
Pathol. 71: 126. 2005.
Authentic material: Not designated.
Original locality: Japan, Hyogo.
Original substrate: Castanea crenata.
Note: This Fusarium epithet is invalid as neither a Latin diagnosis
(Art. 39.1) nor a type designation (Art. 40.1) was included in the
original description.
castaneum Fusarium Lindau (as “(Lib.) Sacc.“), Rabenh. Krypt.Fl. ed. 2, 1(9): 556. 1909.
Synonym: Sclerotium castaneum Lib., in herb. 1832, nom. nud.
(See Fusarium candidum Ehrenb.)
Authentic material: Not located.
Original locality: Belgium, Ardennes.
Original substrate: Brassica oleracea.
cataleptum Fusarium Cooke & Harkn., Grevillea 12: 96. 1884.
(See Fusarium coccophilum)
Holotypus: In K(M), Harkness 1981 fide Index Fungorum.
Type locality: USA, California, San Rafael.
Type substrate: Acacia sp.
Note: Synonym fide Wollenweber & Reinking (1935).
catenatum Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index Fungorum. 440: 1. 2020.
Neocosmospora catenata Sand.-Den. & Crous, Persoonia 41:
115. 2018.
Holotypus: CBS H-23225.
Ex-type culture: CBS 143229 = NRRL 54993 = UTHSC 09-1009.
Type locality: USA, Georgia.
Type substrate: Stegostoma fasciatum.
Descriptions and illustrations: See Sandoval-Denis & Crous
(2018).
Diagnostic DNA barcodes: rpb1: KC808292; rpb2: KC808355;
tef1: KC808214.
cateniforme Fusarium J.W. Xia et al., Persoonia 43: 197. 2019
Holotypus: CBS H-24053.
Ex-type culture: ATCC 11853 = CBS 150.25.
Type locality: Unknown.
Type substrate: Unknown.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170384; tef1: MN170451.
catenulatum Fusarium F.J. Chen, Mycosystema 19: 459. 2000.
Holotypus: HMAS 71749.
Ex-type culture: AS 3.4704.
Type locality: China, Shaanxi, Yangling.
Type substrate: Gossypium hirsutum.
Descriptions and illustrations: See Chen (2000).
caucasicum Fusarium Letov, Mater. Mikol. Fitopatol. 8: 225.
1929.
Holotypus: Not located.
Ex-type culture: CBS 179.35 = IFO 5979 = NRRL 13954.
Type locality: Republic of Azerbaijan.
Type substrate: Gossypium hirsutum.
Descriptions and illustrations: See Gerlach & Nirenberg (1982).
97
CROUS
ET AL.
Notes: Status doubtful/unclear. The ex-type culture (CBS 179.35)
accessioned in CBS appears to be either contaminated or transpositioned by another Fusarium sp. (Sandoval-Denis et al. 2019).
A sequence of the tef1 gene region (DQ247543) from the copy
accessioned at NRRL (NRRL 13954) places this species within
the Neocosmospora falciformis clade (Sandoval-Denis et al.
2019). The status of the copy accessioned at IFO is not known.
caudatum Fusarium Wollenw., J. Agric. Res. 2: 262. 1914.
(See Fusarium scirpi)
Lectotypus (hic designatus, MBT 10000665): USA, South Carolina, Clemson College, Ipomoea batatas, date unknown, Harter
& Field, in Wollenweber, J. Agric. Res. 2: 262, pl. 16, fig. M.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
cavispermum Fusarium Corda, Icon. Fung. 1: 3. 1837.
Cosmosporella cavisperma (Corda) Sand.-Den. et al., Stud.
Mycol. 98 (no. 100116): 44. 2021.
Synonyms: Fusarium aquaeductuum var. cavispermum (Corda)
Raillo, Fungi of the Genus Fusarium: 280. 1950.
Fusarium oxydendri Ellis & Everh., Bull. Torrey Bot. Club 24: 477.
1897.
Fusarium cavispermum var. minus Wollenw., Fusaria Autogr.
Delin. 3: 848. 1930.
Lectotypus: AKJ. Corda, Icon. Fung. 1: pl. I, fig. 58, designated in
this study.
Type locality: Czech Republic.
Type substrate: Resin of Pinus sp.
Epitypus: CBS 172.31 (metabolic inactive specimen) designated
in this study.
Ex-epitype: CBS 172.31 = NRRL 13996.
Epitype locality: Norway.
Epitype substrate: Pinus sylvestris.
Diagnostic DNA barcodes: rpb1: JX171465; rpb2: JX171579.
celosiae Fusarium T. Abe, Mem. Coll. Agric. Kyoto Imp. Univ. 7:
51. 1928.
(See Fusarium fujikuroi)
Holotypus: Not located.
Type locality: Japan.
Type substrate: Living stems and leaves of Celosia cristata.
Note: Synonym fide Wollenweber & Reinking (1935).
celtidicola Fusarium Q.J. Shang et al., Phytotaxa 361: 255.
2018.
Holotypus: MFLU 15-3646.
Ex-type culture: KUMCC 16-0019 = MFLUCC 16-0526.
Type locality: Italy, Forlì-Cesena Province, Forlì, Viale
dell’Appennino.
Type substrate: Celtis australis.
Descriptions and illustrations: See Shang et al. (2018).
Diagnostic DNA barcodes: rpb1: MH576579; rpb2: MH576577;
tef1: MH576581.
celtidis Fusarium Ellis & Tracy, J. Mycol. 6: 76. 1890.
(See Fusarium lateritium)
Syntypes: In BPI, ISC & MICH.
Type locality: USA, Mississippi, Starkville.
Type substrate: Celtis occidentalis.
Note: Synonym fide Wollenweber & Reinking (1935).
98
celtidis Fusarium Pass., Atti Reale Accad. Lincei, Rendiconti Cl.
Sci. Fis., 4 ser. 7: 51. 1891, nom. illegit., Art. 53.1.
Replacing synonym: Fusarium sphaeriiforme Sacc. (as
‘sphaeriaeforme’), Syll. Fung. 10: 723. 1892.
(See Fusarium melanochlorum)
Holotypus: ?PARMA.
Type locality: Italy, Parma, Vigheffio.
Type substrate: Dead branch of Celtis australis.
Note: Synonym fide Wollenweber & Reinking (1935).
cepae Fusarium Hanzawa, Mycol. Centralbl. 5(1): 5. 1914.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000666): Japan, Sapporo,
Allium cepa, 1914, J. Hanzawa, 5(1): 6, fig. 1.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
cerasi Fusarium Rolland & Ferry, in Roumeguere, Rev. Mycol.
(Toulouse) 14: 170. 1892.
?Foveostroma drupacearum (Lev.) DiCosmo, Canad. J. Bot.
56: 1682. 1978.
Basionym: Micropera drupacearum Lev., Ann. Sci. Nat., Bot., ser.
3, 5: 283. 1846.
Synonyms: ?Peziza cerasi Pers., Neues Mag. Bot. 1: 115. 1794.
?Dermea cerasi (Pers.) Fr., Syst. Orb. Veg. 1: 115. 1825.
Syntype: ILL00220294 (Fungi Sel. Gall. Exs. No. 6119).
Type locality: France, Saint-Die-des-Vosges.
Type substrate: Prunus sp. (cherry tree).
Note: This species was excluded from Fusarium by Wollenweber
(1943). Gerlach & Nirenberg (1982) considered this species as a
possible synonym of Micropera drupacearum on which the
present synonymies are based.
cerealis Fusarium (P. Karst.) Gruyter & J.H.M. Schneid., Jaarb.
Plantenziektenk. Dienst 1989/1990, no. 168: 135. 1991, nom.
inval., Art. 41.4.
Gliomastix cerealis (P. Karst.) C.H. Dickinson, Mycol. Pap. 115:
19. 1968.
Basionym: Coniosporium cerealis P. Karst., Meddeland. Soc.
Fauna Fl. Fenn. 14: 109. 1887.
Synonyms: Acremonium cerealis (P. Karst.) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart): 88. 1971.
Gliomastix guttuliformis J.C. Br. & W.B. Kendr., Trans. Brit.
Mycol. Soc. 41: 499. 1958.
Holotypus: In herb. P.A. Karsten in H fide Dickinson (1968).
Type locality: Finland, Mustiala.
Type substrate: Secale cereale.
cerealis Fusarium (Cooke) Sacc., Syll. Fung. 4: 713. 1886.
Basionym: Fusisporium cerealis Cooke, Grevillea 6: 139. 1878.
Synonym: Fusarium culmorum var. cerealis (Cooke) Wollenw.,
Fusaria Autogr. Delin. 3: 946. 1930.
Fusarium roseum f. cerealis (Cooke) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 32: 663. 1945.
Gibberella rosea f. cerealis (Cooke) W.C. Snyder & H.N. Hansen,
Amer. J. Bot. 32: 664. 1945.
Fusarium sambucinum var. cerealis (Cooke) Raillo, Fungi of the
Genus Fusarium: 211. 1950.
Fusarium crookwellense L.W. Burgess et al., Trans. Brit. Mycol.
Soc. 79: 498. 1982.
Holotypus: ?K(M) 133541.
FUSARIUM
Type locality: USA, California, Gainesville.
Type substrate: Zea mays.
Notes: Wollenweber & Reinking (1935) considered F. cerealis as
a variety of F. culmorum, whereas Raillo (1950) considered it as
a variety of F. sambucinum. Gerlach & Nirenberg (1982) applied
a broader concept to F. culmorum that did not separate this
variety in F. culmorum. Nirenberg (1990) recognised F. cerealis
as a species and considered F. crookwellense as a synonym of
F. cerealis. However, Leslie & Summerell (2006) recommend the
use of the name F. crookwellense over F. cerealis, indicating
that no type material is available for F. cerealis. We choose to
follow Nirenberg (1990) to consider F. crookwellense a synonym
under F. cerealis. The material lodged in K(M) requires further
investigation to determine whether epi- or neotypification is
required.
cesatii Fusarium Rabenh., Klotzschii Herb. Viv. Mycol. Cent. 15:
no. 1440. 1850.
Hymenula rubella Fr., Elench. Fung. 2: 38. 1828.
Lectotypus (of Fusarium cesatii, hic designatus, MBT 10000667):
Italy, Vercelli, Carex sp., 1849, collector unknown, Rabenh.,
Klotzschii Herb. Viv. Mycol. Ed. I no. 1440 in HAL.
Note: Synonym fide Wollenweber & Reinking (1935).
cesatii Fusarium Thüm., Pilze Weinst.: 49. 1878, nom. illegit., Art.
53.1, non Fusarium cesatii Rabenh. 1850.
Elsinoe ampelina (de Bary) Shear, Phytopathology 19: 677.
1929.
Basionym: Sphaceloma ampelina de Bary, Ann. Oenol. 4: 165.
1874.
Synonyms: Manginia ampelina (de Bary) Viala & Pacottet, C. r.
hebd. Seanc. Acad. Sci., Paris 139: 88. 1904.
Pionnotes cesatii Sacc., Syll. Fung. 4: 726. 1886.
Ramularia ampelophaga Pass., Bol. Comit. Agric. Parmense 9:
125. 1876.
Gloeosporium ampelophagum (Pass.) Sacc., Michelia 1: 217.
1878.
Authentic material: S-F47363.
Original locality: Italy, Vercelli.
Original substrate: Decaying stump of Vitis vinifera.
Note: Synonym fide Wollenweber & Reinking (1935).
chaetomium Fusarium Wallr., Fl. Crypt. Germ. 2: 242. 1833.
Colletotrichum chaetomium (Wallr.) S. Hughes, Canad. J. Bot.
36: 753. 1958.
Holotypus: ?STR.
Type locality: Germany.
Type substrate: Decaying Cucurbita.
chenopodinum Fusarium (Thüm.) Sacc., Syll. Fung. 4: 701.
1886.
(See Fusarium scirpi)
Basionym: Fusisporium chenopodinum Thüm., Mycoth. Univ.
Cent. 14: no. 1378. 1879.
Syntypes: In BPI, CHRB, ILL, NEB, NY, NYS & PUL.
Type locality: Austria, Niederösterreich, Klosterneuburg.
Type substrate: Chenopodium album.
Note: Synonym fide Wollenweber & Reinking (1935).
chilense Fusarium (Mont.) Sacc., Syll. Fung. 4: 716. 1886.
Gloeosporium chilense (Mont.) Wollenw., Z. Parasitenk.
(Berlin) 3: 496. 1931.
Basionym: Fusisporium chilense Mont., in Gay, Fl. Chil. 8: 25.
1852.
www.studiesinmycology.org
REDELIMITED
Fusisporium argillaceum Mont., Bull. Mass. Agric. Exp. Sta. no.
55. 1842, nom. illegit., Art. 53.1, non Fusarium argillaceum Fr.
1832.
Holotypus: In UPS fide Wollenweber, Fusaria Autogr. Delin. 2:
658.
Type locality: Chile, Juan Fernandez Islands.
Type substrate: Bark of Urtica excelsa.
Note: Synonym fide Wollenweber & Reinking (1935).
chinhoyiense Fusarium Yilmaz & Crous, Persoonia 46: 147.
2021.
Holotypus: PREM 63215.
Ex-type culture: BBA 69031 = DAOM 225149 = Frank
5bCn8 = IMI 375355 = NRRL 25221 = NY007.I2.
Type locality: Zimbabwe, Chinhoyi.
Type substrate: Zea mays.
Descriptions and illustrations: See Yilmaz et al. (2021).
Diagnostic DNA barcodes: rpb1: MW402711; rpb2: MN534262;
tef1: MN534050.
chlamydosporum Fusarium Wollenw. & Reinking, Phytopathology 15: 156. 1925.
Synonyms: Fusarium sporotrichioides var. chlamydosporum
(Wollenw. & Reinking) Joffe, Mycopathol. Mycol. Appl. 53: 211.
1974, nom. inval., Art. 41.1.
Dactylium fusarioides Gonz. Frag. & Cif., Bol. Real Soc. Esp.
Hist. Nat. 27: 280. 1927.
Fusarium fusarioides (Gonz. Frag. & Cif.) C. Booth, The Genus
Fusarium: 88. 1971.
Pseudofusarium purpureum Matsush., Microfungi of the Solomon Islands and Papua New Guinea: 47. 1971.
Neotypus: CBS 145.25 (preserved as metabolically inactive
culture), designated in Lombard et al. (2019a).
Ex-neotype culture: CBS 145.25 = NRRL 26851 = NRRL 26912.
Neotype locality: Honduras, Tela.
Neotype substrate: Musa sapientum.
Descriptions and illustrations: See Wollenweber & Reinking
(1925), Booth (1971), Gerlach & Nirenberg (1982) and Leslie
& Summerell (2006).
Diagnostic DNA barcodes: rpb1: MN120715; rpb2: MN120735;
tef1: MN120754.
cicatricum Fusarium (Berk.) O'Donnell & Geiser, Phytopathology
103: 404. 2013.
Geejayessia cicatricum (Berk.) Schroers, Stud. Mycol. 68: 124.
2011.
Basionym: Sphaeria sanguinea var. cicatricum Berk., Mag. Zool.
Bot. 1: 48. 1837.
Synonyms: Nectria cicatricum (Berk.) Tul. & C. Tul., Select.
Fung. Carpol. 3: 77. 1865.
Sphaeria sanguinea var. cicatricum Haller, Syst. Nat., ed 13, 1:
LII. 1768.
Sphaeria coccinea var. cicatricum Desm., Ann. Sci. Nat., Bot.,
ser. 3, 10: 351. 1848.
Nectria coccinea var. cicatricum (Desm.) Sacc., Syll. Fung. 2:
482. 1883.
Cucurbitaria cicatricum (Desm.) Kuntze, Revis. Gen. Pl. 3: 462.
1898.
Nectria gibbera Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 177. 1870.
Fusarium fuckelii Sacc., Syll. Fung. 4: 695. 1886.
Nectria desmazieri Fuckel ex Sacc., Syll. Fung. 4: 695. 1886,
nom. inval., Art. 36.1(d).
99
CROUS
ET AL.
Lectotypus: K(M) 160064 (MBT 10001323 hic designatus).
Epitypus: CBS H-20374 (MBT 10001324 hic designatus).
Ex-epitype culture: CBS 125549.
Epitype locality: Slovenia, Arboretum Volcji Potok.
Epitype substrate: Decaying twigs of Buxus sempervirens.
Descriptions and illustrations: See Schroers et al. (2011).
Diagnostic DNA barcodes: rpb1: KM232231; rpb2: HM626679;
tef1: HM626643.
Notes: The epitypification in Schroers et al. (2011) was not Code
compliant as neither a supporting holo-, lecto- nor epitype was
cited. The specimen in the Kew herbarium was cited as isotype.
In the protologue a single gathering is mentioned, but an illustration is also cited so a lectotypification is necessary. The epitypification is validated herein.
ciliatum Fusarium (Link) Link, in Willdenow, Sp. Pl., Ed. 4, 6: 105.
1825.
Scolecofusarium ciliatum (Link) L. Lombard et al., Stud. Mycol.
98 (no. 100116): 74. 2021.
Basionym: Atractium ciliatum Link, Mag. Neuesten Entdeck.
Gesammten Naturk. Ges. Naturf. Freunde Berlin 7: 32. 1816.
Synonyms: Microcera ciliata (Link) Wollenw., Fusaria Autogr.
Delin. 1: 435. 1916.
Calonectria ciliata (Link) W.C. Snyder & H.N. Hansen, Amer. J.
Bot. 32: 664. 1945.
Sphaeria agnina Desm., Ann. Sci. Nat., Bot. ser. 3, 6: 72. 1846.
Calonectria agnina (Desm.) Sacc., Michelia 1(3): 311. 1878.
Dialonectria agnina (Desm.) Cooke, Grevillea 12: 111. 1884.
Fusarium peltigerae Westend., Herb. Crypt. Belg. 9: no. 414.
1849.
Fusarium parasiticum Westend., Bull. Seances Cl. Sci. Acad.
Roy. Sci. Belgique, ser. 2, 11: 652. 1861.
Nectria massariae Pass., in Rabenhorst, Fungi Eur. Exs. no.
1827. 1874.
Microcera massariae Sacc., Michelia 1(2): 262. 1878.
Calonectria massariae (Pass.) Sacc., Michelia 1(3): 312. 1878.
Fusisporium filisporum Cooke, Grevillea 8: 8. 1879.
Fusarium filisporum (Cooke) Sacc., Syll. Fung. 4: 708. 1886.
Fusarium scolecoides Sacc. & Ellis, Atti Reale Ist. Veneto Sci.
Lett. Arti, ser. 6, 3: 728. 1885.
Fusarium elongatum Cooke, Grevillea 19: 4. 1890.
Calonectria dearnessii Ellis & Everh., Proc. Acad. Nat. Sci.
Philadelphia 42: 245. 1891.
Neotypus: CBS H-12687 designated in this study.
Ex-neotypus: ATCC 16068 = ATCC 24137 = BBA 9661 = CBS
191.65 = DSM 62172 = IMI 112499 = NRRL 20431.
Neotype locality: Germany.
Neotype substrate: Branch canker of Fagus sylvatica.
Diagnostic DNA barcodes: rpb1: MW834264; rpb2: MW834035;
tef1: MW834296.
cinctum Fusarium Corda, Icon. Fung. 5: 80. 1842.
Striaticonidium cinctum (Corda) L. Lombard & Crous, Persoonia 36: 229. 2016.
Synonyms: Myrothecium cinctum (Corda) Sacc., Syll. Fung. 4:
751. 1886.
?Myrothecium ellipsosporum Fuckel (as ‘ellipsisporium’), Fungi
Rhen. Exs. Cent. 16, no. 1529 (1865).
?Hymenopsis ellipsospora (as ‘ellipsosporum’) (Fuckel) Sacc.,
Syll. Fung. 4: 745. 1886.
Myrothecium striatisporum N.C. Preston, Trans. Brit. Mycol. Soc.
31: 275. 1948.
100
Myrothecium longistriatisporum Matsush., Microfungi Solomon
Isl. Papua-New Guinea: 39. 1971.
Lectotypus: PR 155489, designated in Tulloch (1972).
Epitypus: CBS H-22471, designated in Lombard et al. (2016).
Ex-epitype culture: CBS 932.69 = IMI 145760.
Epitype locality: Netherlands, Eastern Flevoland.
Epitype substrate: Agricultural soil.
Note: The lectotype was cited as holotype in Lombard et al.
(2016) but this is correctable according to Art. 9.10 of the
Code (see also Ex. 11).
cinnabarinum Fusarium (Berk. & M.A. Curtis) Sacc., Syll. Fung.
4: 722. 1886.
(See Fusarium lateritium)
Basionym: Fusisporium cinnabarinum Berk. & M.A. Curtis,
Grevillea 3: 146. 1875.
Syntypes: In PH, Pul & USCH:Fungi (Ellis, N. Amer. F. 3990).
Type locality: USA, Alabama.
Type substrate: Acer negundo.
Note: Synonym fide Wollenweber & Reinking (1935).
circinatum Fusarium Nirenberg & O'Donnell, Mycologia 90:
442. 1998.
Synonyms: Gibberella circinata Nirenberg & O'Donnell, Mycologia 90: 440. 1998, nom. inval., Art. 40.3.
Gibberella circinata Nirenberg & O'Donnell ex Britz et al.,
Sydowia 54: 16. 2002.
Holotypus: B 70 0001693.
Ex-type culture: BBA 69720 = CBS 405.97 = DAOM
225113 = IMI 375321 = NRRL 25331.
Type locality: USA, California.
Type substrate: Pinus radiata.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998).
Diagnostic DNA barcodes: rpb1: JX171510; rpb2: JX171623;
tef1: AF160295.
cirrosum Fusarium Höhn., Sitzungsber. Kaiserl. Akad. Wiss.
Wien, Math.-Naturwiss. Cl., Abt. 1., 116: 153. 1907.
(See Fusarium expansum)
Holotypus: FH00284266.
Type locality: Austria, Niederösterreich, Irenental near
Untertullnerbach.
Type substrate: Parasictic in the acervuli of Steganosporium
pyriforme (syn. Steganosporium ovatum).
Note: Synonym fide Wollenweber & Reinking (1935).
citri Fusarium M.M. Wang et al., Persoonia 43: 79. 2019.
Holotypus: HAMS 248036.
Ex-type culture: CGMCC 3.19467 = LC6896.
Type locality: China, Hunan Province.
Type substrate: Leaves of Citrus reticulata.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289828; rpb2: MK289771;
tef1: MK289617.
citricola Fusarium Guarnaccia et al., Persoonia 40: 12. 2017.
[2018].
Holotypus: CBS H-23020.
Ex-type culture: CBS 142421 = CPC 27805.
Type locality: Italy, Cosenza, Rocca Imperiale.
Type substrate: Citrus reticulata ‘Caffin’.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018a).
FUSARIUM
Diagnostic DNA barcodes: rpb1: LT746290; rpb2: LT746310;
tef1: LT746197.
citriforme Fusarium Jamal., Valt. Maatalousk. Julk. 123: 11.
1943.
(See Fusarium tricinctum)
Lectotypus (hic designatus, MBT 10000668): Finland, Pyhaj€arvi,
Hordeum sativum, 1938, E. Jamalainen, in Valt. Maatalousk.
Julk. 123: 10. 1943, fig. 2.
Ex-type culture: CBS 253.50.
Diagnostic DNA barcodes: rpb1: MW928802; rpb2: MW928823;
tef1: KR071775.
Notes: Jamalainen (1943) cited various specimens in the protologue of F. citriforme, but failed to indicate a holotype. Therefore, a lectotypification is done here to fix the name. Isolate CBS
253.50 was deposited in the public collection of CBS by E.
Jamalainen in 1950. The isolate was indicated as the living extype culture of F. citriforme.
citrinum Fusarium Wollenw., in Lewis, Bull. Maine Agric. Exp.
Sta. 219: 256. 1913.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000669): Germany, Berlin,
Dahlem, rotten fruit of Solanum lycopersicum, Oct. 1910, H.W.
Wollenweber, B70 0100185.
Notes: Synonym fide Wollenweber & Reinking (1935). Only one
specimen located at B matches the original collection event, but
it is not indicated as the type. Therefore B 70 0100185 is
designated as lectotype here.
citrulli Fusarium Taubenh., Bull. Texas Agric. Exp. Sta. 260: 27.
1920.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000670): USA, Texas, Waller
County, seedlings of Citrullus lanatus, 1920, J.J. Taubenhaus, in
Bull. Texas Agric. Exp. Sta. 260: 30, fig. 8h.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
citrulli Fusarium Sartory, Compt. Rend. Hebd. Seances Acad. Sci.
188: 1434. 1929, nom. inval., Art. 35.2; nom. illegit., Art. 53.1.
Neocosmospora martii (Appel & Wollenw.) Sand.-Den. &
Crous, Persoonia 43: 142. 2019.
Basionym: Fusarium martii Appel & Wollenw., Arbeiten Kaiserl.
Biol. Anst. Land-Forstw. 8: 83. 1910.
Synonyms: Fusarium solani var. martii (Appel & Wollenw.)
Wollenw., Fusaria Autogr. Delin. 3: 1034. 1930.
?Selenosporium fuscum Bonord., Handb. Mykol.: 135. 1851.
?Fusarium fuscum (Bonord.) Sacc., Syll. Fung. 4: 699. 1886.
Fusarium citrulli Sartory & J. Mey., Compt. Rend. Soc. Biol. 107:
55. 1931, nom. illegit., Art. 53.1, non Fusarium citrulli Taubenh.
1920.
Neocosmospora croci Guarnaccia et al., Persoonia 40: 17. 2017
[2018].
Authentic material: Not located.
Original locality: France.
Original substrate: Citrullus vulgaris.
Note: Synonyms fide Wollenweber & Reinking (1935) and
Sandoval-Denis et al. (2019).
clavatum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 234. 1915.
(See Fusarium flocciferum)
www.studiesinmycology.org
REDELIMITED
Lectotypus (hic designatus, MBT 10000671): USA, New York,
Castile, rotten tuber of Solanum tuberosum, 1915, C.D. Sherbakoff, in Mem. Cornell Univ. Agric. Exp. Sta. 6: 235, fig. 40.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
clavus Fusarium J.W. Xia et al. (as ‘clavum’), Persoonia 43:
199. 2019.
Holotypus: CBS H-24054.
Ex-type culture: CBS 126202 = RMF N 38.
Type locality: Namibia, northern Karoo, 30 km west of
Maltahöhe.
Type substrate: Desert soil.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170389; tef1: MN170456.
clematidis Fusarium Rolland & Fautrey, Rev. Mycol. (Toulouse)
16: 72. 1894.
Macroconia sphaeriae (Fuckel) Gr€afenhan & Schroers, Stud.
Mycol. 68: 103. 2011.
Basionym: Fusarium sphaeriae Fuckel, Fungi Rhen. Exs. Fasc.
3: no. 212. 1863.
Synonyms: Fusarium sphaeriae var. robustum Davis, Trans.
Wisconsin Acad. Sci. 19: 714. 1919.
Septogloeum robustum (Davis) Wollenw. & Reinking, Fusarien:
336. 1935.
?Nectria leptosphaeriae var. macrospora Wollenw., Angew. Bot.
8: 187. 1926.
Syntype: ILL00220727 (Roumeguere, Fungi Sel. Gall. Exs. no.
6537).
Type locality: France.
Type substrate: Clematis vitalba.
Note: Synonym fide Wollenweber & Reinking (1935).
clypeaster Fusarium (Corda) Sacc., Syll. Fung. 4: 706. 1886.
Septogloeum clypeaster (Corda) Wollenw., Fusarien: 321.
1935.
Basionym: Fusisporium clypeaster Corda, Icon. Fung. 4: 26.
1840.
Lectotypus (hic designatus, MBT 10000672): Czech Republic,
Phragmites, May 1839, A.C.J. Corda, in Icon. Fung. 4, Tab. 6, fig.
82. 1840.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
coccidicola Fusarium Henn. (as ‘coccideicola’), Bot. Jahrb. Syst.
34: 57. 1904.
Microcera diploa (Berk. & M.A. Curtis) Gr€afenhan & Seifert,
Stud. Mycol. 68: 106. 2011.
Basionym: Nectria diploa Berk. & M.A. Curtis, J. Linn. Soc., Bot.
10: 378. 1868.
Synonyms: Cucurbitaria diploa (Berk. & M.A. Curtis) Kuntze,
Revis. Gen. Pl. 3: 461. 1898.
Creonectria diploa (Berk. & M.A. Curtis) Seaver, Mycologia 1:
190. 1909.
Calonectria diploa (Berk. & M.A. Curtis) Wollenw., Angew. Bot. 8:
193. 1926.
Cosmospora diploa (Berk. & M.A. Curtis) Rossman & Samuels,
Stud. Mycol. 42: 121. 1999.
Fusarium derridis Henn., Beibl. Hedwigia 41: (66). 1902.
Fusarium juruanum Henn., Hedwigia 43: 398. 1904.
101
CROUS
ET AL.
Fusarium pentaclethrae Henn., Hedwigia 44: 71. 1905.
Aschersonia henningsii Koord., Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk. 13: 213. 1907.
Microcera henningsii (Koord.) Petch, Ann. Roy. Bot. Gard.
(Peradeniya) 5: 533. 1914.
Pseudomicrocera henningsii (Koord.) Petch, Trans. Brit. Mycol.
Soc. 7: 100. 1921.
Microcera fujikuroi Miyabe & Sawada, J. Fac. Agric. Hokkaido
Imp. Univ. 5: 83. 1913.
Microcera merrillii Syd. & P. Syd., Ann. Mycol. 12(6): 576. 1914.
Pseudomicrocera henningsii var. longispora Petch, Trans. Brit.
Mycol. Soc. 7: 164. 1921.
Fusarium microcera Bilaĭ, Fusarii (Biologija i sistematika): 292.
1955, nom. inval., Art. 39.1.
Holotypus: Zimmerman no. 26 in B fide Hein (1988).
Type locality: Tanzania, East Usambara, Magrotto.
Type substrate: Parasitic on Coccoidea sp. on Camellia sinensis.
coccinellum Fusarium Kalchbr., Flora (Regensburg) 59: 426.
1876.
(See Fusarium coccophilum)
Syntype: ?NY00899913.
Type locality: South Africa, Eastern Cape Province, SomersetEast.
Type substrate: Acacia horrida.
Note: Synonym fide Wollenweber & Reinking (1935).
coccineum Fusarium Schwein., Trans. Amer. Philos. Soc., n.s. 4:
302. 1834.
Holotypus: ?PH00062490.
Type locality: USA, Pennsylvania, Northhampton, Nazareth.
Type substrate: Bark of Castanea sp.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
coccophilum Fusarium (Desm.) Wollenw. & Reinking, Fusarien:
34. 1935.
Microcera coccophila Desm., Ann. Sci. Nat. Bot., ser. 3, 10:
359. 1848.
Synonyms: Tubercularia coccophila (Desm.) Bonord., Abh.
Naturf. Ges. Halle 8: 96. 1864.
Fusarium episphaeria f. coccophilum (Desm.) W.C. Snyder &
H.N. Hansen, Amer. J. Bot. 32: 662. 1945.
Nectria episphaeria f. coccophila (Desm.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 32: 662. 1945.
Fusarium coccinellum Kalchbr., Flora (Regensburg) 56: 426. 1876.
Fusisporium coccinellum (Kalchbr.) Kalchbr., in Thümen, Mycoth.
Univ. no. 782. 1877.
Fusarium cataleptum Cooke & Harkn., Grevillea 12: 96. 1884.
Microcera pluriseptata Cooke & Massee, Grevillea 17: 43. 1888.
Fusarium callosporum Pat., Bull. Soc. Mycol. France 9: 164.
1893.
Fusarium baccharidicola Henn., Hedwigia 48: 20. 1908.
Microcera coccophila var. platyspora Sousa da C^amara, Revista
Agron. (Lisbon): 5 (extr.). 1920.
Lectotypus: K(M) 165807, designated in Gr€afenhan et al. (2011).
Type locality: France, Normandy, near Caen.
Type substrate: Parasitic on Eulecanium tiliae on Salix sp. and
Fraxinus excelsior.
Descriptions and illustrations: See Gr€afenhan et al. (2011).
Notes: No living type material available. Gr€afenhan et al. (2011)
designated a lectotype but did not designate an epitype, which is
still required.
102
coffeatum Fusarium L. Lombard & Crous, Fungal Syst. Evol. 4:
191. 2019.
Replaced synonym: Fusarium chlamydosporum var. fuscum
Gerlach, Phytopathol. Z. 90: 41. 1977.
Holotypus: BBA 62053.
Isotypus: CBS H-631.
Ex-type culture: BBA 62053 = CBS 635.76 = NRRL 20841.
Type locality: South Africa.
Type substrate: Cynodon lemfuensis.
Descriptions and illustrations: See Gerlach (1977a), Gerlach &
Nirenberg (1982) and Xia et al. (2019).
Diagnostic DNA barcodes: rpb1: MN120717; rpb2: MN120736;
tef1: MN120755.
coffeicola Fusarium Henn., Bot. Jahrb. Syst. 22: 82. 1895.
Synonym: Gloeosporium coffeicola (P. Henn.) Wollenw., Fusaria
Autogr. Delin. 1: 493. 1916, nom. illegit., Art. 53.1, non Gloeosporium coffeicola Tassi 1900.
Holotypus: In B fide Hein (1988).
Type locality: Cameroon, Victoria.
Type substrate: Coffea liberica.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
coicis Fusarium Johanssen et al., Fungal Diversity 77: 356.
2015 [2016].
Holotypus: RBG 5368.
Ex-type culture: FRL 19329 = NRRL 66233 = RBG 5368.
Type locality: Australia, Queensland, Mareeba.
Type substrate: Coix gasteenii.
Descriptions and illustrations: See Laurence et al. (2016).
Diagnostic DNA barcodes: rpb1: KP083269; rpb2: KP083274;
tef1: KP083251.
colorans Fusarium (De Jonge) Appel & Wollenw., Arbeiten
Kaiserl. Biol. Anst. Land-Forstw. 8: 39. 1913.
Albonectria rigidiuscula (Berk. & Broome) Rossman & Samuels, Stud. Mycol. 42: 105. 1999.
Basionym: Nectria rigidiuscula Berk. & Broome, J. Linn. Soc.,
Bot. 14: 116. 1873 [1875].
Synonyms: Calonectria rigidiuscula (Berk.
& Broome) Sacc., Michelia 1(3): 313. 1878.
Fusarium rigidiusculum (Berk. & Broome) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 32: 664. 1945.
Calonectria eburnea Rehm, Hedwigia 37: 196. 1898.
Calonectria lichenigena Speg., Bol. Acad. Nac. Ci. Republ.
Argent. 11: 530. 1889.
Calonectria sulcata Starb€ack, Bih. Kongl. Svenska Vetensk.Akad. Handl. 25: 29. 1899.
Fusarium decemcellulare Brick, Jahresber. Vereinigung Angew.
Bot. 6: 227. 1908.
Spicaria colorans De Jonge, Recueil Trav. Bot. Neerl. 6: 48.
1909.
Scoleconectria tetraspora Seaver, N. Amer. Fl. 3: 27. 1910.
Calonectria tetraspora (Seaver) Sacc. & Trotter, Syll. Fung. 22:
487. 1913.
Nectria rigidiuscula f. theobromae E.J. Ford et al., Phytopathology 57: 712. 1967.
Holotypus: Not located.
Type locality: Surinam.
Type substrate: Theobroma cacao.
Notes: Synonym fide Wollenweber & Reinking (1935).
Wollenweber (1916–1935) indicated that cultures and specimens of Spicaria colorans (basionym of F. colorans) were
FUSARIUM
deposited in the Willie Commelin Scholten collection in
Amsterdam. This collection has been accessioned into the CBS
collection (CBS & CBS H). However, no cultures and specimens
or records could be located at CBS.
commune Fusarium K. Skovg. et al., Mycologia 95: 632. 2003.
Holotypus: BBA 71639 in B.
Ex-type culture: AAS 156 = BBA 71639 = CBS 110090 = NRRL
31076.
Type locality: Denmark.
Type substrate: Soil.
Descriptions and illustrations: See Skovgaard et al. (2003).
Diagnostic DNA barcodes: rpb1: MW928803; rpb2: MW934368;
tef1: AF362263.
commutatum Fusarium Sacc., Syll. Fung. 4: 710. 1886.
(See Fusarium solani)
Replaced synonym: Fusisporium candidum Bonord., Handb.
Allg. Mykol.: 96 (1851), nom. illegit., Art. 53.1, non Fusisporium
candidum Link 1824.
Holotypus: Not located.
Type locality: Germany.
Type substrate: Solanum tuberosum.
Note: Synonyms fide Wollenweber & Reinking (1935) and
Sandoval-Denis et al. (2019).
compactum Fusarium (Wollenw.) Raillo, Fungi of the Genus
Fusarium: 180. 1950.
Basionym: Fusarium scirpi var. compactum Wollenw., Fusaria
Autogr. Delin. 3: no. 924. 1930.
Synonym: Fusarium compactum (Wollenw.) Gordon, Canad. J.
Bot. 30: 224. 1952, nom. inval., Art. 53.1.
Lectotypus: Illustration in Wollenweber, Fusaria Autogr. Delin. no.
924 (1930), designated in Xia et al. 2019.
Epitypus: CBS 186.31 (preserved as metabolically inactive culture), designated in Xia et al. (2019).
Ex-epitype culture: CBS 186.31 = NRRL 36323.
Epitype locality: UK, Kew.
Epitype substrate: Cotton thread.
Descriptions and illustrations: See Wollenweber (1916–1935,
no. 924), Raillo (1950), Gordon (1952), Gerlach & Nirenberg
(1982) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb2: GQ505826; tef1: GQ505648.
concentricum Fusarium Nirenberg & O'Donnell, Mycologia 90:
442. 1998.
Holotypus: B 70 0001694.
Ex-type culture: BBA 64354 = CBS 450.97 = DAOM
225146 = IMI 375352 = NRRL 25181.
Type locality: Costa Rica.
Type substrate: Musa sapientum.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998)
and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: LT996192; rpb2: LT575063;
tef1: AF160282.
concolor Fusarium Reinking, Zentralbl. Bakteriol., 2. Abt. 89:
512. 1934.
Synonym: Fusarium polyphialidicum Marasas et al., Mycologia
78: 678. 1986.
Holotypus: IMI 112502.
Ex-type culture: BBA 2607 = BBA 63601 = CBS 183.34 = DAOM
225131 = DSM 62179 = IMI 112502 = NRRL 13994.
www.studiesinmycology.org
REDELIMITED
Type locality: Uruguay, Montevideo.
Type substrate: Hordeum vulgare.
Descriptions and illustrations: See Gerlach & Nirenberg (1982)
and Marasas et al. (1986).
Diagnostic DNA barcodes: rpb1: MH742492; rpb2: MH742569;
tef1: MH742650.
conglutinans Fusarium Wollenw., Ber. Deutsch. Bot. Ges. 31: 34.
1913.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: USA, Wisconsin.
Type substrate: Brassica oleracea var. capitata.
congoense Fusarium Wollenw., Fusaria Autogr. Delin. 1: 307.
1916.
(See Fusarium heterosporum)
Syntype: BPI 451889.
Type locality: Democratic Republic of the Congo.
Type substrate: Bromus willdenowii.
Note: Synonyms fide Wollenweber & Reinking (1935).
coniosporiicola Fusarium Henn., Ann. Mus. Congo Belge, Bot.,
Ser. 5, 2: 106. 1907.
Dendrodochium coniosporiicola (Henn.) Hansf., Proc. Linn.
Soc. London 155: 60. 1943.
Synonym: Fusidium coniosporiicola (Henn.) Wollenw., Fusaria
Autogr. Delin. 1: 477. 1916.
Syntypes: In BR & S.
Type locality: Democratic Republic of the Congo, Gongolo.
Type substrate: Albizia aff. fastigiata.
constrictum Fusarium Penz., Michelia 2: 486. 1882.
Synonym: Ramularia constricta (Penz.) Wollenw., Fusarien: 322.
1935.
Holotypus: Not located; destroyed fide U. Braun.
Type locality: Italy, Padua.
Type substrate: Leaves of Citrus sp.
Notes: Status unclear. Neither Fusarium fide Wollenweber &
Reinking (1935) nor Ramularia (pers. comm. U. Braun).
contaminatum Fusarium L. Lombard & Crous, Persoonia 43:
20. 2018 [2019].
Holotypus: CBS H-23610.
Ex-type culture: CBS 114899.
Type locality: Germany, Schlüchtern.
Type substrate: Pasteurised chocolate milk.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb2: MH484901; tef1: MH484992.
continuum Fusarium X. Zhou et al., Mycologia 108: 677. 2016.
Holotypus: HMNWAFU NX-Ffpl-10-20100851.
Ex-type culture: CBS 140841 = F201030 = NRRL 66286.
Type locality: China, Shaanxi, Fuping, Lei village.
Type substrate: Zanthoxylum bungeanum.
Descriptions and illustrations: See Zhou et al. (2016).
Diagnostic DNA barcodes: rpb1: KM520387; rpb2: KM236782;
tef1: KM236722.
convolutans Fusarium Sand.-Den. et al., MycoKeys 34: 77.
2018.
Holotypus: CBS H-23495.
Ex-type culture: CBS 144207 = CPC 33733.
Type locality: South Africa, Kruger National Park, Skukuza,
Granite Supersite.
103
CROUS
ET AL.
Type substrate: Rhizosphere of Kyphocarpa angustifolia.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018b).
Diagnostic DNA barcodes: rpb1: LT996193; rpb2: LT996141;
tef1: LT996094.
corallinum Fusarium Mattir., Atti Accad. Sci. Ist. Bologna, Cl. Sci.
Fis., Mem. 6: 677. 1897, nom. illegit., Art. 53.1.
(See Fusarium culmorum)
Authentic material: Not located.
Type locality: Italy.
Type substrate: Andropogon sp.
Note: Synonym fide Wollenweber (1931).
corallinum Fusarium Sacc., Nuovo Giorn. Bot. Ital. 8: 196. 1876.
(See Fusarium graminum)
Holotypus: In PAD.
Type locality: Italy, Treviso, Selva.
Type substrate: Cynodon dactylon.
cordae Fusarium Massee, Brit. Fung.-Fl. 3: 481. 1893.
(See Fusarium oxysporum)
Notes: Massee introduced this name to replace F. aurantiacum
Corda, indicating that F. aurantiacum (Link) Sacc., based on
Fusisporium aurantiacum Link (1809), predates Corda's use of
the epithet. However, Corda's use of the epithet in Fusarium
predates Saccardo's recombination into Fusarium.
cortaderiae Fusarium O'Donnell et al., Fungal Genet. Biol. 41:
620. 2004.
Holotypus: BPI 843479.
Ex-type culture: CBS 119183 = ICMP 5435 = NRRL 29297.
Type locality: New Zealand, Auckland, Henderson.
Type substrate: Cortaderia selloana.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: KM361644; rpb2: KM361662;
tef1: AY225885.
crassistipitatum Fusarium Scandiani et al., Mycoscience 53: 171.
2011.
(See Fusarium azukiicola)
Holotypus: BPI 871490.
Ex-type culture: MAFF 239757 = NRRL 36877.
Type locality: Argentina, Santa Fe, Zavalla.
Type substrate: Glycine max.
Descriptions and illustrations: See Aoki et al. (2012a).
Diagnostic DNA barcodes: rpb2: FJ240405; tef1: FJ240351.
crassum Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 1. 2020.
Neocosmospora crassa Sand.-Den. & Crous, Persoonia 43:
122. 2019.
Holotypus: CBS H-23976.
Ex-type culture: CBS 144386 = MUCL 11420.
Type locality: France, Paris.
Type substrate: Unknown.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218109; rpb2: LR583823;
tef1: LR583604.
croceum Fusarium J.W. Xia et al., Persoonia 43: 201. 2019.
Holotypus: CBS H-24055.
Ex-type culture: CBS 131777.
Type locality: Iran, Golestan Province, Gonbad-e Qabus.
Type substrate: Triticum sp.
104
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170396; tef1: MN170463.
croci Fusarium (Guarnaccia, Sand.-Den. & Crous) O'Donnell
et al., Index Fungorum 440: 1. 2020.
Basionym: Neocosmospora croci Guarnaccia, Sand.-Den. &
Crous, Persoonia 40: 17. 2017.
(See Fusarium citrulli Sartory)
Holotypus: CBS H-23022.
Ex-type culture: CBS 142423 = CPC 27186.
Type locality: Italy, Sicily, Catania, Paterno.
Type substrate: Citrus sinensis.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018a).
Diagnostic DNA barcodes: rpb2: LT746329; tef1: LT746216.
cromyophthoron Fusarium Sideris, Phytopathology 14: 212.
1924.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000673): USA, California,
Stockton, roots of Allium sp.,1924, C.P. Sideris, in Phytopathology 14, pl. IX.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
crookwellense Fusarium L.W. Burgess et al., Trans. Brit. Mycol.
Soc. 79: 498. 1982.
(See Fusarium cerealis (Cooke) Sacc.)
Holotypus: FRC R-3090.
Ex-type culture: NRRL 13163.
Type locality: Australia, New South Wales, Crookwell.
Type substrate: Solanum tuberosum tubers.
Descriptions and illustrations: See Burgess et al. (1982).
Note: See Notes under F. cerealis.
cruentum Fusarium Teich, Byull. Sredne-Aziatsk. Gosud. Univ.
19: 178. 1934.
Holotypus: Not located.
Type locality: Uzbekistan, Tashkent.
Type substrate: Roots and stems of Vitis vinifera.
Notes: Status unclear. This species was not treated by any of
Wollenweber & Reinking (1935), Raillo (1950), Bilaĭ (1955),
Booth (1971), Joffe (1974), or Gerlach & Nirenberg (1982).
Furthermore, no additional records could be located.
cryptoseptatum Fusarium (Sand.-Den. & Crous) O'Donnell, Index Fungorum 440: 1. 2020.
Neocosmospora cryptoseptata Sand.-Den. & Crous, Persoonia 43: 122. 2019.
Holotypus: CBS H-23977.
Ex-type culture: BBA 65024 = CBS 145463 = NRRL 22412.
Type locality: French Guiana.
Type substrate: Bark.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834215; rpb2: EU329510;
tef1: AF178351.
cryptum Fusarium McAlpine, Fungus Diseases of Citrus trees in
Australia: 106. 1899.
(See Fusarium larvarum)
Holotypus: VPRI 2557.
Type locality: Australia, South Australia.
Type substrate: Twigs of Citrus limonia.
FUSARIUM
Note: Synonym fide Wollenweber & Reinking (1935).
cubense Fusarium E.F. Sm., Science, N.Y. 31: 754. 1910.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: Cuba.
Type substrate: Musa sp.
Note: Synonym fide Wollenweber & Reinking (1935).
cucumerinum Fusarium Berk. & Broome, Ann. Mag. Nat. Hist.,
ser. 4, 17: 141. 1876.
Holotypus: ?K(M).
Type locality: UK, Northamptonshire, Daventry, Sibbertoft.
Type substrate: Diseased Cucumis sativus.
Notes: Status unclear. Wollenweber & Reinking (1935) synonymised this species under Septomyxa persicina. In contrast, Index
Fungorum indicates that this species is a synonym under
F. oxysporum. The original protologue (Berkeley & Broome 1876)
fits neither S. persicina nor F. oxysporum.
cucurbitae Fusarium Taubenh., Bull. Texas Agric. Exp. Sta. 260:
27. 1920.
Lectotypus (hic designatus, MBT 10000674): USA, Texas, Waller
County, from squash, date unkown, J.J. Taubenhaus, in Bull.
Texas Agric. Exp. Sta. 260: 30, fig. 8j. 1920.
Notes: Based on the description and illustrations provided by
Taubenhaus (1920), this species could represent F. oxysporum.
However, recollection and epitypification are required to confirm
this. No holotype specimen could be located and therefore an
illustration is designated as lectotype.
cucurbitariae Fusarium (Pat.) Sacc., Syll. Fung. 4: 708. 1886.
(See Fusarium avenaceum)
Basionym: Fusisporium cucurbitariae Pat., Rev. Mycol. (Toulouse) 3: 10. 1881.
Holotypus: ?FH01093588.
Type locality: France, Lons-le-Saunier.
Type substrate: Diseased Cucumis sativus.
Note: Synonym fide Wollenweber & Reinking (1935).
cucurbitariae Fusarium Peyronel, Nuovo Giorn. Bot. Ital., n.s. 25:
436. 1918, nom. illegit., Art. 53.1.
Holotypus: ?ROPV.
Type locality: Italy, Piemonte, Riclaretto.
Type substrate: Parasitic on perithecia of Camarosporidiella
laburni (≡ Cucurbitaria laburni).
Notes: Status unclear. Not treated by any of Wollenweber &
Reinking (1935), Booth (1971), or Gerlach & Nirenberg (1982).
cucurbiticola Fusarium O'Donnell et al., Index Fungorum 440: 2.
2020.
Neocosmospora cucurbitae Sand.-Den. et al., Persoonia 43:
125. 2019.
Synonyms: Fusarium solani f. cucurbitae W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 28: 740. 1941.
Fusarium solani f. sp. cucurbitae W.C. Snyder & H.N. Hansen,
Root rots caused by Phycomycetes 28: 740. 1941.
Hypomyces solani f. cucurbitae W.C. Snyder & H.N. Hansen,
Amer. J. Bot. 28: 741. 1941.
Nectria haematococca var. cucurbitae (W.C. Snyder & H.N.
Hansen) Dingley, New Zealand J. Agric. Res. 4: 337. 1961.
Nectria solani f. cucurbitae (W.C. Snyder & H.N. Hansen) G.R.W.
Arnold, Z. Pilzk. 37: 193. 1972.
www.studiesinmycology.org
REDELIMITED
Holotypus: CBS H-23978.
Ex-type culture: BBA 64411 = CBS 616.66 = NRRL 22399.
Type locality: Netherlands.
Type substrate: Cucurbita viciifolia.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834217; rpb2: LR583825;
tef1: DQ247592.
cugenangense Fusarium Maryani et al., Stud. Mycol. 92: 181.
2018 [2019].
Holotypus: InaCC F984 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F984.
Type locality: Indonesia, West Java, Cianjur, Cugenang.
Type substrate: Pseudostem of Musa var. Pisang Kepok.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479560; rpb2: LS479308;
tef1: LS479757.
culmorum Fusarium (Wm.G. Sm.) Sacc., Syll. Fung. 10: 726.
1892.
Basionym: Fusisporium culmorum Wm.G. Sm., Diseases of field
and garden crops, chiefly as are caused by fungi: 209. 1884.
Synonyms: Fusarium schribauxii Delacr., Bull. Soc. Mycol.
France 6: 99. 1890.
Fusarium corallinum Mattir., Atti Accad. Sci. Ist. Bologna, Cl. Sci.
Fis., Mem. 6: 677. 1897, nom. illegit., Art. 53.1.
Fusarium versicolor Sacc., Syll. Fung. 16: 1099. 1902.
Fusarium heidelbergense Sacc., Ann. Mycol. 8: 346. 1910.
?Fusarium neglectum Jacz., Bull. Trimestriel Soc. Mycol.
France 28: 348. 1912.
Fusarium rubiginosum Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land-Forstw. 8: 108. 1910 [1913].
Fusarium culmorum var. leteius Sherb., Mem. Cornell Univ.
Agric. Exp. Sta. 6: 242. 1915.
Fusarium culmorum var. majus Wollenw., Fusaria Autogr. Delin.
2: 613. 1924.
Lectotypus (hic designatus, MBT 10000675): UK, infected ear of
Triticum sp., 1884, W.G. Smith, in Diseases of field and garden
crops, chiefly as are caused by fungi: 210. fig. 92.
Epitypus (hic designatus, MBT 10000676): Denmark, moldy
kernel of Hordeum vulgare, 3 Feb. 1986, U. Thrane, CBS 417.86
(preserved as metabolic inactive culture).
Ex-epitype culture: CBS 417.86 = FRC R-8504 = IMI 309344=
NRRL 25475.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982) and Leslie
& Summerell (2006).
Diagnostic DNA barcodes: rpb1: JX171515; rpb2: JX171628;
tef1: MW233082.
Notes: No holotype specimen could be located. Therefore, an
illustration is designated as lectotype and CBS 417.86 is
designated as epitype as this isolate is commonly used as an
authentic strain for F. culmorum in literature (Ward et al. 2002,
O'Donnell et al. 2013, 2020, Geiser et al. 2021).
cuneiforme Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 129. 1915.
(See Fusarium ventricosum)
Typus: ?CUP-007474.
Type locality: USA, New York.
Type substrate: Solanum tuberosum.
105
CROUS
ET AL.
Notes: Synonym fide Wollenweber & Reinking (1935) and Booth
(1971). Lectotypification pending study of material lodged in
CUP.
cuneirostrum Fusarium O'Donnell & T. Aoki, Mycoscience 46:
170. 2005.
(See Fusarium azukiicola)
Holotypus: BPI 843353.
Ex-type culture: FRC S-1551 = MAFF 239038 = NRRL 31157.
Type locality: USA, Michigan, Presque Isle.
Type substrate: Phaseolus vulgaris.
Descriptions and illustrations: See Aoki et al. (2005).
Diagnostic DNA barcodes: rpb1: KJ511271; rpb2: FJ240389;
tef1: MAEA01003816.
curvatum Fusarium L. Lombard & Crous, Persoonia 43: 21.
2018 [2019].
Holotypus: CBS H-23611.
Ex-type culture: CBS 238.94 = NRRL 26422 = PD 94/184.
Type locality: Netherlands.
Type substrate: Beaucarnea sp.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928804; rpb2: MH484893;
tef1: MH484984.
cuticola Fusarium (R. Blanch.) Gueg., Champ. Paras. Homme:
262. 1904.
(See Fusarium oxysporum)
Basionym: Selenosporium cuticola R. Blanch., Compt. Rend.
Hebd. Seances Acad. Sci. 111: 479. 1890.
Holotypus: Not located.
Type locality: France.
Type substrate: Skin of Chamaeleo vulgaris and Lacerta viridis
(lizards).
Notes: Synonym fide Wollenweber & Reinking (1935). Based on
the substrate, this species could belong to the genus Bisifusarium. However, the protologue is not definitive, and recollection
from type substrate is needed to confirm its taxonomic position.
cyanescens Fusarium (G.A. de Vries et al.) O'Donnell et al.,
Index Fungorum 440: 2. 2020.
Neocosmospora cyanescens (G.A. de Vries et al.) Summerb.
et al., Biology of Microfungi (Cham): 183. 2016.
Basionym: Phialophora cyanescens G.A. de Vries et al., Antonie
van Leeuwenhoek 50: 150. 1984.
Synonyms: Cylindrocarpon cyanescens (G.A. de Vries et al.)
Sigler, J. Clin. Microbiol. 29: 1858. 1991.
Holotypus: CBS 518.82 (maintained as metabolically inactive
culture).
Ex-type culture: CBS 518.82.
Type locality: Netherlands, Groningen Province, Groningen.
Type substrate: Subcutaneous tissue of the right foot of a male
Homo sapiens.
Descriptions and illustrations: See de Vries et al. (1984) and
Zoutman & Sigler (1991).
Diagnostic DNA barcodes: rpb1: MW218110; rpb2: LR583826;
tef1: LR583605.
cyanostomum Fusarium (Sacc. & Flageolet) O'Donnell & Geiser,
Phytopathology 103: 404. 2013.
Cyanonectria cyanostoma (Sacc. & Flageolet) Samuels & P.
Chaverri, Mycol. Progr. 8: 56. 2009.
106
Basionym: Nectria cyanostoma Sacc. & Flageolet, Rendiconti
Congr. Bot. Palermo 1902: 53. 1902.
Lectotypus: BPI 551652, designated in Samuels et al. (2009).
Epitypus: BPI 748307, designated in Samuels et al. (2009).
Ex-epitype culture: BBA 70964 = CBS 101734 = G.J.S. 98-127.
Epitype locality: France.
Epitype substrate: Buxus sempervirens.
Descriptions and illustrations: See Samuels et al. (2009).
Diagnostic DNA barcodes: rpb1: JX171546; rpb2: HQ897759;
tef1: HM626647.
cyclogenum Fusarium Sacc., Nuovo Giorn. Bot. Ital. 8: 197.
1876.
?Gloeosporium orbiculare (Berk.) Berk., Just's Bot. Jahresber.
4: 1274. 1876.
Basionym: Cytospora orbicularis Berk., Ann. Nat. Hist. 1: 207.
1838.
Synonyms: Myxosporium orbiculare (Berk.) Berk., Outl. Brit.
Fungol.: 325. 1860.
Colletotrichum orbiculare (Berk.) Arx, Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk., Sect. 2, 51: 112. 1957, nom. inval., Art.
36.2 (Melbourne).
Sirogloea orbicularis (Berk.) Arx, Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk., Sect. 2, 51: 113. 1957, nom. inval., Art.
36.2 (Melbourne).
Syntypes: In BPI & S.
Type locality: Italy, Treviso, Selva.
Type substrate: Citrullus sp.
Note: Cytospora orbicularis is not a Colletotrichum nor a Fusarium (small ellipsoidal conidia discharged in tendrils) as outlined
in Damm et al. (2013).
cydoniae Fusarium Allesch., Ber. Bot. Vereines Landshut 12:
130. 1892.
(See Fusarium lateritium)
Holotypus: In M.
Type locality: Germany, München.
Type substrate: Cydonia vulgaris.
Note: Synonym fide Wollenweber & Reinking (1935).
cydoniae Fusarium Roum. & Fautrey, Rev. Mycol. (Toulouse) 14:
170. 1892, nom. illegit., Art. 53.1, non Allescher 1892.
(See Fusarium rollandianum)
cydoniae Fusarium (Schulzer) Sacc. & Traverso, Syll. Fung. 19:
724. 1910, nom. illegit., Art. 53.1, non Allescher 1892, nec Roum.
& Fautrey 1892.
Basionym: Selenosporium cydoniae Schulzer, Verhand. K.K.
Zool.-Bot. Ges. Wien 21: 1240. 1871.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Austria, Vienna.
Type substrate: Cydonia vulgaris.
Note: Synonyms fide Wollenweber & Reinking (1935).
cylindricum Fusarium (Mont.) Sacc., Syll. Fung. 4: 720. 1886.
Basionym: Fusisporium cylindricum Mont., Ann. Sci. Nat., Bot.,
ser. 2, 17: 120. 1842.
(See Fusarium candidum (Link) Sacc.)
Holotypus: ?PC.
Type locality: Cuba.
Type substrate: Sarcocarp of unknown fruit.
FUSARIUM
Note: Synonyms fide Wollenweber & Reinking (1935).
cymbiferum Fusarium Berk. & M.A. Curtis, in Berkeley, Grevillea
3: 98. 1875.
Colletotrichum coccodes (Wallr.) S. Hughes, Canad. J. Bot.
36: 754. 1958.
Basionym: Chaetomium coccodes Wallr., Fl. Crypt. Germ. 2:
265. 1833.
Synonyms: Fusarium effusum Schwein., Trans. Amer. Philos.
Soc., n.s. 4: 302. 1832 [1834].
Fusarium georginae Corda, Icon. Fung. 2: 4. 1838.
Vermicularia atramentaria Berk. & Broome, Ann. Mag. Nat. Hist.
5: 378. 1850.
Colletotrichum atramentarium (Berk. & Broome) Taubenh., Mem.
New York Bot. Gard. 6: 554. 1916.
Acrothecium solani Sacc., Michelia 1(3): 74. 1877.
Fusisporium elasticae Thüm., Boll. Soc. Adriat. Sci. Nat. Trieste
3: 440. 1877.
Fusarium elasticae (Thüm.) Sacc., Syll. Fung. 4: 711. 1886.
Gloeosporium elasticae Cooke & Massee, in Cooke, Grevillea
18: 74. 1890.
Fusarium foliicola Allesch., Hedwigia 34: 289. 1895.
Gloeosporium foliicola (Allesch.) Wollenw., Fusarien: 325. 1935,
nom. illegit., Art. 53.1.
Colletotrichum solanicola O'Gara, Mycologia 7: 39. 1915.
Colletotrichum biologicum Chaudhuri, Ann. Bot. 38: 735. 1924.
Holotypus: ?K(M).
Type locality: USA.
Type substrate: Stems of some herbaceous plants.
Note: Synonyms fide Wollenweber & Reinking (1935).
cypericola Fusarium Henn., Hedwigia 48: 116. 1908.
Libertella cypericola (Henn.) Wollenw., Fusaria Autogr. Delin.
1: 486. 1916.
Syntype: In B fide Hein (1988).
Type locality: Brazil, Para.
Type substrate: Cyperus exaltatus.
Note: Synonym fide Wollenweber & Reinking (1935).
dactylidis Fusarium T. Aoki et al., Mycologia 107: 412. 2015.
Holotypus: BPI 892886.
Ex-type culture: CBS 119181 = ICMP 5269 = NRRL 29298.
Type locality: New Zealand, Manawatu, Palmerston North.
Type substrate: Dactylis glomerata.
Descriptions and illustrations: See Aoki et al. (2015).
Diagnostic DNA barcodes: rpb1: KM361654; rpb2: KM361672;
tef1: DQ459748.
decemcellulare Fusarium Brick, Jahresber. Vereinigung Angew.
Bot. 6: 227. 1908.
(See Fusarium colorans)
Holotypus: ?HBG.
Type locality: Cameroon.
Type substrate: Theobroma cacao.
decipiens Fusarium Cooke & Massee, in Cooke, Handb. Austral.
