Abstract
The species-rich Fusarium sambucinum species complex (FSAMSC; Fusarium, Nectriaceae, Hypocreales) is well-known for including devastating plant pathogens and toxigenic species. However, this group of grass-loving fungi also accommodates soil saprobes, endophytes, mycoparasites and rare opportunistic pathogens of humans and other animals. Recent publications have highlighted the vast phylogenetic and biochemical diversity of the FSAMSC, although a large number of taxa in FSAMSC have not been systematically described and still lack Latin binomials. In this study we established the phylogenetic breadth of the FSAMSC using an integrative approach including morphological, multilocus phylogenetic, and coalescence analyses based on five gene regions (calmodulin, RNA polymerase II largest and second largest subunits, translation elongation factor 1-α, and β-tubulin). Results obtained support the recognition of 75 taxa in FSAMSC, including all the currently known species segregates of the Fusarium head-blight pathogen F. graminearum s. lat. Thirty novel species are formally described and illustrated, while four phylogenetic species remain undescribed. An epitype is proposed for the generic type of Fusarium, F. sambucinum, from recently collected material identified by means of morphology, phylogenetics and mating experiments, fixing the phylogenetic application of the name. Additional notes are included on the typification of Fusisporium cerealis (syn. Fusarium cerealis).
Taxonomic novelties: New species: Fusarium agreste Sand.-Den., J.Z. Groenew. & Crous, Fusarium amblysporum Sand.-Den., M.M. Costa, Fusarium bananae Sand.-Den., M.M. Costa, Fusarium bellum Sand.-Den., J.Z. Groenew. & Crous, Fusarium brachypes Sand.-Den., J.Z. Groenew. & Crous, Fusarium carinatum Sand.-Den., J.Z. Groenew. & Crous, Fusarium cultriforme Sand.-Den., M.M. Costa, Fusarium cuspidatum Sand.-Den., J.Z. Groenew. & Crous, Fusarium cygneum Sand.-Den., J.Z. Groenew. & Crous, Fusarium dimorphosporum Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, Fusarium dolichosporum Sand.-Den., J.Z. Groenew. & Crous, Fusarium gladiolum Sand.-Den., J.Z. Groenew. & Crous, Fusarium hamatum Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, Fusarium leptum Sand.-Den., J.Z. Groenew. & Crous, Fusarium longicolle Sand.-Den., J.Z. Groenew. & Crous, Fusarium magnum Sand.-Den., J.Z. Groenew. & Crous, Fusarium mastigosporum Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, Fusarium minutum Sand.-Den., J.Z. Groenew. & Crous, Fusarium mucronatum Sand.-Den., J.Z. Groenew. & Crous, Fusarium parabolicum Sand.-Den., J.Z. Groenew. & Crous, Fusarium platysporum Sand.-Den., J.Z. Groenew. & Crous, Fusarium pratense Sand.-Den., J.Z. Groenew. & Crous, Fusarium procumbens Sand.-Den., J.Z. Groenew. & Crous, Fusarium pseudolongipes Sand.-Den., J.Z. Groenew. & Crous, Fusarium sagittatum Sand.-Den., J.Z. Groenew. & Crous, Fusarium seculiforme Sand.-Den., J.Z. Groenew. & Crous, Fusarium subcylindroides Sand.-Den., J.Z. Groenew. & Crous, Fusarium symmetricum Sand.-Den., J.Z. Groenew. & Crous, Fusarium tropicale Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, Fusarium vermicularioides Sand.-Den., J.Z. Groenew. & Crous. Epitype: Fusarium sambucinum Fuckel.
Citation: Sandoval-Denis M, Costa MM, Broders K, Becker Y, Maier W, Yurkov A, Kermode A, Buddie AG, Ryan MJ, Schumacher RK, Groenewald JZ, Crous PW (2024). An integrative re-evaluation of the Fusarium sambucinum species complex. Studies in Mycology 110: 1–110 doi: 10.3114/sim.2025.110.01
Keywords: Coalescence, fungi, novel species, pathogens, phylogenetics, systematics, taxonomy
INTRODUCTION
The Fusarium sambucinum species complex (FSAMSC) includes a morphologically and phylogenetically diverse, globally distributed assemblage of species. They commonly occur in cultivated and undisturbed soils, but are also known from disparate substrates, including air, animal dung, clinical samples (animals, and rarely from human specimens), food and feed, mostly as storage spoilers, but also used as food itself (O’Donnell et al. 1998a); insects, other fungi, and several plant hosts (Leslie & Summerell 2006, Domsch et al. 2007, Al-Hatmi 2016, O’Donnell et al. 2021). The range of known plant hosts for members of FSAMSC spans over 400 species, distributed in 263 genera, and 79 families; however, a clear predilection for grass hosts (Poaceae) is well documented (Farr et al. 2023). Likewise, species in FSAMSC are associated with a variety of lifestyles including animal associates and rare opportunistic pathogens, endophytes, mycoparasites, plant associates, pathogens and saprobes (Domsch et al. 2007, Hill et al. 2022).
Their role as plant pathogens is certainly the most relevant as FSAMSC includes F. graminearum, the main agent of Fusarium head blight of cereals, associated with epidemics and billions of dollars in wheat and barley yield and quality losses (O’Donnell et al. 2000, McMullen et al. 2012). Other symptoms on plant hosts attributed to species in FSAMSC include canker, branch and stub dieback on woody hosts; brown patch of turf, decline, leaf spots, rots (central bud rot of flowering plants, cob, crown, ear, foot, fruit, head, root, stalk, tuber, and storage rots, including the most threatening agent of potato dry rot, F. sambucinum), seedling blight, silver top or white head of grasses, and wilting (Leslie & Summerell 2006, Domsch et al. 2007, Tiwari et al. 2020).
The agricultural relevance of FSAMSC is not limited to their direct effects on crops, since it also encompasses mycotoxin producers, with well-studied biochemical profiles, and subjected to active surveillance programs to limit its consumption by humans and livestock (Nelson et al. 2004, Desjardin & Proctor 2007, Laraba et al. 2021). Among the fungal genera with known capacity to produce trichothecenes (e.g.,“Cephalosporium”, “Cylindrocarpon”, Fusarium, Myrothecium, Stachybotrys, Trichoderma, Trichothecium, Verticimonosporium, among others; CAST 2003, Polak-Śliwińska & Paszczyk 2021), Fusarium, and particularly the FSAMSC, produces the widest diversity of trichothecenes (i.e., calonectrin and its metabolites; diacetoxyscirpenol, neosolaniol, nivalenol, T2 toxin and derived metabolites; 3-acetyldeoxynivalenol, 7-hydroxy isotrichodermol, isotrichodermol, deoxynivalenol, fusarenon X, iso-neosolaniol, HT-2 toxin, scirpentriol, 4-acetylnivalenol, 4,15-diacetylnivalenol, 4-monoacetoxyscirpenol, 8-acetylneosolaniol, 15-acetoxyscirpenol, 5-monoacetoxyscirpenol, and 15-acetyldeoxynivalenol) (CAST 2003, Desjardin & Proctor 2007, Laraba et al. 2021, and references therein). Other mycotoxin classes produced by FSAMSC include the depsipeptides (beauvericin, and enniatins), lactones (butenolide, zearalenone), polyketides (fusarin C), polyphenols (bikaverin); and the sesquiterpene culmorin and its derivatives (CAST 2003, Desjardin & Proctor 2007, Laraba et al. 2021, and references therein).
By including ubiquitous and devastating plant pathogens, and prolific mycotoxin producers, the FSAMSC stands out as one of the most important species complexes within Fusarium. Moreover, because it contains the conserved generic type of Fusarium (F. sambucinum), and also that of the sexually typified Gibberella (G. pulicaris) (Gams et al. 1997, Gams 1999), the FSAMSC is of central relevance for Fusarium taxonomy. The systematics and taxonomy of the FSAMSC, however, have suffered from the same problems inherent to Fusarium, and serves as a reference frame to understand the conflictive taxonomical history of the genus, as well as an example of the polyphyly of the artificial sectional classification system of the genus Fusarium as demonstrated using molecular phylogenetic methods (O’Donnell 1993, 1996, 1997, O’Donnell et al. 2013).
Currently, the FSAMSC includes elements formerly classified in the Fusarium sections Arthrosporiella, Discolor, Gibbosum, and Sporotrichiella. The largest component of the current FSAMSC belongs to the former Fusarium section Discolor (characterized by thick-walled, distinctly septate macroconidia, with beaked apices and pedicellate basal cells; Wollenweber 1913) which originally included F. rubiginosum (syn. F. culmorum), F. discolor var. sulphureum (syn. F. sambucinum), and F. trichothecioides. The short-lived section Neesiola (Wollenweber 1917), (characterized by pedicellate, slender macroconidia: F. heterosporum and F. reticulatum), was reduced to a subsection of Discolor (Wollenweber 1918); however, both species mentioned above are not related to the FSAMSC, being now classified elsewhere [F. heterosporum SC (FHSC) and F. tricinctum SC (FTSC), respectively; O’Donnell et al. 2013, Crous et al. 2021]. Section Saubinetti (characterized by wide conidia; Wollenweber 1917) included F. culmorum, F. flocciferum (now in FTSC); F. graminearum, the obscure name F. macroceras; F. sambucinum with all its varieties and formae, F. sublunatum (now in F. buharicum SC, FBSC), and F. tumidum (syn. Neocosmospora ambrosia) and were combined with subsection Erumpens (species with tuberculate stroma, covered in sporodochia: F. congoense (syn. F. heterosporum); F. sambucinum var. coeruleum (syn. F. torulosum in the FTSC), and the obscure name F. polymorphum var. pallens), and together with section Trichothecioides (species with non-pedicellate macroconidia: F. bactridioides, currently in the F. fujikuroi SC; and F. trichothecioides (Wollenweber 1917), were also reduced to subsections of Discolor, but were posteriorly merged into subsection Eudiscolor (macroconidia with short apical cells with nipple-like tips) by Raillo (1950), who also included F. buharicum (now in FBSC) and F. gigas (later transferred to section Macroconia, now a synonym of Macroconia papilionacearum; Crous et al. 2021, Visagie et al. 2024), while subsection Saubinetii was redefined to include species with spindle- to sickle-shaped macroconidia with gradual and evenly narrowed apical cells (F. graminearum, F. flocciferum, F. heterosporum, and F. macroceras).
Gordon (1952, 1954a, b), Booth (1971), and Gerlach & Nirenberg (1982) largely followed Wollenweber’s classification. Booth (1971) emended Raillo’s two subsections system, subsection Eudiscolor characterized by broad, curved macroconidia with short apical cells, narrowing to a nipple-like tip, and punctiform sporodochia in culture, which later merged among masses of macroconidia; and subsection Saubinetii with spindle- or sickle-shaped macroconidia, with elongated and evenly narrowed apical cells, mostly borne on lateral conidiophores and sparse sporodochia.
Sections Lateritium (intercalary chlamydospores, pale to rose colonies, and blue erumpent sclerotia; Wollenweber 1917) and Gibbosum (conidiogenous cells are simple phialides, microconidia absent, macroconidia with a tendency to attenuated apical cells, abundant chlamydospores in chains, clumps, or solitary, pale brown at maturity; Wollenweber 1913) were also synonymized with section Discolor (Bilai 1955). However, the former section Lateritium and most species previously accepted in section Gibbosum currently stand on their own ground as the F. lateritium (FLSC) and F. incarnatum-equiseti (FIESC) species complexes, respectively (O’Donnell et al. 2013, Xia et al. 2019, Costa et al. 2024). Nevertheless; F. longipes, previously a synonym of F. equiseti (FIESC) in the sense of Booth (1971), belongs to the FSAMSC (O’Donnell et al, 2013, Crous et al. 2021, Laraba et al. 2021).
Section Sporotrichiella (microconidia globose, pyriform to clavate, sickle-shaped macroconidia and chlamydospores; Lewis 1913) included F. fusarioides (syn. F. chlamydosporum, now in F. chlamydosporum SC, FCSC), F. poae, F. sporotrichioides, and F. tricinctum (now in the FTSC), to which later Bilai (1955) added F. sarcochroum (now in FLSC) and F. sporotrichiella (syn. F. sporotrichioides). While section Arthrosporiella (microconidia short and broad, spindle-shaped; Sherbakoff 1915) included F. anguioides and varieties (currently in the F. concolor SC, FCOSC), F. arthrosporioides (now in FTSC), F. biforme (syn. F. avenaceum, now in FTSC), F. diplosporum (syn. F. sarcochroum), and F. diversisporum (now in FTSC). Booth (1971) reaccommodated the species above according to the nature of their conidiogenous cells into sections Sporotrichiella (monophialides: F. poae and F. tricinctum) and Arthrosporiella [polyblastic conidiogenous cells: F. avenaceum, F. camptoceras (now in FIESC), F. fusarioides (syn. F. chlamydosporum in FCSC), F. semitectum (syn. F. incarnatum, in FIESC), and F. sporotrichioides]. Gerlach & Nirenberg (1982) followed this classification, adding F. concolor (now in FCOSC).
The Snyder & Hansen (1945) system on the other hand, reduced all species and varieties in sections Arthrosporiella, Discolor, and Gibbosum into a single species, F. roseum (Booth 1971).
As obvious from its taxonomical history and diverse phenotypes, there is no single morphological character that defines all species in FSAMSC with respect to other Fusarium species complexes; nevertheless, it is well-defined phylogenetically (O’Donnell et al. 2013, Laraba et al. 2021). In its current phylogenetic framework, the taxonomy of the FSAMSC has been relatively stable for over a decade. The molecular systematics and typification of some important pathogenic or historical species have been recently revised (O’Donnell et al. 2013, Crous et al. 2021, Laraba et al. 2021), although, unresolved taxonomic and nomenclatural issues are yet to be settled, including the epitypification of the generic type, for which no living type material exists; and the circumscription of F. graminearum s. lat. and its 15 segregate species. While a considerable body of literature, particularly taxonomical studies, have followed the segregation of the different lineages of F. graminearum s. lat. (other than F. pseudograminearum) (Aoki & O’Donnell 1999a, b, O’Donnell et al. 2000, 2004, 2008b, Starkey et al. 2007, Yli-Mattila et al. 2009, Sarver et al. 2011, Aoki et al. 2012); the proposed subdivision of F. graminearum s. lat. has not been completely accepted, with disagreements regarding the recognition of F. graminearum s. lat. as a single, although phylogenetically diverse, biological species; or as separate incipient species (Bowden & Leslie 1999, Leslie & Summerell 2006, Leslie et al. 2007, Leslie & Bowden 2008, Trail 2009, McMallum et al. 2012, Gilbert & Haber 2013).
The objectives of this study were thus to revise past and corroborate recent phylogenetic hypotheses in the FSAMSC by providing Latin binomials, descriptions and illustrations for the numerous undescribed phylogenetic species residing therein, but also for poorly illustrated previously named taxa. In addition, the systematics of Fusarium graminearum s. lat. and the complete FSAMSC are tested using an integrative approach, reassessing the taxonomy and nomenclatural status of important pathogens and historical species.
MATERIALS AND METHODS
Fungal strains and fungarium specimens
A total 293 fungal strains were included in this study, obtained from five culture collections, namely the Julius Kühn Institute (JKI), former Biologische Bundesanstalt für Land- und Forstwirtschaft (BBA, Braunschweig, Germany), the Westerdijk Fungal Biodiversity Institute (WI) collection (CBS, Utrecht, The Netherlands), the Centre for Agriculture and Bioscience International (CABI) collection (IMI, Surrey, UK), the U.S. Agricultural Research Service collection (NRRL, Peoria, IL, USA), and the personal collection of P.W. Crous (CPC) held at WI (Table 1). Additionally, fungarium specimens were obtained from the fungarium of the WI (CBS H-, Utrecht, The Netherlands), the Conservatoire et Jardin Botaniques de la Ville de Genève (G, Geneva, Switzerland), and the Royal Botanic Gardens (K, Kew, UK). Fungal host ranges and geographic information for species treated in this study were compiled from data obtained from the US Department of Agriculture fungal databases (Farr et al. 2023), with additional entries obtained from the databases of the WI, Fusarium Research Centre (FRC) and NRRL Fusarium strain collections.
Table 1.
Fusarium cultures, collection details and GenBank accession numbers included in this study.
| Species name1 | Strain2 | Equivalent strain number2 | Substrate | Country | Sequence accession number3 | ||||
|---|---|---|---|---|---|---|---|---|---|
|
| |||||||||
| CaM | rpb1 | rpb2 | tef1 | tub2 | |||||
| F. acaciae-mearnsii | CBS 110253 | FRC R-9628 = MRC 2458 = NRRL 26752, CBS 110255 = FRC R-9630 = MRC 5122 = NRRL 26755 | Acacia mearnsii | South Africa | JAGDVM010000002 | JAGDVM010000003 | JAGDVM010000008 | AF212447 | AF212741 |
| CBS 110254T | FRC R-9629 = MRC 5120 = NRRL 26754 | Acacia mearnsii | South Africa | JAAWUD010000031 | JAAWUD010000100 | JAAWUD010000080 | AF212448 | AF212742 | |
| CBS 110255 | FRC R-9630 = MRC 5122 = NRRL 26755 | Acacia mearnsii | South Africa | JAGDVL010000002 | KM361640 | KM361658 | AF212449 | AF212743 | |
| CBS 123662 | FRC R-4412 = NRRL 34207 | Soil | Australia | JAGDVK010000158 | JAGDVK010000020 | JAGDVK010000031 | JAGDVK010000085 | JAGDVK010000015 | |
| F. aethiopicum | CBS 122858T | FRC R-10088 = NRRL 46726 | Triticum aestivum | Ethiopia | JAGDVJ010000004 | MW233298 | MW233470 | FJ240298 | FJ240288 |
| F. agreste | CBS 120993T | FRC R-8575 = MRC 6971 | Soil debris | South Africa | PQ240129 | PQ280828 | PQ274091 | PQ260805 | PQ273916 |
| NRRL 29296 | FRC R-9861 = ICMP 5173 | Sechium edule | New Zealand | - | MW233269 | MW233441 | MW233097 | - | |
| F. amblysporum | BBA 65418 | DSM 116523 | Solanum tuberosum | South Africa | PQ240130 | PQ280829 | PQ274092 | PQ260806 | PQ273917 |
| BBA 65419 | DSM 116524 | Soil debris | South Africa | PQ240131 | PQ280830 | PQ274093 | PQ260807 | PQ273918 | |
| BBA 71677 | DSM 116910 | Cucumis melo | Hungary | PQ240132 | PQ280831 | PQ274094 | - | PQ273919 | |
| CBS 128.95 | BBA 64568 | Triticum aestivum | Austria | PQ240133 | PQ280832 | PQ274095 | PQ260808 | PQ273920 | |
| CBS 148.95 | BBA 65893 | Spinacia oleracea | Sweden | PQ240134 | PQ280833 | PQ274096 | PQ260809 | PQ273921 | |
| CBS 151809T | BBA 64376 | Cucurbita maxima | New Zealand | PQ240135 | PQ280834 | PQ274097 | PQ260810 | PQ273922 | |
| CBS 151954 | NRRL 66929 = FRC R-8234 | Debris | South Africa | PQ240136 | MW233361 | MW233533 | MW233189 | PQ273923 | |
| NRRL 64078 | FRC R-8227 = KOD 2017 | Debris | South Africa | - | MW233360 | MW233532 | MW233188 | - | |
| F. armeniacum | CBS 485.94T | FRC R-9335 = IMI 352099 = MRC 6230 = NRRL 26908 = NRRL 25141 = NRRL 29133 | Triticum aestivum | Australia | PQ240137 | PQ280835 | PQ274098 | PQ260811 | PQ273924 |
| F. asiaticum | CBS 110256 | ATCC 28114 = FRC R-9620 = ATCC 46780 = NRRL 6101 | Hordeum vulgare | Japan | PQ240138 | PQ280836 | PQ274099 | HM744684 | PQ273925 |
| CBS 110257T | FRC R-5469 = MRC 1963 = NRRL 13818 | Hordeum vulgare | Japan | PQ240139 | MW233240 | JX171573 | AF212451 | AF212745 | |
| CBS 110258 | FRC R-9627 = DAOM 211601 = NRRL 26156 | Triticum sp. | China | JABFEQ010000060 | JABFEQ010000106 | JABFEQ010000057 | AF212452 | AF107856 | |
| CBS 110259 | FRC R-9402 = NRRL 28720 | Zea mays | Nepal | LHTZ01000026 | LHTZ01000010 | LHTZ01000031 | AF212453 | LHTZ01000163 | |
| F. austroamericanum | CBS 110244T | ATCC 46032 = FRC R-9634L = NRRL 2903 | Polypore | Brazil | JAAMOD010000093 | JAAMOD010000230 | JAAMOD010000315 | AF212438 | AF212733 |
| CBS 110245 | BBA 65928 = FRC R-9632 = GJS 90-246 = NRRL 28585 | Herbaceous vine | Venezuela | - | KM361643 | KM361661 | AF212439 | AF212734 | |
| CBS 110246 | FRC R-6964 = NRRL 28718 | Zea mays | Brazil | JAGDVG010000034 | JAGDVG010000002 | JAGDVG010000009 | PQ260812 | JAGDVG010000003 | |
| F. bananae | CBS 151937T | BBA 72296 = DSM 116929 | Musa sp. | Costa Rica | PQ240140 | PQ280837 | PQ274100 | PQ260813 | PQ273926 |
| F. bellum | CBS 151827T | FRC R-9513 = NRRL 66939 | Nematode cyst on Glycine max | China | PQ240141 | MW233389 | MW233561 | MW233217 | PQ273927 |
| NRRL 64095 | FRC R-9121 = KOD 2034 | Soil | China | - | MW233385 | MW233557 | MW233213 | - | |
| F. boothii | CBS 316.73T | ATCC 24372 = FRC R-9631 = IMI 160243 = NRRL 26916 | Zea mays | South Africa | PQ240142 | KM361641 | MN724983 | GQ915503 | GQ915437 |
| CBS 110251 | FRC R-9434 = NRRL 29105 = NRRL 29208 = NRRL 38122 | Zea mays | Nepal | JAGDVE010000007 | JAGDVE010000002 | JAGDVE010000014 | JAGDVE010000001 | JAGDVE010000006 | |
| CBS 119798 | MRC 4712 = NRRL 53893 | Triticum sp. | South Africa | PQ240143 | PQ280838 | PQ274101 | PQ260814 | PQ273928 | |
| CBS 119800 | MRC 6010 = NRRL 53895 | Zea mays | South Africa | PQ240144 | PQ280839 | PQ274102 | PQ260815 | PQ273929 | |
| F. brachygibbosum | BBA 71480 | DSM 116711 | Soil | Sudan | PQ240145 | PQ280840 | PQ274103 | PQ260816 | PQ273930 |
| BBA 72481 | DSM 116936 | Solanum lycopersicon | Sudan | PQ240146 | PQ280841 | PQ274104 | PQ260817 | PQ273931 | |
| CBS 334.75 | FRC R-9859 = NRRL 28448 | Musa sapientum | Turkey | PQ240147 | MW233264 | MW233436 | MW233093 | PQ273932 | |
| CBS 466.92 | NRRL 25806 | Phaseolus vulgare | Sudan | PQ240148 | PQ280842 | PQ274105 | PQ260818 | PQ273933 | |
| CBS 121682 | Stone | India | PQ240149 | PQ280843 | PQ274106 | PQ260819 | PQ273934 | ||
| CBS 131017 | Agropyron sp. | India | PQ240150 | PQ280844 | PQ274107 | PQ260820 | PQ273935 | ||
| CBS 131252 | Triticum sp. | India | PQ240151 | PQ280845 | PQ274108 | PQ260821 | PQ273936 | ||
| CPC 41845 | Soil | Russia | PQ240152 | PQ280846 | PQ274109 | PQ260822 | PQ273937 | ||
| CPC 45794 | W15 | Sea water | Israel | PQ240153 | PQ280847 | PQ274110 | PQ260823 | PQ273938 | |
| CPC 45801 | W53 | Sea water | Israel | PQ240154 | PQ280848 | PQ274111 | PQ260824 | PQ273939 | |
| NRRL 13829 | FRC R-6784 = MRC 2568 | River sediment | Japan | JABFET000000000 | JX171460 | JX171574 | JABFET010000489 | JABFET010000093 | |
| NRRL 20954T | BBA 64691 = DSM 116019 | Sorghum vulgare | India | PQ240155 | MW233246 | MW233418 | MW233075 | PQ273940 | |
| F. brachypes | CBS 151824T | FRC R-8938 = NRRL 66933 | Cultivated soil | Australia | PQ240156 | MW233376 | MW233548 | MW233204 | PQ273941 |
| NRRL 64096 | FRC R-9125 = KOD 2035 | Soil | Australia | - | MW233386 | MW233558 | MW233214 | - | |
| F. brasilicum | CBS 119179 | FRC R-10018 = NRRL 31238 | Hordeum vulgare | Brazil | JAGDVC010000002 | MW233276 | KM361663 | MW233104 | JAGDVC010000016 |
| CBS 119180T | FRC R-10019 = NRRL 31281 | Avena sativa | Brazil | JABCJS010000287 | JABCJS010000032 | JABCJS010000357 | MW233104 | JABCJS010000142 | |
| F. carinatum | CBS 151957T | FRC R-9061 = NRRL 66934 | Sorghum sp. | Nigeria | PQ240157 | MW233380 | MW233552 | MW233208 | PQ273942 |
| F. cerealis | CBS 135.80 | IMI 322101 = NRRL 36243 | Beta vulgaris | Netherlands | PQ240158 | PQ280849 | PQ274112 | PQ260825 | PQ273943 |
| CBS 623.85 | NRRL 36523 | Solanum tuberosum | Netherlands | PQ240159 | PQ280850 | PQ274113 | PQ260826 | PQ273944 | |
| CBS 832.85 | BBA 64545 = NRRL 25794 | Triticum durum | Germany | PQ240160 | PQ280851 | PQ274114 | PQ260827 | PQ273945 | |
| CBS 100101 | NRRL 36070 | Grass | France | PQ240161 | PQ280852 | PQ274115 | PQ260828 | PQ273946 | |
| CBS 110268 | ATCC 66007 = FRC R-09961 = NRRL 13721 | Solanum tuberosum | Poland | PQ240162 | PQ280853 | PQ274116 | PQ260829 | PQ273947 | |
| CBS 119874 | FRC R-09984 = MRC 8399 = KSU 11453 = Univ. Sydney F13169 | Unknown | Unknown | PQ240163 | PQ280854 | PQ274117 | PQ260830 | PQ273948 | |
| NRRL 13163T (ex-type of F. crookwellense) | FRC R-3090 | Soil | Australia | PQ240164 | PQ280855 | PQ274118 | PQ260831 | PQ273949 | |
| F. chaquense | CBS 151952 | NRRL 64072 = FRC R-8102 = KOD2011 = MRC 3936 | Avena sativa | South Africa | PQ240165 | MW233353 | MW233525 | MW233181 | PQ273950 |
| NRRL 64041 | FRC R-3766 = KOD 1980 | Soil | Australia | - | MW233317 | MW233489 | MW233145 | - | |
| NRRL 66748 | RC-J174 | Leersia luziola | Argentina | JAATNT010000291 | JAATNT010000102 | JAATNT010000016 | JAATNT010000651 | JAATNT010000118 | |
| NRRL 66749T | RC-J293 | Sorghastrum setosum | Argentina | JAATNU010000272 | JAATNU010000034 | JAATNU010000127 | JAATNU010000053 | JAATNU010000244 | |
| NRRL 66750 | RC-J1301 | Diplachne sp. | Argentina | JAATNV010000305 | JAATNV010000279 | JAATNV010000019 | JAATNV010000129 | JAATNV010000188 | |
| F. circinatum | CBS 405.97T | BBA 69720 = DAOM 225113 = IMI 375321 = MRC 7541 = NRRL 25331 | Pinus radiata | USA | JAAQPE010000112 | JX171510 | JX171623 | JAAQPE010000100 | JAAQPE010000465 |
| F. cortaderiae | CBS 119183T | FRC R-10016 = ICMP 5435 = NRRL 29297 | Cortaderia selloana | New Zealand | JABCJT010000137 | KM361644 | KM361662 | AY225885 | AH012625 |
| CBS 123655 | FRC R-9682 = ICMP 8998 = NRRL 29306 | Zea mays | New Zealand | PQ240166 | JAGDVA010000005 | JAGDVA010000017 | PQ260832 | JAGDVA010000014 | |
| CBS 123659 | NRRL 31205 | Triticum sp. | Brazil | JAGDUZ010000005 | JAGDUZ010000003 | JAGDUZ010000010 | JAGDUZ010000002 | JAGDUZ010000012 | |
| F. culmorum | CBS 171.28 | IMI 089365 = IMI 089366 = IMI 092033 = NRRL 26854 = NRRL 26915 | Unknown | Canada | PQ240167 | PQ280856 | PQ274119 | PQ260833 | PQ273951 |
| CBS 173.31 | MUCL 783 = NRRL 26853 = NRRL 26914 = NRRL 29139 | Avena sativa | Canada | PQ240168 | PQ280857 | PQ274120 | PQ260834 | PQ273952 | |
| CBS 176.32 | NRRL 29140 | Unknown | Unknown | PQ240169 | PQ280858 | PQ274121 | PQ260835 | PQ273953 | |
| CBS 250.52 | MUCL 555 = NRRL 29291 | Secale cereale | Unknown | PQ240170 | PQ280859 | PQ274122 | PQ260836 | PQ273954 | |
| CBS 251.52 | NRRL 29292 | Triticum aestivum | Unknown | PQ240171 | PQ280860 | PQ274123 | PQ260837 | PQ273955 | |
| CBS 257.51 | NRRL 29142 | Hyacinthus orientalis | Netherlands | PQ240172 | PQ280861 | PQ274124 | PQ260838 | PQ273956 | |
| CBS 416.86 | FRC R-8505 = IMI 309345 = NRRL 36469 | Soil | Denmark | PQ240173 | PQ280862 | PQ274125 | PQ260839 | PQ273957 | |
| CBS 417.86ET | FRC R-8504 = IMI 309344 = NRRL 25475 | Hordeum vulgare | Denmark | PQ240174 | JX171515 | JX171628 | PQ260840 | PQ273958 | |
| CBS 472.95 | NRRL 29144 | Ammophila arenaria | Netherlands | PQ240175 | PQ280863 | PQ274126 | PQ260841 | PQ273959 | |
| CBS 579.97 | NRRL 26241 = NRRL 26602 | Miscanthus sp. | Denmark | PQ240176 | PQ280864 | PQ274127 | PQ260842 | PQ273960 | |
| CBS 597.96 | Unknown | Unknown | PQ240177 | PQ280865 | PQ274128 | PQ260843 | PQ273961 | ||
| CBS 110262 | ATCC 34912 = NRRL 6394 | Millet | Hungary | PQ240178 | PQ280866 | PQ274129 | PQ260844 | PQ273962 | |
| CBS 110565 | NRRL 36195 | Triticum durum | France | PQ240179 | PQ280867 | PQ274130 | PQ260845 | PQ273963 | |
| CBS 110567 | NRRL 36197 | Triticum sp. | France | PQ240180 | PQ280868 | PQ274131 | PQ260846 | PQ273964 | |
| CBS 115698 | Triticum aestivum | Poland | PQ240181 | PQ280869 | PQ274132 | PQ260847 | PQ273965 | ||
| CBS 119869 | MRC 8403 = NRRL 53938 | Unknown | Unknown | PQ240182 | PQ280870 | PQ274133 | PQ260848 | - | |
| CBS 120099 | NRRL 53962 | Secale cereale | Poland | PQ240183 | PQ280871 | PQ274134 | PQ260849 | PQ273966 | |
| CBS 128537 | Triticum aestivum | Belgium | PQ240184 | PQ280872 | PQ274135 | PQ260850 | PQ273967 | ||
| F. cultriforme | CBS 151935T | BBA 65688 = DSM 116528 | Impatiens sp. | Venezuela | PQ240185 | PQ280873 | PQ274136 | PQ260851 | PQ273968 |
| NRRL 36134 | Unknown | Unknown | - | MW233285 | MW233457 | MW233113 | - | ||
| F. cuspidatum | CBS 151953T | FRC R-6269 = NRRL 66923 | Pasture soil debris | USA | PQ240186 | MW233326 | MW233498 | MW233154 | - |
| NRRL 64086 | FRC R-8856 = KOD 2025 | Melon | USA | - | MW233371 | MW233543 | MW233199 | - | |
| F. cygneum | CBS 151805T | IMI 265540 = NRRL 20723 | Unknown | UK | PQ240187 | JX171483 | JX171596 | MW233074 | PQ273969 |
| F. dactylidis | CBS 119181T | ICMP 5269 = NRRL 29298 | Dactylis glomerata | New Zealand | PQ240188 | KM361654 | KM361672 | DQ459748 | DQ459646 |
| CBS 123656 | FRC R-7593 = NRRL 29380 | Dactylis glomerata | USA | PQ240189 | MW233271 | MW233443 | AY452959 | AY452935 | |
| F. dimorphosporum | BBA 64675 | DSM 116017 | Arachis hypogaea | Thailand | PQ240190 | PQ280874 | PQ274137 | PQ260852 | PQ273970 |
| CBS 151959T | FRC R-9230 = NRRL 66938 | Zea mays | Thailand | PQ240191 | MW233388 | MW233560 | MW233216 | PQ273971 | |
| NRRL 64049 | FRC R-6436 = KOD 1988 | Soil | Philippines | - | MW233327 | MW233499 | MW233155 | - | |
| F. dolichosporum | CBS 151817T | FRC R-4069 = MRC 5-B-J = NRRL 66921 | Triticum sp. | South Africa | PQ280985 | MW233318 | MW233490 | MW233146 | PQ280986 |
| NRRL 64077 | FRC R-8203 = KOD 2016 | Debris | South Africa | - | MW233359 | MW233531 | MW233187 | - | |
| F. gerlachii | CBS 119175T | LRG 00-551 = NRRL 36905 | Triticum aestivum | USA | PQ240192 | MW233290 | MW233462 | MW233118 | JAGDUY010000010 |
| CBS 119176 | NRRL 38380 | Arundo donax | USA | JAGDUX010000001 | JAGDUX010000001 | JAGDUX010000006 | JAGDUX010000007 | JAGDUX010000012 | |
| CBS 123666 | NRRL 38405 | Arundo donax | USA | PQ240193 | PQ280875 | PQ274138 | DQ459744 | DQ459642 | |
| F. gladiolum | CBS 151812T | NRRL 54640 = FRC L-247 | Ipomoea batatas | New Guinea | PQ240194 | MW233308 | MW233480 | MW233137 | PQ273972 |
| CBS 151813 | FRC L-252 = NRRL 54683 | Ipomoea batatas | New Guinea | PQ240195 | MW233309 | MW233481 | MW233137 | PQ273973 | |
| NRRL 28066 | FRC R-6767 | Zea mays | Japan | - | MW233262 | MW233434 | MW233091 | - | |
| F. goolgardi | CBS 151815T | KOD 1087 = NRRL 66250 = RBG5411 | Xanthorrhoea glauca | Australia | PQ240196 | PQ280876 | KP083280 | KP101123 | PQ273974 |
| NRRL 66249 | KOD 1090 | Xanthorrhoea glauca | Australia | - | MW233391 | MW233563 | MW233219 | - | |
| F. graminearum | CBS 124.21 | Unknown | Unknown | PQ240197 | PQ280877 | PQ274139 | PQ260853 | PQ273975 | |
| CBS 153.25 | NRRL 29286 | Zea mays | USA | PQ240198 | PQ280878 | PQ274140 | PQ260854 | PQ273976 | |
| CBS 184.32 | NRRL 29288 | Zea mays | Unknown | PQ240199 | PQ280879 | PQ274141 | PQ260855 | PQ273977 | |
| CBS 185.32 | Zea mays | Unknown | PQ240200 | PQ280880 | KT855202 | PQ260856 | PQ273978 | ||
| CBS 189.32 | NRRL 36335 | Zea mays | Unknown | PQ240201 | PQ280881 | PQ274142 | PQ260857 | PQ273979 | |
| CBS 792.70 | BBA 11141 = NRRL 25471 | Zea mays | Iran | PQ240202 | PQ280882 | PQ274143 | PQ260858 | PQ273980 | |
| CBS 110261 | ATCC 46779 = MAFF 237812 = NRRL 5883 = NRRL 38132 | Zea mays | USA | PQ240203 | PQ280883 | PQ274144 | PQ260859 | PQ273981 | |
| CBS 110264 | FRC R-6574 = MAFF 237824 = NRRL 13823 = NRRL 28063 = NRRL 38135 | Zea mays | USA | PQ240204 | PQ280884 | PQ274145 | PQ260860 | PQ273982 | |
| CBS 123657 | NRRL 31084 | Zea mays | USA | PQ240205 | JX171531 | MW233447 | PQ260861 | PQ273983 | |
| CBS 130604 | Triticum sp. | Iran | PQ240206 | PQ280885 | PQ274146 | PQ260862 | PQ273984 | ||
| CBS 130646 | Triticum sp. | Iran | PQ240207 | PQ280886 | PQ274147 | PQ260863 | PQ273985 | ||
| CBS 130666 | Triticum sp. | Iran | PQ240208 | PQ280887 | PQ274148 | PQ260864 | PQ273986 | ||
| CBS 130887 | Triticum sp. | Iran | PQ240209 | PQ280888 | PQ274149 | PQ260865 | PQ273987 | ||
| CBS 130892 | Triticum sp. | Iran | PQ240210 | PQ280889 | PQ274150 | PQ260866 | PQ273988 | ||
| CBS 130914 | Triticum sp. | Iran | PQ240211 | PQ280890 | PQ274151 | PQ260867 | PQ273989 | ||
| CBS 130953 | Triticum sp. | Iran | PQ240212 | PQ280891 | PQ274152 | PQ260868 | PQ273990 | ||
| CBS 130960 | Triticum sp. | Iran | PQ240213 | PQ280892 | PQ274153 | PQ260869 | PQ273991 | ||
| CBS 130963 | Triticum sp. | Iran | PQ240214 | PQ280893 | PQ274154 | PQ260870 | PQ273992 | ||
| CBS 136009ET | Hordeum vulgare | Germany | PQ240215 | PQ280894 | KT855211 | KT855185 | PQ273993 | ||
| CBS 138563 | Triticum sp. | Poland | PQ240216 | PQ280895 | PQ274155 | PQ260871 | PQ273994 | ||
| F. hamatum | BBA 69069 | DSM 116669 | Triticum aestivum | Germany | PQ240217 | - | PQ274156 | PQ260872 | PQ273995 |
| CBS 151936T | BBA 67756 = DSM 116550 | Striga asiatica | Madagascar | PQ240218 | PQ280896 | PQ274157 | PQ260873 | PQ273996 | |
| F. kyushuense | BBA 71679 | Triticum aestivum | Japan | PQ240219 | PQ280897 | PQ274158 | PQ260874 | PQ273997 | |
| CBS 121807 | NRRL 53748 | Unknown | China | PQ240220 | PQ280898 | PQ274159 | PQ260875 | PQ273998 | |
| CBS 151828T | ATCC 56750 = FRC T-346A = MAFF 237645 = MRC 1767 = NRRL 3509 = NRRL A-14732 | Triticum aestivum | Japan | PQ240221 | MW233227 | MW233399 | MW233056 | PQ273999 | |
| NRRL 25349 | Triticum aestivum | Japan | - | - | GQ915492 | GQ915508 | GQ915442 | ||
| F. langsethiae | CBS 113234T | BBA 70945 = DSM 116700 = ITEM 3602 = NRRL 53627 | Avena sativa | Norway | PQ240222 | MW928812 | MW928828 | AB674298 | AB587069 |
| IMI 271486 | Triticum sp. | UK | PQ240223 | PQ280899 | PQ274160 | PQ260876 | - | ||
| IMI 271971 | Triticum sp. | UK | PQ240224 | PQ280900 | PQ274161 | PQ260877 | PQ274000 | ||
| IMI 272890 | Triticum sp. | UK | PQ240225 | PQ280901 | PQ274162 | PQ260878 | PQ274001 | ||
| IMI 272894 | Triticum sp. | UK | PQ240226 | PQ280902 | PQ274163 | PQ260879 | PQ274002 | ||
| IMI 272899 | Triticum sp. | UK | PQ240227 | PQ280903 | PQ274164 | PQ260880 | - | ||
| NRRL 54940 | Avena sp. | Norway | PQ240228 | JX171550 | JX171662 | MW233138 | MW233137 | ||
| F. leptum | CBS 151949T | NRRL 29896 = FRC R-9673 | Unknown | Netherlands | PQ240229 | MW233272 | HQ154449 | HM744660 | HQ141636 |
| NRRL 29897 | FRC R-9674 | Unknown | Netherlands | - | MW233273 | MW233445 | MW233101 | - | |
| F. longicolle | CBS 120991 | FRC R-9983 = MRC 8429 = MRC 8430 = KSU 11429 = KSU 11431 = Univ. Sydney F11993 = Univ. Sydney F5837 | Unknown | Australia | PQ240230 | PQ280904 | PQ274165 | PQ260881 | PQ274003 |
| CBS 151818T | FRC R-6968 = NRRL 66924 | Soil debris | Australia | PQ240231 | MW233332 | MW233504 | MW233160 | PQ274004 | |
| NRRL 64053 | FRC R-6894 = KOD 1992 | Air | Australia | - | MW233331 | MW233503 | MW233159 | - | |
| F. longipes | CBS 120.73 | ATCC 24359 = IMI 136675 = NRRL 20694 = VTT D-81072 | Oryza sativa | Denmark | PQ240232 | PQ280905 | PQ274166 | PQ260882 | PQ274005 |
| CBS 476.77NT | NRRL 20695 | Soil | USA | PXOG01000030 | MW233244 | PXOG01000079 | PXOG01000060 | PXOG01000089 | |
| CBS 739.79 | BBA 62061 = DSM 62061 = FRC R-9842 = NRRL 20696 | Soil | Malaysia | PQ240233 | PQ280906 | PQ274167 | - | PQ274006 | |
| IMI 179815 | NRRL 20725 | Unknown | Malaysia | PQ240234 | PQ280907 | PQ274168 | PQ260883 | PQ274007 | |
| IMI 196466 | NRRL 20724 | Cymbopogon martinii | India | PQ240235 | PQ280908 | PQ274169 | PQ260884 | PQ274008 | |
| IMI 311604 | Psidium guajava | India | PQ240236 | PQ280909 | PQ274170 | PQ260885 | PQ274009 | ||
| IMI 357087 | Acacia mangium | Malaysia | PQ240237 | PQ280910 | PQ274171 | PQ260886 | PQ274010 | ||
| F. louisianense | CBS 127524 | NRRL 54196 | Triticum aestivum | USA | JAGDUW010000014 | JAGDUW010000011 | JAGDUW010000011 | JAGDUW010000004 | JAGDUW010000005 |
| CBS 127525T | NRRL 54197 | Triticum aestivum | USA | JAGDUV010000001 | KM361667 | KM361667 | KM889633 | KM889628 | |
| F. lunulusporum | CBS 636.76T | ATCC 36747 = BBA 62459 = FRC R-5822 = IMI 322097 = NRRL 13393 | Citrus paradisi | South Africa | PQ240238 | KM361637 | KM361655 | AF212467 | PQ274011 |
| F. magnum | CBS 151821T | FRC R-7846 = MRC 3530 = NRRL 66927 | Soil | South Africa | PQ240239 | MW233350 | MW233522 | MW233178 | PQ274012 |
| NRRL 64075 | FRC R-8136 = KOD 2014 | Debris | South Africa | - | MW233356 | MW233528 | MW233184 | - | |
| F. mastigosporum | BBA 62247 | DSM 62247 | Soil | Australia | PQ240240 | PQ280911 | PQ274172 | PQ260887 | PQ274013 |
| CBS 151803 | NRRL 13368 | Soil | Australia | JABFEZ010000001 | MW233231 | MW233403 | JABFEZ010000109 | JABFEZ010000065 | |
| CBS 151825T | FRC R-9077 = NRRL 66935 | Soil debris | Australia | PQ240241 | MW233382 | MW233554 | MW233210 | PQ274014 | |
| F. meridionale | CBS 110247T | FRC R-5329 = NRRL 28436 | Orange twig | New Caledonia | PQ240242 | KM361642 | KM361660 | AF212435 | AF212730 |
| CBS 110248 | FRC R-9438 = NRRL 28723 = NRRL 38118 | Zea mays | Nepal | PQ240243 | - | PQ274173 | PQ260888 | PQ274015 | |
| CBS 110249 | FRC R-4080 = MRC 0856 = NRRL 29010 | Soil | South Africa | JAGDUU010000006 | JAGDUU010000007 | JAGDUU010000019 | AF212437 | AF212732 | |
