Epitypification of Fusarium oxysporum – clearing the taxonomic chaos

Abstract

Fusarium oxysporum is the most economically important and commonly encountered species of Fusarium. This soil-borne fungus is known to harbour both pathogenic (plant, animal and human) and non-pathogenic strains. However, in its current concept F. oxysporum is a species complex consisting of numerous cryptic species. Identification and naming these cryptic species is complicated by multiple subspecific classification systems and the lack of living ex-type material to serve as basic reference point for phylogenetic inference. Therefore, to advance and stabilise the taxonomic position of F. oxysporum as a species and allow naming of the multiple cryptic species recognised in this species complex, an epitype is designated for F. oxysporum. Using multi-locus phylogenetic inference and subtle morphological differences with the newly established epitype of F. oxysporum as reference point, 15 cryptic taxa are resolved in this study and described as species.

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Persoonia 43, 2019: 1– 47 ISSN (Online) 1878-9080
www.ingentaconnect.com/content/nhn/pimj https://doi.org/10.3767/persoonia.2019.43.01
RESEARCH ARTICLE
INTRODUCTION
Fusarium oxysporum is the most economically important and
commonly encountered species of Fusarium. This soil-borne
asexual fungus is known to harbour both pathogenic (plant,
animal and human) and non-pathogenic strains (Leslie & Sum-
merell 2006) and is also ranked fifth on a list of top 10 fungal
pathogens based on scientific and economic importance (Dean
et al. 2012, Geiser et al. 2013). Historically, F. oxysporum has
been defined by the asexual phenotype as no sexual morph
has yet been discovered, even though several studies have
indicated the possible presence of a cryptic sexual cycle (Arie
et al. 2000, Yun et al. 2000, Aoki et al. 2014, Gordon 2017).
This is further supported by phylogenetic studies that place
F. oxysporum within the Gibberella Clade (Baayen et al. 2000,
O’Donnell et al. 2009, 2013). These studies also showed that
F. oxysporum displays a complicated phylogenetic substruc-
ture, indicative of multiple cryptic species within F. oxysporum
(Gordon & Martyn 1997, Laurence et al. 2014). As with other
Fusarium species complexes, the F. oxysporum species com-
plex (FOSC) has suffered from multiple taxonomic/ classification
systems applied in the past.
Diederich F.L. von Schlechtendal first introduced F. oxysporum
in 1824, isolated from a rotten potato tuber (Solanum tubero-
sum) collected in Berlin, Germany. Wollenweber (1913) placed
F. oxysporum within the section Elegans along with eight other
Fusarium species and numerous varieties and forms based
on similarity of the micro- and macroconidial morphology and
dimensions. Snyder & Hansen (1940) later consolidated and
reduced all species within the section Elegans into F. oxysporum
and designated 25 special forms (formae speciales) within this
species. These special forms were further expanded on by Gor-
don (1965) to 66, most of which are still used in literature today.
The use of special forms or formae speciales as subspecific
rank in F. oxysporum classification has become common prac-
tice due to the broad morphological delineation of this species
(Leslie & Summerell 2006). This informal subspecific rank is
defined based on the plant pathogenicity of the particular F. oxy-
sporum strain and excludes both clinical and non-pathogenic
strains (Armstrong & Armstrong 1981, Gordon & Martyn 1997,
Kistler 1997, Baayen et al. 2000, Leslie & Summerell 2006).
Therefore, F. oxysporum strains attacking the same plant host
are generally considered to belong to the same special form.
Although this homologous trait has led to erroneous assump-
tions considering a specific special form to be phylogenetically
monophyletic, several studies (O’Donnell et al. 1998, 2004,
2009, O’Donnell & Cigelnik 1999, Baayen et al. 2000, Lievens
et al. 2009b, Van Dam et al. 2016) have highlighted the para-
and polyphyletic relationships within several F. oxysporum
special forms, e.g., F. oxysporum f. sp. batatas, F. oxysporum
f. sp. cubense and F. oxysporum f. sp. vasinfectum. Addition-
ally, several F. oxysporum special forms are able to infect and
cause disease in more than one (sometimes unrelated) plant
hosts, whereas others are highly specialised to a specific plant
host (Armstrong & Armstrong 1981, Gordon & Martyn 1997,
Kistler 1997, Baayen et al. 2000, Leslie & Summerell 2006,
Fourie et al. 2011).
Naming F. oxysporum special forms are not subject to the Inter-
national Code of Nomenclature for algae, fungi, and plants (ICN;
McNeill et al. 2012, Thurland et al. 2018), and therefore no
diagnosis (in Latin and/or English), nor the deposit of type ma-
terial in a recognised repository is required. This decision was
made due to the difficulty in accepting special forms within the
Code, even though these strains are of great importance to plant
pathologists and breeders (Deighton et al. 1962, Gordon 1965,
Armstrong & Armstrong 1981). Several studies on F. oxysporum
indicate that between 70 to over 150 special forms are known
in F. oxysporum (Booth 1971, Armstrong & Armstrong 1981,
Kistler 1997, Baayen et al. 2000, Leslie & Summerell 2006,
Lievens et al. 2008, O’Donnell et al. 2009, Fourie et al. 2011,
Epitypification of Fusarium oxysporum –
clearing the taxonomic chaos
L. Lombard1, M. Sandoval-Denis1,2, S.C. Lamprecht3, P.W. Crous1,2,4
1 Westerdijk Fungal Biodiversity Institute, Uppsalalaan 8, 3584 CT Utrecht,
The Netherlands;
corresponding author e-mail: l.lombard@westerdijkinstitute.nl.
2 Faculty of Natural and Agricultural Sciences, Department of Plant Sciences,
University of the Free State, P.O. Box 339, Bloemfontein 9300, South Africa.
3 ARC-Plant Health and Protection, Private Bag X5017, Stellenbosch, 7599,
Western Cape, South Africa.
4 Wageningen University and Research Centre (WUR), Laboratory of Phyto-
pathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands.
Key words
cryptic species
diversity
human and plant pathogens
species complex
subspecific classification
Abstract Fusarium oxysporum is the most economically important and commonly encountered species of Fusa-
rium. This soil-borne fungus is known to harbour both pathogenic (plant, animal and human) and non-pathogenic
strains. However, in its current concept F. oxysporum is a species complex consisting of numerous cryptic species.
Identification and naming these cryptic species is complicated by multiple subspecific classification systems and the
lack of living ex-type material to serve as basic reference point for phylogenetic inference. Therefore, to advance
and stabilise the taxonomic position of F. oxysporum as a species and allow naming of the multiple cryptic species
recognised in this species complex, an epitype is designated for F. oxysporum. Using multi-locus phylogenetic
inference and subtle morphological differences with the newly established epitype of F. oxysporum as reference
point, 15 cryptic taxa are resolved in this study and described as species.
Article info Received: 20 June 2018; Accepted: 19 October 2018; Published: 18 December 2018.