Fungi: 388. 1892, nom. inval., Art. 39.1.
(See Fusarium candidum (Link) Sacc.)
Authentic material: ?K(M).
Original locality: Australia, Queensland.
Original substrate: Ficus aspera.
Note: Synonym fide Wollenweber & Reinking (1935).
deformans Fusarium J. Schröt., Jahresber. Schles. Ges. Vaterl.
Cult. 61: 179. 1883.
www.studiesinmycology.org
REDELIMITED
Gloeosporium deformans (J. Schröt.) Lind, Ann. Bot. 7: 19.
1908.
Synonyms: Fusamen deformans (J. Schröt.) P. Karst., Bidrag
K€annedom Finlands Natur Folk 51: 485. 1892.
Calogloeum deformans (J. Schröt.) Nannf., Svensk Bot. Tidskr.
25: 25. 1931.
Platycarpium deformans (J. Schröt.) Petr., Sydowia 7: 296. 1953.
Holotypus: In B fide Wollenweber (1916–1935).
Type locality: Poland, Breslau.
Type substrate: Salix cinerea.
Note: Synonyms fide Wollenweber & Reinking (1935).
delacroixii Fusarium Sacc., Syll. Fung. 10: 725. 1892.
(See Fusarium sambucinum)
Replaced synonym: Fusarium asparagi Delacr., Bull. Soc. Mycol.
France 6: 99. 1890, nom. illegit., Art. 53.1, non Fusarium
asparagi Briard 1890.
Lectotypus (hic designatus, MBT 10000677): France, Paris,
Asparagus officinalis, 1890, M.G. Delaroix, in Bull. Soc. Mycol.
France 6, pl. XV. fig. III.
Notes: Synonyms fide Wollenweber & Reinking (1935). No holotype material is available for the replaced synonym F. asparagi
Delacr. and therefore, an illustration from the original protologue
is designated as lectotype.
delphinoides Fusarium Schroers et al., Mycologia 101: 57. 2009.
Bisifusarium delphinoides (Schroers et al.) L. Lombard &
Crous, Stud. Mycol. 80: 224. 2015.
Holotypus: CBS H-20124.
Ex-type culture: CBS 120718 = NRRL 53290.
Type locality: South Africa, Western Cape Province, Clanwilliam.
Type substrate: Hoodia gordonii stem lesions.
Descriptions and illustrations: See Schroers et al. (2009).
Diagnostic DNA barcodes: rpb1: KM232210; tef1: EU926296.
denticulatum Fusarium Nirenberg & O'Donnell, Mycologia 90:
445. 1998.
Holotypus: B 70 0001691.
Ex-type culture: BBA 67772 = CBS 407.97 = IMI 376115 = NRRL
25311.
Type locality: USA, Louisiana.
Type substrate: Ipomoea batatas.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998)
and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: MT010953; rpb2: MT010970;
tef1: KR909385.
derridis Fusarium Henn., Beibl. Hedwigia 41: (66). 1902.
(See Fusarium coccidicola)
Holotypus: In B fide Hein (1988).
Type locality: Papua New Guinea.
Type substrate: Derris sp.
Note: Synonym fide Wollenweber & Reinking (1935).
desaboruense Fusarium N. Maryani et al., Persoonia 43: 59.
2019.
(See Fusarium sacchari)
Holotypus: InaCC F951 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F951.
Type locality: Indonesia, East Nusa Tenggara, Sikka Flores,
Kecamatan Waigate, Desa Boru.
Type substrate: Musa var. Pisang Kepok.
Descriptions and illustrations: See Maryani et al. (2019b).
107
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ET AL.
Diagnostic DNA barcodes: rpb1: LS479870; rpb2: LS479852.
desciscens Fusarium Oudem., Ned. Kruidk. Arch., 2 ser., 5: 515.
1889.
(See Fusarium sarcochroum)
Holotypus: ?L.
Type locality: Netherlands, Zuid-Holland Province, Den Haag,
Scheveningen.
Type substrate: Sarothamnus vulgaris.
Note: Synonym fide Wollenweber & Reinking (1935).
detonianum Fusarium Sacc. (as ‘de-tonianum’), Syll. Fung. 4:
708. 1886, nom. illegit., Art. 52.1.
(See Fusarium miniatum Sacc.)
Authentic material: Not located.
Original locality: Italy.
Original substrate: Sporangium of Cyathus vernicosa.
dianthi Fusarium Prill. & Delacr., Compt. Rend. Hebd. Seances
Acad. Sci. 129: 745. 1899.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: France, Antibes.
Type substrate: Dianthus caryophyllus.
didymum Fusarium (Harting) Lindau, Rabenh. Krypt.-Fl. Ed. 2,
1(9): 574. 1909.
Basionym: Fusisporium didymum Harting, Nieuwe Verh. Eerste
Kl. Kon. Ned. Inst. Wetensch. Amsterdam 12: 228. 1846.
(See Fusarium eichleri)
Lectotypus (hic designatus, MBT 10000678): Netherlands, Solanum tuberosum, date unknown, Harting, in Nieuwe Verh.
Eerste Kl. Kon. Ned. Inst. Wetensch. Amsterdam 12 (1846), tab.
II, figs 2–4.
Notes: Requires recombination into Neonectria after further
investigation. No preserved specimen could be located and
therefore an illustration is designated as lectotype.
diffusum Fusarium Carmich., Grevillea 16: 81. 1888.
(See Fusarium avenaceum)
Holotypus: ?K(M).
Type locality: UK, Scotland, Appin.
Type substrate: Stems of Asteraceae (thristle).
Note: Synonym fide Wollenweber & Reinking (1935).
dimerum Fusarium Penz., Michelia 2: 484. 1882.
Bisifusarium dimerum (Penz.) L. Lombard & Crous, Stud.
Mycol. 80: 225. 2015.
Synonyms: Fusarium aquaeductuum var. dimerum (Penz.)
Raillo, Fungi of the Genus Fusarium: 279. 1950.
Microdochium dimerum (Penz.) Arx, Trans. Brit. Mycol. Soc. 83:
374. 1984.
?Fusisporium flavum Fr., Syst. Mycol. 3: 444. 1832.
?Pionnotes flava (Fr.) Sacc., Syll. Fung. 4: 726. 1886.
?Fusarium flavum (Fr.) Wollenw., Z. Parasitenk. 3: 305. 1931.
?Fusarium aquaeductuum var. flavum (Fr.) Raillo, Fungi of the
Genus Fusarium: 280. 1950.
Selenosporium aurantiacum Bonord., Abh. Naturf. Ges. Halle 8:
97. 1864, nom. illegit., Art. 53.1.
Fusarium bonordenii Sacc., Syll. Fung. 4: 699. 1886.
Fusarium baptisiae Henn., Notizbl. Bot. Gart. Berlin 2: 383. 1899.
Fusarium subnivale Höhn., in Penther & Zederbauer, Ann. K.K.
Naturhist. Hofmus. 20: 369. 1905.
Fusarium dimerum var. majusculum Wollenw., Fusaria Autogr.
Delin. 1: 90. 1916.
108
?Fusarium pusillum Wollenw., Fusaria Autogr. Delin. 2: 550. 1924.
?Fusarium dimerum var. pusillum (Wollenw.) Wollenw., Fusaria
Autogr. Delin. 3: 851. 1930.
Fusarium dimerum var. violaceum Wollenw., Fusaria Autogr.
Delin. 3: 854. 1930.
Lectotypus: Fig. 1212 in Penzig (1882), designated in Schroers
et al. (2009).
Epitypus: CBS H-20129, designated in Schroers et al. (2009).
Ex-epitype culture: CBS 108944 = NRRL 36140.
Epitype locality: Netherlands.
Epitype substrate: Blood of Homo sapiens with acute myeloid
leukemia.
Descriptions and illustrations: See Schroers et al. (2009).
Diagnostic DNA barcodes: rpb1: KM232212; rpb2: KM232363;
tef1: EU926334.
Note: Synonyms fide Wollenweber & Reinking (1935) and Booth
(1971).
diminutum Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index Fungorum 440: 2. 2020.
Neocosmospora diminuta Sand.-Den. & Crous, Persoonia 43:
127. 2019.
Holotypus: CBS H-23979.
Ex-type culture: CBS 144390 = MUCL 18798.
Type locality: ?Ivory Coast.
Type substrate: Treated wood of Coelocaryon preussii.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834218; rpb2: LR583828;
tef1: LR583607.
dimorphum Fusarium J.V. Almeida & Sousa da C^amara, Revista
Agron. (Lisbon) 1: 306. 1903.
(See Fusarium buxicola)
Holotypus: MA-Funhist:6036-1.
Type locality: Portugal.
Type substrate: Buxus sempervirens.
Note: Synonym fide Wollenweber & Reinking (1935).
diplosporum Fusarium Cooke & Ellis, Grevillea 7: 38. 1878.
(See Fusarium sarcochroum)
Holotypus: ?K(M).
Type locality: USA, New Jersey.
Type substrate: Stems of Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935).
discoideum Fusarium Fautrey & Roum., Rev. Mycol. (Toulouse)
13: 173. 1891.
(See Fusarium lateritium)
Syntype: ILL00220061 (Roumeguere, Fungi Sel. Gall. Exs. no.
5898).
Type locality: France, Noidan.
Type substrate: Sambucus nigra.
Note: Synonym fide Wollenweber & Reinking (1935).
discolor Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol. Anst.
Land-Forstw. 8: 114. 1913.
(See Fusarium sambucinum)
Holotypus: ?S-F45617.
Type locality: Germany, Berlin.
Type substrate: Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935).
diversisporum Fusarium Sherb., Mem. Cornell Univ. Agric.
Exp. Sta. 6: 161. 1915.
FUSARIUM
Typus: ?CUP-007430.
Type locality: USA, New York.
Type substrate: Solanum tuberosum
Descriptions and illustrations: See Sherbakoff (1915) and
Gerlach & Nirenberg (1982).
Notes: This species is recognised by Gerlach & Nirenberg (1982)
who considered isolate CBS 795.70 as authentic for
F. diversisporum. However, typification of F. diversisporum first
requires study of the specimen lodged in CUP.
dlaminii Fusarium Marasas et al., Mycologia 77: 971. 1986
[1985].
Holotypus: DAOM 191112.
Ex-type culture: ATCC 58097 = BBA 69859 = CBS
175.88 = DAOM 191112 = FRC M-1637 = IMI 290241 = MRC
3032 = NRRL 13164.
Type locality: South Africa, Eastern Cape Province, Butterworth.
Type substrate: Plant debris in soil.
Descriptions and illustrations: See Marasas et al. (1985) and
Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: KU171681; rpb2: KU171701;
tef1: KU171721.
domesticum Fusarium (Fr.) H.P. Bachm., LWT – Food Sci.
Technol. 38: 405. 2005, nom. inval., Art. 41.5, See Art. 41.7.
Bisifusarium domesticum (Fr.) L. Lombard & Crous, Stud.
Mycol. 80: 225. 2015.
Basionym: Trichothecium domesticum Fr., Syst. Mycol. 3: 427.
1832.
Neotypus: CBS 434.34 (preserved as metabolically inactive
culture), designated in Bachmann et al. (2005).
Ex-neotype culture: ATCC 13417 = CBS 434.34 = MUCL 9826.
Type locality: Belgium.
Type substrate: Cheese.
Descriptions and illustrations: See Schroers et al. (2009).
dominicanum Fusarium Cif., Sydowia 9: 325. 1955
Holotypus: ?PAV.
Type locality: Dominican Republic, Santo Domingo, Villa
Altagracia.
Type substrate: Byrsonima sp. (between mycelium of Meliola
byrsonimae).
Descriptions and illustrations: See Ciferri (1955).
Notes: Ciferri (1955) considered this a ‘conventional’ species as
the author indicated that more information based on culture
characteristics is required. No living material of this species could
be located and recollection from the type locality is required.
duofalcatisporum Fusarium J.W. Xia et al., Persoonia 43: 201.
2019.
Holotypus: CBS H-24056.
Ex-type culture: CBS 384.92 = NRRL 36448.
Type locality: Sudan, Nile Province.
Type substrate: Seeds of Phaseolus vulgaris.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: GQ505830; tef1: GQ505652.
duoseptatum Fusarium Maryani et al., Stud. Mycol. 92: 181.
2018 [2019].
Holotypus: InaCC F916 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F916.
www.studiesinmycology.org
REDELIMITED
Type locality: Indonesia, Central Kalimantan, Kapuas Timur,
Anjir Serapat Tengah.
Type substrate: Pseudostem of Musa var. Pisang Kepok.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479495; rpb2: LS479239;
tef1: LS479688.
echinatum Fusarium Sand.-Den. & G.J. Marais, Stud. Mycol.
98 (no. 100116): 47. 2021.
Holotypus: CBS H-24658.
Ex-type culture: CAMS 000733 = CBS 146497 = CPC 30815.
Type locality: South Africa.
Type substrate: Unidentified tree.
Descriptions and illustrations: See this study.
Diagnostic DNA barcodes: rpb1: MW834187; rpb2: MW834004;
tef1: MW834273.
echinosporum Fusarium Sibilia, Ann. Reale. Ist. Super. Agrar.
Forest., ser. 2, 1: 77. 1925.
Holotypus: Not located.
Type locality: Italy.
Type substrate: Cedrus deodara.
Descriptions and illustrations: See Sibilia (1925).
Notes: This species is recognised in Petrak's Lists V. 3.
Wollenweber & Reinking (1935) mention this species, but they
did not treat it any further. Booth (1971) considered it a possible
synonym of F. graminearum. Requires recollection from the type
locality and substrate.
effusum Fusarium Schwein., Trans. Amer. Philos. Soc., n.s., 4:
302. 1832 [1834].
(See Fusarium cymbiferum)
Holotypus: PH00062491.
Type locality: USA, Pennsylvania, Northampton, Bethlehem.
Type substrate: Hypericum frondosum.
Note: Synonym fide Wollenweber & Reinking (1935).
eichleri Fusarium Bres., Ann. Mycol. 1: 130. 1903.
(See Fusarium candidum Ehrenb.)
Holotypus: S-F45618.
Type locality: Poland.
Type substrate: Salix caprea.
elaeidis Fusarium L. Lombard & Crous, Persoonia 43: 23. 2018
[2019].
Holotypus: CBS H-23612.
Ex-type culture: CBS 217.49 = NRRL 36358.
Type locality: Zaire.
Type substrate: Elaeis sp.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928805; rpb2: MH484870;
tef1: MH484961.
elasticae Fusarium (Thüm.) Sacc., Syll. Fung. 4: 711. 1886.
Basionym: Fusisporium elasticae Thüm., in Bolle & Thümen,
Boll. Soc. Adriat. Sci. Nat. Trieste 3: 440. 1877.
(See Fusarium cymbiferum)
Lectotypus (hic designatus, MBT 10000679): Italy, Gorizia, Ficus
elastica, 1877, F. de Thümen, in Bolle & Thümen, Boll. Soc.
Adriat. Sci. Nat. Trieste 3, tab. I, fig. 13.
Notes: Synonyms fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
109
CROUS
ET AL.
elegans Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol. Anst.
Land-Forstw. 8: 94. 1913, nom. inval., Art. 36.1(a).
(See Fusarium oxysporum)
Notes: Appel and Wollenweber (l.c.) proposed this name only
provisionally under Fusarium solani. They added an illustration of
conidia on page 38 (fig. 2D).
illegit., Art. 53.1.
(See Fusarium sambucinum)
Authentic material: Not located.
Original locality: USA, Idaho.
Original substrate: Soil.
Note: Synonym fide Wollenweber & Reinking (1935).
elegans Fusarium W. Yamam. & Maeda, Trans. Mycol. Soc.
Japan 3: 115. 1962.
Neocosmospora elegans (W. Yamam. & Maeda) Sand.-Den. &
Crous, Persoonia 43: 127. 2019.
Basionym: Nectria elegans W. Yamam. & Maeda, Hyogo Univ.
Agric. ser. Agric. Biol. 3: 15. 1957.
Synonyms: ?Fusarium solani f. xanthoxyli Y. Sakurai & Matuo,
Ann. Phytopathol. Soc. Japan 26: 117. 1961, nom. inval., Art.
39.1.
?Hypomyces solani f. xanthoxyli Y. Sakurai & Matuo, Ann.
Phytopathol. Soc. Japan 26: 117. 1961, nom. inval., Art. 39.1.
Fusarium yamamotoi O'Donnell et al., Index Fungorum 440: 5.
2020.
Lectotypus: figs 1–9, p. 16, in Yamamoto et al. (1957), designated in Sandoval-Denis et al. (2019).
Epitypus: CBS H-23980, designated in Sandoval-Denis et al.
(2019).
Ex-epitype culture: ATCC 42366 = CBS 144396 = MAFF
238541 = NRRL 22277 = SUF XV-1.
Epitype locality: Japan, Hyogo.
Epitype substrate: Trunk of Zanthoxylum piperitum
Diagnostic DNA barcodes: rpb1: MW218113; rpb2: FJ240380;
tef1: AF178336
Note: This is a valid species name that is not a
homonym since the name F. elegans Appel & Wollenw. is an
invalid name.
elongatum Fusarium Reinking, Zentralbl. Bakteriol. Parasitenk.,
Abt. 2, 89: 511. 1934, nom. illegit., Art. 53.1.
(See Fusarium sublunatum)
Authentic material: B 70 0100189.
Original culture: CBS 190.34 = NRRL 20897.
Original locality: Costa Rica.
Original substrate: Soil from Musa sapientum and Theobroma
cacao plantation.
Diagnostic DNA barcodes: rpb1: KX302927; rpb2: KX302935;
tef1: KX302919.
Note: Synonym fide Wollenweber & Reinking (1935).
eleocharidis Fusarium Rostr. (as ‘heleocharidis’), in Thümen,
Mycoth. Univ., Cent. 22: no. 2185. 1883.
(See Fusarium heterosporum)
Syntypes: In BPI, NEB & S (Mycoth. Univ., Cent. 22: no. 2185).
Type locality: Denmark, Fyn, Langeland.
Type substrate: Eleocharis palustris.
Notes: Synonym fide Wollenweber & Reinking (1935).
elongatum Fusarium Cooke, Grevillea 19: 4. 1890.
(See Fusarium ciliatum)
Holotypus: In K(M), Colenso 538 fide Index Fungorum .
Type locality: New Zealand.
Type substrate: Twigs.
Note: Synonym fide Wollenweber & Reinking (1935).
elongatum Fusarium De Wild., Ann. Soc. Belge Microsc. 17: 42.
1893, nom illegit., Art. 53.1.
Replacing synonym: Fusarium longissimum Sacc. & P. Syd.,
Syll. Fung. 14: 1128. 1899.
Amniculicola longissima (Sacc. & P. Syd.) Nadeeshan & K.D.
Hyde, IMA Fungus 7: 301. 2016.
Synonyms: Anguillospora longissima (Sacc. & P. Syd.) Ingold,
Trans. Brit. Mycol. Soc. 25: 402. 1942.
Holotypus: Not located.
Type locality: Belgium, Brussels, Botanical Garden.
Type substrate: Submerged plant material.
Note: Synonyms fide Rossman et al. (2016).
elongatum Fusarium O.A. Pratt, J. Agric. Res. 13: 84. 1918, nom.
110
ensiforme Fusarium Wollenw. & Reinking, Phytopathology 15:
169. 1925.
Synonym: Fusarium javanicum var. ensiforme (Wollenw. &
Reinking) Wollenw., Z. Parasitenk. 3: 483. 1931.
Fusarium javanicum subsp. ensiforme (Wollenw. & Reinking)
Raillo, Fungi of the Genus Fusarium: 229. 1950.
Holotypus: Not located.
Type locality: Honduras.
Type substrate: Rotten fruit of Ficus sp.
Notes: Synonyms fide Wollenweber & Reinking (1935). Synonym of F. javanicum fide Gerlach & Nirenberg (1982). Status
unclear [see Sandoval-Denis et al. (2019)].
entomophilum Fusarium Petch, Trans. Brit. Mycol. Soc. 11: 260.
1926.
(See Fusarium lateritium)
Holotypus: ?K(M).
Type locality: Sri Lanka, Suduganga.
Type substrate: Clitellaria heminopla.
Note: Synonym fide Wollenweber & Reinking (1935).
epicoccum Fusarium McAlpine, Fungus Diseases of Citrus trees
in Australia: 113. 1899.
(See Fusarium larvarum)
Lectotypus (hic designatus, MBT 10000680): Australia, Victoria,
Melbourne, Aspidiotus aurantium on Citrus deliciosa, 1899, D.
McAlpine, in Fungus Diseases of Citrus trees in Australia, figs
177–180.
Note: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
epimyces Fusarium Cooke, Grevillea 17: 15. 1888, nom. inval.,
Art. 38.1(a).
(See Fusarium azukiicola)
Authentic material: In K(M) fide Index Fungorum.
Original locality: UK, Reading.
Original substrate: Scleroderma sp.
Note: Synonym fide Wollenweber & Reinking (1935).
episphaeria Fusarium (Tode) W.C. Snyder & H.N. Hansen, Amer.
J. Bot. 32: 662. 1945.
Dialonectria episphaeria (Tode) Cooke (as ‘episphærica’),
Grevillea 12: 82. 1884.
FUSARIUM
Basionym: Sphaeria episphaeria Tode, Fung. Mecklenb. Sel. 2:
21. 1791.
Synonyms: Nectria episphaeria (Tode) Fr., Summa Veg. Scand.
2: 388. 1849.
Cucurbitaria episphaeria (Tode) Kuntze, Revis. Gen. Pl. 3: 461.
1898.
Cosmospora episphaeria (Tode) Rossman & Samuels, Stud.
Mycol. 42: 121. 1999.
Hypoxylon phoeniceum Bull., Hist. Champ. France 1: 171. 1791.
Sphaeria sanguinea var. media Fr., Syst. Mycol. 2: 453. 1823.
Nectria episphaeria var. media (Fr.) Sacc., Syll. Fung. 2: 497.
1883.
Dialonectria episphaeria var. verruculosa Cooke, Grevillea 12:
82. 1884.
Nectria episphaeria var. verruculosa (Cooke) Berl. & Voglino,
Syll. Fung., Addit. Vol. 1–4: 203. 1886.
Nectria episphaeria var. kretzschmariae Henn., Bot. Jahrb. Syst.
14: 364. 1891.
Nectria episphaeria var. gregaria Starb€ack, Ark. Bot. 5: 9. 1905.
Lectotypus: L 0112704 (Herb. Lugd. Bat. 910267659 ex Herb.
Persoon), selected in Booth (1959).
Type locality: Unknown.
Type substrate: Partially decorticated twig of Diatrype stigma.
episphaericum Fusarium (Cooke & Ellis) Sacc., Syll. Fung. 4:
708. 1886.
Basionym: Fusisporium episphaericum Cooke & Ellis, Grevillea
5: 50. 1876.
Cosmospora nothepisphaeria (Samuels) Rossman & Samuels, Stud. Mycol. 42: 123. 1999.
Basionym: Nectria nothepisphaeria Samuels, Mycol. Pap. 164:
30. 1991.
Synonyms: Fusarium ciliatum var. episphaericum (Cooke & Ellis)
Wollenw., Fusaria Autogr. Delin. 3: 871. 1930.
Fusarium ciliatum var. majus Wollenw., Fusaria Autogr. Delin. 3:
872. 1930.
Lectotypus (of Fusisporium episphaericum, hic designatus, MBT
10000681): USA, New Jersey, parasitic on Diatrypella sp. on
Corylus avellana, 1876, M.C. Cooke & J.B. Ellis, in Grevillea 5,
pl. 80, fig. 10.
Note: No holotype specimen could be located and therefore an
illustration is designated as lectotype.
epistroma Fusarium (Höhn.) C. Booth (as ‘epistromum’), The
Genus Fusarium: 66. 1971.
Fusicolla epistroma (Höhn.) Gr€afenhan & Seifert, Stud. Mycol.
68: 100. 2011.
Basionym: Dendrodochium epistroma Höhn., Sitzungsber. Kaiserl. Akad. Wiss. Wien. Math.-Naturwiss. Cl., Abt. 1., 118: 424.
1909.
Lectotypus: B 700014042, designated in Gr€afenhan et al. (2011).
Lectotype locality: Germany, Brandenburg, “Schmidt's Grund”
near Tamsel.
Lectotype substrate: Old stromata of Diatrypella favacea.
Epitypus: IMI 85601, designated in Gr€afenhan et al. (2011).
Ex-epitype culture: ATCC 24369 = BBA 62201 = NRRL
20439 = NRRL 20461.
Epitype locality: UK, Yorkshire.
Epitype substrate: Diatrypella on Betula.
Diagnostic DNA barcode: rpb2: HQ897765.
epithele Fusarium McAlpine, Fungus Diseases of Citrus trees in
Australia: 80. 1899.
www.studiesinmycology.org
REDELIMITED
(See Fusarium reticulatum)
Holotypus: VPRI 2563.
Type locality: Australia, New South Wales.
Type substrate: Rotten fruit of Citrus x limon.
Note: Synonym fide Wollenweber & Reinking (1935).
equinum Fusarium Növgaard, Science, N.Y. 14: 899. 1901.
Holotypus: Not located.
Type locality: USA.
Type substrate: Infected skin of Equus sp. (horse).
Notes: Status unclear. Doubtful species fide Wollenweber &
Reinking (1935). Based on the original substrate, this species
might belong to the medically important genus Neocosmospora.
However, recollection is required to confirm its taxonomic
affiliation.
equiseti Fusarium (Corda) Sacc., Syll. Fung. 4: 707. 1886.
Basionym: Selenosporium equiseti Corda, Icon. Fung. 2: 7. 1838.
Synonyms: Fusisporium ossicola Berk. & M.A. Curtis, Grevillea
3: 147. 1875.
Fusarium ossicola (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
714. 1886.
Fusarium nectriae-palmicolae Henn., Bot. Jahrb. Syst. 23: 290.
1896.
Fusarium gibbosum Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land-Forstw. 8: 190. 1910.
Fusarium roseum var. gibbosum (Appel & Wollenw.) Messiaen &
R. Cass., Ann. Inst. Natl. Rech. Agron. Tunisie 19: 435. 1968,
nom. inval., Art. 41.5.
Fusarium roseum var. gibbosum (Appel & Wollenw.) Messiaen &
R. Cass., Agronomie 8: 220. 1988, nom. inval., Art. 41.1.
Fusarium bullatum Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6:
198. 1915.
Fusarium equiseti var. bullatum (Sherb.) Wollenw., Fusaria
Autogr. Delin. 3: 916. 1930.
Fusarium gibbosum var. bullatum (Sherb.) Bilaĭ, Mikrobiol.
Zhurn. 49: 6. 1987.
Fusarium bullatum var. roseum Sherb., Mem. Cornell Univ. Agric.
Exp. Sta. 6: 201. 1915.
Fusarium roseobullatum Wollenw. (as ‘roseo-bullatum'’), Fusaria
Autogr. Delin. 1: 117. 1916.
Fusarium vasinfectum var. pisi Schikora, Arbeiten. Biol. Anst.
Land-Forstwirt. 5: 188, pl. 7. 1906, nom. illegit., Art. 53.1.
Fusarium falcatum Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land-Forstw. 8: 184. 1910.
Fusarium falcatum var. fuscum Sherb., Mem. Cornell Univ. Agric.
Exp. Sta. 6: 138. 1915.
Fusarium equiseti var. crassum Wollenw., Fusaria Autogr. Delin.
3: 921. 1930.
Fusarium terrestre Manns, Bull. North Dakota Agric. Exp. Sta.:
no. 259. 1932.
Gibberella intricans Wollenw., Fusaria Autogr. Delin. 3: 810. 1930.
Fusarium eucheliae Sartory, R. Sartory & J. Mey., Ann. Mycol.
30: 471. 1932.
Fusarium equiseti var. intermedium Saccas, Agron. Trop. (Maracay) 10: 49. 1955, nom. inval., Art. 39.1.
Lectotypus: (hic designatus, MBT 10001325): Czech Republic,
Kuchelbad, near Prague, on stems of Equisetum sp., 1836, AKJ.
Corda. Icon. Fung. 2, tab. IX, fig. 32.
Epitypus (hic designatus, MBT 10000682): Germany,
Braunschweig, Niedersachsen, soil, 3 Aug. 1994, H. I. Nirenberg,
CBS H-5570.
Ex-epitype culture: BBA 68556 = CBS 307.94 = NRRL 26419.
111
CROUS
ET AL.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982), HolubovaJechova et al. (1994) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb2: GQ505777; tef1: GQ505599.
Notes: Holubova-Jechova et al. (1994) incorrectly designated
CBS 307.94 (CBS H-5570) as neotype for Selenosporium
equiseti even though original material was available in PRM as
well as an illustration provided in the protologue. A lectotypification rather than a neotypification was required. Therefore, the
original illustration is selected as lectotype and CBS H-5570 (=
CBS 307.94) is designated as epitype here, superseding the
neotype designation.
equiseticola Fusarium Allesch., Hedwigia 34: 289. 1895.
(See Fusarium scirpi)
Holotypus: In M.
Type locality: Germany, Oberammergau.
Type substrate: Dried stems of Equisetum limosum.
Note: Synonym fide Wollenweber & Reinking (1935).
equisetorum Fusarium Desm., Pl. Crypt. N. France: no. 1546/
1846? 1843.
Basionym: Hymenula equiseti Lib., Pl. Crypt. Arduenna 3: no.
236. 1834.
(See Fusarium oxysporum)
Syntypes: In BPI, BRU, CUP, ISC PH, S & UPS (Pl. Crypt.
Arduenna 3: no. 236).
Type locality: Belgium.
Type substrate: Equisetum limosum.
Notes: Synonym fide Wollenweber & Reinking (1935).
ershadii Fusarium M. Papizadeh et al., Europ. J. Pl. Pathol. 151:
693. 2018, nom. illegit., Art. 52.1.
Basionym: Cylindrocarpon tonkinense Bugnic., Encycl. Mycol.
11: 181. 1939.
(See Fusarium tonkinense)
erubescens Fusarium Berk. & M.A. Curtis, Grevillea 3: 98. 1875.
Synonym: Fusarium alabamense Sacc., Syll. Fung. 4: 722. 1886,
nom. illegit., Art. 52.1.
Holotypus: ?K(M).
Type locality: USA, Alabama, Beaumont.
Type substrate: Dead bark.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
erubescens Fusarium (Durieu & Mont.) Sacc., Syll. Fung. 4: 719.
1886, nom. illegit., Art. 53.1.
Basionym: Fusisporium erubescens Durieu & Mont., Exploration
scientifique de l'Algerie 1–9: 351. 1848.
(See Fusarium bacilligerum)
Holotypus: ?PC.
Type locality: Algeria, Bejaïa.
Type substrate: Rhamnus alaternus.
Note: Synonym fide Wollenweber & Reinking (1935).
erubescens Fusarium Appel & Oven, Landwirtsch. Jahrb. 1905,
nom. illegit., Art. 53.1.
(See Fusarium acuminatum)
Authentic material: Not located.
Original locality: Germany.
Original substrate: Solanum lycopersicum.
Note: Synonym fide Wollenweber & Reinking (1935).
112
eucalypticola Fusarium Henn., Hedwigia 40: 355. 1901.
Holotypus: In B fide Hein (1988).
Type locality: Australia, Western Australia, Cranbrook.
Type substrate: Eucalyptus baxteri (syn. E. santalifolia)
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
eucalyptorum Fusarium Cooke & Harkn., Grevillea 9: 128. 1881.
(See Fusarium oxysporum)
Syntype: BPI 452103.
Type locality: USA, California, San Francisco Masonic Cemetery.
Type substrate: Eucalyptus sp.
Note: Synonym fide Arya & Jain (1962).
eucheliae Fusarium Sartory, R. Sartory & J. Mey., Ann. Mycol.
30: 471. 1932.
(See Fusarium equiseti)
Lectotypus (hic designatus, MBT 10000683): France, digestive
track of living caterpillar, 1932, A. Sartory, R. Sartory & J. Meyer,
in Ann. Mycol. 30: 473, figs 1–13.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
eumartii Fusarium C.W. Carp., J. Agric. Res. 5: 204. 1915.
(See Fusarium solani)
Lectotypus: Illustration Plate XIV, number 4, in Carpenter (1915),
designated in Sandoval-Denis et al. (2019).
Type locality: Unknown.
Type substrate: Solanum tuberosum.
euonymi Fusarium Syd., Beibl. Hedwigia 39: (6). 1900.
(See Fusarium lateritium)
Syntype: S-F45621 (Sydow, Mycoth. March. no. 4896).
Type al locality: Germany, Berlin.
Type substrate: Euonymus bungeanus.
Note: Synonym fide Wollenweber & Reinking (1935).
euonymi-japonici Fusarium Henn., Hedwigia 41: 139. 1902.
(See Fusarium lateritium)
Holotypus: In B fide Hein (1988).
Type locality: Germany, Berlin.
Type substrate: Euonymus japonicus.
Note: Synonym fide Wollenweber & Reinking (1935).
euwallaceae Fusarium S. Freeman et al., Mycologia 105: 1599.
2013.
Neocosmospora euwallaceae (S. Freeman et al.) Sand.-Den.
et al., Persoonia 43: 129. 2019.
Holotypus: BPI 884203.
Ex-type culture: CBS 135854 = NRRL 54722.
Type locality: Israel, central coastal region, Kibbutz Glil Yam.
Type substrate: Euwallacea sp. beetle infecting Persea americana cv. Hass.
Descriptions and illustrations: See Freeman et al. (2013).
Diagnostic DNA barcodes: rpb1: JQ038021; rpb2: JQ038028;
tef1: JQ038007.
expansum Fusarium Schltdl., Fl. Berol. 2: 139. 1824.
Synonym: ?Fusarium carpini Schulzer & Sacc., Hedwigia 23:
128. 1884.
Fusarium socium Sacc., Atti Ist. Veneto Sci. Lett. Arti, ser. 6, 2:
450. 1884.
FUSARIUM
Fusarium cirrosum Höhn., Sitzungsber. Kaiserl. Akad. Wiss.
Wien, Math.-Naturwiss. Cl., Abt. 1., 116: 153. 1907.
Fusarium macounii Dearn., Mycologia 9: 363. 1917.
Holotypus: HAL 1614 F.
Type locality: Germany, Berlin.
Type substrate: Carpinus betulus.
Descriptions and illustrations: See Wollenweber (1916–1935)
and Gerlach & Nirenberg (1982).
Notes: Both Wollenweber & Reinking (1935) and Gerlach &
Nirenberg (1982) recognised this species. This species requires epitypification from the type locality.
fabacearum Fusarium L. Lombard et al., Persoonia 43: 24.
2018 [2019].
Holotypus: CBS H-23613.
Ex-type culture: CBS 144743 = CPC 25802.
Type locality: South Africa, Western Cape Province.
Type substrate: Glycine max.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928806; rpb2: MH484938;
tef1: MH485029.
falcatum Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land-Forstw. 8: 184. 1913.
Replaced synonym: Fusarium vasinfectum var. pisi Schikora,
Arbeiten Biol. Anst. Land-Forstwirt. 5: 188, pl. 7. 1906, nom.
illegit., Art. 53.1
(See Fusarium equiseti)
Holotypus: Not located.
Type locality: Germany, Berlin.
Type substrate: Pisum sativum.
Notes: Synonym fide Wollenweber & Reinking (1935).
falciforme Fusarium (Carrion) Summerb. & Schroers, J. Clin.
Microbiol. 40: 2872. 2002.
Neocosmospora falciformis (Carrion) L. Lombard & Crous,
Stud. Mycol. 80: 227. 2015.
Basionym: Cephalosporium falciforme Carrion, Mycologia 43:
523. 1951.
Synonyms: Acremonium falciforme (Carrion) W. Gams, Cephalosporium-artige Schimmelpilze: 139. 1971.
Fusarium paranaense S.S. Costa et al., Fungal Biology 120: 55.
2015 [2016].
Holotypus: CBS 475.67 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 475.67 = IHM 939 = IMI 268681.
Type locality: Puerto Rico.
Type substrate: Mycetoma from Homo sapiens.
Diagnostic DNA barcodes: rpb1: MW218114; rpb2: LT960558;
tef1: LT906669.
fasciculatum Fusarium J.W. Xia et al., Persoonia 43: 203.
2019.
Holotypus: CBS H-24057.
Ex-type culture: CBS 131382.
Type locality: Australia, Northern Territories, Roper River area.
Type substrate: Stems of Oryza australiensis.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170406; tef1: MN170473.
fautreyi Fusarium Sacc., Syll. Fung. 10: 934. 1892.
Replaced synonym: Fusarium parasiticum Fautrey, Rev. Mycol.
(Toulouse) 11: 153. 1889, nom. illegit., Art. 53.1.
(See Fusarium lateritium)
www.studiesinmycology.org
REDELIMITED
Typus: BR5020140789424.
Type locality: France, Noidan.
Type substrate: Vitis vinifera.
Note: Synonyms fide Wollenweber & Reinking (1935).
ferrugineum Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 2. 2020.
Neocosmospora ferruginea Sand.-Den. & Crous, Persoonia
43: 130. 2019.
Holotypus: CBS H-23981.
Ex-type culture: CBS 109028 = NRRL 32437.
Type locality: Switzerland.
Type substrate: Subcutaneous nodule of Homo sapiens.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: HM347157; rpb2: EU329581;
tef1: DQ246979.
ferruginosum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 190. 1915.
(See Fusarium acuminatum)
Typus: ?CUP-007445.
Type locality: USA, New York, Long Island
Type substrate: Solanum tuberosum
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in CUP.
ficicrescens Fusarium Al-Hatmi et al., Fungal Biol. 120: 274.
2015 [2016].
Holotypus: CBS H-21815.
Ex-type culture: CBS 125178.
Type locality: Iran, Estahban.
Type substrate: Fruit of Ficus carica.
Descriptions and illustrations: See Al-Hatmi et al. (2016).
Diagnostic DNA barcodes: rpb1: MT010950; rpb2: MT010977;
tef1: MT011004.
filiferum Fusarium (Preuss) Wollenw., Fusaria Autogr. Delin. 1:
220. 1916.
Basionym: Fusoma filiferum Preuss, Linnaea 25: 73. 1852.
Synonym: Fusarium scirpi var. filiferum (Preuss) Wollenw.,
Fusaria Autogr. Delin. 3: 936. 1930.
(See Fusarium scirpi)
Holotypus: Not located.
Type locality: Germany.
Type substrate: Bark of Pinus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
filisporum Fusarium (Cooke) Sacc., Syll. Fung. 4: 708. 1886.
Basionym: Fusisporium filisporum Cooke, Grevillea 8: 8. 1879.
(See Fusarium ciliatum)
Holotypus: In K(M), Muller s.n. fide Index Fungorum.
Type locality: UK, Eastbourne.
Type substrate: Orthotrichum sp.
Note: Synonym fide Wollenweber & Reinking (1935).
fissum Fusarium Peyl, Lotos 8: 30. 1858.
(See Fusarium candidum (Link.) Sacc.)
Lectotypus (hic designatus, MBT 10000684): Germany, twigs of
Citrus aurantiacum, 1858, J. Peyl, in Lotos 8, fig. 17.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
flagelliforme Fusarium J.W. Xia et al., Persoonia 43: 204. 2019.
Holotypus: CBS H-24058.
113
CROUS
ET AL.
Ex-type culture: CBS 162.57 = NRRL 36269.
Type locality: Croatia, Zagreb.
Type substrate: Seedlings of Pinus nigra.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: GQ505823; tef1: GQ505645.
flavidum Fusarium (Bonord.) Sacc., Syll. Fung. 4: 698. 1886.
Basionym: Fusisporium flavidum Bonord., Bot. Zeitung (Berlin)
19: 194. 1861.
(See Fusarium reticulatum)
Lectotypus (hic designatus, MBT 10000685): Germany, rotten
tree, 1861, H.F. Bonorden, in Bot. Zeitung (Berlin) 19: tab. VIII,
fig. 3.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
flavum Fusarium (Fr.) Wollenw., Z. Parasitenk. (Berlin) 3: 305.
1931.
Basionym: Fusisporium flavum Fr., Syst. Mycol. 3: 444. 1832.
(See Fusarium dimerum)
Holotypus: Not located.
Type locality: Germany, Bonn.
Type substrate: Aster sp.
Note: Synonym fide Booth (1971).
flocciferum Fusarium Corda, in Sturm, Deutschl. Fl., Abt. 3,
Pilze Deutschl. 2: 17. 1828.
Synonyms: Fusarium vinosum Massee, Brit. Fung.-Fl. 3: 479.
1893.
Fusarium clavatum Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 234. 1915.
Fusarium idahoanum O.A. Pratt, J. Agric. Res. 13: 86. 1918.
Fusarium nigrum O.A. Pratt, J. Agric. Res. 13: 90. 1918.
Lectotypus: (hic designatus, MBT 10001326) Germany, Berlin,
on shell of the fruit of Aesculus hippocastanum. AKJ. Corda,
Sturm, Deutschl. Fl., Abt. 3, Pilze Deutschl. 2, pl. 7.
Epitypus (hic designatus, MBT 10000686): Germany, greenhouse soil, 1966, D. Bredemeier, CBS 821.68 (preserved as
metabolically inactive culture).
Ex-epitype culture: CBS 821.68 = NRRL 28450.
Descriptions and illustrations: See Booth (1971) and Gerlach &
Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: MW928807; rpb2: MW928824;
tef1: MW928837.
Notes: Corda's original illustration of Fusarium flocciferum is here
selected as lectotype. Gerlach & Nirenberg (1982) considered
isolate CBS 821.68, along with CBS 792.70, as good representatives of F. flocciferum. Based on their observations and
collection locality, CBS 821.68 is designated as epitype of
F. flocciferum.
floridanum Fusarium T. Aoki et al., Mycologia 111: 922. 2019.
Neocosmospora floridana (T. Aoki et al.) L. Lombard & Sand.Den., comb. nov. MycoBank MB 837664.
Basionym: Fusarium floridanum T. Aoki et al., Mycologia 111:
922. 2019.
Holotypus: BPI 910972.
Ex-type culture: MAFF 246849 = NRRL 62628.
Type locality: USA, Florida, Gainsville.
Type substrate: Mycangium of Euwallacea interjectus infesting
Acer negundo.
Descriptions and illustrations: See Aoki et al. (2019).
114
Diagnostic DNA barcodes: rpb1: KC691593; rpb2: KC691624,
KC691653; tef1: KC691535.
Notes: A new combination is provided in the genus Neocosmospora based on the phylogenetic relationship (Aoki et al.
2019) of this species to other Neocosmospora spp. in the ambrosia clade.
foeni Fusarium (Berk. & Broome) Sacc., Syll. Fung. 4: 699. 1886.
Basionym: Fusisporium foeni Berk. & Broome, Ann. Mag. Nat.
Hist., ser. 2, 7: 179. 1851.
(See Fusarium merismoides)
Holotypus: ?K(M).
Type locality: UK, Northamptonshire, Apethrope.
Type substrate: A hay stalk.
Note: Synonym fide Wollenweber & Reinking (1935).
foetens Fusarium Schroers et al., Mycologia 96: 398. 2004.
Holotypus: CBS 110286 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 110286 = NRRL 31852 = PD 2001/7244.
Type locality: Netherlands, Zuid-Holland Province, Maasland.
Type substrate: Begonia elatior hybrid.
Descriptions and illustrations: See Schroers et al. (2004) and
Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: MW928808; rpb2: MW928825;
tef1: AY320087.
foliicola Fusarium Allesch., Hedwigia 34: 289. 1895.
(See Fusarium cymbiferum)
Holotypus: In M.
Type locality: Germany, Oberammergau.
Type substrate: Arabis alpina.
Note: Synonym fide Wollenweber & Reinking (1935).
fracticaudum Fusarium Herron et al., Stud. Mycol. 80: 137.
2015.
Holotypus: PREM 60895.
Ex-type culture: CBS 137233 = CMW 25245.
Type locality: Colombia, Risaralda, Angela Maria (Santa Rosa).
Type substrate: Pinus maximinoi.
Descriptions and illustrations: See Herron et al. (2015).
Notes: Comparisons of recently generated sequences for the
living ex-type (CBS 137233 = CMW 25245) of F. fracticaudum
indicate a strain transposition or contamination by another
Fusarium species. Therefore, this species needs to be recollected from the type locality and substrate or sequences need to
be generated from the holotype specimen.
fractiflexum Fusarium T. Aoki et al., Mycoscience 42: 462.
2001.
Holotypus: NIAES 20515.
Ex-type culture: MAFF 237529 = NRRL 28852.
Type locality: Japan, Yamanashi, Enzan.
Type substrate: Cymbidium sp.
Descriptions and illustrations: See Aoki et al. (2001).
Diagnostic DNA barcodes: rpb1: LR792578; rpb2: LT575064;
tef1: AF160288.
fractum Fusarium Sacc. & Cavara, Nuovo Giorn. Bot. Ital., n.s.,
7: 308. 1900.
(See Fusarium candidum (Link) Sacc.)
Holotypus: In PAD.
Type locality: Italy.
Type substrate: Fagus sp.
FUSARIUM
Note: Synonym fide Wollenweber & Reinking (1935).
fragrans Fusarium P. Crouan & H. Crouan, Fl. Finistere: 14.
1867.
(See Fusarium candidum (Link) Sacc.)
Holotypus: ?PC.
Type locality: France.
Type substrate: Salix sp.
Note: Synonym fide Wollenweber & Reinking (1935).
fraxini Fusarium Allesch., Ber. Bot. Vereines Landshut 12: 130.
1892.
(See Fusarium sambucinum)
Holotypus: In M.
Type locality: Germany, München.
Type substrate: Fraxinus excelsior.
Note: Synonym fide Wollenweber & Reinking (1935).
fraxini Fusarium Kabat & Bubak, Fungi Imperf. Exs., no. 900.
1912, nom. illegit., Art. 53.1.
Fusicoccum fraxini Sherb., Phytopathology 18: 148. 1928.
Authentic material: BPI 451324.
Original locality: Czech Republic.
Original substrate: Fraxinus excelsior.
fredkrugeri Fusarium Sand.-Den. et al., MycoKeys 34: 79. 2018.
Holotypus: CBS H-23496.
Ex-type culture: CBS 144209 = CPC 33747.
Type locality: South Africa, Kruger National Park, Skukuza,
Granite Supersite.
Type substrate: Rhizosphere soil of Melhania acuminata.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018b).
Diagnostic DNA barcodes: rpb1: LT996199; rpb2: LT996147;
tef1: LT996097.
fructigenum Fusarium Fr., Syst. Mycol. 3: 471. 1832.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Unknown.
Type substrate: Fruit of Rosa pomifera.
Note: Synonym fide Wollenweber & Reinking (1935).
fuckelii Fusarium Sacc., Syll. Fung. 4: 695. 1886.
Geejayessia desmazieri (Becc. & De Not.) Schroers et al.,
Stud. Mycol. 68: 130. 2011.
Basionym: Nectria desmazieri Becc. & De Not., Schem. di
Classif. Sferiacei: 10. 1863.
Synonyms: Dialonectria desmazieri (Becc. & De Not.) Petch,
Naturalist (London): 281. 1937.
Nectria coccinea var. cicatricum Desm., Ann. Sci. Nat., Bot 10:
351. 1848 (fide Wollenweber & Reinking 1935 and Booth 1971).
Nectria gibbera Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 177. 1870.
Lectotypus: G 00110886 (Fuckel, Fungi Rhen. No. 2357),
designated in Schroers et al. (2011).
Type locality: Germany, Rheingau.
Type substrate: Buxus sempervirens.
fujikuroi Fusarium Nirenberg, Mitt. Biol. Bundesanst. LandForstw. Berlin-Dahlem 169: 32. 1976
Synonyms: Lisea fujikuroi Sawada, Special Bull. Agric. Exp. Sta.
Gov. Formosa 19: 251. 1919.
Gibberella fujikuroi (Sawada) Wollenw., Z. Parasitenk. (Berlin) 3:
514. 1931.
www.studiesinmycology.org
REDELIMITED
?Gibberella fujikuroi var. subglutinans E.T. Edwards, Agric. Gaz.
New South Wales 44: 895. 1933.
?Gibberella subglutinans (E.T. Edwards) P.E. Nelson et al.,
Fusarium species. An illustrated manual for identification (University Park): 135. 1983.
?Oospora cephalosporioides Luchetti & Favilli, Annali Fac. Agrar.
R. Univ. Pisa 1: 399. 1938.
?Gibberella fujikuroi f. oryzae Saccas, Rev. Pathol. Veg. Entomol. Agric. France 30: 77. 1951.
?Gibberella fujikuroi var. intermedia Kuhlman, Mycologia 74:
766. 1982.
Holotypus: IMI 202879.
Ex-type culture: BBA 12428 = BBA 63630 = CBS 221.76 = IHEM
3821 = IMI 196086 = IMI 202879 = NRRL 13620 = NRRL
13998 = NRRL 22174.
Type locality: Taiwan.
Type substrate: Oryza sativa.
Descriptions and illustrations: See Nirenberg (1976), Gerlach &
Nirenberg (1982) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: JX171456; rpb2: JX171570;
tef1: AF160279.
fuliginosporum Fusarium Sibilia, Ann. Reale. Ist. Super. Agrar.
Forest., ser. 2, 1: 77. 1925.
Holotypus: Not located.
Type locality: Italy.
Type substrate: Forest containing mostly Cedrus deodara.
Note: Mentioned by Wollenweber & Reinking (1935), but no
additional records of this species could be located.
fungicola Fusarium (Har. & P. Karst.) Sacc., Syll. Fung. 10: 730.
1892.
?Alysidium hypophleodes (Corda) Bonord., Handb. Allg.
Mykol.: 35. 1851.
Basionym: Fusidium hypophleodes Corda, Icon. Fung. 1: 3, tab.
1, fig. 50. 1837.
Synonym: Fusamen fungicola Har. & P. Karst. (as ‘fungicolum’),
Rev. Mycol. (Toulouse) 12: 129. 1890.
Holotypus: Not located.
Type locality: Finland, Mustiala.
Type substrate: Lenzites betulina.
Note: Synonyms fide Wollenweber & Reinking (1935).
funicola Fusarium Tassi, Bull. Lab. Orto Bot. Reale Univ. Siena 3:
131. 1900.
(See Fusarium graminearum)
Holotypus: ?SIENA.
Type locality: Italy.
Type substrate: Rotten string.
Note: Synonym fide Wollenweber & Reinking (1935).
fusarioides Fusarium (Gonz. Frag. & Cif.) C. Booth, The Genus
Fusarium: 88. 1971.
Basionym: Dactylium fusarioides Gonz. Frag. & Cif., Bol. Real
Soc. Esp. Hist. Nat. 27: 280. 1927.
(See Fusarium chlamydosporum)
Holotypus: ?MA-Funhist: 7609-1.
Type locality: Dominican Republic, Moca.
Type substrate: leaves of Crotalaria sp.
Note: Synonym fide Gerlach & Nirenberg (1982).
fuscum Fusarium (Bonord.) Sacc., Syll. Fung. 4: 699. 1886.
Basionym: Selenosporium fuscum Bonord., Handb. Mykol.: 135.
1851.
115
CROUS
ET AL.
(See Fusarium citrulli Sartory)
Holotypus: Not preserved fide Holubova-Jechova et al. (1994).
Type locality: Germany.
Type substrate: Bark.
Note: Synonym fide Wollenweber & Reinking (1935).
gaditjirrii Fusarium Phan et al., Stud. Mycol. 50: 265. 2004.
Synonym: Gibberella gaditjirrii Phan et al., Stud. Mycol. 50: 264.
2004.
Holotypus: DAR 76663.
Ex-type culture: CBS 116011 = F15048 = NRRL 53678.
Type locality: Australia, Queensland, Walkamin Research
Station.
Type substrate: Heteropogon triticeus.
Descriptions and illustrations: See Phan et al. (2004).
Diagnostic DNA barcodes: rpb2: HQ662690; tef1: AY639636.
gallinaceum Fusarium Cooke & Harkn., Grevillea 9: 8. 1880.
(See Fusarium merismoides)
Holotypus: BPI 452133.
Type locality: USA, California, Sausalito.
Type substrate: Feathers of Gallus sp. (chicken).
Note: Synonym fide Wollenweber & Reinking (1935).
gamsii Fusarium Torbati et al., Mycol. Progr. 18: 127. 2018
[2019].
Holotypus: CBS H-23561.
Ex-type culture: CBS 143610 = CPC 30862.
Type locality: Iran, West Azerbaijan Province, Orumieh-Salmas.
Type substrate: Agaricus bisporus.
Descriptions and illustrations: See Torbati et al. (2019).
Diagnostic DNA barcodes: rpb2: LT970760; tef1: LT970788.
gaudefroyanum Fusarium Sacc., Michelia 2: 132. 1880.
(See Fusarium avenaceum)
Holotypus: In PAD.
Type locality: France, Paris.
Type substrate: Cyperaceae.
Note: Synonym fide Wollenweber & Reinking (1935).
gemmiperda Fusarium Aderh., Z. Pflanzenkrankh. 11: 70. 1901.
(See Fusarium lateritium)
Lectotypus (hic designatus, MBT 10000687): Germany, Prunus
cerasus, 1901, R. Aderhold, in Z. Pflanzenkrankh. 11: pl. II, figs
1–4.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
genevense Fusarium Dasz., Bull. Soc. Bot. Geneve, ser. 2, 4:
305. 1912.
(See Fusarium sambucinum)
Lectotypus (hic designatus, MBT 10000688): Switzerland,
Geneva, from soil, 1912, M. Daszewska, in Bull. Soc. Bot.
Geneve, ser. 2, 4: 306, fig. 27.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
georginae Fusarium Corda, Icon. Fung. 2: 4. 1838.
(See Fusarium cymbiferum)
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, Prague.
Type substrate: Dahlia sp.
116
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in PRM.
gerlachii Fusarium T. Aoki et al., Fungal Genet. Biol. 44: 1202.
2007.
Holotypus: BPI 871657.
Ex-type culture: LRG 00-551 = NRRL 36905.
Type locality: USA, Minnesota, Polk County, Climax.
Type substrate: Triticum aestivum.
Descriptions and illustrations: See Starkey et al. (2007).
Diagnostic DNA barcodes: rpb1: KM361646; rpb2: KM361664;
tef1: DQ459742.
gibbosum Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 190. 1910 [1913].
(See Fusarium equiseti)
Holotypus: ?BPI 452135.
Type locality: Germany, Berlin
Type substrate: Solanum tuberosum
Note: Synonym fide Booth (1971).
gigas Fusarium Speg. Anales Soc. Ci. Argent. 22: 221. 1886.
Holotypus: In LPS fide Farr (1973).
Type locality: Paraguay.
Type substrate: Bambusa sp.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971) and Gerlach & Nirenberg (1982).
Notes: This species requires epitypification. Wollenweber &
Reinking (1935), Booth (1971), and Gerlach & Nirenberg
(1982) accepted this species, although limited information is
available.
glandicola Fusarium Cooke & W.R. Gerard, Grevillea 7: 14.
1878.
Tubercularia glandicola (Cooke & W.R. Gerard) Wollenw. &
Reinking, Fusarien: 325. 1935.
Holotypus: In K(M), Gerard s.n. fide Index Fungorum.
Type locality: USA, New York.
Type substrate: Acorns of Quercus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
glandicola Fusarium Allesch., Ber. Bot. Vereines Landshut 12:
130. 1892, nom. illegit., Art. 53.1.
Replacing synonym. Fusarium allescheri Sacc. & P. Syd., Syll.
Fung. 14: 1128. 1899.
(See Fusarium melanochlorum)
Authentic material: In M.
Original locality: Germany, München.
Original substrate: Fruits of Quercus robur (syn. Q. pedunculata).
Note: Synonyms fide Wollenweber & Reinking (1935).
gleditschiae Fusarium Therry (as ‘gledrischiae’), in Roumeguere,
Fungi Sel. Gall. Exs.: no. 5496. 1890, nom. nud., Art. 38.1(a).
Gloeosporium gleditschiae Therry ex Wollenw., Z. Parasitenk.
(Berlin) 3: 437. 1931.
Note: Synonym fide Wollenweber & Reinking (1935).
gleditschiicola Fusarium Dearn. & Barthol. (as ‘gleditsiaecolum’),
Mycologia 9: 363. 1917.
(See Fusarium lateritium)
Holotypus: JD 4379 in DAOM.
Type locality: USA, Kansas, Stockton.
Type substrate: Gleditsia triacanthos.
Note: Synonym fide Wollenweber & Reinking (1935).
FUSARIUM
globosum Fusarium Rheeder et al., Mycologia 88: 509. 1996.
Holotypus: BPI 802834.
Ex-type culture: CBS 428.97 = DOAM 214966 = FRC M8014 = IMI 375330 = MRC 6647 = NRRL 26131 = PREM 51878.
Type locality: South Africa, Eastern Cape Province, Butterworth
district, Teko Experimental Farm.
Type substrate: Zea mays.
Descriptions and illustrations: See Rheeder et al. (1996) and
Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: KF466396; rpb2: KF466406;
tef1: KF466417.
globulosum Fusarium Pass., in Rabenhorst, Fungi Eur. Exs. no.
2262. 1877.
Syntypes: In BPI, CUP, ILL & S (Fungi Eur. Exs. # 2262).
Type locality: Italy, Parma.
Type substrate: Salvia verticillata.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
gloeosporioides Fusarium Speg. (as ‘gloeosporoide’), Anales
Mus. Nac. Hist. Nat. Buenos Aires 6: 350. 1898 [1899].
(See Fusarium incarnatum)
Holotypus: In LPS fide Farr (1973).
Type locality: Argentina, La Plata.
Type substrate: Fruits of Passiflora tweediana.
Note: Synonym fide Wollenweber & Reinking (1935).
gloeosporioides Fusarium (Speg.) Sacc. & Trotter, Syll. Fung. 22:
1482. 1913, nom. illegit., Art. 53.1.
Basionym: Selenosporium gloeosporioides Speg. (as ‘gloesporioides’), Anales Mus. Nac. Hist. Nat. Buenos Aires 13: 458.
1911.
(See Fusarium lateritium)
Holotypus: In LPS (Myc. Argent. ser. 5, no. 1167) fide Farr
(1973).
Type locality: Argentina, Buenos Aires.
Type substrate: Pircunia dioica.
Note: Synonym fide Wollenweber & Reinking (1935).
glumarum Fusarium Sacc., Syll. Fung. 4: 706. 1886.
Replaced synonym: Fusarium pallens Berk. & M.A. Curtis,
Grevillea 3: 99. 1875, nom. illegit., Art. 53.1, non Fusarium
pallens Nees & T. Nees 1818.
(See Fusarium incarnatum)
Syntype: CBRU00007755.
Type locality: USA.
Type substrate: Juncus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
glycines Fusarium L. Lombard et al., Persoonia 41: 25. 2018
[2019].
Holotypus: CBS H-23614.
Ex-type culture: CBS 144746 = CPC 25808.
Type locality: South Africa, North West Province.
Type substrate: Glycine max.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928809; rpb2: MH484942;
tef1: MH485033.
goolgardi Fusarium D.M. Robinson et al., Fungal Diversity 77:
357. 2015 [2016].
Holotypus: RGB5411.
www.studiesinmycology.org
REDELIMITED
Ex-type culture: NRRL 66250 = RGB5411.
Type locality: Australia, New South Wales, Bungonia State
Conservation Area.
Type substrate: Xanthorrhoea glauca.
Descriptions and illustrations: See Laurence et al. (2016).
Diagnostic DNA barcodes: rpb1: KP083270; rpb2: KP083280;
tef1: KP101123.
gossypinum Fusarium L. Lombard & Crous, Persoonia 41: 26.
2018 [2019].
Holotypus: CBS H-23615.
Ex-type culture: CBS 116613.
Type locality: Ivory Coast, Bouake.
Type substrate: Gossypium hirsutum.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb2: MH484909; tef1: MH485000.
gracile Fusarium McAlpine, Proc. Linn. Soc. New South Wales
28: 554. 1903.
(See Fusarium avenaceum)
Holotypus: VPRI 2564.
Type locality: Australia, Victoria, Sandringham.
Type substrate: Flowering stem of Lobelia gibbosa.
Note: Synonym fide Wollenweber & Reinking (1935).
gracilipes Fusarium J.W. Xia et al., Persoonia 43: 205. 2019.
Holotypus: CBS H-24059.
Ex-type culture: NRRL 43635.
Type locality: USA, Nebraska.
Type substrate: Equus sp. (horse).
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb1: HM347188; rpb2: GQ505840;
tef1: GQ505662.
graminearum Fusarium Schwabe, Fl. Anhalt. 2: 285. 1839.
Synonyms: Sphaeria zeae Schwein., Schriften Naturf. Ges.
Leipzig 1: 48. 1822, non Fusarium zeae (Westend.) Sacc. 1886.
Dothidea zeae (Schwein.) Schwein., Trans. Amer. Philos. Soc.,
n.s., 4: 230. 1832.
Hendersoniopsis zeae (Schwein.) Woron., Fungal and Bacterial
Diseases of Agricultural Plants: 255. 1922.
Gibberella zeae (Schwein.) Petch, Ann. Mycol. 34: 260. 1936.
Fusarium stictoides Durieu & Mont., Explor. Sci. Algerie 1: 334.
1848.
Sphaeria saubinetii Durieu & Mont., Explor. Sci. Algerie 1: 479.
1849.
Gibbera saubinetii (Durieu & Mont.) Mont., Syll. Gen. Sp. Crypt.:
252. 1856.