| CBS 110260 | FRC R-9430 = NRRL 28721 = NRRL 38131 | Zea mays | Nepal | PQ240244 | JAGDUT010000007 | PQ274174 | AF212454 | AF212747 | |
| F. mesoamericanum | CBS 415.86T | FRC R-8506 = IMI 309346 = NRRL 25797 = NRRL 38123 | Musa sp. | Honduras | JAGDUS010000013 | KM361639 | KM361657 | AF212441 | AF006364 |
| CBS 110252 | FRC R-4079 = NRRL 29148 = NRRL 38124 | Cissus rhombifolia | USA | PQ240245 | PQ280912 | PQ274175 | PQ260889 | PQ274016 | |
| F. minutum | CBS 151822 | FRC R-8165 = NRRL 66928 | Olive grove soil | Tunisia | PQ240246 | MW233358 | MW233530 | MW233186 | PQ274017 |
| CBS 151823T | FRC R-8838 = NRRL 66931 | Unknown | Italy | PQ240247 | MW233368 | MW233540 | MW233196 | - | |
| F. mucronatum | CBS 151826T | FRC R-9135 = NRRL 66937 | Pennisetum glaucum | Botswana | PQ240248 | MW233387 | MW233559 | MW233215 | PQ274018 |
| NRRL 64089 | FRC R-8886 = KOD 2028 | Unknown | Nigeria | - | MW233374 | MW233546 | MW233202 | - | |
| F. musarum | BBA 68458 | DSM 116552 | Musa sp. | Costa Rica | PQ240249 | PQ280913 | PQ274176 | PQ260890 | PQ274019 |
| BBA 68461 | DSM 116554 | Musa sp. | Costa Rica | PQ240250 | PQ280914 | PQ274177 | PQ260891 | PQ274020 | |
| BBA 68462 | DSM 116555 | Musa sp. | Costa Rica | PQ240251 | PQ280915 | PQ274178 | PQ260892 | PQ274021 | |
| CBS 151808T | DAOM 225261 = FRC R-9400 = NRRL 28507 = MRC 6240 | Musa sapientum | Panama | PQ240252 | PQ280916 | MW233437 | MW233094 | PQ274022 | |
| F. nepalense | CBS 127503T | NRRL 54222 | Oryza sativa | Nepal | PQ240253 | KM361650 | KM361668 | KM889631 | KM889626 |
| CBS 127669 | NRRL 54220 | Oryza sativa | Nepal | JAGDUR010000004 | JAGDUR010000001 | JAGDUR010000006 | KM889629 | KM889624 | |
| CBS 127943 | NRRL 54221 | Oryza sativa | Nepal | JAGDUQ010000011 | JAGDUQ010000002 | JAGDUQ010000006 | JAGDUQ010000009 | JAGDUQ010000005 | |
| F. nodosum | CBS 201.63T | Arachis hypogaea | Portugal | MN120704 | MN120725 | GQ915484 | MW233117 | JAAVUE010000073 | |
| F. palustre | CBS 120613 | Unknown | Unknown | PQ240254 | PQ280917 | PQ274179 | PQ260893 | PQ274023 | |
| CBS 126796T | NRRL 54056 | Spartina alterniflora | USA | PQ240255 | MW233303 | MW233475 | MW233131 | GQ856983 | |
| NRRL 43289 | Spartina sp. | USA | - | MW233296 | MW233468 | MW233124 | - | ||
| F. parabolicum | CBS 128810 | RMF 8363 | Soil | USA | PQ240256 | PQ280918 | PQ274180 | PQ260894 | PQ274024 |
| CBS 151814T | ATCC 36781 = FRC R-5319 = MRC 1783 = NRRL A-15494 = NRRL 6227 | Festuca arundinacea | USA | PQ240257 | JX171446 | JX171560 | HM744692 | HQ141667 | |
| NRRL 64071 | FRC R-8090 = KOD 2010 = MRC 4027 | Avena sativa | South Africa | PQ240258 | MW233352 | MW233524 | MW233180 | PQ274025 | |
| NRRL 64093 | FRC R-9068 = KOD 2032 | Zea mays | Canada | - | MW233381 | MW233553 | MW233209 | - | |
| F. platysporum | CBS 151810 | FRC R-8755 = NRRL 28725 | Triticum sp. | Argentina | PQ240259 | MW233267 | MH845431 | PQ260895 | - |
| CBS 151816T | KOD 1723 = NRRL 66920 | Glycine max | Ghana | PQ240260 | MW233394 | MW233566 | MW233222 | PQ274026 | |
| NRRL 66922 | FRC R-4932 | Triticum sp. | Brazil | - | MW233321 | MW233493 | MW233149 | - | |
| F. poae | CBS 174.96 | Avena sativa | Norway | PQ240261 | PQ280919 | PQ274181 | PQ260896 | PQ274027 | |
| CBS 175.96 | Avena sativa | Norway | PQ240262 | PQ280920 | PQ274182 | PQ260897 | PQ274028 | ||
| CBS 176.96 | Avena sativa | Norway | PQ240263 | PQ280921 | PQ274183 | PQ260898 | PQ274029 | ||
| CBS 180.96 | Triticum sp. | Norway | PQ240264 | PQ280922 | PQ274184 | PQ260899 | PQ274030 | ||
| CBS 186.96 | Triticum sp. | Poland | PQ240265 | PQ280923 | PQ274185 | PQ260900 | PQ274031 | ||
| CBS 446.67 | BBA 10426 = FRC T-982 = NRRL 25799 | Anthoxanthum odoratum | Germany | PQ240266 | PQ280924 | PQ274186 | PQ260901 | PQ274032 | |
| CBS 623.87 | NRRL 36524 | Hordeum vulgare | Denmark | PQ240267 | PQ280925 | PQ274187 | PQ260902 | PQ274033 | |
| CBS 115696 | Triticum aestivum | Poland | PQ240268 | PQ280926 | PQ274188 | PQ260903 | PQ274034 | ||
| CBS 121297 | NRRL 53735 | Triticum sp. | Switzerland | PQ240269 | - | PQ274189 | PQ260904 | PQ274035 | |
| CBS 128536 | Triticum aestivum | Belgium | PQ240270 | PQ280927 | PQ274190 | PQ260905 | PQ274036 | ||
| CBS 151947 | NRRL 13714 = FRC T-503 = MRC 2181 | Triticum sp. | Canada | PQ240271 | JX171458 | JX171572 | PQ260906 | PQ274037 | |
| CBS 151948ET | NRRL 26941 = FRC T-962 | Hordeum vulgare | USA | JABFFD010000771 | JABFFD010000177 | JABFFD010000819 | JABFFD010000730 | JABFFD010000556 | |
| F. praegraminearum | CBS 141369T | ICMP 8996 = NRRL 39664 = PDD 47563 | Litter | New Zealand | LXHY01000061 | KX260125 | KX260126 | KX260120 | KX260131 |
| F. pratense | CBS 151956T | FRC R-8926 = NRRL 66932 | Wet grassland soil | Australia | PQ240272 | MW233375 | MW233547 | MW233203 | PQ274038 |
| IMI 316488 | Saccharum officinarum | New Guinea | PQ240273 | PQ280928 | PQ274191 | PQ260907 | PQ274039 | ||
| F. procumbens | CBS 121863T | FRC R-9983 = KSU 11431 = MRC 8430 = NRRL 53749 | Unknown | Netherlands | PQ240274 | PQ280929 | PQ274192 | PQ260908 | - |
| F. pseudograminearum | BBA 71458 | DSM 116903 | Hordeum vulgare | Australia | PQ240275 | PQ280930 | PQ274193 | PQ260909 | PQ274040 |
| BBA 71460 | DSM 116904 | Pasture soil | Australia | PQ240276 | PQ280931 | PQ274194 | PQ260910 | PQ274041 | |
| CBS 109956T | FRC R-5291 = NRRL 28062 | Hordeum vulgare | Australia | PQ240277 | PQ280932 | PQ274195 | PQ260911 | PQ274042 | |
| IMI 350956 | Medicago polymorpha | Australia | PQ240278 | PQ280933 | PQ274196 | PQ260912 | PQ274043 | ||
| F. pseudolongipes | CBS 131379 | Oryza australiensis | Australia | PQ240279 | PQ280934 | PQ274197 | PQ260913 | PQ274044 | |
| CBS 131380T | Oryza australiensis | Australia | PQ240280 | PQ280935 | PQ274198 | - | PQ274045 | ||
| F. robustum | CBS 637.76IT | BBA 63667 = FRC R-5821 = IMI 322102 = NRRL 13392 | Araucaria angustifolia | Argentina | PQ240281 | PQ280936 | PQ274199 | PQ260914 | PQ274046 |
| F. sagittatum | CBS 151958T | FRC R-9120 = NRRL 66936 | Soil | China | PQ240282 | MW233384 | MW233556 | MW233212 | - |
| NRRL 64094 | FRC R-9118 = KOD 2033 | Soil | China | - | MW233383 | MW233555 | MW233211 | - | |
| F. sambucinum | CBS 125.95 | BBA 62433 = DSM 62433 | Beta vulgaris | Spain | PQ240283 | PQ280937 | PQ274200 | PQ260915 | PQ274047 |
| CBS 146.95 | BBA 64226 = DSM 115507 = NRRL 20727 = NRRL 22187 = NRRL 36256 | Solanum tuberosum | UK | PQ240284 | PQ280938 | PQ274201 | PQ260916 | PQ274048 | |
| CBS 260.95 | BBA 62397 = DSM 62397 = NRRL 36396 | Solanum tuberosum | Germany | PQ240285 | PQ280939 | PQ274202 | PQ260917 | PQ274049 | |
| CBS 533.96 | Unknown | Unknown | PQ240286 | PQ280940 | PQ274203 | PQ260918 | PQ274050 | ||
| CBS 100056 | NRRL 53596 | Laurus nobilis | France | PQ240287 | PQ280941 | PQ274204 | PQ260919 | PQ274051 | |
| CBS 111114 | NRRL 36200 | Unknown | Netherlands | PQ240288 | PQ280942 | PQ274205 | PQ260920 | PQ274052 | |
| CBS 119802 | FRC R-6380 = MRC 2193 = MRC 6970 = NRRL 13455 | Solanum tuberosum | Germany | PQ240289 | PQ280943 | PQ274206 | PQ260921 | PQ274053 | |
| CBS 119804 | BBA 64995 = FRC R-9153 = MRC 6972 = NRRL 53899 | Brassica oleracea | Netherlands | PQ240290 | PQ280944 | PQ274207 | PQ260922 | PQ274054 | |
| CBS 151939 | CPC 45309 | Sambucus nigra | Germany | PQ240291 | PQ280945 | PQ274208 | PQ260923 | PQ274055 | |
| CBS 151940 | CPC 45310 | Sambucus nigra | Germany | PQ240292 | PQ280946 | PQ274209 | PQ260924 | PQ274056 | |
| CBS 151941 | CPC 45313 | Sambucus nigra | Germany | PQ240293 | PQ280947 | PQ274210 | PQ260925 | PQ274057 | |
| CBS 151802 | CPC 45315 | Sambucus nigra | Germany | PQ240294 | PQ280948 | PQ274211 | PQ260926 | PQ274058 | |
| CBS 151942ET | CPC 45317 | Sambucus nigra | Germany | PQ240295 | PQ280949 | PQ274212 | PQ260927 | PQ274059 | |
| CBS 151943 | CPC 45321 | Sambucus nigra | Germany | PQ240296 | PQ280950 | PQ274213 | PQ260928 | PQ274060 | |
| CBS 151944 | CPC 45322 | Sambucus nigra | Germany | PQ240297 | PQ280951 | PQ274214 | PQ260929 | PQ274061 | |
| CPC 41834 | Solanum tuberosum | Netherlands | PQ240298 | PQ280952 | PQ274215 | PQ260930 | PQ274062 | ||
| CPC 41854 | F80 | Solanum tuberosum | Russia | PQ240299 | PQ280953 | PQ274216 | PQ260931 | PQ274063 | |
| CPC 41888 | F81 | Solanum tuberosum | Russia | PQ240300 | - | PQ274217 | PQ260932 | PQ274064 | |
| NRRL 13394 | BBA 63572 = FRC R-5823 | Unknown | Germany | - | MW233235 | MW233407 | MW233064 | - | |
| NRRL 20663 | IMI 266242 | Pterocarya fraxinifolia | Germany | - | MW233242 | MW233414 | MW233071 | - | |
| NRRL 20666 | IMI 295478 | Solanum tuberosum | UK | - | MW233243 | MW233415 | MW233072 | - | |
| NRRL 31964 | NRRL 39954 = ICMP 11550 = IMI 351949 | Cytisus scoparius | New Zealand | - | MW233278 | MW233450 | MW233106 | - | |
| NRRL 31969 | NRRL 39966 = ICMP 12578 | Ulex europaeus | New Zealand | - | MW233279 | MW233451 | MW233107 | - | |
| F. seculiforme | CBS 151806T | DSM 64262 = NRRL 22189 = NRRL 20445 | Glycine max | Brazil | PQ240301 | MW233247 | MW233419 | MW233076 | U85582 |
| NRRL 64059 | FRC R-7403 = ATCC 52543 = KOD 1998 = MRC 2967 | Solanum tuberosum | South Africa | - | MW233338 | MW233510 | MW233166 | - | |
| F. sibiricum | BBA 71293 | DSM 116899 | Triticum aestivum | Unknown | PQ240302 | PQ280954 | PQ274218 | PQ260933 | PQ274065 |
| BBA 71294 | DSM 116900 | Triticum aestivum | Unknown | PQ240303 | PQ280955 | PQ274219 | PQ260934 | PQ274066 | |
| CBS 140910 | PQ240304 | PQ280956 | HQ154464 | HM744676 | HQ141651 | ||||
| CBS 140915 | MFG 11007 = NRRL 53424 = RCAM 03252 | Hordeum vulgare | Russia | PQ240305 | PQ280957 | HQ154466 | HM744678 | HQ141653 | |
| CBS 140945 | MFG 11014 = NRRL 53431 = RCAM 03252 | Avena sativa | Russia | PQ240306 | PQ280958 | HQ154473 | HM744685 | HQ141660 | |
| CBS 151951T | NRRL 53430 = MFG 11013 | Avena sativa | Russia | PQ240307 | MW233302 | HQ154472 | HM744684 | HQ141659 | |
| CPC 41132 | F12 | Submersible wood | Russia | PQ240308 | PQ280959 | PQ274220 | PQ260935 | PQ274067 | |
| CPC 41881 | F57 | Submersible wood | Russia | PQ240309 | PQ280960 | PQ274221 | PQ260936 | PQ274068 | |
| F. sporotrichioides | BBA 62421 | DSM 115487 | Pinus sylvestris | Germany | PQ240310 | PQ280961 | PQ274222 | PQ260937 | PQ274069 |
| BBA 62422 | DSM 115488 | Malus sp. | Germany | PQ240311 | PQ280962 | PQ274223 | PQ260938 | PQ274070 | |
| BBA 62423 | DSM 115489 | Pinus nigra | Germany | PQ240312 | PQ280963 | PQ274224 | PQ260939 | PQ274071 | |
| CBS 178.64 | MUCL Be-133 = NRRL 36295 | Nicotiana tabacum | Belgium | PQ240313 | PQ280964 | PQ274225 | PQ260940 | PQ274072 | |
| CBS 412.86 | FRC T-822 = IMI 309350 | Juncus sp. | Denmark | PQ240314 | PQ280965 | PQ274226 | PQ260941 | PQ274073 | |
| CBS 448.67 | BBA 10360 = FRC T-0980 = NRRL 25474 | Picea abies | Germany | PQ240315 | PQ280966 | PQ274227 | PQ260942 | PQ274074 | |
| CBS 119839 | DAOM 175513 = MRC 4333 = NRRL 53912 | Zea mays | Canada | PQ240316 | PQ280967 | PQ274228 | AB674304 | AB587075 | |
| CBS 123674 | IBT 1926 | Soil | Netherlands | PQ240317 | PQ280968 | PQ274229 | PQ260943 | PQ274075 | |
| CBS 151950 | NRRL 3299 = NRRL 3287 = ATCC 24631 = FRC T-423 = FRC T-424 = MRC 1768 = MRC 43 | Zea mays | France | PXOF01000117 | JX171444 | HQ154454 | PXOF01000134 | PXOF01000016 | |
| CPC 41121 | F1 | Hordeum vulgare | Russia | PQ240318 | PQ280969 | PQ274230 | PQ260944 | PQ274076 | |
| CPC 41150 | F30 | Pinus sylvestris | Russia | PQ240319 | PQ280970 | PQ274231 | PQ260945 | PQ274077 | |
| CPC 41842 | F97 | Soil | Russia | PQ240320 | PQ280971 | PQ274232 | PQ260946 | PQ274078 | |
| CPC 41885 | F66 | Solanum tuberosum | Russia | PQ240321 | PQ280972 | PQ274233 | PQ260947 | PQ274079 | |
| IMI 322457 | Dianthus sp. | France | PQ240322 | PQ280973 | PQ274234 | PQ260948 | PQ274080 | ||
| IMI 348484 | Hordeum vulgare | UK | PQ240323 | PQ280974 | PQ274235 | PQ260949 | PQ274081 | ||
| F. subcylindroides | CBS 151807T | BBA 70355 = NRRL 26795 | Soil | USA | PQ240324 | MW233258 | MW233430 | MW233087 | PQ274082 |
| F. subflagellisporum | CBS 151955 | FRC R-8774 = NRRL 66930 | Soil | USA | PQ240325 | MW233365 | MW233537 | MW233193 | - |
| FRC R-7642 | KOD 2007 | Sorghum sp. | Puerto Rico | - | MW233347 | MW233519 | MW233175 | - | |
| F. subtropicale | CBS 144706T | NRRL 66764 = UEM 3329 | Hordeum vulgare | Brazil | PQ240326 | MW233312 | MH706973 | MH706974 | MH706968 |
| F. symmetricum | CBS 151804T | NRRL 13465 = NRRL A-26556 | Opuntia aurantiaca | Argentina | PQ240327 | MW233238 | MW233410 | MW233067 | PQ274083 |
| CBS 151819 | FRC R-7071 = MRC 2857 = NRRL 66925 | Opuntia aurantiaca | Argentina | PQ240328 | MW233334 | MW233506 | MW233162 | PQ274084 | |
| F. transvaalense | CBS 119878 | MRC 6142 = NRRL 53947 | Dianthus sp. | South Africa | PQ240329 | PQ280975 | PQ274236 | PQ260950 | PQ274085 |
| CBS 119879 | FRC R-6837 = FRC R-9976 = MRC 2800 | Plant debris | South Africa | PQ240330 | PQ280976 | PQ274237 | PQ260951 | PQ274086 | |
| CBS 144211T | CPC 30923 | Sida cordifolia | South Africa | PQ240331 | PQ280977 | LT996157 | LT996099 | LT996120 | |
| CBS 144212 | CPC 30939 | Kyphocarpa angustifolia | South Africa | PQ240332 | LT996211 | LT996158 | LT996100 | LT996121 | |
| CBS 144214 | CPC 30946 | Sida cordifolia | South Africa | PQ240333 | PQ280978 | LT996160 | LT996101 | LT996123 | |
| CBS 144217 | CPC 30919 | Sida cordifolia | South Africa | PQ240334 | LT996214 | LT996163 | LT996104 | LT996126 | |
| CBS 144218 | CPC 30922 | Sida cordifolia | South Africa | PQ240335 | LT996215 | LT996164 | LT996105 | LT996127 | |
| CBS 144220 | CPC 30927 | Sida cordifolia | South Africa | PQ240336 | PQ280979 | LT996166 | LT996107 | LT996129 | |
| CBS 144222 | CPC 30939 | Kyphocarpa angustifolia | South Africa | PQ240337 | PQ280980 | LT996168 | LT996108 | LT996131 | |
| CBS 144224 | CPC 30928 | Melhania acuminata | South Africa | PQ240338 | LT996219 | LT996170 | LT996110 | LT996133 | |
| NRRL 31008 | BBA 63772 = DSM 115500 = FRC R-9856 | Soil | Australia | - | MW233274 | JX171642 | MW233102 | - | |
| F. tropicale | CBS 151820T | FRC R-7764 = NRRL 66926 | Rice paddy soil | Thailand | PQ240339 | MW233349 | MW233521 | MW233177 | PQ274087 |
| IMI 312359 | Boehmeria nivea | India | PQ240340 | PQ280981 | PQ274238 | PQ260952 | PQ274088 | ||
| F. ussurianum | CBS 123752T | NRRL 45681 = TG-2662/0 | Avena sativa | Russia | JAGDUP010000016 | KM361648 | KM361666 | FJ240301 | FJ240291 |
| CBS 123753 | FRC R-10095 = NRRL 45795 = TG-65202 | Triticum sp. | Russia | PQ240341 | PQ280982 | PQ274239 | FJ240303 | FJ240292 | |
| NRRL 29813 | Triticum sp. | Russia | JAGDUO010000002 | JAGDUO010000001 | JAGDUO010000002 | JAGDUO010000006 | JAGDUO010000004 | ||
| NRRL 58212 | Triticum sp. | Russia | JAGDUM010000011 | JAGDUM010000010 | JAGDUN010000003 | JAGDUN010000012 | JAGDUN010000015 | ||
| F. venenatum | CBS 127.95 | BBA 64478 = NRRL 36230 | Solanum tuberosum | Finland | PQ240342 | PQ280983 | PQ274240 | PQ260953 | PQ274089 |
| CBS 458.93T | BBA 64537 = FRC R-9186 = NRRL 26228 | Triticum aestivum | Austria | KM231392 | MW233256 | MW233428 | MW233085 | KM232079 | |
| CBS 140911 | MFG 58720 | Grass | Russia | PQ240343 | PQ280984 | PQ274241 | PQ260954 | PQ274090 | |
| NRRL 22196 | BBA 65031 = DSM 116036 = FRC R-9187 | Zea mays | Germany | - | JX171494 | JX171607 | MW233078 | - | |
| F. vermicularioides | CBS 151945T | FRC R-5128 = NRRL 13374 | Soil debris | New Guinea | JACCKV010000102 | MW233232 | MW233404 | JACCKV010000130 | JACCKV010000150 |
| F. vorosii | CBS 119177T | FRC R-10011 = NRRL 37605 | Triticum sp. | Hungary | JAGDUL010000040 | JAGDUL010000004 | JAGDUL010000005 | JAGDUL010000032 | JAGDUL010000025 |
| CBS 119178 | FRC R-9999 = NRRL 38208 | Triticum sp. | Japan | JAGDUK010000004 | JAGDUK010000002 | JAGDUK010000007 | JAGDUK010000038 | JAGDUK010000018 | |
| FSAMSC2 | NRRL 54062 | Spartina alternifora | USA | GQ857053 | MW233305 | MW233477 | GQ856948 | GQ856990 | |
| NRRL 64045 | FRC R-5100 = KOD 1984 | Zea mays | Australia | - | MW233323 | MW233495 | MW233151 | - | |
| FSAMSC7 | NRRL 46743 | Triticum sp. | Ethiopia | - | MW233299 | MW233471 | MW233127 | - | |
| NRRL 66736 | KOD 1545 | Glycine max | Ethiopia | - | MW233311 | MW233483 | MW233139 | - | |
| FSAMSC11 | NRRL 22192 | DSM 64918 = FRC R-10045 = NRRL 20447 | Palm tree | Indonesia | - | MW233248 | MW233420 | MW233077 | U85583 |
| FSAMSC13 | NRRL 39635 | ICMP 5288 | Cortaderia sp. | New Zealand | - | MW233292 | MW233464 | MW233120 | - |
| NRRL 39685 | ICMP 8026 | Cortaderia sp. | New Zealand | - | MW233294 | MW233466 | MW233122 | - | |
| FSAMSC31 | NRRL 66919 | KOD 1722 | Glycine max | Ghana | - | MW233393 | MW233565 | MW233221 | - |
1 FSAMSC: Phylogenetic species designation according to Laraba et al. (2021).
2 ATCC: American Type Culture Collection, Manassas, VA, USA; BBA: Collection of the Julius Kühn Institute – Federal Research Centre for Cultivated Plants (former Biologische Bundesanstalt für Land- und Forstwirtschaft) housed at the Institute for Epidemiology and Pathogen Diagnostics, Braunschweig, Germany; CBS: Westerdijk Fungal Biodiverity Institute (WI), Utrecht, The Netherlands; CPC: Collection of P.W. Crous, held at WI; DAOM: Canadian National Mycological Herbarium, Ottawa, Ontario, Canada; DSM: DSMZ-Deutsche Sammlung von Mikroorganismen und Zellkulturen GmbH, Braunschweig, Germany; F: Strains received from I. Pavlov, Laboratory of Reforestation, Mycology and Plant Pathology, V.N. Sukachev Institute of Forest of Russian Academy of Sciences, Krasnoyarsk, Russia; FRC: Fusarium Research Center, Pennsylvannia State University, PA, USA; GJS: Collection of G.J. Samuels, USDA-ARS, USA; IBT: Institut for Bioteknologi, DTU Bioengineering, Lyngby, Denmark; ICMP: International Collection of Microorganisms from Plants. Auckland, New Zealand; IMI: CABI Bioscience, Eggham, UK; ITEM: Agri-Food Toxigenic Fungi Culture Collection, Institute of Sciences of Food Production, Bari, Italy; KOD: Personal collection of Kerry O’Donnell, Agricultural Research Service, USDA, Beltsville, Maryland, USA; KSU: Department of Plant Pathology, Throckmorton Plant Sciences Center, Kansas State University, Manhattan, Kansas, USA; LRG: Personal collection of Liane R. Gale, ARS-USDA, Department of Plant Pathology, University of Minnesota, St. Paul, Minnesota, USA; MAFF: Ministry of Agriculture, Forestry and Fisheries, Tsukuba, Ibaraki, Japan; MFG: VIZR All-Russian Plant Protection Institute collection, St. Petersburg-Pushkin, Russia; MRC: PROMEC, Medical Research Council, Tygerberg, South Africa; MUCL: Mycothèque de l´Université Catholique de Louvain, Louvain-la-Neuve, Belgium; NRRL: Agricultural Research Service Culture Collection, National Center for Agricultural Utilization Research, USDA, Peoria, IL, USA; PDD: Plant Disease Division herbarium, New Zealand Department of Scientific and Industrial Research, Aukland, New Zealand; RBG: Royal Botanic Gardens Trust, Sydney, New South Wales, Australia; RC: Universidad Nacional de Río Cuarto culture (RC) collection, Córdoba, Río Cuarto, Argentina; RCAM: Russian Collection of Agricultural Microorganisms, All-Russia Research Institute for Agricultural Microbiology, St.Petersburg, Russia; RMF: Rocky Mountain Herbarium, Fungi, University of Wyoming, Laramie, WY, USA; UEM: Departamento de Análises Clínicas e Biomedicina, Universidade Estadual de Maringá, Maringá, Brazil; Univ. Sydney: Department of Plant Pathology and Agricultural Entomology, The University of Sidney, Sydney, Australia; VTT: Technical Research Centre of Finland, Espoo, Finland; W: Strains received from O. Yarden, Department of Plant Pathology and Microbiology, The Robert H. Smith Faculty of Agriculture, Food and Environment, The Hebrew University of Jerusalem, Rehovot, Israel; ET: Ex-epitype, IT: Ex-isotype, NT: Ex-neotype, T: Ex-type.
3 CaM: calmodulin; rpb1: RNA polymerase II largest subunit; rpb2: RNA polymerase II second largest subunit; tef1: translation elongation factor 1-α; tub2: β-tubulin; accession numbers with 12 or more digits correspond to genome data; sequences newly generated in this study are shown in bold.
Morphology
Fungal strains were studied according to methods described in Crous et al. (2021). Briefly, in vitro growth rates and colony pigmentation were studied in potato dextrose agar (PDA; Crous et al. 2021), with additional observations made on oatmeal agar (OA; Crous et al. 2021) and synthetic nutrient-poor agar (SNA; Nirenberg 1976), all incubated for 7–14 d at 24 °C in darkness. Micromorphological studies were carried out using sterile distilled water as mounting medium from fungal strains grown on carnation leaf agar (CLA; Fisher et al. 1982), incubated at room temperature (22–24 °C) under a 12/12 h near UV light/dark cycle (Fisher et al. 1982, Leslie & Summerell 2006). Fungal structures from fungarium specimens were rehydrated in 3 % aqueous KOH for a few minutes, and then rinsed by replacing the KOH with sterile distilled water (Samuels 1976). Optical instruments employed included a Nikon Eclipse 80i and a Nikon Eclipse Ni compound microscopes with Differential Interference Contrast (DIC) optics; and a Zeiss Discovery.V20 and a Nikon AZ100 motorized dissecting microscopes. All instruments were equipped with Nikon DS-Ri2 high-definition colour digital cameras. Digital imaging and measurements were done using the Nikon software NIS-Elements v. 5.11.02, and image editing and photoplates were prepared in Adobe Photoshop 2024, release 25.4.0. At least 30 individual measurements were recorded for every fungal structure. Total range (in parentheses), averages and average plus and minus standard deviations values are shown in species descriptions. Colour notations in species descriptions are from Rayner (1970); however, colony colour comparison in taxonomic notes and tables refer to the main shade of colonial pigments and not to a specific colour notation.
DNA extraction, amplification, and sequencing
Fungal isolates were grown on malt extract agar (MEA; Crous et al. 2019) for 3–5 d at 24 °C in darkness. Mycelium was scrapped from the agar surface with the aid of a sterile scalpel and subjected to total genomic DNA extraction using the Wizard® Genomic DNA purification Kit (Promega Corporation, Madison, WI, USA) following the manufacturer’s instructions.
Partial sequences of five loci were amplified according to protocols described elsewhere (Crous et al. 2021) using the primer pairs CAL-228F and CAL-2Rd for calmodulin (CaM; Carbone & Kohn 1999, Quaedvlieg et al. 2011), Fa and G2R or Fa and R9 for the RNA polymerase II largest subunit (rpb1; O’Donnell et al. 2010), RPB2-5f2 and fRPB2-7cr and fRPB2-7cf and RPB2-11ar for two non-contiguous fragments of the RNA polymerase II second largest subunit (rpb2; Liu et al. 1999, Sung et al. 2007), EF-1 and EF-2 for the translation elongation factor 1-α (tef1; O’Donnell et al. 1998b, 2008a), and T1 and TUB4RD for β-tubulin (tub2; O’Donnell & Cigelnik 1997, Woudenberg et al. 2009).
Sequencing was carried out using the same primers as indicated above, plus the internal sequencing primers F6, F7, and R8 from rpb1 (O’Donnell et al. 2010) using a Hitachi 3730xl DNA analyser (Applied Biosystems Inc., Foster City, California, USA). Consensus sequences were assembled from forward and reverse sequences using Geneious Prime v. 2023.2.1 (Biomatters Inc., New Zealand). Sequences generated in this study were deposited in GenBank (Table 1).
Phylogenetic analyses
Single gene alignments were prepared using the MAFFT multiple aligner v. 1.5.0 module (Katoh & Standley 2013) as provided in Geneious Prime. Individual alignments were exported and manually corrected in MEGA v. 6.06 (Tamura et al. 2013), and a 5-gene combined alignment was built using SequenceMatrix v. 1.9 (Vaidya et al. 2011).
Phylogenetic analyses were run using Maximum-Likelihood (ML) and Bayesian inference (BI) algorithms using the software IQ-TREE multicore v. 2.1.2 COVID-edition (Nguyen et al. 2015), and the tools RAxML-HPC2 Workflow on XSEDE v. 8.2.12 (Stamatakis 2006, Silvestro & Michalak 2012) and MrBayes v. 3.2.7a (Ronquist et al. 2012) as implemented in the CIPRES Science Gateway (Miller et al. 2010). For IQ-TREE, the selection of best evolutionary models for each partition was carried out according to the TESTNEW option of ModelFinder (Kalyaanamoorthy et al. 2017) as implemented in IQ-TREE, and branch support was estimated with 1 000 replicates of ultrafast bootstrap approximation (UFBoot; Hoang et al. 2018). RAxML runs included 1 000 non-parametric bootstrap inferences, with free model parameters estimated by the software. Evolutionary models for BI were calculated in MrModelTest v. 2.3 (Posada & Crandall 1998, Nylander 2004), selected according to the Akaike information criterion. The BI analyses were set up to 5 M generations with the stop-rule option on, including four incrementally heated MCMC chains. Trees were sampled every 1 000 generations and runs were checked for convergence by average standard deviation of split frequencies below 0.01 and visualization of trace files using Tracer v. 1.7.2 (Rambaut et al. 2018). Consensus trees (50 %) and posterior probabilities (PP) were calculated after discarding 25 % of the initial samples as the burn-in fraction. Alignments and phylogenetic trees were deposited in www.figshare.com (doi: 10.6084/m9.figshare.27720480).
Determination of mating types and in-vitro mating assays
The mating-type (MAT) was determined for selected, newly collected F. sambucinum isolates and the MAT1-2 tester strain F. sambucinum CBS 119804 (Desjardins & Beremand 1987, Desjardins 2003) by PCR amplification of MAT idiomorphs using the primers pairs fusALPHAfor and fusALPHArev for MAT1-1, and fusHMGfor and fusHMGrev for MAT1-2 following conditions described elsewhere (Kerényi et al. 2004).
To induce the formation of a sexual morph in selected taxa, crossing experiments were carried out using the spermatization method in carrot agar (CA) (Klittich & Leslie 1988, Leslie & Summerell 2006). All crosses were tested in duplicates and checked weekly by visual inspection and using a dissecting microscope. Crosses were considered fertile if ascomata producing viable ascospores were observed. To assess progeny viability, ascospores extruding from fertile perithecia were collected with a sterile needle, suspended in a drop of sterile water and spread evenly on the surface of SNA plates, incubated overnight at 25 °C with alternating 12 h light/darkness, and checked for germination under the dissecting microscope (Leslie & Summerell 2006).
Species delimitation
To test for congruence of molecular species delimitation methods, a series of analyses were carried out following the multi-species coalescent approach. To infer species boundaries within the multi-locus dataset we used Species Tree And Classification Estimation, Yarely (STACEY v. 1.2.5; Jones 2017) as provided in the BEAST v. 2.6.3 package (Bouckaert et al. 2019), Additionally, three single locus delineation methods were employed i.e., Automatic Barcode Gap Discovery (ABGD; Puillandre et al. 2012), Assemble Species by Automatic Partitioning (ASAP; Puillandre et al. 2021), and the Bayesian implementation of the Poisson Tree Process model (bPTP; Zang et al. 2013). Gene datasets were analysed as full alignments (including missing data and all haplotypes), pruned alignments with minimal amount of missing data, and unique haplotype alignments. Alignments were collapsed to haplotypes using ALTER (Glez-Peña et al. 2010).
Input alignments for STACEY were prepared in BEAUTi v. 2.6.7 (Bouckaert et al. 2019). The multi-locus alignment was processed first as a complete dataset, and to ease result visualisation it was divided and analysed independently according to phylogenetic clade distribution as resolved in the 5-gene phylogenetic results. A strict clock was set up with a Yule model of Log Normal Birth and Death,CollapseHeight values between 1.0E-4 and 1.0E-5 were tested (between 1/100–1/1 000 of the expected species tree branch length, as indicated in the software manual, Jones 2019), resulting in similar species delimitations; hence collapseHeight was set to the default value (1.0E-4, roughly 1/100 of the branch length of the species tree). Other parameters were estimated by the software. After a preliminary run for parameter tunning according to software suggestions, the analyses ran for 50 M generations, in duplicates, saving trees every 5 000 generations. Results were checked for convergence on Tracer v. 1.7.2, log and tree files from duplicated analyses were combined using LogCombiner v. 2.6.7 and maximum clade credibility trees were calculated with TreeAnnotator v. 2.6.3, with a burn-in fraction of 10 %. STACEY results were processed with speciesDA, using the same collapseHeight setting as above, a simcutoff 0.95, and burn-in of 1 000. Similarity matrices were visualized from speciesDA outputs in RStudio 2022.12.0 build 353 (Posit team, Boston, MA USA) using the script plot.simmatrix.R (https://github.com/scrameri/smtools/tree/master/SpeciesDelimitation). Single locus delimitation analyses were run on their respective online web servers using alignment files as input for ABGD (https://bioinfo.mnhn.fr/abi/public/abgd/abgdweb.html) with default settings, and ASAP (https://bioinfo.mnhn.fr/abi/public/asap/) under a Jukes-Cantor (JC69) model. Ultrametric input trees for bPTP (https://species.h-its.org/ptp/) were constructed for each gene using BEAST v. 2.3.6, according to priors previously established for phylogenetic inference.
RESULTS
Phylogeny
The combined five gene dataset included 1 377 sequences (CaM 256, rpb1 288, rpb2 293, tef1 290, and tub2 250) from 293 strains, including Fusarium circinatum CBS 405.97 (Fusarium fujikuroi SC) as the outgroup taxon. Including alignment gaps, the multilocus analysis included 4 524 characters (CaM 501, rpb1 1 502, rpb2 1 522, tef1 558, and tub2 441), of which, 2 706 were conserved (CaM 259, rpb1 960, rpb2 996, tef1 251, and tub2 240), 1 782 were variable (CaM 230, rpb1 528, rpb2 525, tef1 303, and tub2 196), 1 486 were phylogenetically informative (CaM 177, rpb1 457, rpb2 442, tef1 265, and tub2 145), and 2 331 were Bayesian unique site patterns (CaM 299, rpb1 694, rpb2 711, tef1 363, and tub2 264).
Best substitution models for ML as calculated by ModelFinder in IQ-TREE were TNe+G4 for CaM and rpb2, TNe+I+G4 for rpb1 and tub2, and TIM2e+I+G4 for tef1; and were codified into the analyses. Independent ML analyses on IQ-TREE and RAxML resulted in identical topologies with best-tree scores of -38115.028 and -38359.969, respectively. Sequences from a single isolate (Fusarium cortaderiae NRRL 29306) failed the composition chi2 test (gap ambiguity 79.13 %, and p-value 1.81 %) as implemented in IQ-TREE.
Bayesian best-fit substitution models selected according to the Akaike criterion in MrModelTest were SYM+I+G for CaM, rpb1 and tub2; and GTR+I+G for rpb2 and tef1. The BI analysis lasted 4.38 M generations, and retained 8 762 trees, of which 6 572 were used to calculate the 50 % majority rule tree and posterior probabilities. The topologies observed for ML and BI analyses were congruent for supported nodes (ML-UFBootBS ≥ 0.95 %, ML-BS ≥ 70 %, and BI-PP ≥ 0.95), thus Fig. 1 shows the IQ-TREE ML topology with ML-UFBootBS, ML-BS, and BI-PP values plotted at the nodes.
Fig. 1.
Maximum likelihood (IQ-TREE) phylogeny of the Fusarium sambucinum species complex (FSAMSC) inferred from combined, partial CaM, rpb1, rpb2, tef1, and tub2 loci. The tree is rooted to Fusarium circinatum CBS 405.97 (F. fujikuroi species complex). Numbers at the nodes are IQ-TREE ultra-fast bootstrap (BS) (UFBoot) values ≥ 95 %, followed by RAxML BS ≥ 70 %, and MrBayes posterior probability (PP) ≥0. 95. Full supported branches (UFBoot and BS = 100 %, and PP = 1) are indicated in bold. Strain codes in bold correspond to type strains; and ex-epitype, ex-isotype, ex-neotype, and ex-type strains are indicated with ET, IT, NT, and T, respectively. Strains accessions are followed by substrate/host information (in red), and geographic origin (in blue). Names of novel taxa described in this paper are in bold.
The combined 5-gene phylogeny resolved 75 species-level monophyletic groups within the ingroup taxa. One recently described additional taxon was excluded from the final analyses (Fusarium guizhouense; He et al. 2024) due to incomplete sequence data and short length of the available sequences; however, based on separate analyses based on available rpb1, rpb2, and tef1 sequences, it resolved as a genetically exclusive clade sister to the unnamed phylogenetic species FSAMSC11 (data not shown). Species in FSAMSC were distributed among six main clades i.e., Brachygibbosum, Gladiolum, Graminearum, Longipes, Sambucinum, and Sporotrichioides; and agreed with terminal clades disposition as in Laraba et al. (2021), differing barely in the position of terminal clades and the presence of additional phylogenetic species.
Phylogenetic position and sexual crosses of Fusarium sambucinum
Fourteen strains morphologically matching F. sambucinum were obtained from four specimens collected from the species type host and country. Isolations were made from conidia and ascospores. All the newly isolated strains clustered within the phylogenetic clade assigned to F. sambucinum with CaM, rpb1, rpb2, tef1, and tub2 single gene and combined datasets. MAT-PCR assays showed the 14 strains to be heterothallic, with eight (57 %) of them carrying the MAT1-1 idiomorph (CBS 151939, CBS 151940, CBS 151942, CPC 45318, and CPC 45320, isolated from sporodochial conidia; and CBS 151944, CPC 45323, and CPC 45324 isolated from ascospores), and six (43 %) carrying the MAT1-2 idiomorph (CBS 151802, CBS 151941, CBS 151943, CPC 45314, CPC 45316, and CPC 45319, all of them isolated from ascospores). The tester strain CBS 119804 was confirmed to carry the MAT1-2 idiomorph and showed high female fertility. Sexual crosses with the tester strain were positive for six strain combinations (CBS 119804 × strains CBS 151940, CBS 151942, CBS 151944, CPC 45320, CPC 45323, and CPC 45324) and confirmed the previous morphological and phylogenetic inferences. Therefore, an epitype was selected (CBS H-25241), with an associated viable ex-type culture (CBS 151942) to fix the application of the name F. sambucinum to a well-defined phylogenetic clade.
Phylogenetic species groups and general morphological aspects
Distinct colonial and conidial morphological characters distinguish the different main clades of FSAMSC. An overview of important morphological features from observations of type material and available bibliographical data is shown in Table S1 and Fig. 2.
Fig. 2.
Distributions of overall macroconidial sizes of species in FSAMSC. Average sizes are indicated by white dashed lines. Typical macroconidial morphology for each species is shown in the right-most column (not up to scale). † Only mesoconidia produced. ‡ No sporodochial conidia produced.
The fully supported (ML-UFBootBS and ML-BS = 100 %, and BI-PP = 1) Brachygibbosum clade included 13 species, 12 of which were highly supported by all three phylogenetic algorithms. The novel species F. mucronatum received low ML support with either IQ-TREE and RAxML analyses; however, it was resolved as a monophyletic clade by BI. Due to lack of strains for morphological analysis, the phylogenetic species FSAMSC31 is not formally described here. Species in the Brachygibbosum clade typically formed red to coral-coloured colonies on PDA, and falcate multiseptate conidia were present in all species, commonly exhibiting slightly elongated apical cells and tapering notoriously, with exception of F. subflagellisporum which formed additional whip-like sporodochial conidia. Microconidia were present either often or rarely in aerial conidiophores with exception of F. cuspidatum, F. hamatum and F. mucronatum.
The Gladiolum clade received full support in all three analyses, and encompassed two fully supported phylogenetic species, one of which (FSAMSC7) was not studied here due to unavailability of living cultures. Fusarium gladiolum was characterized by yellow colonies on PDA, and falcate, robust, gently curved macroconidia without elongated appendages formed on both aerial and sporodochial conidiophores, lacking microconidia.