2Persoonia – Volume 43, 2019
Laurence et al. 2014, Gordon 2017). At present Index Fungo-
rum (http:// www.indexfungorum.org/) lists 124 special forms in
F. oxysporum, whereas MycoBank (http: //www.mycobank.org/)
list 127 special forms. Further careful scrutiny of literature re-
vealed that 144 special forms have been named until February
2018 (Table 1). Although the special forms concept of Snyder
& Hansen (1940) is still applied today, additional subspecific
classification systems for special forms of F. oxysporum have
also been introduced, which include haplotypes, races and
vegetative compatibility groups (VCGs).
The haplotype subspecific classification system was introduced
by Chang et al. (2006) and later expanded upon by O’Donnell
et al. (2008, 2009) to include strains from both the FOSC
and Neocosmospora (formerly the F. solani (FSSC) species
complex). This classification system is based on unique multi-
locus genotypes within the species complex, aimed to resolve
communication problems among public health and agricultural
scientists (O’Donnell et al. 2008). Chang et al. (2006) proposed
a standardised haplotype nomenclature system that depict
the species complex, species and genotype. O’Donnell et al.
(2009) was able to identify 256 unique two-locus haplotypes
from 850 isolates representing 68 special forms of F. oxysporum
as well as environmental and clinical strains. However, this
classification system is not in common use as a reference, and
a continuously updated database is required.
One of the most important subspecific ranks applied to special
forms of F. oxysporum are physiological pathotypes or races.
This classification system is of great importance to plant breed-
ers, especially for resistance breeding. Traditionally, race demar-
cation is based on cultivar specificity linked to specific resistance
genes of the plant host cultivar (Armstrong & Armstrong 1981,
Kistler 1997, Baayen et al. 2000, Roebroeck 2000, Fourie et al.
2011, Epstein et al. 2017). However, race designation has been
inconsistent in the past (Gerlagh & Blok 1988, Correll 1991,
Kistler 1997, Fourie et al. 2011) with several different nomen-
clatural systems being applied (Gabe 1975, Risser et al. 1976,
Armstrong & Armstrong 1981) to further cause confusion (Kistler
1997). With advances in molecular technology, identification
of races has been simplified using sequence-characterised
amplified region (SCAR) primers (Lievens et al 2008, Epstein
et al. 2017, Gilardi et al. 2017). However, time consuming and
laborious pathogenicity tests are still needed to identify new
emerging races and to test whether newly developed plant
cultivars are resistant to known races (Epstein et al. 2017,
Gilardi et al. 2017).
The use of vegetative compatibility (also known as heterokar-
yon compatibility) has formed an integral part of subspecific
classification of F. oxysporum special forms and non-pathogenic
strains. Formation of a stable heterokaryon between two auxo-
trophic nutritional mutants is regulated by several vic or het
incompatibility loci (Correll 1991, Leslie 1993) indicating that
the strains are homogenic at these loci (Correll 1991) and con-
sidered to be part of the same VCG. Therefore, classification
using vegetative compatibility is based on genetic similarity at
specific loci and not pathogenicity, providing a crude marker
for population genetic studies (Correll 1991, Gordon & Martyn
1997, Leslie 1993, Leslie & Summerell 2006). Puhalla (1985),
utilizing nit mutants, was the first to identify VCGs in F. oxyspo-
rum and characterised 16 VCGs in a collection of 21 F. oxyspo-
rum strains. The numbering system applied by Puhalla (1985),
which is still used today, consists of a three-digit numerical
code indicating the special form followed by digit(s) indicating
the VCG (Katan 1999, Katan & Di Primo 1999). Conventional
VCG characterisation is a relatively objective, time consuming
and laborious assay only indicating genetic similarity and not
genetic difference (Kistler 1997). Therefore, several PCR-based
detection methods have been developed to identify economi-
cally important VCGs as diagnostic tool (Fernandez et al. 1998,
Pasquali et al. 2004a, c, Lievens et al. 2008), e.g., F. oxysporum
f. sp. cubense TR4 VCG01213 (Dita et al. 2010).
Until recently, limited knowledge on the genetic premise for host
specificity in F. oxysporum was available (Gordon & Martyn
1997, Kistler 1997, Baayen et al. 2000). However, the discovery
of a lineage-specific chromosome (or transposable/effector/
accessory chromosome) in F. oxysporum f. sp. lycopersici by
Ma et al. (2010), in which the host specific virulence genes lie
(Van der Does et al. 2008, Takken & Rep 2010, Ma et al. 2013),
has provided a new view into the evolution of pathogenicity in
F. oxysporum. In vitro transfer of these accessory chromosomes
into non-pathogenic F. oxysporum strains has converted the
latter strains into host-specific pathogens, providing evidence
that host-specific pathogenicity could be acquired through
horizontal transfer of accessory chromosomes (Takken & Rep
2010, Ma et al. 2010, 2013, Van Dam et al. 2016, Van Dam &
Rep 2017). Therefore, the special form name can be linked to
the accessory chromosome whereas race demarcation can be
linked to the specific virulence genes carried on these acces-
sory chromosomes.
The genetic and functional mechanisms of the infection process
in plants of various special forms of F. oxysporum has been
well documented (Di Pietro et al. 2003, Ma et al. 2013, Upasani
et al. 2016, Gordon 2017). However, these same mechanisms
are still poorly understood in human and animal infections
(O’Donnell et al. 2004, Guarro 2013, Van Diepeningen et al.
2015). Fusarium oxysporum has been linked to fungal keratitis
(Hemo et al. 1989, Chang et al. 2006) and dermatitis (Guarro
& Gene 1995, Romano et al. 1998, Ninet et al. 2005, Cutuli et
al. 2015, Van Diepeningen et al. 2015), and has been isolated
from contaminated hospital water systems (Steinberg et al.
2015, Edel-Hermann et al. 2016) and medical equipment (Bar-
ton et al. 2016, Carlesse et al. 2017) posing a serious threat
to immunocompromised patients. Several recent reports also
indicate that F. oxysporum is able to infect immunocompetent
patients (Jiang et al. 2016, Khetan et al. 2018). In general,
fusariosis is difficult to treat as Fusarium species display a
remarkable resistance to antifungal agents (Guarro 2013, Al-
Hatmi et al. 2018). However, some antimycotics are known to
be effective against F. oxysporum related fusariosis (Al-Hatmi
et al. 2018). Recently, both mycotoxins beauvericin and fusaric
acid, produced by F. oxysporum strains that can infect tomato,
have been shown to be important virulence determinants to
infect immunosuppressed mice (López-Berges et al. 2013,
López-Díaz et al. 2018).
Strains of F. oxysporum are known to produce a cocktail of
polyketide secondary metabolites, some with unknown function
and toxicities (Marasas et al. 1984, Mirocha et al. 1989, Bell
et al. 2003, Desjardins 2006, Manici et al. 2017). Some of the
better-known toxins produced by F. oxysporum include beau-
vericin (Marasas et al. 1984, Logrieco et al. 1998, López-Berges
et al. 2013), fusaric acid (Marasas et al. 1984, López-Díaz et
al. 2018) and fumonisins (Rheeder et al. 2002) to name a few.