Botryosphaeria saubinetii (Durieu & Mont.) Niessl, Verh. Naturf.
Vereins Brünn 10: 195. 1872.
Gibberella pulicaris subsp. saubinetii (Durieu & Mont.) Sacc.,
Michelia 1: 317. 1878.
Gibberella saubinetii (Durieu & Mont.) Sacc., Michelia 1: 513.
1879.
Fusisporium insidiosum Berk., Gard. Chron. 1860: 480. 1860.
Fusarium insidiosum (Berk.) Sacc., Syll. Fung. 4: 707. 1886,
nom. illegit., Art. 53.1.
Gibberella saubinetii var. coronillae Sacc., Michelia 1: 513. 1879.
Fusarium mollerianum Thüm., Inst. Coimbra 28: 263. 1881.
Gibberella saubinetii subsp. pachyspora Sacc., Michelia 2: 74.
1880.
117
CROUS
ET AL.
Gibberella saubinetii var. pachyspora (Sacc.) Sacc., Syll. Fung.
2: 555. 1883.
Fusarium caricis Oudem., Verslagen Meded. Afd. Natuurk. Kon.
Akad. Wetensch., ser. 3, 7: 325. 1890.
Fusarium graminearum var. caricis (Oudem.) Wollenw., Z. Parasitenk. (Berlin) 3: 365. 1931.
?Fusarium rhoicola Fautrey, Rev. Mycol. (Toulouse) 17: 171.
1895.
Fusarium funicola Tassi, Bull. Lab. Orto Bot. Reale Univ. Siena 3:
131. 1900.
Gibberella saubinetii f. acuum Feltgen, Vorstud. Pilzfl. Luxemburg, Nachtr. III: 303. 1903.
Gibberella saubinetii var. acuum (Feltgen) Sacc. & D. Sacc., Syll.
Fung. 17: 813. 1905.
Gibberella saubinetii var. tetraspora Feltgen, Vorstud. Pilzfl.
Luxemburg, Nachtr. III: 302. 1903.
Gibberella saubinetii var. calami Henn., Beibl. Hedwigia 42: (79).
1903.
Gibberella saubinetii var. mate Speg., Anales Mus. Nac. Hist.
Nat. Buenos Aires 17: 129. 1908.
?Selenosporium bufonicola Speg., Anales Mus. Nac. Hist. Nat.
Buenos Aires, ser. 3, 13: 458. 1910.
?Fusarium bufonicola (Speg.) Sacc. & Trotter, Syll. Fung. 22:
1486. 1913.
Fusarium rostratum Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 30. 1910 [1913].
Gibberella saubinetii var. flacca Wollenw., Z. Parasitenk. (Berlin)
3: 433. 1931.
Lectotypus (hic designatus, MBT 10000689): Germany, inflorescence of Triticum sp., 1839, S.H. Schwabe, in Flora Anhaltina
2, tab. VI. fig. 7.
Epitypus (hic designatus, MBT 10000690): Germany, Hordeum
vulgare, 1988, L. Niessen, CBS 136009 (preserved as metabolically inactive culture).
Ex-epitype culture: CBS 136009.
Descriptions and illustrations: See Booth (1971), Gerlach &
Nirenberg (1982) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: MW928810; rpb2: MW928826;
tef1: MW928838.
Notes: This well-known and economically important pathogen of
gramineous hosts has a global distribution and is accepted as
originally circumscribed. However, no type material is available
for taxonomic reference. Therefore, a lectotype based on an
illustration from the original protologue and an epitype is designated here to provide taxonomic stability for this species.
graminum Fusarium Corda, Icon. Fung. 1: 3. 1837.
Synonym: Fusarium herbarum var. graminum (Corda) Wollenw.,
Fusaria Autogr. Delin. 3: 891. 1930.
Fusarium avenaceum var. graminum (Corda) Raillo, Fungi of the
Genus Fusarium: 188. 1950.
Fusarium corallinum Sacc., Nuovo Giorn. Bot. Ital. 8: 196. 1876.
Lectotypus (hic designatus, MBT 10000691): Germany,
gramineous plant, 1837, A.C.J Corda, in Icon. Fung. 1, tab. I, fig.
59.
Descriptions and illustrations: See Wollenweber & Reinking
(1935) and Gerlach & Nirenberg (1982).
Notes: This species is recognised by Wollenweber & Reinking
(1935) and Gerlach & Nirenberg (1982). Recollection from the
type host and locality is required. No holotype specimen could be
located and therefore an illustration is designated as lectotype.
118
granulare Fusarium Kalchbr., Crypt. Austro-Afric., no. 1068.
1874.
(See Fusarium sambucinum)
Holotypus: ?B 70 0100191 (Crypt. Austro-Afric., no. 1068).
Type locality: South Africa, Eastern Cape Province, SomersetEast.
Type substrate: Datura stramonium (syn. Datura tatula).
Note: Synonym fide Wollenweber & Reinking (1935).
granulosum Fusarium Ellis & Everh., Proc. Acad. Nat. Sci.
Philadelphia 45: 466. 1894 [1893].
(See Fusarium avenaceum)
Holotypus: Commons 2091 in NY.
Type locality: USA, Delaware, New Castle, Mount Cuba.
Type substrate: Smilax hispida.
Note: Synonym fide Wollenweber & Reinking (1935).
grosmichelii Fusarium Maryani et al., Stud. Mycol. 92: 176.
2018 [2019].
Holotypus: InaCC F833 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F833.
Type locality: Indonesia, West Java, Bogor, Suakarya
(Megamendung).
Type substrate: Pseudostem of Musa acuminata var. Pisang
Ambon Lumut.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479548; rpb2: LS479295;
tef1: LS479744.
guilinense Fusarium M.M. Wang et al., Persoonia 43: 80. 2019.
Holotypus: HAMS 248037.
Ex-type culture: CGMCC 3.19495 = LC12160.
Type locality: China, Guangxi Province, Guilin.
Type substrate: Leaf of Musa nana.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289831; rpb2: MK289747;
tef1: MK289594.
guttiforme Fusarium Nirenberg & O'Donnell, Mycologia 90:
446. 1998.
Holotypus: B 70 0001690.
Ex-type culture: BBA 69661 = CBS 409.97 = IMI 376113 = NRRL
25295.
Type locality: Brazil.
Type substrate: Ananas comosus.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998).
Diagnostic DNA barcodes: rpb1: MT010938; rpb2: MT010967;
tef1: KC514066.
gymnosporangii Fusarium Jaap, Ann. Mycol. 14: 44. 1916.
Nectria gymnosporangii (Jaap) Rossman, Mycotaxon 8: 515.
1979.
Basionym: Calonectria gymnosporangii Jaap, Ann. Mycol. 14:
10. 1916.
Synonyms: Bactridium gymnosporangii (Jaap) Wollenw., Fusaria
Autogr. Delin. 1: 458. 1916.
Cylindrocarpon gymnosporangii (Jaap) Rossman, Mycol. Pap.
150: 31. 1983.
Holotypus: In HBG fide Rossman (1979).
Type locality: Croatia, Dalmatia, Lapad near Ragusa.
Type substrate: Parasitic on Gymnosporangium confusum on
Juniperus phoenicea branches.
FUSARIUM
haematococcum Fusarium Nalim et al., Mycologia 103: 1322.
2011.
Neocosmospora haematococca (Berk. & Broome) Samuels
et al., Mycologia 103: 1322. 2011.
Basionym: Nectria haematococca Berk. & Broome, J. Linn. Soc.,
Bot. 14: 116. 1875.
Synonyms: Dialonectria haematococca (Berk. & Broome) Cooke,
Grevillea 12: 110. 1884.
Cucurbitaria haematococca (Berk. & Broome) Kuntze, Revis.
Gen. Pl. 3: 461. 1898.
Hypomyces haematococcus (Berk. & Broome) Wollenw., Angew.
Bot. 8: 191. 1926.
Haematonectria haematococca (Berk. & Broome) Samuels &
Nirenberg, Stud. Mycol. 42: 135. 1999.
?Nectria lanata Pat., Bull. Soc. Mycol. France 8: 52. 1892 (fide
Samuels 1976).
?Nectria aurantiella Speg., Anales Mus. Nac. Hist. Nat. Buenos
Aires 6: 287. 1898.
?Nectria episphaerioides Penz. & Sacc., Malpighia 11: 511. 1898
[1897].
?Nectria cinnabarina var. jaraguensis Höhn., Denkschr. Kaiserl.
Akad. Wiss. Wien, Math.-Naturwiss. Kl. 83: 18. 1907.
?Nectria bogoriensis C. Bernard, Bull. Dep. Agric. Indes Neerl.
11: 45. 1907.
?Nectria victoriae Henn., in Rehm, Ann. Mycol. 5: 81. 1907, nom.
inval., Art. 38.1(a).
?Nectria calonectricola Henn., Hedwigia 48: 105. 1908.
?Nectria citri Henn., Hedwigia 48: 104. 1908.
?Nectria luteococcinea Höhn., Sitzungsber. Kaiserl. Akad. Wiss.
Wien, Math.-Naturwiss. Cl., Abt. 1. 118: 299. 1909.
?Nectria bainii var. hypoleuca Sacc., Nuovo Giorn. Bot. Ital. 23:
205. 1916.
?Nectria confluens Seaver, Sci. Surv. Porto Rico & Virgin Islands
8: 44. 1926, nom. illegit., Art. 53.1.
Lectotypus: K(M) 252877, designated in Samuels (1976).
Lectotype locality: Sri Lanka.
Lectotype substrate: Unknown
Epitypus: BPI 871363, designated in Nalim et al. (2011).
Ex-epitype culture: CBS 119600 = FRC S-1832.
Epitype locality: Sri Lanka, Sabaragamuwa Province, Sinharaja
Man and Biosphere Reserve, Morningside, vicinity Bungalow in
forested slope.
Epitype substrate: Dying tree.
Descriptions and illustrations: See Nalim et al. (2011).
Diagnostic DNA barcodes: rpb2: LT960561; tef1: KM231926.
hainanense Fusarium M.M. Wang et al., Persoonia 43: 82.
2019.
Holotypus: HAMS 248038.
Ex-type culture: CGMCC 3.19478 = LC11638.
Type locality: China, Hainan Province.
Type substrate: Stem of Oryza sp.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289833; rpb2: MK289735;
tef1: MK289581.
hakeae Fusarium Henn., Verh. Bot. Vereins Prov. Brandenburg
40: 175. 1899.
Gloeosporium hakeae (Henn.) Wollenw., Fusaria Autogr. Delin.
1: 494. 1916.
Holotypus: In B fide Hein (1988).
Type locality: Germany, Berlin.
www.studiesinmycology.org
REDELIMITED
Type substrate: Hakea salicifolia.
Note: Synonym fide Wollenweber & Reinking (1935).
heidelbergense Fusarium Sacc., Ann. Mycol. 8: 346. 1910.
(See Fusarium culmorum)
Holotypus: In PAD.
Type locality: Germany, Heidelberg.
Type substrate: Cymbidium sp.
Note: Synonym fide Wollenweber & Reinking (1935).
helgardnirenbergiae Fusarium O'Donnell et al., Index Fungorum
440: 2. 2020.
Neocosmospora nirenbergiana Sand.-Den. & Crous, Persoonia 43: 143. 2019.
Holotypus: CBS H-23988.
Ex-type culture: BBA 65023 = CBS 145469 = G.J.S. 87127 = NRRL 22387.
Type locality: French Guiana.
Type substrate: Bark of unidentified tree.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb2: EU329505; tef1: AF178339.
helianthi Fusarium (Schwein.) Wollenw., Fusaria Autogr. Delin. 2:
555. 1924.
Basionym: Vermicularia subeffigurata helianthi Schwein., Trans.
Amer. Philos. Soc., n.s., 4: 228. 1832.
(See Fusarium tricinctum)
Holotypus: PH00078405.
Type locality: Unknown.
Type substrate: Helianthus annuus.
Note: Synonyms fide Wollenweber & Reinking (1935).
helotioides Fusarium Berk. & M.A. Curtis, in Berkeley, Grevillea
3: 98. 1875.
Holotypus: ?K(M).
Type locality: USA, Alabama.
Type substrate: Ilex decidua (syn. Ilex prinoides).
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
hengyangense Fusarium (Z.Q. Zeng & W.Y. Zhuang) O'Donnell
et al., Index Fungorum 440: 2. 2020.
Neocosmospora hengyangensis Z.Q. Zeng & W.Y. Zhuang,
Phytotaxa 319: 179. 2017.
Holotypus: HMAS 254518.
Ex-type culture: HMAS 248884.
Type locality: China, Hunan, Hengyang, Gouloufeng.
Type substrate: Twigs.
Descriptions and illustrations: See Zeng & Zhuang (2017b).
Diagnostic DNA barcodes: tef1: KY829448.
herbarum Fusarium (Corda) Fr., Summa Veg. Scand. 2: 472.
1849.
Basionym: Selenosporium herbarum Corda, Icon. Fung. 3: 34,
tab. 6, fig. 88. 1839.
(See Fusarium avenaceum)
Typus: PRM 155731.
Type locality: Czech Republic, Prague.
Type substrate: Gramineous plant part.
Note: Synonyms fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in PRM.
heteronemum Fusarium Berk. & Broome (as ‘heteronema’), Ann.
Mag. Nat. Hist., ser. 3, 15: 402. 1865.
(See Fusarium candidum (Link) Sacc.)
119
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ET AL.
Holotypus: ?K(M).
Type locality: UK, Batheaston.
Type substrate: Decaying Pyrus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
heterosporioides Fusarium Fautrey, in Roumeguere, Fungi Sel.
Gall. Exs. No. 5399. 1890 and Rev. Mycol. (Toulouse) 12: 126.
1890.
Syntype: ILL00219542 (Roumeguere, Fungi Sel. Gall. Exs no.
5399).
Type locality: France, Charny
Type substrate: Sclerotium clavus on Glyceria fluitans.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
heterosporum Fusarium Nees & T. Nees, Nova Acta Phys.Med. Acad. Caes. Leop.-Carol. Nat. Cur. 9: 235. 1818.
Synonyms: ?Fusarium leucoconium Corda, Icon. Fung. 1: 4.
1837. [fide Booth (1971)].
Sphaeria cyanea Sollm., Bot. Zeitung (Berlin) 21: 193. 1863.
Botryosphaeria cyanea (Sollm.) Weese, Sitzungsber. Kaiserl.
Akad. Wiss. Wien, Math.-Naturwiss. Cl., Abt. 1, 128: 707. 1919.
Gibberella cyanea (Sollm.) Wollenw., Fusaria Autogr. Delin. 1:
39. 1919.
Fusarium secalis Fee, Mem. Soc. Mus. Hist. Nat. Strassbourg 3:
35. 1843.
Fusarium eleocharidis Rostr. (as ‘heleocharidis’), in Thümen,
Mycoth. Univ., Cent. 22, no. 2185. 1883.
Fusisporium lolii Wm.G. Sm., Diseases of field and garden crops,
chiefly as are caused by fungi: 213. 1884.
Fusarium lolii (Wm.G. Sm.) Sacc., Syll. Fung. 11: 652. 1895.
Fusarium heterosporum var. lolii (Wm.G. Sm.) Wollenw., Z.
Parasitenk. (Berlin) 3: 349. 1931.
Fusarium heterosporum f. paspali Ellis & Everh., in Ellis, North
Amer. Fung., Ser. 2, no. 2395. 1886.
Fusarium parasiticum Ellis & Kellerm., J. Mycol. 3: 127. 1887,
nom. illegit., Art. 53.1.
Fusarium pucciniophilum Sacc. & P. Syd., Syll. Fung. 14: 1128.
1899.
Fusarium stromaticum Delacr., Bull. Soc. Mycol. France 9: 186.
1893.
Fusarium paspalicola Henn., Monsunia 1: 38. 1899 [1900].
Fusarium heterosporum var. paspalicola (Henn.) Wollenw., Z.
Parasitenk. (Berlin) 3: 349. 1931.
Fusarium congoense Wollenw., Fusaria Autogr. Delin. 1: 307. 1916.
Fusarium heterosporum var. congoense (Wollenw.) Wollenw., Z.
Parasitenk. (Berlin) 3: 350. 1931.
Fusarium heterosporum f. aleuritis Saccas & Drouillon (as
‘aleuritidis’), Agron. Trop. 6: 251. 1951.
Gibberella gordonii C. Booth, The Genus Fusarium: 177. 1971.
Lectotypus (hic designatus, MBT 10000692): Germany, sclerotium of Claviceps purpurea on a spike of Triticum sp., 1818,
G.C.D. Nees von Esenbeck, in Nova Acta Phys.-Med. Acad.
Caes. Leop.-Carol. Nat. Cur., tab. V. fig. 5.
Epitypus (hic designatus, MBT 10000693): Germany, Rotenburg
near Bremen, sclerotium of Claviceps purpurea on Lolium perenne, Aug. 1967, U.G. Schlösser, CBS 391.68 (preserved as
metabolically inactive culture).
Ex-epitype culture: CBS 391.68 = NRRL 25798.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982) and Leslie
& Summerell (2006).
120
Diagnostic DNA barcodes: rpb1: MW928811; rpb2: MW928827;
tef1: MW928839.
Notes: This species is recognised by Wollenweber & Reinking
(1935), Gerlach & Nirenberg (1982), Booth (1971), and Leslie
& Summerell (2006). Index Fungorum indicates that the correct name for this species is F. lolii. However, this name is not
commonly used and considered as a synonym of
F. heterosporum. Additionally, the epithet ‘heterosporum’ is older
than the epithet ‘lolii’ and should have priority. No holotype
specimen is available and therefore an illustration is designated
as lectotype.
heveae Fusarium Vincens, Bull. Soc. Pathol. Veg. France 2: 19.
1915.
(See Fusarium incarnatum)
Holotypus: ?PC.
Type locality: Brazil, Para.
Type substrate: Hevea brasiliensis.
Note: Synonym fide Wollenweber & Reinking (1935).
hexaseptatum Fusarium Maryani et al., Stud. Mycol. 92: 183.
2018 [2019].
Holotypus: InaCC F866 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F866.
Type locality: Indonesia, West Java, Sukabumi, Parakan Lima.
Type substrate: Musa acuminata var. Pisang Ambon Kuning.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb2: LS479359; tef1: LS479805.
hibernans Fusarium Lindau, Rabenh. Krypt.-Fl., ed. 2, 1(9): 542.
1909, nom. superfl., Art. 52.1.
Basionym: Fusarium nivale Ces. ex Berl. & Voglino, in Saccardo,
Syll. Fung., Addit. I–IV: 390. 1886, non (Fr.) Sorauer, 1901.
(See Fusarium nivale)
Authentic material: Klotzsch, Herb. Viv. Mycol. no. 1439 in HAL.
Original locality: Italy, Vercelli.
Original substrate: Leaves of overwintered crop.
Note: Synonyms fide Wollenweber & Reinking (1935).
hippocastani Fusarium (Corda) Sacc., Syll. Fung. 4: 703. 1886.
Basionym: Selenosporium hippocastani Corda, Icon. Fung. 2: 7.
1838.
(See Fusarium acuminatum)
Lectotypus (hic designatus, MBT 10000694): Czech Republic,
Prague, Aesculus hippocastanum, 1836, A.C.J. Corda, in Icon.
Fung. 2: tab. IX. fig. 31.
Notes: According to Pilat (1938) and Holubova-Jechova et al.
(1994), no material was preserved in PRM. Therefore, an illustration is selected as lectotype.
hoodiae Fusarium L. Lombard et al., Persoonia 41: 27. 2018
[2019].
Holotypus: CBS H-23616.
Ex-type culture: CBS 132474.
Type locality: South Africa, Northern Cape Province, Prieska.
Type substrate: Root of Hoodia gordonii.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb2: MH484929; tef1: MH485020.
hordearium Fusarium Ducomet, Rech. Devel. Champ. Parasit.:
87. 1907.
Holotypus: ?MPA.
Type locality: France.
FUSARIUM
Type substrate: Unknown.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
hordei Fusarium (Wm.G. Sm.) Sacc., Syll. Fung. 11: 652. 1895.
Basionym: Fusisporium hordei Wm.G. Sm., Diseases of field and
garden crops, chiefly as are caused by fungi: 212. 1884.
(See Fusarium sambucinum)
Lectotypus (hic designatus, MBT 10000695): Denmark, Hordeum sp., 1884, W. G. Smith, in Diseases of field and garden
crops, chiefly as are caused by fungi: 211, fig. 94.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype material could be located and therefore an illustration is
designated as lectotype.
hostae Fusarium Geiser & Juba, Mycologia 93: 672. 2001.
Synonym: Gibberella hostae Geiser & Juba, Mycologia 93: 672.
2001.
Holotypus: BPI 748169.
Ex-type culture: FRC O-2074 = NRRL 29889.
Type locality: USA, South Carolina.
Type substrate: Hosta sp.
Descriptions and illustrations: See Geiser et al. (2001).
Diagnostic DNA barcodes: rpb1: JX171527; rpb2: JX171640;
tef1: AY329034.
humi Fusarium (Reinking) Nirenberg & Hagedorn, Nachrichtenbl. Deutsch. Pflanzenschutzdienstes 60: 215. 2008.
Basionym: Fusarium tumidum var. humi Reinking, Zentralbl.
Bakteriol., 2. Abth. 89: 513. 1934.
Lectotypus (hic designatus, MBT 10000706): Honduras, soil,
1931, O.A. Reinking, in Wollenweber's Fusaria Autogr. Delin. no.
1152 of type culture 5236.
Notes: This species is recognised by Wollenweber & Reinking
(1935), Gerlach & Nirenberg (1982), and Nirenberg &
Hagedorn (2008). Recollection from the type host and locality
is required. No holotype material could be located and therefore
an illustration is designated as lectotype.
humicola Fusarium L. Lombard & Crous, Fungal Syst. Evol. 4:
191. 2019.
Holotypus: CBS H-24016.
Ex-type culture: ATCC 24372 = CBS 124.73 = IMI
128101 = NRRL 25535.
Type locality: Pakistan.
Type substrate: Soil.
Descriptions and illustrations: See Lombard et al. (2019a).
Diagnostic DNA barcodes: rpb1: MN120718; rpb2: MN120738;
tef1: MN120757.
humuli Fusarium M.M. Wang et al., Persoonia 43: 83. 2019.
Holotypus: HAMS 248039.
Ex-type culture: CGMCC 3.19374 = CQ1039.
Type locality: China, Jiangsu Province.
Type substrate: Leaves of Humulus scandens.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289840; rpb2: MK289724;
tef1: MK289570.
hydnicola Fusarium Ellis & Everh. (as ‘hydnicolum’), J. Mycol.
4(4–5): 45. 1888.
Alysidium hypophleodes (Corda) Bonord., Handb. Mykol.: 35.
1851.
www.studiesinmycology.org
REDELIMITED
Basionym: Fusidium hypophleodes Corda, Icon. Fung. 1: 3, tab.
1, fig. 50. 1837.
Holotypus: NY (fide Index Fungorum).
Type locality: USA, Missouri, Concordia.
Type substrate: Bark of dead Hydnum membranaceum.
Note: Synonym fide Wollenweber & Reinking (1935).
hymenula Fusarium Pound & Clem., Bot. Surv. Nebraska 4: 7.
1896.
Gloeosporium intermedium var. brevipes Sacc., Syll. Fung. 3:
703. 1884.
Holotypus: NEB0040541.
Type locality: USA, Nebraska.
Type substrate: Helianthus sp.
Notes: Synonym fide Wollenweber & Reinking (1935). The name
is misspelled as ‘lymenula’ in the NEB database.
hyperoxysporum Fusarium Wollenw., J. Agric. Res. 2: 268. 1914.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: USA.
Type substrate: Ipomoea batatas.
Note: Synonym fide Wollenweber & Reinking (1935).
hypocreoideum Fusarium Cooke & Massee, Grevillea 16: 76.
1888.
Aschersonia hypocreoidea (Cooke & Massee) Petch, Ann.
Roy. Bot. Gard. (Peradeniya) 7: 255. 1922.
Holotypus: K(M) 127920.
Type locality: Australia, Queensland.
Type substrate: Ficus aspera.
hypodermium Fusarium (Link) Link, in Willdenow, Sp. Pl., ed. 4,
6: 96. 1825.
Basionym: Fusidium hypodermium Link, Mag. Neuesten
Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 8: 31.
1816 [1815].
Marssonina aurantiaca (Link) Magnus, Hedwigia 45: 90. 1906.
Basionym: Cryptosporium aurantiacum Link, in Willdenow, Sp.
Pl., ed 4, 6: 96. 1825, nom. sanct. (Fries, Syst. Mycol. 3:
481. 1832).
Synonyms: Fusidium aurantiacum (Link) Fr., Syst. Mycol. 3: 481.
1832.
Gloeosporium aurantiacum (Link) Sacc., Syll. Fung. 3: 717.
1884.
Marssonia aurantiaca (Link) Rostr., Bot. Tidsskr. 19: 217. 1895.
Note: Synonyms fide Wollenweber & Reinking (1935).
hypothenemi Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 2. 2020.
Neocosmospora hypothenemi Sand.-Den. & Crous, Persoonia
43: 132. 2019.
Holotypus: CBS H-23982.
Ex-type culture: ARSEF 5878 = CBS 145464 = NRRL 52782.
Type locality: Benin, Niaouli.
Type substrate: Adult Hypothenemus hampei (coffee borer
beetle).
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218117; rpb2: JF741176;
tef1: JF740850.
idahoanum Fusarium O.A. Pratt, J. Agric. Res. 13: 86. 1918.
(See Fusarium flocciferum)
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CROUS
ET AL.
Lectotypus (hic designatus, MBT 10000707): USA, Idaho, soil,
1918, O.A. Pratt, in J. Agric. Res. 13: 87, fig. 2.
Notes: Synonyms fide Wollenweber & Reinking (1935). No holotype material could be located and therefore an illustration is
designated as lectotype.
illosporioides Fusarium Sacc., Harriman Alaska Exped. 5: 15.
1904.
(See Fusarium lateritium)
Holotypus: In PAD.
Type locality: USA, Alaska, Sitka.
Type substrate: Ribes sp.
Note: Synonym fide Wollenweber & Reinking (1935).
illudens Fusarium C. Booth, The Genus Fusarium: 54. 1971.
Neocosmospora illudens (Berk.) L. Lombard & Crous, Stud.
Mycol. 80: 227. 2015.
Basionym: Nectria illudens Berk., in Hooker, Bot. Antarct. Voy. II
(Fl. Nov.-Zel.): 203. 1855.
Synonyms: Cucurbitaria illudens (Berk.) Kuntze, Revis. Gen. Pl.
3: 461. 1898.
Haematonectria illudens (Berk.) Samuels & Nirenberg, Stud.
Mycol. 42: 136. 1999.
Neotypus: PAD S00012, designated in Forin et al. (2020)
Neotype locality: New Zealand.
Neotype substrate: Bark of unknown host plant,
“inaequale Fusarium” Auersw. Bot. Zeitung (Berlin) 8: 439.
1850, typographic error (see Notes).
Ramularia rosea (Fuckel) Sacc., Fungi Ital. Del., Tab. 1001.
1881.
Basionym: Fusidium roseum Fuckel, Fungi Rhen. Fasc. III, no.
219. 1863.
Synonyms: Ovularia rosea (Fuckel) Massee, Brit. Fung.-Fl. 3:
323. 1893.
Cylindrospora rosea (Fuckel) J. Schröt., in Cohn, Krypt.-Fl.
Schles., Pilze II: 493. 1897.
Fusidium inaequale Auersw., in Rabenh., Klotzschii Herb. Viv.
Mycol., Cent. 14: no. 1383. 1850.
Ramularia lucidae Davis, Trans. Wis. Acad. Sci. Art. Lett. 19:
687. 1919.
Authentic material: Rabenh., Klotzschii Herb. Viv. Mycol. 1383 in
HAL.
Original locality: Germany, Leipzig.
Original substrate: Salix amygdalina.
Notes: Not Fusarium fide Wollenweber & Reinking (1935). This
species was first published as Fusidium inaequale Auersw., in
Rabenh., Klotzschii Herb. Viv. Mycol., Cent. 14: no. 1383, 1850.
The description was repeated in Bot. Zeitung 8: 439, 1850 and
Flora 33: 283, 1850 (in the latter publication also under Fusidium), so that in the simultaneous publication in “Botanische
Zeitung” the “F.” was undoubtedly also meant to be Fusidium and
not Fusarium. Syntype material deposited at HAL has recently
been examined, and Fusidium inaequale turned out to be a
heterotypic synonym of Ramularia rosea (Fuckel) Sacc (see
Braun 1998).
incarcerans Fusarium (Berk.) Sacc., Syll. Fung. 4: 713. 1886.
Basionym: Fusisporium incarcerans Berk., Intellectual Observ. 2:
11. 1863.
(See Fusarium avenaceum)
Holotypus: ?K(M).
Type locality: UK, Northamptonshire, Fotheringhay Castle.
Type substrate: Orthotrichum sp.
122
Note: Synonym fide Wollenweber & Reinking (1935).
incarnatum Fusarium (Roberge ex Desm.) Sacc., Syll. Fung. 4:
712. 1886.
Basionym: Fusisporium incarnatum Roberge ex Desm., Ann.
Sci. Nat., Bot., ser. 3, 11: 274. 1849.
Synonyms: Fusarium semitectum Berk. & Ravenel, Grevillea 3:
98. 1875.
Pseudofusarium semitectum (Berk. & Ravenel) Matsush., Icon.
Microfung. Matsush. Lect. (Kobe): 119. 1975.
Fusarium pallens Berk. & M.A. Curtis, Grevillea 3: 99. 1875,
nom. illegit., Art. 53.1, non Fusarium pallens Nees & T. Nees
1818.
Fusarium glumarum Sacc., Syll. Fung. 4: 706. 1886 (nom. nov.
for F. pallens Berk. & M.A. Curtis).
Fusisporium pallidoroseum Cooke, Grevillea 6: 139. 1878.
Fusarium pallidoroseum (Cooke) Sacc., Syll. Fung. 4: 720. 1886.
Fusarium asparagi Briard, Rev. Mycol. (Toulouse) 12: 142. 1890.
Fusarium gloeosporioides Speg. (as ‘gloeosporioide’), Anales
Mus. Nac. Hist. Nat. Buenos Aires 6: 350. 1899.
Fusarium juglandinum Peck, Bull. Torrey Bot. Club 36: 157.
1909.
Fusarium heveae Vincens, Bull. Soc. Pathol. Veg. France 2: 19.
1915.
Fusarium tenuistipes Sacc., Atti Mem. Reale Accad. Sci. Lett.
Arti, Padova 33: 195. 1917.
Fusarium semitectum var. majus Wollenw., Fusaria Autogr.
Delin. 3: 907–910. 1930.
Fusarium semitectum var. violaceum Batikyan & Abramyan (as
‘violaceae’), Biol. Zhurn. Armenii 22: 58. 1969, nom. inval., Art.
39.1.
Lectotypus: (hic designatus, MBT 10001327) France, from
Tagetes erecta, 1848, M. Roberge in Desmazieres, Pl. Crypt. N.
France, ed 2, No. 1303, in PC.
Epitypus: (hic designatus, MBT 10001328) Malawi, on Trichosanthes dioica, date unknown, H.M. Phiri, CBS H-24060.
Ex-epitype culture: ATCC 24387 = CBS 132.73 = IMI
128222 = NRRL 25478.
Descriptions and illustrations: See Booth (1971), Gerlach &
Nirenberg (1982) and Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170409; tef1: MN170476.
Note: The epitypification of Fusarium incarnatum by Xia et al.
(2019) was not effective as the holo- or lectotype was not
correctly indicated (Art. 9.9). Here, a lectotype is selected and
the epitypification is validated.
inflexum Fusarium R. Schneid., in Schneider & Dalchow,
Phytopathol. Z. 82: 80. 1975.
Holotypus: DSM 63203.
Ex-type culture: ATCC 32213 = BBA 63203 = CBS
716.74 = DAOM 225130 = DSM 63203 = IMI 375336 = NRRL
20433.
Type locality: Germany, Hamburg, Vierlanden.
Type substrate: Stem of Vicia faba.
Descriptions and illustrations: See Schneider & Dalchow (1975)
and Gerlach & Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: JX171469; rpb2: JX171583;
tef1: AF008479.
inseptatum Fusarium Schwein., Trans. Amer. Philos. Soc., n.s.,
4: 302. 1832 [1834].
Holotypus: PH00062493.
Type locality: USA, Pennsylvania, Bethlehem.
FUSARIUM
Type substrate: Daphne mezereum.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
insidiosum Fusarium Roum., Michelia 2: 132. 1880.
(See Fusarium lateritium)
Syntypes: In BR, CUP & ILL (Roum., Fungi Sel. Gall. Exs. No.
57).
Type locality: France, Pyrenees-Orientales, Environs de
Perpignan.
Type substrate: Phytolacca decandra.
Note: Synonym fide Wollenweber & Reinking (1935).
insidiosum Fusarium (Berk.) Sacc., Syll. Fung. 4: 707. 1886,
nom. illegit., Art. 53.1.
Basionym: Fusisporium insidiosum Berk., Gard. Chron. 1860:
480. 1860.
(See Fusarium graminearum)
Holotypus: ?K(M).
Type locality: UK.
Type substrate: Agrostis pulchella.
Note: Synonyms fide Wollenweber & Reinking (1935).
ipomoeae Fusarium M.M. Wang et al., Persoonia 43: 83. 2019.
Holotypus: HAMS 248040.
Ex-type culture: CGMCC 3. 19496 = LC12165.
Type locality: China, Jiangsu Province.
Type substrate: Leaves of Ipomoea aquatica.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289859; rpb2: MK289752;
tef1: MK289599.
iranicum Fusarium Torbati et al., Mycol. Progr. 18: 129. 2018
[2019].
Holotypus: CBS H-23560.
Ex-type culture: CBS 143608 = CPC 30860.
Type locality: Iran, West Azerbaijan Province, Orumieh-Salmas.
Type substrate: Agaricus bisporus.
Descriptions and illustrations: See Torbati et al. (2019).
Diagnostic DNA barcodes: rpb2: LT970757; tef1: LT970785.
iridis Fusarium Oudem., Ned. Kruidk. Arch., 2 ser. 5: 515. 1889.
(See Fusarium avenaceum)
Holotypus: ?L.
Type locality: Netherlands.
Type substrate: Iris pseudacorus.
Note: Synonym fide Wollenweber & Reinking (1935).
irregulare Fusarium M.M. Wang et al., Persoonia 43: 84. 2019.
Holotypus: HAMS 248041.
Ex-type culture: CGMCC 3.19489 = LC7188.
Type locality: China, Guangdong Province.
Type substrate: Bambusoideae.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289863; rpb2: MK289783;
tef1: MK289629.
japonicum Fusarium Allesch., Beibl. Hedwigia 36: (164). 1897.
(See Fusarium tortuosum)
Syntype: S-F45631 (Sydow, Mycoth. March. no. 4592).
Type locality: Germany, Berlin.
Type locality: Prunus japonica.
Note: Synonym fide Wollenweber & Reinking (1935).
javanicum Fusarium Koord., Verh. Kon. Akad. Wetensch., Afd.
Natuurk., Sect. 2, 13: 247. 1907.
www.studiesinmycology.org
REDELIMITED
Holotypus: Not located.
Type locality: Indonesia, Central Java, Purworejo.
Type substrate: Ficus elastica.
Note: Status unclear fide Sandoval-Denis et al. (2019).
juglandinum Fusarium Peck, Bull. Torrey Bot. Club 36: 157. 1909.
(See Fusarium incarnatum)
Holotypus: NYSf1607.
Type locality: USA, Kansas, Rooks, Stockton.
Type substrate: Juglans nigra.
Note: Synonym fide Wollenweber & Reinking (1935).
junci Fusarium P. Crouan & H. Crouan, Fl. Finistere: 14. 1867.
Holotypus: ?CO.
Type locality: France, Paris.
Type substrate: Juncus effusus.
Note: ?Fusidium fide Wollenweber & Reinking (1935).
jungiae Fusarium Pat., Bull. Soc. Mycol. France 11: 234. 1895.
(See Fusarium avenaceum)
Holotypus: FH00965356.
Type locality: Argentina, San Jorge.
Type substrate: Parasitic on Puccinia sp. on Jungia sp.
Note: Synonym fide Wollenweber & Reinking (1935).
juruanum Fusarium Henn., Hedwigia 43: 398. 1904.
(See Fusarium coccidicola)
Holotypus: In B.
Type locality: Brazil, Rio Jurua.
Type substrate: Annonaceae sp.
Note: Synonym fide Gerlach & Nirenberg (1982).
kalimantanense Fusarium Maryani et al., Stud. Mycol. 92: 187.
2018 [2019].
Holotypus: InaCC F917 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F917.
Type locality: Indonesia, Central Kalimantan, Katingan, Pulau
Malan.
Type substrate: Musa acuminata var. Pisang Ambon.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479497; rpb2: LS479241;
tef1: LS479690.
kelerajum Fusarium Samuels et al., Mycologia 103: 1326. 2011.
Neocosmospora keleraja Samuels et al., Mycologia 103: 1326.
2011.
Holotypus: BPI 871413.
Ex-type culture: FRC S-1839 = G.J.S. 02-122.
Type locality: Sri Lanka, Minneriya Natl. Forest.
Type substrate: Trunk of Yakuda marang.
Descriptions and illustrations: See Nalim et al. (2011).
Diagnostic DNA barcode: tef1: DQ247518.
keratoplasticum Fusarium Geiser et al., Fung. Gen. Biol. 53: 68.
2013.
Neocosmospora keratoplastica (Geiser et al.) Sand.-Den. &
Crous, Persoonia 41: 120. 2018.
Synonyms: Cephalosporium keratoplasticum T. Morik., Mycopathologia 2: 66. 1939, nom. inval., Art. 39.1.
Hyalopus keratoplasticum T. Morik. ex M.A.J. Barbosa, Subsidios Para o Estudo Parasitologico do Genero Hyalopus Corda,
1838: 19. 1941, nom. inval., Art. 39.1.
Fusarium sedimenticola M.M. Wang et al., Botanica Marina 63:
174. 2020.
123
CROUS
ET AL.
Holotypus: FRC S-2477.
Ex-type culture: CBS 490.63 = FRC S-2477 = NRRL 22661.
Type locality: USA, Virginia, Winchester.
Type substrate: Indoor plumbing.
Descriptions and illustrations: See Nalim et al. (2011).
Diagnostic DNA barcodes: rpb1: MW218121; rpb2: JN235897;
tef1: JN235712.
konzum Fusarium Zeller et al., Mycologia 95: 947. 2003.
Synonym: Gibberella konza Zeller et al., Mycologia 95: 947.
2003.
Holotypus: DAR 76034.
Ex-type culture: CBS 119849 = KSU 10653 = NRRL 53394.
Type locality: USA, Kansas, Manhattan, Konza Praire Biological
Station.
Type substrate: Sorghastrum nutans.
Descriptions and illustrations: See Zeller et al. (2003) and Leslie
& Summerell (2006).
Diagnostic DNA barcodes: rpb1: LT996200; rpb2: LT996148;
tef1: LT996098.
kotabaruense Fusarium Maryani et al., Persoonia 43: 65. 2019.
Holotypus: InaCC F963 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F963.
Type locality: Indonesia, South Kalimantan, Kota Baru, Kecamatan Pamukan Barat, Desa Sungai Birah.
Type substrate: Musa var. Pisang Hawa.
Descriptions and illustrations: See Maryani et al. (2019b).
Diagnostic DNA barcodes: rpb1: LS479875; rpb2: LS479859;
tef1: LS479445.
kuehnii Fusarium (Fuckel) Sacc., Syll. Fung. 4: 714. 1886.
Basionym: Fusisporium kuehnii Fuckel, Fungi Rhen. Exs.,
Suppl., Fasc. 5, no. 1920. 1867.
?Athelia arachnoidea (Berk.) Jülich, Willdenowia 7: 53.
1972. (fide Gerlach & Nirenberg 1982)
Basionym: Corticium arachnoideum Berk., Ann. Mag. Nat. Hist.,
ser. 1, 13: 345. 1844.
Synonym: Fusisporium devastans J.G. Kühn, Krankh. Kulturgew.: 32. 1858, nom. inval., Art. 38.1(a).
Syntype: Fuckel, Fungi Rhen. Exs., Suppl., Fasc. 5, 1920 (e.g.,
HAL).
Type locality: Germany.
Type substrate: Lichens and mosses.
Notes: Status doubtful. Considered a possible synonym of
F. dimerum by Booth (1971).
kurdicum Fusarium Petr., Sydowia 13: 96. 1959.
Cosmospora kurdica (Petr.) Rossman & Samuels, Stud. Mycol.
42: 122. 1999.
Basionym: Calonectria kurdica Petr., Sydowia 13: 95. 1959.
Synonyms: Nectria kurdica (Petr.) Rossman, Mycol. Pap. 150:
35. 1983.
?Stagonopsis sclerotioides Höhn., Ann. K. K. Naturhist. Hofmus.
20: 368. 1905.
?Botryocrea sclerotioides (Höhn.) Petr., Sydowia 3: 141. 1949.
?Fusarium sclerotioides (Höhn.) Samuels & Rossman, Mycol.
Pap. 164: 23. 1991.
Holotypus: K.H. Rechinger, 31 Jul. 1957, in W.
Type locality: Iran, Kurdistan.
Type substrate: Astragalus sp.
Note: Synonyms fide Rossman et al. (1999).
124
kuroshium Fusarium F. Na et al., Plant Disease 102: 1159. 2018,
nom. inval., Art. 40.7.
Neocosmospora kuroshio F. Na et al. ex Sand.-Den. & Crous,
Persoonia 43: 137. 2019.
Holotypus: BPI 910340.
Ex-type culture: CBS 142642 = UCR 3641.
Type locality: USA, California, San Diego, El Cajon.
Type substrate: Euwallacea sp. galleries in Platanus racemosa.
Descriptions and illustrations: See Na et al. (2018).
Diagnostic DNA barcodes: rpb1: KX262236; rpb2: KX262256;
tef1: KX262216.
kurunegalense Fusarium Samuels et al., Mycologia 103: 1323.
2011.
Neocosmospora kurunegalensis Samuels et al., Mycologia
103: 1324. 2011.
Holotypus: BPI 871391.
Ex-type culture: CBS 119599 = G.J.S. 02-94.
Type locality: Sri Lanka, Wagamba Province, Kurunegala.
Type substrate: Recently felled tree.
Descriptions and illustrations: See Nalim et al. (2011).
Diagnostic DNA barcodes: rpb1: MW834228; rpb2: LR583838;
tef1: DQ247511.
kyushuense Fusarium O'Donnell & T. Aoki, Mycoscience 39: 2.
1998.
Holotypus: NIAES99701.
Ex-type culture: ATCC 56750 = FRC T-346A = MAFF
237645 = MRC 1767 = NRRL 3509.
Type locality: Japan, Kumamoto.
Type substrate: Seed of Triticum aestivum.
Descriptions and illustrations: See Aoki & O'Donnell (1998).
Diagnostic DNA barcodes: rpb2: MH582098; tef1: MH582292.
laboulbeniae Fusarium Cepede, Arch. Parasitol. 16: 373. 1914.
(See Fusarium larvarum)
Holotypus: Not located.
Type locality: France, Pas-de-Calais, Wimereux.
Type substrate: Demetrias unipunctata.
Note: Synonym fide Wollenweber & Reinking (1935).
lacertarum Fusarium Subrahm. (as ‘laceratum’), Mykosen 26:
478. 1983.
Holotypus: IMI 300797.
Ex-type culture: ATCC 42771 = CBS 130185 = IMI
300797 = NRRL 20423.
Type locality: India, Poona, Pimpri.
Type substrate: Skin of lizard.
Descriptions and illustrations: See Subrahmanyam (1983).
Diagnostic DNA barcodes: rpb1: JX171467; rpb2: JX171581;
tef1: GQ505593.
lactis Fusarium Pirotta, Arch. Lab. Bot. Crittog. Univ. Pavia 2 &
3: 316. 1879.
Synonyms: ?Fusarium pyrinum Schwein., Trans. Amer. Philos.
Soc., n.s. 4: 302. 1834.
?Fusarium apiogenum Sacc., Syll. Fung. 4: 717. 1886.
Fusarium rubrum Parav., Ann. Mycol. 16: 311. 1918.
Lectotypus: Arch. Lab. Bot. Crittog. Univ. Pavia 2 & 3, Tab. 21,
figs 1–6, designated by Yilmaz et al. (2021).
Lectotype locality: Italy, Pavia.
Lectotype substrate: Clotted milk.
Epitypus: B 70 0001686, designated by Yilmaz et al. (2021).
FUSARIUM
Ex-epitype culture: BBA 68590 = CBS 411.97 = IMI
375351 = NRRL 25200.
Epitype locality: USA, California.
Epitype substrate: Ficus carica.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998)
and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: LT996201; rpb2: LT996149;
tef1: AF160272.
lagenariae Fusarium (Schwein.) Sacc., Syll. Fung. 4: 724. 1886.
Basionym: Fusisporium lagenariae Schwein., Trans. Amer.
Philos. Soc., n.s., 4: 275. 1834.
(See Fusarium oxysporum)
Holotypus: PH00062516
Type locality: USA, Pennsylvania, Bethlehem.
Type substrate: Lagenaria siceraria.
lagenarium Fusarium Pass., Erb. Critt. Ital., ser. 2: no. 148. 1871.
Synonym: Gloeosporium lagenarium (Pass.) Sacc. & Roum.,
Rev. Mycol., Toulouse 2(8): 201. 1880.
(See Fusarium cyclogenum)
Holotypus: In PAD.
Type locality: Italy, Parma.
Type substrate: Lagenaria sp.
Note: Synonym fide Wollenweber & Reinking (1935).
lanceolatum Fusarium O.A. Pratt, J. Agric. Res. 13: 83. 1918.
(See Fusarium acuminatum)
Lectotypus (hic designatus, MBT 10000709): USA, Idaho, from
soil, 1918, O.A. Pratt, in J. Agric. Res. 13: 82, fig. 1A–E.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
langsethiae Fusarium Torp & Nirenberg, Int. J. Food Microbiol.
95: 248. 2004.
Holotypus: B 70 0012234.
Ex-type culture: BBA 70945 = CBS 113234.
Type locality: Norway.
Type substrate: Kernal of Avena sativa.
Descriptions and illustrations: See Torp & Nirenberg (2004).
Diagnostic DNA barcodes: rpb1: MW928812; rpb2: MW928828;
tef1: AB674298.
languescens Fusarium L. Lombard & Crous, Persoonia 43: 28.
2018 [2019].
Holotypus: CBS H-23617.
Ex-type culture: CBS 645.78 = NRRL 36531.
Type locality: Morocco.
Type substrate: Solanum lycopersicum.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928813; rpb2: MH484880;
tef1: MH484971.
laricis Fusarium Sawada, Bull. Gov. Forest Exp. Sta., Meguro
46: 130. 1950.
Holotypus: TFM:FPH 00771.
Type locality: Japan, Aomori, Kamikita, Noheji
Type substrate: Larix kaempferi.
larvarum Fusarium Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 369. 1870.
Microcera larvarum (Fuckel) Gr€afenhan et al., Stud. Mycol. 68:
105. 2011.
Synonyms: Fusarium nivale var. larvarum (Fuckel) Bilaĭ, Fusarii
(Biologija i sistematika): 295. 1955, nom. inval., Art. 41.1
www.studiesinmycology.org
REDELIMITED
Fusarium cryptum McAlpine, Fungus Diseases of Citrus trees in
Australia: 106. 1899.
Fusarium epicoccum McAlpine, Fungus Diseases of Citrus trees
in Australia: 113. 1899.
Microcera parlatoriae Trab., Bull. Agric. Algerie Tunisie 13: 33.
1907.
Microcera curta Sacc., Ann. Mycol. 7: 437. 1909.
Microcera tonduzii Pat., Bull. Soc. Mycol. France 28: 142. 1912.
Fusarium aspidioti Sawada, Bot. Mag. (Tokyo) 28: 312. 1914.
Fusarium laboulbeniae Cepede, Arch. Parasitol. 16: 373. 1914.
Fusarium acremoniopsis Vincens, Bull. Soc. Mycol. France 31:
26. 1915.
?Fusarium meliolicola F. Stevens, Bot. Gaz. 65: 245. 1918.
?Nectria meliolicola F. Stevens, Bot. Gaz. 65: 231. 1918.
Microcera aurantiicola Petch, Trans. Brit. Mycol. Soc. 7: 158.
1921.
Lectotypus: G 00111015, selected in Gr€afenhan et al. (2011).
Lectotype locality: Germany, Hessen, Rheingau, near OestrichWinkel.
Lectotype substrate: Larva cuticles of insects on Malus
domestica.
Epitypus: BBA 62239, designated in Gr€afenhan et al. (2011).
Ex-epitype culture: BBA 62239 = CBS 738.79 = MUCL
19033 = NRRL 20473.
Epitype locality: Iran, Gilan Province, near Rasht.
Epitype substrate: Parasitic on Quadraspidiotus perniciosus
(scale) on Prunus domestica.
Diagnostic DNA barcodes: rpb1: KM232252; rpb2: KM232387;
tef1: KM231957.
lateritium Fusarium Nees, Syst. Pilze: 31. 1817.
Synonyms: Selenosporium lateritium (Nees) Desm., Fl. Cryptog.
Flandres 2: 99. 1867.
Fusarium microsporum Schltdl., Fl. Berol. 2: 139. 1824.
Fusarium fructigenum Fr., Syst. Mycol. 3: 471. 1832.
Fusarium lateritium var. fructigenum (Fr.) Wollenw., Fusaria
Autogr. Delin. 3: 959. 1930.
Sphaeria baccata Wallroth, Fl. Crypt. Germ. 2: 838. 1833.
Gibbera baccata (Wallr.) Fuckel, Jahrb. Nassauischen Vereins
Naturk. 23–24: 167. 1870.
Gibberella pulicaris subsp. baccata (Wallr.) Sacc., Michelia 1 (3):
317. 1878.
Gibberella baccata (Wallr.) Sacc., Syll. Fung. 2: 553. 1883.
Fusarium lateritium var. mori Desm., Ann. Sci. Nat. Bot., ser. 2, 8:
10. 1837.
Selenosporium urticarum Corda (as ‘urticearum’), Icon. Fung. 2:
7. 1838.
Fusarium urticarum (Corda) Sacc., Syll. Fung. 4: 698. 1886.
?Fusarium protractum Lev., Ann. Sci. Nat., Bot., ser. 3, 9: 246.
1848.
Gloeosporium berkeleyi Mont., Ann. Sci. Nat., Bot., ser. 3, 12:
296. 1849.
Fusarium berkeleyi (Mont.) Berk. & Broome, N. Amer. Fung.:
108. 1875.
Botryosphaeria moricola Ces. & De Not., Hedwigia 4: 27.
1865.
Gibberella moricola (Ces. & De Not.) Sacc., Syll. Fung. 2: 553.
1883.
Gibbera euonymi Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 167. 1870.
Gibberella euonymi (Fuckel) Sacc., Michelia 1: 318. 1878.
Hendersonia euonymi (Fuckel) Sacc., Syll. Fung. 2: 556. 1883.
125
CROUS
ET AL.
Selenosporium cydoniae Schulzer, Verh. K.K. Zool.-Bot. Ges.
Wien 21: 1240. 1871.
Fusarium cydoniae (Schulzer) Sacc. & Traverso, Syll. Fung. 19:
724. 1910, nom. illegit., Art. 53.1.
Fusarium sticticum Berk. & M.A. Curtis, Grevillea 3: 99. 1875.
Fusisporium zavianum Sacc., Michelia 1: 83. 1877.
Fusarium zavianum (Sacc.) Sacc., Syll. Fung. 4: 709. 1886.
Fusarium cydoniae Roum. & Fautrey, Rev. Mycol. (Toulouse) 14:
170. 1892, nom. illegit., Art. 53.1.
Fusarium salicis Fuckel, Fungi Rhen. Exs., Suppl., Fasc. 7, no.
2110. 1868.
Fusarium salicis var. minus Wollenw., Fusaria Autogr. Delin. 2:
582. 1924.
Fusarium sambucinum var. minus Wollenw., Fusaria Autogr.
Delin. 3: 941. 1930.
Gibbera mori Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 168. 1870.
Fusarium semitectum Berk. & Ravenel, in Berkeley, Grevillea 3:
98. 1875.
Fusisporium cinnabarinum Berk. & M.A. Curtis, Grevillea 3: 146.
1875.
Fusarium cinnabarinum (Berk. & M.A. Curtis) Sacc., Syll. Fung.
4: 722. 1886.
Fusisporium miniatum Berk. & M.A. Curtis, Grevillea 3: 147.
1875.
Fusarium miniatum (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
722. 1886, nom. illegit., Art. 53.1.
Fusisporium putaminum Thüm., Oesterr. Bot. Z. 27: 272. 1877.
Fusarium putaminum (Thüm.) Sacc., Syll. Fung. 4: 703. 1886.
Fusisporium leguminum Cooke, Grevillea 6: 139. 1878.
Fusarium leguminum (Cooke) Sacc., Syll. Fung. 4: 712. 1886.
Fusarium limonis Briosi, Att. Staz. Chim. Agrar. Rome. 1878.
Fusarium yuccae Cooke, Grevillea 7: 34. 1878, nom. inval., Art.
36.1(a).
Fusisporium azedarachinum Thüm., Mycoth. Univ., cent. 14: no.
1379. 1879.
Fusarium azedarachinum (Thüm.) Sacc., Syll. Fung. 4: 704.
1886.
Fusarium insidiosum Roum., Michelia 2: 132. 1880.
Fusarium roumeguerei Sacc. (as ‘roumegueri’), Syll. Fung. 4:
702. 1886, nom. illegit., Art. 52.1.
Fusarium albertii Roum., Fungi Sel. Gall. Exs., Cent. 19: no.
1867. 1881.
Fusarium rimicola Sacc. (as ‘rimicolum’), Michelia 2: 297. 1881.
Fusarium ziziphinum Pass., Rev. Mycol., (Toulouse) 4: 22. 1882.
Fusarium acaciae Cooke & Harkn., Grevillea 12: 96. 1884.
Fusarium longisporum Cooke & Massee, Grevillea 16: 4. 1887.
Fusarium sphaeroideum Pass., Atti Reale Accad. Lincei, Rendiconti Cl. Sci. Fis., ser. 4, 4: 105. 1888.
Fusarium parasiticum Fautrey, Rev. Mycol. (Toulouse) 11: 153.
1889, nom. illegit., Art. 53.1.
Fusarium fautreyi Sacc., Syll. Fung. 10: 934. 1892.
Fusarium carneoroseum Cooke, Grevillea 19: 4. 1890.
Fusarium celtidis Ellis & Tracy, J. Mycol. 6: 76. 1890.
Fusarium nucicola P. Karst. & Har., Rev. Mycol. (Toulouse) 12:
131. 1890.
Fusarium discoideum Fautrey & Roum., Rev. Mycol. (Toulouse)
13: 173. 1891.
Fusarium cydoniae Allesch., Ber. Bot. Vereines Landshut 12:
130. 1892.
?Fusarium luteum Clem., Bot. Surv. Nebraska 3: 12. 1894.
126
Fusarium asclepiadeum Fautrey, Rev. Mycol. (Toulouse) 18: 68.
1896.
Fusarium samararum Allesch., Ber. Bayer. Bot. Ges. 4: 39. 1896.
Fusarium sophorae Allesch., Beibl. Hedwigia 36: (164). 1897.
Fusarium ailanthinum Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 6: 350. 1899.
Fusarium euonymi Syd., Beibl. Hedwigia 39: (6). 1900.
Fusarium gemmiperda Aderh., Z. Pflanzenkrankh. 11: 70. 1901.
Fusarium euonymi-japonici Henn., Hedwigia 41: 139. 1902.
Fusarium illosporioides Sacc., in Saccardo et al., Harriman
Alaska Expedition 5: 15. 1904.
Fusarium schawrowi Speschnew, Arbeiten Kaukas. Stat. Seidenzucht 10: 30–41. 1906.
Selenosporium gloeosporioides Speg. (as ‘gloesporioides’),
Anales Mus. Nac. Hist. Nat. Buenos Aires 13: 458. 1911.
Fusarium gloeosporioides (Speg.) Sacc. & Trotter, Syll. Fung. 22:
1482. 1913, nom. illegit., Art. 53.1.
Fusarium briosianum Ferraris, Fl. Ital. Crypt. Fungi Fasc. 13:
857. 1912.
Fusarium pseudacaciae Rapaics, Z. Pflanzenkrankh. 25: 208.
1915.
Fusarium gleditschiicola Dearn. & Barthol. (as ‘gleditschicola’),
Mycologia 9: 363. 1917.
Gibberella briosiana Turconi & Maffei, Atti Ist. Bot. Univ. Pavia,
ser. 2, 15: 148. 1918.
Botryosphaeria briosiana (Turconi & Maffei) Weese, Sitzungsber.
Akad. Wiss. Wien, Math.-Naturwiss. Kl., Abt. 1, 128: 708. 1919.
Fusarium uncinatum Wollenw., Ann. Mycol. 15(1/2): 54. 1917.
Fusarium blackmannii W. Br. & A.S. Horne (as ‘blackmanni’),
Ann. Bot. (London) 38: 379. 1924.
Fusarium entomophilum Petch, Trans. Brit. Mycol. Soc. 11: 260.
1925.
Fusarium lateritium var. tenue Wollenw., Fusaria Autogr. Delin. 3:
955. 1930.
Gibberella saubinetii var. flacca Wollenw., Z. Parasitenk. (Berlin)
3: 433. 1931.
Fusarium anisophilum Picado, J. Dept. Agric. Porto Rico 16: 391.
1932.
Lectotypus (hic designatus, MBT 10000710): Germany, unknown host, 1817, G.C.D. Nees von Esenbeck, in System der
Pilze und Schw€amme: 31, tab. 2, fig. 26.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982), Nelson
et al. (1983) and Leslie & Summerell (2006).
Notes: Re-collection from the type host and locality is required.
No holotype specimen could be located and therefore an illustration was designated as lectotype.
laxum Fusarium Peck, Bull. New York State Mus. Nat. Hist. 67:
30. 1903.
(See Fusarium oxysporum)
Holotypus: NYS-F-001667.
Type locality: USA, New York, Albany, Delmar.
Type substrate: Equisetum hyemale.
Note: Synonym fide Wollenweber & Reinking (1935).
leguminum Fusarium (Cooke) Sacc., Syll. Fung. 4: 712. 1886.
Basionym: Fusisporium leguminum Cooke, Grevillea 6: 139.
1878.
(See Fusarium lateritium)
Syntypes: In CUP, ISC, NEB & PH (Fungi Amer. Exs. no. 298).
Type locality: USA, South Carolina, Aiken.
FUSARIUM
Type substrate: Acacia sp.
Note: Synonym fide Wollenweber & Reinking (1935).
leucoconium Fusarium Corda, Icon. Fung. 1: 4. 1837.
(See Fusarium heterosporum and F. reticulatum)
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, Prague.
Type substrate: Rotten plants.
Note: Synonym fide Wollenweber & Reinking (1935) and Booth
(1971). Lectotypification pending study of material lodged in PRM.
libertatis Fusarium L. Lombard & Crous, Persoonia 43: 29.
2018 [2019].
Holotypus: CBS H-23618.
Ex-type culture: CBS 144749 = CPC 28465.
Type locality: South Africa, Western Cape Province, Robben
Island, Van Riebeeck's Quarry.
Type substrate: Rock surface.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb2: MH484944; tef1: MH485035.
lichenicola Fusarium C. Massal., in Maire & Saccardo, Ann.
Mycol. 1: 223. 1903.
Neocosmospora lichenicola (C. Massal.) Sand.-Den. & Crous,
Persoonia 41: 120. 2018.
Synonyms: Bactridium lichenicola (C. Massal.) Wollenw. (‘as
lichenicolum’), Fusaria Autogr. Delin. 1: 456. 1916.
Cylindrocarpon lichenicola (C. Massal.) D. Hawksw., Bull. Brit.
Mus. (Nat. Hist.), Bot. 6: 273. 1979.
Selenosporium lichenicola Speg., Anales Mus. Nac. Hist. Nat.
Buenos Aires, ser. 3, 13: 459. 1911.
Fusarium lichenicola (Speg.) Sacc. & Trotter, Syll. Fung. 22:
1486. 1913, nom. illegit., Art. 53.1.
Monacrosporium tedeschii A. Agostini (as ‘tedeschi’), Atti Ist. Bot.
Univ. Lab. Crittog. Pavia, ser. 3, 4: 195. 1933.
Euricoa dominguiesii Bat. & H. Maia, Anais Soc. Biol. Pernambuco 13: 152. 1955.
Hyaloflorea ramosa Bat. & H. Maia, Anais Soc. Biol. Pernambuco 13: 155. 1955.
Neocosmospora ramosa (Bat. & H. Maia) L. Lombard & Crous,
Stud. Mycol. 80: 227. 2015.
Mastigosporium heterosporum R.H. Petersen, Mycologia 51:
729. 1959.
Holotypus: In PAD.
Epitypus: CBS H-23983, designated in Sandoval-Denis et al.
(2019).
Ex-epitype culture: CBS 623.92.
Epitype locality: Germany, Göttingen.
Epitype substrate: Necrotic wounds of Homo sapiens under
chemotherapy.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb2: LR583845; tef1: LR583620.
limonis Fusarium Briosi, Ann. R. Staz. Chim.-Agrar. Sper. Roma.