The speciose Graminearum clade received partial support (ML-UFBootBS = 100 %, ML-BS <; 70 %, and BI-PP = 0.95) and included 22 species, of which 20 were supported by all three analyses. Fusarium lunulosporum was recovered with confidence only by IQ-TREE ML with ML-UFBootBS = 97 %, while F. aethiopicum was recovered as an unresolved polytomy by ML, and BI analyses of the combined 5-loci, as well as all the single-gene datasets included in this study. Species in this group presented either red, orange or brown pigmentation on PDA, microconidia were rarely present, and form falcate conidia without elongated appendages, either almost straight, equilaterally to slightly asymmetrically curved (segregate species of F. graminearum s. lat. in addition to F. dactylidis, F. praegraminearum, F. pseudograminearum, and F. subtropicale), with a more pronounced curvature and relatively wide for its length (F. culmorum, F. cerealis), or curved and slender (F. lunulosporum).
The fully supported Longipes clade included 10 species, all of which received high support on both ML and BI analyses. Species in this group produced red colonies on PDA, with exception of F. carinatum, F. pratense, and F. vermicularioides which showed yellow pigmentation on PDA; the presence of microconidia and chlamydospores was variable, and with exception of F. carinatum, F. pratense, and F. tropicale (macroconidia falcate with pointy, not elongated apical cells) macroconidia showed moderately to largely elongated, whip-like apical cells.
The fully supported Sambucinum clade encompassed 17 species, 16 of which were supported by all three phylogenetic methods. Fusarium bananae was supported only by BI in the combined analysis. Two phylogenetic species (FSAMSC11 and FSAMSC13) were not studied morphologically due to unavailability of living cultures for this study, and an additional species (F. guizhouense) belonging to the Sambucinum clade was excluded from the analysis (see comment in phylogenetic results above). Species of the Sambucinum clade were morphologically diverse, either in colony pigmentation (red and yellow shades on PDA), presence and absence of microconidia, mesoconidia; and macroconidial shapes, with predominance of falcate, “sambucinum-like” conidia, but also encompassing species with “graminearum-like” conidia or whip-like, elongated apical cells.
The fully supported Sporotrichioides clade included 11 species, 10 of which were supported by ML and BI methods. The ex-type of Fusarium armeniacum (CBS 485.94) was resolved as an unsupported lone lineage in the combined phylogenetic analysis; however, it was confidently resolved in the rpb1, rpb2, tef1 and tub2 single locus analyses. One phylogenetic species (FSAMSC2) was not studied morphologically due to unavailability of living strains for this study. Two distinct morphological groups (clades A and B) were observed in the Sporotrichioides clade, which aligned with the phylogenetic species distribution. Both subclades contained species with yellow or red PDA colonies and produced combinations of micro-, and macroconidia, or mesoconidia (F. sporotrichioides). Species in subclade A, produced falcate to lunate, relatively short macroconidia without elongated apical cells, or completely lacked macroconidia (F. langsethiae and F. sibiricum), while microconidia were always present. Species in subclade B included two distinct types of macroconidia: often falcate, with a slight dorsiventral curve accentuated by a moderately elongated and curved apical cell, accompanied by a shorter type of falcate macroconidia without apical elongation; while the presence of microconidia was variable.
Species delimitation
Analyses were done separately for the full dataset (including all available strains and missing data), a pruned dataset with reduced missing data, and a dataset reduced to unique haplotypes. The results were in general consistent, differing mostly in the over partitioning of taxa with the larger datasets. Maximum clade credibility trees were calculated from 18 000 sampled trees after discarding 2 000 samples as burn-in.
The STACEY coalescent delimitation method resolved 77 groups and was congruent with results from the 5-loci phylogenetic analysis. With exception of phylogenetic species FSAMSC23, included within the circumscription of F. brachygibbosum, and over partitioning of F. graminearum s. str., the analysis supported the retention of all currently known phylogenetic species, plus 30 novel undescribed taxa, and additional undescribed phylogenetic clades (FSAMSC2, FSAMSC7, FSAMSC11, FSAMSC13, and FSAMSC31). An overview of groups obtained by coalescent and phylogenetic methods from a dataset with reduced missing data is shown in Fig. 3, mapped against the results of a 5-gene IQ-TREE multilocus phylogenetic analysis. Cluster analyses of distribution of probabilities from STACEY outputs for separated analyses according to main clades of FSAMSC delineated 13 groups for the Brachygibbosum clade (Fig. 4), 22 groups for the Graminearum clade (Fig. 5), 10 groups for the Longipes clade (Fig. 6), 17 groups for the Sambucinum clade (Fig. 7); and 2 and 11 groups for the Gladiolum and Sporotrichioides clades, respectively (Fig. 8). A noticeable intraspecific genetic structure was observed for F. graminearum s. str., F. longipes, F. pseudograminearum, and strains here assigned to the novel species F. hamatum, which explained the different number of resolved species between phylogenetic and coalescent methods. However, these were all retained as single groups in accordance with combined results from morphological, phylogenetic, and STACEY analyses.
Fig. 3.
Summary of species delimitation results by phylogenetic and coalescent methods of a reduced dataset to minimize missing data. Analyses referred to as “Combined” and STACEY (Species Tree And Classification Estimation) are based on multilocus datasets. Phylo = single-locus phylogeny, ABGD = Automatic Barcode Gap Discovery, ASAP = Assemble Species by Automatic Partitioning, bPTP = Bayesian implementation of the Poisson Tree Process model. Samples are split and colour coded according to hypothetical species and monophyletic clades and mapped against the result of a 5-gene IQ-TREE multilocus phylogenetic analysis. Blank spaces correspond to missing sequence data for the respective strain. Type strains are indicated in bold, followed by ET, IT, NT, and T to denote, respectively, ex-epitype, ex-isotype, ex-neotype, and ex-type strains.
Fig. 4.
Results of STACEY species delimitations of the Brachygibbosum clade of FSAMSC. Similarity matrix showing the probability (Bayesian posterior probabilities, PP) of each strain pair to belong to the same species group. Intensity of coloured blocks is equivalent to PP values, where white = 0 and dark green = 1. The matrix is mapped to a maximum credibility tree of the Brachygibbosum clade generated in BEAST, based on combined CaM, rpb1, rpb2, tef1, and tub2 loci. Values at the nodes indicate PP ≥ 0.95. Branches in bold indicate PP = 1. Ex-type strains are indicated by T and bold font. The inset shows the position of the Brachygibbosum clade with respect to the phylogeny of the entire FSAMSC as shown in Fig. 1
Fig. 5.
Results of STACEY species delimitations of the Graminearum clade of FSAMSC. Similarity matrix showing the probability (Bayesian posterior probabilities, PP) of each strain pair to belong to the same species group. Intensity of coloured blocks is equivalent to PP values, where white = 0 and dark green = 1. The matrix is mapped to a maximum credibility tree of the Graminearum clade generated in BEAST, based on combined CaM, rpb1, rpb2, tef1, and tub2 loci. Values at the nodes indicate PP ≥ 0.95. Branches in bold indicate PP = 1. Ex-epitype, and ex-type strains are indicated by ET, and T, respectively; and bold font. The inset shows the position of the Graminearum clade with respect to the phylogeny of the entire FSAMSC as shown in Fig. 1.
Fig. 6.
Results of STACEY species delimitations of the Longipes clade of FSAMSC. Similarity matrix showing the probability (Bayesian posterior probabilities, PP) of each strain pair to belong to the same species group. Intensity of coloured blocks is equivalent to PP values, where white = 0 and dark green = 1. The matrix is mapped to a maximum credibility tree of the Longipes clade generated in BEAST, based on combined CaM, rpb1, rpb2, tef1, and tub2 loci. Values at the nodes indicate PP ≥ 0.95. Branches in bold indicate PP = 1. Ex-neotype, and ex-type strains are indicated by NT, and T, respectively; and bold font. The inset shows the position of the Longipes clade with respect to the phylogeny of the entire FSAMSC as shown in Fig. 1.
Fig. 7.
Results of STACEY species delimitations of the Sambucinum clade of FSAMSC. Similarity matrix showing the probability (Bayesian posterior probabilities, PP) of each strain pair to belong to the same species group. Intensity of coloured blocks is equivalent to PP values, where white = 0 and dark green = 1. The matrix is mapped to a maximum credibility tree of the Sambucinum clade generated in BEAST, based on combined CaM, rpb1, rpb2, tef1, and tub2 loci. Values at the nodes indicate PP ≥ 0.95. Branches in bold indicate PP = 1. Ex-epitype, ex-isotype, and ex-type strains are indicated by ET, IT, and T, respectively; and bold font. The inset shows the position of the Sambucinum clade with respect to the phylogeny of the entire FSAMSC as shown in Fig. 1.
Fig. 8.
Results of STACEY species delimitations of the Gladiolum and Sporotrichioides clades of FSAMSC. Similarity matrix showing the probability (Bayesian posterior probabilities, PP) of each strain pair to belong to the same species group. Intensity of coloured blocks is equivalent to PP values, where white = 0 and dark green = 1. The matrix is mapped to a maximum credibility tree of the Gladiolum and Sporotrichioides clades generated in BEAST, based on combined CaM, rpb1, rpb2, tef1, and tub2 loci. Values at the nodes indicate PP ≥ 0.95. Branches in bold indicate PP = 1. Ex-epitype, and ex-type strains are indicated by ET, and T, respectively; and bold font. The inset shows the position of the Gladiolum and Sporotrichioides clades with respect to the phylogeny of the entire FSAMSC as shown in Fig. 1.
Results from single locus coalescent analyses were incongruent between methods and gene regions, irrespective of the dataset analysed. The estimated number of minimal clusters ranged from 30 (ASAP-rpb1) to 93 (bPTP-CaM). Lumping or over splitting of taxa was conflictive between partitions and analyses with all datasets studied and were incongruent with species designations according to morphological identification, monophyletic clades and STACEY groups. The putative species are shown in Fig. 3.
TAXONOMY
Fusarium acaciae-mearnsii O’Donnell et al., Fungal Genet. Biol. 41: 619. 2004. Fig. 9.
Fig. 9.
Fusarium acaciae-mearnsii (ex-type CBS 110254). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–K. Conidiophores and conidiogenous cells. L. Macroconidia. Scale bars: E–I, L = 10 μm; J, K = 5 μm .
Descriptions and illustrations: O’Donnell et al. (2004), Aoki et al. (2012).
Typus: South Africa, KwaZulu-Natal, Pietermaritzburg, Acacia mearnsii, 18 Jul. 1997, J. Roux (holotype BPI 843477, ex-type culture CBS 110254 = FRC R-9629 = MRC 5120 = NRRL 26754).
Additional materials examined: Australia, Queensland, from soil, before 1 Feb. 1978, L.W. Burgess, culture CBS 123662 = FRC R-4412 = NRRL 34207. South Africa, unknown location, Acacia mearnsii, before Jul. 1997, J. Roux, cultures CBS 110253 = FRC R-9628 = MRC 2458 = NRRL 26752, CBS 110255 = FRC R-9630 = MRC 5122 = NRRL 26755.
Notes: Fusarium acaciae-mearnsii, together with F. asiaticum, F. austroamericanum, F. boothii, F. brasilicum, F. cortaderiae, F. graminearum s. str., F. meridionale, and F. mesoamericanum are the nine original segregates of F. graminearum s. lat. (O’Donnell et al. 2004), also known as the Fusarium graminearum species complex (Starkey et al. 2007, Aoki et al. 2012). All the above species, plus F. aethiopicum, F. gerlachii, F. louisianense, F. nepalense, F. ussurianum, and F. vorosii represent a homogenous assemblage of cryptic species. They are all agents of Fusarium head blight of cereals, and distinguished primarily by DNA polymorphisms, and minute details in conidial morphology, with documented differences, although with some overlap, in mycotoxin production patterns (Laraba et al. 2021). Most of these species are only known from cryptic morphological descriptions, single nucleotide polymorphism-based descriptions, and line drawings, hence they are illustrated here (Fig. 9) with general morphological features listed in Table S1. It has been suggested that due to known cross-fertility, a wider species delimitation should be favoured for F. graminearum s. lat. (Desjardins 2003, Leslie & Summerell 2006, Leslie & Bowden 2008). Nevertheless, our multilocus phylogeny and coalescence species delimitation results support the retention of these lineages as separate species as well as recent phylogenomic data (Han et al. 2023). Morphological differences, as indicated by O’Donnell et al. (2004) and Aoki et al. (2012), although difficult to assess, are supported by our observations of the ex-types (Table S1, Fig. 2).
Fusarium acaciae-mearnsii differs from other members of F. graminearum s. lat. by forming asymmetrical, gradually curved, 4.5–5 µm wide, 5-septate conidia with narrow apical beaks, which are often wider below the median (O’Donnell et al. 2004, Aoki et al. 2012). This species is known from Australia and South Africa, mostly from Fabaceae and Poaceae hosts, and soil (O’Donnell et al. 2004, Aoki et al. 2012, Farr et al. 2023).
Fusarium aethiopicum O’Donnell et al., Fungal Genet. Biol. 45: 1521. 2008. Fig. 10.
Fig. 10.
Fusarium aethiopicum (ex-type CBS 122858). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophores and conidiogenous cells. G, H. Aerial conidiophores and conidiogenous cells. I. Microconidia. J. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: O’Donnell et al. (2008b), Aoki et al. (2012).
Typus: Ethiopia, Bure district, west Gojam zone of Amhara region, Triticum aestivum stored grain, 26 Feb. 2007, D. Aberra (holotype BPI 878409, ex-type culture CBS 122858 = FRC R-10088 = NRRL 46726).
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Fusarium aethiopicum is morphologically indistinguishable from F. asiaticum, F. graminearum, and F. vorosii; all the mentioned species producing asymmetric, slightly curved 5-septate conidia, 4.5–5 or > 5 µm at its widest point above the median (O’Donnell et al. 2008b, Aoki et al. 2012). Nevertheless, F. aethiopicum has a larger overall main conidial size compared to the species mentioned above (50.8 × 5.1 µm, vs 38.8 × 4.7 µm, 46.5 × 4.7 µm, and 41.6 × 5.1 µm, for F. asiaticum, F. graminearum and F. vorosii, respectively), and although not described in the protologue, 0–3-septate, obovoid to ellipsoidal microconidia were often seen formed on F. aethiopicum aerial conidiophores (Fig. 10). Microconidia were not observed on any of the three species in comparison.
Fusarium aethiopicum is so far known only from Triticum sp., from Ethiopia (Aoki et al. 2012). In contrast, the morphologically comparable species, and particularly important in F. graminearum, have a cosmopolitan distribution and are known from a wider host range, mostly Poaceae, but also Malvaceae, and Solanaceae (Farr et al. 2023).
Fusarium agreste Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855093. Fig. 11.
Fig. 11.
Fusarium agreste (ex-type CBS 120993). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Aerial conidiogenous cells. I. Microconidia. J. Sporodochial conidiophores and conidiogenous cells. K. Macroconidia. Scale bars: G, H = 5 μm; I–K = 10 μm.
Etymology: From Latin agrestis (“rural, rustic, wild”). Referring to the origin of the type.
Sporodochia abundant on the surface of carnation leaves and agar surface, ochreous, luteous to bright orange coloured. Sporodochial conidiophores branching laterally, 16–32 µm tall, bearing terminal whorls of monophialides. Sporodochial conidiogenous cells monophialidic, doliiform to subcylindrical, 8–16.5 × 2.5–5.5 µm, smooth- and thin-walled, with a minute or lacking apical collarette. Sporodochial conidia falcate, dorsiventrally curved, apical cell blunt, slightly curved; basal cell well-developed, foot-shaped, (1–)3–5-septate, (16–)28–41.5(–55) × (4–)4.5–5.5(–6.5) µm (av. 34.6 × 5 μm), mostly 3-septate, (22–)27.5–36(–40) × 4–6.5 µm (av. 31.7 × 4.9 μm). Aerial conidiophores erect or prostrate on substrate mycelium, unbranched, reduced to monophialides forming laterally on aerial mycelium. Aerial conidiogenous cells monophialidic, doliiform, ampulliform to subcylindrical, 8–16.5 × 3.5–5.5 µm, smooth- and thin- walled, apical collarette inconspicuous, or reduced to lateral phialidic pegs, 1–4.5 × 2–3 µm, giving rise to falcate conidia indistinguishable from sporodochial conidia and rarely to microconidia. Microconidia very rare, ovoid, ellipsoidal to allantoid, 0(–1)-septate, 9–11.5 × 3–4 µm. Chlamydospores not formed but swollen cells commonly observed on the remains of senescent conidia.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 6.8–7.8 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface scarlet to rust, felty to woolly, flat, aerial mycelium abundant, ochreous, margin regular and filiform; reverse brick to rust, isabelline at centre. On OA, amber to pure yellow, sulphur yellow at periphery, flat, powdery to velvety, margin regular and filiform; reverse amber to honey.
Typus: South Africa, Eastern Cape Province, Butterworth, from soil debris, before 1 Mar. 1987, collector unknown (holotype designated here CBS H-25432, ex-type culture CBS 120993 = FRC R-8575 = MRC 6971).
Additional material examined: New Zealand, Auckland, from Sechium edule, before 30 Dec. 1999, J.M. Dingley, culture NRRL 29296 = FRC R-9861 = ICMP 5173.
Notes: Fusarium agreste, previously recognized as lineage Fusarium sp. nov. 8 in Laraba et al. (2021), resolved as the closest phylogenetic sibling of F. sambucinum, clustering in an intermediate position between F. sambucinum and F. symmetricum, the three species producing relatively fast growing, red colonies on PDA and up to 5-septate sporodochial conidia of similar width. Nevertheless, phylogenetic analyses, coupled with multilocus and most single-locus coalescent-based analyses support the recognition of this clade as a genetically exclusive novel taxon, which is reinforced by morphological features. Fusarium agreste can be distinguished from F. sambucinum and F. symmetricum by its longer and robust conidia with well-developed foot-cells and by the production of 0–1-septate microconidia on aerial conidiophores. Both F. agreste and F. symmetricum are so far known to occur in the Southern Hemisphere; F. agreste in South Africa and New Zealand, while the latter species is known from Cactaceae in South America. In contrast, F. sambucinum has a worldwide distribution and has been recorded from more than 150 different substrates, including soil, animals (among them insects, and mammals, including human), and more than 130 plant host; mostly from Fabaceae, Poaceae, Pinaceae and Solanaceae (Farr et al. 2023).
Fusarium amblysporum Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855094. Fig. 12.
Fig. 12.
Fusarium amblysporum (ex-type CBS 151809). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G, H. Chlamydospores. I. Aerial conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Greek amblyo (“blunt”). Referring to the macroconidial blunt apical cells.
Sporodochia abundant on the surface of carnation leaves, agar surface, and aerial mycelium, straw, pale luteous to amber coloured. Sporodochial conidiophores branching laterally and irregular, and verticillately, 44–140 µm tall, bearing lateral and terminal monophialides, single or grouped in verticils. Sporodochial conidiogenous cells monophialidic, subcylindrical, 9.5–14.5 × 3.5–5 µm, smooth, thin-walled, with a short and flared apical collarette. Sporodochial conidia falcate, gently dorsiventrally curved, with blunt, sometimes slightly hooked apical cell; basal cell well-developed foot-shaped, (3–)5–6(–7)-septate, (31–)40.5–54(–67) × 4.5–7 µm (av. 47.4 × 5.4 µm), mostly 5-septate, (38–)42–53(–61.5) × (4.5–)5–6 µm (av. 47.6 × 5.4 µm). Aerial conidiophores reduced to monophialides forming laterally on aerial mycelium or submerged in agar, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical or reduced to short phialidic pegs, 4.5–19 × 2–5 µm, giving rise to falcate conidia indistinguishable from sporodochial conidia. Chlamydospores obovate to globose, terminal, or intercalary on hyphae, or lateral on short stipes, 6.5–13 µm diam, solitary or forming short chains.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.9–8.5 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface at first membranous, becoming felty to cottony, white, peach, sulphur yellow or rosy buff, flat, aerial mycelium abundant, moisty at the centre, margin regular, filiform to undulate; reverse sulphur yellow to straw. On OA, primrose to pure yellow, white at centre, flat, cottony to woolly, moisty at the centre, margin regular, filiform to undulate; reverse amber to honey.
Typus: New Zealand, from Cucurbita maxima, before 30 Jun. 1998, G.F. Laundon (holotype designated here CBS H-25428, ex-type culture CBS 151809 = BBA 64376 = NRRL 28571).
Additional materials examined: Austria, from Triticum sativum, 1985, H. Nirenberg, culture CBS 128.95 = BBA 64568. Hungary, from Cucumis melo var. cantalupensis, before 25 Apr. 2016, culture BBA 71677 = DSM 116910. South Africa, Zinggayi, from debris, before 1 Feb. 1986, unknown collector, culture CBS 151954 = NRRL 66929 = FRC R-8234, FRC R-8227 = KOD 2017 = NRRL 64078, from Solanum tuberosum, before 25 Apr. 2016, A. Logrieco, culture BBA 65418 = DSM 116523, from soil debris, before 25 Apr. 2016, A. Logrieco, culture BBA 65419 = DSM 116524. Sweden, from Spinacia oleracea, 1991, M. Larsson, culture CBS 148.95 = BBA 65893.
Notes: Previously assigned to lineage Fusarium sp. nov. 12 (Laraba et al. 2021), F. amblysporum is here described from a set of strains previously stored in culture collections as either F. compactum, F. sambucinum or F. venenatum. Fusarium compactum, a member of the Fusarium incarnatum-equiseti SC produces similar PDA colonies to F. amblysporum, and all three species mentioned above produce wide (averaging > 5.4 µm wide) sporodochial conidia. Conidia of F. amblysporum, F. sambucinum and F. venenatum, however, all differ from those of F. compactum by being more uniform in shape, robust, with a less evident tapering. Fusarium amblysporum can be distinguished from its FSAMSC counterparts above, by its slightly faster growth rate and pigmentation on PDA, which can vary substantially from white to rosy buff, but not reaching the intense red colour observed in F. sambucinum and F. venenatum.
Fusarium armeniacum (G.A. Forbes et al.) L.W. Burgess & Summerell, Mycotaxon 75: 347. 2000. Fig. 13.
Fig. 13.
Fusarium armeniacum (ex-type CBS 485.94). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G. Aerial conidiophore and conidiogenous cells. H. Microconidia. I. Macroconidia. Scale bars = 10 μm.
Basionym: Fusarium acuminatum subsp. armeniacum G.A. Forbes et al., Mycologia 85: 120. 1993.
Synonym: Fusarium armeniacum (G.A. Forbes et al.) L.W. Burgess & Summerell, Stud. Mycol. 98: 86. 2021.
Descriptions and illustrations: Burgess et al. (1993), Burgess & Summerell (2000), and Leslie & Summerell (2006).
Typus: Australia, New South Wales, Edgeroi, Triticum aestivum grain, unknown collection date and collector, dep. P.E. Nelson (holotype DAR 67507, ex-type culture ATCC 90020 = CBS 485.94 = FRC R-9335 = IMI 352099 = MRC 6230 = NRRL 26908 = NRRL 25141 = NRRL 29133).
Notes: Originally described as a subspecies of F. acuminatum (a member of the F. tricinctum SC), differing by longer macroconidia, and faster chlamydospores and colony development (Burgess et al. 1993). Crous et al. (2021) created an isonym when attempting to validate a presumed wrong citation of the basionym (Burgess et al. 1993), which was, however, valid at the time.
Microconidia are cited as not present in F. armeniacum in recent literature (Leslie & Summerell 2006, Nichea et al. 2021), but they were described in the protologue of F. armeniacum subsp. armeniacum (Burgess et al. 1993) which is confirmed here from observations of the ex-type. The absence of microconidia thus, is not a valid character for delimitation of F. armeniacum and its closest phylogenetic relative F. chaquense (Nichea et al. 2021). The two species, however, can be distinguished based on their sporodochial macroconidial shape, both of which present similar overall curvatures, but those of F. armeniacum tend to exhibit an abrupt curvature of it apical cells. Other related species in the Sporotrichioides clade include F. leptum and F. parabolicum, both of which produce yellow PDA colonies (vs red PDA colonies in F. armeniacum) and shorter sporodochial conidia, which shorter apical cells.
Fusarium armeniacum has been isolated mostly from soil, and Poaceae hosts, but has also been described from animals (including humans, Al-Hatmi 2016, O’Donnell et al. 2016), Campanulaceae, Fabaceae, Pinaceae, and Solanaceae in Africa, Australia, China, and North America. Reported host species include Austrostipa aristiglumis, Festuca sp., Glycine max, Platycodon grandiflorus, Pinus elliottii, Solanum lycopersicum, Sorghum sp., Triticum dicoccum, and Zea mays (Farr et al. 2023)
Fusarium asiaticum O’Donnell et al., Fungal Genet. Biol. 41: 619. 2004. Fig. 14.
Fig. 14.
Fusarium asiaticum (ex-type CBS 110257). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F, G. Sporodochial conidiophores and conidiogenous cells. H, I. Aerial conidiogenous cells. J. Macroconidia. Scale bars: F, H, J = 10 μm; G, I = 5 μm.
Descriptions and illustrations: O’Donnell et al. (2004), Aoki et al. (2012).
Typus: Japan, Hordeum vulgare grain, before 1 Jun. 1980, unknown collector (holotype BPI 843478, ex-type culture CBS 110257 = FRC R-5469 = MRC 1963 = NRRL 13818).
Additional materials examined: China, Shanghai Province, from Triticum sp., before 13 Mar. 2000, Y.Z. Wang, culture CBS 110258 = FRC R-9627 = DAOM 211601 = NRRL 26156. Japan, from Hordeum vulgare, before 13 Mar. 2000, unknown collector, dep. K. O’Donnell, culture CBS 110256 = ATCC 28114 = FRC R-9620 = ATCC 46780 = NRRL 6101. Nepal, Ghauli farm, from Zea mays, before 31 Mar. 1993, unknown collector, dep. K. O’Donnell, culture CBS 110259 = FRC R-9402 = NRRL 28720.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Hybridization of F. asiaticum with F. graminearum and F. meridionale has been described (O’Donnell et al. 2000, Jurgenson et al. 2002) which indicates an incomplete reproductive isolation between members of F. graminearum s. lat. (O’Donnell et al. 2004).
Fusarium asiaticum has been reported as indistinguishable from F. aethiopicum, F. graminearum and F. vorosii when 5-septate conidia are compared; all four species being distinguished within the F. graminearum s. lat. clade by its gradually curved, asymmetric conidia, typically wider above the median and lacking narrow apical beaks (Aoki et al. 2012). However, when total ranges of sporodochial conidia are compared, F. asiaticum can be differentiated by having shorter conidia than F. aethiopicum and F. graminearum (av. 38.8 µm long vs 50.8 µm and 46.5 µm for F. aethiopicum and F. graminearum, respectively), which are also narrower than F. aethiopicum and F. vorosii (av. 4.7 µm wide vs 5.1 µm for F. aethiopicum and F. vorosii).
The dataset studied here includes only Asian isolates, where F. asiaticum is the most important agent of fusarium head blight of rice (Aoki et al. 2012). However, F. asiaticum has been also recorded from air, soil, other Poaceae hosts (Hordeum vulgare, Lolium multiflorum, Triticum aestivum, and Zea mays), Fabaceae (Glycine max), Musaceae (Musa nana), Orchidaceae (Bletilla striata), Podocarpaceae (Podocarpus macrophyllus), Rosaceae (Prunus persica), and Rubiaceae (Paederia foetida); with occurrence in Africa, Europe, and North America, most likely as an introduced species (Aoki et al. 2012, Wang et al. 2022, Farr et al. 2023).
Fusarium austroamericanum T. Aoki et al., Fungal Genet. Biol. 41: 617. 2004. Figs 15, 16.
Fig. 15.
Fusarium austroamericanum (ex-type CBS 110244). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–H. Sporodochial conidiophores and conidiogenous cells. I. Macroconidia. Scale bars = 10 μm.
Fig. 16.
Fusarium austroamericanum sexual morph (CBS 110245). A, B. Perithecia. C. Detail of perithecial wall cells. D, E. Asci. F. Ascospores. Scale bars: A, B = 100 μm;, C–F = 10 μm.
Descriptions and illustrations: O’Donnell et al. (2004), Aoki et al. (2012).
Typus: Brazil, unknown host, 1960, J.C. Lewis (holotype BPI 843473, ex-type culture CBS 110244 = ATCC 46032 = FRC R-9634L = NRRL 2903).
Additional materials examined: Brazil, from Zea mays for equine feed, before Sep. 1998, unknown collector, dep. K. O’Donnell, culture CBS 110246 = FRC R-6964 = NRRL 28718. Venezuela, from herbaceous vine, before Jul. 1998, unknown collector, dep. K. O’Donnell, CBS H-25405, culture CBS 110245 = BBA 65928 = FRC R-9632 = GJS 90-246 = NRRL 28585.
Description of the sexual morph: Perithecia gregarious, rarely solitary, globose to subglobose, 190–300 µm diam, non-papillate to slightly papillate, superficial, non-stromatic or seated on a thin stroma; dark purple to black coloured, purple in water, turning blue in 3 % KOH, and red in 100 % lactic acid. Perithecial wall rough, tuberculate, composed of two layers: external layer 20–60 µm thick, composed of globose, thick-walled, pigmented cells, 13–25 µm diam, cell walls 0.5–3.5 µm wide; inner layer 11.5–31 µm thick, composed of compressed, thin-walled, non-pigmented cells. Asci unitunicate, clavate with a short stipe, 8-spored, 52–87.5 × 6.5–13 µm, thin-walled, with simple apex. Ascospores fusoid to ellipsoidal, straight or gently curved, smooth, at first hyaline becoming pale brown after discharge, 3-septate, (19.5–)20.5–26(–30.5) × 5–6.5 µm (av. 23.3 × 5.8 μm), often constricted at the septa.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. In agreement with mating types as determined by O’Donnell et al. (2004), a sexual morph developed homothallically for this species (strain CBS 110245), which is described and illustrated Fig. 16. The sexual morph of F. austroamericanum cannot be confidently distinguished from F. graminearum s. str. on morphological characters; although, at 5–6.5 µm wide, ascospores of F. austroamericanum are consistently wider than those of F. graminearum [3.5–4.5 µm (Samuels 2006) or 3–4 µm (Booth 1971, CABI 1973)]. Nevertheless, a morphological reassessment of sexual morphs of F. graminearum s. lat. is needed following the current taxonomy of this group (O’Donnell et al. 2004). Morphological identification of the asexual morph is possible by its narrow (< 4.5 µm wide) and almost straight, asymmetric conidia, often widest in the median (O’Donnell et al. 2004, Aoki et al. 2012).
Fusarium austroamericanum is known from North and South America, and Nepal; and has been associated mostly with Poaceae (Hordeum vulgare, Triticum sp., and Zea mays) but also reported from an unidentified herbaceous vine, soybean, and insects (Aoki et al. 2012, O’Donnell et al. 2021, Farr et al. 2023).
Fusarium bananae Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855095. Fig. 17.
Fig. 17.
Fusarium bananae (ex-type CBS 151937). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–G. Sporodochia. H–J. Sporodochial conidiophores and conidiogenous cells. K–M. Aerial conidiophores and conidiogenous cells. N. Macroconidia. Scale bars = 10 μm.
Etymology: From Greek banánes (“banana”). Referring to the origin of isolation of the type.
Sporodochia abundant on the surface of carnation leaves and aerial mycelium, erumpent, orange to ochreous coloured. Sporodochial conidiophores branching laterally and profusely, 24–32 µm tall, bearing terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical to navicular, (8–)10.5–15(–17) × 3–5.5 µm, smooth, thin-walled, with a short apical collarette, rarely proliferating percurrently. Sporodochial conidia falcate to lunate, dorsiventrally curved, with blunt or pointed apical cell; basal cell poorly to well-developed foot-shaped, (1–)3–5-septate, (14.5–)34.5–57.5(–66.5) × 4–6 µm (av. 46 × 4.9 μm), mostly 5-septate, (42.5–)47–60.5(–66) × (4–)5–6 µm (av. 53.8 × 4.9 μm). Aerial conidiophores erect, rarely laterally and sparsely branched, 23–58 µm tall, bearing terminal and lateral monophialides, often reduced to monophialides forming laterally on aerial mycelium or to short phialidic pegs. Aerial conidiogenous cells monophialidic, subcylindrical, 10–17(–27) × 3.5–5 µm, phialidic pegs subcylindrical to conical, 2–4.5 × 2–3 µm, often with a vasiform apical collarette, giving rise to falcate conidia indistinguishable from sporodochial conidia and rarely 0-septate, (5.5–)6.5–11(–12) × 3.5–4.5 µm, obovoidal to clavate microconidia. Chlamydospores not observed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.2–7.6 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface luteous, red at centre, with white to peach patches on the periphery, flat, aerial mycelium abundant, felty to velvety, margin regular, filiform to undulate; reverse pale luteous with abundant coral to bay diffusible pigment. On OA, pure yellow, sulphur yellow to amber, with ochreous peripheric ring and scattered flesh to coral patches, flat, woolly to cottony, margin regular, filiform; reverse amber to pure yellow.
Typus: Costa Rica, from Musa sp., before 26 Apr. 2016, A. Felde & L. Pocasangre (holotype CBS H-25434 designated here, ex-type culture CBS 151937 = BBA 72296 = DSM 116929).
Notes: Resolved as a single lineage basal to the main clades in the Sambucinum clade, F. bananae clusters close but separate to F. musarum, also a Central American species known from Musa spp. Both species have similar conidial sizes and morphology. However, unlike the multi-septate conidia of F. musarum (though not true macroconidia, but mesoconidia), those of F. bananae are produced in sporodochia or from aerial monophialides, lacking aerial polyblastic conidiogenous cells. Moreover, both species can be differentiated by their colony pigmentation on PDA (luteous in F. bananae vs red in F. musarum). Another sister species, F. subcylindroides, also has very similar macroconidial sizes to F. bananae; though, apart from differences in conidial shape (less obviously tapered in F. subcylindroides) both species differ in PDA colony pigmentation (luteous in F. bananae vs pink to red in F. subcylindroides), growth rates (av. 7.4 vs 9.1 mm/d in F. bananae and F. subcylindroides, respectively) and in the absence of microconidia in F. subcylindroides.
Fusarium bellum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855096. Fig. 18.
Fig. 18.
Fusarium bellum (ex-type CBS 151827). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G. Aerial conidiophores and conidiogenous cells. H. Microconidia. I. Chlamydospores. J. Macroconidia. Scale bars: F, G, I, J = 10 μm; H = 5 μm.
Etymology: From Latin bellus (“beautiful, pretty, handsome”). In reference to the beautifully shaped macroconidia.
Sporodochia rare on aerial mycelium, moderately abundant on agar surface, transparent to pale saffron coloured. Sporodochial conidiophores branching irregularly laterally and loosely, 33.5–54 µm tall, bearing lateral and terminal verticils of monophialides. Sporodochial conidiogenous cells monophialidic, subulate to subcylindrical, 10.5–23.5 × 2.5–5 µm, smooth- and thin-walled, with an inconspicuous apical collarette. Sporodochial conidia falcate, gently to strongly dorsiventrally curved and tapering abruptly to both extremes, apical cell commonly moderately elongated, rarely short and blunt; basal cell papillate to well-developed, foot-shaped, (1–)3–5(–6)-septate, (14.5–)35–64(–95.5) × 3–6 µm (av. 49.6 × 4.7 μm), mostly 5-septate, (32–)44.5–59.5(–71.5) × 3.5–5.5 µm (av. 52.1 × 4.9 μm). Aerial conidiophores reduced to short, lateral phialidic pegs on agar substrate and aerial mycelium. Aerial conidiogenous cells monophialidic, subulate, 2–5 × 2.5–4 µm, giving rise to micro- and macroconidia. Microconidia ellipsoidal to short falcate, 0–1-septate, 8.5–9.5 × 2.5–4 µm (av. 8.9 × 3.1 μm), aerial macroconidia indistinguishable from sporodochial conidia. Chlamydospores subglobose to globose, smooth- and thick-walled or warted, (7–)9–13.5(–15) µm diam, hyaline to subhyaline, solitary or grouped in short chains.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 9.1–10.3 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface felty to woolly, rosy vinaceous to red, with white, pale luteous to luteous patches, white at centre, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse scarlet to blood, orange to luteous at centre. On OA, at first pure yellow to amber, becoming rosy vinaceous to scarlet, flat, cottony to woolly, with concentric rings of aerial mycelium, margin regular and filiform; reverse luteous to ochreous.
Typus: China, Anda, from nematode cyst on Glycine max root, before 8 Apr. 1997, unknown collector (holotype designated here CBS H-25409, ex-type culture CBS 151827 = FRC R-9513 = NRRL 66939).
Notes: Previously assigned to Fusarium sp. nov. 28 (Laraba et al. 2021), F. bellum clustered as the closest species to F. subflagellisporum, both species known from soil; however, with quite different biogeography, F. bellum known from Asia (China), while F. subflagellisporum has been reported from the Americas and Africa (Crous et al. 2021, Farr et al. 2023). Fusarium bellum differs from F. subflagellisporum by producing only one type of macroconidium (falcate vs falcate plus whip-like conidia in F. subflagellisporum), and faster growth rates on PDA (av. 9.7 vs 6 mm/d in F. subflagellisporum).
The type of F. bellum was originally preserved and received from NRRL as F. acuminatum (a member of the F. tricintum SC). Although Fusarium bellum differs by its blunt apical cells and slower PDA growth rates, both taxa are difficult to tell apart based on morphological features alone, which highlights the difficulties on reconciling the old Fusarium sections system with the current phylogenetic knowledge.
Fusarium boothii O’Donnell et al., Fungal Genet. Biol. 41: 618. 2004. Fig. 19.
Fig. 19.
Fusarium boothii (ex-type CBS 316.73). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F, G. Sporodochial conidiophores and conidiogenous cells. H–J. Aerial conidiogenous cells. K. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: O’Donnell et al. (2004), Aoki et al. (2012).
Typus: South Africa, from Zea mays, 11 Oct. 1997, P. Martin (holotype BPI 843475, ex-type culture CBS 316.73 = ATCC 24372 = FRC R-9631 = IMI 160243 = NRRL 26916).
Additional materials examined: Nepal, Kaski, from Zea mays ear rot, before 1 May 1994, unknown collector, culture CBS 110251 = FRC R-9434 = NRRL 29105 = NRRL 29208 = NRRL 38122. South Africa, Winterton, from Triticum sp. scab, unknown collection date, W.F.O. Marasas, culture CBS 119798 = MRC 4712 = NRRL 53893; unknown location, from Zea mays, 1990, W.F.O. Marasas, culture CBS 119800 = MRC 6010 = NRRL 53895.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Fusarium boothii and F. meridionale can be differentiated from the remaining segregates of F. graminearum s. lat. by its slender (< 4.5 μm wide), curved and symmetrical 5-septate macroconidia, with narrow apical beaks, wider in the mid-region (Aoki et al. 2012). However, these shared characteristics also make these two species morphologically indistinguishable from one other. Nonetheless, as observed here, F. boothii exhibits slightly faster grown rates on PDA (av. 8.3 vs 6.7 mm/d in F. meridionale), while microconidia, which are rarely present in F. meridionale, were not observed in F. boothii. Additionally, both species differ in their mycotoxin chemotypes, F. boothii producing culmorin, lactones and type B trichothecenes; while F. meridionale produces nivalenol and lactones only (Aoki et al. 2004, Laraba et al. 2021).
Both species have also an overlapping geographic distribution, known from the Americas (Argentina, Brazil, Guatemala, Mexico and the USA; F. boothii additionally from Canada), Africa (South Africa; and F. boothii also from Ethiopia and Zambia), and Asia (China, Korea, and Nepal; and F. meridionale additionally from Iran); while F. meridionale was also reported in Oceania (Australia and New Caledonia) (Aoki et al. 2004, Farr et al. 2023). However, F. boothi appears to have a more restricted host range, being reported mostly from Poaceae (Glycine max, Triticum aestivum, and Zea mays) and more uncommonly Solanaceae (Solanum lycopersicum). In contrast, additional hosts of F. meridionale include species in Araceae, Convolvulaceae, Fabaceae, and Myrtaceae (Farr et al. 2023).
Fusarium brachygibbosum Padwick, Mycol. Pap. 12: 11. 1945. Fig. 20.
Fig. 20.
Fusarium brachygibbosum (ex-type NRRL 20954). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–I. Conidiogenous cells. J. Chlamydospore. K. Microconidia. L. Macroconidia. Scale bars: E–G, J–L = 10 μm; H, I = 5 μm.
Descriptions and illustrations: Padwick (1945), Nirenberg (1990), Lombard et al. (2022).
Typus: India, Telangana, Hyderabad, Parbhani, on Sorghum vulgare, Sep. 1940, S. Vaheeduddin (holotype IMI 268019, ex-type culture BBA 64691 = NRRL 20954 = DSM 116019).
Additional materials examined: India, Hampi Temples, Achutaraya temple 1, from stone, unknown collection date and collector, dep. J. Bosselaers, culture CBS 121682. Iran, Ardabil Province, Bilesovar, Djafar Abad, from Agropyron sp., unknown collection date and collector, dep. M. Davari, culture CBS 131017; Gholestan Province, Azadshahr, from Triticum sp., unknown collection date and collector, dep. M. Davari, culture CBS 131252. Israel, Jaffa, from sea water, unknown collection date, O. Yarden, cultures CPC 45794 = W15, CPC 45801 = W53. Japan, from river sediment, before 13 Mar. 1989, unknown collector, dep. P.E. Nelson, culture FRC R-6784 = MRC 2568 = NRRL 13829. Russia, Yalta, from soil on old Vitis vinifera plantation, 2021, I.N. Pavlov, culture CPC 41845 = F43. Sudan, from Phaseolus vulgare dried seed, unknown collection date, M. Eltayeb, culture CBS 466.92 = NRRL 25806; Kassla State, Kassala, Sawagi South, from Solanum lycopersicon, unknown collection date, Elhassan, Raaed & M. Mohieldin, culture BBA 72481 = DSM 116936; Shambat Research Station, from soil, unknown collection date, M.A.M.K. Abdalla, culture BBA 71480 = DSM 116711. Türkiye, from Musa sapientum fruit, before Apr. 1998, unknown collector, dep. Regional Plant Protection Research Institute, Adana, culture CBS 334.75 = FRC R-9859 = NRRL 28448.
Notes: Booth (1971) synonymized F. brachygibbosum under the misapplied name F. semitectum (currently Fusarium incarnatum, in the Fusarium incarnatum-equiseti SC) (Gerlach & Nirenberg 1982, Nirenberg 1990, Xia et al. 2019). Gerlach & Nirenberg (1982) rejected this synonymy, recognizing F. brachygibbosum in section Gibbosum, but acknowledging Padwick (1945) comments on notable differences to other members of that section (i.e., 5-septate macroconidia too short and wide, with less developed foot cells; and chlamydospores both terminal and intercalary). Significant cultural and morphological differences exist between F. brachygibbosum and F. incarnatum including PDA colony pigmentation (red vs yellow to beige/brown in F. incarnatum), and most notably conidial morphology, with F. brachygibbosum macroconidia markedly dorsiventrally curved and conspicuously wider in the mid portion. In addition, F. incarnatum has both phialidic and holoblastic conidiogenesis (mesoconidia) on aerial conidiophores (Pascoe 1990), which has not been described in F. brachygibbosum. Nevertheless, one isolate of F. brachygibbosum examined here (NRRL 13829), although it lacked sporodochia, produced barely curved, spindle shaped, multiseptate aerial conidia with truncate basal cells, which conform to Pascoe’s definition of mesoconidia (Pascoe 1990); although, mesoconidiophores and holoblastic conidiogenous cells were not observed.