Mycotoxicological studies on F. oxysporum has thus far only
focused on a strain to strain basis and therefore no link has
yet been established between special form and/or race and
mycotoxin production capabilities.
In light of the complicated and sometimes confusing classifi-
cation systems applied to F. oxysporum taxonomy and nomen-
clature, the question has risen whether F. oxysporum truly
represent a species (Kistler 1997). Given that F. oxysporum is
a common, widespread, soil-borne fungus, with a global distri-
bution and high economic importance, this question requires
urgent attention. Therefore, to advance and stabilize the taxo-
nomic and nomenclatural position of F. oxysporum and allow

3
L. Lombard et al.: Epitypification of Fusarium oxysporum
adzukicola Kitazawa & Yanagita 1984, 1989 Summerell et al. 2010 Katan & Di Primo 1999
aechmeae Sauthoff & Gerlach 1957, 1958 Fusarium bulbigenum f. aechmeae Gordon 1965, Armstrong & Gherbawy 1999, O’Donnell et al. 2009
Sauthoff & Gerlach, Gratenwelt 57: Armstrong 1968, 1981, Booth 1971,
390. 1957 Summerell et al. 2010
albedinis Sergent & Beguet 1921, Killian & Maire 1930, Cylindrophora albedinis Kill. & Maire, Gordon 1965, Armstrong & Tantaoui et al. 1996, Kistler Tantaoui & Boisson 1991, Tantaoui & Fernandez 1993
Malençon 1934, Louvet & Toutain 1981 Bull. Soc. Hist. Nat. Afrique N. 21: Armstrong 1968, 1981, Booth 1971, et al. 1998, Katan 1999 Tantaoui et al. 1996, Fernandez et al. 1994, 1998,
89 –101. 1930 Summerell et al. 2010 Skovgaard et al. 2001, Mbofung et al. 2007, Lievens
Fusarium albedinis (Kill. & Maire) et al. 2008, O’Donnell et al. 2009, Elliott et al. 2010,
Malençon, Compt. Rend. Acad. Sci. 198: Mirtalebi & Banihashemi 2014
1259–1261. 1930
Fusarium oxysporum var. albedinis
(Kill. & Maire) Malençon, Rev. Mycol.
(Paris) 15: 45– 60. 1950
aleuritis Suelong 1981 Suelong 1981
allii Matuo et al. 1979 Yoo et al. 1993, Katan & Di O’Donnell et al. 2009
Primo 1999
amaranthi Chen & Swart 2001 Summerell et al. 2010 Chen & Swart 2001 Chen & Swart 2001
anethi Janson 1951, Gordon 1965 Gordon 1965, Armstrong &
Armstrong 1968, 1981, Booth 1971,
Summerell et al. 2010
anoectochili Huang et al. 2014 Huang et al. 2014 Huang et al. 2014 Huang et al. 2014
apii Snyder & Hansen 1940 Fusarium apii P.E. Nelson & Sherb., Snyder & Hansen 1940, Gordon Schneider & Norelli 1981, Puhalla Puhalla 1984a, b, Correll Wang et al. 2001, O’Donnell et al. 2009,
Tech. Bull. Mich. Agric. Exp. Sta. 155: 1965, Armstrong & Armstrong 1984a, b, Epstein et al. 2017 et al. 1986, 1987, Toth & Chakrabarti et al. 2011, Epstein et al. 2017
42. 1937 1968, 1981, Booth 1971, Lacy 1991, Kistler et al.
Fusarium oxysporum f. apii (P.E. Summerell et al. 2010 1998, Katan 1999
Nelson & Sherb.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 27: 66. 1940
Fusarium bulbigenum var. apii (P.E.
Nelson & Sherb.) Raillo, Fungi of the
genus Fusarium: 250. 1950
Fusarium apii var. pallidum P.E. Nelson &
Sherb., Tech. Bull. Mich. Agric. Exp.
Sta. 155: 42. 1937
arctii Matuo et al. 1975 Summerell et al. 2010 O’Donnell et al. 2009
asparagi Cohen 1946 Gordon 1965, Armstrong & Blok & Bollen 1997, Elmer Baayen et al. 2000, Mbofung et al. 2007,
Armstrong 1968, 1981, Booth 1971, & Stephens 1989, Yoo et al. O’Donnell et al. 2009, Poli et al. 2012, Mirtalebi &
Summerell et al. 2010 1993, Kistler et al. 1998, Banihashemi 2014
Katan 1999, Katan & Di
Primo 1999
basilica Dzidzariya 1968, Armstrong & Armstrong 1981 Fusarium oxysporum var. basilicum Armstrong & Armstrong 1968, 1981, Elmer et al. 1994, Kistler Chiocchetti et al. 1999, 2001, Pasquali et al. 2006,
Dzidzariya, Pishch. Prom. SSR: Summerell et al. 2010 et al. 1998, Katan 1999, Lievens et al. 2008, O’Donnell et al. 2009
129–140. 1968 Katan & Di Primo 1999
batatas Wollenweber 1914, 1931 Fusarium batatas Wollenw., J. Agric. Snyder & Hansen 1940, Armstrong & Armstrong 1958b, 1968, Katan 1999, Katan & Di O’ Donnell et al. 1998, Kim et al. 2001, Mbofung
Res. 2: 268. 1914 Gordon 1965, Armstrong & Booth 1971 Primo 1999 et al. 2007, Lievens et al. 2009b, O’Donnell et al.
Fusarium bulbigenum var. batatas Armstrong 1968, 1981, Booth 1971, 2009, Pinaria et al. 2015
(Wollenw.) Wollenw., Z. Parasitenk. Summerell et al. 2010
(Berlin) 3: 414. 1931
Fusarium oxysporum f. batatas
(Wollenw.) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 27: 66. 1940
benincasae Gerlagh & Ester 1985 Gerlagh & Blok 1988
betae Stewart 1931 Fusarium conglutinans var. betae Snyder & Hansen 1940, Gordon Armstrong & Armstrong 1976 Harveson & Rush 1997, Cramer et al. 2003, Nitschke et al. 2009, O’Donnell
D. Stewart, Phytopathology 9: 59. 1931 1965, Armstrong & Armstrong 1968, Kistler et al. 1998, et al. 2009, Hill et al. 2011, Covey et al. 2014
Fusarium orthoceras var. betae 1981, Booth 1971, Summerell Webb et al. 2013
Table 1 List of known special forms of Fusarium oxysporum.
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

4Persoonia – Volume 43, 2019
betae (cont.) (D. Stewart) Padwick, Indian J. Agric. et al. 2010
Sci. 10: 282. 1940
Fusarium oxysporum f. betae
(D. Stewart) W.C. Snyder & H.N.