1878.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Italy, Sicily.
Type substrate: Citrus limon.
Notes: Synonym fide Wollenweber & Reinking (1935). Protologue not located.
limosum Fusarium Rostr., Bot. Tidsskr. 22: 263. 1899.
www.studiesinmycology.org
REDELIMITED
(See Fusarium avenaceum)
Holotypus: C-F-111719.
Type locality: Sweden.
Type substrate: Mixture of lime and sugar.
Note: Synonym fide Wollenweber & Reinking (1935).
lineare Fusarium Moesz, Bot. Közlem. 19: 57. 1920.
(See Fusarium obtusisporum)
Holotypus: ?BP.
Type locality: Hungary.
Type substrate: Staphylea pinnata.
Note: Synonym fide Wollenweber & Reinking (1935).
lini Fusarium Bolley, Proc. Annual Meeting Soc. Promot. Agric.
Sci. 22: 42. 1901.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: USA.
Type substrate: Linum usitatissimum.
lini Fusarium Remer, Jahresber. Schles. Ges. Vaterl. Cult. 80: 25.
1903, nom. illegit., Art. 53.1
Holotypus: Not located.
Type locality: Poland.
Type substrate: Linum sp.
liriodendri Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 2. 2020.
Neocosmospora liriodendri Sand.-Den. & Crous, Persoonia
43: 139. 2019.
Holotypus: CBS H-23984.
Ex-type culture: BBA 67587 = CBS 117481 = G.J.S 91148 = NRRL 22389.
Type locality: USA, Maryland.
Type substrate: Liriodendron tulipifera.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218124; rpb2: EU329506;
tef1: AF178340.
loliaceum Fusarium Ducomet, Ann. Ecole
Natl. Agric. Rennes 2:
14. 1909.
(See Fusarium nivale)
Holotypus: ?MPA.
Type locality: France.
Type substrate: Unknown.
Note: Synonym fide Wollenweber & Reinking (1935).
lolii Fusarium (Wm.G. Sm.) Sacc., Syll. Fung. 11: 652. 1895.
Basionym: Fusisporium lolii Wm.G. Sm., Diseases of field and
garden crops, chiefly as are caused by fungi: 213. 1884.
(See Fusarium heterosporum)
Lectotypus (hic designatus, MBT 10000711): UK, Lolium perenne, date unknown, W.G. Smith, in W.G. Smith, Diseases of
field and garden crops, chiefly as are caused by fungi: 213, fig.
96.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
loncheceras Fusarium Sideris, Phytopathology 14: 213. 1924.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000712): USA, California,
Stockton, roots of Allium cepa, 1924, C.P. Sideris, in Phytopathology 14, pl. XI, fig. of F. loncheceras.
127
CROUS
ET AL.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
longicaudatum Fusarium J.W. Xia et al., Persoonia 43: 208.
2019.
Holotypus: CBS H-24061.
Ex-type culture: ATCC 24370 = CBS 123.73 = IMI
160825 = NRRL 25477.
Type locality: Tanzania, Tropical Products Research Inst.
Type substrate: Unknown.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170414; tef1: MN170481.
longicornicola Fusarium Sand.-Den., et al., Persoonia 46: 149.
2021.
Holotypus: CBS H-24661.
Ex-type culture: ARSEF 6455 = CBS 147247 = NRRL 52706.
Type locality: Ethiopia, Kobo, Welo.
Type substrate: Aiolopus longicornis.
Descriptions and illustrations: See Yilmaz et al. (2021).
Diagnostic DNA barcodes: rpb2: JF741114; tef1: JF740788.
longifundum Fusarium J.W. Xia et al., Persoonia 43: 208.
2019.
Holotypus: CBS H-24062.
Ex-type culture: CBS 235.79 = NRRL 36372.
Type locality: Netherlands Antilles, Curaçao.
Type substrate: Air.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: GQ505827; tef1: GQ505649.
longipes Fusarium Wollenw. & Reinking, Phytopathology 15:
160. 1925.
Synonyms: Fusarium scirpi var. longipes (Wollenw. & Reinking)
Wollenw., Z. Parasitenk. (Berlin) 3: 337. 1931.
Fusarium equiseti var. longipes (Wollenw. & Reinking) Joffe,
Mycopathol. Mycol. Appl. 53: 221. 1974.
Neotypus (hic designatus, MBT 10000713): USA, Florida, soil,
1977, W. Gams, CBS 476.77 (preserved as metabolically inactive culture).
Ex-neotype culture: CBS 476.77 = NRRL 20695.
Descriptions and illustrations: See Gerlach & Nirenberg (1982),
Nelson et al. (1983).
Diagnostic DNA barcodes: rpb1: MW233244; rpb2: GQ915493;
tef1: GQ915509.
Notes: This species is recognised by Gerlach & Nirenberg
(1982), Nelson et al. (1983), and Leslie & Summerell (2006).
No holotype specimen could be located and no illustration
accompanied the original protologue. Although an illustration of
the original culture (O.A. Reinking no. R34) is provided in Wollenweber's Fusaria Autogr. Delin. no. 937 (1924), this cannot be
used to designate a lectotype as it does not form part of the
original protologue. Therefore, isolate CBS 476.77 is designated
as neotype here to provide taxonomic stability to this species, as
it appears to have a paraphyletic phylogenetic structure
(O'Donnell et al. 2013).
longisporum Fusarium Cooke & Massee, Grevillea 16: 4. 1887.
(See Fusarium lateritium)
Holotypus: K(M) 159680.
Type locality: Australia, Queensland, Brisbane.
Type substrate: Twigs of Passiflora sp.
Note: Synonym fide Wollenweber & Reinking (1935).
128
longissimum Fusarium Sacc. & P. Syd., Syll. Fung. 14: 1128. 1899.
Replaced synonym: Fusarium elongatum De Wild., Ann. Soc.
Belge Microscop. 17: 43. 1893, nom. illegit., Art. 53.1, non
Fusarium elongatum Cooke 1890.
(See Fusarium elongatum De Wild.)
Holotypus: Not located.
Type locality: Belgium, Brussels, Botanical Garden.
Type substrate: Submerged plant material.
Note: Synonymy fide Rossman et al. (2016).
longum Fusarium (Wallr.) Sacc., Syll. Fung. 4: 719. 1886.
Basionym: Fusisporium longum Wallr., Fl. Crypt. Germ. 2: 283.
1833.
Holotypus: ?STR.
Type locality: Germany, Berlin.
Type substrate: Dead branch.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
louisianense Fusarium L.R. Gale et al., Fungal Genet. Biol. 48:
1105. 2011.
Holotypus: BPI 881005.
Ex-type culture: CBS 127525 = NRRL 54197.
Type locality: USA, Louisiana.
Type substrate: Seeds of Triticum sp.
Descriptions and illustrations: See Sarver et al. (2011).
Diagnostic DNA barcodes: rpb1: KM889655; rpb2: KM889657;
tef1: KM889633.
lucidum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6:
157. 1915.
(See Fusarium avenaceum)
Typus: ?CUP-007473.
Type locality: USA, New York.
Type substrate: Solanum tuberosum.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in CUP.
lucumae Fusarium Henn., Hedwigia 48: 116. 1908.
Ascochyta lucumae (Henn.) Wollenw., Fusaria Autogr. Delin. 1:
504. 1916.
Syntypes: In BPI, ILL, MIN & WIS (Baker 218).
Type locality: Brazil, Para.
Type substrate: Lucuma rivicoa
Note: Synonym fide Wollenweber & Reinking (1935).
luffae Fusarium M.M. Wang et al., Persoonia 43: 85. 2019
Holotypus: HAMS 248042.
Ex-type culture: CGMCC 3.19497 = LC12167.
Type locality: China, Fujian.
Type substrate: Luffa aegyptiaca.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289869; rpb2: MK289754;
tef1: MK289601.
lumajangense Fusarium Maryani et al., Persoonia 43: 59.
2019.
Holotypus: InaCC F872 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F872.
Type locality: Indonesia, East Java, Lumajang, Kecamatan
Senduro, Desa Kandang Kepus.
Type substrate: Musa acuminata var. Pisang Mas Kirana.
Descriptions and illustrations: See Maryani et al. (2019b).
FUSARIUM
Diagnostic DNA barcodes: rpb2: LS479850; tef1: LS479441.
lunatum Fusarium (Ellis & Everh.) Arx, Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk., Sect. 2, 51: 101. 1957.
Bisifusarium lunatum (Ellis & Everh.) L. Lombard & Crous,
Stud. Mycol. 80: 225. 2015.
Basionym: Gloeosporium lunatum Ellis & Everh., Proc. Acad.
Nat. Sci. Philadelphia 43: 82. 1891.
Synonyms: Microdochium lunatum (Ellis & Everh.) Arx, Trans.
Brit. Mycol. Soc. 83: 374. 1984.
Fusarium dimerum var. violaceum Wollenw., Fusaria Autogr.
Delin. 3: 854. 1930.
Holotypus: NY00883039.
Type locality: USA, Texas, San Antonio.
Type substrate: Living leaves of Opuntia sp.
Notes: This species requires epitypification. Gerlach & Nirenberg
(1982) designated CBS 632.76 (= NRRL 20690) as neotype of
F. dimerum var. violaceum, which was originally collected in
Germany. However, Schroers et al. (2009) showed that
F. lunatum is paraphyletic and needs further investigation.
Therefore, CBS 632.76 cannot be designated as epitype for
B. lunatum at this time.
lunulosporum Fusarium Gerlach, Phytopathol. Z. 88: 283.
1977.
Holotypus: BBA 62459.
Ex-type culture: ATCC 36747 = BBA 62459 = CBS 636.76 = IMI
322097 = NRRL 13393.
Type locality: South Africa.
Type substrate: Citrus paradisi.
Descriptions and illustrations: See Gerlach (1977b), Gerlach &
Nirenberg (1982) and Nelson et al. (1983).
Diagnostic DNA barcodes: rpb1: KM361637; rpb2: KM361655;
tef1: AF212467.
luteum Fusarium Clem., Bot. Surv. Nebraska 3: 12. 1894.
(See Fusarium lateritium)
Holotypus: NEB00040542.
Type locality: USA, Nebraska, Lincoln.
Type substrate: Decaying wood.
Note: Synonym fide Wollenweber & Reinking (1935).
luteum Fusarium Parav., Ann. Mycol. 16: 302. 1918, nom. illegit.,
Art. 53.1.
(See Fusarium candidum)
Authentic material: In Ann. Mycol. 16, pl. 4., figs 1–22.
Original locality: Switzerland.
Original substrate: Pyrus sp.
Notes: Synonym fide Wollenweber & Reinking (1935).
lutulatum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 209. 1915.
(See Fusarium oxysporum)
Typus: CUP-007458.
Type locality: USA, Iowa.
Type substrate: Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in CUP.
lyarnte Fusarium J.L. Walsh, Sangal., L.W. Burgess, E.C.Y.
Liew & Summerell, sp. nov. MycoBank MB 837697.
Synonym: Fusarium lyarnte J.L. Walsh, Sangal., L.W. Burgess,
E.C.Y. Liew & Summerell, Fungal Diversity 44: 153. 2010, nom.
inval., Art. 40.7.
www.studiesinmycology.org
REDELIMITED
Etymology. ‘Lyarnte’, meaning circle in eastern and central
Arrernte Aboriginal language (Henderson & Dobson 1994), in
reference to the conspicuous globose microconidia.
For diagnosis see Walsh et al., Fungal Diversity 44: 153. 2010.
Holotypus: CBS 125536 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 125536 = NRRL 54252 = RBG 5331.
Type locality: Australia, Northern Territory, Litchfield.
Type substrate: Soil.
Descriptions and illustrations: See Walsh et al. (2010).
Diagnostic DNA barcodes: rpb1: JX171549; rpb2: JX171661;
tef1: EF107118.
Notes: Walsh et al. (2010) failed to indicate the holotype for
F. lyarnte, thereby rendering the species name invalid (Art. 40.7).
Here we validate the name.
lycopersici Fusarium (Sacc.) Mussat, Syll. Fung. 15: 144. 1901,
nom. inval., Art. 36.1(a), (c).
Basionym: Fusarium oxysporum subsp. lycopersici Sacc., Syll.
Fung. 4: 705. 1886.
(See Fusarium oxysporum)
Authentic material: Not located.
Original locality: Italy.
Original substrate: Solanum lycopersicum.
lycopersici Fusarium Bruschi, Atti Reale Accad. Lincei, Rendiconti Cl. Sci. Fis., ser. 5, 21: 298. 1912.
(See Fusarium oxysporum)
Synonym: Fusarium bulbigenum var. lycopersici (Bruschi) Wollenw. & Reinking, Fusarien: nos. 996–997. 1935.
Holotypus: Not located.
Type locality: Italy.
Type substrate: Solanum lycopersicum.
Note: Synonym fide Wollenweber & Reinking (1935).
lycopersici Fusarium (Sacc.) Wollenw., Phytopathology 3: 29.
1913, nom. illegit., Art. 53.1.
Basionym: Fusarium oxysporum subsp. lycopersici Sacc., Syll.
Fung. 4: 705. 1886.
(See Fusarium oxysporum)
Authentic material: Not located.
Original locality: Italy.
Original substrate: Solanum lycopersicum.
macounii Fusarium Dearn., Mycologia 9: 363. 1917.
(See Fusarium expansum)
Holotypus: DAOM 223428b.
Type locality: Canada, Vancouver Island.
Type substrate: Acer sp.
Note: Synonym fide Wollenweber & Reinking (1935).
macroceras Fusarium Wollenw. & Reinking, Phytopathology
15: 166. 1925.
Holotypus: CBS 146.25 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 146.25 = NRRL 13958.
Type locality: Honduras.
Type substrate: Phaseolus vulgaris.
Descriptions and illustrations: See Wollenweber & Reinking
(1925, 1935) and Gerlach & Nirenberg (1982).
Notes: Phylogenetic inference (not shown) revealed that the extype culture housed at CBS clustered within the N. petroliphila
clade, indicating a possible strain transposition or contamination
of the culture in the past. These species are not morphologically
129
CROUS
ET AL.
conspecific based on the original protologue (Wollenweber &
Reinking 1925) of F. macroceras.
macrosporum Fusarium (Sand.-Den. et al.) O'Donnell et al., Index Fungorum 440: 2. 2020.
Neocosmospora macrospora Sand.-Den. et al., Persoonia 40:
21 2017 [2018].
Holotypus: CBS H-23023.
Ex-type culture: CBS 142424 = CPC 28191.
Type locality: Italy, Sicily, Catania, Guardia.
Type substrate: Citrus sinensis.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018a).
Diagnostic DNA barcodes: rpb1: MW218125; rpb2: LT746331;
tef1: LT746218.
macroxysporum Fusarium Lindf., Meddel. Centralanst. Försöksv€as. Jordbruksomr. Avd. Lantbruksbot. 25: 8. 1922.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: Sweden.
Type substrate: Pinus sylvestris.
Note: Synonym fide Wollenweber & Reinking (1935).
maculans Fusarium Berenger, Atti Riunione Sci. Ital. 6: 474.
1845.
Neophloeospora maculans (Berenger) Videira & Crous, Stud.
Mycol. 87: 338. 2017.
Synonyms: Phloeospora maculans (Berenger) Allesch., Rabenh.
Krypt.-Fl., ed. 2, 1: 935. 1900.
Phloeosporella maculans (Berenger) Ho€hn., Mitt. Bot. Inst.
Techn. Hochsch. Wien 4: 77. 1927.
Cercosporella maculans (Berenger) F.A. Wolf, J. Elisha Mitchell
Sci. Soc. 51: 165. 1935.
Septoria mori Lev., Ann. Sci. Nat., Bot., ser. 3, 5: 279. 1846.
Cheilaria mori (Lev.) Desm., Ann. Sci. Nat., Bot., ser. 3, 8: 27.
1847.
Phloeospora mori (Lev.) Sacc., Michelia 1: 175. 1878.
Septogloeum mori (Lev.) Briosi & Cavara, Fung. Paras. Piante
Colt. Util., Fasc. 1: no. 21. 1888.
Cylindrosporium mori (Lev.) Berl., Riv. Patol. Veg. 5: 205. 1896.
Sphaeria mori Nitschke, Fungi Rhen. Exs. no. 1784. 1866, nom.
inval., Art. 38.1(a).
Sphaerella mori Fuckel, Jahrb. Nassauischen Vereins Naturk.
23–24: 106. 1870.
Mycosphaerella mori (Fuckel) F.A. Wolf, J. Elisha Mitchell Sci.
Soc. 51: 165. 1935.
Sphaerella morifolia Pass., Erb. Critt. Ital., Ser. 2, Fasc. 30, no.
1464. 1885.
Mycosphaerella morifolia (Pass.) Cruchet, Bull. Soc. Vaud. Sci.
Nat. 55: 43. 1923.
Cercospora pulvinulata f. angulosa Savul. & Sandu, Herb. Mycol.
Roman. no. 188. 1931.
Holotypus: Not located.
Type locality: Italy.
Type substrate: Leaves of Morus sp.
madaense Fusarium Ezekiel et al., MycoKeys 67: 112. 2020.
Holotypus: CBS H-24346.
Ex-type culture: CBS 146669 = CPC 38344.
Type locality: Nigeria, Nasarawa, Mada Station.
Type substrate: Arachis hypogaea.
130
Descriptions and illustrations: See Ezekiel et al. (2020).
Diagnostic DNA barcodes: rpb1: LR792575; rpb2: LR792589;
tef1: LR792625.
magnoliae-champaca Fusarium R.H. Perera et al., Mycosphere 11: 2140. 2020.
Holotypus: MFLU 18-2736.
Ex-type culture: MFLUCC 18-0580.
Type locality: Thailand, Chiang Rai, Mae Fah Luang University
garden.
Type substrate: Dried fruits of Magnolia champaca.
Descriptions and illustrations: See Perera et al. (2020).
Diagnostic DNA barcode: rpb2: MT212198.
magnusianum Fusarium Allesch., Fungi Bav. no. 400. 1895.
(See Fusarium aquaeductuum)
Holotypus: In M.
Type locality: Germany, München.
Type substrate: Salix incana.
Note: Synonym fide Wollenweber & Reinking (1935).
mahasenii Fusarium Samuels et al., Mycologia 103: 1325. 2011.
Neocosmospora mahasenii Samuels et al., Mycologia 103:
1325. 2011.
Holotypus: BPI 881228.
Ex-type culture: CBS 119594 = FRC S-1845 = G.J.S. 02-105.
Type locality: Sri Lanka, North Central Province, Giritale. Giritale
Forest Training Center.
Type substrate: Small branch of live tree.
Descriptions and illustrations: See Nalim et al. (2011).
Diagnostic DNA barcodes: rpb1: MW834231; rpb2: LT960563;
tef1: DQ247513.
mali Fusarium Allesch., Ber. Bot. Vereines Landshut 12: 130.
1892.
(See Fusarium candidum)
Holotypus: In M.
Type locality: Germany, München.
Type substrate: Malus pumila.
Note: Synonym fide Wollenweber & Reinking (1935).
malli Fusarium Taubenh., Bull. Texas Agric. Exp. Sta. 273: 25.
1921.
(See Fusarium solani)
Holotypus: ?CUP-011254.
Type locality: USA, Texas, Brazos, College Station.
Type substrate: Allium cepa.
Note: Typification pending study of material lodged in CUP.
malvacearum Fusarium Taubenh., Bull. Texas Agric. Exp. Sta.
260: 27. 1920.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000714): USA, Texas,
Abelmoschus esculentus, 1920, J.J. Taubenhhaus, in Taubenhaus, Bull. Texas Agric. Exp. Sta. 260: 30, fig. 8g.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration
was designated as lectotype.
mangiferae Fusarium Britz et al., Mycologia 94: 725. 2002.
Holotypus: PREM 57299.
Ex-type culture: CBS 120994 = KSU 11781 = MRC
7559 = MUCL 54671 = NRRL 53980.
FUSARIUM
Type locality: Israel, Bet Dagan, Volcani Center.
Type substrate: Mangifera indica.
Descriptions and illustrations: See Britz et al. (2002).
Diagnostic DNA barcodes: rpb1: MW402530; rpb2: LT575059;
tef1: LT574978.
marasasianum Fusarium Herron et al., Stud. Mycol. 80: 146.
2015.
Holotypus: PREM 60899.
Ex-type culture: CBS 137238 = CMW 25261.
Type locality: Colombia, Vivero Pe~nas Negra, Valle del Cauca.
Type substrate: Pinus patula.
Descriptions and illustrations: See Herron et al. (2015).
Notes: Comparisons of recently generated sequences for the
living ex-type (CBS 137238 = CMW 25261) of F. marasasianum
indicate a strain transposition or contamination by another
Fusarium species. Therefore, this species needs to be recollected from the type locality and substrate or sequences need to
be generated from the holotype specimen.
marginatum Fusarium Berk. & M.A. Curtis, Grevillea 3: 97. 1875.
Holotypus: ?K(M).
Type locality: USA, Alabama, Beaumont.
Type substrate: Smilax sp.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
martiellae-discolorioides Fusarium Batikyan, Biol. Zhurn. Armenii
22: 87. 1969, nom. inval., Art. 39.1.
Authentic material: Not located.
Original locality: Armenia.
Original substrate: Soil of wheatfield.
Notes: Published without Latin diagnosis fide Gerlach &
Nirenberg (1982). Also described in Biol. Zhurn. Armenii 26(2):
73. 1973, but also not in Latin.
martii Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol. Anst.
Land- Forstw. 8: 83. 1913.
Neocosmospora martii (Appel & Wollenw.) Sand.-Den. &
Crous, Persoonia 43: 137. 2019.
Synonyms: Fusarium solani var. martii (Appel & Wollenw.)
Wollenw., Fusaria Autogr. Delin. 3: 1034. 1930.
Neocosmospora croci Guarnaccia et al., Persoonia 40: 17. 2017
[2018].
Lectotypus: BPI 452385, selected in Sandoval-Denis et al.
(2019).
Epitypus: CBS H-23986, designated in Sandoval-Denis et al.
(2019).
Ex-epitype culture: CBS 115659 = FRC S-0679 = MRC 2198.
Lecto- and epitype locality: Germany, Berlin.
Lecto- and epitype substrate: Solanum tuberosum.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834232; rpb2: JX435256;
tef1: JX435156.
massalimae Fusarium A.D. Cavalcanti et al., Mycol. Progr. 19:
1137. 2020.
Holotypus: URM 94324.
Ex-type culture: URM 8239.
Type locality: Brazil, Alagoas, Quebrangulo, Pedra Talhada
Biological Reserve.
Type substrate: Handroanthus chrysotrichus.
Descriptions and illustrations: See Cavalcanti et al. (2020).
Diagnostic DNA barcodes: rpb2: MN939767; tef1: MN939763.
matuoi Fusarium Hosoya & Tubaki, Mycoscience 45: 264. 2004.
www.studiesinmycology.org
REDELIMITED
Fusicolla matuoi (Hosoya & Tubaki) Gr€afenhan & Seifert, Stud.
Mycol. 68: 101. 2011.
Synonyms: Fusarium splendens Matuo & Takah. Kobay., Trans.
Mycol. Soc. Japan 2(4): 13. 1960, nom. inval., Art. 39.1.
Cosmospora matuoi Hosoya & Tubaki, Mycoscience 45: 262. 2004.
Holotypus: TNS F-11127.
Ex-type culture: MAFF 410976.
Type locality: Japan, Honshu.
Type substrate: Twigs of Albizia julibrissin.
Descriptions and illustrations: See Hosoya & Tubaki (2004).
mauroi Fusarium Av.-Sacca, Revista Agric. (Piracicaba) 8: 93.
1933.
Macronectria jungneri (Henn.) Salgado & P. Chaverri, Fungal
Diversity 80: 448. 2016.
Basionym: Nectria jungneri Henn., Bot. Jahrb. Syst. 22: 75. 1895.
Synonyms: Cucurbitaria jungneri (Henn.) Kuntze, Revis. Gen. Pl.
3: 461. 1898.
Neonectria jungneri (Henn.) Samuels & Brayford (as ‘Nenectria’),
Mycologia 96: 580. 2004.
Thelonectria jungneri (Henn.) P. Chaverri & Salgado, Stud.
Mycol. 68: 76. 2011.
Nectria eustoma Penz. & Sacc., Malpighia 11: 509. 1898 [1897].
Nectria leucocoma Starb€ack, Bih. Kongl. Svenska Vetensk.Akad. Handl. 25: 28. 1899.
Nectria cinereopapillata Henn. & E. Nyman, inWarburg, Monsunia 1: 161. 1900 [1899].
Nectria striatospora Zimm., Centralbl. Bakteriol. II, 7: 105. 1901.
Nectria theobromae Massee, Bull. Misc. Inform. Kew 1908: 218.
1908.
Cylindrocarpon victoriae Wollenw., Z. Parasitenk. (Berlin) 1: 161.
1928.
Nectria azureo-ostiolata Doi, Mem. Nat. Sci. Mus. Tokyo 10: 23.
1977.
Holotypus: Not located.
Type locality: Brazil.
Type substrate: Caconema radicicola.
Note: Synonyms fide Wollenweber & Reinking (1935) and
Salgado-Salazar et al. (2016).
maydiperdum Fusarium Bubak, Centralbl. Bakteriol. 2. Abth. 31:
497. 1911.
(See Fusarium poae)
Holotypus: BPI 452399.
Type locality: Czech Republic, Tabor.
Type substrate: Seeds of Zea mays.
Note: Synonym fide Wollenweber & Reinking (1935).
maydis Fusarium Kalchbr., Math. Term. Közlem. 3: 285. 1865.
(See Fusarium sambucinum)
Holotypus: BRACR33140.
Type locality: Hungary.
Type substrate: Zea mays.
Note: Synonym fide Wollenweber & Reinking (1935).
melanochlorum Fusarium (Casp.) Sacc., Syll. Fung. 4: 725. 1886.
Basionym: Fusisporium melanochlorum Casp., Ber.
Bekanntm. Verh. Königl. Preuss. Akad. Wiss. Berlin 1855:
309, 314. 1855.
Cosmospora flavoviridis (Fuckel) Rossman & Samuels, Stud.
Mycol. 42: 121. 1999.
Basionym: Sphaerostilbe flavoviridis Fuckel, Jahrb. Nassauischen Vereins Naturk. 25–26: 310. 1871.
131
CROUS
ET AL.
Synonyms: Nectria flavoviridis (Fuckel) Wollenw. Angew. Bot. 8:
186. 1926.
Fusarium celtidis Pass., Atti Reale Accad. Lincei, Rendiconti Cl.
Sci. Fis., 4 ser. 7: 51. 1891, nom. illegit., Art. 53.1.
Fusarium sphaeriiforme Sacc. (as ‘sphaeriaeforme’), Syll. Fung.
10: 723. 1892.
Fusarium glandicola Allesch., Ber. Bot. Vereines Landshut12:
130. 1892, nom. illegit., Art. 53.1, non Cooke & W.R. Gerard,
1878.
Fusarium allescheri Sacc. & P. Syd., Syll. Fung. 14: 1128. 1899.
Holotypus: Not located.
Type locality: Germany, Berlin.
Type substrate: Rotten aquatic plants.
meliolicola Fusarium F. Stevens (as ‘meliolicolum’), Bot. Gaz. 65:
245. 1918.
(See Fusarium larvarum)
Holotypus: ILL00011251.
Type locality: Puerto Rico, Mayagüez.
Type substrate: Parasitic on Meliola paulliniae on Casearia
sylvestris.
Note: Synonym fide Wollenweber & Reinking (1935).
meridionale Fusarium T. Aoki et al., Fungal Genet. Biol. 41:
618. 2004.
Holotypus: BPI 843474.
Ex-type culture: CBS 110247 = FRC R-5329 = NRRL 28436.
Type locality: New Caledonia.
Type substrate: Citrus sinensis.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: KM361642; rpb2: KM361660;
tef1: AF212435.
merismoides Fusarium Corda, Icon. Fung. 2: 4. 1838.
Fusicolla merismoides (Corda) Gr€afenhan et al., Stud. Mycol.
68: 101. 2011.
Synonyms: Fusisporium georginae Klotzsch, Herb. Viv. Mycol.,
Cent. 2: 186. 1832, nom. nud., Art. 38.1 (a).
Fusarium rhizophilum Corda, Icon. Fung. 2: 3. 1838.
Pionnotes rhizophila (Corda) Sacc., Syll. Fung. 4: 727. 1886.
?Fusisporium arachnoideum Corda, Icon. Fung. 1: 11. 1837.
?Fusarium arachnoideum (Corda) Sacc., Syll. Fung. 4: 721.
1886.
?Fusarium biasolettianum Corda, Icon. Fung. 2: 3. 1838.
?Fusisporium biasolettianum (Corda) Sacc., Mycoth. Ven. no.
1040. 1877.
?Pionnotes biasolettiana (Corda) Sacc., Syll. Fung. 4: 725. 1886.
Fusisporium udum Berk., Ann. Mag. Nat. Hist. 6: 438. 1841.
Pionnotes uda (Berk.) Sacc., Syll. Fung. 4: 726. 1886.
Fusarium udum (Berk.) Wollenw., Phytopathology 3: 38. 1913,
nom. illegit., Art. 53.1.
Fusidium udum Berk., in Trotter, Syll. Fung. 25: 979. 1931, nom.
inval., Art. 36.1.
Fusisporium foeni Berk. & Broome, Ann. Mag. Nat. Hist., ser. 2,
7: 179. 1851.
Fusarium foeni (Berk. & Broome) Sacc., Syll. Fung. 4: 699. 1886.
Fusisporium roseolum H.O. Stephens ex Berk. & Broome, Ann.
Mag. Nat. Hist., ser. 2, 7: 178. 1851.
Fusarium roseolum (H.O. Stephens ex Berk. & Broome) Sacc.,
Syll. Fung. 4: 710. 1886.
Fusisporium rimosum Peck, Rep. (Annual) New York State Mus.
Nat. Hist. 30: 58. 1878.
Fusarium rimosum (Peck) Sacc., Syll. Fung. 4: 713. 1886.
132
Fusarium roesleri Thüm., Pilze Weinst.: 51. 1878.
Fusarium arvense Speg., Anales Soc. Ci. Argent. 10: 60. 1880.
Fusarium gallinaceum Cooke & Harkn., Grevillea 9: 8. 1880.
Fusarium nicotianae Oudem., Ned. Kruidk. Arch., ser. 3, 2: 777.
1902.
Fusarium udum var. pusillum Wollenw., Phytopathology 1: 206.
1913, nom. nud.
Fusarium udum var. solani Sherb., Mem. Cornell Univ. Agric.
Exp. Sta. 6: 131. 1915.
Fusarium merismoides f. nicotianae (Oudem.) Subram.,
Hyphomycetes: 676. 1971.
Fusarium oxysporum f. sp. nicotianae (Oudem.) Subram.,
Hyphomycetes: 676. 1971.
Fusarium pelargonii P. Crouan & H. Crouan, Fl. Finistere: 14.
1867.
Fusarium albiziae Woron., Vestn. Tiflissk. Bot Sada 48: 34. 1920.
Fusarium merismoides var. majus Wollenw., Fusaria Autogr.
Delin. 3: 857a. 1930.
Fusarium merismoides var. chlamydosporale Wollenw., Z. Parasitenk. (Berlin) 3: 308. 1931.
Fusarium merismoides var. artocarpi X.H. Fu & Q.T. Chen, Acta
Mycol. Sin. 8: 42. 1989.
Fusarium merismoides var. persicicola X.H. Fu & Q.T. Chen,
Acta Mycol. Sin. 8: 44. 1989.
Typus: PRM 155493.
Type locality: Czech Republic, Prague.
Type substrate: Wet shards of a plant pot.
Note: Lectotypification pending study of material lodged in PRM.
mesentericum Fusarium Cooke & Harkn., Grevillea 9: 128. 1881.
Holotypus: ?K(M).
Type locality: USA, California, San Francisco Masonic Cemetery.
Type substrate: Eucalyptus sp.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
mesoamericanum Fusarium T. Aoki et al., Fungal Genet. Biol.
41: 619. 2004.
Holotypus: BPI 843476.
Ex-type culture: CBS 415.86 = FRC R-8506 = IMI
309346 = NRRL 25797.
Type locality: Honduras.
Type substrate: Musa sp.
Descriptions and illustrations: See O'Donnell et al. (2004).
Diagnostic DNA barcodes: rpb1: KM361639; rpb2: KM361657;
tef1: AF212441.
metachroum Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 141. 1910 [1913].
(See Fusarium avenaceum)
Holotypus: BPI 452408.
Type locality: Poland, Poznan, Slivno Manor.
Type substrate: Triticum aestivum.
Note: Synonym fide Wollenweber & Reinking (1935).
metavorans Fusarium Al-Hatmi et al., Medical Mycol. 56: S147.
2018.
Neocosmospora metavorans (Al-Hatmi et al.) Sand.-Den. &
Crous, Persoonia 41: 121. 2018.
Holotypus: CBS 135789 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 135789.
Type locality: Greece, Athens.
Type substrate: Pleural effusion of Homo sapiens.
FUSARIUM
Descriptions and illustrations: See Al-Hatmi et al. (2018) and
Sandoval-Denis & Crous (2018).
Diagnostic DNA barcodes: rpb1: MW218127; rpb2: LR583849;
tef1: LR583627.
mexicanum Fusarium T. Aoki et al., Phytopathology 100: 1180.
2010.
Holotypus: BPI 879150.
Ex-type culture: NRRL 53147.
Type locality: Mexico, Nueva Italia, Michoacan.
Type substrate: Mangifera indica.
Descriptions and illustrations: See Otero-Colina et al. (2010).
Diagnostic DNA barcodes: rpb1: MG838088; rpb2: MN724973;
tef1: MG838032.
microcera Fusarium Bilaĭ, Fusarii (Biologija i sistematika): 292.
1955, nom. inval., Art. 39.1.
(See Fusarium coccidicola)
Note: This species was invalidly published without Latin
diagnosis.
microconidium Fusarium L. Lombard & Crous, Fungal Syst.
Evol. 4: 192. 2019.
Holotypus: CBS H-24017.
Ex-type culture: CBS 119843 = MRC 8391 = KSU 11396.
Type locality: Unknown.
Type substrate: Unknown.
Descriptions and illustrations: See Lombard et al. (2019a).
Diagnostic DNA barcodes: rpb1: MN120721; tef1: MN120759.
microphlyctis Fusarium Mont., Ann. Sci. Nat., Bot., ser. 3, 12:
297. 1849.
Holotypus: ?PC.
Type locality: France.
Type substrate: Fruit of Olea sp.
Note: Gloeosporium fide Wollenweber & Reinking (1935).
micropus Fusarium Sacc., Philipp. J. Sci. 18: 605. 1921.
Infrafungus micropus (Sacc.) Cif., Mycopathol. Mycol. Appl. 6:
26. 1951.
Holotypus: In PAD.
Type locality: China, Guangdong Province.
Type substrate: Parasitic on Cladosporium herbarum on leaf of
Morus alba.
microspermum Fusarium Berk. & M.A. Curtis, Grevillea 3: 98.
1875.
Holotypus: ?K(M).
Type locality: USA, South Carolina, Santee River.
Type substrate: Ficus sp.
Note: Hymenula fide Wollenweber & Reinking (1935).
microsporum Fusarium Schltdl., Fl. Berol. 2: 139. 1824.
(See Fusarium lateritium)
Holotypus: HAL 1615 F.
Type locality: Germany, Berlin.
Type substrate: Robinia pseudoacaciae.
Note: Synonym fide Wollenweber & Reinking (1935).
mikaniae Fusarium Berk. & M.A. Curtis, Grevillea 3: 98. 1875.
Holotypus: ?K(M).
Type locality: USA, South Carolina, Santee River.
Type substrate: Stems and leaves of Mikania scandens.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
www.studiesinmycology.org
REDELIMITED
mindoanum Fusarium Petr., Sydowia 4: 576. 1950.
Holotypus: In W as no. 03550 (Petrak, Pilzherbarium no. 32229).
Type locality: Ecuador, Pichincha, Mindo.
Type substrate: Dryopteris diplazioides.
Notes: No living material available to confirm taxonomic status.
Requires recollection from type locality and substrate.
miniatulum Fusarium Sacc., Syll. Fung. 10: 727. 1892.
Replaced synonym: Fusarium miniatum Prill. & Delacr., Bull.
Soc. Mycol. France 7: 117. 1891, nom. illegit., Art. 53.1
(See Fusarium nivale)
Holotypus: Not located.
Type locality: France, Paris.
Type substrate: Secale cereale.
Note: Synonym fide Wollenweber & Reinking (1935).
miniatum Fusarium Sacc., Michelia 1: 83. 1877.
Synonym: Fusarium detonianum Sacc., Syll. Fung. 4: 708. 1886,
nom. illegit., Art. 52.1.
Holotypus: In PAD.
Type locality: Italy.
Type substrate: Sporangium of Cyathus vernicosa.
Note: Status unclear. Requires further investigation.
miniatum Fusarium (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
722. 1886, nom. illegit., Art. 53.1.
Basionym: Fusisporium miniatum Berk. & M.A. Curtis, Grevillea
3: 147. 1875.
(See Fusarium lateritium)
Holotypus: ?K(M).
Type locality: USA, North Carolina.
Type substrate: Cornus florida.
Note: Synonym fide Wollenweber & Reinking (1935).
miniatum Fusarium Prill. & Delacr., Bull. Soc. Mycol. France 7:
117. 1891, nom. illegit., Art. 53.1.
Replacing synonym: Fusarium miniatulum Sacc., Syll. Fung. 10:
727. 1892
(See Fusarium nivale)
Authentic material: Not located.
Original locality: France, Paris.
Original substrate: Secale cereale.
Note: Synonym fide Wollenweber & Reinking (1935).
minimum Fusarium Fuckel, Fungi Rhen. Exs., Fasc. 3, no. 213.
1863.
(See Fusarium nivale)
Syntypes: In BPI, MICH, MU & S (Fungi Rhen. Exs., Fasc. 3, no.
213).
Type locality: Germany, Oestrich, Nassau region.
Type substrate: Dry leaves of Poaceae (mainly Zea mays)
Note: Synonym fide Wollenweber & Reinking (1935).
minutissimum Fusarium (Desm.) Sacc., Syll. Fung. 4: 703. 1886.
Passalora minutissima (Desm.) U. Braun & Crous, CBS
Biodiversity Ser. 1: 276. 2003.
Basionym: Selenosporium minutissimum Desm., Pl. Crypt.
France, ed. 3, Fasc. 10: no. 456. 1857.
Phaeoramularia minutissima (Desm.) U. Braun, Nova Hedwigia
55: 214. 1992.
Ramularia geranii-sanguinei C. Massal., Atti Ist. Veneto Sci. Lett.
Arti 59: 688. 1900.
Cercospora geranii-sanguinei Henn., Nytt Mag. Naturvidensk.
42: 33. 1904.
133
CROUS
ET AL.
Lectotypus: Desm., Pl. Crypt. France, Fasc. X, no. 456 in PC fide
Braun (1998).
Lectotype locality: France, Louvigny, Caen.
Lectotype substrate: Geranium molle.
minutulum Fusarium Corda, Icon. Fung. 2: 4. 1838.
?Clonostachys solani (Harting) Schroers & W. Gams, Stud.
Mycol. 46: 111. 2001.
Basionym: Spicaria solani Harting, Nieuwe Verh. Eerste Kl. Kon.
Ned. Inst. Wetensch. Amsterdam, ser. 2, 12: 226. 1846.
Synonyms: ?Gliocladium solani (Harting) Petch, Trans. Brit.
Mycol. Soc. 27: 149. 1945.
?Hypomyces solani Reinke & Berthold, Untersuch. Bot. Lab.
Univ. Göttingen 1: 27. 1879.
?Hypolyssus solani (Reinke & Berthold) Kuntze, Revis. Gen. Pl.
3: 488. 1898.
?Hyphonectria solani (Reinke & Berthold) Petch, Bot. J. (London)
74: 220. 1937 [1936].
?Nectriopsis solani (Reinke & Berthold) C. Booth, Mycol. Pap.
74: 8. 1960.
?Bionectria solani (Reinke & Berthold) Schroers, Stud. Mycol.
46: 111. 2001.
?Gliocladium nigrovirens J.F.H. Beyma, Verh. Kon. Akad.
Wetensch., Afd. Natuurk., Sect. 2, 29: 30. 1931.
?Clonostachys solani f. nigrovirens (J.F.H. Beyma) Schroers,
Stud. Mycol. 46: 115. 2001.
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, Prague.
Type substrate: Wood splinters of Corylus sp.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in PRM.
miscanthi Fusarium W. Gams et al., Mycologia 91: 264. 1999.
Holotypus: CBS H-6063.
Ex-type culture: CBS 577.97 = NRRL 26231.
Type locality: Denmark, Zealand, Højbakkegård Experimental
field.
Type substrate: Miscanthus sinensis.
Descriptions and illustrations: See Gams et al. (1999).
Diagnostic DNA barcodes: rpb1: JX171521; rpb2: JX171634;
tef1: MN193878.
mollerianum Fusarium Thüm., Inst. Coimbra 28: 263. 1881.
(See Fusarium graminearum)
Holotypus: ?S-F45644.
Type locality: Portugal, Coimbra.
Type substrate: Melia azedarach.
Note: Synonym fide Wollenweber & Reinking (1935).
moniliforme Fusarium J. Sheld., Annual Rep. Nebraska Agric.
Exp. Sta. 17: 23. 1904.
(See Fusarium verticillioides)
Syntypes: BPI 452450 & BPI 452452.
Type locality: USA, Nebraska.
Type substrate: Zea mays.
Note: Typification pending further study of the syntypes.
monophialidicum Fusarium J.W. Xia et al., Persoonia 43: 211.
2019.
Holotypus: CBS H-24063.
Ex-type culture: NRRL 54973 = UTHSC 06-1473.
Type locality: USA, Ohio.
Type substrate: Eye of Rhinocerotidae (rhinoceros).
Descriptions and illustrations: See Xia et al. (2019).
134
Diagnostic DNA barcodes: rpb1: KC808299; rpb2: KC808362;
tef1: MN170483.
mori Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 2. 2020.
Neocosmospora mori Sand.-Den. & Crous, Persoonia 43: 143.
2019.
Holotypus: CBS H-23987.
Ex-type culture: ATCC 44934 = CBS 145467 = MAFF
238539 = NRRL 22230.
Type locality: Japan, Miyazaki.
Type substrate: Twigs of Morus alba.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834235; rpb2: EU329499;
tef1: AF178358.
moronei Fusarium Curzi, Revista Biol. (Lisbon) 10: 141. 1928.
(See Fusarium acuminatum)
Holotypus: ?PAV.
Type locality: Italy.
Type substrate: Vesicle on skin of Canis lupus familiaris (dog).
Note: Synonym fide Wollenweber & Reinking (1935).
moschatum Fusarium (Kitasato) Sacc., Syll. Fung. 10: 729.
1892.
Basionym: Fusisporium moschatum Kitasato, Centralbl. Bakteriol. Parasitenk., 1. Abth. 5: 365. 1889.
(See Fusarium aquaeductuum)
Holotypus: Not located.
Type locality: Germany.
Type substrate: Metallic medical equipment.
Note: Synonym fide Wollenweber & Reinking (1935).
mucidum Fusarium J.W. Xia et al., Persoonia 43: 211. 2019.
Holotypus: CBS H-24064.
Ex-type culture: CBS 102395.
Type locality: El Salvador, Cooperacion Coralama.
Type substrate: Mouldy nut of Anacardium occidentale.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170418; tef1: MN170485.
muentzii Fusarium Delacr. (as ‘müntzii’), Bull. Soc. Mycol. France
8: 192. 1892.
(See Fusarium tricinctum)
Lectotypus (hic designatus, MBT 10000715): France, Paris, on
animal waste, May 1891, G. Delacroix, Bull. Soc. Mycol. France
8, pl. XVII, fig. V.
multiceps Fusarium J.W. Xia et al., Persoonia 43: 212. 2019.
Holotypus: CBS H-24065.
Ex-type culture: CBS 130386 = NRRL 43639 = UTHSC 04-135.
Type locality: USA, Florida.
Type substrate: Trichechus sp.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb1: HM347190; rpb2: GQ505844;
tef1: GQ505666.
mundagurra Fusarium M.H. Laurence et al., Fungal Diversity
77: 359. 2015 [2016].
Holotypus: RBG5717.
Ex-type culture: NRRL 66235 = RBG5717.
Type locality: Australia, Queensland, Carnarvon Gorge National
Park.
Type substrate: Soil.
Descriptions and illustrations: See Laurence et al. (2016).
FUSARIUM
Diagnostic DNA barcodes: rpb1: KP083272; rpb2: KP083276;
tef1: KP083256.
musae Fusarium Van Hove et al., Mycologia 103: 579. 2011.
Synonym: Gibberella musae Van Hove et al., Mycologia 103:
577. 2011.
Holotypus: MUCL 52574.
Ex-type culture: CBS 624.87 = MUCL 52574 = NRRL 25059.
Type locality: Honduras.
Type substrate: Musa sp.
Descriptions and illustrations: See Van Hove et al. (2011).
Diagnostic DNA barcodes: rpb1: MW402689; rpb2: FN552108;
tef1: FN552086.
musarum Fusarium Logrieco & Marasas, Mycologia 90: 510.
1998.
Holotypus: BPI 802928.
Ex-type culture: FRC R-9400 = MRC 6240 = NRRL 28507.
Type locality: Panama.
Type substrate: Musa sapientum.
Descriptions and illustrations: See Marasas et al. (1998).
Diagnostic DNA barcodes: rpb1: MW233265; rpb2: MW928829;
tef1: MW233094.
mycophilum Fusarium (P. Karst.) Sacc., Syll. Fung. 10: 730. 1892.
Basionym: Leptosporium mycophilum P. Karst., Meddel. Soc.
Fauna Fl. Fenn. 16: 24. 1888.
Holotypus: ?H.
Type locality: Finland, Merimasku.
Type substrate: Myxogastria.
Note: Hymenula fide Wollenweber & Reinking (1935).
mucophytum Fusarium (W.G. Sm.) Massee, Brit. Fung.-Fl. 3:
483. 1893.
Basionym: Fusisporium mucophytum W.G. Sm., Gard. Chron.
n.s., 22: 245. 1884.
(See Fusarium scirpi)
Holotypus: ?K(M).
Type locality: UK.
Type substrate: Agaricus arvensis.
Note: Synonym fide Wollenweber & Reinking (1935).
myosotidis Fusarium Cooke, Grevillea 16: 49. 1887.
(See Fusarium oxysporum)
Holotypus: In K(M).
Type locality: UK, Forden.
Type substrate: Myosotis sp.
Note: Synonym fide Wollenweber & Reinking (1935).
nanum Fusarium M.M. Wang et al., Persoonia 43: 85. 2019.
Holotypus: HAMS 248043.
Ex-type culture: CGMCC 3.19498 = LC12168.
Type locality: China, Guangxi Province, Guilin.
Type substrate: Leaves of Musa nana.
Descriptions and illustrations: See Wang et al. (2019).
Diagnostic DNA barcodes: rpb1: MK289871; rpb2: MK289755;
tef1: MK289602.
napiforme Fusarium Marasas et al., Mycologia 79: 910. 1988
[1987].
Holotypus: DAOM 196924.
Ex-type culture: BBA 69861 = CBS 748.97 = DAOM
196924 = DAOM 225147 = FRC M-3563 = IMI 375353 = MRC
4144 = NRRL 13604.
Type locality: Namibia, Ovambo.
www.studiesinmycology.org
REDELIMITED
Type substrate: Pennisetum typhoides.
Descriptions and illustrations: See Marasas et al. (1987).
Diagnostic DNA barcodes: rpb1: HM347136; rpb2: EF470117;
tef1: AF160266.
nectriae-palmicolae Fusarium Henn., Bot. Jahrb. Syst. 23: 290.
1896.
(See Fusarium equiseti)
Holotypus: In B fide Hein (1988).
Type locality: Samoa, Upolu.
Type substrate: Leaves of Areca sp.
Note: Synonym fide Wollenweber & Reinking (1935).
nectriae-turraeae Fusarium Henn., Bot. Jahrb. Syst. 22: 82. 1895.
(See Fusarium coccophilum)
Holotypus: In B fide Hein (1988).
Type locality: Tanzania, Marangu.
Type substrate: Turraea volkensii.
Note: Synonym fide Wollenweber & Reinking (1935).
nectricreans Fusarium Kirschst., Ann. Mycol. 37: 138. 1939.
Holotypus: B 70 0100202.
Type locality: Germany, Berlin.
Type substrate: Rotting stem of garden plant.
Note: No living material was available for confirmation of taxonomic status.
nectrioides Fusarium (Wollenw.) Schroers et al., Mycologia 101:
59. 2009.
Bisifusarium nectrioides (Wollenw.) L. Lombard & Crous, Stud.
Mycol. 80: 225. 2015.
Basionym: Fusarium dimerum var. nectrioides Wollenw., Fusaria
Autogr. Delin. 3: 855. 1930.
Lectotypus: No. 855 in Wollenweber, Fusaria Autogr. Delin.
(1930), designated in Schroers et al. (2009).
Ex-type culture: CBS 176.31 = NRRL 20689.
Lectotype and ex-type locality: Honduras.
Lectotype and ex-type substrate: Soil.
Descriptions and illustrations: See Schroers et al. (2009).
Diagnostic DNA barcodes: rpb1: JX171477; rpb2: JX171591;
tef1: EU926312.
neglectum Fusarium Jacz., Bull. Trimestriel Soc. Mycol. France
28: 348. 1912.
(See Fusarium culmorum)
Holotypus: Not located.
Type locality: Ukraine, Poltava.
Type substrate: Zea mays.
Note: Synonym fide Wollenweber & Reinking (1935).
negundinis Fusarium Sherb., in Hubert, J. Agric. Res. 26: 451.
1923.
(See Fusarium reticulatum)
Holotypus: Not located.
Type locality: USA, Wisconsin, Madison.
Type substrate: Acer negundo.
Note: Synonym fide Wollenweber & Reinking (1935).
nelsonii Fusarium Marasas & Logrieco, Mycologia 90: 508.
1998.
Holotypus: BPI 802927.
Ex-type culture: CBS 119876 = FRC R-8670 = MRC
4570 = NRRL 28505 = NRRL 53945.
Type locality: South Africa, Western Cape Province, Malmesbury.
135
CROUS
ET AL.
Type substrate: Plant debris in Triticum soil.
Descriptions and illustrations: See Marasas et al. (1998).
Diagnostic DNA barcodes: rpb1: MN120722; rpb2: GQ505468;
tef1: GQ505404.
nematophilum Fusarium Nirenberg & Hagedorn, Nachrichtenbl.
Deutsch. Pflanzenschutzdienstes 60: 214. 2008.
Luteonectria nematophila (Nirenberg & Hagedorn) Sand.-Den.
& L. Lombard, Stud. Mycol. 98 (no. 100116): 60. 2021.
Holotypus: BBA 72279 in B.
Ex-type culture: BBA 72279 = NRRL 54600.
Type locality: Germany, Berlin.
Type substrate: Isolated from soil with roots of Hedera helix.
Descriptions and illustrations: See Nirenberg & Hagedorn (2008).
Diagnostic DNA barcodes: rpb1: JX171552; rpb2: JX171664;
tef1: JABFFA010003988.
neoceras Fusarium Wollenw. & Reinking, Phytopathology 15:
164. 1925.
(See Fusarium sacchari)
Holotypus: CBS 147.25 (preserved as metabolically inactive
culture).
Ex-type culture: BBA 69863 = CBS 147.25 = DAOM
225410 = IMI 375345= NRRL 20471.
Type locality: Honduras.
Type substrate: Rotting Musa sapientum.
Descriptions and illustrations: See Gerlach & Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: MT010941; rpb2: MT010962;
tef1: MT010988.
neocosmosporiellum Fusarium O'Donnell & Geiser, Phytopathology 103: 405. 2013.
Neocosmospora vasinfecta E.F. Sm., Bull. Div. Veg. Physiol.
Pathol. U.S.D.A. 17: 45. 1899.
Synonyms: Nectriella tracheiphila E.F. Sm., Proc. Amer. Assoc.
Advancem. Sci. 44: 190. 1896, nom. inval. fide Cannon &
Hawksworth 1984.
Neocosmospora vasinfecta var. nivea E.F. Sm., Bull. Div. Veg.
Physiol. Pathol. U.S.D.A. 17: 45. 1899.
Neocosmospora vasinfecta var. tracheiphila E.F. Sm., Bull. Div.
Veg. Physiol. Pathol. U.S.D.A. 17: 45. 1899.
Fusarium tracheiphilum (E.F. Sm.) Wollenw., Phytopathology 3:
29. 1913.
Fusarium vasinfectum var. pisi C.J.J. Hall, Ber. Deutsch. Bot.
Ges. 21: 4. 1903.
Neocosmospora vasinfecta var. pisi (C.J.J. Hall) Sacc., Syll.
Fung. 20: 192. 1911.
Neocosmospora africana Arx, Antonie van Leeuwenhoek 21:
161. 1955.
Neocosmospora vasinfecta var. africana (Arx) P.F. Cannon & D.
Hawksw., Trans. Brit. Mycol. Soc. 82: 676. 1984.
?Pseudonectria ornata Bat. & Maia, Anais Soc. Biol. Pernambuco 13: 74. 1955 (fide Cannon & Hawksworth 1984).
Neocosmospora vasinfecta var. major P. Rama Rao, Mycopathol. Mycol. Appl. 21: 218. 1963.
Neocosmospora ornamentata M.A.F. Barbosa, Garcia de Orta
13: 17. 1965.
Fusarium ornamentatum (M.A.F. Barbosa) O'Donnell et al., Index
Fungorum 440: 3. 2020.
Neocosmospora vasinfecta f. conidiifera Kamyschko, Novosti
Sist. Nizsh. Rast. 1965: 115. 1965.
Neocosmospora boninensis Udagawa et al., Sydowia 41: 350.
1989.
136
Lectotypus: Pl. V, figs 1–2 (Smith, Bull. Div. Veg. Physiol. Pathol.
U.S.D.A. 17, 1899), designated in Sandoval-Denis et al. (2019).
Lectotype locality: USA.
Lectotype substrate: Gossypium sp.
Epitypus: BPI 910920, designated in Aoki et al. (2020).
Ex-epitype culture: ATCC 62199 = NRRL 22166.
Epitype locality: USA, Illinois, southern area.
Epitype substrate: A cyst of Heterodera glycines in a soil sample
from soybean field.
Diagnostic DNA barcodes: rpb1: SSHR01002742; rpb2:
EU329497; tef1: AF178350.
neoscirpi Fusarium L. Lombard et al., Persoonia 43: 213. 2019.
Holotypus: CBS H-24066.
Ex-type culture: CBS 610.95 = NRRL 26861 = NRRL 26922.
Type locality: France.
Type substrate: Soil.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: GQ505779; tef1: GQ505601.
neosemitectum Fusarium L. Lombard et al., Persoonia 43:
214. 2019.
Holotypus: CBS H-24067.
Ex-type culture: CBS 189.60.
Type locality: Democratic Republic of the Congo.
Type substrate: Musa sapientum.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170422; tef1: MN170489.
nepalense Fusarium T. Aoki et al., Fungal Genet. Biol. 48: 1105.
2011.
Holotypus: BPI 881006.
Ex-type culture: CBS 127503 = NRRL 54222.
Type locality: Nepal.
Type substrate: Oryza sativa.
Descriptions and illustrations: See Sarver et al. (2011).
Diagnostic DNA barcodes: rpb1: KM361650; rpb2: KM361668;
tef1: KM889631.
nervisequum Fusarium (Fuckel) Fuckel, Jahrb. Nassauischen
Vereins Naturk. 23–24: 369. 1870.
Basionym: Labrella nervisequa Fuckel, Fungi Rhen. Exs., Fasc.
5, no. 427. 1863.
Apiognomonia platani (Lev.) L. Lombard, comb. nov. MycoBank MB 837698.
Basionym: Hymenula platani Lev., Ann. Sci. Nat., Bot., ser. 3, 9:
128. 1848.
Synonyms: Fusarium platani (Lev.) Mont., Ann. Sci. Nat., Bot.,
ser. 3, 11: 55. 1849.
Fusarium nervisequum f. platani (Lev.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 369. 1870.
Gloeosporidium platani (Lev.) Höhn., Sitzungsber. Kaiserl. Akad.
Wiss. Wien, Math.-Naturwiss. Cl., Abt. 1, 125: 95. 1916.
Myxosporina platani (Lev.) Höhn., Hedwigia 62: 48. 1920, nom.
inval., Art. 35.1.
Gloeosporium nervisequum (Fuckel) Sacc., Syll. Fung. 3: 711. 1884.
Discula nervisequa (Fuckel) M. Morelet, Bull. Soc. Sci. Nat.
Archeol. Toulon & Var 203: 12. 1973.
Gloeosporium platani Oudem., Ned. Kruidk. Arch., ser. 2, 1: 258.
1873.
Laestadia veneta Sacc. & Speg., Michelia 1: 351. 1878.
Carlia veneta (Sacc. & Speg.) Kuntze, Revis. Gen. Pl. 2: 846.
1891.
FUSARIUM
Apiospora veneta (Sacc. & Speg.) Sacc. ex Kleb., Z. Pflanzenkrankh. 12: 258. 1902.
Gnomonia veneta (Sacc. & Speg.) Kleb., Jahrb. Wiss. Bot. 41:
533. 1905, nom. illegit., Art. 53.1.
Gnomonia platani Kleb., Verhandl. Deutsch. Bot. Ges. 1: 28. 1914.
Guignardia veneta (Sacc. & Speg.) Traverso, Fl. Ital. Crypt. 1:
392. 1907.
Apiosporopsis veneta (Sacc. & Speg.) Traverso, Syll. Fung. 22:
78. 1913.
Apiognomonia veneta (Sacc. & Speg.) Höhn., Ann. Mycol. 16:
51. 1918.
Laestadia veneta var. cylindrasca Sacc. & Speg., Michelia 1:
369. 1878.
Laestadia cylindrasca (Sacc. & Speg.) Sacc., Syll. Fung. 1: 422.
1882.
Carlia cylindrasca (Sacc. & Speg.) Kuntze, Revis. Gen. Pl. 2:
846. 1891.
Guignardia cylindrasca (Sacc. & Speg.) Lindau (as ‘cylindracea’), in Engler & Prantl, Nat. Pflanzenfam., Teil. I, 1(1): 422.
1897.
Diaporthe veneta Sacc. & Speg., Michelia 1: 383. 1878.
Discella platani Peck, Rep. (Annual) New York State Mus. Nat.
Hist. 29: 49. 1878, nom. illegit., Art. 53.1.
Discula platani Sacc., Syll. Fung. 3: 674. 1884.
Sporonema platani B€aumler, Oesterr. Bot. Z. 40: 17. 1890.
Placosphaeria platani (B€aumler) Limber, Mycologia 47: 398.
1955.
Myxosporium platanicola Ellis & Everh. (as ‘platanicolum’), Proc.
Acad. Nat. Sci. Philadelphia 46: 372. 1894.
Cryptosporiopsis platanicola (Ellis & Everh.) G.F. Laundon, CBS
List of Cultures (Baarn): (1). 1975.
Gloeosporidina platani Butin & Kehr, Eur. J. Forest Pathol. 28:
299. 1998.
Lectotypus: BPI (Fuckel, Fungi Rhen. 427) of Labrella nervisequa Fuckel, designated in Sogonov et al. (2007).
Lectotype locality: Germany, Reichartshausen.
Lectotype substrate: Plantanus orientalis.
Epitypus: BPI 871953, designated in Sogonov et al. (2007).
Epitype locality: Switzerland, Geneva.
Epitype substrate: Plantanus orientalis.
Notes: Based on priority and synonymies proposed by Sogonov
et al. (2007), the name Hymenula platani Lev. (1848) takes
precedence over Laestadia veneta Sacc. & Speg. (1878).
Therefore, a new combination is proposed here applying the
older name.
newnesense Fusarium M.H. Laurence et al., Fungal Diversity
77: 360. 2015 [2016].
Holotypus: RBG 610.
Ex-type culture: NRRL 66241 = RBG 610.
Type locality: Australia, New South Wales, Newnes State
Forest.
Type substrate: Soil.
Descriptions and illustrations: See Laurence et al. (2016).
Diagnostic DNA barcodes: rpb1: JABCJW010000176; rpb2:
JABCJW010000963; tef1: KP083261.
ngaiotongaense Fusarium O'Donnell et al., Index Fungorum 440:
3. 2020.
Neocosmospora longissima Sand.-Den. & Crous, Persoonia
43: 141 (2019).
www.studiesinmycology.org
REDELIMITED
Holotypus: CBS H-23985.
Ex-type culture: CBS 126407 = G.J.S. 85-72.
Type locality: New Zealand, Russell State Forest, Ngaiotonga
Scenic Reserve.
Type substrate: From tree bark.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834230; rpb2: LR583846;
tef1: LR583621.
nicotianae Fusarium Oudem., Ned. Kruidk. Arch., ser. 3, 2: 777.
1902.
(See Fusarium merismoides)
Holotypus: ?L.
Type locality: Netherlands, Noord-Holland Province, Bussum.
Type substrate: Nicotiana tabacum.
nigrum Fusarium O.A. Pratt, J. Agric. Res. 13: 90. 1918.
(See Fusarium flocciferum)
Lectotypus (hic designatus, MBT 10000716): USA, Idaho, from
soil, 1918, O.A. Pratt, in J. Agric. Res. 13: 82, fig. 1J–L.