Closely related species to F. brachygibbosum include F. cuspidatum, F. minutum and F. sagittatum, all of them showing distinctive morphological characters. Fusarium brachygibbosum is distinguished from F. minutum and F. sagitattum by its typical hyperbolic curvature (vs a more regular curvature in F. minutum, and almost straight conidia in F. sagitattum); and differs from F. cuspidatum by having smaller macroconidia (av. 34.3 × 4.9 vs 42 × 5.5 µm in F. cuspidatum), while microconidia are absent in the latter species. Microconidia, originally not observed in the protologue of F. brachygibbosum (Padwick 1945), were observed here in several strains, including the ex-type.
Fusarium brachygibbosum has a wide distribution, being known from Africa, Asia, Europe, North America, and Oceania (EFSA PLH Panel et al. 2021, Farr et al. 2023). It is a saprobe, but can also act as a plant pathogen, associated with canker; crown, ear, root, stalk, and tuber rot; decline, dieback, leaf spot, and wilting on a variety of hosts. It was recently evaluated as a potential quarantine pest in the European Union; although it does satisfy all the criteria to be regarded as such a pest, the high uncertainty regarding its distribution in the Union and its potential impact in the Union has not resulted in its inclusion in the EU Commision Implementing Regulation 2019/2072 at this time (EFSA PLH Panel et al. 2021). It has been reported from sediments, soil, humans, insects, water, and 25 plant host genera from 16 families, including Allium cepa, Agropyron sp., Arachis hypogaea, Beta vulgaris, Cannabis sativa, Citrullus lanatus, Citrus limettioides, Euphorbia larica, Glycine max, Gossypium hirsutum, Helianthus annuus, Musa sapientum, Nerium oleander, Nicotiana tabacum, Phaseolus vulgare, Phoenix dactylifera, Prunus dulcis, Sansevieria trifasciata, Solanum tuberosum, Sorghum spp., Tectona grandis, Trifolium subterraneum, Triticum spp., Vitis vinifera. and Zea mays (Al-Hatmi 2016, EFSA PLH Panel et al. 2021, Farr et al. 2023, O’Donnell et al. 2012, Dhlamini et al. 2024).
Fusarium brachypes Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855097. Fig. 21.
Fig. 21.
Fusarium brachypes (ex-type CBS 151824). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Cluster of macroconidia on agar surface. F–I. Aerial conidiophores and conidiogenous cells. J. Microconidia. K. Macroconidia. Scale bars = 10 μm.
Etymology: From Greek brachy (“short”) and pes (“foot”). Referring to the poorly developed macroconidial foot cells.
Sporodochia not seen, but radiate groups of macroconidia commonly observed on agar surface. Aerial conidiophores scantly produced, erect or prostrate on substrate mycelium, simple to sparsely branched, 14.5–59 µm tall, often reduced to single monophialides or phialidic pegs forming laterally on aerial mycelium or submerged in agar. Aerial conidiogenous cells monophialidic, subulate to subcylindrical, smooth- and thin-walled, 7.5–23.8 × 2.5–5.5 µm, with inconspicuous apical collarette, or reduced to lateral phialidic pegs, 2–2.5 × 3–5 µm, giving rise to micro- and macroconidia. Microconidia ellipsoidal to allantoid, 0(–3)-septate, (6–)8–17.5(–37.5) × (2.5–)3–5 µm (av. 12.9 × 3.5 μm); macroconidia falcate, moderately dorsiventrally curved, with a curved or almost straight ventral line, apical cell blunt, basal cell papillate or poorly- to well developed-foot-shaped, (1–)2–3-septate, (22–)26.5–35.5(–40) × 3.5–5 µm (av. 31.2 × 4.3 μm), mostly 3-septate, 28–36.5(–40) × 3.5–5 µm (av. 32.4 × 4.3 μm). Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.0–9.1 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface at first membranous, becoming cottony to woolly at periphery, sulphur yellow, rosy buff to red at periphery, white at centre, flat with radial patches, aerial mycelium abundant at periphery, margin regular and filiform; reverse sulphur yellow to amber, flesh to red at periphery. On OA, at first amber to ochreous, later developing white to sulphur yellow mycelium, flat, at first membranous, becoming cottony to woolly, moist at centre, with regular and filiform margin; reverse honey to sienna.
Typus: Australia, Northern Territory, from cultivated soil, before 1 Nov. 1988, unknown collector (holotype designated here CBS H-25408, ex-type culture CBS 151824 = FRC R-8938 = NRRL 66933).
Notes: Previously assigned to Fusarium sp. nov. 33 (Laraba et al. 2021), F. brachypes is so far known only from soil in Australia. Macroconidia were only produced on aerial mycelium, while sporodochia were not produced. It is not clear is this is a result of strain degeneration or a characteristic of the species. Conidial size of F. brachypes is in the range of the phylogenetically distant species F. culmorum, F. cultriforme, and F. sporotrichioides. Fusarium brachypes differs from F. culmorum by its slender conidia (≤ 5 µm vs up to 6.5 µm wide in F. culmorum), differs from F. cultriforme by its macroconidial shape, with less pronounced dorsiventral curvature, often straight ventral line, and blunt apical cell; and faster PDA growth rates (av. 8.6 vs 4.3 mm/d in F. cultriforme), and differs from F. sporotrichioides by lacking pyriform microconidia and mesoconidia; and by its almost straight and not as markedly tapering macroconidia. Additionally, F. brachypes produces yellow pigmentation on PDA, which further distinguishes this species from the red PDA colonies of F. culmorum and F. sporotrichioides.
Fusarium brachypes can be recognized from its closest phylogenetic siblings i.e., F. dimorphosporum and F. mucronatum by its smaller macroconidia (av. 31.2 × 4.3 vs 47.7 × 6 and 45.3 × 6.2 µm, respectively), which lacks elongated apical cells.
Fusarium brasilicum T. Aoki et al., Fungal Genet. Biol. 41: 620. 2004.
Descriptions and illustrations: O’Donnell et al. (2004), Aoki et al. (2012).
Typus: Brazil, on Avena sativa, 2000, J. Martinelli (holotype BPI 843480, ex-type culture CBS 119180 = FRC R-10019 = NRRL 31281).
Additional material examined: Brazil, Rio Grande do Sul, Pelotas, from Hordeum vulgare, before Jun. 2001, J. Fernandes, culture CBS 119179 = FRC R-10018 = NRRL 31238.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum.
Cultures preserved at CBS, including the ex-type strain, are degenerated and no longer sporulate, hence the species was not studied morphologically. Fusarium brasilicum was resolved as a genetically exclusive clade in the combined phylogeny and single locus analyses of rpb1, rpb2 and tub2; as well as in the STACEY analyses. This species cannot be morphologically distinguished from F. cortaderiae, its closest phylogenetic relative (O’Donnell et al. 2004, Aoki et al. 2012, Laraba et al. 2021); both species having asymmetric, straight to curved, 4.5–5 µm wide 5-septate conidia, with narrow apical beaks and wider below the median (Aoki et al. 2012).
Fusarium brasilicum is known from Poaceae (i.e., Avena sativa, Hordeum vulgare, and Triticum sp.), reported in Brazil and South Africa (Farr et al. 2023).
Fusarium carinatum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855098. Fig. 22.
Fig. 22.
Fusarium carinatum (ex-type CBS 151957). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G. Sporodochial conidiophores and conidiogenous cells. H, I. Aerial conidiophores, conidiogenous cells and microconidia. J. Aerial macroconidia. K. Chlamydospores. L. Sporodochial macroconidia. Scale bars = 10 μm.
Etymology: From Latin carīnātus (“shaped like a keel”). In reference to the macroconidial dorsal curvature, resembling a ship’s full keel.
Sporodochia scant on the surface of carnation leaves, abundant on agar surface and aerial mycelium, transparent white, sulphur yellow to straw coloured. Sporodochial conidiophores sparingly branching laterally, compact, 21–31 µm tall, bearing terminal monophialides. Sporodochial conidiogenous cells monophialidic, doliiform to subcylindrical, 7.5–15 × 3.5–5.5 µm, smooth- and thin-walled, with a short, flared apical collarette. Sporodochial conidia falcate, gently dorsiventrally curved to almost straight with an almost straight ventral line, often widest at or above the mid-region, apical cell tapering and pointy; basal cell papillate to well-developed, foot-shaped, (3–)5–8-septate, (37.5–)49–57(–60.5) × (4.5–)5.5–6.5 µm (av. 52.9 × 5.8 μm), mostly 6-septate, (47–)51.5–57.5(–59.5) × 5–6.5 µm (av. 54.7 × 5.8 μm). Aerial conidiophores prostrate, sparsely branched, 21–24 µm tall, commonly reduced to monophialides forming laterally on aerial mycelium or submerged in agar and rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical, 5–21.5 × 2.5–6 µm, smooth- and thin-walled, giving rise to micro- and macroconidia. Microconidia rare, clavate to falcate, 0–1-septate (11.5–)14–21(–24.5) × 3–5 µm (av. 17.2 × 3.8 μm), smooth- and thin-walled; aerial macroconidia, fusiform, slender, straight to barely dorsiventrally curved, smooth- and thin-walled, 1–5-septate, (19–)22–38(–48) × (3.5–)4–5(–5.5) µm (av. 30.2 × 4.3 μm), mostly 3-septate, (22–)25–36.5(–39.5) × 3.5–5.5 µm (av. 30.8 × 4.3 μm). Chlamydospores globose, smooth- and thick-walled, hyaline, 3.5–5.5 µm diam, formed in clusters, intercalary on hyphae and macroconidia, or laterally in chains on short stipes.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.7–8 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface velvety to dusty, primrose to sulphur yellow, flat with abundant radial patches, aerial mycelium scant, margin regular and filiform; reverse sulphur yellow to pale luteous. On OA, sulphur yellow to pale luteous, flat, membranous to slimy, with regular and undulate margin; reverse pale luteous to amber.
Typus: Nigeria, Niger State, from Sorghum sp. soil debris, before 11 May 1988, unknown collector (holotype designated here CBS H-25419, ex-type culture CBS 151957 = FRC R-9061 = NRRL 66934).
Notes: Originally assigned to Fusarium sp. nov. 22 in Laraba et al. (2021), the production of yellow PDA colonies quicky distinguishes F. carinatum from most species in the Longipes clade, except for F. pratense and F. vermicularioides. Both F. carinatum and F. pratense differ from all other species in the Longipes clade by its falcate conidia with pointy, but not elongated apical cells; while all other currently described species in the Longipes clade exhibit whip-like conidia, although, with different degrees of apical cell elongation. Fusarium carinatum differs from F. pratense by its larger (av. 52.9 × 5.8 vs 44.9 × 4.1 µm in F. pratense) and often more frequently septate conidia (up to 8, commonly 6 septa, vs rarely up to 6, commonly 5-septate in F. pratense) sporodochial conidia; and by forming a secondary type of falcate macroconidia on its aerial mycelium. In addition, both species differ in their biogeography, F. carinatum known so far from Nigeria (on Sorghum spp., soil debris), and F. pratense from Oceania (from soil and Poaceae).
Fusarium cerealis (Cooke) Sacc., Syll. Fung. 4: 713. 1886. Figs 23, 24.
Fig. 23.
Fusarium cerealis (neotype K(M)133541). A. Specimen label. B. Specimen. C, D. Macroconidia from specimen as depicted in Nirenberg (1990). E. Microphotograph of macroconidia. F. Microphotograph of sporodochial conidiophore and conidiogenous cells. G. Macroconidia. Scale bar = 10 μm.
Fig. 24.
Fusarium cerealis (CBS 832.85). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiophores. J. Macroconidia. Scale bars = 10 μm.
Basionym: Fusisporium cerealis Cooke, Grevillea 6: 139. 1878.
Synonyms: Fusarium culmorum var. cerealis (Cooke) Wollenw., Fusaria Autogr. Delin. 3: 946. 1930. (Pro parte).
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.
Descriptions and illustrations: Nirenberg (1990), Leslie & Summerell (2006).
Typus: USA, California, on stem of Zea mays, unknown collection date, HW. Harkness (Harkness 1493) (neotype of Fusisporium cerealis K(M)133541, designated in Nirenberg 1990).
Additional materials examined: France, Montours, Fougéres, from grass 1993, C. Snijders, culture CBS 100101 = NRRL 36070. Germany, Stuttgart, from Triticum durum seed, 1985, H. Nirenberg, culture CBS 832.85 = BBA 64545 = NRRL 25794. Netherlands, from Beta vulgaris, J.W. Veenbaas-Rijks, culture CBS 135.80 = IMI 322101 = NRRL 36243, Haren, from Solanum tuberosum tuber buried in soil, unknown collection date and collector, isol. H. Reinartz, H.R. 11, culture CBS 623.85 = NRRL 36523. Poland, Posnan, from Solanum tuberosum, unknown date, P. Golinski, culture CBS 110268 = ATCC 66007 = FRC R-09961 = NRRL 13721. Unknown, unknown collection data, culture CBS 119874 = FRC R-09984 = MRC 8399 = KSU 11453 = Univ. Sydney F13169.
Notes: The holotype citation in Nirenberg (1990), and Crous et al. (2021) is incorrect. The material hosted at Kew [K(M)133541 = “Harkness 1493”], is labelled “holotype” and “type” in several instances; however, is not the holotype of Fusisporium cerealis. It consists of plant material, apparently stems of Zea mays, two slide preparations, and two microphotographs (Fig. 23). Discordant collection data between this specimen and the protologue of the species support an annotation by K.A. Seifert (Dec. 2006) indicating that this specimen is not the holotype of F. cerealis, as indicated in Seifert et al. (2009). However, the stamp on the specimen indicates this specimen is authentic for the name, and former part of M.C. Cooke’s fungarium (K.A. Seifert, pers. comm.). Nirenberg (1990) cited “type, No. 1493. Harkness, K” as the holotype of F. cerealis. In the absence of original material for the species, as confirmed in Seifert et al. (2009), the above type citation constitutes an effective, though unintended, neotypification. Although referred to as holotype in Nirenberg (1990), this is correctable according to Art. 9.10.
Previously reduced to a variety of F. culmorum by Wollenweber (1930). Several authors retained F. cerealis as either a variety or a synonym of F. culmorum (Gordon 1952, Booth 1971, Gerlach & Nirenberg 1982, Nelson et al. 1983), or a variety of F. sambucinum (Raillo 1950). The modern concepts of F. cerealis and F. culmorum rejects the previous synonymies (Nirenberg 1990, Leslie & Summerell 2006, Domsch et al. 2007) which is supported by morphological and phylogenetic evidence. An investigation of documents accompanying strain deposits at CBS, revealed that at least one authentic strain deposited by Wollenweber as F. culmorum var. cerealis was indeed F. culmorum (CBS 173.31, received as strain 2544 and illustrated in Fus. Aut. Del. 946; Wollenweber 1930), which indicates that the original synonymy of F. cerealis and F. culmorum was based on observations of mixed isolates from both species. The two taxa are phylogenetic and morphological siblings, although they can be distinguished morphologically; F. cerealis being characterized by slender and longer macroconidia with beaked apical cells, versus the robust, shorter and broader macroconidia with almost pointy apical cells in F. culmorum.
Fusarium cerealis known substrates include soil, and several plant host species, mostly in Poaceae (Anthoxanthum aristatum, Dactylis glomerata, Hordeum vulgare, Paspalum sp., Secale cereale, Triticum spp., and Zea mays), but also Amaranthaceae (Beta vulgaris), Araliaceae (Panax ginseng), Caryophyllaceae (Dianthus sp.), Ericaceae (Rhododendron sp.), Fabaceae (Glycine max, Medicago sativa, and Trifolium pratense), Iridaceae (Iris hollandica), Lauraceae (Persea americana), Nymphalidae (Cirsium sp.), Oleaceae (Syringa sp.), Pinaceae (Pinus radiata), Rosaceae (Prunus persica), and Solanaceae (Solanum tuberosum); and it has been recorded from Asia (China, Israel), Europe (Austria, Finland, France, Germany, Italy, Poland, Russia, Spain, Switzerland, and Yugoslavia), North and South America (Argentina, Canada, Colombia, and the USA), and Oceania (Australia) (Farr et al. 2023).
Fusarium chaquense M.J. Nichea et al., Mycologia 114: 54. 2021. Fig. 25.
Fig. 25.
Fusarium chaquense (NRRL 64072). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiophores, conidiogenous cells and microconidia. J. Macroconidia. Scale bars = 10 μm.
Description and illustration: Nichea et al. (2021).
Typus: Argentina, Chaco Province, Wetlands, Ramsar site no. 1366, from Sorghastrum setosum, Jul. 2011, M.L. Ramirez (holotype RCVC 9613, ex-type culture NRRL 66749 = RC-J293).
Additional materials examined: Argentina, Chaco Province, from Leersia luziola, Jul. 2011, M.L. Ramirez, culture NRRL 66748 = RC-J174, from Diplachne sp., Feb 2014, M.L. Ramirez, culture NRRL 66750 = RC-J1301. Australia, from soil, before 1 Oct. 1976, unknown collector, dep. L.W. Burgess, culture NRRL 64041 = FRC R-3766 = KOD 1980. South Africa, Free State, Frankfort, from Avena sativa, before 1 Oct. 1985, unknown collector, culture CBS 151952 = CBS H-25404 = NRRL 64072 = FRC R-8102 = KOD2011 = MRC 3936.
Notes: This recently described species is presently only known from Poaceae (on species of the genera Avena, Chloris, Cynodon, Dichantium, Diplachne, Elionurus, Eragrostis, Eriochloa, Hymenachme, Leersia, Panicum, Paspalidium, Paspalum, and Sorghastrum) and soil in Argentina (Nichea et al. 2021), Australia and South Africa (this paper). Fusarium chaquense (previously Fusarium sp. nov. 1 in Laraba et al. 2021) is the closest morphological and phylogenetic sibling of F. armeniacum, from which it can be distinguished by its conidial shape and biogeography; the separation of both taxa according to the presence of microconidia does not stand according to the observation of the ex-type of the latter species (see notes under F. armeniacum). The description and measurements of microconidia in the protologue of F. chaquense includes a mixture of elements that are here separated into microconidia and aerial macroconidia (Table S1).
Fusarium cortaderiae O’Donnell et al., Fungal Genet. Biol. 41: 620. 2004. Fig. 26.
Fig. 26.
Fusarium cortaderiae (ex-epitype CBS 119183). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiogenous cells. G, H. Aerial conidiophores and conidiogenous cells. I. Macroconidia. Scale bars = 10 μm.
Description and illustration: O’Donnell et al. (2004), Aoki et al. (2012).
Typus: New Zealand, Auckland, Henderson, from Cortaderia selloana, Aug. 1965, J. Dingley (holotype BPI 843479, ex-type culture CBS 119183 = FRC R-10016 = ICMP 5435 = NRRL 29297).
Additional materials examined: Brazil, Triticum sp., variety-CD105, before Jun. 2001, J. Martinelli, culture CBS 123659 = NRRL 31205. New Zealand, Palmerston North, from Zea mays, before Dec. 1999, H. Hussein, culture CBS 123655 = FRC R-9682 = ICMP 8998 = NRRL 29306.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Although resolved by phylogenetic and STACEY analyses, F. cortaderiae is morphologically indistinguishable from its closest phylogenetic sibling F. brasilicum, both species forming asymmetric, straight to curved, 4.5–5 µm wide, 5-septate conidia, with narrow apical beaks, and wider below the median (Aoki et al. 2012). Moreover, both species overlap in their host preferences and partially in their biogeographic patterns; however, F. cortaderiae has a wider geographical distribution, with additional records in Europe (Italy), Oceania (Australia and New Zealand) and South America (Argentina, Brazil, and Uruguay), apart from Africa (South Africa) (Farr et al. 2023). In addition, unlike F. brasilicum, known only from a restricted number of Poaceae hosts (Avena sativa, Hordeum vulgare, and Triticum sp.), F. cortaderiae is known from Asteraceae (Gerbera sp.), Caryophyllaceae (Dianthus sp.), Fabaceae (Glycine max), Poaceae (Cortaderia selloana, Hordeum vulgare, Lolium multiflorum, Oryza sativa, Triticum aestivum, Triticum durum, and Zea mays), and soil (Farr et al. 2023).
Fusarium culmorum (Wm.G. Sm.) Sacc., Syll. Fung. 10: 726. 1892. Figs 27, 28.
Fig. 27.
Fusarium culmorum (lectotype of Fusisporium culmorum). Reproduction from the original publication by Smith (1884). A–C. Conidiogenous cells and macroconidia. D. Host cells (wheat).
Fig. 28.
Fusarium culmorum (ex-epitype CBS 417.86). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiogenous cells. I, J. Aerial conidiogenous cells. K. Macroconidia undergoing microcyclic conidiogenesis. L. Macroconidia. Scale bars: G–J = 5 μm; K, L = 10 μm.
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].
Fusoma tenue Grove, J. Bot. 50: 16. 1912.
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.
Fusarium culmorum var. cerealis (Cooke) Wollenw., Fusaria Autogr. Delin. 3: 946. 1930. (pro parte).
Fusarium culmorum f. 1 Wollenw., Z. Parasitenk. 3: 361. 1931.
Descriptions and illustrations: Wollenweber & Reinking (1935), CABI (1964), Booth (1971), Gerlach & Nirenberg (1982), Leslie & Summerell (2006).
Typus: UK, on infected ear of Triticum sp., 1884, W.G. Smith (lectotype fig. 92, page 210 in Diseases of field and garden crops, chiefly such as are caused by fungi, designated by Crous et al. 2021). Denmark, mouldy kernel of Hordeum vulgare, 3 Feb. 1986, U. Thrane [epitype CBS 417.86 (preserved as metabolic inactive culture) designated by Crous et al. 2021], ex-epitype culture CBS 417.86 = FRC R-8504 = IMI 309344 = NRRL 25475).
Additional materials examined: Belgium, Sorinnes, from Triticum aestivum seed, 2004, A. Chandelier, culture CBS 128537. Canada, unknown host, H.W. Wollenweber, culture CBS 171.28 = IMI 089365 = IMI 089366 = IMI 092033 = NRRL 26854 = NRRL 26915; Saskatchewan, Saskatoon, from Avena sativa hypocotyl, 1927, P.M. Simmonds, culture CBS 173.31 = MUCL 783 = NRRL 26853 = NRRL 26914 = NRRL 29139. Denmark, from Miscanthus sp. chopped stems, buried in soil, 1996, M. Klamer No S-1296-56-1, culture CBS 579.97 = NRRL 26241 = NRRL 26602; Sjaelland, Flakkebjerg, from soil from barley field, Aug. 1985, S. Elmholt No. SEF 15, culture CBS 416.86 = FRC R-8505 = IMI 309345 = NRRL 36469. France, from Triticum durum, 1998, B. Bènèdicte, culture CBS 110565 = NRRL 36195; from Triticum sp., 1998, B. Bènèdicte, culture CBS 110567 = NRRL 36197. Hungary, from millet, before 24 Apr. 1978, unknown collector, culture CBS 110262 = ATCC 34912 = NRRL 6394. Netherlands, Terschelling, from Ammophila arenaria root, H.F. Kauffman, culture CBS 472.95 = NRRL 29144; Lisse, from Hyacinthus orientalis root, 1 Jun. 1943, A. Slootweg No. 1056, culture CBS 257.51 = NRRL 29142. Poland, Powisle region, from Triticum aestivum, 2004, unknown collector, dep. T. Kulik, culture CBS 115698; Lublin region, rhizosphere soil of Secale cereale, unknown collection date, Department of Environmental Microbiology, Fungal Collection (DEM), culture CBS 120099 = NRRL 53962. Unknown, unknown collection data, culture CBS 176.32 = NRRL 29140, V. Edel, culture CBS 597.96, CBS 119869 = MRC 8403 = NRRL 53938; from Triticum aestivum grain “Koga”, Jul. 1952, G. Dinger, culture CBS 251.52 = NRRL 29292; from Secale cereale seed, 1952, G. Dinger, culture CBS 250.52 = MUCL 555 = NRRL 29291.
Notes: Crous et al. (2021) recently fixed the typification of F. culmorum, the lectotype, an original drawing from its protologue is reproduced here (Fig. 27), and additional barcodes were sequenced for the ex-epitype (CBS 417.86).
Fusarium culmorum can be easily recognized by its robust, falcate macroconidia, which are quite broad for its length. These characteristics, plus the absence of an apical beak differentiates F. culmorum from its closest morphological and phylogenetic sibling F. cerealis. For additional comments about the taxonomy of F. culmorum and F. cerealis, see notes under F. cerealis. Fusarium cultriforme, a morphologically similar species can be differentiated by the presence of microconidia, the formation of yellow PDA colonies, and less frequently septate, slender macroconidia on the latter species.
Fusarium culmorum is a cosmopolitan species present in dry, temperate, and tropical climates which has been recorded from over 50 countries in Africa, Asia, Europe, Oceania; and Central, North and South America (Farr et al. 2023). This is a soil fungus, and apart from aquatic environments, insects, and salt marshes (Domsch et al. 2007, O’Donnell et al. 2012), it can also be found either as endophyte or pathogen in more than 140 plant host species, from 85 genera, mostly in Poaceae, followed by Fabaceae, and 25 additional families (Farr et al. 2023). It is an agent of crown, foot, and root rots; storage rots, and head and seedling blight (CABI 1964, Leslie & Summerell 2006, Domsch et al. 2007).
Fusarium cultriforme Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855099. Fig. 29.
Fig. 29.
Fusarium cultriforme (ex-type CBS 151935). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Aerial conidiophores and conidiogenous cells. I. Sporodochial conidiophore and conidiogenous cells. J. Microconidia. K. Macroconidia. Scale bars: G–I, K = 10 μm; J = 5 μm.
Etymology: From Latin culter, cultri (“knife”) and -form. In reference to the shape of the macroconidia, resembling a pruning knife.
Sporodochia abundant on the surface of carnation leaves and later on aerial mycelium, pale luteous, luteous to orange coloured. Sporodochial conidiophores branching multiple times laterally and verticillately, 40–55 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical, 11–21 × 2.5–4.5 µm, smooth, thin-walled, with a minute apical collarette. Sporodochial conidia falcate, dorsiventrally curved, with blunt to hooked apical cell; basal cell papillate or with a well-developed foot-cell, (1–)2–5-septate, (23.5–)28–33(–42) × 4–5 µm (av. 30.2 × 4.9 μm), mostly 3-septate, (24.5–)27–32(–35.5) × 4–5.5 µm (av. 29.8 × 4.8 μm). Aerial conidiophores erect, laterally branched multiple times, 34–220 µm tall, forming laterally from substrate mycelium, proliferating percurrently. Aerial conidiogenous cells mono- and polyphialidic, subcylindrical, 9–27 × 2.5–4.5 µm, giving rise to falcate conidia indistinguishable from sporodochial conidia, and rarely to 0–1(–2)-septate, (9.5–)10–17.5 × 3–5 µm ellipsoidal microconidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 4–4.5 mm/d, 27–31 mm diam after 7 d. Surface primrose to luteous with abundant straw concentric rings, felty, flat to slightly raised, margin irregular, filiform to undulate; reverse pale luteous. On OA, saffron to pale ochreous, flat, cottony to woolly, margin regular, entire to undulate; reverse pale luteous to pale saffron.
Typus: Venezuela, from Impatiens sp., before 25 Apr. 2016, unknown collector (holotype designated here CBS H-25407, ex-type culture CBS 151935 = BBA 65688 = DSM 116528).
Additional material examined: Unknown, unknown collection data, culture NRRL 36134.
Notes: A member of the Sambucinum clade of FSAMSC, and previously assigned to Fusarium sp. nov. 14 in Laraba et al. (2021), macroconidia of F. cultriforme, resemble those of F. culmorum, although, differing by being commonly less septate and slender in F. cultriforme. Additionally, both species can be differentiated by the presence (however rare) of microconida in F. cultriforme and its yellow, slow growing PDA colonies (Table S1). Closest phylogenetic relatives include F. kyushuense and F. venenatum, both species characterized by red PDA colonies and larger and more frequently septate macroconidia (av. 30.2 × 4.9 µm and up to 5-septate in F. cultriforme, vs 40.5 × 4.1 µm and 7-septate, and 44.5 × 5.4 µm and 9-septate in F. kyushuense and F. venenatum, respectively). Additionally, F. cultriforme differs from F. venenatum by forming microconidia, its slower growth rates on PDA (av. 4.3 mm/d vs 7.6 mm/d in the latter species), and lacking terminal chains of chlamydospores; and from F. kyushuense by lacking clavate, multiseptate aerial conidia (Aoki & O’Donnell 1998).
Fusarium cuspidatum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855100. Fig. 30.
Fig. 30.
Fusarium cuspidatum (ex-type CBS 151953). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin cuspidatus (“having a cusp” or “tapering to a sharp point”). In reference to the pointy apex of the macroconidia.
Sporodochia erumpent, abundant on the surface of carnation leaves and aerial mycelium, rare on agar surface, transparent white, orange to sienna coloured. Sporodochial conidiophores densely packed, branching laterally and verticillately, 25.5–44 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, ampulliform to subcylindrical, 8–17.5 × 2.5–5 µm, smooth, thin-walled, with an inconspicuous to absent apical collarette. Sporodochial conidia falcate, dorsiventrally curved, with an almost straight ventral line, wider to somewhat swollen on or above the median and tapering abruptly towards both ends, apical cell conical to pointy; basal cell well-developed, foot-shaped, (1–)3–6-septate, (32.5–)38.5–45.5(–52) × (4.5–)5–6.5 µm (av. 42 × 5.5 μm), mostly 5-septate, (32.5–)38.5–46(–51.5) × 4.5–6.5 µm (av. 42.2 × 5.5 μm). Aerial conidiophores erect on substrate and aerial mycelium or below the agar surface, simple, reduced to solitary lateral phialides or phialidic pegs. Aerial conidiogenous cells monophialidic, subcylindrical or short subulate, 3–10 × 2–3 µm, mostly reduced to phialidic pegs with an inconspicuous apical collarette, giving rise to falcate conidia undistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.7–9.7 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface at first membranous, becoming velvety with woolly patches, red to scarlet, with white, pale luteous to pure yellow mycelial ropes, flat, aerial mycelium scant, margin regular and filiform; reverse peach to scarlet, amber at centre. On OA, amber to pure yellow, with scarlet to coral radial stripes at periphery and small patches and ropes of white, dense mycelium, flat, membranous, moisty at the centre, margin regular and filiform; reverse pale luteous to honey.
Typus: USA, North Dakota, from pasture soil debris, before 8 Oct. 2019, unknown collector (holotype designated here CBS H-25412, ex-type culture CBS 151953 = FRC R-6269 = NRRL 66923).
Notes: A close phylogenetic relative of F. brachygibbosum, F. minutum and F. sagittatum in the Brachygibbosum clade of FSAMSC; F. cuspidatum (previously Fusarium sp. nov. 26 in Laraba et al. 2021) is distinguished from F. minutum and F. sagittatum by its gibbosum-like conidia (macroconidia changing abruptly in diameter, especially at the median, with elongated apical cells; Wollenweber 1918); and differs from F. brachygibbosum by lacking microconidia and having larger macroconidia (av. 42 × 5.5 µm vs 34.3 × 4.9 µm in F. brachygibbosum) which present a more regular curvature on its dorsal face, and tapers abruptly towards both ends.
Other species with similar macroconidial morphology include F. dimorphosporum, F. hamatum and F. platysporum in the Brachygibbosum clade; and F. pratense, and F. tropicale in the Longipes clade. Fusarium cuspidatum differs from F. dimorphosporum, F. platysporum, and F. pratense by the absence of microconidia; while its red PDA colonies further differentiate F. cuspidatum from the yellow PDA colonies of F. pratense. Macroconidia of F. cuspidatum (av. 42 × 5.5 µm) are also smaller than those of F. dimorphosporum (av. 47.7 × 6 µm), F. hamatum (47.4 × 6.2 µm), and F. platysporum (48 × 6 µm); while contrasting with all three latter species in addition to F. tropicale, F. cuspidatum lacks chlamydospores.
Fusarium cygneum Sand.-Den., J.Z. Groenew. & Crous, sp. nov., MycoBank MB 855101. Fig. 31.
Fig. 31.
Fusarium cygneum (ex-type CBS 151805). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin cygnus (“swan”). In reference to the curved shape adopted by the elongated macroconidial apical cells, resembling a swan’s neck.
Sporodochia abundant on the surface of carnation leaves and agar surface, pale yellow, luteous to orange coloured. Sporodochial conidiophores branching laterally and verticillately, 16.5–40 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical to subulate, 10.5–20 × 2.5–5 µm, smooth- and thin-walled, with a conspicuous apical collarette. Sporodochial conidia whip-like, robust, strongly dorsiventrally curved and recurved, apical cell strongly elongated with rounded end; basal cell well-developed foot-shaped and elongated, 5–9(–11)-septate, (79.5–)111–146(–157) × 3.5–5.5 µm (av. 128.7 × 4.4 μm), mostly 6-septate, (81.5–)110–137(–141.5) × 3.5–5 µm (av. 123.5 × 4.4 μm). Aerial conidiophores erect or prostrate on substrate mycelium, rarely branched, 26–54 µm tall, commonly reduced to monophialides forming laterally on aerial mycelium or submerged in agar, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, ampulliform to subcylindrical, 8.5–20 × 3–5.5 µm, giving rise to whip-like conidia undistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 3.4–5.4 mm/d, 23–37 mm diam after 7 d. Surface scarlet to intense red, velvety, with scarlet radial patches, umber to blood coloured at centre, flat, aerial mycelium scarce and short, margin regular, filiform to undulate; reverse saffron to scarlet, umber at centre. On OA, amber to pure yellow, flat, membranous to slimy at centre, velvety at periphery, margin regular, filiform to undulate; reverse amber to luteous.
Typus: UK, Wiltshire, from unknown host, 24 Feb. 1982, unknown collector (holotype designated here CBS H-25421, ex-type culture CBS 151805= IMI 265540 = NRRL 20723).
Notes: Previously assigned to lineage F. longipes 3 (O’Donnell et al. 2013, Laraba et al. 2021), F. cygneum is one of six species in the FSAMSC (in addition to F. dolichosporum, F. longicolle, F. magnum, F. mastigosporum, and F. procumbens) characterized by large (average length close or above 100 μm), whip-like macroconidia with remarkably elongate apical cells, resembling, although to a greater degree, members of the Fusarium incarnatum-equiseti SC (Xia et al. 2019), F. aywerte (in the F. aywerte SC, Sangalang et al. 1995), or F. nurragi (a lone lineage basal to the F. heterosporum SC, O’Donnell et al. 2013). This macroconidial type is present, but not exclusively, in the Longipes and Sambucinum clades of FSAMSC. Fusarium cygneum, together with F. longicolle, F. mastigosporum and F. procumbens are all segregates of F. longipes s. lat., the latter species here confined to a narrower phylogenetic and morphological circumscription. A slower growth rate on PDA (less than 4.6 mm/d) distinguishes F. cygneum, F. longicolle, and F. procumbens, from F. dolichosporum, F. magnum, and F. mastigosporum (all above 7.9 mm/d). In addition, F. cygneum lacks microconidia and does not produce secondary, falcate aerial macroconidia; which further differentiates this species from F. dolichosporum and F. mastigosporum, respectively. Fusarium cygneum growth on CLA was almost pionnotal, only a few erect conidiophores were rarely observed. This separates F. cygneum from F. longicolle and F. procumbens, the two latter species also showing shorter and less septate macroconidia (av. length 128.7 µm, and up to 11-septate vs 78 µm, up to 9-septate; and 106.1 µm, up to 8-septate in F. longicolle and F. procumbens, respectively).
Fusarium dactylidis T. Aoki et al., Mycologia 107: 412. 2015. Fig. 32.
Fig. 32.
Fusarium dactylidis (ex-type CBS 119181). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–L. Conidiophores and conidiogenous cells. M. Macroconidia. Scale bars = 10 μm.
Description and illustration: Aoki et al. (2015).
Typus: New Zealand, Manawatu, Palmerston North, from Dactylis glomerata, 30 Oct. 1958, J.M. Dingley No. 5941 (holotype BPI 892886, ex-type culture CBS 119181 = ICMP 5269 = NRRL 29298).
Additional material examined: USA, Oregon, from Dactylis glomerata, 1983, R.E. Welty 2A-1, culture CBS 123656 = FRC R-7593 = NRRL 29380.
Notes: A member of the Graminearum clade of FSAMSC; however, not included in the F. graminearum s. lat. clade (also termed F. graminearum SC). Fusarium dactylidis, known from forage grasses in New Zealand and USA (Aoki et al. 2015, Farr et al. 2023), is a nivalenol and zearalonone producer, and a mild inducer of head blight of wheat under experimental conditions (Aoki et al. 2015).
A sister species to the crown pathogen F. pseudograminearum (Leslie & Summerell 2006), F. dactylidis can be distinguished by its shorter and wider conidia, and the absence of chlamydospores. Microconidia (aseptate in F. dactylidis, and 0–2-septate in F. pseudograminearum), absent in the protologue of both species, were seldomly seen in the ex-types of both F. dactylidis and F. pseudograminearum. In addition to morphological differences, the known biogeographical and host distribution of F. dactylidis is reduced compared to that of F. pseudograminearum, known from Africa, Europe, North America and Oceania; mostly in Poaceae crops, Fabaceae, Rosaceae, and soil (see additional details under F. pseudograminearum) (Farr et al. 2023).
Fusarium dimorphosporum Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, sp. nov., MycoBank MB 855102. Fig. 33.
Fig. 33.
Fusarium dimorphosporum (ex-type CBS 151959). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, G, H, J. Conidiophores and conidiogenous cells. F. Chlamydospores. I. Microconidia. K. Aerial macroconidia. L. Sporodochial macroconidia. Scale bars = 10 μm.
Etymology: From Latin dimorphus (“existing in two different forms”) and spora (“spore”). In reference to the two different types of macroconidia.
Sporodochia not seen. Aerial conidiophores erect on aerial mycelium and prostrate or submerged on substrate mycelium, simple or sparsely branched, 14–150 µm tall, mostly reduced to monophialides on aerial mycelium and phialidic pegs on substrate mycelium. Aerial conidiogenous cells monophialidic, subulate to subcylindrical or doliiform, smooth- and thin-walled, 7.5–16.5 × 3–5.5 µm, with short conspicuous apical collarette, giving rise to microconidia and falcate conidia; phialidic pegs 1.5–5.5 × 1.5–2.5 µm, often with a flared apical collarette, giving rise to falcate conidia. Microconidia ellipsoidal to short falcate, 0–1-septate, (4.5–)11–22(–23) × (2.5–)3–4 µm (av. 16.7 × 3.3 μm); falcate conidia of two types: a) produced below and on the agar surface, from phialidic pegs, straight to moderately dorsiventrally curved, often with a straight ventral line, widest above the median, apical cell straight to barely curved, blunt, rarely pointy; basal cell papillate to poorly developed foot-shaped, 1–3(–6)-septate, (17.5–)24–42(–58) × (2.5–)3.5–5(–6) µm (av. 33 × 4 μm), mostly 3-septate, (22.5–)26–37.5(–44) × 2.5–4.5 µm (av. 32.1 × 3.8 μm); b) produced on phialides on aerial mycelium, robust, almost straight to moderately curved, with an almost straight ventral line and typically widest to somewhat swollen at the median, tapering abruptly to both ends, septa thick and conspicuous, apical cell moderately elongated and curved to a blunt to pointy apex; basal cell well-developed foot-shaped, (3–)5–6(–7)-septate, (20.5–)41.5–54(–58.5) × (3.5–)5–7 µm (av. 47.7 × 6 μm), mostly 5-septate, (38.5–)43.5–52.5(–55) × (4–)5–7 µm (av. 48 × 6 μm). Chlamydospores subglobose, smooth-walled, (5.5–)7–9.5(–10.5) µm, singly or in pairs borne on short stipes.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.8–9.4 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface membranous to velvety, rosy buff, scarlet to coral at centre, with patches of white mycelium and luteous to pure yellow mycelial ropes at centre, flat, aerial mycelium scant, margin regular and filiform; reverse sulphur yellow to pale luteous, scarlet to coral at centre. On OA, pure yellow to luteous, flat, dusty to cottony, with regular and filiform margin; reverse honey to cinnamon.
Typus: Thailand, Tak Fa, Nakhon Sawan, from Zea mays, before 18 Mar. 1991, unknown collector (holotype designated here CBS H-25420, ex-type culture CBS 151959 = FRC R-9230 = NRRL 66938).
Additional materials examined: Thailand, from Arachis hypogaea, before 25 Apr. 2016, U. Thrane, culture BBA 64675 = DSM 116017. Philippines, Tarlac, from grassland soil, before 1 Aug. 1981, unknown collector, culture NRRL 64049 = FRC R-6436 = KOD 1988.
Notes: Previously assigned to Fusarium sp. nov. 32 in Laraba et al. (2021), F. dimorphosporum is characterized by gibbosum-like sporodochial macroconidia, similar to those of F. cuspidatum, F. hamatum and F. platysporum (Brachygibbosum clade); and F. pratense, and F. tropicale (Longipes clade). Fusarium dimorphosporum, as indicated by its epithet, produces two types of macroconidia (gibbosum-like sporodochial conidia; and falcate, almost straight aerial macroconidia) in addition to aerial microconidia. This combination of features differentiates F. dimorphosporum from all the similar species mentioned above.
Fusarium dolichosporum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855103. Fig. 34.
Fig. 34.
Fusarium dolichosporum (ex-type CBS 151817). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–H. Conidiophores and conidiogenous cells. I. Microconidia. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin dolicho- (“long”) and spora (“spore”). In reference to its macroconidial length.
Sporodochia not seen. Aerial conidiophores prostrate, emerging from substrate mycelium, rarely erect on aerial mycelium, reduced to solitary lateral monophialides or branched laterally two or three times at the apical end, 33–278 µm tall. Conidiogenous cells monophialidic, subcylindrical, 2.5–22 × 2.5–5 µm. Macroconidia, whip-like, rarely falcate, robust, thick-walled, strongly dorsiventrally curved or straight; apical cell tapering, slightly curved, hooked to strongly elongated, basal cell well-developed foot-shaped, elongated with a tapering, somewhat pointy appendage, (0–)3–11(–17)-septate, (31.5–)61–160(–209) × 3.5–6 µm (av. 110.9 × 4.6 µm), commonly 10-septate (111.5–)117–161(–173) × 4–5.5 µm (av. 139 × 4.8 µm). Microconidia, fusiform, 0–2-septate, (14.5–)15.5–21(–23) × 2.5–3.5 µm, rarely produced intermixed with macroconidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 9.4–10.6 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface velvety, pale luteous to pale coral with abundant white to pale grey woolly aerial mycelium, flat, margin regular and filiform; reverse pale luteous with peach to scarlet or coral patches. On OA, amber to honey with white woolly patches, flat, membranous at centre, cottony to woolly at periphery, margin regular, undulate; reverse cinnamon to ochreous.
Typus: South Africa, from Triticum sp., before 1 May 1977, unknown collector (holotype designated here CBS H-25418, ex-type culture CBS 151817 = FRC R-4069 = MRC 5-B-J = NRRL 66921).
Additional material examined: South Africa, Transkei, debris, before 1 Jan. 1986, unknown collector, culture FRC R-8203 = KOD 2016 = NRRL 64077.
Notes: Previously termed as Fusarium sp. nov. 15 (Laraba et al. 2021), Fusarium dolichosporum presents remarkably long, whip-like conidia, comparable to those of F. cygneum, F. longicolle, F. magnum, F. mastigosporum, and F. procumbens. See additional comments under F. cygneum. Fusarium dolichosporum, together with F. magnum and F. mastigosporum, can be separated by their fast-growing PDA colonies (all above 7.9 mm/d). Moreover, F. dolichosporum presents the greatest maximum conidial size (209 µm long); however, the fourth largest average conidial size (110.9 × 4.6) after F. cygneum (128.7 × 4.4 µm), F. mastigosporum (127.7 × 4.3 µm), and F. magnum (117.5 × 5.3 µm); and in addition to the lack of sporodochia, among the species mentioned above, only F. dolichosporum produces microconidia and luteous PDA colonies.