Hansen, Amer. J. Bot. 27: 66. 1940
Fusarium oxysporum var. orthoceras
(Appel & Wollenw.) Bilaǐ, The Fusaria:
282. 1955
bouvardiae Marziano et al. 1987 O’Donnell et al. 2009
brassica Williams et al. 2016 Williams et al. 2016
callistephi Beach 1918 Fusarium conglutinans var. callistephi Snyder & Hansen 1940, Gordon Armstrong & Armstrong 1971 Mbofung et al. 2007, O’Donnell et al. 2009, Poli et al.
Beach, Rep. Michigan Acad. Sci. 1965, Armstrong & Armstrong 1968, 2012
29: 297. 1918 1981, Booth 1971, Summerell
Fusarium orthoceras var. callistephi et al. 2010
(Beach) Padwick, Indian J. Agric. Sci.
10: 283. 1940
Fusarium oxysporum f. callistephi
(Beach) W.C. Snyder & H.N. Hansen,
Amer. J. Bot. 27: 66. 1940
Fusarium conglutinans var. majus
Wollenw., Fusaria Autographica
Delineata 3: 981. 1930
canariensis Mercier & Louvet 1973, Feather et al. 1979 Summerell et al. 2010 Katan 1999, Pyler et al. Pyler et al 2000, Gunn & Summerell 2002, Mbofung
2000, Gunn & Summerell et al. 2007, Lievens et al. 2009b, Elliott et al. 2010,
2002 Laurence et al. 2015, Pinaria et al. 2015
cannabis Noviello & Snyder 1962 Gordon 1965, Armstrong & Armstrong O’Donnell et al. 2009
1968, 1981 Booth 1971
capsici Black et al. 1993
carthami Klisiewicz & Houston 1963 Gordon 1965, Armstrong & Arm- Klisiewicz & Thomas 1970a, b, Shende et al. 2015
strong 1968, 1981, Booth 1971, Klisiewicz 1975
Summerell et al. 2010
cassiae Armstrong 1954, Gordon 1965 Gordon 1965, Armstrong & Arm- O’Donnell et al. 2009
strong 1968, 1981, Booth 1971,
Summerell et al. 2010
cattleyae Foster 1955 Gordon 1965, Armstrong & Arm- Baayen & Kleijn 1989 O’Donnell et al. 2009
strong 1968, 1981, Booth 1971,
Summerell et al. 2010
cepae Hanzawa 1914 Fusarium cepae Hanzawa, Snyder & Hansen 1940, Gordon Molnár et al. 1990, Yoo et al. Gherbawy 1999, Mbofung et al. 2007, Galván et al.
Mykol. Zentbl. 5: 5. 1914 1965, Armstrong & Armstrong 1968, 1993, Katan & Di Primo 2008, O’Donnell et al. 2009, Bayraktar et al. 2010,
Fusarium oxysporum f. cepae 1981, Booth 1971, Summerell 1999, Swift et al. 2002, Lin et al. 2010, Southwood et al. 2012, Mirtalebi &
(Hanzawa) W.C. Snyder & H.N. et al. 2010 Widodo et al. 2008, Banihashemi 2014, Taylor et al. 2016
Hansen, Amer. J. Bot. 27: 66. 1940 Bayraktar et al. 2010,
Fusarium oxysporum var. cepae Southwood et al. 2012
(Hanzawa) Raillo, Fungi of the genus
Fusarium: 253. 1950
chrysanthemi Armstrong et al. 1970 Armstrong & Armstrong 1968, Huang et al. 1992, Troisi et al. 2013 Puhalla 1985, Correll et al. Kim et al. 2001, Pasquali et al. 2003, 2004a, b, c,
1981, Booth 1971, 1987, Kistler et al. 1998, Bogale et al. 2007, Lievens et al. 2008, O’Donnell
Summerell et al. 2010 Katan 1999, Pasquali et al. et al. 2009, Li et al. 2010, Lin et al. 2010, Troisi et al.
2004c 2010, 2013
ciceris Padwick 1940, Erwin 1958, Matuo & Sato 1962 Fusarium orthoceras var. ciceri Armstrong & Armstrong 1968, Haware & Nene 1982, Barve et al. Kistler et al. 1998 Kelly et al. 1994, 1998, García- Pedrasjas et al. 1999,
Padwick, Indian J. Agr. Sci. 10: 1981, Booth 1971 2001, Jiménez-Gasco et al. 2001, Barve et al. 2001, Jiménez-Gasco et al. 2001, 2002,
241–284. 1940 2004a, b, Jiménez-Gasco & 2004a, b, Jiménez-Gasco & Jiménez-Díaz 2003,
Fusarium lateritium f. ciceri (Padwick) Jiménez-Díaz 2003, Sharma et al. 2004, 2014, 2016, Honnareddy & Dubey
Erwin, Phytopathology 48: 500. 1958 Sharma et al. 2004, Honnareddy & 2006, Bayraktar et al. 2008, Dubey & Singh 2008,
Dubey 2006, Gurjar et al. 2009, Gurjar et al. 2009, Dubey et al. 2012, Demers et al.
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

5
L. Lombard et al.: Epitypification of Fusarium oxysporum
ciceris (cont.) Dubey et al. 2012, Demers et al. 2014, 2014, Ghosh et al. 2015, Upasani et al. 2016, Williams
Upasani et al. 2016 et al. 2016
cichorii Poli et al. 2012 Poli et al. 2012
citri Timmer et al. 1979, Timmer 1982 Hannachi et al. 2015
coffeae Alvarez 1945, Wellman 1954 Fusarium bulbigenum var. coffeae Gordon 1965, Armstrong & Armstrong
Álv. García, J. Agric. Univ. Puerto 1968, 1981, Booth 1971, Summerell
Rico 29: 8. 1945 et al. 2010
colocasiae Nishimura & Kudo 1994 Hirano & Arie 2009, Poli et al. 2013
conglutinans Wollenweber 1913, Padwick 1940 Fusarium conglutinans Wollenw., Snyder & Hansen 1940, Gordon Ramirez-Villupadua et al. 1985, Puhalla 1985, Bosland & Bosland & Williams 1987, Kistler et al. 1987, Kistler
Phytopathology 3 (1): 30. 1913 1965, Armstrong & Armstrong 1968, Armstrong & Armstrong 1952, Williams 1987, Correll et al. & Benny 1989, Crowhurst et al. 1995, Gherbawy
Fusarium orthoceras var. conglutinans 1981, Booth 1971, Summerell 1953, 1966 1987, Correll 1991, Kistler 1999, Kim et al. 2001, Bogale et al. 2007, Hirano &
(Wollenw.) Padwick, Indian J. Agric. et al. 2010 et al. 1998, Katan 1999, Arie 2009, O’Donnell et al. 2009, Srinivasan et al.
Sci. 10: 282. 1940 Katan & Di Primo 1999 2010, Poli et al. 2012, Covey et al. 2014, Zang et al.