Notes: Synonym fide Wollenweber & Reinking (1935). As the
holotype specimen was not located, an illustration accompanying
the original protologue is designated here as lectotype.
nirenbergiae Fusarium L. Lombard & Crous, Persoonia 43: 29.
2018 [2019].
Holotypus: CBS H-23619.
Ex-type culture: CBS 840.88.
Type locality: Netherlands, Noord-Holland Province, Aalsmeer.
Type substrate: Dianthus caryophyllus.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb2: MH484887; tef1: MH484978.
nisikadoi Fusarium T. Aoki & Nirenberg, Mycoscience 38: 330.
1997.
Holotypus: BBA 69015 in B.
Ex-type culture: BBA 69015 = CBS 456.97 = MAFF
237506 = NRRL 25205 = NRRL 25308.
Type locality: Japan, Oita, Hita.
Type substrate: Triticum aestivum.
Descriptions and illustrations: See Nirenberg & Aoki (1997).
Diagnostic DNA barcodes: rpb1: MG282391; rpb2: MG282421;
tef1: KR909358.
nitidum Fusarium Berk. & M.A. Curtis, Grevillea 3: 98. 1875.
Holotypus: ?K(M).
Type locality: USA, Pennsylvania, Michener.
Type substrate: Aralia spinosa.
Note: Doubtful species fide Wollenweber & Reinking (1935).
nivale Fusarium Ces. ex Berl. & Voglino, Syll. Fung., Addit. I–IV:
390. 1886.
Microdochium nivale (Fr.) Samuels & I.C. Hallett, Trans. Brit.
Mycol. Soc. 81: 479. 1983.
Basionym: Lanosa nivalis Fr., Summa Veg. Scand. 2: 495. 1849.
Synonyms: Fusarium nivale (Fr.) Sorauer, Z. Pflanzenkrankh. 11:
220. 1901, nom. illegit., Art. 53.1.
Fusarium hibernans Lindau, Rabenh. Krypt.-Fl., ed. 2, 1(9): 542.
1909, nom. superfl., Art. 52.1.
Gerlachia nivalis (Ces. ex Berl. & Voglino) W. Gams & E. Müll.,
Netherlands J. Pl. Pathol. 86: 49. 1980.
Fusarium minimum Fuckel, Fungi Rhen. Exs., Fasc. 3, no. 213.
1863.
137
CROUS
ET AL.
Fusarium ustilaginis Rostr., Bot. Foren. Festskr. 54: 137. 1890,
nom. illegit., Art. 53.1.
Fusarium miniatum Prill. & Delacr., Bull. Soc. Mycol. France 7:
117. 1891, nom. illegit., Art. 53.1.
Fusarium tritici Erikss., Fungi Paras. Scand. Exs. no. 400. 1891,
nom. illegit., Art. 53.1.
Fusarium miniatulum Sacc., Syll. Fung. 10: 727. 1892.
Nectria pseudograminicola Weese, Ann. Mycol. 8: 466. 1910,
nom. inval., Art. 38.1.
Fusarium loliaceum Ducomet, Ann. Ecole
Natl. Agric. Rennes 2:
14. 1909.
Fusarium secalis Jacz., Bull. Trimestriel Soc. Mycol. France 28:
346. 1912, nom. illegit., Art. 53.1.
Sphaerulina divergens Rehm, Ann. Mycol. 11: 397. 1913.
Monographella divergens (Rehm) Petr., Ann. Mycol. 22: 144. 1924.
Calonectria nivalis Schaffnit, Mycol. Centralbl. 2: 257. 1913.
Griphosphaeria nivalis (Schaffnit) E. Müll. & Arx, Phytopathol. Z.
24: 356. 1955.
Micronectriella nivalis (Schaffnit) C. Booth, The Genus Fusarium:
42. 1971.
Monographella nivalis (Schaffnit) E. Müll., Rev. Mycol. (Paris) 41:
132. 1977.
Calonectria graminicola F. Stevens, Bot. Gaz. 65: 232. 1918,
nom. illegit., Art. 53.1.
Melioliphila graminicola Speg., Bol. Acad. Ci. (Cordoba) 26: 344.
1921.
Calonectria graminicola var. neglecta Krampe, Angew. Bot. 8:
252. 1926.
Monographella nivalis var. neglecta (Krampe) Gerlach,
Netherlands J. Pl. Pathol. 86: 49. 1980.
Fusarium nivale var. oryzae Zambett., Mitt. Inst. Colombo-Aleman Invest. Ci. 30: 489. 1950, nom. inval., Art. 39.1.
Syntypes: In HAL & ILL [Rabenhorst, Klotzschii Herb. Viv. Mycol.
no. 1439 (sub F. oxysporum)].
Type locality: Italy.
Type substrate: Poaceae.
niveum Fusarium E.F. Sm., Proc. Amer. Assoc. Advancem. Sci.
43: 289. 1894, nom. inval., Art. 36.1(a).
(See Fusarium oxysporum)
Authentic material: Not located.
Original locality: USA.
Original substrate: Citrullus vulgaris.
niveum Fusarium McAlpine, Australas. J. Pharm. 17: 3. 1902.
Note: Unable to locate protologue.
nodosum Fusarium L. Lombard & Crous, Fungal Syst. Evol. 4:
193. 2019.
Holotypus: CBS H-24018.
Ex-type culture: CBS 201.63.
Type locality: Portugal, Lisbon.
Type substrate: Seed of Arachis hypogaea.
Descriptions and illustrations: See Lombard et al. (2019a).
Diagnostic DNA barcodes: rpb1: MN120725; rpb2: MN120743;
tef1: MN120763.
noneumartii Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 3. 2020.
Neocosmospora noneumartii Sand.-Den. & Crous, Persoonia
43: 145. 2019.
Holotypus: CBS H-23989.
Ex-type culture: CBS 115658 = FRC S-0661.
Type locality: Israel, Palestine.
138
Type substrate: Solanum tuberosum.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218129; rpb2: MW446618;
tef1: LR583630.
nucicola Fusarium P. Karst. & Har., Rev. Mycol. (Toulouse) 12:
131. 1890.
(See Fusarium lateritium)
Holotypus: ?UPS fide Wollenweber, Fusaria Autogr. Delin. 1:
236. 1916.
Type locality: France.
Type substrate: Epicarp of nut.
Note: Synonym fide Wollenweber & Reinking (1935).
nurragi Fusarium (Summerell & L.W. Burgess) Benyon et al.,
Mycol. Res. 104: 1171. 2000.
Basionym: Fusarium avenaceum subsp. nurragi Summerell &
L.W. Burgess, Mycol. Res. 99: 289. 1995.
Holotypus: DAR 69502.
Ex-type culture: CBS 393.96 = DAR 69501 = F10108 = F11121.
Type locality: Australia, Victoria, Wilson's Promontory National
Park.
Type substrate: Soil.
Descriptions and illustrations: See Sangalang et al. (1995).
Diagnostic DNA barcodes: rpb1: MW928814; rpb2: MW928830;
tef1: MW928840.
nygamai Fusarium L.W. Burgess & Trimboli, Mycologia 78: 223.
1986.
Synonym: Gibberella nygamai Klaasen & P.E. Nelson, Mycologia
88: 967. 1997.
Holotypus: FRC-M-1375.
Ex-type culture: ATCC 58555 = BBA 69862 = CBS 749.97 = FRC
M-1375 = IMI 375354 = NRRL 13448.
Type locality: Australia, New South Wales, Narrabri.
Type substrate: Necrotic roots of Sorghum sp.
Descriptions and illustrations: See Burgess & Trimboli (1986).
Diagnostic DNA barcodes: rpb1: LT996202; rpb2: KU604262;
tef1: MT011009.
obliquiseptatum Fusarium, T. Aoki et al., Mycologia 111: 929.
2019.
Neocosmospora obliquiseptata (T. Aoki et al.) L. Lombard &
Sand.-Den., comb. nov. MycoBank MB 837699.
Basionym: Fusarium obliquiseptatum, T. Aoki et al., Mycologia
111: 929. 2019.
Holotypus: BPI 910970.
Ex-type culture: MAFF 246845 = NRRL 62611.
Type locality: Australia, Queensland, Beerwah.
Type substrate: A gallery wall of an ambrosia beetle (Euwallacea
sp.) infecting Persea americana.
Descriptions and illustrations: See Aoki et al. (2019).
Diagnostic DNA barcodes: rpb1: KC691606; rpb2: KC691637,
KC691666; tef1: KC691535.
Note: A new combination is provided in the genus Neocosmospora based on the phylogenetic relationship (Aoki et al.
2019) and morphology.
oblongum Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index Fungorum 440: 3. 2020.
Neocosmospora oblonga Sand.-Den. & Crous, Persoonia 43:
148. 2019.
Holotypus: CBS H-23990.
Ex-type culture: CBS 130325 = CDC B-4701= NRRL 28008.
FUSARIUM
Type locality: USA.
Type substrate: Eye of Homo sapiens.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834239; rpb2: LR583853;
tef1: LR583631.
obtusatum Fusarium Corda, Icon. Fung. 1: 3. 1837.
(See Fusarium tortuosum)
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, Liberec (Reichenberg).
Type substrate: Branches of trees and shrubs.
Note: Not Fusarium fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in PRM.
obtusisporum Fusarium Cooke & Harkn., Grevillea 12: 97. 1884.
Neonectria obtusispora (Cooke & Harkn.) Rossman et al.,
Phytopathol. Medit. 53: 529. 2014.
Synonyms: Cylindrocarpon obtusisporum (Cooke & Harkn.)
Wollenw., Fusaria Autogr. Delin. 1: 465. 1916.
Ramularia obtusispora (Cooke & Harkn.) Wollenw., Fusaria
Autogr. Delin. 1: 465. 1916.
Fusarium lineare Moesz, Bot. Közlem. 19: 57. 1920.
Holotypus: K(M) 128869.
Type locality: USA, California.
Type substrate: Twigs of Acacia sp.
obtusiusculum Fusarium Sacc., Michelia 2: 297. 1881.
(See Fusarium candidum Ehrenb.)
Holotypus: In PAD.
Type locality: Italy, Padua.
Type substrate: Nelumbium sp.
obtusum Fusarium (Cooke) Sacc., Syll. Fung. 4: 708. 1886.
Basionym: Fusisporium obtusum Cooke, Grevillea 5: 58. 1876.
Mycogloea macrospora (Berk. & Broome) McNabb, Trans. Brit.
Mycol. Soc. 48: 187. 1965.
Basionym: Dacrymyces macrosporus Berk. & Broome, Ann.
Mag. Nat. Hist., ser. 4, 11: 343. 1873.
Holotypus: In K(M) fide Index Fungorum.
Type locality: UK, Scotland, Forres.
Type substrate: Diatrype sp.
ochraceum Fusarium (Mont.) Sacc., Syll. Fung. 4: 722. 1886.
Basionym: Fusisporium ochraceum Mont., Ann. Sci. Nat., Bot.,
ser. 2, 3: 355. 1835.
Holotypus: In ?PC.
Type locality: Chile, Juan Fernandez Islands.
Type substrate: Bark.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
odoratissimum Fusarium Maryani et al., Stud. Mycol. 92: 159.
2019.
Synonym: Fusarium purpurascens Maryani et al., Stud. Mycol.
92: 160. 2018 [2019a].
Holotypus: InaCC F822 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F822.
Type locality: Indonesia, East Kalimantan, Kampung Salak
Martadinata.
Type substrate: Musa sp. cv. Pisang Kepok.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479618; rpb2: LS479386;
tef1: LS479828.
Notes: Re-analysis of the sequence data set of Maryani et al.
(2019a) revealed that the ex-type strain of F. purpurascens
www.studiesinmycology.org
REDELIMITED
(InaCC F971) clustered within the F. odoratissimum clade.
Therefore, we consider F. purpurascens a synonym of
F. odoratissimum.
oidioides Fusarium Speg., Rev. Mycol. (Toulouse) 8: 183. 1886.
Holotypus: In LPS (Fungi Japon. No. 2) fide Farr (1973).
Type locality: Japan, Tokyo.
Type substrate: Fallopia multiflora.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
oligoseptatum Fusarium T. Aoki et al., Fung. Syst. Evol. 1: 29.
2018.
Neocosmospora oligoseptata (T. Aoki et al.) Sand.-Den. &
Crous, Persoonia 43: 149. 2019.
Holotypus: BPI 910525.
Ex-type culture: CBS 143241 = FRC S-2581 = MAFF
246283 = NRRL 62579.
Type locality: USA, Pennsylvania, Dauphin.
Type substrate: From a live female ambrosia beetle (Euwallacea
validus), extracted from a gallery in a tree-of-heaven (Ailanthus
altissima).
Descriptions and illustrations: See Aoki et al. (2018).
Diagnostic DNA barcodes: rpb1: KC691596; rpb2: KC691627,
KC691656; tef1: KC691538.
ophioides Fusarium A. Jacobs, et al., Persoonia 46: 149. 2021.
Holotypus: CBS H-24659.
Ex-type culture: CBS 118512 = FCC 2979 = FCC 2980 = MRC
6744.
Type locality: South Africa, Mpumulanga Province, Ngodwana.
Type substrate: Panicum maximum.
Descriptions and illustrations: See Yilmaz et al. (2021).
Diagnostic DNA barcodes: rpb2: MN534303; tef1: EU921239.
opuli Fusarium Oudem., Hedwigia 37: 318. 1898.
Holotypus: ?L.
Type locality: Netherlands, Gelderland Province, Nunspeet.
Type substrate: Viburnum opulus.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
opuntiarum Fusarium Speg., Anales Mus. Nac. Hist. Nat. Buenos
Aires 6: 350. 1898 [1899].
(See Fusarium oxysporum)
Holotypus: In LPS (Fungi Argent. n.v.c. no. 866) fide Farr (1973).
Type locality: Argentina, La Plata.
Type substrate: Branches of Opuntia sp.
Note: Synonym fide Wollenweber & Reinking (1935).
orchidis Fusarium Petch, Ann. Roy. Bot. Gard. (Peradeniya) 6:
256. 1917.
(See Fusarium reticulatum)
Holotypus: PDA 4798.
Type locality: Sri Lanka.
Type substrate: Leaves of Orchidaceae.
Note: Synonym fide Wollenweber & Reinking (1935).
ornamentatum Fusarium (M.A.F. Barbosa) O'Donnell et al., Index
Fungorum 440: 3. 2020.
(See Fusarium neocosmosporiellum)
Holotypus: CBS 562.70 (preserved as metabolically inactive
culture).
Ex-type culture: ATCC 32363 = CBS 562.70 = IMI 251387.
Type locality: Guinea-Bissau.
Type substrate: Stored nuts of Arachis hypogaea.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
139
CROUS
ET AL.
Diagnostic DNA barcodes: rpb2: LR583901; tef1: DQ247606.
Note: Synonym fide Sandoval-Denis et al. (2019).
orobanches Fusarium Jacz., Ezhegodnik Svedeniy Boleznykh i
Povrezhdeniyakh Kult'turnykh i Dikorastushchikh Poleznykh
Rasteniy. Pertograd. 6: 190. 1910 [1912].
Holotypus: Not located.
Type locality: Russia, Saratov.
Type substrate: Orobanche sp.
Notes: Status unclear. Could be a synonym of F. oxysporum.
orthoceras Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 155. 1910.
(See Fusarium oxysporum)
Syntypes: B 70 0100192 & B 70 0100193.
Type locality: Germany, Berlin, Dahlem.
Type substrate: Solanum tuberosum.
Note: Typification pending further study of the syntypes in B.
orthoconium Fusarium Wollenw., Fusaria Autogr. Delin. 2: 637.
1926.
Mycogloea orthospora (Syd.) McNabb ex Dingley, Mem. New
York Bot. Gard. 49: 206. 1989.
Basionym: Microcera orthospora Syd., Ann. Mycol. 22: 317.
1924, non Fusarium orthosporum Sacc. 1902.
Synonyms: Fusarium microcera var. orthoconium (Wollenw.)
Bilaĭ, Mikrobiol. Zhurn. 49: 7. 1987, nom. inval., Arts. 35.1, 41.4.
Holotypus: Not located.
Type locality: New Zealand, Wellington, York Bay.
Type substrate: Nothofagus sp.
orthosporum Fusarium Sacc. & P. Syd., Syll. Fung. 16: 1100.
1902.
Cylindrodendrum orthosporum (Sacc. & P. Syd.) L. Lombard,
comb. nov. MycoBank MB 837700.
Basionym: Fusarium orthosporum Sacc. & P. Syd., Syll. Fung.
16: 1100. 1902.
Synonyms: Cylindrocarpon orthosporum (Sacc. & P. Syd.)
Wollenw., Fusaria Autogr. Delin. 1: 462. 1916.
Ramularia orthospora (Sacc. & P. Syd.) Wollenw., Fusaria
Autogr. Delin. 1: 462. 1916.
Neonectria hubeiensis W.Y. Zhuang et al., Fungal Diversity 24:
351. 2007.
Ilyonectria hubeiensis (W.Y. Zhuang et al.) Z.Q. Zeng & W.Y.
Zhuang, Phytotaxa 85: 17. 2013.
Cylindrodendrum hubeiense (W.Y. Zhuang et al.) L. Lombard &
Crous, Phytopathol. Medit. 53: 523. 2014.
Holotypus: In PAD.
Type locality: France.
Type substrate: Juglans nigra.
Descriptions and illustrations: See Zhuang et al. (2007) and
Lombard et al. (2014).
Notes: The epithet of Fusarium orthosporum Sacc. & P. Syd
(1902) predates that of Neonectria hubeiensis W.Y. Zhuang et al.
(2007). Therefore, a new combination is proposed here with the
older epithet.
oryzae Fusarium Vincens, Rev. Pathol. Veg. Entomol. Agric.
France 10: 126. 1923.
Holotypus: ?PC.
Type locality: Vietnam.
Type substrate: Oryza sativa.
Notes: Status unclear. Could be a synonym of F. fujikuroi.
osiliense Fusarium Bres. & Vestergr., Bot. Not. 1900: 33. 1900.
140
Septogloeum oxysporum Sacc. et al., Bull. Soc. Roy. Bot.
Belgique 29: 294. 1890.
Syntypes: In BPI, NEB, S & UPS.
Type locality: Estonia, Osilia.
Type substrate: Briza media.
Notes: Synonym fide Wollenweber & Reinking (1935). Typification pending further study of the syntypes.
ossicola Fusarium (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
714. 1886.
Basionym: Fusisporium ossicola Berk. & M.A. Curtis, Grevillea 3:
147. 1875.
(See Fusarium equiseti)
Holotypus: ?K(M).
Type locality: USA.
Type substrate: Old decaying bones.
Note: Synonyms fide Wollenweber & Reinking (1935).
osteophilum Fusarium Speg., Anales Soc. Ci. Argent. 10: 60.
1880.
(See Fusarium scirpi)
Holotypus: In LPS (Fungi Argent. pug. 2, no. 155) fide Farr
(1973).
Type locality: Argentina, Rio de la Plata, La Recoleta.
Type substrate: Decayed bones of Gallus sp. (chicken).
Note: Synonym fide Wollenweber & Reinking (1935).
otomycosis Fusarium Y.N. Ming & T.F. Yu, Acta Microbiol. Sin.
12: 178. 1966.
Holotypus: Not located.
Type locality: China, Beijing.
Type substrate: Ear of Homo sapiens.
Notes: Status unclear. Requires further investigation.
oxydendri Fusarium Ellis & Everh., Bull. Torrey Bot. Club 24: 477.
1897.
(See Fusarium cavispermum)
Syntypes: In BPI, BRU, CLEM, CUP, F, FLAS, ILL, ILLS, ISC,
MICH, MSC, MU, NEB, OSC, PH, PUL, UC & WIS.
Type locality: USA, West Virginia.
Type substrate: Oxydendrum arboreum.
Notes: Synonym fide Wollenweber & Reinking (1935). Typification pending further study of the syntypes.
oxysporum Fusarium Schltdl., Fl. Berol. 2: 139. 1824.
Synonyms: Fusisporium aurantiacum Link, Mag. Ges. Naturf.
Freunde Berlin 3: 19. 1809.
Fusarium aurantiacum (Link) Sacc., Syll. Fung. 4: 720. 1886,
nom. illegit., Art. 53.1.
Fusarium aurantiacum Corda, in Sturm, Deutschl. Fl., 3 Abt.
(Pilze Deutschl.) 2: 19. 1829.
Fusarium oxysporum var. aurantiacum (Corda) Rabenh.,
Deutschl. Krypt.-Fl., 1: 51. 1844.
Atractium aurantiacum (Corda) Bonord., Abh. Naturf. Ges. Halle
8: 135. 1851.
Fusisporium lagenariae Schwein., Trans. Amer. Philos. Soc., n.s.
4: 275. 1834.
Fusarium lagenariae (Schwein.) Sacc., Syll. Fung. 4: 724. 1886.
Hymenula equiseti Lib., Pl. Crypt. Arduenna Fasc. 3: no. 236.
1834.
Fusarium equisetorum (Lib.) Desm., Pl. Crypt. N. France no.
1546/1846? 1843.
Fusarium parasiticum Thüm., Nuovo Giorn. Bot. Ital. 12: 198.
1880, nom. illegit., Art. 53.1.
FUSARIUM
Fusarium thuemenii Sacc., Syll. Fung. 4: 722. 1886.
Fusisporium calcareum Thüm., Inst. Coimbra 28: 262. 1881.
Fusarium calcareum (Thüm.) Sacc., Syll. Fung. 4: 712. 1886.
Fusarium eucalyptorum Cooke & Harkn., Grevillea 9: 128.
1881.
Fusarium oxysporum f. eucalypti (Cooke & Harkn.) Arya & G.L.
Jain, Phytopathology 52: 641. 1962.
Fusarium oxysporum f. lycopersici Sacc., Syll. Fung. 4: 705. 1886.
Fusarium lycopersici (Sacc.) Mussat, Syll. Fung. 15: 144. 1901,
nom. inval., Art. 36.1(a), (c).
Fusarium lycopersici (Sacc.) Wollenw., Phytopathology 3: 29.
1913, nom. illegit., Art. 53.1.
Fusarium bulbigenum Cooke & Massee, Grevillea 16: 49. 1887.
Fusarium myosotidis Cooke, Grevillea 16: 49. 1887.
Leptosporium mycophilum P. Karst., Meddel. Soc. Fauna Fl.
Fenn. 16: 24. 1888.
Fusarium mycophilum (P. Karst.) Sacc., Syll. Fung. 10: 730. 1892.
?Selenosporium cuticola R. Blanch., Compt. Rend. Hebd.
Seances Acad. Sci., Ser. D. 111: 479. 1890.
?Fusarium cuticola (R. Blanch.) Gueg., Champ. Paras. Homme:
262. 1904.
Fusarium sclerodermatis Peck, Rep. (Annual) Regents Univ.
State New York New York State Mus. 43: 77. 1890, nom. illegit.,
Art. 53.1.
Fusarium peckii Sacc., Syll. Fung. 10: 727. 1892, nom. illegit.,
Art. 53.1 [pro. p. fide Wollenweber & Reinking (1935)].
Fusarium saccardoanum P. Syd., Syll. Fung. 14: 1128. 1899.
Fusarium vasinfectum G.F. Atk., Bull. Alabama Agric. Exp. Sta.
41: 28. 1892.
Fusarium cordae Massee, Brit. Fung.-Fl. 3: 481. 1893.
Fusarium niveum E.F. Sm., Proc. Amer. Assoc. Advancem. Sci.
43: 289. 1894, nom. inval., Art. 36.1(a).
Fusarium bulbigenum var. niveum E.F. Sm. ex Wollenw.,
Fusarien: 117. 1931.
Fusarium blasticola Rostr. (as ‘blasticolum’), Gartn.-Tidende
1895: 122. 1895.
Fusoma blasticola (Rostr.) Sacc. & Traverso, Syll. Fung. 20:
1241. 1911.
Fusarium bulbigenum var. blasticola (Rostr.) Wollenw., Z. Parasitenk. (Berlin) 3: 412. 1931.
Fusarium beticola A.B. Frank, Kampfbuch gegen die Sch€adlinge
unserer Feldfrüchte: 137. 1897.
Fusarium dianthi Prill. & Delacr., Compt. Rend. Hebd. Seances
Acad. Sci. 129: 745. 1899.
Fusarium oxysporum f. dianthi (Prill. & Delacr.) W.C. Snyder &
H.N. Hansen, Amer. J. Bot. 27: 66. 1940.
Fusarium oxysporum var. dianthi (Prill. & Delacr.) Raillo, Fungi of
the Genus Fusarium: 255. 1950.
Fusarium opuntiarum Speg., Anales Mus. Nac. Hist. Nat. Buenos
Aires 6: 350. 1898 [1899].
Fusoma pini Hartig, Lehrb. Pflanzenkrankh., Bot., Forstl., Landw.
G€art.: 116. 1900.
Fusarium laxum Peck, Bull. New York State Mus. Nat. Hist. 67:
30. 1903.
Fusarium lini Bolley, Proc. Annual Meeting Soc. Promot. Agric.
Sci. 22: 42. 1902.
Fusarium oxysporum f. lini (Bolley) W.C. Snyder & H.N. Hansen,
Amer. J. Bot. 27: 66. 1940.
Fusarium tabacivorum Delacr., Ann. Inst. Natl. Rech. Agron., ser.
2, 5: 207. 1906.
Fusarium candidulum Sacc., Ann. Mycol. 6: 567. 1908.
Fusarium cubense E.F. Sm., Science, N.Y. 31: 754. 1910.
www.studiesinmycology.org
REDELIMITED
Fusarium oxysporum var. cubense (E.F. Sm.) Wollenw., Fusarien: 119. 1935.
Fusarium oxysporum f. cubense (E.F. Sm.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 27: 66. 1940.
Fusarium orthoceras Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 155. 1910.
Fusarium oxysporum var. orthoceras (Appel & Wollenw.) Bilaĭ,
Microbiol. Zhurn. 49: 7. 1987.
?Fusarium violae F.A. Wolf, Mycologia 2: 21. 1910.
Fusarium albidoviolaceum Dasz. (as ‘albido-violaceum’), Bull.
Soc. Bot. Geneve, ser. 2, 4: 293. 1912.
Fusarium orthoceras var. albidoviolaceum (Dasz.) Wollenw.,
Fusaria Autogr. Delin. 1: 361. 1916.
Fusarium lycopersici Bruschi, Atti Reale Accad. Lincei, Rendiconti Cl. Sci. Fis., ser. 5, 21: 298. 1912.
Fusarium bulbigenum var. lycopersici (Bruschi) Wollenw. &
Reinking, Fusarien: 114. 1935.
Fusarium citrinum Wollenw., in Lewis, Bull. Maine Agric. Exp.
Sta. 219: 256. 1913.
Fusarium conglutinans var. citrinum (Wollenw.) Wollenw., Z.
Parasitenk. (Berlin) 3: 407. 1931.
Fusarium conglutinans Wollenw., Ber. Deutsch. Bot. Ges. 31: 34.
1913.
Fusarium orthoceras var. conglutinans (Wollenw.) Padwick, Indian J. Agric. Sci. 10: 282. 1940.
Fusarium oxysporum f. conglutinans (Wollenw.) W.C. Snyder &
H.N. Hansen, Amer. J. Bot. 27: 66. 1940.
Fusarium elegans Appel & Wollenw., Arbeiten Kaiserl. Biol. Anst.
Land- Forstw. 8: 94. 1913, nom. inval., Art. 36.1(a) (non Fusarium elegans W. Yamam. & Maeda 1962).
Fusarium batatas Wollenw. (as ‘batatae’), J. Agric. Res. 2: 268.
1914.
Fusarium bulbigenum var. batatas (Wollenw.) Wollenw., Z. Parasitenk. (Berlin) 3: 414. 1931.
Fusarium oxysporum f. batatas (Wollenw.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 27: 66. 1940.
Fusarium cepae Hanzawa, Mycol. Centralbl. 5(1): 5. 1914.
Fusarium oxysporum f. cepae (Hanzawa) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 27: 66. 1940.
Fusarium oxysporum var. cepae (Hanzawa) Raillo, Fungi of the
Genus Fusarium: 253. 1950.
Fusarium hyperoxysporum Wollenw., J. Agric. Res. 2: 268. 1914.
Fusarium angustum Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 203. 1915.
Fusarium lutulatum Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 209. 1915.
Fusarium vasinfectum var. lutulatum (Sherb.) Wollenw., Fusaria
Autogr. Delin. 3: 1019. 1930.
Fusarium lutulatum var. zonatum Sherb., Mem. Cornell Univ.
Agric. Exp. Sta. 6: 214. 1915.
Fusarium zonatum (Sherb.) Wollenw., Fusaria Autogr. Delin. 1:
392. 1916.
Fusarium vasinfectum var. zonatum (Sherb.) Wollenw., Fusaria
Autogr. Delin. 3: 1020. 1930.
Fusarium oxysporum var. asclerotium Sherb., Mem. Cornell
Univ. Agric. Exp. Sta. 6: 222. 1915.
Fusarium asclerotium (Sherb.) Wollenw., Fusaria Autogr. Delin.
1: 364. 1916.
Fusarium sclerotioides Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 214. 1915.
Fusarium sclerotioides var. brevius Sherb., Mem. Cornell Univ.
Agric. Exp. Sta. 6: 218. 1915.
141
CROUS
ET AL.
Fusarium trifolii Jacz., Jahrb. Pflanzenkrankh. Russl. VII-VIII:
Abt. 6. 1917.
Fusarium citrulli Taubenh., Bull. Texas Agric. Exp. Sta. 260: 27.
1920.
Fusarium malvacearum Taubenh., Bull. Texas Agric. Exp. Sta.
260: 27. 1920.
Fusarium poolense Taubenh., Bull. Texas Agric. Exp. Sta. 260:
27. 1920.
Fusarium macroxysporum Lindf., Meddel. Centralanst. Försöksv€as. Jordbruksomr. Avd. Lantbruksbot. 25: 8. 1922.
Fusarium spinaciae Hungerf., Phytopathology 13: 209. 1923.
Fusarium cromyophthoron Sideris, Phytopathology 14: 212.
1924.
Fusarium loncheceras Sideris, Phytopathology 14: 213. 1924.
Fusarium loncheceras var. microsporon Sideris, Phytopathology
14: 213. 1924.
Fusarium rhizochromatistes Sideris, Phytopathology 14: 212.
1924.
Fusarium sclerostromaton Sideris, Phytopathology 14: 213.
1924.
Fusarium zonatum f. 1 Link & Bailey, J. Agric. Res. 33: 941.
1926.
Fusarium zonatum f. 2 Link & Bailey, J. Agric. Res. 33: 941.
1926.
Fusarium conglutinans var. betae D. Stewart, Phytopathology 21:
67. 1931.
Fusarium oxysporum f. betae (D. Stewart) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 27: 66. 1940.
Fusarium oxysporum f. 7 Wollenw., Fusaria Autogr. Delin. 4:
1176. 1935
Fusarium apii P.E. Nelson & Sherb., Techn. Bull. Michigan Agric.
Exp. Sta 155: 42. 1937.
Fusarium orthoceras var. apii (R. Nelson & Sherb.) Wollenw. &
Reinking, Fusarien: 112. 1935.
Fusarium oxysporum f. apii (R. Nelson & Sherb.) W.C. Snyder &
H.N. Hansen, Amer. J. Bot. 27: 66. 1940.
Fusarium apii var. pallidum R. Nelson & Sherb., Techn. Bull.
Michigan Agric. Exp. Sta. 155: 42. 1937.
Fusarium bulbigenum var. apii (R. Nelson & Sherb.) Raillo, Fungi
of the Genus Fusarium: 251. 1950.
Cylindrophora albedinis Kill. & Maire, Bull. Soc. Hist. Nat. Afrique
N. 21: 97. 1930, nom. inval., Art. 36.1(b).
Fusarium oxysporum var. albedinis Kill. & Maire ex
Malençon, Rev. Mycol. (Paris) 15: 45– 60. 1950, nom. inval., Art.
36.1(b).
Fusarium oxysporum f. sp. albedinis Kill. & Maire ex W.L. Gordon, Canad. J. Bot. 43: 1310. 1965.
Fusarium perniciosum Hepting, Circ. U.S.D.A. 535: 7. 1939.
Fusarium oxysporum f. perniciosum (Hepting) Toole, Phytopathology 31: 599. 1941.
Fusarium vasinfectum var. perniciosum (Hepting) Carrera, Rev.
Fac. Agron. Buenos Aires 13(3): 483 1955
?Fusarium retusum Wellman, Phytopathology 33: 957. 1943.
Holotypus: HAL 1612 F.
Epitypus: CBS H-23620, designated in Lombard et al. (2019b).
Ex-epitype culture: CBS 144134.
Type locality: Germany, Berlin.
Type substrate: Solanum tuberosum.
Descriptions and illustrations: See Lombard et al. (2019b)
Diagnostic DNA barcodes: rpb2: MH484953; tef1: MH485044.
142
palczewskii Fusarium Jacz., Bull. Soc. Mycol. France 28: 345.
1912.
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000717): Russia, Ussuriysk, Primorsky krai (Far East Territory), grain of Lolium sp.,
1912, A.A. Jaczewski, in Bull. Soc. Mycol. France 28: 345,
fig. 1.
Notes: Synonyms fide Wollenweber & Reinking (1935). As no
holotype specimen could be located; an illustration accompanying the original protologue is designated here as lectotype.
pallens Fusarium Berk. & M.A. Curtis, Grevillea 3: 99. 1875,
nom. illegit., Art. 53.1.
Replacing synonym: Fusarium glumarum Sacc., Syll. Fung. 4:
706. 1886.
(See Fusarium incarnatum)
Authentic material: Car. Inf. no. 3799, in K(M).
Original locality: USA.
Original substrate: Juncus sp.
Note: Synonyms fide Wollenweber & Reinking (1935).
pallens Fusarium (Nees & T. Nees) Link, Sp. pl. 6(2): 104. 1825.
Basionym: Atractium pallens Nees & T. Nees, Nova Acta Phys.Med. Acad. Caes. Leop.-Carol. Nat. Cur. 9: 237. 1818.
Synonyms: Volutella pallens (Nees & T. Nees) Fr., Syst. Mycol. 3:
468. 1832.
Selenosporium pallens (Nees & T. Nees) Corda, Icon. Fung. 1: 7.
1837.
Holotypus: In B.
Type locality: Germany.
Type substrate: Fallen branch.
Notes: The type material of Atractium pallens is deposited at B
and examined by Gr€afenhan et al. (2011), identifying it as a
coelomycete.
pallidoroseum Fusarium (Cooke) Sacc., Syll. Fung. 4: 720. 1886.
Basionym: Fusisporium pallidoroseum Cooke, Grevillea 6: 139.
1878.
(See Fusarium incarnatum)
Holotypus: S. Car. no. 2279 in ?K(M).
Type locality: USA, South Carolina, Aiken.
Type substrate: Chenopodium anthelminticum.
Note: Synonyms fide Wollenweber & Reinking (1935).
pallidulum Fusarium Sacc. & Trotter, Syll. Fung. 22: 1483. 1913.
Replaced synonym: Atractium pallidum Bonord., Handb. Mykol.:
135. 1851.
Synonym: Fusarium pallidum (Bonord.) Sacc. & Traverso, Syll.
Fung. 19: 727. 1910, nom. illegit., Art. 53.1.
Lectotypus (hic designatus, MBT 10000718): Germany, decaying bark, 1913, H.F. Bonorden, in Handb. Mykol., tab. 10, fig.
219.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935). As no holotype specimen could be located,
an illustration accompanying the original protologue is designated here as lectotype.
pallidum Fusarium Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10:
359. 1869.
Holotypus: In K(M).
Type locality: Cuba.
Type substrate: Dead twigs.
FUSARIUM
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
palustre Fusarium W.H. Elmer & Marra, sp. nov. MycoBank
MB 837702.
Synonym: Fusarium palustre W.H. Elmer & Marra, Mycologia
103(4): 815. 2011, nom. inval., Art. 40.7.
Etymology. ‘palustre’, from Latin palus, referring to marsh habitat
in which this fungus is found.
For diagnosis see Elmer & Marra, Mycologia 103(4): 815.
2011.
Holotypus: CBS 126795 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 126796 = NRRL 54056.
Type locality: USA, Connecticut, Madison, Hammonasset Beach
State Park.
Type substrate: Spartina alterniflora.
Descriptions and illustrations: See Elmer & Marra (2011).
Diagnostic DNA barcodes: rpb1: KT597718; rpb2: KT597731;
tef1: GQ856941.
Notes: Elmer & Marra (2011) failed to indicate the holotype for
F. palustre, rendering the species name invalid (Art. 40.7). Here
we validate the name.
pampini Fusarium Thüm. & Pass., Pilze Weinst.: 50. 1878.
Gloeosporium physalosporae Cavara, Rev. Mycol. (Toulouse)
10: 99. 1888.
Holotypus: Not located.
Type locality: Italy, Parma.
Type substrate: Vitis vinifera.
Note: Synonym fide Wollenweber & Reinking (1935).
pandani Fusarium (Corda) Sacc., Syll. Fung. 4: 724. 1886.
Basionym: Fusisporium pandani Corda, Icon. Fung. 1: 11. 1837.
Lectotypus (hic designatus, MBT 10000719): Czech Republic,
Liberec (Reichenberg), Pandanus sp., 1837, A.C.J. Corda, in
Icon. Fung. 1, tab. 2, fig. 162.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935). As no holotype specimen could be located,
an illustration accompanying the original protologue is designated here as lectotype.
pannosum Fusarium Massee, Bull. Misc. Inform. Kew 1898: 117.
1898.
(See Fusarium sambucinum)
Holotypus: K(M) 191093.
Type locality: India, Punjab.
Type substrate: Cornus macrophylla.
Note: Synonym fide Wollenweber & Reinking (1935).
paraeumartii Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 3. 2020.
Neocosmospora paraeumartii Sand.-Den. & Crous, Persoonia
43: 149. 2019.
Holotypus: CBS H-23991.
Ex-type culture: BBA 62215 = CBS 487.76 = NRRL 13997.
Type locality: Argentina.
Type substrate: Decaying stem base of Solanum tuberosum.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834240; rpb2: LR583855;
tef1: DQ247549.
paranaense Fusarium S.S. Costa et al., Fungal Biology 120: 55.
2015 [2016].
(See Fusarium falciforme)
www.studiesinmycology.org
REDELIMITED
Holotypus: CML 1830.
Ex-type culture: CBS 141593 = CML 1830.
Type locality: Brazil, Goias State, Cristalina.
Type substrate: Diseased tissue of Glycine max.
Descriptions and illustrations: See Costa et al. (2016).
Diagnostic DNA barcodes: rpb2: KF680011; tef1: KF597797.
Note: Synonym fide Sandoval-Denis et al. (2019).
parasiticum Fusarium Westend., Bull. Acad. Roy. Sci. Belgique,
Cl. Sci., ser. 2, 11: 652. 1861.
(See Fusarium ciliatum)
Holotypus: BR5020140791441.
Type locality: Belgium, Louette-Saint-Pierre.
Type substrate: Sphaeria gigaspora.
Note: Synonym fide Wollenweber & Reinking (1935).
parasiticum Fusarium Thüm., Nuovo Giorn. Bot. Ital. 12: 198.
1880, nom. illegit., Art. 53.1.
Replacing synonym: Fusarium thuemenii Sacc., Syll. Fung. 4:
722. 1886.
(See Fusarium oxysporum)
Authentic material: Not located.
Original locality: Russia, Orenburg.
Original substrate: Betula pendula.
Note: Synonyms fide Wollenweber & Reinking (1935).
parasiticum Fusarium Ellis & Kellerm., J. Mycol. 3: 127. 1887,
nom. illegit., Art. 53.1.
Replacing synonym: Fusarium pucciniophilum Sacc. & P. Syd.,
Syll. Fung. 14: 1128. 1899.
(See Fusarium heterosporum)
Authentic material: Kellerman & Swingle 1104 in NY.
Original locality: USA, Manhattan.
Original substrate: Parasitic on Puccinia seymeriae on Swietenia
macrophylla.
Note: Synonyms fide Wollenweber & Reinking (1935).
parasiticum Fusarium Fautrey, Rev. Mycol. (Toulouse) 11: 153.
1889, nom. illegit., Art. 53.1.
Replacing synonym: Fusarium fautreyi Sacc., Syll. Fung. 10:
934. 1892.
(See Fusarium lateritium)
Authentic material: BR5020140789424.
Original locality: France, Noidan.
Original substrate: Vitis vinifera.
Note: Synonyms fide Wollenweber & Reinking (1935).
parceramosum Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 3. 2020.
Neocosmospora parceramosa Sand.-Den. & Crous, Persoonia
43: 151. 2019.
Holotypus: CBS H-23992.
Ex-type culture: CBS 115695 = CPC 1246.
Type locality: South Africa.
Type substrate: Soil.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb2: JX435249; tef1: JX435149.
parvisorum Fusarium Herron et al., Stud. Mycol. 80: 146. 2015.
Holotypus: PREM 60897.
Ex-type culture: CBS 137236 = CMW 25267.
Type locality: Colombia, Vivero, Pe~nas Negra, Valle del Cauca.
Type substrate: Pinus patula.
Descriptions and illustrations: See Herron et al. (2015).
143
CROUS
ET AL.
Notes: Comparisons of recently generated sequences for the
living ex-type (CBS 137236 = CMW 25267) of F. parvisorum
indicate a strain transposition or contamination by another Fusarium species. Therefore, this species needs to be recollected from
the type locality and substrate or sequences need to be generated
from the holotype specimen to confirm that it is indeed distinct.
paspali Fusarium Henn., Bot. Jahrb. Syst. 38: 129. 1905.
(See Fusarium avenaceum)
Syntype: In B as Zenker, Georg August, no. 2152 fide Hein
(1988).
Type locality: Cameroon, Bipindi.
Type locality: Paspalum sp.
Notes: Synonym fide Wollenweber & Reinking (1935). Typification pending further study of the syntype in B.
paspalicola Fusarium Henn., in Warburg, Monsunia 1: 38. 1899
[1900].
(See Fusarium heterosporum)
Holotypus: In B fide Wollenweber, Fusaria Autogr. Delin. 1: 299.
(1916) & Hein (1988).
Type locality: Philippines, Mindanao, Davao.
Type substrate: Paspalum sp.
Note: Synonym fide Wollenweber & Reinking (1935).
patouillardii Fusarium Sacc. (as ‘patouillardi’), Syll. Fung. 10:
729. 1892.
Replaced synonym: Fusarium uredinicola Pat. & Gaillard, Bull.
Soc. Mycol. France 4: 127. 1888, nom. illegit., Art. 53.1.
Holotypus: ?PC or FH.
Type locality: Venezuela, Caracas.
Type substrate: Parasitic on Sphaerellopsis filum on Puccinia
pallidissima.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
peckii Fusarium Sacc., Syll. Fung. 4: 713. 1886.
Replaced synonym: Fusisporium parasiticum Peck, Rep.
(Annual) New York State Mus. Nat. Hist. 29: 53. 1878, non
Fusarium parasiticum Westend. 1861.
Holotypus: NYSf2260.
Type locality: USA, New York, Albany.
Type substrate: Sphaeria collinsii.
Notes: Status unclear. Not treated by Wollenweber & Reinking
(1935) or Booth (1971).
peckii Fusarium Sacc., Syll. Fung. 10: 727. 1892, nom. illegit.,
Art. 53.1.
Replaced synonyms: Fusarium sclerodermatis Peck, Rep.
(Annual) Regents Univ. State New York New York State Mus. 43:
77. 1890, nom. illegit., Art. 53.1, non Fusarium sclerodermatis
Oudem. 1889.
Fusarium saccardoanum Syd., Syll. Fung. 13: 1130. 1898.
(See Fusarium oxysporum pr. p. & Fusarium avenaceum pr.
p.)
Authentic material: NYSf2731.
Original locality: USA, New York, Suffolk.
Original substrate: Scleroderma vulgaris.
Note: Synonyms fide Wollenweber & Reinking (1935).
pelargonii Fusarium P. Crouan & H. Crouan, Fl. Finistere: 14.
1867.
(See Fusarium merismoides)
Holotypus: ?PC.
Type locality: France, Finistere.
144
Type substrate: Pelargonium sp.
Note: Synonym fide Wollenweber & Reinking (1935).
peltigerae Fusarium Westend., Herb. Crypt. Belg. Fasc. 9: no.
414. 1849.
(See Fusarium ciliatum)
Syntypes: In BR & PH (Herb. Crypt. Belg. 9: no. 414).
Type locality: Belgium.
Type substrate: Peltigera rufescens.
Notes: Synonym fide Wollenweber & Reinking (1935). Typification pending further study of the syntypes.
penicillatum Fusarium (Harz) Sacc., Syll. Fung. 4: 710. 1886.
Basionym: Menispora penicillata Harz, Bull. Soc. Imp. Naturalistes Moscou 44: 127. 1871.
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000720): Germany, Berlin,
decaying Sclerotium clavus, 1886, C. Harz, in Bull. Soc. Imp.
Naturalistes Moscou 44, tab. 1, fig. 4.
Notes: Synonym fide Wollenweber & Reinking (1935). As
no holotype specimen could be located, an illustration accompanying the original protologue is designated here as lectotype.
pentaclethrae Fusarium Henn., Hedwigia 44: 71. 1905.
(See Fusarium coccidicola)
Syntype: In B (Ule no. 3011) fide Hein (1988).
Type locality: Brazil, Manaus, Rio Nigro.
Type substrate: Leaves of Pentaclethra sp.
Notes: Synonym fide Wollenweber & Reinking (1935). Typification pending further study of the syntype in B.
penzigii Fusarium Schroers et al., Mycologia 101: 61. 2009.
Bisifusarium penzigii (Schroers et al.) L. Lombard & Crous,
Stud. Mycol. 80: 225. 2015.
Holotypus: CBS H-20125.
Ex-type culture: CBS 317.34 = NRRL 22109.
Type locality: UK, Surrey.
Type substrate: Decayed wood of Fagus sylvatica.
Descriptions and illustrations: See Schroers et al. (2009).
Diagnostic DNA barcodes: rpb1: KM232211; rpb2: KM232362;
tef1: EU926324.
pernambucanum Fusarium A.C.S. Santos et al., Mycologia
111: 253. 2019.
Holotypus: URM 91193.
Ex-type culture: MUM 1862 = URM 7559.
Type locality: Brazil, Pernambuco, Paudalho.
Type substrate: Aleurocanthus woglumi.
Descriptions and illustrations: See Santos et al. (2019).
Diagnostic DNA barcodes: rpb1: MH668869; rpb2: LS398519;
tef1: LS398489.
perniciosum Fusarium Hepting, Circul. U.S.D.A. 535: 7. 1939.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: USA.
Type substrate: Albizia julibrissin.
persicae Fusarium (Sacc.) G.F. Atk., J. Elisha Mitchell Sci. Soc.
8: 41. 1892.
Basionym: Cercospora persicae Sacc. (as ‘persica’), Hedwigia
15: 119. 1876.
Mycosphaerella pruni-persicae Deighton, Trans. Brit. Mycol.
Soc. 50: 328. 1967.
FUSARIUM
Synonyms: Cercosporella persicae (Sacc.) Sacc. (as ‘persica’),
Michelia 2: 20. 1880.
Clasterosporium persicae (Sacc.) Tsuji, Ann. Phytopathol. Soc.
Japan 1(2): 33. 1919.
Miuraea persicae (Sacc.) Hara, Byogaichu-Hoten (Manual of
Pests and Diseases): 224. 1948.
Mycosphaerella persicae B.B. Higgins & F.A. Wolf (as ‘persica’),
Phytopathology 27: 695. 1937.
Syntype: In HAL, ILL & NEB (Saccardo, Mycoth. Ven. no. 598).
Type locality: Italy.
Type substrate: Prunus persica.
persicinum Fusarium J.W. Xia et al., Persoonia 43: 215 2019.
Holotypus: CBS H-24068.
Ex-type culture: CBS 479.83.
Type locality: Unknown.
Type substrate: Unknown.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170428; tef1: MN170495.
personatum Fusarium Cooke, in Harkness, Grevillea 7: 12. 1878.
(See Fusarium allescherianum)
Holotypus: ?K(M).
Type locality: USA, California.
Type substrate: Oreodaphne californica.
Note: Synonym fide Wollenweber & Reinking (1935).
perseae Fusarium (Sand.-Den. & Guarnaccia) O'Donnell et al.,
Index Fungorum 440: 3. 2020.
Neocosmospora perseae Sand.-Den. & Guarnaccia, Fungal
Syst. Evol. 1: 136. 2018.
Holotypus: CBS H-23433.
Ex-type culture: CBS 144142 = CPC 26829.
Type locality: Italy, Catania, San Leonardello.
Type substrate: Trunk canker lesions on Persea americana.
Descriptions and illustrations: See Guarnaccia et al. (2018).
Diagnostic DNA barcodes: rpb1: MW218130; rpb2: LT991909;
tef1: LT991902.
peruvianum Fusarium L. Lombard & Crous, Fungal Syst. Evol.
4: 194. 2019.
Holotypus: CBS H-24019.
Ex-type culture: CBS 511.75.
Type locality: Peru.
Type substrate: Seedlings of Gossypium sp.
Descriptions and illustrations: See Lombard et al. (2019a).
Diagnostic DNA barcodes: rpb1: MN120728; rpb2: MN120746;
tef1: MN120767.
pestis Fusarium Sorauer, Atlas Pfl.-Krankh. 4: 19, pl. XXV. 1890.
(See Fusarium azukiicola)
Holotypus: Not located.
Type locality: Germany.
Type substrate: Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935).
petersiae Fusarium L. Lombard, Persoonia 39: 457. 2017.
Holotypus: CBS H-23233.
Ex-type culture: CBS 143231.
Type locality: Netherlands, Gelderland Province, Arnhem.
Type substrate: Soil.
Descriptions and illustrations: See Crous et al. (2017).
Diagnostic DNA barcodes: rpb1: MG386139; rpb2: MG386150;
tef1: MG386160.
www.studiesinmycology.org
REDELIMITED
petroliphilum Fusarium (Q.T. Chen & X.H. Fu) Geiser et al.,
Fungal Genet. Biol. 53: 69. 2013.
Neocosmospora petroliphila (Q.T. Chen & X.H. Fu) Sand.Den. & Crous, Persoonia 41: 121. 2018.
Basionym: Fusarium solani var. petroliphilum Q.T. Chen & X.H.
Fu, Acta Mycol. Sin., Suppl. 1: 330. 1987.
Synonyms: Fusarium solani f. sp. cucurbitae (Race 2) W.C.
Snyder & H.N. Hansen, Amer. J. Bot. 28: 740. 1941.
Holotypus: HMAS 43748.
Ex-type culture: FRC S-2176 = NF4475 = NRRL 22268.
Type locality: China, Beijing.
Type substrate: Deteriorated petroleum.
Descriptions and illustrations: See Sandoval-Denis & Crous
(2018).
peziziforme Fusarium Berk. & M.A. Curtis (as ‘pezizaeforme’), J.
Linn. Soc., Bot. 10: 360. 1869.
Holotypus: In K(M).
Type locality: Cuba.
Type substrate: Poaceae.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
pezizoides Fusarium Desm., Ann. Sci. Nat., Bot., ser. 3, 18: 373.
1852.
Trochila craterium (DC.) Fr., Summa Veg. Scand. 2: 367. 1849.
Basionym: Sphaeria craterium DC., Fl. Franç., ed. 3, 2: 298.
1805.
Synonyms: Phacidium craterium (DC.) Gillet, Champ. France
Discomyc. (7): 167. 1886.
Sphaeria punctiformis var. hederae Pers., Syn. Meth. Fung. 1:
90. 1801.
Myxosporium paradoxum De Not., Mem. Reale Accad. Sci.
Torino, ser. 2, 3: 81. 1841.
Gloeosporium paradoxum (De Not.) Mont., in Berkeley &
Broome, Ann. Mag. Nat. Hist. 5: 455. 1850.
Gloeosporidium paradoxum (De Not.) Petr., Ann. Mycol. 20: 14.
1922.
Cryptocline paradoxa (De Not.) Arx, Verh. Kon. Ned. Akad.
Wetensch., Afd. Natuurk. 51: 115. 1957.
Gloeotrochila paradoxa (De Not.) Petr., Sydowia 1: 50. 1947.
Trochila craterium var. nucleata Rehm, Ber. Bayer. Bot. Ges. 13:
125. 1912.
Ceuthospora hederae Grove, Bull. Misc. Inform. Kew 1923: 355.
1923.
Holotypus: ?PC.
Type locality: France.
Type substrate: Peziza insidiosa.
Note: Synonyms fide Wollenweber & Reinking (1935).
pezizoideum Fusarium (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
711. 1886.
Basionym: Fusisporium pezizoideum Berk. & M.A. Curtis, Grevillea 3: 147. 1875.
(See Fusarium sambucinum)
Holotypus: ?K(M).
Type locality: USA, Pennsylvania.
Type substrate: Stems of herbaceous plants.
Note: Synonyms fide Wollenweber & Reinking (1935).
phacidioideum Fusarium Dearn., Mycologia 21: 331. 1929.
Holotypus: JD 4303 in DAOM.
Type locality: Canada, Vancouver, Stanley Park.
Type substrate: Dead branches of Pseudotsuga taxifolia.
145
CROUS
ET AL.
Note: Status unclear; requires recollection from type locality and
substrate.
pharetrum Fusarium L. Lombard & Crous, Persoonia 43: 32.
2018 [2019].
Holotypus: CBS H-23621.
Ex-type culture: CBS 144751 = CPC 30824.
Type locality: South Africa.
Type substrate: Aloidendron dichotomum.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928815; rpb2: MH484952;
tef1: MH485043.
phaseoli Fusarium (Burkh.) T. Aoki & O'Donnell, Mycologia 95:
671. 2003.
Basionym: Fusarium martii f. phaseoli Burkh., Mem. Cornell Univ.
Agric. Exp. Sta. 26: 1007. 1919.
(See Fusarium azukiicola)
Lectotypus (hic designatus, MBT 10000721): USA, New York,
roots of Phaseolus vulgaris, 1919, W.H. Burkholder, in Mem.
Cornell Univ. Agric. Exp. Sta. 26: 1009, fig. 134.
Notes: Synonym fide Sandoval-Denis et al. (2019). Although
Burkholder deposited several specimens in CUP, none are
directly linked to the original protologue (Burkholder 1919).
Several of these specimens appear to have been isolated from
greenhouse assays undertaken by Burkholder. Therefore, an
illustration accompanying the original protologue is designated
here as lectotype.
phialophorum Fusarium Maryani et al., Stud. Mycol. 92: 169.
2018 [2019].
Holotypus: InaCC F971 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F971.
Type locality: Indonesia, South Kalimantan, Tanah Bumbu,
Kampung Betung.
Type substrate: Musa var. Pisang Awak.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479545; rpb2: LS479292;
tef1: LS479741.
phormii Fusarium Henn., Verh. Bot. Vereins Prov. Brandenburg
40: 175. 1898 [1899].
Colletotrichum phormii (Henn.) D.F. Farr & Rossman, Mycol.
Res. 110: 1403. 2006.
Synonym: Gloeosporium phormii (Henn.) Wollenw., Fusaria
Autogr. Delin. No. 498. 1916, nom. illegit., Art. 53.1, non
Gloeosporium phormii Sacc. 1915.
Holotypus: B 70 0005220.
Epitypus: CBS H-20720, designated in Damm et al. (2012).
Ex-epitype: A.R. 3546 = CBS 118194.
Type locality: Germany, Berlin.
Type substrate: Phormium tenax.
phragmiticola Fusarium Kirschst., Ann. Mycol. 34: 183. 1936,
nom. inval., Art. 39.1.
Authentic material: B 70 0100199, B 70 0100200, B 70010020.
Original locality: Germany.
Original substrate: Phragmites communis.
phragmitis Fusarium Matsush., Icon. Microfung. Matsush. Lect.:
72. 1975, nom. inval., Art. 40.1.
Authentic material: Not indicated.
Original locality: Japan.
Original substrate: Rotten wood of Fagus crenata.
146
phyllachorae Fusarium Henn., in de Wildeman, Mission E.
Laurent, Fasc. 4: 363. 1907.
Syntype: Laurent in B fide Hein (1988).
Type locality: Democratic Republic of Congo, between Kinshasa and Kwamouth.
Type substrate: Panicum maximum.
Notes: Not Fusarium fide Wollenweber & Reinking (1935).
Typification pending further study of the syntype in B.
phyllogenum Fusarium (Cooke & Peck) Sacc., Syll. Fung. 4: 703.
1886.
Basionym: Fusisporium phyllogenum Cooke & Peck, Rep.
(Annual) New York State Mus. Nat. Hist. 29: 53. 1878.
Syntype: NYSf2335.
Type locality: USA, New York, Albany, Bethlehem.
Type substrate: Erigeron annuum.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935). Typification pending further study of the syntype in NYS.
phyllophilum Fusarium Nirenberg & O'Donnell, Mycologia 90:
447. 1998.
Holotypus: IMI 202874.
Ex-type culture: BBA 63625 = CBS 216.76 = DAOM
225132 = IMI 375338 = NRRL 13617.
Type locality: Italy.
Type substrate: Dracaena deremensis.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998).
Diagnostic DNA barcodes: rpb1: KF466399; rpb2: KF466410;
tef1: KF466421.
phyllostachydicola Fusarium W. Yamam., Trans. Mycol. Soc.
Japan 3: 118. 1962.
Basionym: Gibberella phyllostachydicola W. Yamam., Hyogo
Univ. Agric. ser. Agric. Biol. 3: 15. 1957.
Lectotypus (hic designatus, MBT 10000722): Japan, Tamba,
Sasayama-cho, culms of Phyllostachys bambusoides, 31 Aug.
1956, W. Yamamoto, in Hyogo Univ. Agric. ser. Agric. Biol. 3: 17,
figs 16–18.
Descriptions and illustrations: See Yamamoto et al. (1957).
Notes: This species requires recollection from the type host and
locality. As no holotype specimen could be located, an illustration
accompanying the original protologue is designated here as
lectotype.
pilosicola Fusarium Yilmaz et al., Persoonia 46: 152. 2021.
Holotypus: PREM 63216.
Ex-type culture: CMWF 1183 = NRRL 29124 = NY007.H7.
Type locality: USA, Florida.
Type substrate: Bidens pilosa.
Descriptions and illustrations: See Yilmaz et al. (2021).
Diagnostic DNA barcodes: rpb2: MN534248; tef1: MN534055.
pininemorale Fusarium Herron et al., Stud. Mycol. 80: 146.
2015.
Holotypus: PREM 60901.
Ex-type culture: CBS 137240 = CMW 25243.
Type locality: Colombia, Risaralda, Angela Maria (Santa Rosa).
Type substrate: Pinus tecunumanii.
Descriptions and illustrations: See Herron et al. (2015).
Notes: Comparisons of recently generated sequences from the
living ex-type (CBS 137240 = CMW 25243) of F. pininemorale
indicate a strain transposition or contamination by another
Fusarium species. Therefore, this species needs to be
FUSARIUM
recollected from the type locality and substrate or sequences
need to be generated from the holotype specimen to confirm its
phylogenetic affiliation.
piperis Fusarium (F.C. Albuq.) O'Donnell et al., Index Fungorum
440: 3. 2020.
Neocosmospora piperis (F.C. Albuq.) Sand.-Den. & Crous,
Persoonia 43: 152. 2019.
Basionym: Fusarium solani f. piperis F.C. Albuq., Circ. Inst.
Agron. N. 5: 19. 1961.
Holotypus: IAN 825 in the herbarium of Embrapa Amazo^nia
Oriental.
Epitypus: CBS H-23993, designated in Sandoval-Denis et al.
(2019).
Ex-epitype culture: CBS 145470 = CML 1888 = G.J.S. 8914 = NRRL 22570.
Type locality: Brazil.
Type substrate: Piper nigrum.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834241; rpb2: EU329513;
tef1: AF178360.
sic et al., Sci. Rep. 8(no. 1252):
pisi Fusarium (F.R. Jones) A. Si
2. 2018, nom. inval., Art. F.5.1.
Neocosmospora pisi (F.R. Jones) Sand.-Den. & Crous, Persoonia 43: 154. 2019.
Basionym: Fusarium martii var. pisi F.R. Jones, J. Agric. Res. 26:
459. 1923.
Synonyms: Fusarium solani f. pisi (F.R. Jones) W.C. Snyder &
H.N. Hansen, Amer. J. Bot. 28: 740. 1941.
Fusarium vanettenii O'Donnell et al., Index Fungorum 440: 5.
2020.
Fusarium solani var. martii ‘f 2’ Wollenw., Z. Parasitenk. (Berlin)
3: 290. 1931.
Hypomyces solani f. sp. pisi Reichle, W.C. Snyder & Matuo,
Nature 203: 664. 1964.
Lectotypus: Jones (1923; fig. 1 on p. 463), designated in
Sandoval-Denis et al. (2019).
Epitypus: CBS H-23994, designated in Sandoval-Denis et al.
(2019).
Ex-epitype culture: ATCC MYA-4622 = CBS 123669 = NRRL
45880 = Vanetten 77-13-4.
Type locality: USA.
Type substrate: Sexual cross of parents from Pisum sativum and
soil from a potato field.
sic et al. (2018b) and
Descriptions and illustrations: See Si
Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: JX171543; rpb2: EU329640;
tef1: FJ240352.
plagianthi Fusarium (Dingley) O'Donnell & Geiser, Phytopathology 103: 404. 2013.