Fusarium gerlachii T. Aoki et al., Fungal Genet. Biol. 44: 1202. 2007. Fig. 35.
Fig. 35.
Fusarium gerlachii (ex-type CBS 119175). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–G. Sporodochia. H–J. Sporodochial conidiophores and conidiogenous cells. K. Aerial conidiophore and conidiogenous cell. L. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: Starkey et al. (2007), Aoki et al. (2012).
Typus: USA, Minnesota, Polk County, Climax, from Triticum aestivum, 25 Jul. 2000, L. Gale (holotype BPI 871657, ex-type culture CBS 119175 = LRG 00-551 = NRRL 36905).
Additional materials examined: USA, Wisconsin, from Arundo donax, unknown collection date, L. Gale, culture CBS 119176 = NRRL 38380; North Dakota, from Triticum sp., unknown collection date, L. Gale, culture CBS 123666 = HK-3ND-7-17 = NRRL 38405.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. In comparison to other F. graminearum s. lat. segregates, F. gerlachii has a reduced host and geographical range, being known from Poaceae (Arundo donax, Dactylis glomerata, and Triticum aestivum) in the USA and New Zealand (Aoki et al. 2012, Farr et al. 2023). It is morphologically distinguished by producing 4.5–5 µm wide, gradually curved, asymmetric 5-septate macroconidia, with beaked apical cells (Aoki et al. 2012).
Fusarium gladiolum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855104. Fig. 36.
Fig. 36.
Fusarium gladiolum (ex-type CBS 151812). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F, G. Sporodochial conidiophores and conidiogenous cells. H, I. Aerial conidiophores and conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin gladiolus (“little sword”). Referring to the macroconidial shape, resembling a sword.
Sporodochia abundant on the surface of carnation leaves, agar surface and aerial mycelium, erumpent, buff, honey to luteous coloured, sporodochia on aerial mycelium lighter, pale pinkish orange. Sporodochial conidiophores densely packed, branching irregularly, 24–98 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, subulate to subcylindrical, 7–23.5 × 2.5–5 µm, smooth- and thin-walled, with a minute or lacking apical collarette, proliferating apically. Sporodochial conidia falcate, gently dorsiventrally curved to almost straight, widest at or above the median, apical cell slightly curved, blunt; basal cell well-developed foot-shaped, (1–)3–5(–8)-septate, (28.5–)41.5–57(–68) × (3–)5–6(–7) µm (av. 49.3 × 5.6 μm), mostly 5-septate, (36–)47–58(–63) × 5–6.5 µm (av. 52.5 × 5.6 μm). Aerial conidiophores prostrate on substrate mycelium, scarce, sparsely branched, 25–41 µm tall, mostly reduced to monophialides forming laterally on substrate mycelium, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subulate to subcylindrical, 10–24 × 4–5.5 µm, smooth- and thin-walled, with a minute apical collarette, proliferating apically, giving rise to falcate conidia undistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 6.0–7.7 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface velvety, dusty to felty, amber, pure yellow to coral with white patches of aerial mycelium, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse coral to rust, bay to brown vinaceous at centre. On OA, amber to pure yellow with patches of white or coral mycelium, flat, velvety to felty, with regular and undulate margin; reverse ochreous to amber.
Typus: New Guinea, Papua, from Ipomoea batatas stem, before 1 Nov. 1988, unknown collector (holotype designated here CBS H-25417, ex-type culture CBS 151812 = NRRL 54640 = FRC L-247).
Additional material examined: New Guinea, Papua, from Ipomoea batatas stem, before 1 Nov. 1988, unknown collector, CBS H-25400, culture CBS 151813 = FRC L-252 = NRRL 54683.
Notes: This phylogenetic species was previously assigned to Fusarium sp. nov. 6 in Laraba et al. (2021). Fusarium gladiolum, together with an unnamed phylogenetic species (FSAMSC7) form a phylogenetically distinctive group here termed the Gladiolum clade of FSAMSC, for which no morphologically comparable sister species exists. The two strains examined here and assigned to F. gladiolum, were originally identified as F. lateritium, isolated from rotten sweet potato stems. The original misidentification makes sense considering the yellowish PDA colonies and the falcate, almost straight macroconidia that resemble those originally reported for F. hanswilhelmii (syn. F. lateritium var. longum) or F. lateritium (Wollenweber 1931, Costa et al. 2024). Nevertheless, the two latter species are distantly related phylogenetically, residing in the FLSC (Costa et al. 2024). Moreover, in addition to its smaller macroconidia, F. gladiolum differs from F. lateritium by lacking microconidia and from both F. hanswilhelmii and F. lateritium by lacking chlamydospores. Among the FSAMSC, conidia of F. gladiolum are in the size and septation range of F. longipes and F. vermicularioides; however, contrary to F. gladiolum, the two latter species produce chlamydospores, microconidia, and two types of macroconidia (falcate on aerial conidiophores, and whip-like on sporodochia).
Fusarium goolgardi D.M. Robinson et al., Fungal Diversity 77: 357. 2015 [2016]. Fig. 37.
Fig. 37.
Fusarium goolgardi (ex-type CBS 151815). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G, H. Aerial conidiophores and conidiogenous cells. I. Microconidia. J. Macroconidia. Scale bars = 10 μm.
Description and illustration: Laurence et al. (2015).
Typus: Australia, New South Wales, Bungonia State Conservation Area, from Xanthorrhoea glauca, Jan. 2010, D.M. Robinson & M.H. Laurence (holotype RBG5411, ex-type culture CBS 151815 = KOD 1087 = NRRL 66250 = RBG5411).
Additional material examined: Australia, New South Wales, Bungonia State Conservation Area, from Xanthorrhoea glauca, Jan. 2010, D.M. Robinson & M.H. Laurence, culture NRRL 66249 = KOD 1090.
Notes: Fusarium goolgardi was originally described causing Xanthorrhoea decline in Australia, with additional isolates reported from soil in the USA (Laurence et al. 2015, Farr et al. 2023). The protologue of this species indicates morphological affinities with F. armeniacum, F. langsethiae, F. sambucinum, F. sporotrichioides and F. venenatum in the FSAMSC and with F. acuminatum and F. avenaceum in the Fusarium tricinctum SC (Laurence et al. 2015). However, there is very little overlap in conidial morphology between F. goolgardi and F. acuminatum, F. armeniacum, F. avenaceum, or F. langsethiae (Fig. 2). Moreover, apart from a close phylogenetic relationship to F. goolgardi, F. langsethiae and its closest sibling F. sibiricum are characterized by not producing macroconidia under any condition, and only producing napiform to globose microconidia, which are not produced by F. goolgardi. Macroconidia of F. goolgardi are larger than those of F. sambucinum and F. sporotrichioides (av. 38.9 × 4.7 µm vs 28 × 5.3 µm and 30.9 × 4.4 µm, respectively), but smaller than those of F. venenatum (av. 44.5 × 5.4 µm) which also present a more pronounced curvature. Additionally, F. goolgardi lacks mesoconidia, which distinguishes it from F. sporotrichioides. Moreover, F. goolgardi is unique among the species mentioned above, also including its closest phylogenetic relative F. nodosum, by the formation of sympodially proliferating and branched aerial monophialides (Laurence et al. 2015).
Fusarium guizhouense Lin Huang et al., MycoKeys 101: 64. 2024.
Description and illustration: He et al. (2024).
Typus: China, Guizhou Province, Qiandongnan Miao and Dong Autonomous Prefecture, Cengong County, Kelou, from Cunninghamia lanceolata leaf spots, May 2017, Wen-Li Cui (holotype CFCC 57575, ex-type culture GZ7-20-1).
Notes: The closest phylogenetic relative of the unnamed phylogenetic species FSAMSC11, F. guizhouense, described as a pathogen of Cunninghamia lanceolata (Cupressaceae, He et al. 2024), was not included in the phylogenetic and morphological analyses due to scarcity of DNA sequences for comparison and unavailability of the ex-type strain. Currently available sequences for the species (partial rpb1, rpb2 and tef1) are considerably shorter than the average sequence length included here. However, a separate analysis with trimmed sequences (not shown) confirmed the phylogenetic position of this species as indicated in the protologue (He et al. 2024). For a morphological comparison with related FSAMSC taxa see He et al. (2024).
Fusarium graminearum Schwabe, Fl. Anhalt. 2: 285. 1839. Figs 38, 39.
Fig. 38.
Fusarium graminearum (lectotype). Reproduction from the original publication by Schwabe (1884). A, B. Macroconidia.
Fig. 39.
Fusarium graminearum (ex-epitype CBS 136009). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiogenous cell. J. Macroconidia. Scale bars = 10 μm.
Synonyms: Sphaeria zeae Schwein., Schriften Naturf. Ges. Leipzig 1: 48. 1822.
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 gynerii Cooke & Harkn. ex Sacc., Syll. fung. (Abellini) 4: 713. 1886.
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.
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.
Pionnotes flavicans Sacc. & D. Sacc., Atti Reale Ist. Veneto: 720. 1902.
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 discolor var. majus Wollenw., Bull. Maine Agric. Exp. Sta. 219: 255. 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.
Descriptions and illustrations: Booth (1971), Gerlach & Nirenberg (1982), and Leslie & Summerell (2006), Domsch et al. (2007), Aoki et al. (2012).
Typus: Germany, from inflorescence of Triticum sp., 1839, S.H. Schwabe [lectotype tab. VI, fig. 7 in Schwabe (1839), Flora Anhaltina 2, designated in Crous et al. (2021)]; from Hordeum vulgare grain, 1988, L. Niessen [epitype CBS 136009, preserved as metabolically inactive culture, designated by Crous et al. (2021), ex-epitype culture CBS 136009].
Additional materials examined: Iran, from Zea mays, 1968, W. Gerlach, culture CBS 792.70 = BBA 11141 = NRRL 25471; from Triticum sp. cultivar Falat, unknown collection date and collector, culture CBS 130963; from Triticum sp. cultivar Tajan, Apr. 2010, M. Davari, culture CBS 130666; from Triticum sp., cultivar N 8020, Apr. 2010, M. Davari, culture CBS 130887; Bilesovar, Babak, from Triticum sp. cultivar Koohdasht, unknown collection date and collector, dep. M. Davari, culture CBS 130953; Mohan R. Company, from Triticum sp. cultivar 8019, Apr. 2010, M. Davari, culture CBS 130646; from Triticum sp. cultivar Chamran, unknown collection date and collector, dep. M. Davari, culture CBS 130960; Parsabad, from Triticum sp., Apr. 2010, M. Davari, culture CBS 130914; college, from Triticum sp., cultivar N8019, Apr. 2010, M. Davari, culture CBS 130892; Govashlou, from Triticum sp., cultivar Atila 50, 2010, M. Davari, culture CBS 130604. Poland, damaged Triticum sp., Kernel, unknown collection date and collector, dep. T. Kulik & A. van Diepeningen, culture CBS 138563. Unknown, before 18 Jan. 1921, unknown collector, dep. L.L. Harter, culture CBS 124.21 = NRRL 29285; from Zea mays stalk, before 15 May 1932, unknown collector, dep. E.C. Stakman, culture CBS 184.32 = NRRL 29288, CBS 185.32, CBS 189.32 = NRRL 36335. USA, before 28 Feb. 1925, unknown collector, dep. L.H. Leonian, culture CBS 153.25 = NRRL 29286; Ohio, from Zea mays, unknown collection date and collector, culture CBS 110261 = ATCC 46779 = MAFF 237812 = NRRL 5883 = NRRL 38132; Michigan, from Zea mays stalk, before 1 Oct. 1981, L.P. Hart, culture CBS 110264 = FRC R-6574 = MAFF 237824 = NRRL 13823 = NRRL 28063 = NRRL 38135; from Zea mays, unknown collection date, H. Kistler, culture CBS 123657 = NRRL 31084.
Notes: Fusarium graminearum is the main cause of fusarium head blight of cereals, associated with epidemics in the USA during the 1990s (O’Donnell et al. 2000), it is also known causing cob, foot, head, root or stalk rots; and branch or stub dieback (Leslie & Summerell 2006, Domsch et al. 2007). This species is known from soil, insects (O’Donnell et al. 2012), and is a colonizer or parasite of diverse plant hosts, recorded from over 100 host species, from 81 different genera, mainly in Poaceae, but also in Amaranthaceae, Amaryllidaceae, Apiaceae, Arecaceae, Asparagaceae, Asteraceae, Cannabaceae, Caryophyllaceae, Cucurbitaceae, Dryopteridaceae, Fabaceae, Lauraceae, Linaceae, Lythraceae, Malvaceae, Musaceae, Myrtaceae, Orchidaceae, Pinaceae, Polygonaceae, Rubiaceae, Salicaceae, Solanaceae, Vitaceae, and Zygophyllaceae (Farr et al. 2023). Nevertheless, host records for F. graminearum need re-evaluation according to the current taxonomy of this group.
Originally thought to be a single species, F. graminearum s. lat. has recently been split into several taxa, with narrow biological, ecological, and pathological characteristics (O’Donnell et al. 2004, 2008, Leslie & Summerell 2006, Domsch et al. 2007). First, a heterothallic population known as F. graminearum Group 1 was separated as F. pseudograminearum, its sexual morph described as G. coronicola (Aoki & O’Donnell 1999a, 1999b). Later, a succession of studies resulted in the split of the homothallic population known as F. graminearum Group 2, into 15 phylogenetically and biogeographically distinct species i.e., F. graminearum s. str., F. gerlachii, F. cortaderiae, F. brasilicum, F. austroamericanum, F. meridionale, F. louisianense, F. boothii, F. acaciae-mearnsii, F. vorosii, F. ussurianum, F. asiaticum, F. aethiopicum, F. nepalense, and F. mesoamericanum; in addition to a number of undescribed lineages (O’Donnell et al. 2004, 2008, Starkey et al. 2007, Yli-Mattila et al. 2009, Sarver et al. 2011, Aoki et al. 2012). Our phylogenetic and species delimitation analyses uncovered a wide intraspecific diversity within F. graminearum s. str. The nomenclature of F. graminearum was recently stabilized by lecto- and epitypification by Crous et al. (2021): the lectotype, an original illustration by S.H. Schwabe, is reproduced here (Fig. 38).
Fusarium graminearum is morphologically undistinguishable from F. asiaticum and F. aethiopicum, the tree species producing 4.5–5 µm, curved and asymmetric, 5-septate macroconidia without apical beaks and wider above the median (Aoki et al. 2012), but see additional notes under the respective species.
Fusarium hamatum Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855105. Fig. 40.
Fig. 40.
Fusarum hamatum (ex-type CBS 151936). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F, H. Aerial conidiophores and conidiogenous cells. G. Chlamydospores. I. Sporodochial conidiophore and conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin hāmātus (“hooked, crooked”). In reference to the macroconidial shape.
Sporodochia abundant on the surface of carnation leaves, agar surface and aerial mycelium, cinnamon, luteous to orange coloured. Sporodochial conidiophores branching laterally irregularly and verticillately, 24.5–65 µm tall, bearing terminal monophialides. Sporodochial conidiogenous cells monophialidic, ampulliform to subcylindrical, 16.5 × 3–5 µm, smooth, thin-walled, with a minute apical collarette, proliferating apically. Sporodochial conidia falcate, gently to strongly dorsiventrally curved, widest at or above the median, apical cell pyramidal or barely to moderately elongated with blunt apex; basal cell well-developed foot-shaped, 5–6-septate, (34–)41.5–53(–63) × 5–7 µm (av. 47.4 × 6.2 μm), mostly 5-septate, (34–)41.5–53(–60.5) × 5–7 µm (av. 47.1 × 6.2 μm). Aerial conidiophores erect or prostrate on substrate mycelium, mostly as simple phialides borne laterally on hyphae, with or without a short stipe, 18–47 µm tall, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical, smooth- and thin-walled, 7–16.5 × 2.5–5 µm, giving rise to falcate conidia undistinguishable from sporodochial conidia. Chlamydospores globose, smooth- and thick-walled, 5.5–13.5 µm diam, at first hyaline, becoming pale to golden brown, commonly grouped in chains, rarely solitary or in clusters.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.2–10.6 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface velvety to woolly, scarlet to red, flat, aerial mycelium abundant, margin regular and filiform; reverse scarlet to coral. On OA, pure yellow to amber with peach to coral patches, rosy buff at periphery, flat, cottony to woolly, with regular and filiform margin; reverse amber to scarlet.
Typus: Madagascar, from Striga asiatica, before 25 Apr. 2016, J. Kroschel (holotype designated here CBS H-25422, ex-type culture CBS 151936 = BBA 67756 = DSM 116550).
Additional material examined: Germany, Brandenburg, from Triticum aestivum, before 25 Apr. 2016, U. Damm, culture BBA 69069 = DSM 116669.
Notes: Fusarium hamatum was resolved as the closest phylogenetic relative to FSAMSC31 (Figs 1, 3). The latter phylogenetic species was not studied morphologically, thus further comparisons are not possible. Macroconidia of F. hamatum shows similar characteristics to those of F. cuspidatum, F. dimorphosporum, F. platysporum, F. pratense, and F. tropicale; all six species producing gibbosum-like, abruptly tapered conidia, often with slightly elongated apical cells. Fusarium hamatum differs from F. dimorphosporum, F. platysporum, and F. pratense by the absence of microconidia; and additionally, differs from F. dimorphosporum by lacking a secondary macroconidial type, and from F. pratense by its red PDA colonies (yellow in F. pratense). Conidial dimensions largely overlap between F. hamatum, F. cuspidatum and F. tropicale; however, F. tropicale is phylogenetically distant, and does not belong to the Brachygibbosum clade, but to the Longipes clade. In contrast, conidia of F. hamatum often present a more elongated and curved apical cell, with a stronger overall dorsiventral curvature, and pigmented chlamydospores are rapidly produced.
Fusarium kyushuense O’Donnell & T. Aoki, Mycoscience 39: 2. 1998. Fig. 41.
Fig. 41.
Fusarium kyushuense (ex-type CBS 151828). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–I. Conidiophores and conidiogenous cells. J. Microconidia. K. Macroconidia. Scale bars = 10 μm.
Description and illustration: Aoki & O’Donnell (1998).
Typus: Japan, Kyushu, Kumamoto, from Triticum aestivum seed, 1963, H. Tsunoda (holotype NIAES 99701, ex-type culture CBS 151828 = ATCC 56750 = FRC T-346A = MAFF 237645 = MRC 1767 = NRRL 3509 = NRRL A-14732).
Additional materials examined: China, Guizhou province, Fuquan, unknown host and collection date, J. Bangming, culture CBS 121807 = NRRL 53748. Japan, from Triticum aestivum, unknown collection data, culture NRRL 25349; Kuyshu, Kumamoto, from Triticum aestivum, unknown collection date, H. Tsunoda, culture BBA 71679.
Notes: Fusarium kyushuense has characteristic sympodially and percurrently proliferating holoblastic conidiogenous cells on aerial conidiophores, producing clavate, obovate to subglobose microconidia, and mesoconidia; comparable to those produced by F. sporotrichioides. Fusarium kyushuense is distinguished from F. sporotrichioides by the absence of chlamydospores, and slower growth rates on PDA (av. 3.8 mm/d vs 6.8 mm/d in F. sporotrichioides). Additional morphological comparisons with distantly related species outside of FSAMSC are included in Aoki & O’Donnell (1998). As mentioned in the protologue of the species, although not illustrated, phialidic conidiogenous cells are also present on aerial conidiophores, producing basipetal chains of conidia which are shown here (Fig. 41). Originally described from Japan, additional isolations have been made in China and England, mainly from Poaceae (Hordeum vulgare, Oryza sativa, Triticum aestivum, and Zea mays), but also from Ericaceae (Chamaedaphne calyculata), Fagaceae (Lithocarpus glabra), Musaceae (Musa nana), and Solanaceae (Solanum tuberosum) (Wang et al. 2022, Farr et al. 2023).
Fusarium langsethiae Torp & Nirenberg, Int. J. Food Microbiol. 95: 248. 2004. Fig. 42.
Fig. 42.
Fusarium langsethiae (ex-type CBS 113234). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–H. Conidiophores and conidiogenous cells. I. Microconidia. Scale bars = 10 μm.
Description and illustration: Torp & Nirenberg (2004).
Typus: Norway, from kernel of Avena sativa, 1998, M. Torp (holotype B 700012234, ex-type culture CBS 113234 = BBA 70945 = DSM 116700 = ITEM 3602 = NRRL 53627).
Additional materials examined: Norway, from Avena sp., 1998, M. Torp, culture NRRL 54940. UK, Berkshire, from Triticum sp., 5 Nov. 1982, E. Niles, cultures IMI 272890, IMI 272894, and IMI 272899; Derbyshire, from Triticum sp., 30 Sep. 1982, J. Ibbotson, culture IMI 271486; Kent, from Triticum sp., 18 Oct. 1982, M. Bateson, culture IMI 271971.
Notes: Although a weak pathogen of Avena sativa and Hordeum vulgare, F. langsethiae is, however, the main cause of trichothecene contamination of small grain cereals in Europe (Lysøe et al. 2016), with reports mostly from Europe (Finland, Italy, Norway, Poland, Russia, and the UK) and Syria (Farr et al. 2023); and has been associated with human infections (Al-Hatmi 2016).
Originally compared to F. sporotrichioides, due to sharing similar mycotoxin profiles and features of its microconidia (Torp & Nirenberg 2004), both taxa are phylogenetically closely related within the Sporotrichioides clade of FSAMSC. Several morphological differences separate these two species including the abundant polyblastic conidiogenous cells and mesoconidia of F. sporotrichioides; the napiform to globose, but never fusiform conidia of F. langsethiae, which does not produce typical Fusarium multiseptate conidia, and slow growing (av. 3.8 mm/d) PDA colonies on the latter species, which lack red diffusible pigments (vs av. 6.8 mm/d red PDA colonies in F. sporotrichioides). As predicted in the protologue of F. langsethiae (Torp & Nirenberg 2004), most strains studied here were received as F. poae, a distant species in the Sambucinum clade, which differ from F. langsethiae by producing, although not constant, typical fusarioid sporodochial conidia and fast-growing (av. 9.7 mm/d), red PDA colonies.
Fusarium leptum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855106. Fig. 43.
Fig. 43.
Fusarium leptum (ex-type CBS 151949). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochium. F, G. Sporodochial conidiophores and conidiogenous cells. H–J. Aerial conidiophores and conidiogenous cells. K. Macroconidia. Scale bars: F, G, K = 10 μm; I, J = 5 μm.
Etymology: From Greek leptós (“thin, slender”). Referring to the slender macroconidia formed by this species.
Sporodochia punctiform, erumpent, abundant on the surface of carnation leaves and agar surface, buff to luteous coloured. Sporodochial conidiophores sparingly irregularly branching, 39.5–48 µm tall, bearing solitary lateral and terminal, and terminal whorls of monophialides. Sporodochial conidiogenous cells monophialidic, subulate to subcylindrical, 7.5–19 × 2–4.5 µm, smooth- and thin-walled, with short, inconspicuous apical collarette. Sporodochial conidia falcate, dorsiventrally curved with an almost straight ventral line, tapering towards both ends, apical cell elongated with a blunt to pointy apex, basal cell well-developed foot-shaped, rarely papillate, 1–3(–4)-septate, (28.5–)36.5–46(–53.5) × 3–5 µm (av. 41.5 × 4 μm), mostly 3-septate, (28.5–)37–46.5(–53.5) × 3–5 µm (av. 41.7 × 4 μm). Aerial conidiophores erect or prostrate on substrate mycelium, simple or sparsely branched, 18–50 µm tall, mostly reduced to monophialides forming laterally on aerial mycelium. Aerial conidiogenous cells monophialidic, subcylindrical to clavate, 6.5–15 × 3–4 µm, smooth- and thin-walled, with a minute apical collarette, giving rise to falcate conidia undistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.2–8.7 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface velvety to woolly, straw to sulphur yellow, orange at centre due to accumulation of sporodochia, flat, aerial mycelium scarce as white ropes irregularly distributed, margin regular, filiform to undulate; reverse straw to orange at centre. On OA, luteous to orange, flat, velvety becoming membranous to slimy due to accumulation of sporodochia, margin regular and filiform, reverse pale luteous to pure yellow.
Typus: Netherlands, unknown host and collection data (holotype designated here CBS H-25416, ex-type culture CBS 151949 = NRRL 29896 = FRC R-9673).
Additional material examined: Netherlands, unknown host and collection data, culture NRRL 29897 = FRC R-9674.
Notes: Previously assigned to Fusarium sp. nov. 4 in Laraba et al. (2021), F. leptum is currently only known from the Netherlands; however, host data are not available. Macroconidia of F. leptum are in the size range of the distantly related species F. goolgardi (Sporotrichioides clade), and F. sagittatum (Brachygibbosum clade). Conidia of F. leptum differ by being slender and conspicuously wider at the lower portion of the conidium (length wise). Additionally, F. leptum lacks microconidia, which further distinguishes this species from F. goolgardi, F. sagittatum, and all the phylogenetically close species in the Sporotrichioides B clade (i.e., F. armeniacum, F. chaquense, F. palustre, and F. parabolicum).
Fusarium longicolle Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855107. Fig. 44.
Fig. 44.
Fusarium longicolle (ex-type CBS 151818). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G. Sporodochial conidiophores and conidiogenous cells. H, I. Aerial conidiophores and conidiogenous cells. J. Macroconidia. Scale bars: G, H, J = 10 μm; I = 5 μm.
Etymology: From Latin longus (“long, extended, prolonged”) and collum (“neck”). Referring to the long apical extensions of the macroconidia.
Sporodochia abundant on the surface of carnation leaves and agar surface, erumpent, orange, pale luteous, luteous to honey coloured. Sporodochial conidiophores branching laterally, 23.5–38 µm tall, bearing lateral and terminal monophialides, single or grouped in verticils. Sporodochial conidiogenous cells monophialidic, subcylindrical, (8.5–)11.5–16.5(–18) × (2.5–)3–4(–5) µm, smooth, thin-walled, with a minute apical collarette. Sporodochial conidia whip-like, robust, strongly dorsiventrally curved or recurved, with an elongated apical cell; basal cell well-developed, foot-shaped and elongated, (3–)5–7(–9)-septate, (37.5–)62–94(–109) × (3–)4–5(–6) µm (av. 78 × 4.6 μm), mostly 6-septate, (58–)63.5–96.5(–107) × 4–5(–6) µm (av. 79.9 × 4.6 μm). Aerial conidiophores rare, scattered, mostly between sporodochial conidiophores or prostate on substrate mycelium, sparsely branched, 11–21.5 µm tall, often reduced to monophialides forming laterally on aerial mycelium or submerged in agar. Aerial conidiogenous cells monophialidic, ampulliform to subcylindrical, 9–13 × 3–5 µm, giving rise to conidia undistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 3.6–4.6 mm/d, 50–65 mm diam after 7 d. Surface velvety, dusty to woolly, red to scarlet, flesh to coral at centre, flat, aerial mycelium abundant at centre, margin regular, filiform to undulate; reverse coral to scarlet, citrine to isabelline at centre. On OA, pale luteous, luteous to ochreous, flat, velvety with abundant dusty radial patches, margin regular, filiform to undulate; reverse pale luteous to luteous.
Typus: Australia, Rockhampton, from soil debris, before 1 Oct. 1982, unknown collector (holotype designated here CBS H-25406, ex-type culture CBS 151818 = FRC R-6968 = NRRL 66924).
Additional materials examined: Australia, Northern Territory, unknown substrate, 1997, K. Gott, CBS H-25402, culture CBS 120991 = FRC R-9983 = MRC 8429 = MRC 8430 = KSU 11429 = KSU 11431 = Univ. Sydney F11993 = Univ. Sydney F5837; Queensland, Rockhampton, from air, before 1 Aug. 1982, unknown collector, culture FRC R-6894 = KOD 1992 = NRRL 64053.
Notes: Fusarium longicolle was previously assigned to Fusarium sp. nov. 18 in Laraba et al. (2021). This species, together with F. cygneum, F. dolichosporum, F. magnum, F. mastigosporum, and F. procumbens, are all characterized by forming large, whip-like macroconidia with exceptionally long apical cells (see additional comments under F. cygneum). An initial distinction based on their comparatively slower growth rates (less than 4.5 mm/d on average) separates F. longicolle, together with F. cygneum and F. procumbens, from the faster growing (averaging above 8.3 mm/d) F. dolichosporum, F. magnum, and F. mastigosporum. Additionally, conidia of F. longicolle are the shortest among the three closest species mentioned above, both in overall size (av. 78 × 4.6 μm vs 128.7 × 4.4 μm, and 106.1 × 4.3 μm, for F. cygneum and F. procumbens, respectively), and 6-septate conidia (av. 79.9 × 4.6 μm vs 123.5 × 4.4 μm and 112.8 × 4.4 μm, for F. cygneum and F. procumbens, respectively), the most common degree of septation observed for all three species.
Fusarium longipes Wollenw. & Reinking, Phytopathology 15: 160. 1925. Fig. 45.
Fig. 45.
Fusarium longipes (ex-neotype CBS 476.77). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G. Sporodochial conidiophores and conidiogenous cells. H–J. Aerial conidiogenous cells. K. Aerial macroconidia. L. Sporodochial macroconidia. Scale bars: G, K, L = 10 μm; H–J = 5 μm.
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. Nom. inval., Art. 41.1 (Melbourne).
Descriptions and illustrations: Reinking & Wollenweber (1927), Wollenweber (1930), Gerlach & Nirenberg (1982), Nelson et al. (1983).
Typus: USA, Florida, from soil, 1977, W. Gams [neotype CBS 476.77, preserved as metabolically inactive culture, designated by Crous et al. (2021), ex-neotype culture CBS 476.77 = FRC R-09841 = NRRL 20695].
Additional materials examined: Denmark, from Oryza sativa grain, 18 Dec. 1968, unknown collector, dep. C. Booth, culture CBS 120.73 = ATCC 24359 = IMI 136675 = NRRL 20694. India, from Cymbopogon martinii, culture IMI 196466 = NRRL 20724; Uttar Pradesh, Varanasi, from Psidium guajava root, Nov. 1986, unknown collector, culture IMI 311604. Malaysia, from soil, unknown collection date, W. Gerlach, culture CBS 739.79 = BBA 62061 = DSM 62061 = FRC R-9842 = NRRL 20696; Borneo, Sabah, from Oryza sativa leaf, unknown collection date and collector, culture IMI 179815 = NRRL 20725; Sandakan, Kolapis, from Acacia mangium decaying or rotted heartwood, unknown collection date and collector, culture IMI 357087.
Notes: The traditional, morphology-based concept of F. longipes (F. longipes s. lat.) is paraphyletic, delineating a species complex with significant genetic, morphological and biochemical diversity (O’Donnell et al. 2012, Crous et al. 2021, Laraba et al. 2021). Crous et al. (2021) fixed the application of the name by neotypification, confining F. longipes to a discrete phylogenetic clade (recognized as F. longipes 4 in Laraba et al. 2021). Nine additional segregate species are recognized and described here in the Longipes clade.
Species in the Longipes clade can be recognized by the typical shape of their macroconidia (whip-like, prominently curved, changing abruptly in width towards both extremes, often with prominently extended apical and foot cells); although, with three exceptions (i.e., F. carinatum, F. pratense, and F. tropicale) which do not produce extended apical and basal cells. However, F. longipes, together with F. mastigosporum and F. vermicularioides can produce a secondary type of falcate macroconidium on aerial conidiophores, which lacks extended apical cells. Additionally, comparable, large, whip-like macroconidia can be found in the distant Sambucinum clade (F. dolichosporum and F. magnum). The presence of two macroconidial types, however, differentiates F. longipes, F. mastigosporum and F. vermicularioides from F. cygneum, F. longicolle, F. magnum, F. procumbens, and F. pseudolongipes, while F. dolichosporum produces very long, whip-like macroconidia, from aerial conidiophores only. Furthermore, F. longipes differs from F. mastigosporum by the absence of chlamydospores and microconidia, and significantly larger macroconidia in the latter species (av. 45.1 × 4.5 µm in F. longipes vs 127.7 × 4.3 µm in F. mastigosporum); and differs from F. vermicularioides by its fast growing, red PDA colonies (av. 13 mm/d in F. longisporum, vs yellow colonies, averaging 6.9 mm/d in F. vermicularioides).
Fusarium longipes is known from subtropical and tropical regions, including Africa (Nigeria, South Africa, and Zambia), Central and North America (Puerto Rico and USA), Asia (China, India, Indonesia, Malaysia, Philippines, Singapore and Thailand), and Oceania (Australia, Micronesia, and Papua-New Guinea), with additional reports from Europe (UK and Denmark) (Gerlach & Nirenberg 2982, Wang et al. 2022, Farr et al. 2023). It is often recorded from undisturbed and agricultural soil and soil debris, but also from plant hosts mainly in Poaceae (genera Cymbopogon, Oriza, Paspalum, and Triticum), but also in Araceae (Cyrtosperma chamissonis), Fabaceae (Acacia, and several unidentified bean species), and Myrtaceae (Psidium guajava). However, known host ranges must be re-assessed according to current species concepts.
Fusarium louisianense L.R. Gale et al., Fungal Genet. Biol. 48: 1105. 2011. Fig. 46.
Fig. 46.
Fusarium louisianense (ex-type CBS 127525). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G, H. Aerial conidiophores and conidiogenous cells. I. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: Sarver et al. (2011), Aoki et al. (2012).
Typus: USA, Louisiana, Jefferson Davis Parish Lou, from Triticum aestivum seed, 2007, L.R. Gale (holotype BPI 881005, ex-type culture CBS 127525 = NRRL 54197).
Additional material examined: USA, Louisiana, Jefferson Davis Parish Lou, from Triticum aestivum head, 2007, L.R. Gale, culture CBS 127524 = NRRL 54196.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Fusarium louisianense, so far only known from Triticum spp. in the USA (Aoki et al. 2012, Farr et al. 2023), is morphologically distinguishable from other segregate species of F. graminearum s. lat., by its non-beaked, 4–4.5 µm wide, curved and asymmetric 5-septate conidia, which are wider at the median (Aoki et al. 2012).
Fusarium lunulosporum Gerlach, Phytopathol. Z. 88: 283. 1977. Fig. 47.
Fig. 47.
Fusarium lunulosporum (ex-type CBS 636.76). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochium. F, G. Sporodochial conidiophores and conidiogenous cells. H. Aerial conidiogenous cells. I. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: Gerlach (1977b), Gerlach & Nirenberg (1982) and Nelson et al. (1983).
Typus: South Africa, from Citrus paradisi, May 1968, R. Schwarz (holotype BBA 62459, ex-type culture CBS 636.76 = ATCC 36747 = BBA 62459 = FRC R-5822 = IMI 322097 = NRRL 13393).
Notes: Fusarium lunulosporum belongs to the Graminearum clade of FSAMSC. However, it does not pertain to the F. graminearum s. lat. clade, but resolved as a basal clade to the sister species F. cerealis and F. culmorum. These three species share common morphological characteristics, including red PDA colonies, lack of microconidia, and falcate, predominantly 5-septate macroconidia of similar lengths. With its slender, and more prominently curved conidia, F. lunulosporum is significantly different not only from its closest siblings (F. cerealis and F. culmorum), but from all other species in the Graminearum clade. Fusarium lunulosporum is an infrequent species, known from Triticum aestivum (Poaceae) and Citrus maxima in South Africa (Gerlach 1977, Farr et al. 2023).
Fusarium magnum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855108. Fig. 48.
Fig. 48.
Fusarium magnum (ex-type CBS 151821). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–G. Sporodochia. H. Sporodochial conidiophore and conidiogenous cells. I. Aerial conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin magnus (“great”). In reference to the large-sized macroconidia.
Sporodochia abundant on the surface of carnation leaves, agar surface and aerial mycelium, pale orange, sienna to hazel coloured. Sporodochial conidiophores profusely branched laterally and verticillately, 28–100 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical, (9–)12.5–19(–23.5) × 3–5.5 µm, smooth, thin-walled, with a conspicuous apical collarette. Sporodochial conidia whip-like and robust, rarely falcate, thick-walled, strongly dorsiventrally curved or flexuous, apical cell strongly elongated and tapering; basal cell well-developed to elongated, foot-shaped, (0–)3–13(–17)-septate, (23–)78–157(–179) × (4–)4.5–6(–6.5) µm (av. 117.5 × 5.3 μm), mostly 5–9-septate, (43–)70–149(–179) × (4–) 4.5–6(–6.5) µm (av. 109.3 × 5.2 μm). Aerial conidiophores erect or prostrate on aerial and substrate mycelium, 11–166 µm tall, simple, reduced to single monophialides forming laterally on mycelium or borne on short stipes, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical, (8–)11.5–17(–23) × 3–5 µm, giving rise to whip-like conidia undistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 9.7–11.2 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface woolly to cottony, flesh, rosy vinaceous to coral, flat, aerial mycelium abundant on periphery, flesh coloured, margin regular, filiform to undulate; reverse rosy buff, brick to coral, vinaceous to dark brick at centre. On OA, woolly to cottony, sulphur to pure yellow, flat, aerial mycelium abundant on periphery, straw to sulphur yellow, margin regular and filiform; reverse straw.
Typus: South Africa, from soil, before 1 Dec. 1985, unknown collector (holotype designated here CBS H-25431, ex-type culture CBS 151821 = FRC R-7846 = MRC 3530 = NRRL 66927).
Additional material examined: South Africa, Eastern Cape Province, Transkei, from debris, before 12 Jan. 1985, unknown collector, culture FRC R-8136 = KOD 2014 = NRRL 64075.
Notes: Previously assigned to Fusarium sp. nov. 16 in Laraba et al. (2021), F. magnum shares a similar conidial morphology (large whip-like macroconidia with remarkably elongate apical cells) with F. cygneum, F. dolichosporum, F. longicolle, F. mastigosporum, and F. procumbens (see additional comments under F. cygneum). Both F. magnum and F. dolichosporum, however, are genetically distant to the remaining four species above, belonging to the Sambucinum clade of FSAMSC. Fusarium magnum is morphologically and phylogenetically closest to F. dolichosporum. It differs by the red pigmentation of its PDA colonies (luteous in the latter species), the absence of microconidia, and by producing a single macroconidial morphotype. This species is so far only known from soil, soil debris and Triticum sp. in South Africa (Laraba et al. 2023).
Fusarium mastigosporum Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855109. Fig. 49.
Fig. 49.
Fusarium mastigosporum (ex-type CBS 151825). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochium. F. Sporodochial conidiophores and conidiogenous cells. G–I. Aerial conidiophores and conidiogenous cells. J. Aerial macroconidia. K. Sporodochial macroconidia. Scale bars: F, H, J, K = 10 μm; I = 5 μm.
Etymology: From Greek mastigio (“whip”) and spora (“spore”). In reference to the macroconidia, shaped like a whip.
Sporodochia abundant and erumpent on the surface of carnation leaves, aerial mycelium, agar surface, and submerged on agar, luteous to bright orange coloured, turning pale luteous when old. Sporodochial conidiophores densely packed, branching copiously laterally and verticillately, 27–56 µm tall, bearing terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical, 9.5–20 × 2–4.5 µm, smooth- and thin-walled, with an inconspicuous apical collarette, often proliferating. Sporodochial conidia whip-like, strongly dorsiventrally curved or recurved, apical cell largely elongated; basal cell well-developed, foot-shaped, often elongated downwards, (4–)5–9-septate, (44.5–)100–155(–166) × (3.5–)4–5 µm (av. 127.7 × 4.3 μm), mostly 8-septate, (113–)128.5–154(–166) × (3.5–)4–5 µm (av. 141.4 × 4.3 μm). Aerial conidiophores erect or prostrate, sparsely branched, 15–22 µm tall, or reduced to monophialides forming laterally on aerial mycelium or submerged in agar. Aerial conidiogenous cells monophialidic, subcylindrical, 7.5–18 × 3–5 µm, smooth- and thin-walled, with or without apical collarette, or reduced to lateral 2.5–3 × 3–5 µm phialidic pegs. Aerial conidia, falcate, robust, moderately to strongly dorsiventrally curved, apical cell tapering with round apex, basal cell papillate to well-developed, foot-shaped, (1–)3–5(–6)-septate, (20.5–)33–50(–68.5) × (3.5–)4–6 µm (av. 41.4 × 4.7 μm), mostly 5-septate, (38.5–)40–50(–59) × 4–5(–6) µm (av. 44.5 × 4.8 μm). Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.9–8.8 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface felty to cottony, woolly at centre, rosy buff to peach with white to salmon patches and abundant scarlet to red excretions, sulphur yellow to pale luteous at centre, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse scarlet to dark brick. On OA, primrose to rosy buff at periphery; white, pure yellow to scarlet at centre, flat, cottony to woolly, with regular, filiform or undulate margin; reverse pure yellow to amber, brick at centre.
Typus: Australia, Darwin, from soil debris, before 7 Nov. 1989, unknown collector (holotype designated here CBS H-25433, ex-type culture CBS 151825 = FRC R-9077 = NRRL 66935).
Additional materials examined: Australia, from soil, before 28 Jan. 1985, unknown collector, CBS H-25398, culture CBS 151803 = NRRL 13368; unknown collection data, culture BBA 62247 = DSM 62247.
Notes: Fusarium mastigosporum is one of nine segregates of F. longipes recognized here (see additional notes under F. cygneum and F. longipes). Previously assigned to Fusarium longipes 1 in Laraba et al. (2021), it produces large whip-like macroconidia, common, but not exclusive to the Longipes clade of FSAMSC, and observed also in F. cygneum, F. dolichosporum, F. longicolle, F. magnum, and F. procumbens. Fusarium mastigosporum is distinguished by a combination of characters i.e., very long (av. 127.7 × 4.3 µm), robust, whip-like sporodochial conidia, and the presence of a secondary, falcate macroconidial morphotype on aerial conidiophores.
This species has been recorded on soil, other fungi (Daldinia sp.), and plant hosts in Poaceae (Sporobolus africanus and Triticum sp.), Rosaceae (Rubus sp.), and Solanaceae (Solanum tuberosum), in North America (US), Europe (Germany), and Oceania (Australia and New Zealand) (Farr et al. 2023).
Fusarium meridionale T. Aoki et al., Fungal Genet. Biol. 41: 618. 2004. Fig. 50.
Fig. 50.
Fusarium meridionale (ex-type CBS 110247). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G–I. Aerial conidiophores and conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: O’Donnell et al. (2004), Aoki et al (2012).
Typus: New Caledonia, from orange twig, before 10 Jun. 1998, unknown collector (holotype BPI 843474, ex-type culture CBS 110247 = FRC R-5329 = NRRL 28436).
Additional materials examined: Nepal, Kaski, from Zea mays, before 5 Jan. 1994, unknown collector, culture CBS 110260 = FRC R-9430 = NRRL 28721 = NRRL 38131; Lalitpur, from Zea mays, CBS 110248 = FRC R-9438 = NRRL 28723 = NRRL 38118. South Africa, Transkei, from soil, before 1 May 1977, unknown collector, culture CBS 110249 = FRC R-4080 = MRC 0856 = NRRL 29010.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Fusarium meridionale is practically indistinguishable morphologically from F. boothii, both species segregated from F. graminearum s. lat., and distinguished from all other species in that clade by forming slender (< 4.5 μm wide), curved and symmetric 5-septate macroconidia, with narrow apical beaks and wider in the median (Aoki et al. 2012), but see additional comments about morphological, biochemical and biogeographical differences under F. boothii.
Recorded substrates of F. meridionale include soil, insects, and plant hosts, mainly in Poaceae (Hordeum vulgare, Oryza sativa, Triticum aestivum, and Zea mays), Araceae (Amorphophallus konjac), Convolvulaceae (Ipomoea batatas), Fabaceae (Glycine max), Musaceae (Musa sp.), Myrtaceae (Eucalyptus grandis, and Psidium spp.), Rubiaceae (Coffea sp.), Rutaceae (Citrus sp.), and Solanaceae (Solanum lycopersicum) (O’Donnell et al. 2012, Wang et al. 2022, Farr et al. 2023).