Fusarium oxysporum f. conglutinans 2014, Hansen et al. 2015, Kashiwa et al. 2016, Li et
(Wollenw.) W.C. Snyder & H.N. Hansen, al. 2015, 2016, Taylor et al. 2016, Van Dam & Rep
Amer. J.Bot. 27: 66. 1940 2017
coriandrii Booth 1971, Armstrong & Armstrong 1981 Armstrong & Armstrong 1968, 1981,
Booth 1971, Summerell et al. 2010
crassulae Ortu et al. 2013 Ortu et al. 2013
croci Boerema & Hamers 1989 Roebroeck 2000 Roebroeck 2000 Roebroeck 2000, Palmero et al. 2014
crotalariae Kulkarni 1934, Gupta 1974 Fusarium vasinfectum var. crotalariae Armstrong & Armstrong 1968, 1981
Kulk., Indian J. Agric. Sci 4: 994. 1934
Fusarium udum f.sp. crotalariae (Kulk.)
Subram., The genus Fusarium: 114. 1971
cubense Smith 1910, Brandes 1919 Fusarium cubense E.F. Sm., Science, Snyder & Hansen 1940, Gordon See review by Fourie et al. 2011 See review by Fourie et al. See review by Fourie et al. 2011, Ploetz 2015 and
N.S. 31: 755. 1910 1965, Armstrong & Armstrong and Ploetz 2015 2011 and Ploetz 2015, Lin & Shen 2017, Mostert et al. 2017, Aguayo et al.
Fusarium cubense var. inodoratum E.W. 1968, 1981, Booth 1971, Mostert et al. 2017 2017, Van Dam & Rep 2017, Czislowski et al. 2017
Brandes, Phytopathology 9: 374. 1919 Summerell et al. 2010
Fusarium oxysporum var. cubense
(E.F. Sm.) Wollenw., Die Fusarien,
ihre Beschreibung, Schadwirkung und
Bekämpfung: 119. 1935
Fusarium oxysporum f. cubense
(E.F. Sm.) W.C. Snyder & H.N. Hansen,
Amer. J. Bot. 27: 66. 1940
cucumerinum Owen 1956 Gordon 1965, Armstrong & Arm- Armstrong & Armstrong 1978b, Ahn et al. 1998, Kistler et al. Namiki et al. 1994, Vakalounakis & Fragkiadakis 1999,
strong 1968, 1981, Booth 1971, Armstrong et al. 1978, Gerlagh & 1998, Katan 1999, Kim et al. 2001, Skovgaard et al. 2001, Wang et al.
Summerell et al. 2010 Blok 1988 Katan & Di Primo 1999, 2001, Vakalounakis et al. 2004, Lievens et al. 2007,
Vakalounakis & Fragkiadakis 2008, Hirano & Arie 2009, O’Donnell et al. 2009, Lin
1999, Vakalounakis et al. et al. 2010, Poli et al. 2013, Scarlett et al. 2013,
2004 Mirtalebi & Banihashemi 2014, Bertoldo et al. 2015
cucurbitacearum Gerlagh & Blok 1988 Gerlagh & Blok 1988 Bogale et al. 2007, O’Donnell et al. 2009, Bennett
et al. 2013
cumini Patel et al. 1957 Summerell et al. 2010 Talaviya et al. 2014, Nawade et al. 2017
cyclaminis Gerlach 1954 Gordon 1965, Armstrong & Arm- Woudt et al. 1995, Kistler Woudt et al. 1995, Gherbawy 1999, Kim et al. 2001,
strong 1968, 1981, Booth 1971, et al. 1998, Katan 1999, O’Donnell et al. 2009, Lecomte et al. 2016
Summerell et al. 2010 Lori et al. 2012
dahliae Summerell et al. 2010 Summerell et al. 2010
delphinii Laskaris 1949 Gordon 1965, Armstrong & Arm- Kondo et al. 2013
strong 1968, 1981, Booth 1971,
Summerell et al. 2010
dianthi Snyder & Hansen 1940 Fusarium dianthi Prill. & Delacr., Compt. Snyder & Hansen 1940, Gordon Hood & Stewart 1957, Garibaldi Puhalla 1985, Correll et al. Manicom et al. 1990, Manicom & Baayen 1993,
Rend. Acad. Sci.: 744–745. 1899 1965, Armstrong & Armstrong 1968, 1975, 1977, 1983, Baayen et al. 1988, 1987, Hadar et al. 1989, Manulis et al. 1994, Crowhurst et al. 1995, Baayen
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

6Persoonia – Volume 43, 2019
dianthi (cont.) Fusarium oxysporum f. dianthi (Prill. & 1981, Booth 1971, Summerell Aloi & Baayen 1993, Summerell et al. Molnár et al. 1990, Manicom et al. 1997, 2000, Gherbawy 1999, Kim et al. 2001,
Delacr.) W.C. Snyder & H.N. Hansen, et al. 2010 2010 et al. 1990, Aloi & Baayen Skovgaard et al. 2001, Bogale et al. 2007, Lievens
Amer. J. Bot. 27: 66. 1940 1993, Baayen et al. 1997, et al. 2008, Hirano & Arie 2009, O’Donnell et al. 2009,
Fusarium oxysporum f. sp. barbati Kistler et al. 1998, Katan Poli et al. 2013, Bertoldo et al. 2015, Pinaria et al.
W.C. Snyder, Phytopathology 31: 1056. 1941 1999, Katan & Di Primo 2015, Koyyappurath et al. 2016, Taylor et al. 2016
Fusarium oxysporum var. dianthi (Prill. & 1999
Delacr.) Raillo, Fungi of the genus Fusarium:
255. 1950
dioscoreae Wellman 1972
echeveriae Ortu et al. 2015a Ortu et al. 2015a
elaeagni Armstrong & Armstrong 1968 Fusarium oxysporum var. orthoceras Armstrong & Armstrong 1968, 1981,
(Appel & Wollenw.) Bilaǐ, The Fusaria: Booth 1971, Summerell et al. 2010
282. 1955
elaeidis Gordon 1965 Gordon 1965, Booth 1971, See Flood 2006 for prior See Flood 2006 for prior publications; Bogale et al.