Neocosmospora plagianthi (Dingley) L. Lombard & Crous,
Stud. Mycol. 80: 227. 2015.
Basionym: Nectria plagianthi Dingley, Trans. Roy. Soc. New
Zealand 79: 196. 1951.
?Nectria pulverulenta Dingley, Trans. Roy. Soc. New Zealand 83:
657. 1956.
Holotypus: PDD 10916.
Type locality: New Zealand, Fiordland, Hollyford Valley.
Type substrate: Plagianthus betulinus.
Descriptions and illustrations: See Dingley (1951) and Samuels
& Brayford (1994).
www.studiesinmycology.org
REDELIMITED
platani Fusarium (Lev.) Mont., Ann. Sci. Nat., Bot., ser. 3, 11: 55.
1849.
Basionym: Hymenula platani Lev., Ann. Sci. Nat., Bot., ser. 3, 9:
128. 1848
(See Fusarium nervisequum)
Holotypus: ?PC.
Type locality: France.
Type substrate: Platanus orientalis.
platanoidis Fusarium Oudem., Ned. Kruidk. Arch., ser. 3, 2: 1131.
1904.
Holotypus: ?L.
Type locality: Netherlands, Gelderland Province, Nunspeet.
Type substrate: Acer platanoides.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
poae Fusarium (Peck) Wollenw., in Lewis, Bull. Maine. Agric.
Exp. Sta. 219: 254. 1913 [1914].
Basionym: Sporotrichum poae Peck, Bull. New York State Mus.
67: 29. 1904 [1903].
Synonyms: Fusarium tricinctum f. poae (Peck) W.C. Snyder &
H.N. Hansen, Amer. J. Bot. 32: 663. 1945.
Fusarium sporotrichiella var. poae (Peck) Bilaĭ, Yadovitye griby
na zerne khlebnykh zlakov (Poisonous fungi on cereal seed): 86.
1953, nom. inval., Art. 39.1.
Fusarium sporotrichiella var. poae (Peck) Bilaĭ, Microbiol. Zhurn.
49: 6. 1987, nom. inval., Arts. 35.1, 41.4.
Sporotrichum anthophilum Peck, Bull. New York State Mus. 105:
28. 1906.
Fusarium maydiperdum Bubak, Centralbl. Bakteriol. Parasitenk.,
2. Abth. 31: 497. 1911.
Holotypus: NYSf2393.
Type locality: USA, New York, Geneva.
Type substrate: Sheaths and culms of Poa pratensis.
Epitypus (hic designatus, MBT 10000723): USA, North Dakota,
Minot, from infected barley kernel, date and collector unknown,
NRRL 26941 (preserved as metabolically inactive culture).
Ex-epitype culture: NRRL 26941.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982) and Leslie
& Summerell (2006).
Diagnostic DNA barcodes: rpb1: KU171686; rpb2: KU171706;
tef1: JABFFD010000730.1
Note: No living material linked to the holotype is available for this
important mycotoxin producing species, and therefore, an epitype is designated here to provide taxonomic stability for this
species.
poincianae Fusarium Pass., Atti Reale Accad. Lincei, Rendiconti
Cl. Sci. Fis., ser. 4, 4: 105. 1888.
Holotypus: Not located.
Type locality: Italy, Parma.
Type substrate: Poinciana gilliesii.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
polymorphum Fusarium Matr., Rech. Devel. Muced.: 84. 1892.
(See Fusarium sambucinum)
Lectotypus (hic designatus, MBT 10000724): France, horse
dung, 1892, L. Matruchot, in Rech. Devel. Muced., Pl. 7, figs
6–14.
Notes: Synonym fide Wollenweber & Reinking (1935). As no
holotype specimen could be located, an illustration accompanying the original protologue is designated here as lectotype.
147
CROUS
ET AL.
polymorphum Fusarium Marchal, Bull. Soc. Roy. Bot. Belgique
34: 145. 1895, nom. illegit., Art. 53.1.
(See Fusarium aderholdii)
Authentic material: Not located.
Original locality: Belgium, Brussels.
Original substrate: Homo sapiens.
Notes: Synonym fide Wollenweber & Reinking (1935).
polyphialidicum Fusarium Marasas et al., Mycologia 78: 678. 1986.
(See Fusarium concolor)
Holotypus: DAOM 192986.
Ex-type culture: ATCC 60096 = CBS 961.87 = DAR
52851 = FRC M-2405 = MRC 3389 = NRRL 13459.
Type locality: South Africa, Mpumalanga Province, Nelspruit.
Type substrate: Plant debris in soil.
Descriptions and illustrations: See Marasas et al. (1986).
Diagnostic DNA barcodes: rpb1: JX171455; rpb2: JX171569;
tef1: MH742681.
poncetii Fusarium Guiart (as ‘ponceti’), Compt.-Rend. Seances
Mem. Soc. Biol. 73: 271. 1912, nom. inval., Art. 36.1(a).
Authentic material: Not located.
Original locality: ?France.
Original substrate: Homo sapiens granuloma teleangiectaticum.
Notes: Status unclear. Not treated by any of Wollenweber &
Reinking (1935), Booth (1971), or Gerlach & Nirenberg (1982).
Fusarium proliferatum var. minus Nirenberg, Mitt. Biol. Bundesanst. Land- Forstw. 169: 43. 1976. nom. inval., Art. 41.3.
Lectotypus: Microfungi of the Solomon Islands and Papua-New
Guinea: 11, fig 121.2, designated by Yilmaz et al. (2021).
Lectotype locality: Papua New Guinea.
Lectotype substrate: Forest soil.
Epitypus: CBS 480.96 (preserved as metabolically inactive culture), designated by Yilmaz et al. (2021).
Epitype locality: Papua New Guinea, Morobe Province, Bulolo.
Epitype substrate: Forest soil.
Ex-epitype culture: CBS 480.96 = IAM 14682 = NRRL
26427 = NY007.B6.
Descriptions and illustrations: See Matsushima (1971), Yilmaz
et al. (2021).
Diagnostic DNA barcodes: rpb2: MN534272; tef1: MN534059.
protoensiforme Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 3. 2020.
Neocosmospora protoensiformis Sand.-Den. & Crous, Persoonia 43: 156. 2019.
Holotypus: CBS H-23995.
Ex-type culture: CBS 145471 = G.J.S. 90-168 = NRRL 22178.
Type locality: Venezuela.
Type substrate: Bark of dicot tree.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834244; rpb2: EU329498;
tef1: AF178334.
poolense Fusarium (as ‘poolensis’) Taubenh., Bull. Texas Agric.
Exp. Sta. 260: 27. 1920.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000725): USA, Citrullus
lanatus, 1920, J.J. Taubenhaus, in Bull. Texas Agric. Exp. Sta.
260: 30, fig. 8i.
Notes: Synonym fide Wollenweber & Reinking (1935). As no
holotype specimen could be located, an illustration accompanying the original protologue is designated here as lectotype.
protractum Fusarium Lev., Ann. Sci. Nat., Bot., ser. 3, 9: 246.
1848.
(See Fusarium lateritium)
Holotypus: ?PC.
Type locality: France, Romainville.
Type substrate: Dead shoots of Solanum dulcamara.
Note: Synonym fide Wollenweber & Reinking (1935).
praegraminearum Fusarium Gr€afenhan & O'Donnell, Mycologia 108: 1232. 2016.
Holotypus: PDD 47563.
Ex-type culture: CBS 141369 = ICMP 8996 = NRRL 39664.
Type locality: New Zealand, North Island, Levin (near
Wellington).
Type substrate: Litter in maize paddock.
Descriptions and illustrations: See Gr€afenhan et al. (2016).
Diagnostic DNA barcodes: rpb1: KX260125; rpb2: KX260126;
tef1: KX260120.
prunorum Fusarium McAlpine, Fungus Diseases of stone-fruit
trees in Australia: 91. 1902.
(See Fusarium candidum (Link) Sacc.)
Lectotypus (hic designatus, MBT 10000726): Australia, Victoria,
Melbourne, Burnley, from shriveled and blackened apricot fruit,
Jun. 1900, D. McAlpine, in Fungus Diseases of stone-fruit trees
in Australia (1902), pl. XX, fig. 42.
Notes: Synonym fide Wollenweber & Reinking (1935). As no
holotype specimen could be located, an illustration accompanying the original protologue is designated here as lectotype.
prieskaense Fusarium G.J. Marais & Sand.-Den., Stud. Mycol.
98 (no. 100116): 50. 2021.
Holotypus: CBS H-24660.
Ex-type culture: CAMS 001176 = CBS 146498 = CPC 30826.
Type locality: South Africa, Northern Cape Province, Prieska.
Type substrate: Prunus spinosa.
Descriptions and illustrations: See this study.
Diagnostic DNA barcodes: rpb1: MW834190; rpb2: MW834007;
tef1: MW834275.
pseudacaciae Fusarium Rapaics, Z. Pflanzenkrankh. 25: 208.
1915.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Hungary, Debrecen.
Type substrate: Robinia pseudoacaciae.
Note: Synonym fide Wollenweber & Reinking (1935).
proliferatum Fusarium (Matsush.) Nirenberg ex Gerlach &
Nirenberg, Mitt. Biol. Bundesanst. Land- Forstw. 209: 309. 1982.
Basionym: Cephalosporium proliferatum Matsush., Microfungi of
the Solomon Islands and Papua-New Guinea: 11. 1971.
Synonyms: Fusarium proliferatum (Matsush.) Nirenberg, Mitt.
Biol. Bundesanst. Land- Forstw. 169: 38. 1976, nom. inval., Art.
41.3.
148
pseudensiforme Fusarium Samuels et al., Mycologia 103: 1323.
2011.
Neocosmospora pseudensiformis Samuels et al., Mycologia
103: 1323. 2011.
Holotypus: BPI 881226.
Ex-type culture: CBS 125729 = FRC S-1834 = G.J.S 0295 = G.J.S 9318 = NRRL 46517.
Type locality: Sri Lanka, Wagamba, Kurunegala, Arangakele.
Type substrate: Bark of tree.
FUSARIUM
Descriptions and illustrations: See Nalim et al. (2011).
Diagnostic DNA barcodes: rpb1: KC691615; rpb2: KC691645;
tef1: KC691555.
pseudoanthophilum Fusarium Nirenberg et al., Mycologia 90:
461. 1998.
Holotypus: In B.
Ex-type culture: BBA 69002 = CBS 414.97 = IMI 376112 = NRRL
25211.
Type locality: Zimbabwe, Gambiza.
Type substrate: Zea mays.
Descriptions and illustrations: See Nirenberg et al. (1998).
Diagnostic DNA barcodes: rpb1: MT010949; rpb2: MT010980;
tef1: MK639073.
pseudocircinatum Fusarium O'Donnell & Nirenberg, Mycologia 90: 448. 1998.
Holotypus: B 70 0001689.
Ex-type culture: BBA 69636 = CBS 126.73= CBS
449.97 = DAOM 225117 = IMI 375316 = NRRL 22946.
Type locality: Ghana.
Type substrate: Solanum sp.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998).
Diagnostic DNA barcodes: rpb1: MG838070; rpb2: MN724939;
tef1: MG838023.
pseudoeffusum Fusarium Murashk., Proc. Siberian Agric. Acad.
Omsk 3: 106. 1924.
(See Fusarium acuminatum)
Holotypus: Not located.
Type locality: Russia, Siberia.
Type substrate: Triticum polonicum.
Note: Synonym fide Wollenweber & Reinking (1935).
pseudograminearum Fusarium O'Donnell & T. Aoki, Mycologia
91: 604. 1999.
Holotypus: BPI 746087.
Ex-type culture: FRC R-5291 = NRRL 28062.
Type locality: Australia, New South Wales, Young.
Type substrate: Hordeum vulgare.
Descriptions and illustrations: See Aoki & O'Donnell (1999).
Diagnostic DNA barcodes: rpb1: JX171524; rpb2: JX171637;
tef1: AF212468.
pseudoheterosporum Fusarium Jacz., Bull. Soc. Mycol. France
28: 347. 1912.
(See Fusarium avenaceum)
Holotypus: Not located.
Type locality: France.
Type substrate: Lolium sp. and Triticum sp.
Note: Synonym fide Wollenweber & Reinking (1935).
pseudonectria Fusarium Speg., Anales Mus. Nac. Hist. Nat.
Buenos Aires 6: 351. 1898 [1899].
(See Fusarium avenaceum)
Holotypus: In LPS (Fungi Argent. n.v.c. no. 867) fide Farr (1973).
Type locality: Ecuador, San Salvador Island.
Type substrate: Dead culms of Poaceae.
Note: Synonym fide Wollenweber & Reinking (1935).
pseudonygamai Fusarium O'Donnell & Nirenberg, Mycologia
90: 449. 1998.
Holotypus: B 70 0001688.
Ex-type culture: BBA 69552 = CBS 417.97 = DAOM
225136 = FRC M-1166 = IMI 375342 = NRRL 13592.
www.studiesinmycology.org
REDELIMITED
Type locality: Nigeria.
Type substrate: Pennisetum typhoides.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998).
Diagnostic DNA barcodes: rpb1: LT996205; rpb2: LT996152;
tef1: AF160263.
pseudoradicicola Fusarium (Sand.-Den. & Crous) O'Donnell
et al., Index Fungorum 440: 3. 2020.
Neocosmospora pseudoradicicola Sand.-Den. & Crous, Persoonia 43: 157. 2019.
Holotypus: CBS H-23996.
Ex-type culture: ARSEF 2313 = CBS 145472 = NRRL 25137.
Type locality: Papua New Guinea, East New Britain, Keravat,
Lowlands Agricultural Experiment Station.
Type substrate: Diseased pods of Theobroma cacao.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218133; rpb2: JF741084;
tef1: JF740757.
pseudotonkinense Fusarium (Sand.-Den. & Crous) O'Donnell
et al., Index Fungorum 440: 3. 2020.
Neocosmospora pseudotonkinensis Sand.-Den. & Crous,
Persoonia 43: 159. 2019.
Holotypus: CBS H-23997.
Ex-type culture: CBS 143038.
Type locality: Netherlands, Zuid-Holland Province, Leiden.
Type substrate: Cornea of Homo sapiens.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb2: LR583867; tef1: LR583640.
pteridis Fusarium Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 45: 466. 1894.
Septogloeum pteridis (Ellis & Everh.) Wollenw., Fusaria Autogr.
Delin. 1: 446. 1916.
Syntypes: In BPI, BRU, CUP, FLAS, ILL, ISC, MICH, MSC, MU,
NEB, NY, OSC, PH, PUL, WIS & WSP.
Type locality: USA, New Jersey, Gloucester, Newfield.
Type substrate: Phyllachora flabella on Pteris aquilina.
pucciniophilum Fusarium Sacc. & P. Syd., Syll. Fung. 14: 1128.
1899.
Replaced synonym: Fusarium parasiticum Ellis & Kellerm., J.
Mycol. 3 (11): 127. 1887, nom. illegit., Art. 53.1.
(See Fusarium heterosporum)
Holotypus: Kellerman & Swingle no. 1104 in NY.
Type locality: USA, Kansas, Manhattan.
Type substrate: Parasitic on Puccinia seymeriae on leaves of
Solidago macrophylla.
Note: Synonym fide Wollenweber & Reinking (1935).
pulvinatum Fusarium (Link) Nees, Syst. Pilze: 32. 1817.
Basionym: Atractium pulvinatum Link, Mag. Ges. Naturf.
Freunde Berlin 8: 32. 1816.
Holotypus: Not located.
Type locality: Poland, Wrocław.
Type substrate: Hanging scrub branches.
Notes: Status unclear. Not treated by any of Wollenweber &
Reinking (1935), Booth (1971), or Gerlach & Nirenberg (1982).
pulvinatum Fusarium (Berk. & Broome) Sacc., Syll. Fung. 4: 699.
1886, nom. illegit., Art. 53.1.
Basionym: Fusisporium pulvinatum Berk. & Broome, J. Linn.
Soc., Bot. 14: 102. 1873 [1875].
(See Fusarium sambucinum)
Holotypus: In K(M).
149
CROUS
ET AL.
Type locality: Sri Lanka.
Type substrate: Bark.
Note: Synonym fide Wollenweber & Reinking (1935).
punctiforme Fusarium Durieu & Mont., Expl. Sci. Algerie 1: 335.
1848.
(See Fusarium reticulatum)
Holotypus: Not located.
Type locality: Algeria.
Type substrate: Citrus aurantium.
Note: Synonym fide Wollenweber & Reinking (1935).
purpurascens Fusarium Maryani et al., Stud. Mycol. 92: 160.
2018 [2019].
(see Fusarium odoratissimum)
Holotypus: InaCC F886 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F886.
Type locality: Indonesia, East Kalimantan, Kampung Salak
Martadinata.
Type substrate: Musa var. Pisang Kepok.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb2: LS479385; tef1: LS479827.
pusillum Fusarium Wollenw., Fusaria Autogr. Delin. 2: 550. 1924.
(See Fusarium dimerum)
Lectotypus (hic designatus, MBT 10000727): Germany, Solanum tuberosum, 1919, H.W. Wollenweber, in Fusaria Autogr.
Delin. 2: 550. 1924.
Note: As no holotype specimen could be located, an illustration
accompanying the original protologue is designated here as
lectotype.
putaminum Fusarium (Thüm.) Sacc., Syll. Fung. 4: 703. 1886.
Basionym: Fusisporium putaminum Thüm., Oesterr. Bot. Z. 27:
272. 1877.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Austria, Klosterneuburg.
Type substrate: Prunus domestica.
Note: Synonyms fide Wollenweber & Reinking (1935).
putrefaciens Fusarium Osterw., Mitth. Thurgauischen Naturf.
Ges. 16: 123. 1904.
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000728): Switzerland,
Zürich, fruit and seeds of Pyrus sp., 1904, collector unknown, in
Osterwalder, Mitth. Thurgauischen Naturf. Ges. 16, tab. 2, figs
10–30.
Notes: Synonym fide Wollenweber & Reinking (1935). As no
holotype specimen could be located, an illustration accompanying the original protologue is designated here as lectotype.
pyrinum Fusarium Schwein., Trans. Amer. Philos. Soc., n.s. 4:
302. 1834, unavailable, see Art. F.3.4.
(See Fusarium lactis)
pyrinum Fusarium (Fr.) Sacc., Syll. Fung. 4: 720. 1886.
Basionym: Fusisporium pyrinum Fr., Syst. Mycol. 3: 445. 1832,
nom. sanct.
(See Fusarium avenaceum)
Holotypus: Not located.
Type locality: Sweden.
Type substrate: Rotten fruit of Pyrus communis.
Note: Synonym fide Wollenweber & Reinking (1935).
150
pyrochroum Fusarium (Desm.) Sacc., Michelia 1: 534. 1879.
Calonectria pyrochroa (Desm.) Sacc., Michelia 1: 308. 1878.
Basionym: Selenosporium pyrochroum Desm., Ann. Sci. Nat.,
Bot., ser. 3, 14: 111. 1850.
Synonyms: Nectria pyrochroa Desm., Pl. Crypt. N. France, ed. 2:
no. 372. 1856.
Calonectria daldiniana De Not., Comment. Soc. Crittog. Ital. 2:
477. 1867.
Fusarium pyrochroum var. diatrypellicola P. Syd., Mycoth.
March., Cent. 41: no. 4063. 1893.
Nectria abnormis Henn., Hedwigia 36: 219. 1897.
Holotypus: In ?PAD or PC.
Type locality: France.
Type substrate: Sambucus nigra.
quercicola Fusarium Oudem., Ned. Kruidk. Arch., ser. 3, 2: 777.
1902.
Holotypus: ?L.
Type locality: Netherlands, Noord-Holland Province, Bussum.
Type substrate: Quercus rubra.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
quercinum Fusarium O'Donnell et al., Index Fungorum 440: 4.
2020.
Neocosmospora quercicola Sand.-Den. & Crous, Persoonia
43: 159. 2019.
Holotypus: CBS H-23998.
Ex-type culture: CBS 141.90 = NRRL 22652.
Type locality: Italy.
Type substrate: Quercus cerris.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834247; rpb2: LR583869;
tef1: DQ247634.
radicicola Fusarium Wollenw., J. Agric. Res. 2: 257. 1914.
(See Fusarium solani)
Lectotypus: Plate XVI, fig. K, in Wollenweber (1914), designated
in Sandoval-Denis et al. (2019).
Lectotype locality: USA, Washington.
Lectotype substrate: Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935) & SandovalDenis et al. (2019).
ramigenum Fusarium O'Donnell & Nirenberg, Mycologia 90:
451. 1998.
Holotypus: B 70 0001687.
Ex-type culture: BBA 68592 = CBS 418.97 = DAOM
225137 = IMI 375343 = NRRL 25208.
Type locality: USA, California.
Type substrate: Ficus carica.
Descriptions and illustrations: See Nirenberg & O'Donnell (1998).
Diagnostic DNA barcodes: rpb1: KF466401; rpb2: KF466412;
tef1: AF160267.
ramosum Fusarium (Batista & H. Maia) O'Donnell et al., Index
Fungorum 440: 4. 2020.
Basionym: Hyaloflorea ramosa Bat. & H. Maia, Anais Soc. Biol.
Pernambuco 13: 155. 1955.
Synonyms: Neocosmospora ramosa (Bat. & H. Maia) L.
Lombard & Crous, Stud. Mycol. 80: 227. 2015.
(See Fusarium lichenicola C. Massal.)
Holotypus: IMUR 410.
Ex-type culture: CBS 509.63 = IMUR 410 = MUCL 8050.
Type locality: Brazil.
FUSARIUM
Type substrate: Air.
Diagnostic DNA barcodes: rpb2: LR583843; tef1: LR583618.
Note: Synonymies fide Sandoval-Denis & Crous (2018).
ramulicola Fusarium Sawada, Special Publ. Coll. Agric. Natl.
Taiwan Univ. 8: 228. 1959, nom. inval., Art. 39.1.
Authentic material: Not located.
Original locality: Taiwan.
Original substrate: Branches of Citrus tankan f. koshotankan.
Note: This name is invalid because of missing Latin diagnosis.
rectiphorum Fusarium Samuels et al. (as ‘rectiphorus’), Mycologia 103: 1324. 2011.
Neocosmospora rectiphora Samuels et al., Mycologia 103:
1324. 2011.
Neocosmospora bomiensis Z.Q. Zeng & W.Y. Zhuang, Phytotaxa
319: 177. 2017.
Holotypus: BPI 881229.
Ex-type culture: CBS 125727 = FRC S-1831 = G.J.S. 02-89.
Type locality: Sri Lanka, Wagamba Province, vic. Kurunegala,
Arangakele.
Type substrate: Bark.
Descriptions and illustrations: See Nalim et al. (2011).
Diagnostic DNA barcodes: rpb1: MW834249; rpb2: LR583871;
tef1: LR583641.
redolens Fusarium Wollenw., Phytopathology 3: 29. 1913 and
Ber. Deutsch. Bot. Ges. 31: 31. 1913.
Synonyms: Fusarium oxysporum var. redolens (Wollenw.) W.L.
Gordon, Canad. J. Bot. 30: 238. 1952.
Fusarium solani var. redolens (Wollenw.) Bilaĭ, Fusarii (Biologija i
sistematika): 288. 1955.
?Fusarium retusum Wellman, Phytopathology 33: 957. 1943.
Holotypus: Not located.
Type locality: Unknown.
Type substrate: Pisum sativum.
Lectotypus (hic designatus, MBT 10000729): Unknown, Pisum
sativum, 1913, H.W. Wollenweber, in Phytopathology 3: 31, fig.
E.
Epitypus (hic designatus, MBT 10000730): Germany, BerlinDahlem, vascular bundle of Dianthus caryophyllus, 16 May 1959,
D. Hantschke & W. Gerlach, CBS 360.87 (preserved as metabolically inactive culture).
Ex-epitype culture: ATCC 16067 = BBA 9526 = CBS
248.61 = CBS 360.87 = DSM 62390 = NRRL 20426 = NRRL
25600.
Descriptions and illustrations: See Gerlach & Pag (1961),
Gerlach & Nirenberg (1982) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: MT409433; rpb2: MT409443;
tef1: MT409453.
Notes: As both protologue publications occurred more or less
simultaneously for F. redolens, we select the illustration provided
in Phytopathology as lectotype, since no holotype material could
be located. Gerlach & Nirenberg (1983) considered CBS 248.61
(= CBS 360.87) a good representative of F. redolens, which was
initially designated by Gerlach & Pag (1961) as representative of
F. redolens f. sp. dianthi. Therefore, an epitype is designated
here to provide taxonomic stability for this species.
regulare Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 4. 2020.
Neocosmospora regularis Sand.-Den. & Crous, Persoonia 43:
162. 2019.
Holotypus: CBS H-23999.
www.studiesinmycology.org
REDELIMITED
Ex-type culture: CBS 230.34
Type locality: Netherlands, Zeeland Province, Zuid Beveland,
near Kloetinge.
Type substrate: Pisum sativum.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb2: LR583873; tef1: LR583643.
rekanum Fusarium Lynn & Marinc., Antonie van Leeuwenhoek
113: 816. 2020.
Neocosmospora rekana (Lynn & Marinc.) L. Lombard & Sand.Den., comb. nov. MycoBank MB 837706.
Basionym: Fusarium rekanum Lynn & Marinc., Antonie van
Leeuwenhoek 113: 816. 2020.
Holotypus: PREM 62333.
Ex-type culture: CMW 52862 = PPRI 27163.
Type locality: Indonesia, Sumatra, Riau, Pelalawan.
Type substrate: Acacia crassicarpa infested with Euwallacea
perbrevis.
Descriptions and illustrations: See Lynn et al. (2020).
Diagnostic DNA barcodes: rpb2: MN249137, MN249108; tef1:
MN249151.
Note: Based on the phylogenetic position of this species related
to the ‘ambrosia’ clade as illustrated by Lynn et al. (2020), we
provide a new combination in the genus Neocosmospora.
reticulatum Fusarium Mont., Ann. Sci. Nat., Bot., ser. 2, 20:
379. 1843.
Synonyms: ?Fusarium leucoconium Corda, Icon. Fung. 1: 4.
1837. (fide Wollenweber & Reinking 1935).
?Fusarium punctiforme Durieu & Mont., Expl. Sci. Algerie 1: 335.
1848.
Fusisporium flavidum Bonord., Bot. Zeitung (Berlin) 19: 194. 1861.
Fusarium flavidum (Bonord.) Sacc., Syll. Fung. 4: 698. 1886.
Fusarium ampelodesmi Fautrey & Roum., in Roumeguere, Rev.
Mycol. (Toulouse) 13: 82. 1891.
Fusarium epithele McAlpine, Fungus Diseases of Citrus trees in
Australia: 80. 1899.
Fusarium orchidis Petch, Ann. Roy. Bot. Gard. (Peradeniya) 6:
256. 1917.
Fusarium negundinis Sherb., in Hubert, J. Agric. Res. 26: 451.
1923.
Fusarium reticulatum var. negundinis (Sherb.) Wollenw., Z.
Parasitenk. (Berlin) 3: 351. 1931.
Fusarium heterosporum var. negundinis (Sherb.) Raillo, Fungi of
the Genus Fusarium: 217. 1950.
Fusarium reticulatum var. medium Wollenw., Z. Parasitenk.
(Berlin) 3: 358. 1931.
Lectotypus (hic designatus, MBT 10000731): France, NouvelleAquitaine, Saint-Sever, Citrullus sp., 1843, L. Dufour, in Montagne, Ann. Sci. Nat., Bot., 2 ser. 20: 379: pl. 16, fig. 3.
Epitypus (hic designatus, MBT 10000732): Germany, Rellingen/
Holstein, bark lesion of Sophora japonica, Jun. 1976, R.
Schwarz, CBS 473.76 (preserved as metabolically inactive).
Ex-epitype culture: BBA 63657 = CBS 473.76 = NRRL 20684.
Descriptions and illustrations: See Gerlach & Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: MW928816; tef1: MW928841.
Notes: Gerlach & Nirenberg (1983) considered CBS 473.76 a
good representative of F. reticulatum. As no holotype specimen
could be located, an illustration is designated as lectotype here
and an epitype is designated to provide taxonomic stability for
this species.
retusum Fusarium Wellman, Phytopathology 33: 957. 1943.
151
CROUS
ET AL.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: USA, Indiana.
Type substrate: Solanum lycopersicum.
rhabdophorum Fusarium Berk. & Broome, Ann. Mag. Nat. Hist.,
ser. 4, 17: 142. 1876.
Holotypus: In K(M).
Type locality: UK, Scotland, Forres.
Type substrate: Dead sticks.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
rhizochromatistes Fusarium Sideris, Phytopathology 14: 212.
1924.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000733): USA, California,
Stockton, roots of Allium cepa, 1924, C.P. Sideris, in Phytopathology 14, pl. XI.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration
was designated as lectotype.
rhizogenum Fusarium Pound & Clem., Bot. Surv. Nebraska 3:
12. 1894.
(See Fusarium candidum Ehrenb.)
Holotypus: NEB0040548.
Type locality: USA, Lincoln.
Type substrate: Roots of Malus domestica seedlings.
Note: Synonym fide Wollenweber & Reinking (1935).
rhizogenum Fusarium Aderh., Centralbl. Bacteriol. Parasitenk.,
1. Abth., 6: 623. 1900, nom. illegit., Art. 53.1.
(See Fusarium aderholdii)
Authentic material: Not located.
Original locality: Germany.
Original substrate: Malus domestica.
Notes: Synonym fide Wollenweber & Reinking (1935). The
original publication could not be checked but Sorauer (1923)
clearly stated that Aderhold only used the name Fusarium rhizogenum Pound & Clem. to describe a disease using the latter
name.
rhizophilum Fusarium Corda, Icon. Fung. 2: 3. 1838.
Synonym: Fusisporium georginae Klotzsch, Herb. Viv. Mycol.,
Cent. 2: 186. 1832, nom. nud., Art. 38.1(a).
(See Fusarium merismoides)
Lectotypus (hic designatus, MBT 10000734): Czech Republic,
Prague, roots of garden plants, 1838, A.C.J. Corda, in Icon.
Fung. 2, Tab. VIII, fig. 15.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
rhizophorae Fusarium (Dayar.) O'Donnell et al., Index Fungorum
440: 4. 2020.
Neocosmospora rhizophorae Dayar., Mycosphere 11: 112.
2020.
Holotypus: MFLU 17-2588.
Ex-type culture: MFLUCC 17-2461.
Type locality: Thailand, Krabi Province, Phang Nga.
Type substrate: Submerged wood of Rhizophora.
Descriptions and illustrations: See Dayarathne et al. (2020)
152
rhodellum Fusarium McAlpine, Proc. Linn. Soc. New South
Wales 24: 122. 1899.
Lectotypus (hic designatus, MBT 10000735): Kerguelen
Islands, Pringlea antiscorbutica, 1899, D. McAlpine, in Proc.
Linn. Soc. New South Wales 24: Pl. XIII, Fig. 7.
Notes: Not Fusarium fide I. Pascoe. No holotype specimen could
be located and therefore an illustration is designated as
lectotype.
rhoicola Fusarium Fautrey, Rev. Mycol. (Toulouse) 17: 171.
1895.
(See Fusarium graminearum)
Holotypus: ?PC.
Type locality: France via USA.
Type substrate: Rhus toxicodendron.
Note: Synonym fide Wollenweber & Reinking (1935).
ricini Fusarium (Berenger) Bizz., Fl. Ven. Critt. 1: 539. 1885.
Basionym: Fusisporium ricini Berenger, Mem. Accad. Agric.
Verona 44: 257. 1866.
(See Fusarium sambucinum)
Holotypus: Not located.
Type locality: Italy.
Type substrate: Ricinus communis.
Note: Synonym fide Wollenweber & Reinking (1935).
rigidiusculum Fusarium (Berk. & Broome) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 32: 664. 1945.
Basionym: Nectria rigidiuscula Berk. & Broome, J. Linn. Soc.,
Bot. 14: 116. 1873 [1875].
(See Fusarium colorans)
Holotypus: ?K(M).
Type locality: ?Sri Lanka.
Type substrate: Bark.
Note: Synonym fide Wollenweber & Reinking (1935).
rimicola Fusarium Sacc. (as ‘rimicolum’), Michelia 2: 297. 1881.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Italy, Padua.
Type substrate: Erythrina crista-galli.
Note: Synonym fide Wollenweber & Reinking (1935).
rimosum Fusarium (Peck) Sacc., Syll. Fung. 4: 713. 1886.
Basionym: Fusisporium rimosum Peck, Rep. (Annual) New York
State Mus. Nat. Hist. 30: 58. 1878.
(See Fusarium merismoides)
Holotypus: NYSf2609.
Type locality: USA, New York, Albany.
Type substrate: Cut ends of stalks of Zea mays.
Note: Synonym fide Wollenweber & Reinking (1935).
riograndense Fusarium Dalle Rosa et al., J. Mycol. Med. 28: 33.
2018.
Neocosmospora riograndensis (Dalle Rosa et al.) Sand.-Den.
& Crous, Persoonia 43: 165. 2019.
Holotypus: UFMG-CM F12570.
Ex-type culture: UFMG-CM F12570 = URM-7361.
Type locality: Brazil, Rio Grande do Sul, Porto Alegre, Hospital
de Clínicas de Porto Alegre.
Type substrate: Nasal cavity of Homo sapiens.
Descriptions and illustrations: See Dalle Rosa et al. (2018).
Diagnostic DNA barcodes: rpb2: KX534003; tef1: KX534002.
FUSARIUM
robiniae Fusarium Pass., Atti Reale Accad. Lincei, Rendiconti Cl.
Sci. Fis., ser. 4, 7: 51. 1891.
(See Fusarium sarcochroum)
Holotypus: ?PARMA.
Type locality: Italy, Padua.
Type substrate: Robinia pseudoacacia.
Note: Synonym fide Wollenweber & Reinking (1935).
robustum Fusarium Gerlach, Phytopathol. Z. 88: 36. 1977.
Holotypus: In B.
Isotypus: CBS H-629.
Ex-type culture: BBA 63667 = CBS 637.76 = FRC R-5821 = IMI
322102 = NRRL 13392.
Type locality: Argentina.
Type substrate: Araucaria angustifolia.
Descriptions and illustrations: See Gerlach (1977c).
Diagnostic DNA barcodes: rpb2: MW928831; tef1: MW928842.
roesleri Fusarium Thüm., Pilze Weinst.: 51. 1878.
(See Fusarium merismoides)
Lectotypus (hic designatus, MBT 10000736): Austria, Klosterneuburg, Vitis vinifera, 1878, K.A.E.J. Thümen, in Pilze
Weinst. Tab. 3, fig. 7.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
rollandianum Fusarium Sacc., Syll. Fung. 11: 650. 1895.
Replaced synonym: Fusarium cydoniae Roum. & Fautrey, Rev.
Mycol. (Toulouse) 14: 170. 1892, nom. illegit., Art. 53.1, non
Allescher 1892.
Syntype: ILL00220295 (Fautrey, Fungi Sel. Gall. Exs. No. 6120).
Type locality: France.
Type substrate: Fruit of Cydonia vulgaris.
Notes: Not Fusarium fide Wollenweber & Reinking (1935).
Typification pending further study of the syntype lodged in ILL.
rosae Fusarium (Preuss) Sacc., Syll. Fung. 4: 697. 1886.
Basionym: Selenosporium rosae Preuss, Linnaea 24: 150. 1851.
Holotypus: Not located; not preserved in B fide HolubovaJechova et al. (1994).
Type locality: Germany, Hoyerswerda.
Type substrate: Rosa sp.
Notes: Status unclear. Not treated by any of Wollenweber &
Reinking (1935), Booth (1971), or Gerlach & Nirenberg (1982).
roseobullatum Fusarium Wollenw. (as ‘roseo-bullatum’), Fusaria
Autogr. Delin. 1: 117. 1916.
Basionym: Fusarium bullatum var. roseum Sherb., Mem. Cornell
Univ. Agric. Exp. Sta. 6: 201. 1915.
(See Fusarium equiseti)
Holotypus: ?CUP-007433.
Type locality: USA, Iowa.
Type substrate: Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935).
roseolum Fusarium (H.O. Stephens ex Berk. & Broome) Sacc.,
Syll. Fung. 4: 710. 1886.
Basionym: Fusisporium roseolum H.O. Stephens ex Berk. &
Broome, Ann. Mag. Nat. Hist., ser. 2, 7: 178. 1851.
(See Fusarium merismoides)
Holotypus: ?K(M).
www.studiesinmycology.org
REDELIMITED
Type locality: UK, Bristol.
Type substrate: Decayed Solanum tuberosum.
Note: Synonyms fide Wollenweber & Reinking (1935).
roseum Fusarium Link, Mag. Ges. Naturf. Freunde Berlin 3: 10.
1809, nom. rej.
(See Fusarium sambucinum)
Lectotypus: In B, selected in Gams et al. (1997).
Type locality: Germany.
Type substrate: Malvaceae.
Notes: Gams et al. (1997) proposed that the name, F. roseum be
rejected due to ambiguity surrounding the type of this species,
with F. sambucinum taking preference. This proposal was
accepted in 1999 (see Gams 1999).
rostratum Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 30. 1910 [1913].
(See Fusarium graminearum)
Lectotypus (hic designatus, MBT 10000737): Germany, Berlin,
Triticum aestivum, 1913, O.A. Appel & H.W. Wollenweber, in
Arbeiten Kaiserl. Biol. Anst. Land- Forstw. 8: 30, Abb. 1, figs
E1–E13.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
roumeguerei Fusarium Sacc. (as ‘roumegueri’), Syll. Fung. 4:
702. 1886, nom. illegit., Art. 52.1.
Replaced synonym: Fusarium insidiosum Roum., Michelia 2 (6):
132. 1880.
(See Fusarium lateritium)
Type material: See Fusarium insidiosum.
Note: Synonym fide Wollenweber & Reinking (1935).
ruberrimum Fusarium Delacr., Bull. Soc. Mycol. France 6: 139.
1890.
(See Fusarium avenaceum)
Holotypus: ?PC.
Type locality: France, Paris.
Type substrate: Onobrychis viciifolia.
Note: Synonym fide Wollenweber & Reinking (1935).
rubi Fusarium (G. Winter) Berl. & Voglino, Add. Syll. Fung. 1–4:
391. 1886.
Basionym: Fusisporium rubi G. Winter, in Rabenh., Fungi Eur.
Extraeur Exs., Ed. Nov., Ser. Sec., Cent. 13 (resp. 33): 3280.
1885.
Synonym: Ramularia rubi (G. Winter) Wollenw., Fusaria Autogr.
Delin. 1: 470. 1916.
Cercosporella rubi (G. Winter) Plakidas, J. Agricultural Research
54: 275. 1937.
Syntypes: In BPI, CHRB, CUP, F, HAL, ISC, LSUM, MSC, MU,
NEB & PH (Fungi Eur. Extraeur. Exs. no. 3280).
Type locality: USA, Illinois, Cobden
Type substrate: Rubus villosus
Note: Status unclear fide Braun (1998).
rubicolor Fusarium Berk. & Broome, Trans. Linn. Soc. London,
Bot. 2: 68. 1883.
Holotypus: ?K(M).
Type locality: Australia, Queensland, Brisbane.
Type substrate: Leaves of Eucalyptus sp.
153
CROUS
ET AL.
Note: Not Fusarium fide Wollenweber & Reinking (1935).
rubiginosum Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 108. 1910 [1913].
(See Fusarium culmorum)
Lectotypus (hic designatus, MBT 10000738): Germany, Solanum tuberosum, 1913, O.A. Appel & H.W. Wollenweber, in
Arbeiten Kaiserl. Biol. Anst. Land- Forstw. 8: Tab. I, figs 31–48.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
rubrum Fusarium Parav., Ann. Mycol. 16: 311. 1918.
(See Fusarium lactis)
Lectotypus (hic designatus, MBT 10000739): Germany, core of
Malus domestica fruit, 1918, E. Paravicini, in Ann. Mycol. 16, pl.
4, figs 23–33.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
rusci Fusarium (Sacc.) O'Donnell & Geiser, Phytopathology 103:
404. 2013.
Basionym: Fusarium roseum var. rusci Sacc., Michelia 2: 294.
1881.
Synonyms: Trichofusarium rusci (Sacc.) Bubak, Bull. Herb.
Boissier, ser. 2, 6: 488. 1906.
Pycnofusarium rusci D. Hawksw. & Punith., Trans. Brit. Mycol.
Soc. 61: 63. 1973.
Syntype: BPI 453152.
Type locality: Italy, Selva.
Type substrate: Ruscus aculeatus.
Notes: Examination of the syntype (BPI 453152) revealed that
this species does not belong to the genus Fusarium, having a
myrothecium-like morphology. Also see notes under Nothofusarium devonianum.
russianum Fusarium Manns, Bull. North Dakota Agric. Exp. Sta.
259: 34. 1932.
(See Fusarium acuminatum)
Holotypus: Not located.
Type locality: USA, North Dakota.
Type substrate: Linum usitatissimum.
Note: Synonym fide Wollenweber & Reinking (1935).
ruticola Fusarium Fautrey & Roum. (as ‘rutaecola’), Rev. Mycol.
(Toulouse) 13: 82. 1891.
(See Fusarium avenaceum)
Syntype: ?PC (Fungi Sel. Gall. Exs. No. 5686).
Type locality: France, Noidan.
Type substrate: Ruta graveolens.
Note: Synonym fide Wollenweber & Reinking (1935).
saccardoanum Fusarium P. Syd., Syll. Fung. 14: 1128. 1899.
Replaced synonym: Fusarium sclerodermatis Peck, Rep.
(Annual) Regents Univ. State New York New York State Mus. 43:
77. 1890, nom. illegit., Art. 53.1, non Fusarium sclerodermatis
Oudem. 1889.
(See Fusarium oxysporum)
Holotypus: NYSf2731.
Type locality: USA, New York, Suffolk, Manor, Long Island.
Type substrate: Scleroderma vulgaris.
Note: Synonym fide Wollenweber & Reinking (1935).
154
sacchari Fusarium (E.J. Butler) W. Gams, Cephalosporiumartige Schimmelpilze: 218. 1971.
Basionym: Cephalosporium sacchari E.J. Butler, Mem. Dept.
Agric. India, Bot. Ser. 6: 185. 1913.
Synonyms: Fusarium neoceras Wollenw. & Reinking, Phytopathology 15: 164. 1925.
Gibberella sacchari Summerell & J.F. Leslie, Mycologia 97: 719.
2005, nom. illegit., Art. 53.1, non Gibberella sacchari Speg. 1896.
Fusarium desaboruense N. Maryani et al., Persoonia 43: 59.
2019.
Lectotypus: In Mem. Dept. Agric. India, Bot. Ser. 6: 185, pl. II, figs
1–13. 1913, designated by Yilmaz et al. (2021).
Epitypus: CBS 223.76 (preserved as metabolically inactive culture), designated by Yilmaz et al. (2021).
Ex-epitype culture: BBA 63340 = CBS 223.76 = DAOM
225138 = IMI 202881 = NRRL 13999.
Lectotype and epitype locality: India.
Lectotype and epitype substrate: Saccharum officinarum.
Descriptions and illustrations: See Butler & Khan (1913), Gams
(1971), Gerlach & Nirenberg (1982), Leslie et al. (2005) and
Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: JX171466; rpb2: JX171580;
tef1: AF160278.
salicicola Fusarium Allesch. (as ‘salicicolum’), Ber. Bayer. Bot.
Ges. 4: 39. 1896.
(See Fusarium avenaceum)
Holotypus: In M.
Type locality: Germany, München, forest near Großhesselohe.
Type substrate: Dead branch of Salix caprea.
Note: Synonym fide Wollenweber & Reinking (1935).
salicinum Fusarium Corda, Icon. Fung. 3: 33. 1839.
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, near Prague.
Type substrate: Thin branches of Salix sp.
Notes: Not Fusarium fide Wollenweber & Reinking (1935).
Lectotypification pending study of material lodged in PRM.
salicis Fusarium Fuckel, Fungi Rhen. Exs., Suppl., Fasc. 7, no.
2110. 1868.
(See Fusarium lateritium)
Syntype: S-F267709 (Fungi Rhen. Exs. no. 2110).
Type locality: Germany, Hessen, Münchau, near Hattenheim
Type substrate: Dry branches of Salix triandra.
Notes: Synonym fide Wollenweber & Reinking (1935). Typification pending further study of the syntype lodged in S.
salinense Fusarium Sand.-Den. et al., Persoonia 40: 15. 2017
[2018].
Holotypus: CBS H-23019.
Ex-type culture: CBS 142420 = CPC 26973.
Type locality: Italy, Sicily, Messina, Leni.
Type substrate: Twigs of Citrus sinensis.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018a).
Diagnostic DNA barcodes: rpb1: LT746286; rpb2: LT746306;
tef1: LT746193.
salmonicolor Fusarium Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10:
359. 1868 [1869].
Synonym: Fusidium salmonicolor (Berk. & M.A. Curtis) Wollenw.,
Fusaria Autogr. Delin. 1: 478. 1916.
FUSARIUM
Holotypus: In K(M).
Type locality: Cuba.
Type substrate: Dead twigs of unknown host.
Notes: Synonym fide Wollenweber & Reinking (1935). This taxon
needs to be recombined into the genus Neonectria but requires
further investigation.
samararum Fusarium Allesch., Ber. Bayer. Bot. Ges. 4: 39. 1896.
(See Fusarium lateritium)
Holotypus: In M.
Type locality: Germany, München, Starnberg.
Type substrate: Fallen fruits of Fraxinus excelsior.
Note: Synonym fide Wollenweber & Reinking (1935).
sambucinum Fusarium Fuckel, Fungi Rhen. Exs., Fasc. 3, no.
211. 1863, nom. cons.
Synonyms: Fusarium roseum Link, Mag. Ges. Naturf. Freunde
Berlin 3: 10. 1809, nom. rej.
Fusidium roseum (Link) Link, Mag. Ges. Naturf. Freunde Berlin
8: 31. 1815 [1816].
Gibberella rosea (Link) W.C. Snyder & H.N. Hansen, Amer. J.
Bot. 32: 664. 1945.
Sphaeria pulicaris Fr., Mykol. Hefte 2: 37. 1823.
Gibbera pulicaris (Fr.) Fr., Summa Veg. Scand. 2: 402. 1849.
Botryosphaeria pulicaris (Fr.) Ces. & De Not., Comment. Soc.
Crittog. Ital. 1: 212. 1863.
Nectria pulicaris (Fr.) Tul. & C. Tul., Select. Fung. Carpol. 3: 63.
1865.
Cucurbitaria pulicaris (Fr.) Quel., Mem. Soc. Emul.
Montbeliard,
ser. 2, 5: 511. 1875.
Gibberella pulicaris (Fr.) Sacc., Michelia 1: 43. 1877.
Fusarium sulphureum Schltdl., Fl. Berol. 2: 139. 1824, nom. rej.
Fusidium sulphureum (Schltdl.) Link, in Willdenow, Sp. Pl. ed. 4,
6: 98. 1825.
Fusarium discolor var. sulphureum (Schltdl.) Appel & Wollenw.,
Arbeiten Kaiserl. Biol. Anst. Land- Forstw. 8: 115. 1910 [1913].
Sphaeria cyanogena Desm., Ann. Sci. Nat., Bot., ser. 3, 10: 352.
1848.
Botryosphaeria cyanogena (Desm.) Niessl, Verh. Naturf. Vereins
Brünn 10: 197. 1872.
Gibberella cyanogena (Desm.) Sacc., Syll. Fung. 2: 555. 1883.
Calonectria cyanogena (Desm.) Lar.N. Vassiljeva, Nizshie Rasteniya, Griby i Mokhoobraznye
Dalnego Vostoka Rossii, Griby. Tom 4. Pirenomitsety i
Lokuloaskomitsety: 169. 1998.
Fusarium maydis Kalchbr., Math. Term. Közlem. 3: 285. 1865,
nom. rej.
Fusisporium ricini Berenger, Mem. Accad. Agric. Verona 44: 257.
1866, nom. rej.
Fusarium ricini (Berenger) Bizz., Fl. Ven. Critt. 1: 539. 1885.
Fusarium subcarneum P. Crouan & H. Crouan, Fl. Finistere: 14.
1867, nom. rej.
Fusarium violaceum P. Crouan & H. Crouan, Fl. Finistere: 14.
1867, nom. illegit., Art. 53.1.
Fusisporium pezizoideum Berk. & M.A. Curtis, Grevillea 3: 147.
1875.
Fusarium pezizoideum (Berk. & M.A. Curtis) Sacc., Syll. Fung. 4:
711. 1886.
Fusisporium pulvinatum Berk. & Broome, J. Linn. Soc., Bot. 14:
102. 1873 [1875].
Fusarium pulvinatum (Berk. & Broome) Sacc., Syll. Fung. 4: 699.
1886, nom. illegit., Art. 53.1.
Fusarium roseum var. buxi Sacc., Michelia 2: 294. 1881.
www.studiesinmycology.org
REDELIMITED
Fusarium roseum var. calystegiae Sacc., Michelia 2: 294. 1881.
Fusarium roseum var. cucubali-bacciferi Sacc., Michelia 2: 295.
1881.
Fusarium roseum var. dulcamarae Sacc., Michelia 2: 295. 1881.
Fusarium roseum var. filicis Sacc., Michelia 2: 295. 1881.
Fusarium roseum var. fraxini Therry, Cryptog. Lyonn.: 5717.
1881.
Fusarium roseum var. helianti Sacc., Michelia 2: 295. 1881.
Fusarium roseum var. maydis Sacc., Michelia 2: 295. 1881.
Fusarium roseum var. phytolaccae Sacc., Michelia 2: 294. 1881.
Fusarium roseum var. rosae Sacc., Michelia 2: 295. 1881.
Fusarium roseum var. vitalbae Sacc., Michelia 2: 294. 1881.
Fusarium granulare Kalchbr., Crypt. Austro-Afric., no. 1068.
1874.
Fusarium roseum var. dracaenae Roum., Fungi Sel. Gall. Exs.,
Cent. 19: 1869. 1882.
Fusisporium tenuissimum Peck, Rep. (Annual) New York State
Mus. Nat. Hist. 34: 48. 1883.
Fusarium tenuissimum (Peck) Sacc., Syll. Fung. 4: 711. 1886.
Fusisporium hordei Wm.G. Sm., Diseases of field and garden
crops, chiefly as are caused by fungi: 212. 1884.
Fusarium hordei (Wm.G. Sm.) Sacc., Syll. Fung. 11: 652. 1895.
Gibberella pulicaris f. robiniae P. Syd., Mycoth. March., Cent. 14:
1544. 1887.
Fusarium tenellum Sacc. & Briard, Rev. Mycol. (Toulouse) 7:
212. 1885.
Fusarium asparagi Delacr., Bull. Soc. Mycol. France 6: 99. 1890,
nom. illegit., Art. 53.1.
Fusarium delacroixii Sacc., Syll. Fung. 10: 725. 1892.
Fusarium fraxini Allesch., Ber. Bot. Vereines Landshut 12: 130.
1892.
Fusarium polymorphum Matr., Rech. Devel. Muced.: 84. 1892.
Fusarium roseum var. lonicerae Allesch., Ber. Bayer. Bot. Ges. 5:
22. 1897.
Fusarium roseum f. visci Brunaud, Actes Soc. Linn. Bordeaux
52: 149. 1897.
Fusarium pannosum Massee, Bull. Misc. Inform. Kew 1898: 117.
1898.
Gibberella pulicaris var. subtropica Rehm, in Theissen, Ann.
Mycol. 9: 63. 1911.
Gibberella subtropica (Rehm) Wollenw., Fusaria Autogr. Delin. 1:
38. 1916.
Botryosphaeria subtropica (Rehm) Weese, Sitzungsber. Akad.
Wiss. Wien, Math.-Naturwiss. Cl., Abt. 1, 128: 708. 1919.
Fusarium genevense Dasz., Bull. Soc. Bot. Geneve, ser. 2, 4:
305. 1912.
Fusarium discolor Appel & Wollenw., Arbeiten Kaiserl. Biol. Anst.
Land- Forstw. 8: 114. 1913.
Fusarium subpallidum Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 230. 1915.
Fusarium roseum var. phaseoli Gonz. Frag., Trab. Mus. Nac.
Cienc. Nat., Ser. Bot. 10: 173. 1916.
Fusarium aridum O.A. Pratt, J. Agric. Res. 13: 89. 1918.
Fusarium elongatum O.A. Pratt, J. Agric. Res. 13: 84. 1918, nom.
illegit., Art. 53.1.
Fusarium roseum var. zeae Cif., Bull. Soc. Bot. Ital. 1921: 73. 1921.
Fusarium sambucinum var. medium Wollenw., Z. Parasitenk.
(Berlin) 3: 358. 1931.
Fusarium sambucinum f2 Wollenw., Z. Parasitenk. (Berlin) 3:
357. 1931.
Fusarium sambucinum f3 Wollenw., Z. Parasitenk. (Berlin) 3:
357. 1931.
155
CROUS
ET AL.
Fusarium sambucinum f4 Wollenw., Z. Parasitenk. (Berlin) 3:
357. 1931.
Fusarium sambucinum f6 Wollenw., Z. Parasitenk. (Berlin) 3:
358. 1931.
Gibberella pulicaris var. minor Wollenw., Z. Parasitenk. (Berlin) 3:
356. 1931.
Fusarium roseum f. phaseoli N. Barros, Revista Inst. Colomb.
Agropecu. 1: 80. 1966.
Fusarium roseum f. compactum Tivoli, Agronomie 8: 220. 1988,
nom. inval., Arts. 35.1, 39.1.
Fusarium roseum var. lavaterae-arboreae Thüm., Mycoth. Univ.
Cent. 11: no. 1084. 1878.
Lectotypus: G00266369.
Type locality: Germany, Hessen.
Type substrate: Dead branches of Sambucus nigra.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Gerlach & Nirenberg (1982), Nelson
et al. (1983), and Leslie & Summerell (2006).
Notes: The taxonomy of F. sambucinum, the type species of the
genus Fusarium, is confusing. Divergent species concepts have
been derived from multiple taxonomic systems and the conflicting application of the older name F. roseum (Gams et al.
1997, Leslie & Summerell 2006). After examination of the type
material, a proposal to conserve F. sambucinum against several
earlier names was presented (Gams et al. 1997) and unanimously accepted by the committee for fungal taxonomy (Gams
1999). Further older valid synonymous names are in need to
be rejected, notably Sphaeria pulicaris and Sphaeria cyanogena.
samoense Fusarium Gehrm., Arbeiten Kaiserl. Biol. Anst. LandForstw. 9: 24. 1913.
(See Fusarium verticillioides)
Lectotypus (hic designatus, MBT 10000740): Samoa, cortex of
Theobroma cacao, 1913, K. Gehrmann, in Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 9: Abb. 6, figs 1–3.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
sampaioi Fusarium Gonz. Frag., Bol. Soc. Brot. 2: 50. 1924.
Synonym: Illosporium corallinum Roberge, in Desmazieres, Pl.
Crypt. N. France, ed. 1, Fasc. 32: no. 1551. 1847 (pr. p. fide
Hawksworth 1979).
Marchandiomyces corallinus (Roberge) Diederich & D. Hawksw.,
Mycotaxon 37: 312. 1990 (pr. p. fide Diederich 1990).
Aegerita physciae Vouaux, Bull. Trimestriel Soc. Mycol. France
30: 314. 1914.
Holotypus: Not indicated. Several syntypes fide Hawksworth
(1979).
Type locality: Portugal, near Gaia, Alto da Bandeira; and near
Tabuaço.
Type substrate: Lichen thallus (on Lasallia pustulata, Parmelia
saxatilis, P. soredians and P. exasperata; Physcia semipinnata,
P. tenella, Phaeophyscia orbicularis and Physconia grisea).
Notes: Hawksworth (1979), after examination of a syntype,
concluded that the Fusarium name should be rejected since the
studied material was based on discordant elements. Nevertheless, examination of all available syntypes is required to confirm
these observations or otherwise, to fix the use of this name by
lectotypification.
samuelsii Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 4. 2020.
156
Neocosmospora samuelsii Sand.-Den. & Crous, Persoonia 43:
165. 2019.
Holotypus: CBS H-24001.
Ex-type culture: CBS 114067 = G.J.S. 89-70.
Type locality: Guyana, Mount Wokomung, on ridge leading NW
toward summit, 0.5–1 h walk from Base Camp.
Type substrate: Bark.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834252; rpb2: LR583874;
tef1: LR583644.
sangayamense Fusarium Maryani et al., Stud. Mycol. 92: 187.
2018 [2019].
Holotypus: InaCC F960 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F960.
Type locality: Indonesia, South Kalimantan, Kota Baru,
Sengayam.
Type substrate: Pseudostem of Musa var. Pisang Kepok.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479537; rpb2: LS479283;
tef1: LS479732.
sanguineum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 193. 1915.
(See Fusarium acuminatum)
Typus: ?CUP-007444.
Type locality: USA, New York, Ithaca
Type substrate: Solanum tuberosum.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in CUP.
sapindophilum Fusarium Speg., Anales Mus. Nac. Hist. Nat.
Buenos Aires 6: 351. 1898 [1899].
Synonym: Cercoseptoria sapindophila (Speg.) Cif., Mycopathol.
Mycol. Appl. 6: 26. 1951.
Holotypus: In LPS (Fungi Argent. n.v.c. no. 868).
Type locality: Argentina, near Tucuman.
Type substrate: Living leaves of unknown climbing Sapindaceae.
Note: Synonym fide Wollenweber & Reinking (1935).
sarcochroum Fusarium (Desm.) Sacc., Michelia 1: 534. 1879.
Basionym: Selenosporium sarcochroum Desm., Ann. Sci. Nat.,
Bot., ser. 3, 14: 112. 1850.
Synonyms: Fusarium diplosporum Cooke & Ellis, Grevillea 7: 38.
1878.
Fusarium desciscens Oudem., Ned. Kruidk. Arch., ser. 2, 5: 515.
1889.
Fusarium robiniae Pass., Atti Reale Accad. Lincei, Rendiconti Cl.
Sci. Fis., ser. 4, 7: 51. 1891.
Fusarium sarcochroum var. robiniae (Pass.) Wollenw., Z. Parasitenk. (Berlin) 3: 388. 1931.Fusarium sarcochroum f. polygalaemyrtifoliae Henn., Verh. Bot. Vereins Prov. Brandenburg 40: 174.
1898 [1899].
Fusarium sarcochroum var. casei Loubiere, Rech. Mucedinees:
53. 1924.
Gibberella pseudopulicaris Wollenw., Z. Parasitenk. (Berlin) 3:
387. 1931.
Neotypus (hic designatus, MBT 10000741): Switzerland, Viscum album, 1977, W. Gerlach, CBS 745.79 (preserved as
metabolically inactive culture).
Ex-neotype culture: BBA 63714 = CBS 745.79 = NRRL 20472.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Raillo (1950), Bilaĭ (1955), Gerlach & Nirenberg (1982).
FUSARIUM
Diagnostic DNA barcodes: rpb1: JX171472; rpb2: JX171586;
tef1: JABEXW010000634.
Notes: No type material could be located. Therefore, CBS 745.79
is designated as neotype here. Both Gerlach & Nirenberg (1982)
and O'Donnell et al. (2013) considered this isolate an authentic
representation of this species.
schawrowii Fusarium Speschnew (as ‘schawrovi‘), Arbeiten
Kaukas. Stat. Seidenzucht 10: 1906.
(See Fusarium lateritium)
Holotypus: Not located.
Type locality: Turkey, Anatolia.
Type substrate: Branch of Morus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
schiedermayeri Fusarium (Thüm.) Sacc., Syll. Fung. 4: 712.
1886.
Basionym: Fusisporium schiedermayeri Thüm., Fungi Austr. Exs.
Cent. 1: no. 78. 1871.
(See Fusarium avenaceum)
Syntypus: In HAL.
Type locality: Austria, Linz.
Type substrate: Ovaries of Luzula pilosa, in association with
Ustilago luzulae.
Note: Synonyms fide Wollenweber & Reinking (1935).
schnablianum Fusarium Allesch., Hedwigia 34: 289. 1895.
(See Fusarium avenaceum)
Holotypus: In M.
Type locality: Germany, Großhesselohe, near München.
Type substrate: Decorticated branch of Acer pseudoplatanus.
Note: Synonym fide Wollenweber & Reinking (1935).
schribauxii Fusarium Delacr., Bull. Soc. Mycol. France 6: 99.
1890.
(See Fusarium culmorum)
Holotypus: ?PC.
Type locality: France.
Type substrate: Seeds of Triticum sativum, in association with
Trichothecium roseum.
Note: Synonym fide Wollenweber & Reinking (1935).
schweinitzii Fusarium Ellis & Harkn., Bull. Torrey Bot. Club 8: 27.
1881.
Colletotrichum crassipes (Speg.) Arx, Verh. Kon. Akad.
Wetensch., Afd. Natuurk., Sect. 2, 51: 77. 1957.
Basionym: Gloeosporium crassipes Speg., Rivista Vitic. Enol. 2:
405. 1878.
Syntypes: In CHRB, CUP, ILL, MICH, MU, NEB, NYS, PH, PUL &
WIS (Ellis, N. Amer. Fungi no. 539).
Type locality: USA, New Jersey, Newfield.
Type substrate: Vitis sp. vine
Note: Synonym fide Wollenweber & Reinking (1935).
scirpi Fusarium Lambotte & Fautrey, Fautrey, Fungi Sel. Gall.