Fusarium mesoamericanum T. Aoki et al., Fungal Genet. Biol. 41: 619. 2004. Fig. 51.
Fig. 51.
Fusarium mesoamericanum (ex-type CBS 415.86). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G. Sporodochial conidiophore and conidiogenous cells. H. Aerial conidiophore. I. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: O’Donnell et al. (2004), Aoki et al. (2012).
Typus: Honduras, from Musa sp. fruit, 4 Mar. 1986, K. Alsøe (holotype BPI 843476, ex-type culture CBS 415.86 = FRC R-8506 = IMI 309346 = NRRL 25797 = NRRL 38123).
Additional material examined: USA, Pennsylvania, from Cissus rhombifolia, before 1 May 1977, unknown collector, culture CBS 110252 = FRC R-4079 = NRRL 29148 = NRRL 38124.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Fusarium mesoamericanum is morphologically recognizable among segregates of F. graminearum s. lat. by producing 4–4.5 µm wide, straight, asymmetric, 5-septate conidia, which do not show an apical beak and are widest above the median (Aoki et. al. 2012).
Reported from Canada, Honduras and the USA, the currently known host range of F. measoamericanum [Musaceae (Musa sp.), Pinaceae (Pinus ponderosa), and Vitaceae (Cissus spp., and Parthenocissus tricuspidata)] differ from those of sibling species in F. graminearum s. lat., commonly isolated from Poaceae hosts (Aoki et al. 2012, Farr et al. 2023). In vitro pathogenicity of F. mesoamericanum towards wheat has been demonstrated (O’Donnell et al. 2000).
Fusarium minutum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855110. Fig. 52.
Fig. 52.
Fusarium minutum (ex-type CBS 151823). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–I. Conidiophores and conidiogenous cells. J. Microconidia. K. Chlamydospores. L. Macroconidia. Scale bars: E, H–L = 10 μm; F, G = 5 μm.
Etymology: From Latin minūtus (“diminished”). Referring to the small sized macroconidia.
Sporodochia not seen. Aerial conidiophores erect or prostrate on substrate and mycelium, or submerged in agar, sparsely branched, 2.5–30 µm tall, often reduced to monophialides forming laterally on aerial mycelium or submerged in agar. Aerial conidiogenous cells monophialidic, subcylindrical to subulate, 4–19 × 2.5–5.5 µm, or reduced to lateral phialidic pegs, 2.5–7 × 2.5–5.5 µm, with conspicuously flared apical collarette, giving rise to micro- and macroconidia. Microconidia ovoid, ellipsoidal to short falcate, 0–1-septate, (6–)8.5–13.5(–17) × 2.5–4 µm (av. 11 × 3.4 μm), often grouped in small false heads. Macroconidia falcate, strongly dorsiventrally curved and tapering, apical cell slightly elongated and pointy to somewhat hooked; basal cell papillate or well-developed foot-shaped, 1–5-septate, (9.5–)16–29.5(–42) × (2.5–)3.5–4.5(–5.5) µm (av. 22.7 × 4 μm), mostly 3-septate, (15.5–)19.5–28(–34) × 3–5 µm (av. 23.6 × 4.1 μm). Chlamydospores globose to subglobose, smooth- and thick-walled, hyaline to subhyaline, 8.5–13.5 µm diam intercalary on hyphae and conidia or borne laterally on short stipes.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.5–9.3 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface at first membranous, becoming felty, scarlet to rosy buff, sulphur yellow at periphery, flat, aerial mycelium abundant, glabrous and moisty at centre, margin regular and filiform; reverse scarlet to coral, sulphur yellow at periphery. On OA, white, straw to sulphur yellow, flat, membranous to velvety, margin regular and filiform; reverse sulphur yellow to honey.
Typus: Italy, unknown substrate, before 1 May 1988, P. Gambogi (holotype designated here CBS H-25429, ex-type culture CBS 151823 = FRC R-8838 = NRRL 66931).
Additional material examined: Tunisia, Mahdes, from olive grove soil, before 1 Jan. 1986, unknown collector, culture CBS 151822 = FRC R-8165 = NRRL 66928.
Notes: Fusarium minutum was previously informally assigned to Fusarium sp. nov. 25 in Laraba et al. (2021), and it is currently only known from soil and an undetermined substrate in Italy and Tunisia.
Fusarium minutum exhibits macroconidia that are quite short for its width (averaging 22.7 × 4 μm) being the shortest macroconidia measured in FSAMSC. There is a slight overlap in conidial length between F. minutum and several other short-spored species, including F. brachygibbosum, F. poae, F. sambucinum, F. subtropicale, among others. However, all the species above form conidia averaging > 25 μm long (34.3, 26.2, 28, and 27.9 μm long, respectively). Fusarium minutum can be differentiated from its closest phylogenetic relatives F. cuspidatum and F. sagittatum by its smaller, and more prominently curved conidia. In contrast, both F. cuspidatum and F. sagittatum form macroconidia with almost straight ventral lines, while F. cuspidatum also lacks microconidia.
Fusarium mucronatum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855111. Fig. 53.
Fig. 53.
Fusarium mucronatum (ex-type CBS 151826). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiophore and conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin mūcrōnātus (“pointy”). In reference to the pointy macroconidial apical cells.
Sporodochia abundant on the surface of carnation leaves, scarce on agar surface, buff, honey to hazel coloured. Sporodochial conidiophores branching laterally, 27–75 µm tall, bearing terminal groups or solitary lateral monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical to subulate, 9–20 × 3.5–5.5 µm, smooth- and thin-walled, with a minute apical collarette. Sporodochial conidia falcate, gently to strongly dorsiventrally curved, tapering abruptly towards the apex, with almost straight ventral line, often conspicuously widest at or below the median, apical cell moderately elongate and hooked; basal cell often well-developed foot-shaped, 1–5-septate, (36–)40–51(–64.5) × 5.5–7 µm (av. 45.3 × 6.2 μm), mostly 5-septate, (36.5–)40–51(–65) × 5.7–7 µm (av. 45.7 × 6.2 μm). Aerial conidiophores rare, mostly reduced to solitary phialides borne laterally on hyphae, less often sparingly laterally branched, bearing terminal monophialides, 18–37 µm tall, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, ampulliform, subcylindrical to subulate, 10.5–17 × 3.5–5 µm, giving rise to falcate conidia indistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 10.4–11.7 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface felty to woolly, rosy vinaceous to red, rust and moist at the centre, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse luteous to red. On OA, pure yellow to amber becoming white to rosy vinaceous, flat, at first membranous becoming velvety to cottony, with regular and undulate margin; reverse pure yellow to luteous.
Typus: Botswana, Pandamatenga, from Pennisetum glaucum, before 12 Jul. 1990, unknown collector (holotype designated here CBS H-25435, ex-type culture CBS 151826 = FRC R-9135 = NRRL 66937).
Notes: Fusarium mucronatum is here described from a strain previously assigned to Fusarium sp. nov. 30 in Laraba et al. (2021). This species differs from all close phylogenetic relatives in the Brachygibbosum clade (F. brachypes, F. dimorphosporum, F. hamatum, F. platysporum, and F. transvaalense) by lacking microconidia and chlamydospores, while producing only a single macroconidial morphotype. Macroconidia of F. mucronatum are similar in size to those of F. carinatum, a distant species in the Longipes clade; however, the two species show obvious differences in conidial shape, with those of F. mucronatum showing a more defined curvature, with moderately elongated apical cells and widest below the median. In contrast, sporodochial conidia of F. carinatum are almost straight, without elongated apical cells, and are often widest above the median.
Fusarium musarum Logrieco & Marasas, Mycologia 90: 510. 1998. Fig. 54.
Fig. 54.
Fusarium musarum (ex-type CBS 151808). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–H. Conidiophores and conidiogenous cells. I. Mesoconidium undergoing microcyclic conidiogenesis. J. Microconidia. K. Mesoconidia. Scale bars: E, F = 20 μm; G–K = 10 μm.
Description and illustrations: Marasas et al. (1998), Leslie & Summerell (2006).
Typus: Panama, from Musa sapientum fruit, 1991, A. Logrieco (holotype BPI 802928, ex-type culture CBS 151808 = DAOM 225261 = FRC R-9400 = NRRL 28507 = MRC 6240).
Additional materials examined: Costa Rica, from Musa sp. fruit, before 25 Apr. 2016, unknown collector, cultures BBA 68458 = DSM 116552, BBA 68461 = DSM 116554, BBA 68462 = DSM 116555.
Notes: Fusarium musarum is a Central American species, originally described as a segregate of F. camptoceras (FIESC), together with F. nelsonii (F. chlamydosporum SC). The three species were assigned to section Arthrosporiella, and are characterized by the production of mesoconidia from polyblastic conidiogenous cells (Marasas et al. 1998), but are currently known to be far related genetically, each species allocated in distant species complexes; F. musarum resolved as the basal-most species in the Sambucinum clade, however, with significant morphological differences. Fusarium musarum is uncommon within FSAMSC by producing polyblastic aerial conidiogenous cells, while lacking sporodochia and true macroconidia. Additionally, microcyclic conidiogenesis was frequently observed here in the ex-type strain.
Fusarium nepalense T. Aoki, et al., Fungal Genet. Biol. 48: 1105. 2011.
Descriptions and illustrations: Sarver et al. (2011), Aoki et al. (2012).
Typus: Nepal, Lamjung, from Oryza sativa seed, 1997, A. Desjardins (holotype BPI 881006, ex-type culture CBS 127503 = NRRL 54222).
Additional materials examined: Nepal, Lamjung, from Oryza sativa, 1997, unknown collector, cultures CBS 127669 = NRRL 54220, CBS 127943 = NRRL 54221.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum. Strains of F. nepalense studied here, including the ex-type strain, are degenerated and although they still produce typical pigments on culture, we were unable to induce sporulation; hence, listed morphological features are based entirely on published descriptions. Fusarium nepalense is characterized by 4.5–5 μm wide, gradually curved, asymmetric, 5-septate conidia, with narrow apical beaks, being wider above the median (Sarver et al. 2011); this combination of characters differentiates F. nepalense from all formally described segregates of F. graminearum s. lat., yet is not enough to distinguish this species from known but currently undescribed lineages (Fusarium sp. NRRL 34461, and the “Gulf Coast population” of F. graminearum s. lat.; Aoki et al. 2012).
Fusarium nepalense is known from Musaceae (Musa nana), Poaceae (Oryza sativa), and Theaceae (Camellia sinensis), from China and Nepal (Aoki et al. 2012, Wang et al. 2022, Farr et al. 2023).
Fusarium nodosum L. Lombard & Crous, Fungal Syst. Evol. 4: 193. 2019.
Description and illustrations: Lombard et al. (2019).
Typus: Portugal, Lisbon, from Arachis hypogaea seed, 1961, unknown collector, dep. C.M. Baeta Neves (holotype CBS H-24018, ex-type culture CBS 201.63).
Notes: Fusarium nodosum was recently described from a set of isolates originally identified as F. chlamydosporum (F. chlamydosporum SC), and reidentified as members of the FSAMSC (Lombard et al. 2019). This species is the closest phylogenetic relative to F. goolgardi (Laurence et al. 2016, Han et al. 2023), from which it differs by forming red PDA colonies (vs orange in F. goolgardi), while lacking sympodially proliferating and branching aerial monophialides, in contrast producing aerial polyphialides (Lombard et al. 2019).
Fusarium nodosum is chiefly known from Europe (France, Portugal, and Türkiye) and Asia (Iran), isolated from Fabaceae (Arachis hypogaea), Poaceae (Arundo donax, Triticum spp.), and Solanaceae (Solanum lycopersicum) (Lombard et al. 2019, Farr et al. 2023).
Fusarium palustre W.H. Elmer & Marra, Stud. Mycol. 98: 143. 2021. Fig. 55.
Fig. 55.
Fusarium palustre (ex-type CBS 126796). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochium and chlamydospores. F, G. Chlamydospores. H. Conidiogenous cell and microconidia. I–K. Conidiogenous cells. L. Macroconidia. Scale bars = 10 μm.
Synonym: Fusarium palustre W.H. Elmer & Marra, Mycologia 103: 815. 2011, nom. inval., Art. 40.7.
Description and illustration: Elmer & Marra (2011).
Typus: USA, Connecticut, Madison, Hammonasset Beach State Park, from Spartina alterniflora, Jun. 2006, W.H. Elmer [holotype CBS 126796 (preserved as metabolically inactive culture), ex-type culture CBS 126796 = NRRL 54056].
Additional materials examined: Unknown, unknown collection data, culture CBS 120613. USA, from Spartina sp., rhizosphere, before 1 Jan. 2003, W. Elmer, culture NRRL 43289.
Notes: This species was originally invalidly published because of an incorrect citation of the holotype (Elmer & Marra 2011). When validating the species name (Crous et al. 2021), an erroneus specimen number was cited for the holotype (CBS 126795 instead of CBS 126796) which is incorrect for the type collection data. This is corrected here according to Art. 9.2.
Fusarium palustre resolved as the basal-most species in the Sporotrichioides B clade, differing from its closest phylogenetic siblings (F. chaquense, F. leptum, and F. parabolicum) by its orange, and faster growing (av. 13.9 mm/d) PDA colonies (vs red or yellow PDA colonies, averaging < 10 mm/d in the other three species). The ex-type culture of F. palustre seems deteriorated, and sporulates poorly. Microconidia were not present in the protologue of the species; however, aseptate, ovoidal to ellipsoidal microconidia measuring (7.5–)8–15(–16.5) × 3–4.5 were produced (Table S1, Fig. 55), although rarely, by the ex-type.
Fusarium palustre is pathogenic to Spartina alterniflora and has been isolated from Spartina plants and associated soil in the USA (Elmer & Marra 2011).
Fusarium parabolicum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855112. Fig. 56.
Fig. 56.
Fusarium parabolicum (ex-type CBS 151814). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–G. Sporodochia. H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiophore and conidiogenous cells. J. Chlamydospores. K. Microconidia. L. Macroconidia. Scale bars = 10 μm.
Etymology: From Greek parabolikós (“in the shape of a parabola or paraboloid”). Referring to the dorsal curvature of its macroconidia.
Sporodochia rarely formed on the surface of carnation leaves and agar surface, pale luteous, orange to ochreous coloured. Sporodochial conidiophores sparingly branching irregularly and laterally, 26–62 µm tall, bearing solitary lateral and terminal verticils of monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical, (9–)11–15(–18) × 2.5–5 µm, smooth-and thin-walled, apical collarette inconspicuous to absent. Sporodochial conidia falcate, moderately to strongly dorsiventrally curved and tapering towards the apical end, widest at or below the median, apical cell elongated and tapering to a blunt or slightly hooked, somewhat spathulate end; basal cell well-developed foot-shaped, 1–5(–6)-septate, (19.5–)36–54(–62) × (3–)4–5 µm (av. 44.9 × 4.1 μm), mostly 3-septate, (31.5–)33–47.5(–57.5) × 3–5 µm (av. 40.1 × 4.1 μm). Aerial conidiophores erect or prostrate on substrate mycelium, reduced to solitary monophialides borne laterally on hyphae, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical, 5–15 × 2.5–5 µm, giving rise to macroconidia indistinguishable from sporodochial macroconidia, and rarely to microconidia. Microconidia ellipsoidal, 0–1-septate, (8.5–)10–16(–17.5) × 3.5–4.5 µm (av. 13 × 3.9 μm). Chlamydospores subglobose, smooth- and thick-walled, 8.5–15 µm diam, subhyaline, solitary, in pairs or short chains, often borne laterally on short stipes.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 9–10.2 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface at first pale luteous and membranous, slimy at centre, becoming cottony to woolly, white to pale buff, flat, aerial mycelium abundant in irregular patches, margin regular, filiform to undulate; reverse sulphur to pure yellow. On OA, ochreous to apricot, flat, membranous to woolly, with irregular patches of white to pale luteous aerial mycelium, margin regular and undulate; reverse sulphur yellow to ochreous.
Typus: USA, Missouri, from Festuca arundinacea, before 24 May 1967, unknown collector (holotype designated here CBS H-25410, ex-type culture CBS 151814 = ATCC 36781 = FRC R-5319 = MRC 1783 = NRRL A-15494 = NRRL 6227).
Additional material examined: USA, Kansas, Konza Prairie Research Natural Area, Long-term Ecological Research site (LTER), near Manhattan, from soil, 1986, unknown collector, culture CBS 128810 = RMF 8363.
Notes: Formerly Fusarium sp. nov. 3 in Laraba et al. (2021), Fusarium parabolicum is known from soil, and Poaceae hosts (Avena sativa, Festuca arundinacea and Zea mays) (Laraba et al. 2021, Farr et al. 2023). This species belongs to the Sporotrichioides B clade, and it is distinguished by its macroconidial shape, which contrary to its closest siblings (F. armeniacum, F. chaquense, F. leptum, and F. palustre, exhibits a regular curvature, with less pronounced tapering and shorter apical cells. Additionally, it differs from F. armeniacum, F. chaquense, and F. palustre by lacking an extra apical curvature, and by forming yellow PDA colonies (vs red PDA colonies in F. armeniacum and F. chaquense, and orange in F. palustre). Fusarium leptum produces similar yellow colonies, but F. parabolicum differs by its less conspicuous conidial foot cells and by producing microconidia.
Fusarium platysporum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855113. Fig. 57.
Fig. 57.
Fusarium platysporum (ex-type CBS 151816). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G. Sporodochial conidiophores and conidiogenous cells. H. Aerial conidiophore and conidiogenous cell. I. Microconidia. J. Chlamydospores. K. Macroconidia. Scale bars: G, H, J, K = 10 μm; I = 5 μm.
Etymology: Greek platýs (“wide”) and spora (“seed”). Referring to the wide sporodochial conidia.
Sporodochia erumpent and abundant on the surface of carnation leaves, agar surface and aerial mycelium, primrose, sulphur to pure yellow coloured. Sporodochial conidiophores branching irregularly laterally and verticillately, 35–88 µm tall, bearing lateral and terminal verticils of monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical to subulate, 9–19.5 × 2.5–5 µm, smooth, thin-walled, with a short, flared apical collarette. Sporodochial conidia falcate, gently to moderately dorsiventrally curved with an almost straight ventral line, widest to somewhat swollen above the median and tapering abruptly upwards, apical cell tapering, conical to barely elongated, pointy to hooked; basal cell papillate to well-developed, foot-shaped, (2–)3–7(–9)-septate, (24–)38.5–57.5(–87) × (4–)5–7 µm (av. 48 × 6 μm), mostly 5-septate, (34.5–)39–53(–87) × (4.5–)5.5–6.5(–7) µm (av. 45.8 × 6.1 μm). Aerial conidiophores erect or prostrate on substrate and aerial mycelium, unbranched, 13–18 µm tall, reduced to monophialides forming laterally on aerial mycelium or submerged in agar, with or without a short stipe. Aerial conidiogenous cells monophialidic, ampulliform to subulate, 4.5–17.5 × 1.5–5.5 µm, giving rise to macroconidia indistinguishable from sporodochial conidia, and more rarely to microconidia. Microconidia ovoid, ellipsoidal to somewhat reniform, 0(–1)-septate, (7–)8–13.5(–16.5) × 2.5–3.5 µm (av. 52.1 × 5.6 μm). Chlamydospores subglobose to globose, thick- and rough-walled, (6.3–)8–12(–14.5) µm diam, hyaline at first, becoming pale to dark golden brown.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.6–11.3 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface felty to woolly, rosy vinaceous to red, with pale luteous to pure yellow patches or mycelial ropes, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse coral to red, chestnut to brown vinaceous at centre. On OA, orange to scarlet, with white, pure yellow or red patches, flat, felty to cottony, with regular and undulate margin; reverse honey.
Typus: Ghana, near Tamale, from Glycine max root, before 8 Oct. 2019, unknown collector (holotype designated here CBS H-25413, ex-type culture CBS 151816 = KOD 1723 = NRRL 66920).
Additional materials examined: Argentina, from Triticum sp., before 4 Aug. 1998, Lopez, CBS H-25399, culture CBS 151810 = FRC R-8755 = NRRL 28725. Brazil, from Triticum sp. sub-crown internode, before 1 Jan. 1979, unknown collector, culture NRRL 66922 = FRC R-4932
Notes: Originally referred to as Fusarium sp. nov. 29 in Laraba et al. (2021), F. platysporum has similar sporodochial conidia to several other species in FSAMSC, i.e., F. cuspidatum, F. dimorphosporum, F. hamatum, F. pratense, and F. tropicale. The presence of microconidia differentiates F. platysporum from F. cuspidatum, F. hamatum, and F. tropicale, and the presence of chlamydospores and red PDA colonies distinguishes F. platysporum from F. pratense. In addition, by forming a single type of macroconidium, F. platysporum can be distinguished from F. dimorphosporum.
Fusarium platysporum has been recorded from soil, soil debris, and plant hosts in Fabaceae (Glycine max), Poaceae (Sorghum sp., Triticum sp.) in Africa (Ghana and Nigeria), North America (USA) and South America (Argentina and Brazil) (Farr et al. 2023).
Fusarium poae (Peck) Wollenw., 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 Bubák, Centralbl. Bakteriol. Parasitenk., 2. Abth. 31: 497. 1911.
Descriptions and illustrations: Wollenweber & Reinking (1935), Booth (1971), Gerlach & Nirenberg (1982), Nelson et al. (1983), and Leslie & Summerell (2006), and Domsch et al. (2007).
Typus: USA, New York, Ontario, Geneva, from Poa pratensis sheaths and culms, Jun. 1902, F.C. Stewart (holotype NYSf2393); North Dakota, Minot, from Hordeum vulgare infected kernel, before 21 Nov. 1997, unknown collector [epitype NRRL 26941 (preserved as metabolically inactive culture), designated by Crous et al. (2021), ex-epitype culture CBS 151948 = NRRL 26941 = FRC T-962].
Additional materials examined: Belgium, Tubize, from Triticum aestivum seed, 2005, A. Chandelier, culture CBS 128536. Canada, from overwintered Triticum sp., before 18 Mar. 1988, unknown collector, dep. P.E. Nelson, culture CBS 151947 = NRRL 13714 = FRC T-503 = MRC 2181. Denmark, from Hordeum vulgare straw, 30 Sep. 1985, O. Filtenborg, culture CBS 623.87 = NRRL 36524. Germany, Berlin, from Anthoxanthum odoratum inflorescence, 1965, unknown collector, dep. E. Seemüller, culture CBS 446.67 = BBA 10426 = FRC T-982 = NRRL 25799. Norway, Aas, from Avena sativa, unknown collection date and collector, dep. L. Sundheim, culture CBS 174.96, from Hordeum vulgare, unknown collection date and collector, dep. L. Sundheim, cultures CBS 175.96, CBS 176.96; Oslo, from Triticum sp., unknown collection date and collector, dep. L. Sundheim, culture CBS 180.96. Poland, from Triticum sp., unknown collection date and collector, dep. L. Sundheim, culture CBS 186.96; Zulawy region, from Triticum aestivum, before 20 Feb. 2004, unknown collector, dep. T. Kulik, culture CBS 115696. Switzerland, Canton Bern, Belpberg, Triticum sp. kernels, 2003, I. Bänziger, culture CBS 121297 = NRRL 53735.
Notes: Fusarium poae was recently epitypified by Crous et al. (2021). This is an easily recognizable species due to its abundant globose microconidia, and seldom present sickle-shaped, short macroconidia, which together with the production of fast growing (av. 9.7 mm/d), red PDA colonies distinguish this species from the morphologically similar, albeit phylogenetically distant species F. langsethiae and F. sibiricum (av. growth rates < 6 mm/d, and yellow colonies).
Fusarium poae is a cosmopolitan species and a weak plant pathogen (Leslie & Summerell 2006), present in soil and aqueous environments (Domsch et al. 2007), animal feed and even human specimens (Leslie & Summerell 2006). Known plant hosts include over 80 species distributed in an extensive list of genera, mostly in Poaceae (Anthoxanthum, Aristida, Austrostipa, Avena, Cynodon, Dactylis, Elymus, Festuca, Hordeum, Muhlenbergia, Oryza, Panicum, Pennisetum, Phleum, Phragmites, Poa, Saccharum, Secale, Setaria, Sitanion, Triticum, and Zea), but also in Amaranthaceae (Celosia), Arecaceae (Phoenix), Asteraceae (Helianthus), Brassicaceae (Brassica), Caryophyllaceae (Dianthus), Cucurbitaceae (Cucumis, Cucurbita), Ericaceae (Calluna), Fabaceae (Glycine, Lupinus, Macroptilium, Phaseolus, Pisum, Trifolium, Vicia), Liliaceae (Tulipa), Linaceae (Linum), Malvaceae (Gossypium), Moraceae (Morus), Rosaceae (Prunus), Solanaceae (Solanum), Theaceae (Camellia), and Vitaceae (Vitis); and has been recorded in Africa (South Africa), Asia (Bahrain, China, India, Iraq, Pakistan, Turkey, and Uzbekistan), North and South America (Argentina, Brazil, Canada, Paraguay, and USA), Europe (Austria, Belgium, Denmark, England, Estonia, Finland, France, Germany, Italy, Norway, Poland, Russia, Spain, Sweden, Switzerland, Ukraine, United Kingdom, and Yugoslavia), and Oceania (Australia and New Zealand) (Domsch et al. 2007, Wang et al. 2022, Farr et al. 2023).
Fusarium praegraminearum Gräfenhan & O’Donnell, Mycologia 108: 1232. 2016. Fig. 58.
Fig. 58.
Fusarium praegraminearum (ex-type CBS 141369). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G–I. Aerial conidiophores and conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: Gräfenhan et al. (2016).
Typus: New Zealand, North Island, Levin, near Wellington, from litter in Zea mays field, May 1984, H. Hussein (holotype PDD 47563, ex-type culture CBS 141369 = ICMP 8996 = NRRL 39664 = PDD 47563).
Notes: A rare species known from New Zealand, F. praegraminearum was shown to induce moderate head blight symptoms on wheat under in vitro conditions (Gräfenhan et al. 2016).
Morphologically, F. praegraminearum has been compared to F. dactylidis, F. graminearum, F. pseudograminearum (all three in the Graminearum clade) and F. acuminatum (FTSC), being distinguished by a combination of colony characteristics and its conidial shape. For a detailed morphological comparison see Gräfenhan et al. (2016).
Fusarium pratense Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855114. Fig. 59.
Fig. 59.
Fusarium pratense (ex-type CBS 151956). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G–J. Aerial conidiophores and conidiogenous cells. K. Microconidia. L. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin prātēnse (“pasture-dwelling”). In reference to the origin of the type, from grassland.
Sporodochia abundant on the surface of carnation leaves and agar surface, saffron to luteous coloured and transparent. Sporodochial conidiophores sparingly branched laterally, 20–75 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, doliiform, ampulliform to subcylindrical, 9–18 × 2.5–5 µm, smooth- and thin-walled, lacking apical collarette. Sporodochial conidia falcate, almost straight or moderately dorsiventrally curved, with an almost straight ventral line and tapering towards both ends, with blunt and tapering apical cell; basal cell papillate or poorly- to well-developed foot-shaped, (0–)1–5(–6)-septate, (19.5–)36–54(–62) × 3–5 µm (av. 44.9 × 4.1 μm), mostly 5-septate, (39.5–)43.5–57(–62) × 3.5–5 µm (av. 50.4 × 4.3 μm). Aerial conidiophores erect or prostrate on substrate mycelium, reduced to monophialides forming laterally on aerial mycelium or submerged in agar. Aerial conidiogenous cells monophialidic, doliiform, ampulliform to subcylindrical, 8–12.5 × 3.5–5 µm, smooth- and thin-walled, giving rise to falcate conidia indistinguishable from sporodochial conidia and rarely to 0–1-septate, ellipsoidal to falcate microconidia, (8.5–)10–15(–17) × 3.5–4.5 µm. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.2–8.6 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface at first membranous becoming velvety and moisty, sulphur yellow to straw, flat, aerial mycelium scant, margin regular and filiform; reverse sulphur yellow to straw. On OA, sulphur yellow to straw, flat, membranous to slimy, with regular and undulate margin; reverse sulphur yellow to pure yellow.
Typus: Australia, Northern Territory, from wet grassland soil, before 1 Nov. 1988, unknown collector (holotype designated here CBS H-25426, ex-type culture CBS 151956 = FRC R-8926 = NRRL 66932).
Additional material examined: New Guinea, from Saccharum officinarum leaf, unknown collection date and collector, culture IMI 316488.
Notes: Formerly the Fusarium sp. nov. 19 in Laraba et al. (2021), F. pratense differs from all its closest phylogenetic relatives in the Longipes clade of FSAMSC, except for F. carinatum, by forming yellow PDA colonies and forming falcate conidia with pointy apices instead of elongated, whip-like conidia. Fusarium pratense differs from F. carinatum by producing a single macroconidial morphotype and ellipsoidal to falcate, but not clavate microconidia. Other species with similar macroconidial morphology are allocated in the distant Brachygibbosum clade (F. cuspidatum, F. dimorphosporum, F. hamatum, and F. platysporum), although they all differ by producing red pigments on PDA, in addition to the lack of microconidia (F. cuspidatum and F. hamatum), producing ovate to reniform, but not falcate microconidia (F. platysporum); or exhibiting two distinct macroconidial morphologies (F. dimorphosporum).
Fusarium procumbens Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855115. Fig. 60.
Fig. 60.
Fusarium procumbens (ex-type CBS 121863). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Aerial conidiophores and conidiogenous cells. I. Sporodochial conidiophore and conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin prōcumbēns (“falling forwards”). In reference to the curvature of the macroconidia.
Sporodochia abundant on the surface of carnation leaves and agar surface, erumpent, pale buff, bright orange to luteous coloured. Sporodochial conidiophores sparingly branching laterally, 35–115 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, subulate to subcylindrical, 9–19.5 × 2.5–5 µm, smooth, thin-walled, lacking apical collarette. Sporodochial conidia whip-like, rarely falcate, strongly dorsiventrally curved to recurved, apical cell strongly elongate, tapering to a blunt end, rarely slightly to moderately elongated and somewhat hooked; basal cell well-developed foot-shaped and elongated, (2–)3–8-septate, (34–)81–131.5(–149) × (3–)4–4.5(–6) µm (av. 106.1 × 4.3 μm), mostly 6-septate, (79–)100.5–125(–143) × (3.5–)4–5 µm (av. 112.8 × 4.4 μm). Aerial conidiophores erect or prostrate on substrate mycelium, 30–145 µm tall, often as solitary lateral phialides on aerial mycelium, later confluent, rarely sparsely branched bearing terminal and lateral phialides. Aerial conidiogenous cells monophialidic, subulate to subcylindrical, 9.5–21.5 × 2.5–4.5 µm, giving rise to whip-like conidia indistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 3.9–5.1 mm/d, 27.3–35.5 mm diam after 7 d. Surface powdery to velvety, coral to red with peach to flesh patches forming concentric rings, flat, aerial mycelium abundant and short, margin regular, filiform to undulate; reverse coral, red to bay at centre. On OA, amber to luteous with peach to red concentric rings, pale luteous at periphery, flat, powdery to cottony, membranous to slimy at centre, margin regular, filiform to undulate; reverse luteous to amber.
Typus: Netherlands, unknown substrate, before 1 Dec. 2004, W.F.O. Marasas (holotype designated here CBS H-25430, ex-type culture CBS 121863 = FRC R-9983 = KSU 11431 = MRC 8430 = NRRL 53749).
Notes: One of six species (together with F. cygneum, F. dolichosporum, F. longicolle, F. magnum, and F. mastigosporum) producing large (average length close or above 100 μm), whip-like macroconidia with remarkably elongate apical cells (see additional comments under F. cygneum). Its slow growing colonies on PDA (av. < 4.6 mm/d) differentiates F. procumbens, together with F cygneum and F. longicolle, from the remaining morphologically similar species. There is a significant conidial size overlap between F. cygneum, F. longicolle and F. procumbens, however, with an average length of 106.1 µm, and up to 8-septate, macroconidia of F. procumbens tend to have an intermediate size between the longer macroconidia of F. cygneum (av. length 128.7 µm, and up to 11-septate) and the shorter macroconidia of F. longicolle (av. 78 µm long, and up to 9-septate).
Fusarium pseudograminearum O’Donnell & T. Aoki, Mycologia 91: 604. 1999.
Descriptions and illustrations: Aoki & O’Donnell (1999a, b).
Typus: Australia, New South Wales, Darling Downs, 43 km south of Young, from Hordeum vulgare crown, 1980, L.W. Burgess (holotype BPI 746087, ex-type culture CBS 109956 = FRC R-5291 = NRRL 28062).
Additional materials examined: Australia, Darling Downs, from Hordeum vulgare, before 25 Apr. 2016, unknown collector, culture BBA 71458 = DSM 116903; New South Wales, Breeza, from pasture soil, before 25 Apr. 2016, unknown collector, culture BBA 71460 = DSM 116904; Western Australia, Cunderdin, from Medicago polymorpha dead fallen pod, Feb. 1989, unknown collector, culture IMI 350956.
Notes: Fusarium pseudograminearum, formerly known as F. graminearum Group 1 was the first segregate population of F. graminearum s. lat. to be formally described as a different taxon (see additional comments under F. graminearum) (Aoki & O’Donnell 1999a, b). Phylogenetically, it resolves as the nearest relative to F. dactylidis, from which it differs by producing chlamydospores and larger multiseptate conidia.
Fusarium pseudograminearum is a soil-borne fungus and causes Fusarium crown rot and occasionally Fusarium head blight on small grain cereals, and has been recently evaluated as a potential quarantine pest in the EU (EFSA Panel on Plant Health et al. 2022). Although the outcome of the pest categorization indicated that it is a potential Union quarantine pest, it has not been included in the EU Commission Implementing Regulation 2019/2072 yet. It is known to affect several hosts in Fabaceae (Glycine max, and Medicago spp.), Poaceae (Aegilops tauschii, Ammophila sp., Austrostipa aristiglumis, Avena sp., Dactylis glomerata, Hordeum spp., Lolium sp., Panicum virgatum, Triticum spp., and Zea mays), and Rosaceae (Malus domestica); and experimentally proven to also infect Brassicaceae (Brassica napus), Fabaceae (Cicer arietinum), Poaceae (Oryza sativa, Secale cereale, Sorghum sp., and Triticosecale rimpaui) (Akinsanmi et al. 2007, EFSA Panel on Plant Health et al. 2022, Farr et al. 2023). This is a globally distributed species reported from Asia (Azerbaijan, China, Iran, Malaysia, Syria, Iraq, and Türkiye), Africa (Algeria, Morocco, South Africa, and Tunisia), Europe (Italy, Netherlands, and Spain), North and South America (Argentina, Canada and USA), and Oceania (Australia, New Zealand) (EFSA Panel on Plant Health et al. 2022, Farr et al. 2023).
Fusarium pseudolongipes Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855116. Fig. 61.
Fig. 61.
Fusarium pseudolongipes (ex-type CBS 131380). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Microconidia. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Greek pseudo- (“false, fake”). In reference to the species morphological similarity to Fusarium longipes.
Sporodochia abundant on the surface of carnation leaves, aerial mycelium and agar surface, primrose to bright luteous coloured. Sporodochial conidiophores branching laterally and verticillately, 27–46.5(–53) µm tall, bearing terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical to doliiform, 7.5–11 × 13.5–17 µm, smooth, thin-walled, apical collarettes inconspicuous or absent. Sporodochial conidia falcate, dorsiventrally curved, widest above the median, with an elongated, whip-like apical cell; basal cell elongate, foot-shaped, (1–)2–5-septate, (20–)39.5–61(–71) × (3.5–)4.5–5(–6) µm (av. 50.4 × 4.9 μm), mostly 5-septate, (42.5–)48–61(–71) × 4–6 µm (av. 54.6 × 5 μm). Aerial conidiophores reduced to single lateral monophialides quickly converging into sporodochia on aerial mycelium or emerging as solitary phialides from the agar surface. Aerial conidiogenous cells monophialidic, identical to sporodochial phialides, giving rise to falcate conidia indistinguishable from sporodochial conidia and rarely to 0–1(–2)-septate, (9.5–)11–16.5(–17) × 3.5–4.5 µm, obovoid to subcylindrical microconidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 5.5–6.5 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface cottony to woolly, peach to coral with amber to luteous patches, pale peach at periphery, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse scarlet to rust, umber at centre. On OA, pure yellow to amber with coral to flesh concentric rings, flat, glabrous at centre, with regular and filiform to undulate margin; reverse amber to pure yellow.
Typus: Australia, Northern Territories, Roper River area, from Oryza australiensis stem, Apr. 2009, unknown collector, dep. P. Harvey (holotype designated here CBS H-25414, ex-type culture CBS 131380).
Additional material examined: Australia, Northern Territories, Roper River area, from Oryza australiensis stem, Apr. 2009, unknown collector, dep. P. Harvey, CBS H-25403, culture CBS 131379.
Notes: A novel lineage identified here, so far only known from Australia, F. pseudolongipes is morphologically very similar to F. longipes. Both species differ significantly in their conidial shape and dimensions from the long-spored species in the Longipes clade (F. cygneum, F. longicolle, F. mastigosporum, and F. procumbens). The whip-like conidia differentiate F. longipes and F. pseudolongipes from close relatives forming falcate macroconidia (i.e., F. carinatum, F. pratense, and F. tropicale), while red PDA colonies distinguish both species from F. vermicularioides. Additionally, F. pseudolongipes differs from F. longipes by forming a single macroconidial morphotype and by its slower growth rate on PDA (av. 6 mm/d vs 13 mm/d in F. longipes), requiring also longer incubation times to achieve sporulation with sporodochia requiring up to 3 wk to be formed.
Fusarium robustum Gerlach, Phytopathol. Z. 88: 36. 1977. Fig. 62.
Fig. 62.
Fusarium robustum (ex-isotype CBS 637.76). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I. Aerial conidiophore. J. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: Gerlach & Nirenberg (1982), Nelson et al. (1983).
Typus: Argentina, from Araucaria angustifolia rotten stem, Mar. 1973, W. Radtke (holotype in B, isotype CBS H-629, ex-isotype culture CBS 637.76 = BBA 63667 = FRC R-5821 = IMI 322102 = NRRL 13392).
Notes: A rarely reported species, F. robustum is known from Argentina and Germany on Araucariaceae (Araucaria angustifolia), Poaceae (Triticum sp.) and from soil. This species has been compared morphologically with the genetically distant species F. cerealis and F. graminearum (Gerlach & Nirenberg 1982, Nelson et al. 1983); however, averaging 79.3 µm long, conidia of F. robustum are significantly longer than those of the species mentioned above, while they also differ in shape, being straighter, and more robust than in F. cerealis and F. graminearum, and tend to present a hooked apical cell and conspicuously well-developed foot cells.
Fusarium sagittatum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855117. Fig. 63.
Fig. 63.
Fusarium sagittatum (ex-type CBS 151958). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–G. Conidiophores and conidiogenous cells. H. Chlamydospores. I. Microconidia. J. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin sagittātus (“like a spear” or (“like an arrow”). In reference to the macroconidial shape.
Sporodochia not seen. Aerial conidiophores erect and prostrate, abundant on agar surface and aerial mycelium, reduced to solitary monophialides borne laterally on hyphae, commonly grouping in palisades, proliferating percurrently or laterally from below the conidiogenous opening. Conidiogenous cells monophialidic, subcylindrical to ampulliform, smooth- and thin-walled, 8–19 × 3.5–6.5 µm, or often reduced to a short phialidic peg, 1.5–4.5 × 1.5–3 µm, with conspicuous flared apical collarette, giving rise to micro- and macroconidia. Microconidia ellipsoidal to short falcate and slightly curved, 0–1-septate, (12.5–)13–18.5(–21.5) × 3–4.5 µm (av. 15.7 × 3.5 μm). Macroconidia falcate, almost straight with moderate apical curvature, tapering towards a blunt to pointy apical cell; basal cell papillate to poorly-developed, foot-shaped, (1–)3–5-septate, (19–)30–44.5(–54) × (3.5–)4–5.5 µm (av. 37.3 × 4.3 μm), mostly 3-septate, (26–)30.5–40(–45.5) × (3.5–)4–5 µm (av. 35.2 × 4.2 μm). Chlamydospores subglobose to globose, smooth- and thick-walled, hyaline, 7–14 µm diam, solitary or grouped in short chains, intercalary on hyphae or borne on short lateral stipes.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.1–9.2 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface velvety, dusty to woolly, red to coral with white to saffron ropes and patches, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse scarlet to coral with peach patches, red at centre. On OA, orange to scarlet with patches of primrose to straw mycelium, flat, velvety, dusty to cottony, with regular and undulate margin; reverse honey to amber.
Typus: China, from soil, before 13 June 1990, unknown collector (holotype designated here CBS H-25415, ex-type culture CBS 151958 = FRC R-9120 = NRRL 66936).
Notes: A soilborne species known from China, F. sagittatum, formerly the Fusarium sp. nov. 24 in Laraba et al. (2021) is described here from a strain originally received as F. compactum (FIESC; Xia et al. 2019). Nevertheless, apart from being phylogenetically distant, several morphological features distinguish these two species including the shape of the macroconidia, which are often straight, barely pedicellate, and not as evidently tapered as in F. compactum; and its conidiogenous cells, which are more elongated, and not forming clusters as in F. compactum (Gerlach & Nirenberg 1982).
The characteristics of the macroconidia also distinguishes F. sagittatum from the typically hyperbolically curved conidia of F. brachygibbosum; and the regularly curved conidia of F. cuspidatum and F. minutum; the latter species, all close phylogenetic relatives of F. sagittatum.
Fusarium sambucinum Fuckel, Fungi Rhen. Exs., Fasc. 3, no. 211. 1863, nom. cons. Figs 64, 65.
Fig. 64.
Fusarium sambucinum (lectotype G00266369). A–C. Herbarium specimen. D. Sporodochium. E–H. Sporodochial conidiophores and conidiogenous cells. I. Macroconidia. Scale bars: D = 20 μm; E, F, I = 10 μm; G, H = 5 μm.
Fig. 65.
Fusarium sambucinum (ex-epitype CBS 151942). A, B. Perithecia on host. C, D. Perithecia in culture. E–G. Detail of perithecial wall cells (G in 100 % lactic acid). H. Detail of perithecial anatomy (in 100 % lactic acid). I. Detail of ostiolar region (in 100 % lactic acid). J–L. Asci. M. Ascospores. N–P. Sporodochia on host. Q. Sporodochial conidiophores and conidiogenous cells. R. Micro- and macroconidia. S. Macroconidia. Scale bars: E (applies to F and G); H–K, Q = 20 μm; L, M, R, S = 10 μm.
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].
Fusarium sambucinum Brondeau, Arch. Fl. France Allemagne 1: 170 (1855)
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.) Quél., Mém. Soc. Émul. Montbéliard, sér. 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., sér. 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 Bérenger, Mem. Accad. Agric. Verona 44: 257. 1866, nom. rej.
Fusarium ricini (Béerenger) Bizz., Fl. Ven. Critt. 1: 539. 1885.
Fusarium subcarneum P. Crouan & H. Crouan, Fl. Finistére: 14. 1867, nom. rej.
Fusarium violaceum P. Crouan & H. Crouan, Fl. Finistére: 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.
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 roseum var. lavaterae-arboreae Thüm., Mycoth. Univ. Cent. 11: no. 1084. 1878.
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. Dével. Mucéd.: 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. Genéve, sér. 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 f.2 Wollenw., Z. Parasitenk. (Berlin) 3: 357. 1931.
Fusarium sambucinum f.3 Wollenw., Z. Parasitenk. (Berlin) 3: 357. 1931.
Fusarium sambucinum f.4 Wollenw., Z. Parasitenk. (Berlin) 3: 357. 1931.
Fusarium sambucinum f.6 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.
Descriptions and illustrations: Wollenweber (1931, 1945), Gerlach & Nirenberg (1982), Nelson et al. (1983), Samuels et al. (1990, 2006), Nirenberg (1995), Leslie & Summerell (2006), Perera et al. (2023).