Armstrong & Armstrong 1981, publications 2007, O’Donnell et al. 2009, Elliott et al. 2010
Summerell et al. 2010
erucae Chatterjee & Rai 1974
erythroxyli Sands et al. 1997 Summerell et al. 2010 Sands et al. 1997, Kistler Sands et al. 1997, Lievens et al. 2009b, O’Donnell
et al. 1998, Katan 1999, et al. 2009
Katan & Di Primo 1999
eucalypti Arya & Jain 1962 Gordon 1965, Armstrong & Arm-
strong 1968, 1981, Booth 1971,
Summerell et al. 2010
eustomae Raabe 1985a Bertoldo et al. 2015
fabae Yu & Fang 1948 Gordon 1965, Armstrong & Arm- Mbofung et al. 2007, O’Donnell et al. 2009, Srinivasan
strong 1968, 1981, Booth 1971 et al. 2010, Mirtalebi & Banihashemi 2014
fatshederae Triolo & Lorenzini 1983 O’Donnell et al. 2009
foli see Hirooka et al. 2008 Hirooka et al. 2008
fragariae Winks & Williams 1965 Armstrong & Armstrong 1968, 1981, Katan & Di Primo 1999, Kim et al. 2001, Nagarajan et al. 2004, 2006, Hirano
Booth 1971, Summerell et al. 2010 Nagarajan et al. 2006 & Arie 2009, O’Donnell et al. 2009, Chakrabarti et
al. 2011, Fang et al. 2013, Poli et al. 2013, Suga et
al. 2013, Bertoldo et al. 2015, Czislowski et al. 2017,
Henry et al. 2017
freesia Taylor et al. 2016
garlic Matuo et al. 1986 Yoo et al. 1993, Katan &
Di Primo 1999
gerberae Von Arx 1952, Gordon 1965 Gordon 1965, Armstrong & Armstrong 1968,
1981, Booth 1971, Summerell et al. 2010
gladioli Massey 1926, Snyder & Hansen Fusarium oxysporum var. gladioli Snyder & Hansen 1940, Gordon Roebroeck & Mes 1992, Mes Molnár et al. 1990, Mes et Mes et al. 1994, Crowhurst et al. 1995, Baayen et al.
1940, Buxton 1955 Massey, Phytopathology 16: 511. 1926 1965, Armstrong & Armstrong et al. 1994, De Haan et al. 2000 al. 1994, Kistler et al. 1998, 2000, De Haan et al. 2000, Kim et al. 2001, Bogale
Fusarium oxysporum f. gladioli 1968, 1981, Booth 1971, Summerell Katan 1999, Katan & Di et al. 2007, O’Donnell et al. 2009, Elliott et al. 2010,
(Massey) W.C. Snyder & H.N. Hansen, et al. 2010 Primo 1999, Di Primo et al. Lin et al. 2010, Pinaria et al. 2015, Van Dam & Rep
Amer. J. Bot. 27: 66. 1940 2002 2017
Fusarium orthoceras var. gladioli
L. McCulloch, Phytopathology 34:
280. 1944
glycines Armstrong & Armstrong 1965 Armstrong & Armstrong 1968, 1981, Lievens et al. 2009b, O’Donnell et al. 2009, Pinaria
Booth 1971, Summerell et al. 2010 et al. 2015, Koyyappurath et al. 2016
hebes Raabe 1985b Gordon 1965, Armstrong & Armstrong
1968, 1981, Booth 1971, Summerell
et al. 2010
heliconiae Waite 1963 (see Ploetz 2006)
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

7
L. Lombard et al.: Epitypification of Fusarium oxysporum
heliotropae Netzer & Weintal 1987 Mbofung et al. 2007, O’Donnell et al. 2009
herbemontis Gordon 1965 Fusarium oxysporum var. herbemontis Gordon 1965, Armstrong & Armstrong
Tochetto, Revta Agron., Porto Alegre: 1968, 1981, Booth 1971, Summerell
82– 89. 1954 et al. 2010
iridiacearum Roebroeck 2000 Roebroeck 2000 Roebroeck 2000 Roebroeck 2000
koae Gardner 1980 Shiraishi et al. 2012 O’Donnell et al. 2009, Shiraishi et al. 2012
laciniati Pandotra et al. 1971 Summerell et al. 2010
lactucae Matuo & Motohashi 1967, Summerell et al. 2010 Fujinaga et al. 2001, 2003, 2005, Kistler et al. 1998, Katan Fujinaga et al. 2005, 2014, Shimazu et al. 2005,
Hubbard & Gerik 1993 2014, Yamauchi et al. 2001, 2004, 1999, Ogiso et al. 2002, Mbofung et al. 2007, Pasquali et al. 2007, 2008,
Ogiso et al. 2002, Shimazu et al. Yamauchi et al. 2004, Lievens et al. 2008, Hirano & Arie 2009, O’Donnell
2005, Pasquali et al. 2007, 2008, Pasquali et al. 2005, 2008, et al. 2009, Lin et al. 2010, 2014, Mbofung & Pryor
Lin et al. 2014, Gilardi et al. 2017 Pintore et al. 2017 2010, Poli et al. 2012, 2013, Mirtalebi & Banihas-
hemi 2014, Bertoldo et al. 2015, Gilardi et al. 2017
lagenariae Matuo & Yamamoto 1967 Armstrong & Armstrong 1968, Armstrong & Armstrong 1978b Katan & Di Primo 1999 Okuda et al. 1998, Kim et al. 2001, Galván et al.
1981, Booth 1971, Summerell 2008, Hirano & Arie 2009, O’Donnell et al. 2009,
et al. 2010 Poli et al. 2013
lathyri Bhide & Uppal 1948 Fusarium oxysporum var. lathyri Gordon 1965,Armstrong & Arm-
V.P. Bhide & Uppal, Phytopathology 38: strong 1968, 1981, Booth 1971,
560– 567. 1948 Summerell et al. 2010
lentis Vasudeva & Srinivasan 1952 Fusarium orthoceras var. lentis Gordon 1965, Armstrong & Arm- Pouralibaba et al. 2016, 2017 Belabid & Fortas 2002 Belabid et al. 2004, O’Donnell et al. 2009, Taheri et
Vasudeva & Sriniv., Indian Phytopathol. strong 1968, 1981, Booth 1971, al. 2010, Datta et al. 2011, Mohammadi et al. 2011,
5: 28. 1953 Summerell et al. 2010 Rafique et al. 2015, Al-Husien et al. 2017, Nourollahi
& Madahjalai 2017
lilii Imle 1942 Gordon 1965, Armstrong & Arm- Löffler & Rumine 1991, Baayen et al. 1998, 2000, Kim et al. 2001, Skovgaard
strong 1968, 1981, Booth 1971, Baayen et al. 1998, Kistler et al. 2001, Wang et al. 2001, O’Donnell et al. 2009,
1981, Summerell et al. 2010 et al. 1998, Katan 1999, Lin et al. 2010, Baysal et al. 2013, Van Dam & Rep
Katan & Di Primo 1999 2017
lini Bolley 1901 Fusarium lini Bolley, Proc. Ann. Snyder & Hansen 1940, Gordon Katan & Di Primo 1999, Baayen et al. 2000, Bogale et al. 2007, O’Donnell
Meeting Soc. Prom. Agr. Sci. 22: 42. 1965, Armstrong & Armstrong 1968, Baayen et al. 2000 et al. 2009, Pinaria et al. 2015, Taylor et al. 2016
1901 1981, Booth 1971, Summerell
Fusarium oxysporum f. lini (Bolley) et al. 2010
W.C. Snyder & H.N. Hansen, Amer.