Exs. no. 6540. 1893.
Synonyms: ?Fusoma helminthosporii Corda, Icon. Fung. 1: 7.
1837.
?Fusoma filiferum Preuss, Linnaea 25: 73. 1852.
?Fusarium filiferum (Preuss) Wollenw., Fusaria Autogr. Delin. 1:
220. 1916.
?Fusarium scirpi var. filiferum (Preuss) Wollenw., Fusaria Autogr.
Delin. 3: 936. 1930.
?Fusisporium chenopodinum Thüm., Mycoth. Univ., Cent. 14: no.
1378. 1879.
www.studiesinmycology.org
REDELIMITED
?Fusarium chenopodinum (Thüm.) Sacc., Syll. Fung. 4: 701.
1886.
?Fusarium aloes Kalchbr. & Cooke (as ‘alo€es’), Grevillea 9: 23.
1880.
?Fusarium osteophilum Speg., Anales Soc. Ci. Argent. 10: 60. 1880.
?Fusisporium mucophytum W.G. Sm., Gard. Chron. n.s. 22: 245.
1884.
?Fusarium mucophytum (W.G. Sm.) Massee, Brit. Fung.-Fl. 3:
483. 1893.
Fusarium equiseticola Allesch., Hedwigia 34: 289. 1895.
Fusarium sclerotium Wollenw., Ber. Deutsch. Bot. Ges. 31: 301.
1913.
Fusarium caudatum Wollenw., J. Agric. Res. 2: 262. 1914.
Fusarium sclerodermatis var. lycoperdonis Picb., Bull. Ecol. Sup.
Agron., Brno, R.C.S. Fac. Silvicult. 13: 27. 1929.
Fusarium scirpi var. comma Wollenw., Fusaria Autogr. Delin. 3:
922. 1930.
Fusarium scirpi var. nigrantum F.T. Benn. (as ‘nigrans’), Ann.
Appl. Biol. 19: 26. 1932.
Fusarium scirpi var. pallens F.T. Benn., Ann. Appl. Biol. 19: 21.
1932.
Lectotypus (hic designatus, MBT 10000742): France, Schoenoplectus lacustris (= Scirpus lacustris), 1893, F. Fautrey,
ILL00220730 (Fautrey, Fungi Sel. Gall. Exs. No. 6540).
Epitypus (hic designatus, MBT 10000743): Australia, New
South Wales, near Broken Hill, pasture soil, 1981, P.E. Nelson,
CBS H-24069.
Ex-epitype culture: CBS 447.84 = FRC R-6252 = NRRL 36478.
Descriptions and illustrations: See Wollenweber (1916–1935),
Wollenweber & Reinking (1935), Burgess et al. (1985) and Leslie
& Summerell (2006).
Diagnostic DNA barcodes: rpb2: GQ505832; tef1: GQ505654.
Notes: The epitypification of Fusarium scirpi by Xia et al. (2019)
was not Code compliant as the holo- or lectotype was not
correctly indicated (Art. 9.9). Here, the lectotype is clearly indicated, making the epitypification valid.
sclerodermatis Fusarium Oudem., Ned. Kruidk. Arch., ser. 2, 5:
516. 1889.
(See Fusarium torulosum)
Holotypus: ?L.
Type
locality:
Netherlands,
Zuid-Holland
Province,
Scheveningen.
Type substrate: Rotten Scleroderma vulgaris.
Note: Synonym fide Nirenberg (1995).
sclerodermatis Fusarium Peck, Rep. (Annual) Regents Univ.
State New York New York State Mus. 43: 77. 1890, nom. illegit.,
Art. 53.1.
(See Fusarium oxysporum)
Authentic material: NYSf2731.
Original locality: USA, New York, Suffolk.
Original substrate: Peridium of Scleroderma vulgaris.
Notes: A later homonym of F. sclerodermatis Oudem. Saccardo
(1892) published F. peckii as a replacement name which was
again an illegitimate homonym; the taxon was later synonymised
with F. oxysporum var. aurantiacum (Wollenweber & Reinking
1935).
sclerostromaton Fusarium Sideris, Phytopathology 14: 213.
1924.
(See Fusarium oxysporum)
Holotypus: Not located.
157
CROUS
ET AL.
Type locality: USA, California, Delta, near Stockton.
Type substrate: Roots of Allium sp. with symptoms of pink root
disease.
Note: Synonym fide Wollenweber & Reinking (1935).
sclerotioides Fusarium Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. 6: 214. 1915.
(See Fusarium oxysporum)
Typus: ?BPI 452971.
Type locality: USA, New York, Ithaca.
Type substrate: Solanum tuberosum.
Notes: Synonym fide Wollenweber & Reinking (1935). Typification pending further study of the specimen lodged in BPI.
sclerotioides Fusarium (Höhn.) Samuels & Rossman, Mycological Papers 164: 23. 1991, nom. illegit., Art. 53.1
Basionym: Stagonopsis sclerotioides Höhn., in Penther &
Zederbauer, Ann. K. K. Naturhist. Hofmus. 20: 368. 1905.
(See Fusarium kurdicum)
Holotypus: FH00965353.
Type locality: Turkey, near Erciyes Dagı.
Type substrate: Thin twigs of Astragalus sp.
sclerotium Fusarium Wollenw., Ber. Deutsch. Bot. Ges. 31: 30.
1913.
(See Fusarium scirpi)
Holotypus: Not located.
Type locality: USA.
Type substrate: Citrullus vulgaris and Lycopersicon esculentum.
Note: Synonym fide Nirenberg (1995).
scolecoides Fusarium Sacc. & Ellis, Miscellanea Mycologia 2:
18. 1885.
(See Fusarium ciliatum)
Holotypus: In PAD.
Type locality: USA, Pennsylvania, Bethlehem.
Type substrate: Branch of Robinia sp.
Note: Synonym fide Wollenweber & Reinking (1935).
secalis Fusarium Fee, Mem. Soc. Mus. Hist. Nat. Strassbourg 3:
35. 1843.
(See Fusarium heterosporum)
Holotypus: Not located.
Type locality: France.
Type substrate: Spikes of Secale cereale.
Note: Synonym fide Wollenweber & Reinking (1935).
secalis Fusarium Jacz., Bull. Trimestriel Soc. Mycol. France 28:
346. 1912, nom. illegit., Art. 53.1.
(See Fusarium nivale)
Authentic material: Not located.
Original locality: Russia, near Moscow.
Original substrate: Grain of Secale sp.
Note: Synonym fide Wollenweber & Reinking (1935).
secorum Fusarium Secor et al., Fungal Biology 118: 767. 2014.
Holotypus: BPI 892692.
Ex-type culture: NRRL 62593.
Type locality: USA, Minnesota, Sabin.
Type substrate: Root of Beta vulgaris.
Descriptions and illustrations: See Secor et al. (2014).
Diagnostic DNA barcodes: rpb1: JABEEM010001657; rpb2:
JABEEM010001483; tef1: KJ189225.
sedimenticola Fusarium M.M. Wang et al., Botanica Marina 63:
174. 2020.
158
(See Fusarium keratoplasticum)
Holotypus: HAMS 248044.
Ex-type culture: CGMCC 3.19499 = LC12845.
Type locality: China, South-West Indian Ocean.
Type substrate: Deep-sea sediments.
Descriptions and illustrations: See Jones et al. (2020).
Diagnostic DNA barcodes: rpb2: MK190729; tef1: MK190727.
Notes: Fusarium sedimenticola was recently introduced by Jones
et al. (2020) in the FSSC (=Neocosmospora) isolated from deepsea sediment in the Indian Ocean. However, based on comparisons of both protologues and sequences using a larger
sampling of N. keratoplastica isolates (results not shown), we
consider F. sedimenticola a synonym under N. keratoplastica.
seemenianum Fusarium Henn., in Seemen, Allg. Bot. Z. Syst. 2:
83. 1896.
(See Fusarium avenaceum)
Holotypus: B 70 0100194.
Type locality: Germany, Borkum.
Type substrate: Leaves of Platanthera bifolia var. robusta.
Note: Synonym fide Wollenweber & Reinking (1935).
semitectum Fusarium Berk. & Ravenel, Grevillea 3: 98. 1875.
(See Fusarium incarnatum)
Holotypus: ?K(M).
Type locality: USA, Pennsylvania, Philadelphia.
Type substrate: Petioles of Musa sp.
serjaniae Fusarium Syd. & P. Syd., Beibl. Hedwigia 40: (2). 1901.
Synonym: Cercospora serjaniae (Syd. & P. Syd.) Wollenw., Z.
Parasitenk. (Berlin) 3: 496. 1931.
Holotypus: S-F45658.
Type locality: Mexico, Puebla, Tehuacan.
Type substrate: Leaves of Serjania racemosa.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935) and not Cercospora fide Crous & Braun (2003).
serpentinum Fusarium J.W. Xia et al., Persoonia 43: 217.
2019.
Holotypus: CBS H-24070.
Ex-type culture: BBA 62209 = CBS 119880 = MRC 1813.
Type locality: Unknown.
Type substrate: Unknown.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170432; tef1: MN170499.
setosum Fusarium Nirenberg & Samuels, Canad. J. Bot. 67:
3372. 1989.
Setofusarium setosum (Samuels & Nirenberg) Sand.-Den. &
Crous, Stud. Mycol. 98 (no. 100116): 75. 2021.
Synonym: Nectria setofusarii Samuels & Nirenberg (as ‘setofusariae’), Canad. J. Bot. 67: 3372. 1989.
Holotypus: NY00927992.
Type locality: French Guiana, Piste de Saint-Elie, Km 16 on
road between Sinnamary and St. Elie, ORSTOM research area,
"ECEREX".
Type substrate: Bark of living liana.
Epitypus: CBS H-24723, designated in this study.
Ex-epitype culture: CBS 635.92 = G.J.S. 88-12 = NRRL 36526.
Epitype locality: French Guiana, Cayenne, 15 km from Remise,
trail to Vidal-old farm, secondary forest.
Epitype substrate: Bark.
Descriptions and illustrations: See Samuels & Nirenberg (1989).
Diagnostic DNA barcodes: rpb1: JX171539; rpb2: JX171651;
tef1: MW834294.
FUSARIUM
sibiricum Fusarium Gagkaeva et al., Int. J. Food Microbiol. 147:
64. 2011.
Holotypus: LEP 12652.
Ex-type culture: MFG 11013 = NRRL 53430.
Type locality: Russia, Khabarovsk.
Type substrate: Grain of Avena sativa.
Descriptions and illustrations: See Yli-Mattila et al. (2011).
Diagnostic DNA barcodes: rpb1: MW233302; rpb2: HQ154472;
tef1: HM744684.
siculi Fusarium Sand.-Den. et al., Persoonia 40: 17. 2017
[2018].
Holotypus: CBS H-23021.
Ex-type culture: CBS 142422 = CPC 27188.
Type locality: Italy, Sicily, Catania, Paterno.
Type substrate: Citrus sinensis.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018a).
Diagnostic DNA barcodes: rpb1: LT746299; rpb2: LT746327;
tef1: LT746214.
silvicola Fusarium (Sand.-Den. & Crous) O'Donnell et al., Index
Fungorum 440: 4. 2020.
Neocosmospora silvicola Sand.-Den. & Crous, Persoonia 43:
167. 2019.
Synonyms: Fusarium solani f. robiniae Matuo & Y. Sakurai, Ann.
Phytophathol. Soc. Japan 30: 35. 1965.
Hypomyces solani f. robiniae Matuo & Y. Sakurai, Ann. Phytophathol. Soc. Japan 30: 35. 1965.
Nectria solani f. robiniae (Matuo & Y. Sakurai) G.R.W. Arnold, Z.
Pilzk. 37: 193. 1972.
Holotypus: CBS H-24002.
Ex-type culture: CBS 123846 = G.J.S. 04-147.
Type locality: USA, Tennessee, Great Smoky Mountains National
Park.
Type substrate: Fallen trunk of Liriodendron tulipifera.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834254; rpb2: LR583876;
tef1: LR583646.
sinense Fusarium Z.H. Zhao & G.Z. Lu (as ‘sinensis’), Mycologia 100: 747. 2008.
Holotypus: IBE 000007.
Ex-type culture: CBS 122710.
Type locality: China, Shandong Province, Jinan.
Type substrate: Seed of Triticum aestivum.
Descriptions and illustrations: See Zhao & Lu (2008).
Diagnostic DNA barcode: tef1: EF531235.
socium Fusarium Sacc., Atti Ist. Veneto Sci. Lett. Arti, ser. 6, 2:
450. 1884.
(See Fusarium expansum)
Holotypus: Not located.
Type locality: France, Troyes.
Type substrate: Cortex of Carpinus sp. in association with Stilbospora sp. and Nectria stilbosporae.
Note: Synonym fide Wollenweber & Reinking (1935).
solani Fusarium (Mart.) Sacc., Michelia 2: 296. 1881.
Neocosmospora solani (Mart.) L. Lombard & Crous, Stud.
Mycol. 80: 228. 2015.
Basionym: Fusisporium solani Mart., Die Kartoffel-Epidemie der
letzten Jahre oder die Stockf€aule und R€aude der Kartoffeln: 20.
1842.
www.studiesinmycology.org
REDELIMITED
Synonyms: Fusisporium solani-tuberosi Desm., Ann. Sci. Nat.,
Bot., ser. 3, 3: 359. 1845.
Fusarium solani-tuberosi (Desm.) Sacc., Syll. Fung. 4: 189.
1886.
Pionnotes solani-tuberosi (Desm.) Sacc., Syll. Fung. 4: 727.
1886.
Fusisporium rhizophilum var. solani-tuberosi (Desm.) Westend.,
Bull. Acad. Roy. Sci. Belgique, Cl. Sci. 18(2): 413. 1852.
Fusisporium candidum Bonord., Handb. Allg. Mykol.: 96. 1851,
nom. illegit., Art. 53.1, non Fusarium candidum (Link) Sacc. 1886
Fusarium commutatum Sacc., Syll. Fung. 4: 710. 1886.
Fusarium allii-sativi Allesch., Ber. Bot. Vereines Landshut 12:
131. 1892.
Hymenula affinis (Fautrey & Lambotte) Wollenw., Fusaria Autogr.
Delin. 1: 484. 1916 [pr. p. fide Booth (1971)].
Pionnotes viridis Lechmere, Compt. Rend. Hebd. Seances Acad.
Sci. 155: 178. 1912.
Fusarium viride (Lechmere) Wollenw., Fusaria Autogr. Delin. 1:
418. 1916.
Fusarium radicicola Wollenw., J. Agric. Res. 2: 257. 1914.
Fusarium javanicum var. radicicola (Wollenw.) Wollenw., Z.
Parasitenk. (Berlin) 3: 286. 1931.
Fusarium solani f. radicicola (Wollenw.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 28: 740. 1941.
Fusarium eumartii C.W. Carp., J. Agric. Res. 5: 204. 1915.
Fusarium solani var. eumartii (C.W. Carp.) Wollenw., Z. Parasitenk. (Berlin) 3: 452. 1931.
Fusarium solani f. eumartii (C.W. Carp.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 28: 740. 1941.
Fusarium malli Taubenh., Bull. Texas Agric. Exp. Sta. 273: 25. 1921.
Fusarium alluviale Wollenw. & Reinking, Phytopathology 15: 167.
1925.
Fusarium aduncisporum Weimer & Harter, J. Agric. Res. 32: 312.
1926.
Fusarium solani var. aduncisporum (Weimer & Harter) Wollenw.,
Fusaria Autogr. Delin. 3: 1035. 1930.
Neocosmospora rubicola L. Lombard & Crous, Stud. Mycol. 80:
227. 2015.
Lectotypus: Illustration tab. III, fig. 29 in von Martius (1842),
designated in Schroers et al. (2016).
Epitypus: CBS H-22335, designated in Schroers et al. (2016).
Ex-epitype culture: CBS 140079 = FRC S-2364 = NRRL 66304.
Epitype locality: Slovenia, Doljenska, Radohova.
Epitype substrate: Rotten tuber of Solanum tuberosum.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Leslie & Summerell (2006), and Schroers et al. (2016).
Diagnostic DNA barcodes: rpb1: MW218134; rpb2: KT313623;
tef1: KT313611.
solani-melongenae Fusarium O'Donnell et al., Index Fungorum
440: 4. 2020.
Neocosmospora ipomoeae (Halst.) L. Lombard & Crous, Stud.
Mycol. 80: 227. 2015.
Basionym: Nectria ipomoeae Halst., Rep. (Annual) New Jersey
Agric. Exp. Sta. 12: 281. 1891.
Synonyms: Cucurbitaria ipomoeae (Halst.) Kuntze, Revis. Gen.
Pl. 3: 461. 1898.
Creonectria ipomoeae (Halst.) Seaver, N. Amer. Fl. 3: 22. 1910.
Hypomyces ipomoeae (Halst.) Wollenw., Phytopathology 3: 34.
1913.
Haematonectria ipomoeae (Halst.) Samuels & Nirenberg, Stud.
Mycol. 42: 136. 1999.
159
CROUS
ET AL.
Nectria ipomoeae f. ipomoeae Halst., Rep. (Annual) New Jersey
Agric. Exp. Sta. 12: 281. 1891.
Nectria ipomoeae var. ipomoeae Halst., Rep. (Annual) New
Jersey Agric. Exp. Sta. 12: 281. 1891.
Hypomyces ipomoeae var. ipomoeae (Halst.) Wollenw., Phytopathology 3: 34. 1913.
Hypomyces ipomoeae var. major Wollenw., Fusaria Autogr.
Delin. 3: 826. 1930.
?Fusarium striatum Sherb., Cornell Univ. Agric. Exp. Sta. Mem.
6: 255. 1915
?Fusarium solani var. striatum (Sherb.) Wollenw., Z. Parasitenk.
(Berlin) 3: 451. 1931.
Holotypus: BPI 552416.
Type locality: USA, New Jersey, Mickleton.
Type substrate: Solanum melongena.
Note: This species requires epitypification from the type locality
and host.
solani-tuberosi Fusarium (Desm.) Sacc., Syll. Fung. 4: 189.
1886.
Basionym: Fusisporium solani-tuberosi Desm., Ann. Sci. Nat.,
Bot., ser. 3, 3: 359. 1845.
(See Fusarium solani)
Holotypus: ?PC.
Type locality: France.
Type substrate: Rotten tuber of Solanum tuberosum.
Note: Synonyms fide Wollenweber & Reinking (1935).
sophorae Fusarium Allesch., Beibl. Hedwigia 36: (164). 1897.
(See Fusarium lateritium)
Holotypus: In M.
Type locality: Germany, Berlin, Sp€ath'sche Baumschulen.
Type substrate: Sophora japonica.
Note: Synonym fide Wollenweber & Reinking (1935).
sorghi Fusarium Henn., Ann. Mus. Congo Belge, Bot., ser. 5, 2:
105. 1907.
(See Fusarium avenaceum)
Syntype: Vanderyst 171 in B fide Hein (1988).
Type locality: Democratic Republic of the Congo, Kisantu.
Type substrate: Spikelet of Sorghum vulgare (= Sorghum
bicolor).
Note: Synonym fide Wollenweber & Reinking (1935).
sororula Fusarium Herron et al., Stud. Mycol. 80: 146. 2015.
Holotypus: PREM 60903.
Ex-type culture: CBS 137242 = CMW 40578.
Type locality: Colombia, Risaralda, Angela Maria (Santa Rosa).
Type substrate: Stem cankers of Pinus patula.
Descriptions and illustrations: See Herron et al. (2015).
Notes: Comparisons of recently generated sequences from the
living ex-type culture (CBS 137242 = CMW 40578) of F. sororula
indicate a strain transposition or contamination by another
Fusarium species. Therefore, this species needs to be recollected from the type locality and substrate or sequences need to
be generated from the holotype specimen.
spartinae Fusarium Ellis & Everh., J. Mycol. 8: 14. 1902.
Septogloeum spartinae (Ellis & Everh.) Wollenw. & Reinking,
Fusarien: 336. 1935.
Holotypus: NY (fide Index Fungorum).
Type locality: USA, California, Pacific Grove.
Type substrate: Leaves of Spartina stricta.
Note: Synonym fide Wollenweber & Reinking (1935).
160
spartum Fusarium S. Gargouri et al., Mycologia 112: 799. 2020.
Holotypus: BPI 911207.
Ex-type culture: NRRL 66896.
Type locality: Tunisia, Kasserine Governorate.
Type substrate: Rhizosphere of Macrochloa tenacissima.
Descriptions and illustrations: See Gargouri et al. (2020).
Diagnostic DNA barcodes: rpb1: MT409439; rpb2: MT409449;
tef1: MT409459.
spathulatum Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 4. 2020.
Neocosmospora spathulata Sand.-Den. & Crous, Persoonia
43: 171. 2019.
Holotypus: CBS H-24003.
Ex-type culture: CBS 145474 = NRRL 28541 = UTHSC 98-1305.
Type locality: USA, New England.
Type substrate: Synovial fluid from Homo sapiens.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218137; rpb2: EU329542;
tef1: DQ246882.
speiranthae Fusarium Henn. (as ‘speiranthis’), Verh. Bot. Vereins
Prov. Brandenburg 40: 174. 1898.
Colletotrichum dematium (Pers.) Grove, J. Bot. 56: 341. 1918.
Basionym: Sphaeria dematium Pers., Syn. Meth. Fung.: 88.
1801.
Synonyms: Exosporium dematium (Pers.) Link, in Willdenow, Sp.
pl., Ed. 4, 6: 122. 1825.
Vermicularia dematium (Pers.) Fr., Syst. Mycol. 3: 255. 1829.
Lasiella dematium (Pers.) Quel., Mem. Soc. Emul.
Montbeliard
ser. 2, 5: 518. 1875.
Gloeosporium speiranthae (Henn.) Wollenw., Fusaria Autogr.
Delin. 1: 500. 1916.
Holotypus: In B fide Hein (1988).
Type locality: Germany, Berlin, botanical garden.
Type substrate: Leaves of Speirantha convallarioides.
Notes: Wollenweber (1916) studied and illustrated authentic
material of this species, recombining it in Gloeosporium. The
shape of the conidia is similar to species in the Colletotrichum
dematium species complex. However, the conidia are slightly
broader than those of the ex-type strain of C. dematium (3.5–4.5
vs 3–4 μm, Damm et al. 2009); the synonymy needs to be
confirmed. Publication data cited in name repositories (Allg. Bot.
Z. Syst. 2: 83. 1896.) are incorrect and instead refer to the
protologue of F. seemenianum (syn. F. avenaceum), an unrelated
taxon.
speiseri Fusarium Lindau, Rabenh. Krypt.-Fl. Ed. 2, 1(9): 580.
1909.
(See Fusarium avenaceum).
Holotypus: B 70 0100195.
Type locality: Poland, Karthaus, Nýdek.
Type substrate: Dead Auchenorrhyncha (cicada).
Note: Synonym fide Wollenweber & Reinking (1935).
spermogoniopsis Fusarium Jul. Müll., Ber. Deutsch. Bot. Ges. 3:
394. 1885.
Hymenella spermogoniopsis (Jul. Müll.) L. Lombard & Sand.Den., comb. nov. MycoBank MB 837721.
Basionym: Fusarium spermogoniopsis Jul. Müll., Ber. Deutsch.
Bot. Ges. 3: 394. 1885.
Synonym: Hymenula spermogoniopsis (Jul. Müll.) Wollenw.,
Fusaria Autogr. Delin. 1: 483. 1916.
Syntypes: ?B 70 0100196, B 70 0100197 & B 700100198.
FUSARIUM
Type locality: Germany
Type substrate: Sporocarps of Phragmidium subcorticium (=
Phragmidium mucronatum) and on the uredo- and teliospores of
Phragmidium rubi (= Phragmidium barclayi).
Notes: Wollenweber (1916) provided a new combination for
F. spermogoniopsis in the genus Hymenula. However, the
generic name Hymenella (1822) predates the generic name
Hymenula (1828) and therefore we provide a new combination in
the latter genus.
sphaeriae Fusarium Fuckel, Fungi Rhen. Exs., Fasc. 3, no. 212.
1863.
(See Fusarium clematidis)
Lectotypus: G00111017, designated in Gr€afenhan et al. (2011).
Lectotype locality: Germany, Hessen, Reichartshausen near
Oestrich-Winkel.
Lectotype substrate: Parasitic on Leptosphaeria (Sphaeria) dioica, on Urtica dioica.
sphaeriiforme Fusarium Sacc. (as ‘sphaeriaeforme’), Syll. Fung.
10: 723. 1892.
Replaced synonym: Fusarium celtidis Pass., Atti Reale Accad.
Lincei, Rendiconti Cl. Sci. Fis., ser. 4, 7: 51. 1891, nom. illegit.,
Art. 53.1.
(See Fusarium melanochlorum)
Holotypus: ?PARMA.
Type locality: Italy, Parma, Vigheffio.
Type substrate: Dead branches of Celtis australis.
Note: Synonyms fide Wollenweber & Reinking (1935).
sphaeroideum Fusarium Pass., Atti Reale Accad. Lincei, Rendiconti Cl. Sci. Fis., ser. 4, 4: 105. 1888.
(See Fusarium lateritium)
Holotypus: ?PARMA.
Type locality: Italy, Parma.
Type substrate: Branch of Ficus carica.
Note: Synonym fide Wollenweber & Reinking (1935).
sphaerosporum Fusarium Q.T. Chen & X.H. Fu, Acta Mycol. Sin.,
Suppl. 1: 331. 1987.
Neocosmospora sphaerospora (Q.T. Chen & X.H. Fu) Sand.Den. & Crous, Persoonia 43: 173. 2019.
Holotypus: HMAS 43749.
Ex-type culture: NF 5840.
Type locality: China, Guangdong Province, Maoming.
Type substrate: Water from underground pipes of oilfield.
Descriptions and illustrations: See Chen et al. (1987).
spinaciae Fusarium Hungerf., Phytopathology 13: 209. 1923.
(See Fusarium oxysporum)
Lectotypus (hic designatus, MBT 10000744): USA, Idaho, roots
of Spinacia oleracea, 1923, C.W. Hungerford, in Phytopathology
13: 208, fig. 4.
Notes: Synonym fide Booth (1971). No holotype specimen could
be located and therefore an illustration was designated as
lectotype.
spinosum Fusarium L. Lombard et al., Fungal Syst. Evol. 4:
195. 2019.
Holotypus: CBS H-24020.
Ex-type culture: CBS 122438.
Type locality: Brazil.
Type substrate: Galia melon imported into the Netherlands.
Descriptions and illustrations: See Lombard et al. (2019a).
www.studiesinmycology.org
REDELIMITED
Diagnostic DNA barcodes: rpb1: MN120729; rpb2: MN120747;
tef1: MN120768.
spinulosum Fusarium (Pfenning) O'Donnell et al., Index Fungorum 440: 4. 2020.
Neocosmospora spinulosa Pfenning, Sydowia 47: 66. 1995.
Holotypus: CBS H-5452a.
Ex-type culture: CBS 321.93.
Type locality: Brazil, Para, Capit~ao Poço.
Type substrate: Soil under Theobroma cacao.
Descriptions and illustrations: See Pfenning (1995).
splendens Fusarium Matuo & Takah. Kobay., Trans. Mycol. Soc.
Japan 2(4): 13. 1960, nom. inval., Art. 39.1.
(See Fusarium matuoi)
Authentic material: Not located.
Original locality: Japan.
Original substrate: Twigs of Albizzia julibrissin.
Descriptions and illustrations: See Matuo & Kobayashi (1960)
and Hosoya & Tubaki (2004).
sporodochiale Fusarium L. Lombard & Crous, Fungal Syst.
Evol. 4: 196. 2019.
Holotypus: CBS H-12681.
Ex-type culture: ATCC 14167 = CBS 220.61 = MUCL
8047 = NRRL 20842.
Type locality: South Africa, Gauteng Province, Johannesburg.
Type substrate: Soil.
Descriptions and illustrations: See Lombard et al. (2019a).
Diagnostic DNA barcodes: rpb1: MN120731; rpb2: MN120749;
tef1: MN120770.
sporotrichiella Fusarium Bilaĭ, Yadovitye griby na zerne khlebnykh zlakov: 86. 1953, nom. inval., Art. 39.1.
(See Fusarium sporotrichioides)
Authentic material: Not located.
Original locality: Ukraine.
Original substrate: Unknown.
Descriptions and illustrations: See Bilaĭ (1955).
Notes: This taxon was published as a new name for all the taxa
in section Sporotrichiella. However, it is invalid as no type and
Latin diagnosis were provided. Synonym fide Gerlach &
Nirenberg (1982).
sporotrichioides Fusarium Sherb., Mem. Cornell Univ. Agric.
Exp. Sta. 6: 183. 1915.
Synonyms: Fusarium sporotrichiella var. sporotrichioides
(Sherb.) Bilaĭ, Yadovitye griby na zerne khlebnykh zlakov
(Poisonous fungi on cereal seed), Kiev: 87. 1953, nom. inval.,
Art. 39.1.
Fusarium sporotrichiella Bilaĭ, Yadovitye griby na zerne khlebnykh zlakov (Poisonous fungi on cereal seed), Kiev: 86. 1953,
nom. inval., Art. 39.1.
Fusarium sporotrichioides var. minus Wollenw., Fusaria Autogr.
Delin. 3: 886. 1930.
Fusarium sporotrichioides subsp. minus (Wollenw.) Raillo, Fungi
of the Genus Fusarium: 196. 1950.
Lectotypus (hic designatus, MBT 10000745): USA, New York,
rotten tubers of Solanum tuberosum, together with F. solani and
F. oxysporum, 1915, C.D. Sherbakoff, in Mem. Cornell Univ.
Agric. Exp. Sta. 6: 184, fig. 22.
Notes: This economically important species requires epitypification from the type locality and substrate. No holotype
161
CROUS
ET AL.
specimen could be located and therefore an illustration was
designated as lectotype.
staphyleae Fusarium Samuels & Rogerson, Brittonia 36: 84.
1984.
Geejayessia atrofusca (Schwein.) Schroers & Gr€afenhan, Stud.
Mycol. 68: 126. 2011.
Basionym: Sphaeria atrofusca Schwein., Trans. Amer. Philos.
Soc., n.s. 4: 206. 1832.
Synonyms: Valsaria atrofusca (Schwein.) Cooke ex Sacc., Syll.
Fung. 9: 759. 1891.
Nectria atrofusca (Schwein.) Ellis & Everh., N. Amer. Pyren.: 99.
1892.
Pseudodiplodia atrofusca (Schwein.) Starb€ack, Bih. Kongl.
Svenska Vetensk.-Akad. Handl. 19: 94. 1894.
Cucurbitaria atrofusca (Schwein.) Kuntze, Revis. Gen. Pl. 3: 460.
1898.
Creonectria atrofusca (Schwein.) Seaver, Mycologia 1: 186.
1909.
Holotypus: In NY.
Ex-type culture: ATCC 66906 = CBS 502.94 = IMI
345891 = NRRL 22120.
Type locality: USA, Massachusetts, Berkshire, south of Ashley
Falls, Bartholomew's Cobble.
Type substrate: Branches of Staphylea trifolia.
Descriptions and illustrations: See Samuels & Rogerson (1984)
and Schroers et al. (2011).
sic et al., Antonie van Leeuwenhoek 111:
stercicola Fusarium Si
1793. 2018.
sic et al.) Sand.-Den. & Crous,
Neocosmospora stercicola (Si
Persoonia 43: 173. 2019.
Synonyms: Fusarium martii var. viride Sherb., Mem. Cornell
Univ. Agric. Exp. Sta. 6: 247. 1915.
Fusarium solani var. martii ‘f. 1’ Wollenw., Z. Parasitenk. (Berlin)
3: 290. 1931.
sic et al., Antonie van LeeuFusarium witzenhausenense Si
wenhoek 111: 1795. 2018.
Fusarium xiangyunense F. Zhang et al. (as ‘xiangyunensis’),
Phytotaxa 450: 278. 2020. nom. inval., Art. 40.8.
Holotypus: CBS H-23352.
Ex-type culture: CBS 142481 = DSM 106211 = FS 89.
Type locality: Germany, Niedersachsen, Hannover.
Type substrate: Compost yard waste plant debris.
sic et al. (2018a).
Descriptions and illustrations: See Si
Diagnostic DNA barcodes: rpb1: MW834255; rpb2: LR583887;
tef1: LR583658.
stercorarium Fusarium Rostr., Meddel. Grønland 18: 74. 1894.
Holotypus: C-F-92401
Type locality: Greenland, Vestfjord.
Type substrate: Dung of Rangifer tarandus (reindeer).
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
stercoris Fusarium Fuckel, Fungi Rhen. Exs., Suppl., Fasc. 5: no.
1921. 1867 [and Jahrb. Nassauischen Vereins Naturk. 23–24:
369. 1870].
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000746): Germany, Hessen,
Oestrich-Winkel, soil next to Peziza stercoraria, date unknown,
K.W.G.L. Fuckel, Fungi Rhen. Exs., Suppl., Fasc. 5: no. 1921 in
HAL.
162
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore the exsiccate
lodged in HAL is designated as lectotype.
sterilihyphosum Fusarium Britz et al., Mycologia 94: 726.
2002.
Holotypus: PREM 57302.
Ex-type culture: NRRL 25623.
Type locality: South Africa, Limpopo Province, Tzaneen, Letsitele area.
Type substrate: Malformed inflorescence of Mangifera indica.
Descriptions and illustrations: See Britz et al. (2002) and Leslie &
Summerell (2006).
Diagnostic DNA barcodes: rpb1: MN193925; rpb2: MN193897;
tef1: MN193869.
sticticum Fusarium Berk. & M.A. Curtis, in Berkeley, Grevillea 3:
99. 1875.
(See Fusarium lateritium)
Holotypus: ?K(M).
Type locality: USA, South Carolina.
Type substrate: Twigs of Prunus persica.
Note: Synonym fide Wollenweber & Reinking (1935).
stictoides Fusarium Durieu & Mont., Explor. Sci. Algerie 1: 334.
1848.
(See Fusarium graminearum)
Holotypus: ?PC.
Type locality: Algeria.
Type substrate: Branch of flowering Agave sp.
Note: Synonym fide Wollenweber & Reinking (1935).
stilbaster Fusarium (Link) Link, Sp. pl., Ed. 4, 6: 106. 1825.
Atractium stilbaster Link, Mag. Ges. Naturf. Freunde, Berlin 3:
10. 1809.
Synonyms: Atractium fuscum Sacc., Syll. Fung. 2: 514. 1883.
Stilbella fusca (Sacc.) Seifert, Stud. Mycol. 27: 77. 1985.
Atractium flavoviride Sacc., Syll. Fung. 2: 514. 1883.
Stilbum madidum Peck, Rep. (Annual) New York State Mus. Nat.
Hist. 46: 115. 1894.
Didymostilbe eichleriana Bres. & Sacc., Atti Congr. Bot. Palermo:
59. 1903.
Didymostilbe obovoidea Matsush., Icon. Microfung. Matsush.
lect.: 60. 1975.
Lectotypus: Illustration published in Mag. Ges. Naturf. Freunde,
Berlin 3, tab. I, fig. 11, designated in Gr€afenhan et al. (2011).
Epitypus: CBS 410.67 (preserved as metabolically inactive culture), designated in Gr€afenhan et al. (2011).
Ex-epitype culture: CBS 410.67.
Epitype locality: Germany, Bayerischer Wald, Rachelseewand.
Epitype substrate: Bark.
Descriptions and illustrations: See Seifert (1985) and Gr€afenhan
et al. (2011).
Diagnostic DNA barcodes: rpb1: KM232206; tef1: KM231920.
stilboides Fusarium Wollenw., Fusaria Autogr. Delin. 2: 615.
1924.
Synonyms: Fusarium lateritium var. stilboides (Wollenw.) Bilaĭ,
Fusarii (Biologija i sistematika): 266. 1955, nom. inval., Art. 41.5.
Fusarium lateritium var. stilboides (Wollenw.) Bilaĭ, Mikrobiol.
Zhurn. 49: 6. 1987.
Fusarium lateritium var. longum Wollenw., Fusaria Autogr. Delin.
1: 385. 1916.
FUSARIUM
Fusarium fructigenum var. minus Wollenw., Z. Parasitenk.
(Berlin) 3: 386. 1931.
Fusarium stilboides var. minus (Wollenw.) Wollenw., Z. Parasitenk. (Berlin) 3: 333. 1931.
Fusarium stilboides ‘f. 1’ Raillo, Fungi of the Genus Fusarium:
271. 1950.
Gibberella stilboides W.L. Gordon ex C. Booth, The Genus
Fusarium: 119. 1971.
Lectotypus (hic designatus, MBT 10000747): Philippines, Los
Ba~nos, living twigs of Citrus sp., invaded by coccids, 1917, O.A.
Reinking, in Fusaria Autogr. Delin. 2: 615.
Epitypus (hic designatus, MBT 10000748): Cook Islands, Citrus
sp., Sep. 1978, G.F. Laundon, CBS 746.79 (preserved as
metabolically inactive culture).
Ex-epitype culture: BBA 63887 = CBS 746.79 = ICMP
10624 = NRRL 25485.
Descriptions and illustrations: See Wollenweber (1924, 1930),
Wollenweber & Reinking (1935), Doidge (1938), Raillo (1950),
Booth (1971), and Gerlach & Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: MW928817; rpb2: MW928832;
tef1: MW928843.
Note: No holotype specimen could be located and therefore an
illustration was designated as lectotype.
stillatum Fusarium De Not. ex Sacc., in Berlese & Voglino, Syll.
Fung., Addit. I–IV: 390. 1886.
Myxosporium stillatum (De Not. ex Sacc.) Wollenw., Fusaria
Autogr. Delin. 1: 490. 1916.
Lectotypus (hic designatus, MBT 10000749): Italy,
Valle Intrasca, at the bridge on Possaccio, dried stems of Genista
tinctoria, 1862, G. de Notaris, S-F45664 [Baglietto, Cesati &
Notaris, Erb. Critt. Ital. Ser. I no. 148 (1148)].
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore the exsiccate
lodged in S is designated as lectotype.
stoveri Fusarium C. Booth, The Genus Fusarium: 37. 1971.
Microdochium stoveri (C. Booth) Samuels & I.C. Hallett, Trans.
Brit. Mycol. Soc. 81: 481. 1983.
Basionym: Micronectriella stoveri C. Booth, Mycol. Pap. 94: 3.
1964.
Synonym: Monographella stoveri (C. Booth) Samuels & I.C.
Hallett, Trans. Brit. Mycol. Soc. 81: 473. 1983.
Holotypus: IMI 92905.
Type locality: Honduras.
Type substrate: Leaf of Musa sp.
Descriptions and illustrations: See Booth (1964, 1971), Gerlach
& Nirenberg (1982) and Samuels & Hallet (1983).
striatum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6:
255. 1915.
(See Fusarium solani-melongenae)
Typus: ?CUP-007460.
Type locality: USA, Colorado
Type substrate: Solanum tuberosum.
Notes: Synonym fide Nirenberg & Brielmaiers-Liebetanz, 1996
and Sandoval-Denis et al. (2019). Lectotypification pending
study of material lodged in CUP.
strobilinum Fusarium Corda, Icon. Fung. 1: 4. 1837.
Sirococcus conigenus (Pers.) P.F. Cannon & Minter, Taxon 32:
577. 1983.
Basionym: Hysterium conigenum Pers., Ann. Bot. (Usteri) 15: 30.
1795.
www.studiesinmycology.org
REDELIMITED
Synonyms: Hypoderma conigenum (Pers.) DC., Fl. Franç., ed. 3,
2: 305. 1805.
Hypodermopsis conigena (Pers.) Kuntze, Revis. Gen. Pl. 3: 487.
1898.
Discella conigena (Pers.) Höhn., Mitt. Bot. Inst. T. H. Wien 6: 120.
1929.
Ascochyta strobilina (Corda) Wollenw., Fusaria Autogr. Delin. 1:
505. 1916.
Sphaeria strobilina Holl & J.C. Schmidt, Deutschl. Schw€amme,
Erste Lieferung: 4. 1815, nom. inval., Art. 38.1(a).
Sphaeria strobilina Holle & J.C. Schmidt ex Fr., Syst. Mycol. 2:
495. 1823.
Dichaena strobilina (Holle & J.C. Schmidt ex Fr.) Fr., Summa
Veg. Scand. 2: 403. 1849.
Sporonema strobilinum Desm., Ann. Sci. Nat., Bot., ser. 3, 18:
368. 1852.
Plenodomus strobilinus (Desm.) Höhn., Sitzungsber. Kaiserl.
Akad. Wiss. Wien, Math.-Naturwiss. Cl., Abt. 1, 119: 647. 1910.
Discella strobilina (Desm.) Died., Krypt.-Fl. Brandenburg 9: 752.
1914.
Sirococcus strobilinus (Desm.) Petr., Sydowia 1: 155. 1947, nom.
illegit., Art. 53.1.
Sirococcus strobilinus Preuss, Linnaea 26: 716. 1855.
Phoma conigena P. Karst., Rev. Mycol. (Toulouse) 7: 106. 1885.
Septoria parasitica R. Hartig, Z. Forst- Jagdwesen 1890: 1. 1890.
Diplodina parasitica (R. Hartig) Prill., Maladies des Plantes
Agricoles 2: fig. 365. 1897.
Ascochyta parasitica Fautrey, Rev. Mycol. (Toulouse) 13: 79.
1891.
Ascochyta piniperda Lindau, Nat. Pflanzenfam., Teil. I, 1: 367.
1900.
Phoma conigena var. abieticola Sacc., Ann. Mycol. 3: 233. 1905.
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, near Liberec (Reichenberg).
Type substrate: Rotten cone scales of Pinus sp.
Note: Typification pending study of material lodged in PRM.
stromaticola Fusarium Henn., Bot. Jahrb. Syst. 28: 280. 1900.
Dialonectria volutella (Ellis & Everh.) L. Lombard & Sand.Den., comb. nov. MycoBank MB 837722.
Basionym: Fusarium volutella Ellis & Everh., Proc. Acad. Nat.
Sci. Philadelphia 43: 93. 1891.
Synonyms: Fusarium aquaeductuum var. medium Wollenw.,
Fusaria Autogr. Delin. 3: 844. 1930.
Fusarium aquaeductuum subsp. medium (Wollenw.) Raillo,
Fungi of the Genus Fusarium: 278. 1950.
Dialonectria ullevolea Seifert & Gr€afenhan, Stud. Mycol. 68: 97.
2011.
Holotypus: In B fide Hein (1988).
Type locality: Japan, Tokyo.
Type substrate: Old stroma of Dothideaceae, on Bambusa sp.
branches with Zythia stromaticola.
Notes: Synonym fide Wollenweber & Reinking (1935) and
Gr€afenhan et al. (2011). The older epithet ‘volutella’ (1891) supersedes the epithet ‘ullevolea’ (2011) and, therefore, a new
combination is provided.
stromaticum Fusarium Delacr., Bull. Soc. Mycol. France 9: 186.
1893.
(See Fusarium heterosporum)
Holotypus: ?PC.
Type locality: France, overseas department of Mayotte, Mayotte
islands.
163
CROUS
ET AL.
Type substrate: Seeds of unknown Poaceae (= Gramineae).
Note: Synonym fide Wollenweber & Reinking (1935).
IMB 5238), which represents the ex-type culture (Gerlach &
Nirenberg 1982), is designated as lectotype.
subcarneum Fusarium P. Crouan & H. Crouan, Fl. Finistere: 14.
1867, nom. rej.
(See Fusarium sambucinum)
Authentic material: ?PC.
Original locality: France, Brittany, Finistere, marshes.
Original substrate: Twigs and dead leaves of Ulex sp.
Note: Synonym fide Wollenweber & Reinking (1935).
subnivale Fusarium Höhn., in Penther & Zederbauer, Ann. K. K.
Naturhist. Hofmus. 20: 369. 1905.
(See Fusarium dimerum)
Holotypus: FH00965354.
Type locality: Turkey, Anatolia.
Type substrate: Stems and leaves of decayed Astragalus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
subcorticale Fusarium Oudem., Ned. Kruidk. Arch., ser. 3, 3:
135. 1898.
(See Fusarium buxicola)
Holotypus: ?L.
Type locality: Netherlands, Zuid-Holland Province, Zorgvliet.
Type substrate: Buxus sempervirens.
Note: Synonym fide Wollenweber & Reinking (1935).
subpallidum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp.
Sta. Mem. 6: 230. 1915.
(See Fusarium sambucinum)
Typus: ?CUP-007480.
Type locality: USA, Louisiana, Edgerton.
Type substrate: Solanum tuberosum.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in CUP.
subglutinans Fusarium (Wollenw. & Reinking) P.E. Nelson
et al., Fusarium species. An illustrated manual for identification:
135. 1983.
Basionym: Fusarium moniliforme var. subglutinans Wollenw. &
Reinking, Phytopathology 15: 163. 1925.
Synonyms: Fusarium moniliforme f. subglutinans (Wollenw. &
Reinking) C. Moreau, Rev. Mycol. (Paris) 17: 23. 1952.
Fusarium sacchari var. subglutinans (Wollenw. & Reinking) Nirenberg, Mitt. Biol. Bundesanst. Land- Forstw. 169: 53. 1976.
Gibberella fujikuroi var. subglutinans (Wollenw. & Reinking) E.T.
Edwards, Agric. Gaz. New South Wales 44: 895. 1933 (Art. F.8.1,
Note 2, Exs. 2).
Gibberella subglutinans (Wollenw. & Reinking) P.E. Nelson et al.,
Fusarium species. An illustrated manual for identification (University Park): 135. 1983.
Neotypus: CBS 747.97 (preserved as metabolically inactive
culture), designated by Yilmaz et al. (2021).
Ex-neotype culture: BBA 62451 = CBS 747.97 = DAOM
225141 = FRC M-36 = MRC 8554 = NRRL 22016 = NRRL
22114.
Neotype locality: USA, Illinois, Saint Elmo.
Neotype substrate: Zea mays.
Descriptions and illustrations: See Booth (1971), Nirenberg
(1976, 1981), Nelson et al. (1983), Pascoe (1990), Leslie &
Summerell (2006).
Diagnostic DNA barcodes: rpb1: JX171486; rpb2: JX171599;
tef1: HM057336.
sublunatum Fusarium Reinking, Zentralbl. Bakteriol., 2. Abt.
89: 510. 1934.
Synonyms: Fusarium elongatum Reinking, Zentralbl. Bakteriol.
Parasitenk., Abt. 2, 89: 511. 1934, nom. illegit., Art. 53.1.
Fusarium sambucinum var. sublunatum (Reinking) Bilaĭ, Mikrobiol. Zhurn. 49: 6. 1987, nom. inval., Art. 41.4, Note 1.
Lectotypus (hic designatus, MBT 10000750): Costa Rica,
Limon, soil in Musa sapientum plantation, 1933, O.A. Reinking,
CBS 189.34 (preserved as metabolically inactive culture).
Ex-type culture: BBA 62431 = CBS 189.34 = DSM
62431 = NRRL 13384 = NRRL 20840.
Descriptions and illustrations: See Gerlach & Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: JX171451; rpb2: KX302935;
tef1: KX302919.
Notes: No holotype specimen could be located for F. sublunatum
and therefore the metabolically inactive culture CBS 189.34 (=
164
subtectum Fusarium Roberge ex Desm., Pl. Crypt. N. France,
ed. 1, Fasc. 29, no. 1428. 1845.
Rhodesia subtecta (Roberge ex Desm.) Grove, British Stemand Leaf-Fungi (Coelomycetes) 2: 205. 1937.
Synonyms: Myxosporina subtecta (Roberge ex Desm.) Höhn., in
Weese, Ber. Deutsch. Bot. Ges. 37: 155. 1919, nom. inval., Art.
35.1.
Myxosporina subtecta (Roberge ex Desm.) Höhn., Mitt. Bot. Inst.
Tech. Hochsch. Wien 4: 74. 1927.
Hainesia subtecta (Roberge ex Desm.) Grove, J. Bot. 70: 4.
1932.
Hymenula psammae Oudem., Ned. Kruidk. Arch., ser. 3, 1: 533.
1898. (fide Wollenweber & Reinking 1935).
Syntypes: Pl. Crypt. N. France no. 1428 in ?BRU, PC & PH.
Type locality: France.
Type substrate: Dead leaves of Arundo arenaria.
subtropicale Fusarium C. Pereira et al., Mycologia 110: 864.
2018.
Holotypus: BPI 910644.
Ex-type culture: CBS 144706 = NRRL 66764.
Type locality: Brazil, Parana State, Guarapuava.
Type substrate: Hordeum vulgare.
Descriptions and illustrations: See Pereira et al. (2018).
Diagnostic DNA barcodes: rpb1: MH706972; rpb2: MH706973;
tef1: MH706974.
subulatum Fusarium Appel & Wollenw., Arbeiten Kaiserl. Biol.
Anst. Land- Forstw. 8: 131. 1913.
Replaced synonym: Fusarium roseum var. lupini-albi Sacc.,
Michelia 2: 295. 1881.
(See Fusarium avenaceum)
Holotypus: Not located.
Type locality: Italy, Selva.
Type substrate: Lupinus albus.
Note: Synonyms fide Wollenweber & Reinking (1935).
subviolaceum Fusarium Roum. & Fautrey, Fungi Sel. Gall. Exs.
no. 6022. 1892.
(See Fusarium avenaceum)
Syntype: ILL0020193 (Fungi Sel. Gall. Exs. no. 6022).
Type locality: France, Jardin de Noidan.
Type substrate: Dry stems of Asparagus officinalis.
Note: Synonym fide Wollenweber & Reinking (1935).
FUSARIUM
succisae Fusarium Schröt. ex Sacc., Syll. Fung. 10: 724. 1892.
Synonym: Fusisporium succisae J. Schröt., Hedwigia 13: 180.
1874, nom. inval., Art. 36.1(a).
Lectotypus: ILL00076313 (Thümen, Mycoth. Univ. no. 675),
designated by Yilmaz et al. (2021).
Lectotype locality: Germany, Bavaria, Borussia.
Lectotype substrate: Succisa pratensis.
Epitypus: IMI 202876, designated by Yilmaz et al. (2021).
Ex-epitype culture: BBA 12287 = BBA 63627 = CBS
219.76 = DAOM 225142 = IMI 202876 = IMI 375347 = NRRL
13613.
Epitype locality: Germany.
Epitype substrate: Succisa pratensis.
Descriptions and illustrations: See Nirenberg (1976), Gerlach &
Nirenberg (1982).
Diagnostic DNA barcodes: rpb1: LT996207; rpb2: LT970764.
sudanense Fusarium S.A. Ahmed et al., Antonie van Leeuwenhoek 110: 826. 2017.
Holotypus: CBS H-22547.
Ex-type culture: CBS 454.97.
Type locality: Sudan.
Type substrate: Plant debris of Striga hermonthica.
Descriptions and illustrations: See Moussa et al. (2017).
Diagnostic DNA barcodes: rpb1: LT996208; rpb2: LT996155;
tef1: KU711697.
sulawesiense Fusarium Maryani et al. (as ‘sulawense’), Persoonia 43: 65. 2019.
Holotypus: InaCC F940 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F940.
Type locality: Indonesia, South Sulawesi, Bone, Kecamatan
Bengo, Desa Selli.
Type substrate: Infected pseudostem of Musa acuminata var.
Pisang Cere (AAA).
Descriptions and illustrations: See Maryani et al. (2019b).
Diagnostic DNA barcodes: rpb2: LS479855; tef1: LS479443.
sulphureum Fusarium Schltdl., Fl. Berol. 2: 139. 1824, nom. rej.
(See Fusarium sambucinum)
Holotypus: HAL 1613 F.
Type locality: Germany, Berlin.
Type substrate: Rotting tuber of Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935).
suttonianum Fusarium (Sand.-Den. & Crous) O'Donnell et al.,
Index Fungorum 440: 4. 2020.
Neocosmospora suttoniana Sand.-Den. & Crous, Persoonia
41: 123. 2018.
Holotypus: CBS H-23224.
Ex-type culture: CBS 143214 = FRC S-1423 = NRRL 32858.
Type locality: USA, Louisiana.
Type substrate: Homo sapiens.
Descriptions and illustrations: See Sandoval-Denis & Crous
(2018).
Diagnostic DNA barcodes: rpb1: MW218138; rpb2: EU329630;
tef1: DQ247163.
tabacinum Fusarium (J.F.H. Beyma) W. Gams, Persoonia 5: 179.
1968.
Basionym: Cephalosporium tabacinum J.F.H. Beyma, Zentralbl.
Bakteriol. 2. Abt. 89: 240. 1933.
www.studiesinmycology.org
REDELIMITED
Plectosphaerella cucumerina (Lindf.) W. Gams, in Domsch &
Gams, Fungi in Agricultural Soils: 160. 1972.
Basionym: Venturia cucumerina Lindf., Meddn Centralanst.
Försksv. Jordbruksomr. Bot. Avd. 17: 7. 1919.
Synonyms: Monographella cucumerina (Lindf.) Arx, Trans. Brit.
Mycol. Soc. 83: 374. 1984.
Microdochium tabacinum (J.F.H. Beyma) Arx, Trans. Brit. Mycol.
Soc. 83: 374. 1984.
Plectosporium tabacinum (J.F.H. Beyma) M.E. Palm, W. Gams &
Nirenberg, Mycologia 87: 399. 1995.
Plectosphaerella cucumeris Kleb., Phytopathol. Z. 1: 43. 1929.
Micronectriella cucumeris (Kleb.) C. Booth, The Genus Fusarium: 39. 1971.
Cephalosporium ciferrii Verona, Studio sulle cause microbiche
che danneggiano la carta ed i libri: 30. 1939.
Cephalosporiopsis imperfecta M. Moreau & Moreau, Rev. Mycol.
(Paris) 6: 67. 1941, nom. inval., Art. 39.1.
Neotypus: CBS H-7656, designated in Palm et al. (1995).
Ex-neotype culture: CBS 137.33 = MUCL 9701 = NRRL 22455.
Neotype locality: UK, England, Bristol.
Neotype substrate: Stems of Nicotiana tabacum.
Descriptions and illustrations: See Domsch et al. (2007), Carlucci
et al. (2012), Giraldo & Crous (2019).
tabacivorum Fusarium Delacr., Ann. Inst. Natl. Agron., ser. 2, 5:
207. 1906.
(See Fusarium oxysporum)
Holotypus: ?PC.
Type locality: France, Perigueux, Razac-sur-l'Isle.
Type substrate: Nicotiana tabacum.
Note: Synonym fide Wollenweber & Reinking (1935).
tanahbumbuense Fusarium Maryani et al., Persoonia 43: 63.
2019.
Holotypus: InaCC F965 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F965.
Type locality: Indonesia, South Kalimantan, Tanah Bumbu,
Kecamatan Kusan Hilir, Desa Betung.
Type substrate: Pseudostem of Musa var. Pisang Hawa.
Descriptions and illustrations: See Maryani et al. (2019b).
Diagnostic DNA barcodes: rpb1: LS479877; rpb2: LS479863;
tef1: LS479448.
tardichlamydosporum Fusarium Maryani et al., Stud. Mycol.
92: 181. 2018 [2019].
Holotypus: InaCC F958 (preserved as metabolically inactive
culture).
Ex-type culture: InaCC F958.
Type locality: Indonesia, East Nusa Tenggara, Sikka Flores,
Desa Kota Uneng Kecamatan Alok.
Type substrate: Pseudostem of Musa acuminata var. Pisang
Barangan.
Descriptions and illustrations: See Maryani et al. (2019a).
Diagnostic DNA barcodes: rpb1: LS479534; rpb2: LS479280;
tef1: LS479729.
tardicrescens Fusarium Maryani et al., Persoonia 43: 69. 2019.
Synonym: Fusarium tardicrescens Maryani et al., Stud. Mycol.
92: 185. 2018 [2019], nom. inval., Art. 40.7.
Holotypus: CBS 102024 (preserved as metabolically inactive
culture).
165
CROUS
ET AL.
Ex-type culture: CBS 102024 = NRRL 36113.
Type locality: Malawi, Karonga, Misuku Hills.
Type substrate: Musa sapientum cv. Harare.
Descriptions and illustrations: See Maryani et al. (2019b).
Diagnostic DNA barcodes: rpb1: LS479474; rpb2: LS479217;
tef1: LS479665.
tasmaniense Fusarium (McAlpine) Rossman (as ‘tasmanica’),
Mycol. Pap. 150: 54. 1983.
Basionym: Microcera tasmaniensis McAlpine, J. Dept. Agric.
Victoria 2: 647. 1904.
Synonyms: Discofusarium tasmaniense (McAlpine) Petch,
Trans. Brit. Mycol. Soc. 7: 143, 165. 1921.
Microcera myrtilaspis McAlpine, J. Dept. Agric. Victoria 2: 647.
1904.
Calonectria coccidophaga Petch, Trans. Brit. Mycol. Soc. 7: 161.
1921.
Nectria coccidophaga (Petch) Rossman, Mycotaxon 8: 499.
1979.
Holotypus: VPRI 2744.
Type locality: Australia, Tasmania.
Type substrate: Parasitic on Aspidiotus sp. (scale) on Eucalyptus
bark.
Descriptions and illustrations: See Rossman (1983).
Notes: Status unclear. Rossman (1983) studied the specimen in
K(M) and recombined the asexual morph name in Fusarium,
which is not supported by the features of the sexual-morph. This
species most likely belongs to Microcera as originally specified
by McAlpine (1904).
temperatum Fusarium Scaufl. & Munaut, Mycologia 103: 593.
2011.
Holotypus: MUCL 52463-H.
Ex-type culture: MUCL 52463.
Type locality: Belgium, Waals-Brabant Province, Chastre.
Type substrate: Zea mays.
Descriptions and illustrations: See Scauflaire et al. (2011).
Diagnostic DNA barcode: tef1: KM487197.
tenellum Fusarium Sacc. & Briard, Rev. Mycol. (Toulouse) 7:
212. 1885.
(See Fusarium sambucinum)
Holotypus: Not located.
Type locality: France, Troyes.
Type substrate: Rotten stem of Brassica oleracea.
Note: Synonym fide Wollenweber & Reinking (1935).
tenue Fusarium Corda, Icon. Fung. 1: 3. 1837.
(See Fusarium avenaceum)
Typus: In PRM fide Pilat (1938).
Type locality: Czech Republic, near Prague.
Type substrate: Rotting stem of an unidentified host.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of material lodged in PRM.
tenuicristatum Fusarium (S. Ueda & Udagawa) O'Donnell et al.,
Index Fungorum 440: 4. 2020.
Basionym: Neocosmospora tenuicristata S. Ueda & Udagawa,
Mycotaxon 16: 387. 1983.
Synonym: Acremonium tenuicristatum S. Ueda & Udagawa,
Mycotaxon 16: 387. 1983.
Holotypus: NHL 2911.
Type locality: Japan, Nagasaki.
Type substrate: Marine sludge.
166
Descriptions and illustrations: See Ueda & Udagawa (1983).
Notes: Status unclear. See Sandoval-Denis et al. (2019).
tenuissimum Fusarium (Peck) Sacc., Syll. Fung. 4: 711. 1886.
Basionym: Fusisporium tenuissimum Peck, Rep. (Annual) New
York State Mus. Nat. Hist. 34: 48. 1883. 1881.
(See Fusarium sambucinum)
Holotypus: NYSf3163.
Type locality: USA, New York, Schenectady.
Type substrate: Dead stem of unidentified host.
Note: Synonyms fide Wollenweber & Reinking (1935).
tenuistipes Fusarium Sacc., Atti Mem. Reale Accad. Sci. Lett.
Arti, Padova 33: 195. 1917.
(See Fusarium incarnatum)
Holotypus: In PAD.
Type locality: Unknown.
Type substrate: Pennisetum spicatum.
Note: Synonym fide Wollenweber & Reinking (1935).
terrestre Fusarium Manns, Bull. North Dakota Agric. Exp. Sta.:
no. 259. 1932.
(See Fusarium equiseti)
Holotypus: Not located.
Type locality: USA, North Dakota.
Type substrate: Soil.
Note: Synonym fide Wollenweber & Reinking (1935).
terricola Fusarium Al-Hatmi et al., Antonie van Leeuwenhoek
110: 826. 2017.
Holotypus: CBS H-22548.
Ex-type culture: CBS 483.94.
Type locality: Australia, Queensland.
Type substrate: Desert soil.
Descriptions and illustrations: See Moussa et al. (2017).
Diagnostic DNA barcodes: rpb1: LT996209; rpb2: LT996156;
tef1: KU711698.
thapsinum Fusarium Klittich et al., Mycologia 89: 644. 1997.
Synonym: Gibberella thapsina Klittich et al., Mycologia 89: 643.
1997.
Holotypus: BPI 737885.
Ex-type culture: ATCC 200522 = CBS 777.96 = FRC M-6564.
Type locality: USA, Kansas.
Type substrate: Stalk of Sorghum sp.
Descriptions and illustrations: See Klittich et al. (1997).
Diagnostic DNA barcodes: rpb1: MW928818; rpb2: MW928833;
tef1: MW928844.
theobromae Fusarium Appel & Strunk, Centralbl. Bacteriol., 2.
Abth., 11: 635. 1904.
Neocosmospora theobromae (Appel & Strunk) Sand.-Den. &
Crous, Persoonia 43: 174. 2019.
Synonyms: Fusarium javanicum var. theobromae (Appel &
Strunk) Wollenw., Z. Parasitenk. (Berlin) 3: 483. 1931.
Neotypus: BPI 453072, designated in Sandoval-Denis et al.