Typus: Germany, Hessen, from Sambucus nigra dry branches, unknown collection date, Fuckel (lectotype G00266369, designated by Gams et al. 1997); near Berlin, from Sambucus nigra attached, dead and corticated branch, 2 Dec. 2022, R.K. Schumacher, RKS 1176 (epitype CBS H-25241 designated here, MBT 10021761; ex-epitype culture CBS 151942 = CPC 45317).
Additional materials examined: France, Ardèche, Pléout, from Laurus nobilis with scale insects, 8 May 1996, M. Molterbeer, culture CBS 100056 = NRRL 53596. Germany, from Pterocarya fraxinifolia, 9 Mar. 1982, unknown collector, dep. H.I. Nirenberg, culture NRRL 20663 = IMI 266242; from Solanum tuberosum, 1969, Leiber, culture CBS 260.95 = BBA 62397 = DSM 62397 = NRRL 36396; unknown collection date and collector, dep. W.F.O. Marasas, culture CBS 119802 = FRC R-6380 = MRC 2193 = MRC 6970 = NRRL 13455; Berlin, unknown host, before 1981, unknown collector, dep. W. Gerlach, culture NRRL 13394 = BBA 63572 = FRC R-5823; near Berlin, from Sambucus nigra attached, dead and corticated branch, isolated from conidia, 2 Dec. 2022, R.K. Schumacher, RKS 1174, CBS H-25241, culture CBS 151939 = CPC 45309 (isolated from conidia), CBS 151940 = CPC 45310 (from conidia), R.K. Schumacher, RKS 1178, culture CBS 151941 = CPC 45313 (from ascospores), R.K. Schumacher, RKS 1176, culture CBS 151802 = CPC 45315 (from ascospores), R.K. Schumacher, RKS 1175, culture CBS 151943 = CPC 45321 (from conidia), CBS 151944 = CPC 45322 (from ascospores). Netherlands, unknown substrate, before 2001, unknown collector, dep. H. Regeer, culture CBS 111114 = NRRL 36200; from Brassica oleracea, before 1 Nov. 1990, unknown collector, dep. W.F.O. Marasas, culture CBS 119804 = BBA 64995 = FRC R-9153 = MRC 6972 = NRRL 53899; Utrecht, Overvecht, from Solanum tuberosum rotten tuber, 2021, M. Sandoval-Denis, culture CBS 151938 = CPC 41834. New Zealand, Taupo, Hatepe, from Cytisus scoparius, before 1 Sep. 1991, unknown collector, dep. P.R. Johnston, culture NRRL 31964 = NRRL 39954 = ICMP 11550 = IMI 351949; Westland, Haast Beach, from Ulex europaeus lesions on stem and spines, before 1 Mar. 1992, dep. M. Fletcher, culture NRRL 31969 = NRRL 39966 = ICMP 12578. Russia, Siberia, Krasnoyarsk Territory, Minino, from Solanum tuberosum tuber, 2021, Y.A. Litovka, culture CPC 41854 = F80; I.N. Pavlov, culture CPC 41888 = F81. Spain, from Beta vulgaris, 1968, unknown collector, dep. H.I. Nirenberg, culture CBS 125.95 = BBA 62433 = DSM 62433. UK, from Solanum tuberosum tuber, 1982, unknown collector, dep. H.I. Nirenberg, culture CBS 146.95 = BBA 64226 = DSM 115507 = NRRL 20727 = NRRL 22187 = NRRL 36256; 22 Apr. 1985, unknown collector, culture NRRL 20666 = IMI 295478. Unknown, unknown collection data, dep. V. Edel, culture CBS 533.96.
Notes: The original description of Fusarium by Link (1809), with a single species, F. roseum, was based on elements of different fungi i.e., F. avenaceum, F. sambucinum and F. sporotrichioides, in their current concepts (Wollenweber 1916, Gams et al. 1997). The generic name was later sanctioned by Fries (1832), citing only a malvaceous host, hence determining the application of the name (Domsch et al. 2006). The posterior inaccurate use of the name F. roseum by Snyder & Hansen (1945) for an assemblage of over 20 different taxa which excluded the original concept of F. roseum, rendered the latter name ambiguous and unapplicable (Gams et al. 1997). Thus, a proposal to conserve the name F. sambucinum over F. roseum and earlier synonyms was adopted (Gams et al. 1997, Gams 1999). The lectotype of F. sambucinum is shown in Fig. 64.
An earlier, often neglected homonym, F. sambucinum Brondeau (Brondeau 1855) is unavailable because of the conserved status of F. sambucinum Fuckel. The latter name is conserved against all names listed as rejected, and against all combinations of the rejected names (Art. 14.4), as well as against all earlier homonyms (Art 14.10). The current taxonomic position of F. sambucinum Brondeau is unknown; however, with “short, obtuse sporidia” it probably corresponds to a species different than the present concept of F. sambucinum.
Several specimens of F. sambucinum were recently collected from its original host (Sambucus nigra) in Germany, from which seven monosporic cultures, isolated from either ascospores or sporodochial conidia were included here. Their identity was confirmed by morphological and phylogenetic analyses, and sexual crosses with a known F. sambucinum (G. pulicaris) tester strain. To fix the application of the name F. sambucinum to a defined phylogenetic clade, an epitype was selected here (CBS H-25241), and an ex-epitype culture (CBS 151942) was made available to facilitate further research (Fig. 65). The phylogenetic analysis presented in He et al. (2024) indicates CBS 146.95 as “ex-holotype” of F. sambucinum, which is incorrect in every aspect (wrong collection year, collector, country, and substrate).
Fusarium sambucinum resides in the Sambucinum clade of FSAMSC, being morphologically and phylogenetically close to F. agreste, F. seculiforme and F. symmetricum. Shorter macroconidia (av. 28 µm long) differentiate F. sambucinum from F. agreste (av. 34.6 µm long) and differs from F. seculiforme and F. symmetricum by having distinctly apically curved (asymmetrical) conidia, often with a pointy apex, and wider above the median. Additional species commonly compared with F. sambucinum are F. torulosum and F. venenatum (Nirenberg 1995, Leslie & Summerell 2006, Domsch et al. 2007). Fusarium torulosum is genetically unrelated, allocated in the FTSC, and differs by its slow growing colonies, narrower macroconidia, and long chains of chlamydospores (Nirenberg 1995, Leslie & Summerell 2006), while F. venenatum commonly presents slightly larger conidia (av. 44.5 µm long vs 30 µm in F. sambucinum) with up to 9 septa (up to 5 septa in F. sambucinum), and characteristic chlamydospores arranged in curved, terminal chains (Nirenberg 1995, Leslie & Summerell 2006).
Reports of F. sambucinum exist from all over the world, from more than 60 countries in all continents, except Antarctica. This species has been isolated from soil, insects, mammals and human specimens; and it is associated with over 130 plant hosts, either as an endophyte, saprophyte, or inducing canker, die-back or rot symptoms (fruit and root rot, potato dry rot, and storage rot) in more than 100 genera in 45 different families (Farr et al. 2023). However, reports and diseases attributed to F. sambucinum before 1995 should be verified since they could include isolates now identifiable as F. torulosum or F. venenatum (Leslie & Summerell 2006).
Fusarium seculiforme Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855118. Fig. 66.
Fig. 66.
Fusarium seculiforme (ex-type CBS 151806). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F, G. Sporodochial conidiophores and conidiogenous cells. H. Aerial conidiophores and conidiogenous cells. I. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin seculiformis (“sickle-shape, crescent-shape”). In reference to the macroconidial shape.
Sporodochia abundant on the surface of carnation leaves and agar surface, erumpent, transparent, pale luteous to bright orange coloured. Sporodochial conidiophores branching laterally, 26–62 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical, 9–19.5 × 2.5–5 µm, smooth, thin-walled, with a minute or lacking apical collarette. Sporodochial conidia falcate, gently dorsiventrally curved, with blunt apical cell; basal cell papillate or with a well-developed foot-cell, 2–5-septate, (23–)27–35.5(–43.5) × (4–)5(–6) µm (av. 31.3 × 5 μm), mostly 3-septate, (23–)26–32.5(–41) × 4–6 µm (av. 29.4 × 4.9 μm). Aerial conidiophores erect or prostrate on substrate or aerial mycelium, sparsely branched, 26–54 µm tall, often reduced to monophialides forming laterally on aerial mycelium or submerged in agar, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical, 10–12 × 3.5–5 µm, giving rise to falcate conidia indistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.3–8.1 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface cottony at the margin, floccose at centre, scarlet to rust, flat, aerial mycelium short and abundant, moisty at the centre, margin regular, filiform to undulate; reverse rust to pale bay. On OA, luteous to amber, flat, floccose to woolly, with regular and undulate margin, scarlet to rust coloured patches often present; reverse pale luteous to luteous.
Typus: Brazil, from Glycine max, before 31 Jan. 1990, unknown collector, dep. H. Nirenberg (holotype designated here CBS H-25425, ex-type culture CBS 151806 = DSM 64262 = NRRL 22189 = NRRL 20445).
Additional material examined: South Africa, from Solanum tuberosum, before 1 Oct. 1983, unknown collector, culture FRC R-7403 = ATCC 52543 = KOD 1998 = MRC 2967 = NRRL 64059.
Notes: Fusarium seculiforme is here described from strains previously assigned to Fusarium sp. nov. 10 in the Sambucinum clade of FSAMSC (Laraba et al. 2021). Conidial morphology resembles that of the phylogenetically closely related species F. agreste, F. sambucinum and F. symmetricum. Fusarium seculiforme differs from F. agreste by its shorter and more evidently tapered conidia, and from F. sambucinum by its more regularly curved conidia, which lack a distinct apical curvature, showing blunt apices. In addition, it differs from F. venenatum by its shorter conidia (av. 31.3 × 5 µm vs 44.5 × 5.4 µm in F. venenatum). However, due to significant overlap in all morphological characters it is not possible to differentiate this taxon from F. symmetricum.
Fusarium sibiricum Gagkaeva et al., Int. J. Food Microbiol. 147: 64. 2011. Fig. 67.
Fig. 67.
Fusarium sibiricum (ex-type CBS 151951). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E–H. Conidiophores and conidiogenous cells. I. Microconidia. Scale bars = 10 μm.
Descriptions and illustrations: Yli-Mattila et al. (2011).
Typus: Russia, Far East, Khabarovsk, from Avena sativa grain, 2003, L.S. Malinovskaya & E.A. Pirjazeva (holotype LEP12652, ex-type culture CBS 151951 = NRRL 53430 = MFG 11013).
Additional materials examined: Russia, Far East, Blagoveshchensk, from Avena sativa grain, 2001, unknown collector, dep. T. Yu. Gagkaeva & A. van Diepeningen, culture CBS 140945 = MFG 11014 = NRRL 53431 = RCAM 03252; Siberia, Buryatia, from Hordeum vulgare grain, 2002, unknown collector, dep. T.Yu. Gagkaeva & A. van Diepeningen, culture CBS 140910 = MFG 11005 = NRRL 53422 = RCAM 00585; Krasnoyarsk, from Hordeum vulgare grain, 2000, unknown collector, dep. T.Yu. Gagkaeva & A. van Diepeningen, culture CBS 140915 = MFG 11007 = NRRL 53424 = RCAM 03252; The Republic of Khakassia, Salt Lake Tus, from submersible wood, unknown collection date and collector, dep. I.N. Pavlov, culture CPC 41132 = F12; 2019, I.N. Pavlov, culture CPC 41881 = F57. Unknown, from Triticum aestivum, unknown collection data, culture BBA 71293 = DSM 116899, BBA 71294 = DSM 116900.
Notes: A sister species to F. sporotrichioides, F. sibiricum can be recognized by lacking polyphialides, mesoconidia, macroconidia and chlamydospores. The lack of polyphialides and red pigments, plus, its long (> 100 µm), nodose conidiophores, differentiates F. sibiricum from F. langsethiae, another genetically close species in the Sporotrichioides clade lacking macroconidia (Yli-Mattila et al. 2011). Morphological differentiation of F. sibiricum from F. poae can be very difficult when strains of the latter species lack macroconidia and red pigments; however, conidiophores of F. poae are short and branched (Leslie & Summerell 2006), while both species are phylogenetically distant, F. poae being allocated in the Sambucinum clade of FSAMSC. For details regarding hosts and biogeography see Yli-Mattila et al. (2011).
Fusarium sporotrichioides Sherb., Mem. Cornell Univ. Agric. Exp. Sta. 6: 183. 1915. Figs 68, 69.
Fig. 68.
Fusarium sporotrichioides (lectotype). Reproduction from the original publication by Sherbakoff (1915). A, E. Macroconidia. B, J, M, N. Aerial conidiophores. C, G. Balls of conidia. D. Germinating conidium. F. Chlamydospores. H. Sporodochial conidiophore. I, K, L. Microconidia. O. Plectenchymic stroma.
Fig. 69.
Fusarium sporotrichioides (CBS 448.67). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochium. F. Sporodochial conidiophore and conidiogenous cells. G–J. Aerial conidiophores and conidiogenous cells. K. Microconidia. L. Macroconidia. Scale bars = 10 μm.
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.
Descriptions and illustrations: Sherbakoff (1915), Wollenweber (1916), Wollenweber & Reinking (1935), Gordon (1952), Subramanian (1971), Booth (1971), Gerlach & Nirenberg (1982), Nelson et al. (1983), Leslie & Summerell (2006), and Domsch et al. (2007).
Typus: USA, New York, from Solanum tuberosum rotten tuber, 1915, C.D. Sherbakoff [lectotype fig. 22, page 184 in Fusaria of potatoes. Mem. Cornell Univ. Agric. Exp. Sta. 6, designated by Crous et al. (2021)].
Additional materials examined: Belgium, Heverlee, from Nicotiana tabacum seedling root, in greenhouse, before 9 Mar 1964, unknown collector, isol. G.L. Hennerbert, culture CBS 178.64 = MUCL Be-133 = NRRL 36295. Canada, Ontario, from Zea mays kernel, 1986, G. Neish, culture CBS 119839 = DAOM 175513 = MRC 4333 = NRRL 53912. Denmark, Rørvig, Skansehage, from Juncus sp., stipe, Nov. 1984, U. Thrane, culture CBS 412.86 = FRC T-822 = IMI 309350. UK, Hertfordshire, Harpenden, from Hordeum vulgare grain, 1974, unknown collector, culture IMI 348484. France, from Zea mays, before 15 Dec. 1967, W. Snyder, culture CBS 151950 = NRRL 3299 = NRRL 3287 = ATCC 24631 = FRC T-423 = FRC T-424 = MRC 1768 = MRC 43; Channel Island, from Dianthus sp. stem, 25 Feb. 1988, unknown collector, culture IMI 322457. Germany, from Malus sp., unknown collection date and collector, culture BBA 62422 = DSM 115488; from Pinus nigra, unknown collection date and collector, culture BBA 62423 = DSM 115489; Berlin, Grunewald, from Pinus sylvestris, unknown collection date and collector, culture BBA 62421 = DSM 115487; Niedersachsen, from Picea abies seed, 1965, E. Seemüller, culture CBS 448.67 = BBA 10360 = FRC T-0980 = NRRL 25474. Netherlands, Knardijk, from soil under nettles, unknown collection date, U. Thrane, culture CBS 123674 = IBT 1926. Russia, Northwestern Federal District, St. Petersburg, Botanical Garden, from soil under Aesculus hippocastanum, 2021, I.N. Pavlov, culture CPC 41842 = F97; Siberia, Krasnoyarsk Territory, from Hordeum vulgare seed, unknown collection date and collector, dep. I.N. Pavlov, culture CPC 41121 = F1; Dolgolostovsky Forestry, from Pinus sylvestris seed, unknown collection date and collector, dep. I.N. Pavlov, culture CPC 41150 = F30; Pugachevo, from Solanum tuberosum tuber, 2019, Y.A. Litovka, culture CPC 41885 = F66.
Notes: Fusarium sporotrichioides was recently lectotypified by Crous et al. (2021), the lectotype, an original drawing by C.D. Sherbakoff is reproduced here (Fig. 68). Epitypification, however, is still pending the selection of a suitable specimen from the correct host and country.
Fusarium sporotrichioides produces pyriform microconidia, which can be confused with the napiform to globose microconidia of its closest molecular and morphological sibling species F. langsethiae and F. sibiricum; and, with the globose to pyriform microconidia of the distantly related F. poae (Sambucinum clade). The presence of polyphialides distinguishes F. sporotrichioides from F. poae. This same feature and its red PDA colonies distinguishes F. sporotrichioides from F. sibiricum. Lastly, the presence of either aerial or sporodochial macroconidia and mesoconidia separates F. sporotrichioides from F. langsethiae.
Unlike morphologically similar species (F. langsethiae, F. poae, and F. sibiricum), F. sporotrichioides is common in soil, in temperate and tropical climates (Leslie & Summerell 2006, Domsch et al. 2007). It has also been reported from animal feed, insects, as a human pathogen, and from mushrooms, and it is a potent mycotoxin (trichothecenes) producer (O’Donnell et al. 2012, Al-Hatmi 2016, Farr et al. 2023). Known plant hosts include over 70 genera, mostly in Poaceae, but also Sapindaceae, Apiaceae, Aquifoliaceae, Araliaceae, Arecaceae, Asteraceae, Betulaceae, Cactaceae, Cannabaceae, Caryophyllaceae, Cucurbitaceae, Ericaceae, Fabaceae, Iridaceae, Juncaceae, Lamiaceae, Linaceae, Lythraceae, Malvaceae, Musaceae, Nymphalidae, Pinaceae, Rosaceae, Rutaceae, Salicaceae, Sclerotiniaceae, Solanaceae, Tortricidae, Typhaceae, and Verbenaceae (Farr et al. 2023); described from Africa (Nigeria, South Africa, and Tanzania), Asia (Bahrain, China, Iran, Japan, Malaysia, Oman, and Türkiye), Europe (Austria, Belgium, Bulgaria, Croatia, Denmark, England, Finland, France, Germany, Hungary, Italy, Norway, Poland, Russia, UK, and the former Yugoslavia), North and South America (Barbados, Brazil, Canada, Chile, Colombia, Mexico, Puerto Rico, and the USA); and Oceania (Australia, and New Zealand). However, due to inaccurate morphological determinations in published literature, host and geographic records need to be reassessed (Domsch et al. 2007).
Fusarium subcylindroides Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855119. Fig. 70.
Fig. 70.
Fusarium subcylindroides (ex-type CBS 151807). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G–I. Sporodochial conidiophores and conidiogenous cells. J. Aerial conidiogenous cell. K. Macroconidia. Scale bars = 10 μm.
Etymology: From Latin sub- (“less than completely”) and cylindrus (“a cylinder”). In reference to the shape of the macroconidia.
Sporodochia abundant on the surface of carnation leaves, agar surface and aerial mycelium, sulphur yellow, luteous to pale citrine coloured. Sporodochial conidiophores branching irregularly laterally, 40.5–79 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, subcylindrical, (10.5–)12.5–17.5(–22) × 3.5–6 µm, smooth- and thin-walled, with or without a minute apical collarette. Sporodochial conidia falcate, robust, gently dorsiventrally curved, with blunt apical cell; basal cell papillate, rarely with a well-developed foot-cell, (1–)3–5-septate, (28.5–)33–39.5(–46.5) × 4–6.5 µm (av. 36.4 × 5.1 μm), mostly 3-septate, (28.5–)32–37(–39) × 4–6 µm (av. 34.7 × 4.9 μm). Aerial conidiophores erect or prostrate on substrate mycelium, sparsely laterally branched, 30–69 µm tall, often reduced to monophialides forming laterally on aerial mycelium or submerged in agar, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical, (8.5–)9.5–19(–22) × 3.5–5.5 µm, smooth- and thin-walled, giving rise to falcate conidia indistinguishable from sporodochial conidia. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 8.9–9.2 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface felty to cottony, buff to rosy buff, red to scarlet at centre, flat, aerial mycelium abundant, margin regular and filiform; reverse scarlet to red, blood at centre. On OA, sulphur yellow to buff with scarlet to red excretions, pale luteous to pure yellow at periphery, flat, felty to woolly, with regular and filiform margin; reverse pale luteous to amber.
Typus: USA, from soil, before 27 Aug. 1997, unknown collector (holotype designated here CBS H-25424, ex-type culture CBS 151807 = BBA 70355 = NRRL 26795).
Notes: Fusarium subcylindroides is described here from a strain isolated from soil in the USA, originally identified as F. torulosum and later determined as Fusarium sp. nov. 17 in Laraba et al. (2021). Apart from being genetically very distant, several features differentiate F. subcylindroides from F. torulosum (a member of the FTSC), including the often 3-septate macroconidia with pointy apical cells (vs commonly 5-septate, with rounded apices in F. torulosum), the lack of microconidia and chlamydospores, the later typically formed in long chains in F. torulosum; and a faster growth rate on PDA (av. 9.1 mm/d vs 2.2 mm/d in F. torulosum; Nirenberg 1995). Due to its poorly developed basal cells and because of its gentle curvature and tapering, conidia of F. subcylindroides adopt a somewhat cylindrical appearance, which stands out against the remaining species in the FSAMSC.
Fusarium subflagellisporum T.F. Nóbrega & R.W. Barreto, Persoonia 47: 313. 2021.
Description and illustration: Crous et al. (2021).
Typus: Brazil, Pernambuco, Petrolina, Special Fruit Farm, from Mangifera indica hypertrophied floral and vegetative branches, 30 Aug. 2019, T.F. Nóbrega (holotype VIC 47377, ex-type culture COAD 2989).
Materials examined: Puerto Rico, from Sorghum sp. debris, before 4 Jan. 1984, culture FRC R-7642 = KOD 2007. USA, Illinois, from soil, before 8 Oct. 2019, unknown collector, culture CBS 151955 = FRC R-8774 = NRRL 66930.
Notes: One of the basal-most clades in FSAMSC, F. subflagellisporum, originally discovered as the phylogenetic species Fusarium sp. nov. 27 by Laraba et al. (2021), is characterized by producing falcate aerial macroconidia, but also whip-like sporodochial macroconidia, which contrasts with all its closest molecular siblings in the Brachygibbosum clade (F. bellum, F. brachygibbosum, F. cuspidatum, F. minutum, and F. sagittatum). Additionally, F. subflagellisporum produces abundant chains of pigmented chlamydospores and has been reported with reduced growth rates on PDA (av. 6 mm/d vs above 7.4 mm/d for the aforementioned species) (Crous et al. 2021).
This species is known from Africa (Nigeria and Zimbabwe); and South and North America (Brazil, Puerto Rico, and the USA), mostly from agricultural soil debris, but also from Mangifera sp. (Anacardiaceae) and Arachis hypogaea (Fabaceae) (Crous et al. 2021, Laraba et al. 2021, Farr et al. 2023).
Fusarium subtropicale C.B. Pereira et al., Mycologia 110: 864. 2018. Fig. 71.
Fig. 71.
Fusarium subtropicale (ex-type CBS 144706). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochium. F. Sporodochial conidiophore and conidiogenous cells. G–J. Aerial conidiophores and conidiogenous cells. K. Macroconidia. Scale bars: F = 5 μm; G–K = 10 μm.
Description and illustration: Pereira et al. (2018).
Typus: Brazil, Parana, Guarapuava, from Hordeum vulgare root, 2013, H.R. Feksa, F1478 (holotype BPI 910644, ex-type culture CBS 144706 = NRRL 66764 = UEM 3329).
Notes: Resolved as the closest phylogenetic lineage to F. praegraminearum, both species are only known from a single strain each (Pereira et al. 2018). However, both taxa were previously resolved as distinct lineages using a 10-gene dataset (Pereira et al. 2018), which matches our phylogenetic and coalescent results (Figs 1, 3). For a detailed discussion on morphological differentiation see Pereira et al. (2018).
Fusarium symmetricum Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855120. Fig. 72.
Fig. 72.
Fusarium symmetricum (ex-type CBS 151804). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochium. F, G. Sporodochial conidiophores and conidiogenous cells. H–J. Aerial conidiophores and conidiogenous cells. K. Macroconidia. Scale bars: F, G, I, K = 10 μm; J = 5 μm.
Etymology: From Latin symmētricus (“symmetrical”). In reference to the symmetrically curved macroconidia.
Sporodochia abundant on the surface of carnation leaves and agar surface, ochreous, luteous to bright orange coloured. Sporodochial conidiophores densely packed, branching irregularly laterally and verticillately, 26–82 µm tall, bearing lateral and terminal monophialides and apical whorls of monophialides. Sporodochial conidiogenous cells monophialidic, ampulliform to subcylindrical, 8.5–19 × 2.5–6 µm, smooth- and thin-walled, apical collarette minute or lacking. Sporodochial conidia falcate, moderately dorsiventrally curved, with blunt apical cell; basal cell well-developed foot-shaped, (0–)1–5-septate, (16.5–)25.5–34.5(–41.5) × (3.5–)5–6(–6.5) µm (av. 30 × 5.3 μm), mostly 3-septate, (21.5–)25.5–32.5(–36.5) × 4–6 µm (av. 29.1 × 5.2 μm). Aerial conidiophores erect or prostrate on substrate mycelium, simple to sparsely branched irregularly and verticillately, 15–40 µm tall, or reduced to solitary lateral monophialides and later converging into sporodochia. Aerial conidiogenous cells monophialidic, ampulliform, subulate to subcylindrical, 5.5–24 × 3.5–5.5 µm, smooth- and thin-walled, with a minute or lacking apical collarette, giving rise to falcate conidia indistinguishable from sporodochial conidia grouped in small false heads. Chlamydospores not formed.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.2–8.6 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface at first cottony to woolly, rust to coral with sulphur yellow to apricot aerial mycelium, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse rust to red, isabelline at centre. On OA, sulphur to pure yellow with patches or irregular rings of white, rosy vinaceous to amber mycelium, flat, felty to woolly, with regular, filiform to undulate margin; reverse pale luteous to ochreous.
Typus: Argentina, Opuntia aurantiaca, before 9 Apr. 1985, unknown collector (holotype designated here CBS H-25411, ex-type culture CBS 151804 = NRRL 13465 = NRRL A-26556).
Additional material examined: Argentina, Opuntia aurantiaca, before 1 Dec. 1982, unknown collector, CBS H-25401, culture CBS 151819 = FRC R-7071 = MRC 2857 = NRRL 66925.
Notes: Previously as Fusarium sp. nov. 9 in Laraba et al. (2021), the lineage here described as F. symmetricum is, so far, only known from Argentina.
Fusarium symmetricum produces fast growing, red PDA colonies and up to 5-septate sporodochial conidia of similar width to those of F. agreste, F. sambucinum, and F. seculiforme, its closest phylogenetic siblings. Apart from its more prominently tapered, symmetrical and regularly curved macroconidia, the lack of microconidia, and the lack of both microconidia and chlamydospores distinguishes F. symmetricum from F. agreste and F. sambucinum, respectively. On the other hand, F. symmetricum and F. seculiforme cannot be separated with confidence based on morphological characters only.
Fusarium transvaalense Sand.-Den., Crous & W.J. Swart, MycoKeys 34: 82. 2018.
Description and illustration: Sandoval-Denis et al. (2018).
Typus: South Africa, Kruger National Park, Skukuza, Granite Supersite, from Sida cordifolia rhizosphere, 23 Mar. 2015, W.J. Swart (holotype CBS H-23497, ex-type culture CBS 144211 = CPC 30923).
Additional materials examined: Australia, from soil, 1975, unknown collector, culture NRRL 31008 = BBA 63772 = DSM 115500 = FRC R-9856. South Africa, Kruger National Park, Skukuza, Granite Supersite, from Kyphocarpa angustifolia rhizosphere, 23 Mar. 2015, W.J. Swart, culture CBS 144222 = CPC 30939; from Melhania acuminata rhizosphere, 23 Mar. 2015, W.J. Swart, cultures CBS 144212 = CPC 30929, CBS 144224 = CPC 30928; from Sida cordifolia rhizosphere, 23 Mar. 2015, W.J. Swart, cultures CBS 144214 = CPC 30946, CBS 144217 = CPC 30919, CBS 144218 = CPC 30922, CBS 144220 = CPC 30927; Kwazulu-Natal province, from plant debris in soil, 1982, W.F.O. Marasas, culture CBS 119879 = FRC R-6837 = FRC R-9976 = MRC 2800; Western Cape, Philadelphia, from Dianthus sp. roots, 1990, S. Lamprecht, culture CBS 119878 = MRC 6142 = NRRL 53947.
Notes: Originally described from the rhizosphere of three native African herbaceous plants i.e., Melhania acuminata and Sida cordifolia (Malvaceae), and Kyphocarpa angustifolia (Amaranthaceae) in South Africa (Sandoval-Denis et al. 2018), the host and geographic range of this species have been recently expanded to include isolates from soil, Avena sativa horse feed (Poaceae), and Medicago sativa (Fabaceae); recorded also from Australia and the USA (Wang et al. 2022, Farr et al. 2023).
Fusarium transvaalense is distinguished from its sibling species in the Brachygibbosum clade (F. brachypes, F. dimorphosporum, F. hamatum, F. mucronatum, and F. platysporum) by a combination of morphological features including the presence of clavate microconidia, two macroconidial morphotypes, and although with some overlap, the reduced size of its sporodochial macroconidia (av. 33.2 × 5.1 µm vs > 45 µm long, and ≥ 6 µm wide on average for all the above mentioned species).
Fusarium tropicale Sand.-Den., M.M. Costa, J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855121. Fig. 73.
Fig. 73.
Fusarium tropicale (ex-type CBS 151820). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G, H. Sporodochial conidiophores and conidiogenous cells. I, J. Aerial conidiophores and conidiogenous cells. K. Chlamydospores. L. Macroconidia. Scale bars: G, H, L = 10 μm; I–K = 5 μm.
Etymology: From Greek tropikós (“tropic, tropical”). In reference to the geographical origin of the type.
Sporodochia abundant and erumpent on the surface of carnation leaves, agar surface, substrate and aerial mycelium, rosy buff, pale peach to orange coloured. Sporodochial conidiophores sparingly branching laterally and irregularly, 28–37 µm tall, bearing lateral monophialides or groups of terminal monophialides. Sporodochial conidiogenous cells monophialidic, doliiform to subcylindrical, 7–15 × 3–5 µm, smooth- and thin-walled, lacking apical collarette. Sporodochial conidia falcate, gently to moderately dorsiventrally curved with an almost straight ventral line, commonly widest above the median, apical cell tapering, blunt to pointy; basal cell well-developed, foot-shaped, rounded or slightly elongated, (1–)3–5-septate, (21.5–)40–49(–52) × (3.5–)5–6(–6.5) µm (av. 44.3 × 5.5 μm), mostly 5-septate, (38.5–)42.5–48(–52) × 4.5–6.5 µm (av. 44.8 × 5.6 μm). Aerial conidiophores erect or prostrate on substrate mycelium, simple to sparsely laterally branched, reaching 20–45 µm tall, often reduced to monophialides forming laterally on aerial mycelium or submerged in agar, and rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, subcylindrical to ampulliform, 10–12 × 3.5–5 µm, smooth- and thin-walled, with a minute to absent apical collarette, giving rise to falcate conidia identical to sporodochial conidia. Chlamydospores globose to somewhat obovate, smooth- and thick-walled, hyaline, 5–9.5 µm diam, grouping in chains or clusters formed intercalary or lateral on hyphae.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 7.7–8.2 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface felty to woolly, peach to red with white, rosy vinaceous to amber patches and ropes, flat, aerial mycelium abundant, margin regular, filiform to undulate; reverse scarlet to red, grey olivaceous to chestnut at centre. On OA, felty, cottony to woolly, rosy buff to saffron with white, peach to rose, and pale luteous to pure yellow patches, flat, with regular and filiform margin; reverse pale luteous to honey.
Typus: Thailand, Ubon, from rice paddy soil, before 1 Aug. 1984, unknown collector (holotype designated here CBS H-25427, ex-type culture CBS 151820 = FRC R-7764 = NRRL 66926).
Additional material examined: India, Assam, Pin, from Boehmeria nivea, 10 Dec. 1986, unknown collector, culture IMI 312359.
Notes: Fusarium tropicale, formerly Fusarium sp. nov. 20 (Laraba et al. 2021), exhibits similar overall morphological features to F. cuspidatum, F. dimorphosporum, F. hamatum, and F. platysporum (Brachygibbosum clade); and F. pratense (Longipes clade). The latter species, together with F. carinatum and F. tropicale are also morphologically odd species in the Longipes clade, which instead of the strongly apically elongated macroconidia common to this clade, present a morphology more closely related to species in the Brachygibbosum clade. The lack of microconidia differentiates F. tropicale from F. dimorphosporum, F. platysporum, and F. pratense, while its red colonies further differentiate F. tropicale from F. pratense. The lack of chlamydospores separates F. tropicale from F. cuspidatum, and with macroconidia averaging (44.3 × 5.5 µm), F. tropicale differs from the shorter conidia of F. cuspidatum (av. 42 × 5.5 µm) and the wider conidia of F. hamatum (av. 47.4 × 6.2 µm).
Fusarium ussurianum T. Aoki et al., Mycologia 101: 847. 2009. Fig. 74.
Fig. 74.
Fusarium ussurianum (ex-type CBS 123752). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G. Sporodochial conidiophore and conidiogenous cells. H, I. Aerial conidiogenous cells. J. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: Yli-Mattila et al. (2009), Aoki et al. (2012).
Typus: Russia, Primorsky krai (Far East territory), field near the city of Ussuriysk, from Avena sativa seed, 2002, T. Gagkaeva (holotype BPI 878845, ex-type culture CBS 123752 = NRRL 45681 = TG-2662/0).
Additional materials examined: Russia, from Triticum sp., unknown collection date and collector, culture NRRL 28813; Primorsky Krai, Triticum sp., unknown collection date and collector, culture NRRL 58212; Kamen-Ribolov, Triticum sp. seed, 2006, T. Gagkaeva, culture CBS 123753 = FRC R-10095 = NRRL 45795 = TG-65202.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum.
Fusarium ussurianum is known from Poaceae (Avena sativa, Hordeum vulgare, and Triticum sp.), and Cyperaceae (Rhynchospora sp.), and has been reported from China and Russia (Yli-Mattila et al. 2009, Aoki et al. 2012, Wang et al. 2022, Farr et al. 2023).
This species is the closest phylogenetic relative to F. acaciae-mearnsii, F. asiaticum and F. vorosii in Fusarium graminearum s. lat. It can be recognized among F. graminearum segregates by producing > 5 µm, symmetrically curved and apically narrowly beaked 5-septate macroconidia, which are wider above the median. These are unique characters for this species (Aoki et al. 2012). In contrast, the phylogenetically closest species mentioned above all produce asymmetric 5-septate conidia, which are either 4–5 µm wide (F. acaciae-mearnsii and F. asiaticum), lack narrow apical beaks (F. asiaticum and some strains of F. vorosii), or are wider below the median (F. acaciae-mearnsii) (Aoki et al. 2012).
Fusarium venenatum Nirenberg, Mycopathologia 129: 136. 1995. Fig. 75.
Fig. 75.
Fusarium venenatum (ex-type CBS 458.93). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G–I. Aerial conidiophores and conidiogenous cells. J. Chlamydospores. K. Macroconidia. Scale bars: F–H, J, K = 10 μm; I = 5 μm.
Descriptions and illustrations: Nirenberg (1995), O’Donnell et al. (1998a), Leslie & Summerell (2006).
Typus: Austria, Rohrau, from Triticum aestivum culm, before 1 Feb. 1991, H.I. Nirenberg (holotype CBS 458.93, a metabolically inactive culture, ex-type strain CBS 458.93 = BBA 64537 = FRC R-9186 = NRRL 26228).
Additional materials examined: Finland, from Solanum tuberosum plant, 1984, Seppänen, culture CBS 127.95 = BBA 64478 = NRRL 36230. Germany, from Zea mays, before 2 Feb. 1992, unknown collector, dep. H.I. Nirenberg, culture NRRL 22196 = BBA 65031 = DSM 116036 = FRC R-9187. Russia, Moscow, Central part, from unidentified grass, 2009, unknown collector, dep. T.Yu. Gagkaeva & A. van Diepeningen, culture CBS 140911 = MFG 58720.
Notes: Fusarium venenatum was described as a segregate species from F. sambucinum, together with F. torulosum, the three species originally distinguished by their macroconidial size, presence of microconidia and chlamydospores, and growth rates on PDA (Nirenberg 1995). Fusarium torulosum, differing by slower growth rates, red/purple PDA pigmentation, and narrower macroconidia (Nirenberg 1995, Leslie & Summerell 2006, Domsch et al. 2007) was later reaccommodated in the FTSC, hence it is genetically distant to F. sambucinum and F. venenatum (O’Donnell et al. 2013, Crous et al. 2021, Laraba et al. 2022). Both Fusarium sambucinum and F. venenatum are nested within the Sambucinum clade of FSAMSC but are well-resolved phylogenetically. With up to 9-septate macroconidia averaging 44.5 µm long, its typical terminal chains of chlamydospores, and lacking microconidia, F. venenatum differs from F. sambucinum (macroconidia av. 30 µm long, up to 5-septate) and the closest phylogenetic siblings F. amblysporum (macroconidia av. 47.4 µm long, up to 7-septate) and F. cultriforme (macroconidia av. 30.2 µm long, up to 5-septate). Additionally, the growth rates and pigmentation of F. venenatum on PDA (intense red colonies, av. 7.6 mm/d) differ from those of F. amblysporum (white to rosy buff, av. 8.2 mm/d), and F. cultriforme (yellow, av. 4.3 mm/d).
Fusarium venenatum is known from soil, soil debris, and several plant hosts in Amaranthaceae (Beta vulgaris), Asteraceae (Tanacetum cinerariifolium), Cannabaceae (Humulus spp.), and Poaceae (Hordeum vulgare, Triticum spp., and Zea mays), mostly from Europe (Austria, England, Finland, France, Germany, Norway, Poland, Russia, and Spain), but also in Oceania (Australia, New Zealand, Tasmania) and the USA (Farr et al. 2023). The strains involved in the production of the commercial mycoprotein Quorn, originally assigned to F. graminearum, were later identified as F. venenatum by O’Donnell et al. (1998a).
Fusarium vermicularioides Sand.-Den., J.Z. Groenew. & Crous, sp. nov. MycoBank MB 855122. Fig. 76.
Fig. 76.
Fusarium vermicularioides (ex-type CBS 151945). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E, F. Sporodochia. G. Sporodochial conidiophores and conidiogenous cells. H. Aerial conidiogenous cell. I. Aerial macroconidia and microconidium (arrow). J. Chlamydospores. K. Sporodochial macroconidia. Scale bars: G, I–K = 10 μm; H = 5 μm.
Etymology: From Latin vermis (“worm, snake”) and -oīdēs (“-like, -oid”). Referring to the macroconidial shape, resembling a worm.
Sporodochia abundant on the surface of carnation leaves and agar surface, ochreous, luteous to pale sienna coloured. Sporodochial conidiophores sparingly branched laterally, 15–45 µm tall, bearing lateral and terminal monophialides. Sporodochial conidiogenous cells monophialidic, ampulliform to subcylindrical, 6–17 × 2.5–5.5 µm, smooth- and thin-walled, with a flared apical collarette. Sporodochial conidia falcate to whip-like, gently to strongly dorsiventrally curved and tapering abruptly towards both ends, apical cell moderately elongate; basal cell papillate to well-developed foot-shaped and elongate, 4–6-septate, (38.5–)47.5–63(–77) × 3–6 µm (av. 55.4 × 4.5 μm), mostly 5-septate, (39.5–)48–63(–77) × 3–6 µm (av. 55.6 × 4.5 μm). Aerial conidiophores erect or prostrate on substrate mycelium, unbranched, reaching 10–25 µm tall, often reduced to monophialides forming laterally on aerial mycelium or submerged in agar or on short stipes, rapidly converging onto sporodochia. Aerial conidiogenous cells monophialidic, ampulliform to subcylindrical, 6–10.5 × 3.5–5 µm, with short flared apical collarettes, giving rise to falcate conidia and very rarely to microconidia. Aerial falcate conidia almost straight to moderately dorsiventrally curved, apical cell blunt, basal cell blunt to papillate, (2–)3–5(–6)-septate, (21.5–)30–47(–61.5) × 3.5–5 µm (av. 38.3 × 4.2 μm), mostly 5-septate, (32.5–)34.5–50(–61.5) × 3.5–5 µm (av. 42.3 × 4.3 μm); microconidia ellipsoidal, 0–1-septate, (7.5–)8.5–14.5(–15.5) × 3.5–4 µm. Chlamydospores subglobose to globose smooth- and thick-walled, 12.5–17 µm diam, subhyaline, solitary or in short chains.
Culture characteristics: Colonies on PDA at 25 °C growing in the dark with an average radial growth rate of 6.7–7.1 mm/d, filling an entire 9-cm-diam Petri dish after 7 d. Surface membranous to velvety, sulphur yellow to honey, flat, aerial mycelium scant, moisty at the centre, margin regular and filiform; reverse sulphur yellow to buff. On OA, sulphur yellow to pale luteous, flat, membranous to slimy, dusty at periphery, margin regular, filiform to undulate; reverse pale luteous to honey.
Typus: New Guinea, from soil debris, before 1 May 1979, unknown collector (holotype designated here CBS H-25423, ex-type culture CBS 151945 = FRC R-5128 = NRRL 13374).
Notes: Previously assigned to Fusarium longipes 2 (O’Donnell et al. 2013, Laraba et al. 2021), F. vermicularioides is one of the segregated lineages of F. longipes s. lat., elevated here to species level. Fusarium vermicularioides, together with F. carinatum and F. pratense are distinguished from all remaining species in the Longipes clade by forming yellow colonies on PDA. Its slower growth rate (up to 7.1 mm/d), presence of ellipsoidal but neither clavate or falcate microconidia, and formation of whip-like macroconidia differentiates F. vermicularioides from F. carinatum and F. pratense (PDA growth rates above 7.5 mm/d for the two latter species).
Fusarium vorosii B. Tóth et al., Fungal Genet. Biol. 44: 1202. 2007. Fig. 77.
Fig. 77.
Fusarium vorosii (ex-type CBS 119177). A, B, D. Colonies on PDA, OA, and SNA respectively. C. Reverse of colonies on PDA (up), and OA (down). E. Sporodochia. F. Sporodochial conidiophore and conidiogenous cells. G, H. Aerial conidiogenous cells. I. Macroconidia. Scale bars = 10 μm.
Descriptions and illustrations: Starkey et al. (2007), Aoki et al. (2012).
Typus: Hungary, Pest County, Ipolydamásd, from Triticum sp. spikelet, Dec. 2002, G. Giczey & L.L. Hornok (holotype BPI 871658, ex-type culture CBS 119177 = FRC R-10011 = NRRL 37605).
Additional material examined: Japan, Hokkaido, from Triticum sp., head, before 2005, unknown collector, culture CBS 119178 = FRC R-9999 = NRRL 38208.
Notes: For comments about the F. graminearum s. lat. clade see notes under F. acaciae-mearnsii and F. graminearum.
Fusarium vorosii is known from Poaceae (Hordeum vulgare, Oryza sativa, Triticum spp., and Zea mays), from Asia (Japan and Korea) and Europe (Hungary, Netherlands and Russia) (Aoki et al. 2012, Farr et al. 2023).
Morphologically, this species is hardly distinguishable from F. graminearum, F. asiaticum and F. aethiopicum, all of which are characterized by asymmetric, gradually curved 5-septate macroconidia, without narrow apical beaks, and being wider above the mid-region. Nevertheless, 5-septate conidia of F. vorosii are often > 5 μm wide and its morphology is not as consistent since straight, apically beaked conidia can also be present (Starkey et al. 2007, Aoki et al. 2012).