J. Bot. 27: 66. 1940
loti Bergstrom & Kalb 1995 Wunsch et al. 2009 Galván et al. 2008, O’Donnell et al. 2009, Wunsch
et al. 2009
luffae Kawai et al. 1958 Summerell et al. 2010 Armstrong & Armstrong 1978b Kim et al. 1993, Wang et al. 2001, Lin et al. 2010
lupini Snyder & Hansen 1940 Snyder & Hansen 1940, Gordon Richter 1941, Armstrong & Armstrong Kistler et al. 1998, Katan Bogale et al. 2007, O’Donnell et al. 2009
1965, Armstrong & Armstrong 1968, 1964, Rataj-Guranowska et al. 1984 1999, Katan & Di Primo
1981, Booth 1971, Summerell 1999
et al. 2010
lycopersici Wollenweber 1913 Fusarium oxysporum subsp. lycopersici Snyder & Hansen 1940, Gordon Alexander & Tucker 1945, Gerdemann Puhalla 1985, Correll et al. Elias & Schneider 1992; Elias et al. 1993, Crowhurst
Sacc., Syll. Fung. 4: 705. 1886 1965, Armstrong & Armstrong 1968, & Finley 1951, Gabe 1975, Elias & 1987, Hadar et al. 1989, et al. 1995, Marlatt et al. 1996, Mes et al. 1998,
Fusarium lycopersici Bruschi, Rc. 1981, Booth 1971, Summerell Schneider 1992, Elias et al. 1993, Molnár et al. 1990, Correll Gherbawy 1999, Kim et al. 2001, Bao et al. 2002, Cai
Accad. Naz. Lincei: 298. 1912 et al. 2010 Marlatt et al. 1996, Mes et al. 1998, 1991, Elias & Schneider et al. 2003, Hirano & Arie 2006, 2009, Bogale et al.
Fusarium lycopersici (Sacc.) Wollenw., Cai et al. 2003, Hirano & Arie 2006, 1991, 1992, Marlatt et al. 2007, Mbofung et al. 2007, Lievens et al. 2009a, b,
Phytopathology 3 (1): 29. 1913 Lievens et al. 2009a 1996, Kistler et al. 1998, O’Donnell et al. 2009, Elliott et al. 2010, Inami et al.
Fusarium oxysporum f. lycopersici Mes et al. 1998, Katan 2010, Ma et al. 2010, See review by Takken & Rep
(Sacc.) W.C. Snyder & H.N. Hansen, 1999, Katan & Di Primo 2010, Chakrabarti et al. 2011, Poli et al. 2012, 2013,
Amer. J. Bot. 27: 66. 1940 1999, Cai et al. 2003 Thatcher et al. 2012, Baysal et al. 2013, Bennett et
al. 2013, Covey et al. 2014, Gawehns et al. 2014,
Mirtalebi & Banihashemi 2014, Bertoldo et al. 2015,
Hansen et al. 2015, Nirmaladevi et al. 2016, Taylor
et al. 2016, Williams et al. 2016, Bilju et al. 2017, Van
Dam & Rep 2017, Jelinski et al. 2017
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

8Persoonia – Volume 43, 2019
magnoliae Lin & Chen 1994
matthiolae Baker 1948 Booth 1971, Summerell et al. 2010 Correll 1991, Kistler et al. Kistler et al. 1987, Mbofung et al. 2007, O’Donnell
1998, Katan 1999 et al. 2009, Srinivasan et al. 2010, Poli et al. 2012
medicaginis Weimer 1928 Fusarium oxysporum var. medicaginis Snyder & Hansen 1940, Gordon Puhalla 1985, Correll et al. Mbofung et al. 2007, O’Donnell et al. 2009, Srinivasan
Weimer, J. Agric. Res. 37: 425. 1928 1965, Armstrong & Armstrong 1987, Molnár et al. 1990, et al. 2010, Poli et al. 2012, Mirtalebi & Banihashemi
Fusarium oxysporum f. medicaginis 1968, 1981, Booth 1971, Kistler et al. 1998, Katan 2014, Thatcher et al. 2016, Williams et al. 2016,
(Weimer) W.C. Snyder & H.N. Hansen, Summerell et al. 2010 1999 Czislowski et al. 2017
Amer. J. Bot. 27: 66. 1940
melongenae Matuo & Ishigami 1958 Gordon 1965, Armstrong & Hadar et al. 1989, Kistler Crowhurst et al. 1995, Kim et al. 2001, Hirano & Arie
Armstrong 1968, Booth 1971, et al. 1998, Katan 1999, 2009, O’Donnell et al. 2009, Altinok & Can 2010,
1981, Summerell et al. 2010 Katan & Di Primo 1999, Baysal et al. 2010, Bennett et al. 2013, Poli et al.
Altinok & Can 2010, Altinok 2013, Bertoldo et al. 2015, Dong et al. 2017
2013, Altinok et al. 2013
melonis Leach & Currence 1938, Fusarium bulbigenum var. niveum Snyder & Hansen 1940, Gordon Risser & Mas 1965, Risser et al. Correll et al. 1987, Jacobson & Gordon 1990b, Kim et al. 1993, 2001,
Snyder & Hansen 1940 Leach & Curr., Minnisota Agric. Exp. 1965, Armstrong & Armstrong 1976, Armstrong & Armstrong 1978b, Jacobson & Gordon 1988, Crowhurst et al. 1995, Namiki et al. 1998, 2001,
Sta. Tech. Bull. 129: 1–32. 1938 1968, 1981, Booth 1971, Gerlagh & Blok 1988, Katan et al. 1994, 1990a, Hadar et al. 1989, Gherbawy 1999, Skovgaard et al. 2001, Mbofung et
Summerell et al. 2010 Luongo et al. 2014, Mirtalebi & Correll 1991, Katan et al. al. 2007, Hirano & Arie 2009, Lievens et al. 2009b,
Banihashemi 2014, Sebastiani et 1994, Kistler et al. 1998, O’Donnell et al. 2009, Lin et al. 2010, Bennett et al.
al. 2017 Katan 1999, Katan & Di 2013, Poli et al. 2013, Covey et al. 2014, Gawehns
Prim o 199 9, Mir talebi & et al. 2014, Luongo et al. 2014, Ma et al. 2014, Mirtalebi
Banihashemi 2014 & Banihashemi 2014, Bertoldo et al. 2015, Hansen
et al. 2015, Pinaria et al. 2015, Schmidt et al. 2016,
Taylor et al. 2016, Williams et al. 2016, Van Dam &
Rep 2017, Sebastiani et al. 2017
meniscoideum (var.) Bugnicourt 1939 Gerlach & Nirenberg 1982 O’Donnell et al. 2009
momordicae Sun & Huang 1983 Skovgaard et al. 2001, O’Donnell et al. 2009, Lin et
al. 2010, Bennett et al. 2013, Chen et al. 2015
mori Pastrana et al. 2017 Pastrana et al. 2017 Pastrana et al. 2017
narcissi Wollenweber & Reinking 1935, Snyder & Hansen 1940, Gordon Linfield 1993, Crowhurst et al. 1995, O’Donnell et al.