(2019).
Type locality: Cameroon, Victoria.
Type substrate: Fruits and seeds of Theobroma cacao.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcode: tef1: LR583660.
Notes: This Fusarium name was recently resurrected, neotypified, and transferred to Neocosmospora by Sandoval-Denis
et al. (2019). DNA barcodes were generated from the neotype
FUSARIUM
specimen; however,
epitypification.
fresh
collections
are
needed
for
theobromae Fusarium M.L. Lutz, Bull. Soc. Bot. France 53: L.
1907 [1906], nom. illegit., Art. 53.1.
Diplocladium theobromae Sacc. & Trotter, Syll. Fung. 22:
1309. 1913.
Authentic material: Not located.
Original locality: Democratic Republic of S~ao Tome and
Príncipe.
Original substrate: Fermented beans of Theobroma cacao.
Note: Originally erroneously assigned to the genus Fusarium.
thevetiae Fusarium Tassi, Atti Reale Accad. Fisiocrit. Siena, ser.
4, 8: 238. 1897.
Holotypus: ?SIENA.
Type locality: India.
Type substrate: Thevetia venenifera.
Notes: Status unclear. A doubtful species fide Wollenweber &
Reinking (1935).
thuemenii Fusarium Sacc., Syll. Fung. 4: 722. 1886.
Replaced synonym: Fusarium parasiticum Thüm., Nuovo Giorn.
Bot. Ital. 12: 198. 1880, nom. illegit., Art. 53.1.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: Russia, Orenburg.
Type substrate: Rotten branches of Betula verrucosa (= Betula
pendula).
Note: Synonym fide Wollenweber & Reinking (1935).
tjaetaba Fusarium T.T.H. Vu et al., Fungal Diversity 77: 361.
2015 [2016].
Holotypus: RBG 5361.
Ex-type culture: FRL14350 = NRRL 66243 = RBG 5361.
Type locality: Australia, Northern Territory, Litchfield National
Park.
Type substrate: Sorghum interjectum.
Descriptions and illustrations: See Laurence et al. (2016).
Diagnostic DNA barcodes: rpb1: KP083267; rpb2: KP083275;
tef1: KP083263.
tjaynera Fusarium J.L. Walsh et al., Fungal Diversity 77: 361.
2015 [2016].
Holotypus: RBG 5367.
Ex-type culture: NRRL 66246 = RBG 5367.
Type locality: Australia, Northern Territory, Litchfield National
Park.
Type substrate: Triodia microstachya.
Descriptions and illustrations: See Laurence et al. (2016).
Diagnostic DNA barcodes: rpb1: KP083268; rpb2: KP083279;
tef1: EF107152.
tomentosum Fusarium Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10:
359. 1868 [1869].
Holotypus: In K(M).
Type locality: Cuba.
Type substrate: Dead sticks.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
tonkinense Fusarium (Bugnic.) O'Donnell et al., Index Fungorum
440: 4. 2020.
Neocosmospora tonkinensis (Bugnic.) Sand.-Den. & Crous,
Persoonia 41: 126. 2018.
www.studiesinmycology.org
REDELIMITED
Basionym: Cylindrocarpon tonkinense Bugnic., Encyclop.
Mycol.11: 181. 1939.
Synonym: Fusarium ershadii M. Papizadeh et al., Europ. J. Pl.
Pathol. 151: 693. 2018, nom. illegit., Art. 52.1.
Holotypus: IMI 113868.
Ex-type culture: CBS 115.40 = IMI 113868.
Type locality: Vietnam, Tonkin.
Type substrate: Musa sapientum.
Diagnostic DNA barcodes: rpb1: MW218140; rpb2: LT960564;
tef1: LT906672.
torreyae Fusarium T. Aoki et al., Mycologia 105: 314. 2013.
Holotypus: BPI 884050.
Ex-type culture: CBS 133858 = MAFF 243468 = NRRL 54151.
Type locality: USA, Florida, Liberty County, Torreya State Park,
Aspalaga Tract.
Type substrate: Stem tissue of diseased Torreya taxifolia.
Descriptions and illustrations: See Aoki et al. (2013).
Diagnostic DNA barcodes: rpb1: MW928819; rpb2: MW928834;
tef1: MW928845.
tortuosum Fusarium Thüm. & Pass., Pilze Weinst.: 51. 1878.
Neofabraea vagabunda (Desm.) P.R. Johnst., IMA Fungus 5:
103. 2014.
Basionym: Phlyctema vagabunda Desm., Ann. Sci. Nat., Bot.,
ser. 3, 8: 16. 1847.
Synonyms: Rhabdospora vagabunda (Desm.) Zerov, Viznachnik
gribiv Ukraini. T. 3. Nezaversheni gribi: 501. 1971, nom. inval.,
Art. 41.1.
Rhabdospora vagabunda (Desm.) R.S. Mathur, Coelomycetes of
India: 234. 1979.
Gloeosporium tortuosum (Thüm. & Pass.) Sacc., Michelia 2: 117.
1880.
Myxosporium tortuosum (Thüm. & Pass.) Allesch., Rabenh.
Krypt.-Fl., Ed. 2, 1(7): 534. 1903.
?Fusarium obtusatum Corda, Icon. Fung. 1: 3. 1837.
Fusarium bipunctatum Preuss, Linnaea 25: 741. 1852.
Lituaria riessii Schulzer, Verh. K. K. Zool.-Bot. Ges. Wien 21:
1241. 1871.
Gloeosporium riessii (Schulzer) Schulzer & Sacc., Hedwigia 23:
110. 1884.
Gloeosporium tineum Sacc., Michelia 1: 219. 1878.
Gloeosporium frigidum Sacc., Michelia 2: 168. 1880.
Cylindrosporium frigidum (Sacc.) Vassiljevsky, Fungi Imperfecti
Parasitici 2: 515. 1950.
Gloeosporium pyrenoides Sacc. & Malbr., in Saccardo, Michelia
2: 633. 1882.
Gloeosporium phillyreae Pass., Atti Reale Accad. Lincei, Rendiconti Cl. Sci. Fis., ser. 4, 4: 103. 1888.
Gloeosporium allantosporum Fautrey, Rev. Mycol. (Toulouse) 14:
97. 1892.
Gloeosporium allantoideum Peck, Rep. (Annual) Regents Univ.
State New York New York State Mus. 45: 81. 1893.
Gloeosporium alutaceum Sacc., Malpighia 11: 317. 1897.
Allantozythia alutacea (Sacc.) Höhn., Ann. Mycol. 22: 203. 1924.
Phlyctema alutacea (Sacc.) Petr., Ann. Mycol. 27: 370. 1929.
Fusarium japonicum Allesch., Beibl. Hedwigia 36: (164). 1897.
Gloeosporium unedonis Traverso, R.C. Congr. Bot. Palermo,
1902: 3 (extr.). 1902.
Trichoseptoria fructigena Maubl., Bull. Trimestriel Soc. Mycol.
France 21: 95. 1905.
Gloeosporium beguinotii Sacc., in Potebnia, Ann. Mycol. 5: 20.
167
CROUS
ET AL.
1907.
Cylindrosporium olivae Petri, Ann. Mycol. 5: 324. 1907.
Gloeosporium olivae (Petri) Foschi, Ann. Sperim. Agrar, n.s. 9:
911. 1955.
Gloeosporium album Osterw., Centralbl. Bacteriol. Parasitenk.,
2. Abth., 18: 826. 1907.
Gloeosporium diervillae Grove, J. Bot. 60: 145. 1922.
Pezicula alba E.J. Guthrie, Trans. Brit. Mycol. Soc. 42: 504. 1959.
Neofabraea alba (E.J. Guthrie) Verkley, Stud. Mycol. 44: 125.
1999.
Holotypus: ?PARMA.
Type locality: Italy, Parma.
Type substrate: Dry twigs of Vitis vinifera.
Note: Synonyms fide Wollenweber & Reinking (1935).
torulosum Fusarium (Berk. & M.A. Curtis) Gruyter & J.H.M.
Schneid., Jaarb. Plantenziektenkundige Dienst, Wageningen
1989/1990, no. 168: 135. 1991, nom. inval., Art. 41.4.
Basionym: Fusidium torulosum Berk. & M.A. Curtis, Grevillea 3:
112. 1875.
(See Fusarium torulosum (Berk. & M.A. Curtis) Nirenberg)
torulosum Fusarium (Berk. & M.A. Curtis) Nirenberg, Mycopathologia 129: 136. 1995.
Basionym: Fusidium torulosum Berk. & M.A. Curtis, Grevillea 3:
112. 1875.
Synonyms: Fusoma torulosum (Berk. & M.A. Curtis) Sacc., Syll.
Fung. 4: 220. 1886.
Fusarium torulosum (Berk. & M.A. Curtis) Gruyter & J.H.M.
Schneid., Jaarboek. Plantenziektenkundige Dienst. Wageningen
1989/1990 no. 168: 135. 1991, nom. inval., Art. 41.4.
Fusarium sclerodermatis Oudem., Nederl. Kruidk. Arch. ser. 2, 5:
516. 1889.
Fusarium sambucinum var. coeruleum Wollenw., Ann. Mycol. 15:
55. 1917.
?Gibberella pulicaris var. minor Wollenw., Z. Parasitenk. (Berlin)
3: 356. 1931.
Syntype: ?Car Inf. no. 6034. in K(M).
Type locality: USA, Pennsylvania, Michener.
Type substrate: Decaying Brassica stalks or Pinus.
Descriptions and illustrations: See Nirenberg (1995).
toxicum Fusarium L. Lombard & J.W. Xia, Persoonia 43: 220.
2019.
Holotypus: CBS H-24071.
Ex-type culture: CBS 406.86 = FRC R-8507 = IMI
309347 = NRRL 25796.
Type locality: Germany, Berlin.
Type substrate: Soil.
Descriptions and illustrations: See Xia et al. (2019).
Diagnostic DNA barcodes: rpb2: MN170441; tef1: MN170508.
tracheiphilum Fusarium (E.F. Sm.) Wollenw., Phytopathology 3:
29. 1913.
Basionym: Neocosmospora vasinfecta var. tracheiphila E.F. Sm.,
Bull. Div. Veg. Physiol. Pathol. U.S.D.A. 17: 45. 1899.
(See Fusarium neocosmosporiellum)
Syntypes: IN BPI, F, FLAS, ISC, MICH, PUL, UC & WSP.
Type locality: USA, South Carolina, James Island.
Type substrate: Dead stem of Vigna sinensis.
Note: Published as a new name for the sporodochial morph
found on the authentic material of N. vasinfecta var. tracheiphila.
168
translucens Fusarium Berk. & Broome, Ann. Mag. Nat. Hist., ser.
4, 17: 141. 1876.
Holotypus: ?K(M).
Type locality: UK, Scotland, Glamis.
Type substrate: Wood.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
transvaalense Fusarium Sand.-Den. et al., MycoKeys 34: 82.
2018.
Holotypus: CBS H-23497.
Ex-type culture: CBS 144211.
Type locality: South Africa, Kruger National Park, Skukuza,
Granite Supersite.
Type substrate: Rhizosphere of Sida cordifolia.
Descriptions and illustrations: See Sandoval-Denis et al.
(2018b).
Diagnostic DNA barcodes: rpb1: LT996210; rpb2: LT996157;
tef1: LT996099.
tremelloides Fusarium Grev., Scott. Crypt. Fl. 1: 10. 1822.
Calloria tremelloides (Grev.) L. Lombard, comb. nov. MycoBank MB 837723.
Basionym: Fusarium tremelloides Grev., Scott. Crypt. Fl. 1: 10.
1822.
Synonyms: Peziza fusarioides Berk., Mag. Zool. Bot. 1: 46. 1837.
Calloria fusarioides (Berk.) Fr., Summa Veg. Scand. 2: 359. 1849.
Callorina fusarioides (Berk.) Korf, Phytologia 21: 203. 1971.
Peziza neglecta Lib., Pl. Crypt. Arduenna Fasc. 1: no. 29. 1830.
Calloria neglecta (Lib.) B. Hein, Beih. Willdenowia 9: 54. 1976.
Holotypus: Not located.
Type locality: UK, Scotland, near Edinburg.
Type substrate: Dead stems of Urtica dioica.
Notes: Synonyms fide Wollenweber & Reinking (1935). As the
epithet of F. tremelloides (1822) takes priority above the epithet
of C. neglecta (1830), a new combination is introduced here.
trichothecioides Fusarium Wollenw., J. Wash. Acad. Sci. 2: 147.
1912.
Synonyms: Fusarium sambucinum var. trichothecioides (Wollenw.) Bilaĭ, Fusarii (Biologija i sistematika): 268. 1955, nom.
inval., Art. 41.1.
Fusarium tuberivorum Wilcox & G.K. Link, Res. Bull. Nebraska
Agric. Exp. Sta. 1: 48. 1913.
Lectotypus (hic designatus, MBT 10000751): USA, rotten tuber
of Solanum tuberosum, Aug. 1912, H.W. Wollenweber, in J.
Wash. Acad. Sci. 2: 150, figs A–F.
Descriptions and illustrations: See Booth (1971) and Gerlach &
Nirenberg (1982).
Notes: A putative synonym of F. sulphureum (Gordon 1959,
Subramanian 1971, Gerlach & Nirenberg 1982) or
F. sambucinum (Nelson et al. 1983, Nirenberg 1995). The taxonomy of this potato pathogen has not yet been resolved. As no
holotype specimen was preserved (Gerlach & Nirenberg 1982),
the figures accompanying the original protologue are designated
as lectotype here.
tricinctum Fusarium (Corda) Sacc., Syll. Fung. 4: 700. 1886.
Basionym: Selenosporium tricinctum Corda, Icon. Fung. 2: 7.
1838.
Synonyms: Fusarium sporotrichioides var. tricinctum (Corda)
Raillo, Fungi of the Genus Fusarium: 197. 1950.
FUSARIUM
Fusarium sporotrichiella var. tricinctum (Corda) Bilaĭ, Yadovitye
griby na zerne khlebnykh zlakov. Kiev: 87. 1953, nom. inval., Art.
39.1.
Fusarium sporotrichiella var. tricinctum (Corda) Bilaĭ, Mikrobiol.
Zhurn. 49: 7. 1987, nom. inval., Art. 35.1.
?Vermicularia subeffigurata γ helianthi Schwein., Trans. Amer.
Philos. Soc., n.s. 4: 228. 1832 [1834].
?Fusarium helianthi (Schwein.) Wollenw., Fusaria Autogr. Delin.
2: 555. 1924.
Fusarium muentzii Delacr. (as ‘müntzii’), Bull. Soc. Mycol. France
8: 192. 1892.
Fusarium citriforme Jamal., Valt. Maatalousk. Julk. 123: 11. 1943.
Gibberella tricincta El-Gholl et al., Canad. J. Bot. 56: 2206. 1978.
Lectotypus: PRM 155623 (designated in Holubova-Jechova et al.
1994).
Type locality: Czech Republic, near Prague, Chuchle,
Vyskocilka.
Type substrate: Stem of Umbelliferae.
Epitypus: In PRM, designated in Holubova-Jechova et al. (1994).
Ex-epitype culture: BBA 64485 = CBS 393.93 = NRRL 25481.
Epitype locality: Germany, Berlin.
Epitype substrate: Culm base of Triticum aestivum.
Descriptions and illustrations: See Holubova-Jechova et al.
(1994) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: JX171516; rpb2: JX171629;
tef1: MH582379.
trifolii Fusarium Jacz., Jahrb. Pflanzenkrankh. Russlands. VIIVIII: Abt. 6. 1917.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: Russia, St. Petersburg.
Type substrate: Root crown of Trifolium sp.
Note: Synonym fide Wollenweber & Reinking (1935).
triseptatum Fusarium L. Lombard & Crous, Persoonia 43: 34.
2018 [2019].
Holotypus: CBS H-23622.
Ex-type culture: CBS 258.50 = NRRL 36389.
Type locality: USA.
Type substrate: Ipomoea batatas.
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb1: MW928820; rpb2: MH484873;
tef1: MH484964.
tritici Fusarium Liebman bis, Tidsskr. Landoekon., n.s., 2: 515.
1840.
(See Fusarium avenaceum)
Lectotypus (hic designates, MBT 10000752): Denmark, Triticum
sp., in Tidsskr. Landoekon., n.s., 2: figs B, 1, 2.
Notes: Synonymy fide Rostrup (1894). No holotype specimen
could be located and therefore an illustration is designated as
lectotype.
tritici Fusarium Erikss., Fungi Paras. Scand. Exs. no. 400. 1891,
nom. illegit., Art. 53.1.
(See Fusarium nivale)
Authentic material: CHRB-F-0007556.
Original locality: Sweden, Stockholm.
Original substrate: Triticum durum.
Note: Synonym fide Wollenweber & Reinking (1935).
truncatum Fusarium Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 155. 1915.
www.studiesinmycology.org
REDELIMITED
(See Fusarium avenaceum)
Typus: ?CUP-007429.
Type locality: USA, New York.
Type substrate: Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of the material lodged in CUP.
tuaranense Fusarium T. Aoki et al., Mycologia 111: 926. 2019.
Neocosmospora tuaranensis (T. Aoki et al.) L. Lombard &
Sand.-Den., comb. nov. MycoBank MB 837724.
Basionym: Fusarium tuaranense T. Aoki et al., Mycologia 111:
926. 2019.
Holotypus: BPI 910971.
Ex-type culture: ATCC 16563 = MAFF 246842 = NRRL 22231.
Type locality: Malaysia, Sabah State, Tuaran.
Type substrate: Hevea brasiliensis damaged by an unknown
ambrosia beetle.
Descriptions and illustrations: See Aoki et al. (2019).
Diagnostic DNA barcodes: rpb1: KC691600; rpb2: KC691660,
KC691631; tef1: KC691542.
Note: A new combination is provided in the genus Neocosmospora based on the phylogenetic relationship and
morphology of this species (Aoki et al. 2019).
tubercularioides Fusarium (Corda) Sacc., Syll. Fung. 4: 697.
1886.
Basionym: Selenosporium tubercularioides Corda, Icon. Fung. 1:
7. 1837.
(See Fusarium avenaceum)
Typus: PRM 155625.
Type locality: Czech Republic, Liberec, Hamrstejn (as ‘Sudetenland, Reichenberg, Hammerstein’).
Type substrate: Dead branches of Rubus idaeus.
Descriptions and illustrations: See Holubova-Jechova et al.
(1994).
Note: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of the material lodged in PRM.
tuberis Fusarium Preuss, Linnaea 24: 148. 1851.
Holotypus: In B fide Jülich (1974).
Type locality: Germany, Hoyerswerda.
Type substrate: Tuber of Dahlia sp.
Note: Status unclear. Not Fusarium fide Wollenweber & Reinking
(1935).
tuberivorum Fusarium Wilcox & G.K. Link, Res. Bull. Nebraska
Agric. Exp. Sta. 1: 48. 1913.
(See Fusarium trichothecioides)
Lectotypus (hic designates, MBT 10000753): USA, Nebraska,
Solanum tuberosum, in Res. Bull. Nebraska Agric. Exp. Sta. 1,
Pl. 24.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
tucumaniae Fusarium T. Aoki et al., Mycologia 95: 664. 2003.
(See Fusarium azukiicola)
Holotypus: BPI 841955.
Ex-type culture: MAFF 238418 = MJ-172 = NRRL 31096.
Type locality: Argentina, Tucuman, San Agustin.
Type substrate: Glycine max.
Descriptions and illustrations: See Aoki et al. (2003).
Diagnostic DNA barcodes: rpb1: MAED01000445; rpb2:
EU329557; tef1: GU170636.
169
CROUS
ET AL.
tumidum Fusarium Sherb., Phytopathology 18: 148. 1928.
Synonym: Gibberella tumida P.G. Broadh. & P.R. Johnst., Mycol.
Res. 98: 730. 1994.
Syntypes: Krieger, Fungi Saxon. Exs. no. 2499 in BPI & HAL.
Type locality: Germany.
Type substrate: Heads of Sarothamnus scoparius.
Note: Typification pending further study of the syntypes.
tupiense Fusarium C.S. Lima et al., Mycologia 104: 1414. 2012.
Holotypus: CMB-UB 22068.
Ex-type culture: CML 262 = CMM 3655 = KSU 16195 = NRRL
53984.
Type locality: Brazil, Minas Gerais, Lavras.
Type substrate: Diseased tissue of Mangifera indica.
Descriptions and illustrations: See Lima et al. (2012).
Diagnostic DNA barcodes: rpb1: LR792583; rpb2: LR792619;
tef1: GU737404.
udum Fusarium E.J. Butler, Mem. Dept. Agric. India, Bot. Ser.
2(9): 54. 1910.
Synonyms: Fusarium oxysporum f. sp. udum (E.J. Butler) W.C.
Snyder & H.N. Hansen, Amer. J. Bot. 24: 66. 1940.
Fusarium butleri Wollenw., Phytopathology 3: 38. 1913, nom.
illegit., Art. 52.1.
Fusarium lateritium var. uncinatum (Wollenw.) Wollenw., Z.
Parasitenk. (Berlin) 3: 375. 1931.
Fusarium vasinfectum var. crotalariae Kulkarni, Indian J. Agric.
Sci. 4: 994. 1934.
Fusarium udum f. sp. crotalariae (Kulkarni) Subram., The Genus
Fusarium: 114. 1971.
Fusarium udum var. cajani Padwick, Indian J. Agric. Sci. 10: 878.
1940.
Fusarium lateritium f. cajani (Padwick) W.L. Gordon, Canad. J.
Bot. 30: 232. 1952.
Fusarium udum var. crotalariae Padwick, Indian J. Agric. Sci. 10:
877. 1940.
Fusarium lateritium f. crotalariae (Padwick) W.L. Gordon, Canad.
J. Bot. 30: 232. 1952.
Gibberella indica B. Rai & R.S. Upadhyay, Mycologia 74: 343.
1982.
Lectotypus: Butler (1910), Pl. IV, fig. 4, designated in Pfenning
et al. (2019).
Epitypus: UB23905, designated in Pfenning et al. (2019).
Ex-epitype culture: BBA 65058 = CML 3238 = NRRL 25199.
Type locality: India.
Type substrate: Cajanus cajan.
Descriptions and illustrations: See Wollenweber & Reinking
(1935), Booth (1971), Subramanian (1971), Booth (1978),
Gerlach & Nirenberg (1982) and Pfenning et al. (2019).
Diagnostic DNA barcodes: rpb2: KY498875; tef1: MK639096.
udum Fusarium (Berk.) Wollenw., Phytopathology 3: 38. 1913,
nom. illegit., Art. 53.1.
Basionym: Fusisporium udum Berk., Ann. Mag. Nat. Hist. 6: 438.
1841.
(See Fusarium merismoides)
Holotypus: ?K(M).
Type locality: UK, King's Cliffe.
Type substrate: Unidentified tree.
Note: Synonyms fide Wollenweber & Reinking (1935).
ulmi Fusarium P. Crouan & H. Crouan, Fl. Finistere: 14. 1867.
(See Fusarium candidum (Link) Sacc.)
Holotypus: ?PC.
170
Type locality: France, Finistere, edge of a stream.
Type substrate: Roots of Ulmus sp.
Note: Synonym fide Wollenweber & Reinking (1935).
ulmicola Fusarium Dearn. & House, Circ. New York Stat. Mus.
24: 60. 1940, nom. inval., Art. 39.1.
Authentic material: NYSf3256.
Original locality: USA, New York, Albany, Ravena.
Original substrate: Dead branches of Ulmus thomasii.
Notes: Lacks a Latin diagnosis. Requires further investigation to
confirm its taxonomic affiliation.
uncinatum Fusarium Wollenw., Ann. Mycol. 15: 54. 1917.
(See Fusarium udum)
Holotypus: Not located.
Type locality: India, Dehli, Pusa.
Type substrate: Dried stem of Cajanus indicus.
Note: Synonym fide Wollenweber & Reinking (1935) and Gerlach
& Nirenberg (1982).
uniseptatum Fusarium Höhn., Ann. Mycol. 1: 409. 1903.
Synonyms: Cylindrocarpon uniseptatum (Höhn.) Wollenw.,
Fusaria Autogr. Delin. 2: 646. 1924.
Ramularia uniseptata (Höhn.) Wollenw., Fusaria Autogr. Delin. 2:
646. 1924.
Holotypus: Not located.
Type locality: Austria, Vienna.
Type substrate: Rotten Gleditsia triacanthos.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935) and not Ramularia fide Braun (1998).
uredinicola Fusarium Jul. Müll., Ber. Deutsch. Bot. Ges. 3: 395.
1885.
(See Fusarium avenaceum)
Holotypus: Not located.
Type locality: Germany.
Type substrate: Aecidium of Phragmidium subcorticium (=
Phragmidium mucronatum) and Phragmidium rubi (= Phragmidium barclayi).
Note: Synonym fide Wollenweber & Reinking (1935).
uredinicola Fusarium Pat. & Gaillard, Bull. Soc. Mycol. France 4:
127. 1888, nom. illegit., Art. 53.1.
Synonym: Fusarium patouillardii Sacc. (as ‘patouillardi’), Syll.
Fung. 10: 729. 1892.
Authentic material: Not located.
Original locality: Venezuela, Caracas.
Original substrate: Parasitic on the bottom of spots of Puccinia
pallidissima, between the perithecia of Darluca filum parasitised
by the Puccinia sp.
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
uredinicola Fusarium Petch, Ann. Roy. Bot. Gard. (Peradeniya)
6: 256. 1917, nom. illegit., Art. 53.1.
Authentic material: PDA 4731.
Original locality: Sri Lanka, Hakgala.
Original substrate: Parasitic on Uredo microglossa on leaves of
Microglossa zeylanica.
Notes: Status unclear. A probable synonym of F. solani var.
minus (syn. Neocosmospora brevicona) according to
Wollenweber & Reinking (1935).
uredinophilum Fusarium Speg. (as ‘urediniphilum’), Anales Mus.
Nac. Hist. Nat. Buenos Aires 31: 445. 1922.
FUSARIUM
Holotypus: In LPS (Fungi Parag. pp. 93–94, no. 262).
Type locality: Paraguay, near Puerto Sajonia.
Type substrate: Parasitic on the acervuli of Uredo cyclotrauma,
on leaving leaves of Pithecellobium cauliflorum.
Notes: Status unclear. Not treated by any of Wollenweber &
Reinking (1935), Booth (1971), or Gerlach & Nirenberg (1982).
uredinum Fusarium Ellis & Everh., N. Amer. Fungi, Ser. II, no.
2799. 1890, nom. inval., Art. 38.1(a).
Ramularia uredinis (W. Voss) Sacc., Syll. Fung. 4: 199. 1886.
Basionym: Cylindrosporium uredinis W. Voss, Verh. Zool.-Bot.
Ges. Wien 29: 684. 1879.
Synonym: Ramularia nambuana Henn., Hedwigia 43: 146. 1904.
Authentic material: NY00928692.
Original locality: USA, Wisconsin, Racine.
Original substrate: Parasitic on uredinia of Melampsora salicina,
on leaf of Salix sp.
Notes: Wollenweber & Reinking (1935) considered F. uredinum a
synonym of Cladosporium herbarum. It is quite possible that this
common saprobic Cladosporium species also occurred on
uredinia in N. Am. Fungi 2799, but it can be ruled out that Ellis &
Everhard confused this dematiaceous hyphomycete characterised by having long conidiophores with thickened and darkened conidiogeneous loci and large catenate conidia with a
colourless Fusarium. Davis (1915) found Ramularia uredinis, a
common mucedinacous hyphomycete on Melampsora spp. on
Populus and Salix, in material authentic for this name. This is
undoubtedly correct.
urticearum Fusarium (Corda) Sacc., Syll. Fung. 4: 698. 1886.
Basionym: Selenosporium urticearum Corda, Icon. Fung. 2: 7.
1838.
(See Fusarium lateritium)
Lectotypus (hic designatus, MBT 10000754): Czech Republic,
Prague, dead branches of Ficus elastica and Morus nigra, 1838.
A.C.J. Corda, in Icon. Fung. 2, Tab. 9, fig. 30.
Notes: Synonym fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
ussurianum Fusarium T. Aoki et al., Mycologia 101: 847. 2009.
Holotypus: BPI 878845.
Ex-type culture: CBS 123752 = NRRL 45681 = TG-2662/0.
Type locality: Russia, Ussuriysk, Primorsky krai (Far East territory), agricultural field near the city Ussuriysk.
Type substrate: Seed of Avena sativa.
Descriptions and illustrations: See Yli-Mattila et al. (2009).
Diagnostic DNA barcodes: rpb1: KM361648; rpb2: KM361666;
tef1: FJ240301.
ustilaginis Fusarium Kellerm. & Swingle, Rep. (Annual) Kansas
Agric. Exp. Sta. 2: 285. 1890 [1889].
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000755): USA, Kansas,
Manhattan, on Ustilago avenae, on Avena sativa, 1890, W.A.
Kellerman & W.T. Swingle, in Rep. (Annual) Kansas Agric. Exp.
Sta. 2, pl. IX, figs 1–13.
Note: Synonym fide Wollenweber & Reinking (1935).
ustilaginis Fusarium Rostr., Bot. Foren. Festskr. 54: 137. 1890,
nom. illegit., Art. 53.1.
(See Fusarium nivale)
Authentic material: C-F-125286.
Original locality: Denmark, Jutland, near Viborg.
www.studiesinmycology.org
REDELIMITED
Original substrate: Parasitic on Ustilago grandis on Phragmites
communis.
Note: Synonym fide Wollenweber & Reinking (1935).
vanettenii Fusarium O'Donnell et al., Index Fungorum 440: 5.
2020.
Basionym: Fusarium martii var. pisi F.R. Jones, J. Agric. Res. 26:
459. 1923.
(See Fusarium pisi)
vasinfectum Fusarium G.F. Atk., Bull. Alabama Agric. Exp. Sta.
41: 28. 1892.
(See Fusarium oxysporum)
Holotypus: ?CUP-A-(0100)#1.
Type locality: USA, Alabama, Montgomery, Mathews.
Type substrate: Gossypium herbaceum.
Note: Synonym fide Wollenweber & Reinking (1935).
venenatum Fusarium Nirenberg, Mycopathologia 129: 136.
1995.
Misapplied
names:
Fusarium
sambucinum
var.
coeruleum Wollenw. sensu Booth, The Genus Fusarium:
171–172. 1971.
Fusarium sambucinum var. coeruleum Wollenw. sensu Gerlach
& Nirenberg, Mitt. Biol. Bundesanst. Land.- Forstw. 209:
213–216. 1982.
Holotypus: CBS 458.93 (preserved as metabolically inactive
culture).
Ex-type culture: BBA 64537 = CBS 458.93 = NRRL 26228.
Type locality: Austria.
Type substrate: Culm of Triticum aestivum.
Descriptions and illustrations: See Nirenberg (1995).
Diagnostic DNA barcodes: rpb2: KM232382; tef1: KM231942.
venerorum Fusarium Dounin & Goldmacher, Index of the plant
diseases in the U.S. 5: 284–298. 1927.
(See Fusarium avenaceum)
Holotypus: Not located.
Type locality: Unknown.
Type substrate: Unknown.
Note: Synonym fide Wollenweber & Reinking (1935).
venezuelense Fusarium O'Donnell et al., Index Fungorum 440:
5. 2020.
Neocosmospora robusta Sand.-Den. & Crous, Persoonia 43:
165. 2019, non Fusarium robustum Gerlach 1977.
Holotypus: CBS H-24000.
Ex-type culture: BBA 65682 = CBS 145473 = NRRL 22395.
Type locality: Venezuela.
Type substrate: Bark.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW834251; rpb2: EU329507;
tef1: AF178341.
ventricosum Fusarium Appel & Wollenw., Phytopathology 3: 32.
1913.
Rectifusarium ventricosum (Appel & Wollenw.) L. Lombard &
Crous, Stud. Mycol. 80: 229. 2015.
Synonyms: Fusarium solani var. ventricosum (Appel & Wollenw.)
Joffe, Pl. & Soil 38: 440. 1973.
Fusarium cuneiforme Sherb., Mem. Cornell Univ. Agric. Exp. Sta.
6: 129. 1915.
Hypomyces solani Reinke & Berth., Untersuch. Bot. Lab. Univ.
Göttingen 1: 27. 1879.
171
CROUS
ET AL.
Hyponectria solani (Reinke & Berth.) Petch, J. Bot. 75. 220.
1937.
Nectriopsis solani (Reinke & Berth.) C. Booth, Mycol. Pap. 74: 8.
1960.
Nectria ventricosa Booth, The Genus Fusarium: 55. 1971.
Holotypus: B 70 0021849.
Epitypus: CBS H-21947, designated in Lombard et al. (2015).
Ex-epitype culture: BBA 62452 = CBS 748.79 = NRRL
20846 = NRRL 22113.
Type locality: Germany, Berlin.
Type substrate: Tuber of Solanum tuberosum.
Descriptions and illustrations: See Wollenweber (1917), Booth
(1971) and Lombard et al. (2015).
Diagnostic DNA barcodes: rpb1: JX171484; rpb2: JX171597;
tef1: KM231924.
Notes: Contrary to Wollenweber & Reinking (1935), Booth (1971)
considered this species as different from F. argillaceum, which
was later confirmed by Lombard et al. (2015). The same authors
designated an epitype for this taxon and transferred it to the
genus Rectifusarium as R. ventricosum.
veratri Fusarium (Allesch.) Höhn., in Kabat & Bubak, Fungi
Imperf. Exs. No. 349. 1906.
Gloeosporium veratri (Allesch.) Höhn., Mitt. Bot. Inst. Tech.
Hochsch. Wien 4: 112. 1927.
Basionym: Fusoma veratri Allesch., Ber. Bayer. Bot. Ges. 2: 19.
1892.
Synonym: Septogloeum veratri (Allesch.) Wollenw., Fusaria
Autogr. Delin. 1: 439. 1916.
Holotypus: ?M.
Type locality: Germany, Bavaria, Oberammergau.
Type substrate: Leaves of Veratrum lobelianum.
Notes: This species produces acervuli and 1-septate conidia with
truncate basal cells. Therefore, it was transferred to Gloesporium
(Helotiales, Dermataceae).
verrucosum Fusarium (Pat.) O'Donnell & Geiser, Phytopathology
103: 404. 2013.
Albonectria verrucosa (Pat.) Rossman & Samuels, Stud.
Mycol. 42: 108. 1999.
Basionym: Calonectria verrucosa Pat., Bull. Soc. Mycol. France
11: 228. 1895.
Synonym: Nectria astromata Rossman, Mycotaxon 8: 550. 1979,
non N. verrucosa (Schwein.) Sacc.
Holotypus: In FH fide Rossman et al. (1999).
Type locality: Ecuador, San Jorge.
Type substrate: Chusquea sp.
Descriptions and illustrations: See Rossman (1983) and
Rossman et al. (1999).
Notes: Although recently recombined in Fusarium (Geiser et al.
2013), the taxonomy of this species is uncertain. With 5–9(–13)septate ascospores, this species cannot be a member of
Fusarium s. str., and the identity of the isolates included in recent
phylogenetic estimates (CBS 102163, originally identified as
F. concolor and NRRL 22566) cannot be confirmed at this stage.
versicolor Fusarium Sacc., Syll. Fung. 16: 1099. 1902.
(See Fusarium culmorum)
Holotypus: In PAD.
Type locality: France, C^ote-d’Or.
Type substrate: Cortex of Cucurbita sp.
Note: Synonym fide Wollenweber & Reinking (1935).
versiforme Fusarium Kabat & Bubak, Hedwigia 44: 358. 1905.
172
Holotypus: BPI 453128.
Type locality: Czech Republic, Bohemia, Turnov.
Type substrate: Living leaves of Hosta sieboldii (syn. Hosta
albomarginata).
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
verticillioides Fusarium (Sacc.) Nirenberg, Mitt. Biol. Bundesanst. Land- Forstw. 169: 26. 1976.
Basionym: Oospora verticillioides Sacc., Fung. Ital., Fasc.
17–28: pl. 879. 1881.
Synonyms: Alysidium verticillioides (Sacc.) Kuntze, Revis. Gen.
Pl. 3: 442. 1898.
Fusarium moniliforme J. Sheld., Annual Rep. Nebraska Agric.
Exp. Sta. 17: 23. 1904.
Gibberella moniliformis Wineland, J. Agric. Res. 28: 909. 1924.
Lectotypus: Pl. 879 in Saccardo, Fung. Ital. (1881), designated
by Yilmaz et al. (2021).
Epitypus: CBS 218.76 (preserved as metabolically inactive culture), designated by Yilmaz et al. (2021).
Ex-epitype culture: BBA 11782 = CBS 218.76 = DSM
62264 = IMI 202875 = NRRL 13993.
Epitype locality: Germany.
Epitype substrate: Zea mays.
Descriptions and illustrations: See Nirenberg (1976, 1981),
Gerlach & Nirenberg (1982) and Leslie & Summerell (2006).
Diagnostic DNA barcodes: rpb1: MW402638; rpb2: MW928835;
tef1: KF499582.
veterinarium Fusarium L. Lombard & Crous, Persoonia 43: 35.
2018 [2019].
Holotypus: CBS H-23623.
Ex-type culture: CBS 109898 = NRRL 36153.
Type locality: Netherlands.
Type substrate: Peritoneum of Selachimorpha (shark).
Descriptions and illustrations: See Lombard et al. (2019b).
Diagnostic DNA barcodes: rpb2: MH484899; tef1: MH484990.
victoriae Fusarium Henn., in herb., fide Wollenweber, Fusaria
Autogr. Delin. 1: 66. 1916.
Macronectria jungneri (Henn.) C. Salgado & P. Chaverri,
Fungal Diversity 80: 448. 2016. Basionym: Nectria jungneri
Henn., Bot. Jahrb. Syst. 22: 75. 1895.
Synonyms: Nectria eustoma Penz. & Sacc., Malpighia 11: 509.
1898.
Nectria leucocoma Starb€ack, Bih. Kongl. Svenska Vetensk.Akad. Handl. 25: 28. 1899.
Nectria cinereopapillata Henn. & E. Nyman, Monsunia 1: 161.
1900.
Nectria striatospora Zimm., Centralbl. Bakteriol. Abt. 1, 7: 105.
1901.
Cylindrocarpon victoriae Wollenw., Z. Parasitenk. (Berlin) 1: 161.
1928.
Nectria azureo-ostiolata Yoshim. Doi, Mem. Nat. Sci. Mus. Tokyo
10: 23. 1977.
Authentic material: In B fide Wollenweber, Fusaria Autogr. Delin.
1: 66. 1916.
Original locality: Cameroon.
Original substrate: Trunk of an unknown tree.
vinosum Fusarium Massee, Brit. Fung.-Fl. 3: 479. 1893.
(See Fusarium flocciferum)
Holotypus: ?K(M).
Type locality: UK.
FUSARIUM
REDELIMITED
Type substrate: Decaying mast manufactured from Fagus
sylvatica.
Note: Synonym fide Wollenweber & Reinking (1935).
Notes: Synonyms fide Wollenweber & Reinking (1935). No holotype specimen could be located and therefore an illustration is
designated as lectotype.
vinosum Fusarium Greco, Origine des Tumeurs (Etiologie du
Cancer. etc.) et Observations de Mycoses (Blastomycoses. etc.)
Argentines (Buenos Aires): 670. 1916, nom. illegit., Art. 53.1.
Authentic material: Not located.
Original locality: Argentina.
Original substrate: Homo sapiens.
Note: A late homonym of F. vinosum Massee.
vogelii Fusarium Henn., Z. Pflanzenkrankh. 12: 16. 1902.
Synonyms: Septosporium curvatum Rabenh. & A. Braun,
Krankh. Pfl.: 14. 1854.
Septoria curvata (Rabenh. & A. Braun) Sacc., Syll. Fung. 3: 484.
1884.
Cercospora curvata (Rabenh. & A. Braun) Wollenw., Fusaria
Autogr. Delin. 1: 451. 1916.
Holotypus: In B (Kabat & Bubak, Fungi Imp. Exs. 248) fide Hein
(1988).
Type locality: Poland, Dąbroszyn (former Tamsel).
Type substrate: Leaf of Robinia pseudoacacia.
Notes: Status unclear. Neither Fusarium fide Wollenweber &
Reinking (1935) nor Cercospora fide Chupp (1954).
violaceum Fusarium P. Crouan & H. Crouan, Fl. Finistere: 14.
1867, nom. illegit., Art. 53.1.
(See Fusarium sambucinum)
Authentic material: ?PC.
Original locality: France, Brittany, Finistere, marshes.
Original substrate: Bark of unknown tree.
Notes: An illegitimate homonym of F. violaceum Fuckel (1863).
Synonym fide Gams et al. (1997).
violaceum Fusarium Fuckel, Fungi Rhen. Exs. No. 209. 1863.
(See Fusarium caeruleum)
Syntypes: In BPI, F, HAL, MICH, S & WSP (Fuckel, Fungi Rhen.
Exs. No. 209).
Type locality: Germany, Hessen, Oestrich.
Type substrate: Solanum tuberosum.
Note: Synonym fide Wollenweber & Reinking (1935) and Booth
(1971).
violae Fusarium F.A. Wolf, Mycologia 2: 21. 1910.
(See Fusarium oxysporum)
Holotypus: Not located.
Type locality: USA, Nebraska, Lincoln.
Type substrate: Stems and roots of Viola tricolor.
Note: Synonym fide Wollenweber & Reinking (1935).
virguliforme Fusarium O'Donnell & T. Aoki, Mycologia 95: 667.
2003.
(See Fusarium azukicola)
Holotypus: BPI 841956.
Ex-type culture: MAFF 238553 = NRRL 31041 = Shuxian Li # 95.
Type locality: USA, Illinois.
Type substrate: Glycine max.
Descriptions and illustrations: See Aoki et al. (2003).
Diagnostic DNA barcodes: rpb1: JX171530; rpb2: JX171643;
tef1: AY220193.
viride Fusarium (Lechmere) Wollenw., Fusaria Autogr. Delin. 1:
418. 1916.
Basionym: Pionnotes viridis Lechmere, Compt. Rend. Hebd.
Seances Acad. Sci. 155: 178. 1912.
(See Fusarium solani)
Holotypus: Not located.
Type locality: Ivory Coast.
Type substrate: Undetermined wood.
Note: Synonyms fide Wollenweber & Reinking (1935).
viticola Fusarium Thüm. (as ‘viticolum’), Pilze Weinst.: 52. 1878.
Synonym: Fusarium herbarum var. viticola (Thüm.) Wollenw.,
Fusaria Autogr. Delin. 3: 898. 1930.
(See Fusarium avenaceum)
Lectotypus (hic designatus, MBT 10000756): Italy, Liguria,
Genoa, Rapallo, dry twigs of Vitis vinifera, Jul. 1876, G. Passerini, in Thümen, Pilze Weinst. 1878: pl. 3, fig. 3.
www.studiesinmycology.org
volatile Fusarium Al-Hatmi et al., Fungal Syst. Evol. 4: 174.
2019.
Holotypus: CBS H-24004.
Ex-type culture: CBS 143874.
Type locality: French Guiana, Cayenne.
Type substrate: Bronchoalveolar lavage effusion from Homo
sapiens with lung infection.
Descriptions and illustrations: See Al-Hatmi et al. (2019).
Diagnostic DNA barcodes: rpb2: LR596006; tef1: LR596007.
volutella Fusarium Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 43: 93. 1891.
(See Fusarium stromaticum)
Holotypus: Langlois 1505 in NY fide Index Fungorum.
Type locality: USA, Louisiana, Saint Martinsville.
Type substrate: Dead twigs of Nekemias arborea (syn. Ampelopsis arborea).
Note: Synonym fide Wollenweber & Reinking (1935) and
Gr€afenhan et al. (2011).
vorosii Fusarium B. Toth et al., Fungal Genet. Biol. 44: 1202.
2007.
Holotypus: BPI 871658.
Ex-type culture: NRRL 37605.
Type locality: Hungary, Pest, Ipolydamasd.
Type substrate: Spikelet of Triticum aestivum.
Descriptions and illustrations: See Starkey et al. (2007).
Diagnostic DNA barcodes: rpb1: KM361647; rpb2: KM361665;
tef1: DQ459745.
waltergamsii Fusarium O'Donnell et al., Index Fungorum 440: 5.
2020.
Neocosmospora gamsii Sand.-Den. & Crous, Persoonia 41:
116. 2018.
Holotypus: CBS H-23226.
Ex-type culture: CBS 143207 = NRRL 32323 = UTHSC 99-250.
Type locality: USA, Pennsylvania.
Type substrate: Bronchoalveolar lavage fluid from Homo
sapiens.
Descriptions and illustrations: See Sandoval-Denis & Crous
(2018).
Diagnostic DNA barcodes: rpb1: MW834223; rpb2: KM361665;
tef1: DQ246951.
werrikimbe Fusarium J.L. Walsh, L.W. Burgess, E.C.Y. Liew &
B.A. Summerell, sp. nov. MycoBank MB 837725.
173
CROUS
ET AL.
Synonym: Fusarium werrikimbe J.L. Walsh, L.W. Burgess, E.C.Y.
Liew & B.A. Summerell, Fungal Diversity 44: 155. 2010, nom.
inval., Art. 40.7.
Etymology: In reference to Werrikimbe National Park, the
geographic origin of the isolates first recognised as belonging to
this species.
For diagnosis see Walsh et al., Fungal Diversity 44: 155. 2010.
Holotypus: CBS 125535 (preserved as metabolically inactive
culture).
Ex-type culture: CBS 125535 = F19350 = RBG 5332.
Type locality: Australia, New South Wales, Werrikimbe National
Park.
Type substrate: Sorghum leiocladum.
Descriptions and illustrations: See Walsh et al. (2010).
Diagnostic DNA barcodes: rpb1: MW928821; rpb2: MN534304;
tef1: MW928846.
Notes: Walsh et al. (2010) did not indicate the holotype for
F. werrikimbe, rendering the name invalid (Art. 40.7). Here we
validate the name.
willkommii Fusarium Lindau, Rabenh. Krypt.-Fl. ed. 2, 1(9): 551.
1910.
Replaced synonym: Fusarium candidum Sacc. & D. Sacc., Syll.
Fung. 18: 674. 1906, nom. illegit., Art. 53.1, non Fusarium
candidum Ehrenb. 1818.
Lectotypus (hic designatus, MBT 10000757): Germany, Saxony,
Fagus sylvatica, 1866, M. Willkomm, in Die mikroskopischen
Feinde des Waldes 1, Tab. VI, figs 11–12.
Notes: Lindau's description of F. willkommii was based on
Willkomm's (1866: 103) description and illustration under the
name Fusidium candidum Link as well as Saccardo's (l.c.)
description under Fusarium candidum. Therefore, the illustration
by Willkomm (1866) is designated as lectotype.
sic et al., Antonie van Leeuwitzenhausenense Fusarium Si
wenhoek 111: 1795. 2018.
(See Fusarium stercicola)
Holotypus: CBS H-23351.
Ex-type culture: CBS 142480 = DSM 106212.
Type locality: Germany, Hessen, Witzenhausen, NeuEichenberg.
Type substrate: Branch of Hibiscus sp.
sic et al. (2018a).
Descriptions and illustrations: See Si
Diagnostic DNA barcodes: rpb1: MG237865; rpb2: LR583886;
tef1: KY556525.
wolgense Fusarium Rodigin, Trudy Bashkir. Sel’. Khoz. Inst. 3:
101. 1942.
Holotypus: Not located.
Type locality: Russia, Volgograd (formerly Stalingrad).
Type substrate: Fruit of Citrullus lanatus (syn. Citrullus vulgaris).
Notes: Status unclear. Not treated by either of Booth (1971) and
Gerlach & Nirenberg (1982).
wollenweberi Fusarium Raillo, Fungi of the Genus Fusarium:
189. 1950, nom. illegit., Art. 52.1.
(See Fusarium anthophilum)
Authentic material: Not located.
Original locality: Azerbaijan.
Original substrate: Seeds and stems of Gossypium sp.
Descriptions and illustrations: See Raillo (1950).
Notes: Fusarium wollenweberi was published as a new combination, but no basionym was indicated. As a nomen novum, it
can only be based on F. anthophilum, the only cited name, which
174
is a valid name. Therefore, F. wollenweberi would be illegitimate
(nom. superfl., Art. 52.1.). Additionally, the condition for the
introduction of a new species is also not met as a Latin diagnosis, necessary in 1950, is lacking.
xiangyunense Fusarium F. Zhang et al. (as ‘xiangyunensis’),
Phytotaxa 450: 278. 2020, nom. inval., Art. 40.8.
(See Fusarium stercicola)
Authentic material: DLU11-1, School of Agriculture and Biology,
Dali University, China.
Authentic culture: CGMCC 3.19676.
Original locality: China, Yunnan, Xiangyun, Dali, Da-bo-na hotspring.
Original substrate: Waterlogged soil.
Descriptions and illustrations: See Zhang et al. (2020).
Diagnostic DNA barcodes: rpb1: MH999281; tef1: MH992629.
Note: Based on phylogenetic and morphological evidence provided by Zhang et al. (2020), this invalid name (Art. 40.8) belongs
to the genus Neocosmospora and is a synonym of N. stercicola.
xylarioides Fusarium Steyaert, Bull. Soc. Roy. Bot. Belgique
80: 42. 1948.
Synonyms: Gibberella xylarioides (Steyaert) R. Heim & Saccas,
Rev. Mycol. (Paris) 15 (Suppl. Colon.): 97. 1950.
Fusarium oxysporum f. xylarioides (Steyaert) Delassus, Bull. sci.
Minist. Colon., Sect. Agric. trop. 5: 347. 1954.
Lectotypus (hic designatus, MBT 10000758): Central African
Republic, Bangui, trunk of Coffea excelsa, 1939, H. Frederic, in
Steyaert, Bull. Soc. Roy. Bot. Belgique 80, pl. I, fig. 8.
Epitypus (hic designatus, MBT 10001275): Ivory Coast, on trunk
of Coffea sp., Feb. 1951, C. & M. Moreau, CBS 258.52 (preserved as metabolically inactive culture).
Ex-epitype culture: CBS 258.52 = NRRL 25486.
Descriptions and illustrations: See Steyaert (1948), Booth
(1971), Gerlach & Nirenberg (1982) and Geiser et al. (2005).
Diagnostic DNA barcodes: rpb1: JX171517; rpb2: JX171630;
tef1: AY707136.
Notes: A lectotype is designated here based on an illustration
provided by Steyaert (1948) accompanying the original protologue. All attempts to locate the holotype specimen lodged at the
Universite de Bangui (BANG), Central African Republic, as
indicated by Steyaert (1948), failed. In addition, an epitype (CBS
258.52) is designated here to provide taxonomic stability for this
important species.
xyrophilum Fusarium I. Laraba et al., Mycologia 112: 45. 2019
[2020].
Holotypus: BPI 910919.
Ex-type culture: FRC M-8921 = NRRL 62721.
Type locality: Guyana, Cuyuni-Mazaruni, Kamakusa Mountain.
Type substrate: Xyris surinamensis.
Descriptions and illustrations: See Laraba et al. (2020).
Diagnostic DNA barcodes: rpb1: MN193933; rpb2: MN193905;
tef1: MN193877.
yamamotoi Fusarium O'Donnell et al., Index Fungorum 440: 5.
2020.
Replaced synonym: Nectria elegans W. Yamam. & Maeda,
Hyogo Univ. Agric. ser. Agric. Biol. 3: 15. 1957, non Fusarium
elegans Appel & Wollenw. 1910.
Neocosmospora elegans (W. Yamam. & Maeda) Sand.-Den. &
Crous, Persoonia 43: 127. 2019.
Lectotypus: Figs 1–9, page 16, in Yamamoto et al. (1957),
designated in Sandoval-Denis et al. (2019).
FUSARIUM
Epitypus: CBS H-23980, designated in Sandoval-Denis et al.
(2019).
Ex-epitype culture: ATCC 42366 = CBS 144396 = MAFF
238541 = NRRL 22277 = SUF XV-1.
Type locality: Japan.
Type substrate: Twigs and trunks of Zanthoxylum piperitum.
Descriptions and illustrations: See Sandoval-Denis et al. (2019).
Diagnostic DNA barcodes: rpb1: MW218113; rpb2: FJ240380;
tef1: AF178336.
yuccae Fusarium Cooke, Grevillea 7: 34. 1878, nom. inval., Art.
36.1(a).
(See Fusarium lateritium)
Authentic material: BPI 453149.
Original locality: USA, South Carolina, Aiken.
Original substrate: Yucca aloifolia.
Note: Synonym fide Wollenweber & Reinking (1935).
zanthoxyli Fusarium X. Zhou et al., Mycologia 108: 675. 2016.
Holotypus: HMNWAFU XZ-Fyzs133-20130408
Ex-type culture: CBS 140838 = NRRL 66285.
Type locality: China, Shaanxi, Tongchuan, Yaozhou, Sunyuan.
Type substrate: Zanthoxylum bungeanum.
Descriptions and illustrations: See Zhou et al. (2016).
Diagnostic DNA barcodes: rpb1: KM520383; rpb2: KM236763;
tef1: KM236703.
zavianum Fusarium (Sacc.) Sacc., Syll. Fung. 4: 709. 1886.
Basionym: Fusisporium zavianum Sacc., Michelia 1: 83. 1877.
(See Fusarium lateritium)
Holotypus: In PAD.
Type locality: Italy, Vittorio.
Type substrate: Vitis vinifera.
Note: Synonyms fide Wollenweber & Reinking (1935).
zeae Fusarium (Westend.) Sacc., Syll. Fung. 4: 713. 1886.
Basionym: Fusisporium zeae Westend., Bull. Acad. Roy. Sci.
Belgique, Cl. Sci. 18: 414. 1852. (non Fusisporium zeae Roum.,
Rev. Mycol. (Toulouse) 6: 163. 1884).
(See Fusarium avenaceum)
Holotypus: BR5020141668483.
Type locality: Belgium, Kortrijk railway station.
Type substrate: Rotting stalks of Zea mays.
Note: Synonyms fide Wollenweber & Reinking (1935).
zealandicum Fusarium Nirenberg & Samuels, Canad. J. Bot. 78:
1483. 2000.
Geejayessia zealandica (Cooke) Schroers, Stud. Mycol. 68:
133. 2011.
Basionym: Nectria zealandica Cooke, Grevillea 8: 65. 1879.
Synonyms: Cucurbitaria zelandica (Cooke) Kuntze, Revis. Gen.
Pl. 3: 462. 1898.
Cosmospora zealandica (Cooke) Samuels & Nirenberg, Canad.
J. Bot. 78: 1483. 2000.
Holotypus: BPI 747915.
Ex-type culture: BBA 64792 = CBS 111.93.
Type locality: New Zealand, Auckland, Waitakere Ranges
Regional Park, Cascades Kauri.
Type substrate: Bark of Hoheria populnea.
Descriptions and illustrations: See Nirenberg & Samuels (2000).
Diagnostic DNA barcodes: rpb2: HM626684; tef1: HQ728148.
ziziphinum Fusarium Pass., Erb. Critt. Ital. ser. 2 no. 1084. 1881.
(See Fusarium lateritium)
Syntype: F 982523 ( Erb. Critt. Ital. no. 1048).
www.studiesinmycology.org
REDELIMITED
Type locality: Italy.
Type substrate: Twigs of Ziziphus sinensis (syn. Ziziphus jujuba).
Note: Synonym fide Wollenweber & Reinking (1935).
zonatum Fusarium (Sherb.) Wollenw., Fusaria Autogr. Delin. 1:
392. 1916.
Basionym: Fusarium lutulatum var. zonatum Sherb., Mem.
Cornell Univ. Agric. Exp. Sta. 6: 214. 1915.
(See Fusarium oxysporum)
Typus: ?CUP-007453.
Type locality: USA, New York, Ithaca.
Type substrate: Solanum tuberosum.
Notes: Synonym fide Wollenweber & Reinking (1935). Lectotypification pending study of the material lodged in CUP.
zygopetali Fusarium Delacr., Bull. Soc. Mycol. France 13: 103.
1897.
Holotypus: ?PC.
Type locality: France, Paris, Luxembourg gardens.
Type substrate: Leaves of Zygopetalum maculatum (syn.
Zygopetalum mackayi).
Notes: Status unclear. Not Fusarium fide Wollenweber &
Reinking (1935).
CONCLUSIONS
The present study is the first to provide an up-to-date morphological, biochemical, and phylogenetic overview of the 20
fusarioid genera that are presently recognised in Nectriaceae.
Morphological species recognition frequently fails to distinguish
fusarioid taxa that have been described based on genealogical
concordance phylogenetic species recognition (GCPSR sensu
Taylor et al. 2000). To address this issue, we have established a
new database, Fusarioid-ID, with accurate names for species
and genera of fusarioid taxa. Although the phylogenetically most
informative genes remain tef1, rpb1 and rpb2, additional markers
such as act1, CaM, tub2, ITS and LSU are also incorporated.
These genetic fragments can be amplified by PCR and
sequenced using the primers indicated in Table 2. In the future,
new species and other phylogenetically informative orthologous
genes, will be added to resolve isolates at species and genus
level. Researchers interested in obtaining reference strains
should contact the Westerdijk Fungal Biodiversity Institute
(https://wi.knaw.nl/page/Collection), which houses a large
collection of phylogenetically diverse fusarioid taxa.
As we have shown here, the phylogenetically derived argument that species under the node F1 should be considered
members of “Fusarium” is not practical, as this circumscription
would lead to a genus without apparent synapomorphies, as
lineages outside the genus would also share its characteristics.
However, the F3 node (corresponding to Fusarium s. str.) is
resolved by all genetic markers so far analysed (e.g., see Geiser
et al. 2021) and delineates the morphologically, ecologically, and
biochemically well-delineated genus Fusarium.
Fusarium s. str. does not have different sexual morphs, other
than Gibberella. Fusarioid genera are not only morphologically
distinct, but as we have shown in this study, correlate to different
monophyletic groups and also differ in their biology and mycotoxin profiles.
One of the reasons for the desire to classify any species
producing conidia with foot-shaped basal cells into a single
genus could be that plant pathologists and clinicians typically
175
CROUS
ET AL.
isolate conidia or obtain cultures from vegetative mycelium that
inhabits their specimens. Also, Wollenweber and his successors
may have primarily worked with vegetatively proliferating materials, although it was also Wollenweber (1924, 1926) who produced the first general synopsis of holomorphs in the
Hypocreales. However, mainly Joan M. Dingley (1951, 1957),
Colin Booth (1959), and especially Gary J. Samuels (Samuels
1976a, b, 1978, 1988, Samuels et al. 1991) significantly
changed our points of view by systematically isolating ascospores obtained from ascomata, of which a vast majority were
not gathered in agricultural fields but from woody or herbaceous
substrata in forests of pantropical, species-rich regions. The
result of their taxonomic considerations was an infrageneric
subgrouping system in Nectria that was based on sexual and
asexual connections. The classification of species according to
morphological similarities in sexual morphs allowed understanding patterns of asexual characteristics that are unique for
the sexually defined subgroups and eventually correlating sexual
groupings with Wollenweber's section system. The diversity of
nectria-like species Samuels looked at is huge and was eventually interpreted on the level of families, within which numerous
genera were recognised or newly described (Rossman et al.
1999) with infrageneric, informal species groups of Nectria
accepted at the genus level (e.g., see Chaverri et al. 2011 and
subsequent studies). Applying the generic level to the numerous
nectria-like subgroups producing fusarioid conidia is therefore
another small but unavoidable step towards a taxonomic system
that allows distinguishing natural diversity above the species
level based on morphologically and phylogenetically well-defined
units.
When Colin Booth delivered his Presidential address to the
British Mycological Society in 1977, he chose the title “Do you
believe in genera?”. He addressed this topic based on his
interpretation of Nectriaceae (Booth 1978). Booth subsequently
showed that several “groups” of species formed fusarioid
asexual morphs, namely Gibberella (now Fusarium s. str.),
Haematonectria (now Neocosmospora), Nectria episphaeria
(now Cosmosporella and Dialonectria), and Calonectria rigidiuscula (now Albonectria). Booth concluded that the “fusarium
morphs” reflected “terms of convenience” rather than genealogical relationships. In moving to the one fungus = one name
nomenclature (Hawksworth et al. 2011, Wingfield et al. 2012),
Fusarium s. str. was chosen over Gibberella (Gr€afenhan et al.
2011, Schroers et al. 2011, Rossman et al. 2013). As the genus
Fusarium was thus clearly well-defined, other Nectriaceae lineages with a fusarium-like morphology were recognised
(Gr€afenhan et al. 2011, Schroers et al. 2011, Lombard et al.
2015, Lechat & Fournier 2015). As we have shown here,
taxa are constantly being newly collected and added to the
phylogeny of Nectriaceae. The only stable option forward is to
apply and use the genus name Fusarium (= Gibberella) as
more precisely defined based on its own monophyletic node as
presented here (F3), supported by morphology, biochemistry,
and biology.
DISCLAIMER
The present paper represents a separate initiative to Geiser et al.
(NSF 1655980): A phylogenetic revisionary monograph of the
genus Fusarium.
176
ACKNOWLEDGEMENTS
The authors are grateful for the open discussions with David Geiser and Kerry
O'Donnell who have also shared their datasets for comparison in this study, as
well as several authors listed on their recent papers. Uwe Braun is thanked for his
contributions in compiling the list of Fusarium names, and checking the names
published on numerous old fungarium exsiccatae sheets.
APPENDIX A. SUPPLEMENTARY DATA
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.simyco.2021.100116.
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