DISCUSSION
In this study we tested recent phylogenetic circumscriptions in FSAMSC using multigene phylogenetics, traditional cultural and morphological analyses, and coalescent-based species delimitation methods using a large set of strains from four culture collections (i.e., BBA, CBS, IMI and NRRL) plus recent isolations from diverse substrates and locations. With more than 75 species [75 species resolved here, plus F. guizhuense (He et al. 2024), additional undescribed lone lineages in F. graminearum s.l. (O’Donnell et al. 2018), and an unpublished African novel lineage sister to F. brachygibbosum (A. Jacobs, pers. comm.)], the FSAMSC is one of the most speciose groups in Fusarium together with the F. fujikuroi SC (FFSC) and the FIESC with ca. 90 and 60 spp., respectively (www.fusarium.org, accessed 6 Nov. 2024), and one of the most relevant in terms of plant pathogenicity and mycotoxin production (Dean et al. 2012, Bhunjun et al. 2024).
In contrast to other recently revised Fusarium species complexes (e.g., FCSC, Lombard et al. 2019; FLSC, Costa et al. 2024), recent studies had already partially dealt with several matters on the FSAMSC, including building a phylogenetic backbone (O’Donnell et al. 2013, Crous et al. 2021, Laraba et al. 2021), mycotoxin profiling (O’Donnell et al. 2018, Laraba et al. 2021), and fixing taxonomical issues (Crous et al. 2021). Nevertheless, these aspects had not been assembled into a cohesive picture, with over 35 phylogenetic lineages awaiting formal recognition and unsettled typification for historically relevant taxa. While efforts were made to include type material for all known species of FSAMSC in the analyses, five phylogenetic species (FSAMSC2, FSAMSC7, FSAMSC11, FSAMSC13, and FSAMSC31) could not be formally introduced in this study because the original strains were unavailable at the time of writing this manuscript and conspecific isolates were not available to us. Moreover, even when nomenclatural types exist for all species in FSAMSC, we were unable to analyse any type material for F. sporotrichioides, since Sherbakoff’s original specimens for this species are not present in CUP (T. Iturriaga, pers. comm.); hence, the species was lectotypified with an illustration (Crous et al. 2021), while we were unable to locate a proper strain to serve as epitype (USA, New York, from Solanum tuberosum). A similar situation occurred for F. cerealis, for which a neotype is available [K(M)133541, Nirenberg 1990] and was examined here, although, no suitable epitype could be designated from the living material studied. Conversely, efforts to collect appropriate material for epitypification of F. sambucinum were successful, resulting in the unambiguous genetic and biological allocation of the generic type to a discrete phylogenetic clade.
The taxonomical history of the FSAMSC illustrates the non-monophyletic nature of the deprecated sectional taxonomical system of Fusarium (O’Donnell 1993, 1996, 1997, O’Donnell et al. 2013). Moreover, unlike genetically close species complexes (e.g., FCSC, FFSC, FOSC, and FTSC), morphological heterogeneity prevents us to draw a clear phenotypic definition for the FSAMSC. In fact, as shown here, relying on morphological aspects only, allocation of many FSAMCS taxa within the complex is challenging, but possible through a detailed inspection under standard culture conditions (Crous et al. 2021). Morphological identification can indeed be achieved for many species treated here, but it is quite impractical, especially considering the amount of curated DNA data currently available for comparison on public nucleotide depositories. The latter is also valid for other recently revised Fusarium SCs (FCSC, FFSC, FIESC, FLSC, and FRSC) and Neocosmospora (Sandoval-Denis et al. 2019, Crous et al. 2021). Nevertheless, an accurate morphological analysis will provide a confirmation for DNA-based identifications and contribute useful means for strain characterization when dealing with lesser studied species complexes (e.g., the FTSC) for which, despite having a large amount of DNA data available, their taxonomy is still unrevised, and assignment of species to phylogenetic clades remains unclear (Laraba et al. 2022).
Multilocus sequence typing is, arguably, the most accurate, fastest and hence recommended identification procedure, especially for groups rich in cryptic diversity as the FSAMSC. Several DNA markers are suitable for this purpose, and 850 novel gene sequences were generated in this study including sequences for the highly efficient and commonly employed Fusarium markers rpb2 and tef1 (O’Donnell et al. 2013, Crous et al. 2021). Sequences for these markers are currently available for all ex-types in FSAMSC, many of which were newly generated here (Table 1, but also see Crous et al. 2021, Laraba et al. 2021), Single gene analyses based on rpb2 or tef1 sequences outperform any other available DNA marker for the FSAMSC, resolving 61 and 62 of the currently known species, respectively; however, these markers are not suitable, either alone or in combination, to resolve all segregate species of F. graminearum s.l., nor all the cryptic species in the Longipes clade. Similarly, analyses based on rpb1 confidently identified 58 of the known FSAMSC species, and sequences for this marker were also made available here for all the ex-types (Table 1). However, rpb1 PCR amplification and sequencing can be problematic for this marker, hence making it less suitable for routine identification purposes. Calmodulin and tub2 sequences were incorporated here, although achieving considerably less resolution than the above-mentioned loci, identifying 38 and 41 species, respectively. Nevertheless, unlike the rpb1, rpb2, or tef1 datasets, the CaM and tub2 alignments in this study contained a proportion of missing data due to inaccessibility to living strains, which partially explains this lower resolution. Both CaM and tub2 are easily amplified and aligned, but analogous to other Fusarium species complexes (i.e., FCSC and FIESC, O’Donnell et al. 2009) and Neocosmospora (O’Donnell 2000, O’Donnell et al. 2008a), putative paralogous tub2 gene sequences were randomly amplified in FSAMSC, which can be partially overcome with the use of the alternative forward PCR primer BT2a (Glass & Donaldson 1995). Unfortunately, this will impact on sequence informativeness by eliminating two informative intron regions. Han et al. (2023), recommended histone 3 gene (his3 or H3) sequences to be incorporated in phylogenetic analyses of the FSAMSC. Although this marker was not tested in our dataset, it has shown good performance for resolving species in Nectriaceae, including fusarioid fungi (Lombard et al. 2015); however, his3 sequences are scarce for Fusarium spp. in public databases, including FSAMSC ex-type strains.
Most species in FSAMSC are proven producers of mycotoxins of diverse classes, mainly a variety of trichothecenes, lactones and sesquiterpenes. This is of particular importance for the FSAMSC due to its marked fondness for cereal hosts, and the deleterious influence of these compounds over the food and feed supply (Nelson et al. 2004, Desjardin & Proctor 2007). Mycotoxin profiling may also provide additional taxonomic information, as for instance, among all Fusarium species complexes, trichothecene biosynthetic gene clusters have only been reported from FSAMSC and FIESC (O’Donnell et al. 2018, and references therein), unlike morphologically similar species in the FTSC (O’Donnell et al. 2018). Reports of trichothecene production by members of FTSC (F. acuminatum, Ueno et al. 1975, Logrieco et al. 1992) are thought to refer to the former subspecies F. armeniacum (Fusarium acuminatum subsp. armeniacum), now allocated to the FSAMSC (O’Donnell et al. 2018, Laraba et al. 2022). Within the FSAMSC, mycotoxin profiles are available for 72 species, including the undescribed phylogenetic species FSAMSC2, FSAMSC7, FSAMSC11, FSAMSC13, and FSAMSC31, and have been discussed extensively elsewhere (Thrane & Hansen 1995, O’Donnell et al. 2018, Laraba et al. 2021). Laraba et al. (2021) demonstrated that, with the exception of F. lunulosporum, for which only the lactone zearalenone has been reported, all species in the Graminearum clade, including all segregates of F. graminearum s.l. produce at least one type of tricothecene, mostly nivalenol (type A trichothecene) and derivatives of deoxynivalenol (type B trichothecenes). With the exception of F. lousianense (only nivalenol reported), these tricothecenes can be accompanied by either a sesquiterpene (culmorin and derivatives), a lactone (zearalenone or butenolide), a depsipeptide (beauvericin, reported for F. graminearum), a polyphenol (bikaverin, reported for F. culmorum), and a polyketide (fusarin C) or a enniatin (enniatin A1) as reported for F. subtropicale. In contrast, for most of the remaining species in FSAMSC, type A trichothecenes (particularly diacetoxyscirpenol, HT-2 and T-2 toxins; or neosolaniol), while a variety oflactones, sesquiterpenes, depsipeptides, polyketides and enniatins have been reported. Only a few known species are regarded as non-toxigenic (i.e., F. carinatum, F. cygneum, F. longicolle, F. pratense, and F. vermicularioides), all of which cluster whitin the Longipes clade. However, it is currently unknown whether the three additional lineages described here (F. bananae, F. procumbrens, and F. pseudolongipes) and the recently described F. guizhouense (He et al. 2024) possess the necessary mycotoxin biosynthetic capability. Although their significance as mycotoxigenic species remains to be tested, F. bananae and F. guizhuense are both members of the Sambucinum clade of FSAMSC, and are likely to produce type A trichothecenes, a common feature of species in this clade, while depsipeptides (beauvericin), enniatins, lactones, and sesquiterpene mycotoxins are also widely distributed (Laraba et al. 2021). This includes the undescribed lineage FSAMSC11, a T-2 toxin producing species and the closest phylogenetic relative of F. guizhuense. However, FSAMSC11 is so far known from a single strain (Thrane & Hansen 1995, Laraba et al. 2021), while F. guizhuenze has not been tested for this matter (Laraba et al. 2021). Conversely, F. pseudolongipes and F. procumbens belong to the Longipes clade, where most species do not produce mycotoxins (Laraba et al. 2021). Exceptions include the trichothecene producers F. longipes, F. mastigosporum, which also produces beauvericin; and F. tropicale (Laraba et al. 2021). In general, it is not known if any of the species formally named here pose an immediate risk for human and animal health, but it is expected that with improved means for identification and communication it will become easier to record and track this in future.
Often, species of Fusarium fail to satisfy all species concepts (morphological, biological, ecological or phylogenetic) (Leslie & Bowden 2008), and hence some degree of discordance can be expected among the different methods. Here, we introduced coalescent species delimitation methods in Fusarium taxonomy, following up on recent integration of these methods to fungal taxonomy (Løken et al. 2019, Bian et al. 2022, Glässnerová et al. 2022, Réblová et al. 2022, Sklenář et al. 2022). Nevertheless, coalescent-based species delimitation methods alone are not expected to fully discern linage diversification processes, and results should be weighted with other sources of data (Fujita et al. 2012, Luo et al. 2018). Therefore, we incorporated these analyses to provide additional measurements to further support or fine-tune species hypotheses in a particularly complex group of fungi which has suffered from nearly century-old conflictive taxonomical approaches (Wollenweber 1913, 1945, Snyder & Hansen 1945, O’Donnell 1993, 1996, 1997). Final species delimitations in this work relied on the congruence of data from all sources, including phylogenetic, morphological, and coalescent-based algorithms. Single-gene coalescent species delimitation methods have been found to reflect locus variability or informativeness, with more variable markers resulting in more species groups (Vitecek et al. 2017). Moreover, groups obtained by these methods may not be statistically demonstrable to originate due to population isolation or real species boundaries (Sukumaran & Knowles 2017), and may be affected by gene flow between distant species, a known phenomenon in Fusarium (Leslie & Summerell 2006, Leslie & Bowden 2008, Luo et al. 2018). Although some degree of congruence with phylogenetically and morphologically identifiable species was found in this study, the sometimes inconsistent over-partitioning (e.g., F. graminearum, F. poae, F. sambucinum, F. sporotrichioides, among others) and over-aggregation (e.g., F. culmorum and F. cerealis, F. graminearum segregates, F. palustre and F. mastigosporum, F. sporotrichioides and F. sibiricum, among others) of species groups; plus the considerable difference of number of species groups obtained between methods, reduced the taxonomical informativeness of single-gene coalescent species delimitation analyses. Model-based, multi-locus coalescent methods (i.e., STACEY) integrate all available gene information and should be preferred (Rannala 2015, Manzo et al. 2024). In our dataset, STACEY analyses suggested 77 groups, 75 of which were equivalent to the 75 species as determined by phylogenetic and morphological analyses. Interestingly, STACEY results also suggested additional species groups within F. graminearum s. str. Although there is evidence of additional undescribed lineages in Fusarium graminearum s.lat. (O’Donnell et al. 2018, Laraba et al. 2021), we refrained to take any taxonomical action in this regard, following the morphological and phylogenetic evidence. Additionally, STACEY results provided supplementary evidence to support the status of species for lineages represented by single isolates (i.e., F. bananae, F. carinatum, F. cygneum, F. subcylindroides, F. vermicularioides, and the undescribed phylogenetic species FSAMSC11 and FSAMSC31). It is expected that STACEY may inflate the number of species groups when conditioned by a limited number of individuals (Toprak et al. 2016, Sukumaran & Knowles 2017, Løken et al. 2019). However, the same groups were also confidently determined in this study by both phylogenetic and morphological analyses.
Several phylogenetic lineages in this study are known from single (see above) or very few observations. Hence, their currently known biogeographic patterns, host distributions, and mycotoxigenic profiles are most likely incomplete. This is vital information for the design of quarantine policies, trade regulations, and management strategies. Especially for the grain-loving taxa in FSAMSC. We therefore urge researchers and citizen scientists to collect fresh, well-annotated living cultures and to deposit living material and DNA sequence data in public databases and collections.
ACKNOWLEDGEMENTS
We want to thank the curators and personnel of the following fungaria and culture collections for sending specimens and help locating specimen data: M. Hernández-Restrepo, T. Merkx, A. van Iperen and G. Verkley (CBS); M. Price, J.C. Zamora and I. Valette (herbarium G); L. Davies and R. Woods (herbarium KEW); T. Iturriaga (CUP), T.W. Adkins, J. Swezey and T. Ward (NRRL). We thank I. Pavlov and O. Yarden for sending fungal strains. And we thank K. Bensch and K.A. Seifert for commenting on nomenclatural issues. This work was in part supported by the U.S. Department of Agriculture – Agricultural Research Service and the Dutch NWO Roadmap grant agreement No. 2020/ENW/00901156, project “Netherlands Infrastructure for Ecosystem and Biodiversity Analysis – Authoritative and Rapid Identification System for Essential biodiversity information” (acronym NIEBA-ARISE).
COMPETING INTERESTS
The authors have declared that no competing interests exist. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the U.S. Department of Agriculture. USDA is an equal opportunity provider and employer.
Supplementary Material: https://studiesinmycology.org/
Morphological characteristics of Fusarium species included in this study.
REFERENCES
- Akinsanmi OA, Chakraborty S, Backhouse D, et al. (2007). Passage through alternative hosts changes the fitness of Fusarium graminearum and Fusarium pseudograminearum. Environmental Microbiology 9: 512–520. [DOI] [PubMed] [Google Scholar]
- Al-Hatmi A. (2016). Phylogeny, diagnostics and antifungal susceptibility of clinically relevant Fusarium species. Ph.D. dissertation. Faculty of Science, Mathematics and Computer Science, University of Amsterdam, The Netherlands.
- Aoki T, O’Donnell K. (1998). Fusarium kyushuense sp. nov. from Japan. Mycoscience 39: 1–6. [Google Scholar]
- Aoki T, O’Donnell K. (1999a). Morphological and molecular characterization of Fusarium pseudograminearum sp. nov., formerly recognized as the Group 1 population of F. graminearum. Mycologia 91: 597–609. [Google Scholar]
- Aoki T, O’Donnell K. (1999b). Morphological characterization of Gibberella coronicola sp. nov., obtained through mating experiments of Fusarium pseudograminearum. Mycoscience 40: 443–453. [Google Scholar]
- Aoki T, Ward TJ, Kistler HC, et al. (2012). Systematics, phylogeny and trichothecene mycotoxin potential of Fusarium head blight cereal pathogens. JSM Mycotoxins 62: 91–102. [Google Scholar]
- Bhunjun CS, Chen YJ, Phukhamsakda C, et al. (2024). What are the 100 most cited fungal genera? Studies in Mycology 108: 1–411. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Bian C, Kusuya Y, Sklenář F, et al. (2022). Reducing the number of accepted species in Aspergillus series Nigri. Studies in Mycology 102: 95–132. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Bilai VI. (1955). The fusaria (biology and systematics). Akademia Naukwe Ukrainskii, SSR, Kiev. [Google Scholar]
- Booth C. (1971). The genus Fusarium. Commonwealth Mycology Institute, Kew, England. [Google Scholar]
- Bouckaert R, Vaughan TG, Barido-Sottani J, et al. (2019). BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Computational Biology 15: e1006650. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Bowden RL, Leslie JF. (1999). Sexual recombination in Gibberella zeae (Fusarium graminearum). Phytopathology 89: 182–188. [DOI] [PubMed] [Google Scholar]
- Brondeau L. (1855). Observations pour servir á la classification des genres Fusarium et Fusidium de Link. Archives de Flore de France et D’Allemagne 1: 167–171. [Google Scholar]
- Burgess LW, Forbes GA, Windels C, et al. (1993). Characterization and distribution of Fusarium acuminatum subsp. armeniacum subsp. nov. Mycologia 85: 119–124. [Google Scholar]
- CAB International, Commonwealth Mycological Institute (CMI) (1964). Fusarium culmorum. Descriptions of Pathogenic Fungi and Bacteria. No. 26, Kew Surrey, UK. [Google Scholar]
- CAB International, Commonwealth Mycological Institute (CMI) (1973). Gibberella zeae. Descriptions of Pathogenic Fungi and Bacteria No. 384, Kew Surrey, UK. [Google Scholar]
- Carbone I, Kohn LM. (1999). A method for designing primer sets for speciation studies in filamentous ascomycetes. Mycologia 91: 553–556. [Google Scholar]
- Costa MM, Sandoval-Denis M, Moreira GM, et al. (2024). Known from trees and the tropics: new insights into the Fusarium lateritium species complex. Studies in Mycology 109: 403–450. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Council for Agricultural Science and Technology (CAST) (2003). Mycotoxins: risks in plant, animal, and human systems, task force report No. 139, CAST, Ames, Iowa, USA, 2003. [Google Scholar]
- Crous PW, Lombard L, Sandoval-Denis M, et al. (2021). Fusarium: more than a node or a foot-shaped basal cell. Studies in Mycology 98: 100116. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Crous PW, Osieck ER, Jurjevi Ž, et al. (2021). Fungal Planet description sheets: 1284–1382. Persoonia 47: 178–374. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Crous PW, Verkley GJM, Groenewald JZ, et al. (2019). Westerdijk Laboratory Manual Series 1: Fungal Biodiversity. Westerdijk Fungal Biodiversity Institute, Utrecht, The Netherlands. [Google Scholar]
- Dean R, Van Kan JAL, Pretorius ZA, et al. (2012). The top 10 fungal pathogens in molecular plant pathology. Molecular Plant Pathology 13: 414–430. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Desjardinds AE. (2003). Gibberella from A (venaceae) to Z (eae). Annual Review of Phytopathology 41: 177–198. [DOI] [PubMed] [Google Scholar]
- Desjardins AE, Beremand M. (1987). A genetic system for trichothecene toxin production in Gibberella pulicaris (Fusarium sambucinum). Phytopathology 77: 678–863. [Google Scholar]
- Desjardins AE, Proctor RH. (2003). Molecular biology of Fusarium mycotoxins. International Journal of Food Microbiology 119: 47–50. [DOI] [PubMed] [Google Scholar]
- Dhlamini L, Jacobs A, van der Merwe NA, et al. (2024). The population structure of the Fusarium brachygibbosum species complex in the grassland biome of South Africa. Australasian Plant Pathology 53: 31–39. [Google Scholar]
- Domsch KH, Gams W, Anderson TH. (2007). Compendium of soil fungi, ed 2. IHW Verlag, Eching. [Google Scholar]
- EFSA Panel on Plant Health (PLH) Bragard C, Baptista P, et al. (2022). Scientific opinion on the pest categorisation of Fusarium pseudograminearum. EFSA Journal 20: 7399. [DOI] [PMC free article] [PubMed] [Google Scholar]
- EFSA PLH Panel (EFSA Panel on Plant Health) Bragard C, Di Serio F, et al. (2021). Scientific opinion on the pest categorisation of Fusarium brachygibbosum. EFSA Journal 19: 6887. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Elmer WH, Marra RE. (2011). New species of Fusarium associated with dieback of Spartina alterniflora in Atlantic salt marshes. Mycologia 103: 806–819. [DOI] [PubMed] [Google Scholar]
- Farr DF, Rossman AY, Castlebury LA. (2023). United States National Fungus Collections fungus-host dataset. https://agdatacommons.nal.usda.gov/articles/dataset/United_States_National_Fungus_Collections_Fungus-Host_Dataset/24855585/1. Accessed December 2023.
- Fisher NL, Burgess LW, Toussoun TA, et al. (1982). Carnation leaves as a substrate and for preserving cultures of Fusarium species. Phytopathology 72: 151–153. [Google Scholar]
- Fries EM. (1832). Systema Mycologicum 3 (2): i-ii, 261-524. Greifswald; Sumptibus Ernesti Mauritii. [Google Scholar]
- Fujita MK, Leaché AD, Burbrink FT, et al. (2012). Coalescent-based species delimitation in an integrative taxonomy. Trends in Ecology and Evolution 27: 480–488. [DOI] [PubMed] [Google Scholar]
- Gams W. (1999). Report of the committee for fungi: 8. Taxon 48: 807–810. [Google Scholar]
- Gams W, Nirenberg HI, Seifert KA, et al. (1997). (1275) Proposal to conserve the name Fusarium sambucinum (Hyphomycetes). Taxon 46: 111–113. [Google Scholar]
- Gilbert J, Haber S. (2013). Overview of some recent research developments in fusarium head blight of wheat. Canadian Journal of Plant Pathology 35: 149–174. [Google Scholar]
- Glass NL, Donaldson GC. (1995). Development of primer sets designed for use with the PCR to amplify conserved genes from filamentous ascomycetes. Applied and Environmental Microbiology 61: 1323–1330. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Glässnerová K, Sklenář F, Jurjević Ž, et al. (2022). A monograph of Aspergillus section Candidi. Studies in Mycology 102: 1–51. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Glez-Peña D, Gómez-Blanco D, Reboiro-Jato M, et al. (2010). ALTER: program-oriented conversion of DNA and protein alignments. Nucleic Acids Research 38: W14–8. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Gordon WL. (1952). The occurrence of Fusarium species in Canada. II. Prevalence and taxonomy of Fusarium species in cereal seeds. Canadian Journal of Botany 30: 209–251. [Google Scholar]
- Gordon WL. (1954a). The occurrence of Fusarium species in Canada III. Taxonomy of Fusarium species in the seed of vegetable forage, and miscellaneous crops. Canadian Journal of Botany 32: 576–590. [Google Scholar]
- Gordon WL. (1954b). The occurrence of Fusarium species in Canada IV. Taxonomy and prevalence of Fusarium species in the soil of cereal plots. Canadian Journal of Botany 32: 622–629. [Google Scholar]
- Gräfenhan T, Johnston PR, Vaughan MM, et al. (2017). Fusarium praegraminearum sp. nov., a novel nivalenol mycotoxin-producing pathogen from New Zealand can induce head blight on wheat. Mycologia 108: 1229–1239. [DOI] [PubMed] [Google Scholar]
- Han SL, Wang MM, Ma ZY, et al. (2023). Fusarium diversity associated with diseased cereals in China, with an updated phylogenomic assessment of the genus. Studies in Mycology 104: 87–148. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Hill R, Buggs RJA, Vu DT, et al. (2022). Lifestyle transitions in fusarioid fungi are frequent and lack clear genomic signatures. Molecular Biology and Evolution 39: msac085. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Hoang DT, Chernomor O, von Haeseler A, et al. (2018). UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35: 518–522. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Jones G. (2017). Algorithmic improvements to species delimitation and phylogeny estimation under the multispecies coalescent. Journal of Mathematical Biology 74: 447–467. [DOI] [PubMed] [Google Scholar]
- Jones G. (2019). STACEY package documentation: species delimitation and species tree estimation with BEAST2. Operative manual distributed with the software.
- Jurgenson JE, Bowden RL, Zeller KA, et al. (2002). A genetic map of Gibberella zeae (Fusarium graminearum). Genetics 160: 1451–1460. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Kalyaanamoorthy S, Minh BQ, Wong TKF, et al. (2017). ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Katoh K, Standley DM. (2013). MAFFT Multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Klittich C, Leslie JF. (1988). Nitrate reduction mutants of Fusarium moniliforme. Genetics 118: 417–423. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Laraba I, Busman M, Geiser DM, et al. (2022). Phylogenetic diversity and mycotoxin potential of emergent phytopathogens within the Fusarium tricinctum species complex. Phytopathology 112: 1284–1298. [DOI] [PubMed] [Google Scholar]
- Laraba I, McCormick SP, Vaughan MM, et al. (2021). Phylogenetic diversity, trichothecene potential, and pathogenicity within Fusarium sambucinum species complex. PLoS ONE 16: e0245037. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Laurence MH, Walsh JL, Shuttleworth LA, et al. (2015). Six novel species of Fusarium from natural ecosystems in Australia. Fungal Diversity 77: 349–366. [Google Scholar]
- Leslie JF, Anderson LL, Bowden RL, et al. (2007). Inter- and intra-specific genetic variation in Fusarium. International Journal of Food Microbiology 119: 25–32. [DOI] [PubMed] [Google Scholar]
- Leslie JF, Bowden RL. (2008). Fusarium graminearum: when species concepts collide. Cereal Research Communications 36: 609–615. [Google Scholar]
- Leslie JF, Summerell BA. (2006). The Fusarium laboratory manual. Blackwell Publishing, Ames. [Google Scholar]
- Lewis CE. (1913). Comparative studies of certain disease-producing species of Fusarium. Bulletin of the Maine Agricultural Experimental Station 219: 203–258. [Google Scholar]
- Link HF. (1809). Observationes in ordines plantarum naturales. Dissertatio Ima. Magazin für die Neuesten Entdeckungen in der Gesammten Naturkunde, Gesellschaft Naturforschender Freunde zu Berlin 3: 3–42. [Google Scholar]
- Liu YJ, Whelen S, Hall BD. (1999). Phylogenetic relationships among ascomycetes: evidence from an RNA polymerase II subunit. Molecular Biology and Evolution 16: 1799–1808. [DOI] [PubMed] [Google Scholar]
- Logrieco A, Altomare C, Moretti A, et al. (1992). Cultural and toxigenic variability in Fusarium acuminatum. Mycological Research 96: 518–523. [Google Scholar]
- Løken SB, Skrede I, Schumacher T. (2019). The Helvella corium species aggregate in Nordic countries – phylogeny and species delimitation. Fungal Systematics and Evolution 5: 169–186. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Lombard L, van Doorn R, Crous PW. (2019). Neotypification of Fusarium chlamydosporum - a reappraisal of a clinically important species complex. Fungal Systematics and Evolution 4: 183–200. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Lombard L, van der Merwe NA, Groenewald JZ, et al. (2015). Generic concepts in Nectriaceae. Studies in Mycology 80: 189–245. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Lombard L, van Doorn R, Groenewald JZ, et al. (2022). Fusarium diversity associated with the Sorghum-Striga interaction in Ethiopia. Fungal Systematics and Evolution 10: 177–215. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Luo A, Ling C, Ho SYW, et al. (2018). Comparison of methods for molecular species delimitation across a range of speciation scenarios. Systematic Biology 67: 830–846. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Lysøe E, Frandsen RJN, Divon HH, et al. (2016). Draft genome sequence and chemical profiling of Fusarium langsethiae, an emerging producer of type A trichothecenes. International Journal of Food Microbiology 221: 29–36. [DOI] [PubMed] [Google Scholar]
- Manzo E, Nillies S, Gutiérrez DG, et al. (2024). Species delimitation in Xanthium sect. Acanthoxanthium (Heliantheae, Asteraceae) and the neglected species Xanthium argenteum. Taxon 73: 1251–1271. [Google Scholar]
- Marasas WFO, Rheeder JP, Logrieco A, et al. (1998). Fusarium nelsonii and F. musarum: two new species in Section Arthrosporiella related to F. camptoceras. Mycologia 90: 505–513. [Google Scholar]
- McMullen M, Bergstrom G, De Wolf E, et al. (2012). A unified effort to fight an enemy of wheat and barley: Fusarium head blight. Plant Disease 96: 1712–1728. [DOI] [PubMed] [Google Scholar]
- Miller MA, Pfeiffer W, Schwartz T. (2010). Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, LA pp. 1–8.
- Nelson PE, Dignani MC, Anaissie EJ. (1994). Taxonomy, biology, and clinical aspects of Fusarium species. Clinical Microbiology Reviews 7: 479–504. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Nelson PE, Toussoun TA, Marasas WFO. (1983). Fusarium species: an illustrated manual for identification. Pennsylvania State University Press, USA. [Google Scholar]
- Nguyen LT, Schmidt HA, von Haeseler A, et al. (2015). IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32: 268–274. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Nichea MJ, Proctor RH, Probyn CE, et al. (2021). Fusarium chaquense, sp. nov., a novel type A trichothecene–producing species from native grasses in a wetland ecosystem in Argentina. Mycologia 114: 46–62. [DOI] [PubMed] [Google Scholar]
- Nirenberg HI. (1976). Untersuchungen über die morphologische und biologische Differenzierung in der Fusarium-Sektion Liseola. Mitteilungen der Biologischen Bundesanstalt für Land- und Forstwirtschaft Berlin-Dahlem 169: 1–117. [Google Scholar]
- Nirenberg HI. (1990). Recent advances in the taxonomy of Fusarium. Studies in Mycology 32: 91–101. [Google Scholar]
- Nirenberg HI. (1995). Morphological differentiation of Fusarium sambucinum Fuckel sensu stricto, F. torulosum (Berk. & Curt.) Nirenberg comb. nov. and F. venenatum Nirenberg sp. nov. Mycopathologia 129: 131–141. [DOI] [PubMed] [Google Scholar]
- Nylander JAA. (2004). MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala University. [Google Scholar]
- O’Donnell K. (1993). Fusarium and its near relatives. In: The fungal holomorph: mitotic, meiotic and pleomorphic speciation in fungal systematics (Reynolds DR, Taylor JW, eds.). CAB International, Wallingford. [Google Scholar]
- O’Donnell K. (1996). Progress towards a phylogenetic classification of Fusarium. Sydowia 48: 57–70. [Google Scholar]
- O’Donnell K. (1997). Phylogenetic evidence indicates the important mycotoxigenic strains Fn-2, Fn-3, Fn-2B and Fn-M represent a new species of Fusarium. Mycotoxins 45: 1–10. [Google Scholar]
- O’Donnell K. (2000). Molecular phylogeny of the Nectria haematococca-Fusarium solani species complex. Mycologia 92: 919–938. [Google Scholar]
- O’Donnell K, Cigelnik E, Casper HH. (1998. a). Molecular phylogenetic, morphological, and mycotoxin data support reidentification of the Quorn mycoprotein fungus as Fusarium venenatum. Fungal Genetics and Biology 23: 57–67. [DOI] [PubMed] [Google Scholar]
- O’Donnell K, Sutton DA, Fothergill A, et al. (2008a). Molecular phylogenetic diversity, multilocus haplotype nomenclature, and in vitro antifungal resistance within the Fusarium solani species complex. Journal of Clinical Microbiology 46: 2477–2490. [DOI] [PMC free article] [PubMed] [Google Scholar]
- O’Donnell K, Cigelnik E. (1997). Two divergent intragenomic rDNA ITS2 types within a monophyletic lineage of the fungus Fusarium are nonorthologous. Molecular Phylogenetics and Evolution 7: 103–116. [DOI] [PubMed] [Google Scholar]
- O’Donnell K, Humber RA, Geiser DM, et al. (2012). Phylogenetic diversity of insecticolous fusaria inferred from multilocus DNA sequence data and their molecular identification via FUSARIUM-ID and Fusarium MLST. Mycologia 104: 427–445. [DOI] [PubMed] [Google Scholar]
- O’Donnell K, Kistler HC, Cigelnik E. (1998b). Multiple evolutionary origins of the fungus causing Panama disease of banana: Concordant evidence from nuclear and mitochondrial gene genealogies. Proceedings of the National Academy of Sciences of the USA 95: 2044–2049. [DOI] [PMC free article] [PubMed] [Google Scholar]
- O’Donnell K, Kistler HC, Tacke BK, et al. (2000). Gene genealogies reveal global phylogeographic structure and reproductive isolation among lineages of Fusarium graminearum, the fungus causing wheat scab. Proceedings of the National Academy of Sciences of the United States of America 97: 7905–7910. [DOI] [PMC free article] [PubMed] [Google Scholar]
- O’Donnell K, McCormick SP, Busman M, et al. (2018). Marasas et al. 1984 “toxigenic Fusarium species: identity and mycotoxicology” revisited. Mycologia 110: 1058–1080. [DOI] [PubMed] [Google Scholar]
- O’Donnell K, Rooney AP, Proctor RH, et al. (2013). Phylogenetic analyses of RPB1 and RPB2 support a middle Cretaceous origin for a clade comprising all agriculturally and medically important fusaria. Fungal Genetics and Biology 52: 20–31. [DOI] [PubMed] [Google Scholar]
- O’Donnell K, Sutton DA, Rinaldi MG, et al. (2009). Novel multilocus sequence typing scheme reveals high genetic diversity of human pathogenic members of the Fusarium incarnatum-F. equiseti and F. chlamydosporum species complexes within the United States. Journal of Clinical Microbiology 47: 3851–3861. [DOI] [PMC free article] [PubMed] [Google Scholar]
- O’Donnell K, Sutton DA, Rinaldi MG. (2010). Internet-accessible DNA sequence database for identifying fusaria from human and animal infections. Journal of Clinical Microbiology 48: 3708–3718. [DOI] [PMC free article] [PubMed] [Google Scholar]
- O’Donnell K, Sutton DA, Wiederhold N, et al. (2016). Veterinary fusarioses within the United States. Journal of Clinical Microbiology 54: 2813–2819. [DOI] [PMC free article] [PubMed] [Google Scholar]
- O’Donnell K, Ward TJ, Aberra D, et al. (2008b). Multilocus genotyping and molecular phylogenetics resolve a novel head blight pathogen within the Fusarium graminearum species complex from Ethiopia. Fungal Genetics and Biology 45: 1514–1522. [DOI] [PubMed] [Google Scholar]
- O’Donnell K, Ward TJ, Geiser DM, et al. (2004). Genealogical concordance between the mating type locus and seven other nuclear genes supports formal recognition of nine phylogenetically distinct species within the Fusarium graminearum clade. Fungal Genetics and Biology 41: 600–623. [DOI] [PubMed] [Google Scholar]
- Padwick GW. (1945). Notes on Indian fungi III. Mycological Papers 12: 1–15. [Google Scholar]
- Pascoe IG. (1990). Fusarium morphology I: identification and characterisation of a third conidial type, the mesoconidium. Mycotaxon 27: 121–160. [Google Scholar]
- Pereira CB, Ward TJ, Tessmann DJ, et al. (2018). Fusarium subtropicale, sp. nov., a novel nivalenol mycotoxin–producing species isolated from barley (Hordeum vulgare) in Brazil and sister to F. praegraminearum. Mycologia 110: 860–871. [DOI] [PubMed] [Google Scholar]
- Perera RH, Hyde KD, Jones EBG, et al. (2023). Profile of Bionectriaceae, Calcarisporiaceae, Hypocreaceae, Nectriaceae, Tilachlidiaceae, Ijuhyaceae fam. nov., Stromatonectriaceae fam. nov. and Xanthonectriaceae fam. nov. Fungal Diversity 118: 98–271. [Google Scholar]
- Polak-Śliwińska M, & Paszczyk B. (2021). Trichothecenes in food and feed, relevance to human and animal health and methods of detection: a systematic review. Molecules 26: 454. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Posada D, Crandall KA. (1998). MODELTEST: testing the model of DNA substitution. Bioinformatics 14: 817–818. [DOI] [PubMed] [Google Scholar]
- Puillandre N, Brouillet S, Achaz G. (2021). ASAP: assemble species by automatic partitioning. Molecular Ecology Resources 21: 609–620. [DOI] [PubMed] [Google Scholar]
- Puillandre N, Lambert A, Brouillet S, et al. (2012). ABGD, automatic barcode gap discovery for primary species delimitation. Molecular Ecology 21: 1864–1877. [DOI] [PubMed] [Google Scholar]
- Quaedvlieg W, Kema GHJ, Groenewald JZ. (2011). Zymoseptoria gen. nov.: a new genus to accommodate septoria-like species occurring on graminicolous hosts. Persoonia 26: 57–69. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Raillo A. (1950). Fungi of the genus Fusarium. State publishing house of agricultural literature, Moscow, USSR.
- Rannala B. (2015). The art and science of species delimitation. Current Zoology 61: 846–853. [Google Scholar]
- Rayner RW. (1970). A mycological colour chart. CMI and British Mycological Society, Kew, Surrey, UK. [Google Scholar]
- Rambaut A, Drummond AJ, Xie D, et al. (2018). Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67: 901–904. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Réblová M, Hernández-Restrepo M, Sklenář F, et al. (2022). Consolidation of Chloridium: new classification into eight sections with 37 species and reinstatement of the genera Gongromeriza and Psilobotrys. Studies in Mycology 103: 87–212. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Reinking OA, Wollenweber HW. (1927). Tropical fusaria. The Philippine Journal of Science 32: 103–253. [Google Scholar]
- Ronquist F, Teslenko M, van der Mark P, et al. (2012). MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Samuels GJ. (1976). A revision of the fungi formerly classified in Nectria subg. Hyphonectria. Memoires of the New York Botanical Garden 26: 1–126. [Google Scholar]
- Samuels GJ, Doi Y, Rogerson CT. (1990). Hypocreales. Memoirs of the New York Botanical Garden 59: 6–108. [Google Scholar]
- Samuels GJ, Rossman AY, Chaverri P, et al. (2006). Hypocreales of the Southeastern United States: an identification guide. Centrallbureau voor Schimmelcultures, Utrecht, The Netherlands.
- Sandoval-Denis M, Swart WJ, Crous PW. (2018). New Fusarium species from the Kruger National Park, South Africa. MycoKeys 34: 63–92. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Sangalang AE, Backhouse D, Burgess LW. (1995). Survival and growth in culture of four Fusarium species in relation to occurrence in soils from hot climatic regions. Mycological Research 99: 529–533. [Google Scholar]
- Sarver AJ, Ward TJ, Gale LR, et al. (2011). Novel Fusarium head blight pathogens from Nepal and Louisiana revealed by multilocus genealogical concordance. Fungal Genetics and Biology 48: 1096–1107. [DOI] [PubMed] [Google Scholar]
- Schwabe SH. (1839). Flora anhaltina, Tomus II. Apud Ge. Reimerum. Berlin, Germany. [Google Scholar]
- Seifert KA, Grafenhan T, Redhead SA. (2009). Typification of Fusarium cerealis and Fusarium crookwellense. Inoculum 60: 40. [Google Scholar]
- Sherbakoff CD. (1915). Fusaria of potatoes. Memoirs of the Cornell University Agricultural Experimental Station 6: 87–270. [Google Scholar]
- Silvestro D, Michalak I. (2012). RaxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution 12: 335–337. [Google Scholar]
- Sklenář F, Glässnerová K, Jurjević Ž, et al. (2022). Taxonomy of Aspergillus series Versicolores: species reduction and lessons learned about intraspecific variability. Studies in Mycology 102: 53–93. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Smith WG. (1884). Diseases of field and garden crops, chiefly such as are caused by fungi. Macmillan and co. London, UK. [Google Scholar]
- Snyder WC, Hansen HN. (1945). The species concept in Fusarium with reference to Discolor and other sections. American Journal of Botany 32: 657–666. [Google Scholar]
- Stamatakis A. (2006). RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2688–2690. [DOI] [PubMed] [Google Scholar]
- Starkey DA, Ward TJ, Aoki T, et al. (2007). Global molecular surveillance reveals novel Fusarium head blight species and trichothecene toxin diversity. Fungal Genetics and Biology 44: 1191–1204. [DOI] [PubMed] [Google Scholar]
- Subramanian CV. (1971). Hyphomycetes. An account of Indian species, except Cercosporae. Indian Council of Agricultural Research, New Delhi, India. [Google Scholar]
- Sukumaran J, Knowles LL. (2017). Multispecies coalescent delimits structure, not species. Proceedings of the National Academy of Sciences of the United States of America 114: 1607–1612. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Tamura K, Stecher G, Peterson D, et al. (2013) MEGA6: molecular genetics analysis version 6.0. Molecular Biology and Evolution 30: 2725–2729. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Thrane U, Hansen U. (1995) Chemical and physiological characteristics of taxa in the Fusarium sambucinum complex. Mycopathologia 129: 183–190. [DOI] [PubMed] [Google Scholar]
- Tiwari RK, Kumar R, Sharma S, et al. (2020). Potato dry rot disease: current status, pathogenomics and management. 3 Biotech 10: 503. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Toprak Z, Pfeil BE, Jones G, et al. (2016). Species delimitation without prior knowledge: DISSECT reveals extensive cryptic speciation in the Silene aegyptiaca complex (Caryophyllaceae). Molecular Phylogenetics and Evolution 102: 1–8. [DOI] [PubMed] [Google Scholar]
- Torp M, Nirenberg HI. (2004). Fusarium langsethiae sp. nov. on cereals in Europe. International Journal of Food Microbiology 95: 247–256. [DOI] [PubMed] [Google Scholar]
- Trail F. (2009) For blighted waves of grain: Fusarium graminearum in the postgenomics era. Plant Physiology 149: 103–110. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Ueno Y, Sawano M, Ishii K. (1975). Production of trichothecene mycotoxins by Fusarium species in shake culture. Applied Microbiology 30: 4–9. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Vaidya G, Lohman DJ, Meier R. (2011). SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics 27: 171–180. [DOI] [PubMed] [Google Scholar]
- Visagie CM, Yilmaz N, Allison JD, et al. (2024). New and interesting fungi. 7. Fungal Systematics and Evolution 13: 441–494. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Vitecek S, Kučinić M, Previšić A, et al. (2017). Integrative taxonomy by molecular species delimitation: multi-locus data corroborate a new species of Balkan Drusinae micro-endemics. BMC Evolutionary Biology 17: 129. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Wang MM, Crous PW, Sandoval-Denis M, et al. (2022). Fusarium and allied genera from China: species diversity and distribution. Persoonia 48: 1–53. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Wollenweber HW. (1913). Studies on the Fusarium problem. Phytopathology 3: 24–50. [Google Scholar]
- Wollenweber HW. (1916). Fusaria autographice delineata 1: 1–509. Berlin. Germany. [Google Scholar]
- Wollenweber HW. (1917). Fusaria autographice delineata. Annales Mycologici 15: 1–56. [Google Scholar]
- Wollenweber HW. (1918). Conspectus analyticus fusariorum. Berichte der Deutschen Botanischen Gesellschaft 35: 732–742. [Google Scholar]
- Wollenweber HW. (1930). Fusaria autographice delineata 3: 660–1100. Berlin. Germany. [Google Scholar]
- Wollenweber HW. (1931). Fusarium – monographie. Fungi parasitici et saprophytici Zeitschrift für Parasitenkunde 3: 260–516. [Google Scholar]
- Wollenweber HW, Reinking OA. (1935). Die fusarien. Verlagsbuchandlung Paul Parey, Berlin, Germany. [Google Scholar]
- Woudenberg JHC, Aveskamp MM, de Gruyter J. (2009). Multiple Didymella teleomorphs are linked to the Phoma clematidina morphotype. Persoonia 22: 56–62. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Xia JW, Sandoval-Denis M, Crous PW, et al. (2019). Numbers to names - restyling the Fusarium incarnatum-equiseti species complex. Persoonia 43: 186–221. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Yli-Mattila T, Ward TJ, O’Donnell K, et al. (2011). Fusarium sibiricum sp. nov., a novel type A trichothecene-producing Fusarium from northern Asia closely related to F. sporotrichioides and F. langsethiae. International Journal of Food Microbiology 147: 58–68. [DOI] [PubMed] [Google Scholar]
- Zhang J, Kapli P, Pavlidis P, et al. (2013). A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29: 2869–2876. [DOI] [PMC free article] [PubMed] [Google Scholar]
Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Morphological characteristics of Fusarium species included in this study.