Snyder & Hansen 1940 1965, Armstrong & Armstrong 2009, Taylor et al. 2016, Van Dam & Rep 2017
1968, 1981, Booth 1971,
Summerell et al. 2010
nelumbicola Gordon 1965 Fusarium bulbigenum var. nelumbicola Snyder & Hansen 1940, Gordon
Y. Nisik. & Kyoto Watan., Ber. Ohara 1965, Armstrong & Armstrong
Inst. Landw. Biol. Okayama Univ.: 3. 1968, 1981, Booth 1971,
1953 Summerell et al. 2010
nicotianae Johnson 1921 Fusarium oxysporum var. nicotianae Booth 1971, Summerell et al. 2010 Bogale et al. 2007, O’Donnell et al. 2009
J. Johnson, J. Agric. Res. 20: 525. 1921
niveum Wollenweber & Reinking 1935 Fusarium niveum E.F. Sm., Bull. Snyder & Hansen 1940, Gordon Reid 1958, Crall 1963, Netzer 1976, Puhalla 1985, Correll et al. Kim et al. 1993, 2001, Crowhurst et al. 1995, Zhang
U.S.D.A. 1894 1965, Armstrong & Armstrong Armstrong & Armstrong 1978b, 1987, Hadar et al. 1989, et al. 2005, Bogale et al. 2007, Hirano & Arie 2009,
Fusarium bulbigenum var. niveum 1968, 1981, Booth 1971, Martyn 1987, Gerlagh & Blok 1988, Larkin et al. 1988, 1990, O’Donnell et al. 2009, Lin et al. 2010, Chakrabarti et
(E.F. Sm.) Wollenw., Die Fusarien: Summerell et al. 2010 Martyn & Bruton 1989, Larkin et al. Correll 1991, Kistler et al. al. 2011, Poli et al. 2013, Gawehns et al. 2014, Mirtalebi
117. 1935 1990, Zhou et al. 2010 1998, Katan 1999, Katan & Banihashemi 2014, Bertoldo et al. 2015, Ren et al.
Fusarium oxysporum f. niveum & Di Primo 1999 2015, Van Dam & Rep 2017, Czislowski et al. 2017
(E.F. Sm.) W.C. Snyder & H.N. Hansen,
Amer. J. Bot. 27: 66. 1940
opuntiarum Gordon 1965 Fusarium oxysporum var. opuntiarum Gordon 1965, Armstrong & Katan & Di Primo 1999 Baayen et al. 2000, Mbofung et al. 2007, O’Donnell
Pettinari, Annali Sper. Agr.: 1419. 1951 Armstrong 1968, 1981, et al. 2009, Ortu et al. 2013, Pinaria et al. 2015,
Booth 1971, Summerell et al. 2010 Koyyappurath et al. 2016, Bertetti et al. 2017
orthoceras Bilaǐ 1955
oxysporum (var.) Von Schlechtendahl 1824 Gerlach & Nirenberg 1982
palmarum Elliott et al. 2010 O’Donnell et al. 2009, Elliott et al. 2010, 2017,
Giesbrecht et al. 2013
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

9
L. Lombard et al.: Epitypification of Fusarium oxysporum
papaveris Ortu et al. 2015b Summerell et al. 2010 Katan 1999 Bertetti et al. 2014, Ortu et al. 2015b
passiflorae Gordon 1965 Gordon 1965, Armstrong & Arm- Gherbawy 1999, Bogale et al. 2007, Lievens et al.
strong 1968, 1981, Booth 1971, 2009b, O’Donnell et al. 2009, Chakrabarti et al. 2011,
Summerell et al. 2010 Dos Santos Silva et al. 2013, Gawehns et al. 2014,
Pinaria et al. 2015, Koyyappurath et al. 2016, Czislowski
et al. 2017
perillae Kim et al. 2002
perniciosum Toole 1941 Fusarium perniciosum Hepting, Gordon 1965, Armstrong & Arm- Toole 1952 Crowhurst et al. 1995, Bogale et al. 2007, Mbofung
Circ. U.S.D.A.: 7. 1939 strong 1968, 1981, Booth 1971, et al. 2007, Lievens et al. 2009b, O’Donnell et al.
Fusarium oxysporum f. perniciosum Summerell et al. 2010 2009, Elliott et al. 2010, Bennett et al. 2013, Pinaria
(Hepting) Toole, Phytopathology 31: et al. 2015
599. 1941
Fusarium vasinfectum var. perniciosum
(Hepting) Carrera, Monatsh. Landw.:
483. 1955
phaseoli Kendrick & Snyder 1942b Gordon 1965, Armstrong & Arm- Ribeiro 1977, Ribeiro & Hagedorn Woo et al. 1996, Kistler Woo et al. 1996, Cramer et al. 2003, Zanotti et al.
strong 1968, 1981, Booth 1971, 1979, Salgado & Schwartz 1993, et al. 1998, Katan 1999, 2006, Alves-Santos et al. 2002b, Bogale et al. 2007,
Summerell et al. 2010 Woo et al. 1996, Alves-Santos et al. Katan & Di Primo 1999, Mbofung et al. 2007, Hirano & Arie 2009, O’Donnell
2002a, Cramer et al. 2003, Henrique Alves-Santos et al. 2002a et al. 2009, De Vega-Bartol et al. 2011, Baysal et al.
et al. 2015 2013, Poli et al. 2013, Mirtalebi & Banihashemi 2014,
Da Silva et al. 2014, Bertoldo et al. 2015, De Sousa
et al. 2015
phormii Wager 1947 Gordon 1965, Armstrong & Arm-
strong 1968, 1981, Booth 1971,
Summerell et al. 2010
pini Hartig 1892, Snyder & Hansen 1940 Fusisporium aurantiacum Link, Mag. O’Donnell et al. 2009
Ges. Naturf. Freunde Berlin 3: 19. 1809
Fusoma pini Hartig, Forstl.-Naturwiss.
Z. 1: 432– 436. 1892
Fusarium blasticola Rostr., Gartner-
Tidende 1895: 122. 1895
Fusarium oxysporum f. pini (Hartig) W.C.
Snyder & H.N. Hansen, Amer. J. Bot. 27: 66. 1940
Fusarium oxysporum f. sp. blasticola Bilaǐ,
Fusarii: 281. 1955
pisi Van Hall 1903, Snyder & Hansen 1940 Fusarium vasinfectum var. pisi C.J.J. Snyder & Hansen 1940, Gordon Snyder & Walker 1935, Snyder & Puhalla 1985, Correll et al. Coddington et al. 1987, Kistler et al. 1991, Whitehead
Hall, Ber. Deutsch. Bot. Ges. 21: 4. 1903 1965, Armstrong & Armstrong Hansen 1940, Schreuder 1951, 1987, Correll 1991, White- et al. 1992, Grajal-Martin et al. 1993, Gherbawy 1999,
Fusarium orthoceras var. pisi Linford, 1968, 1981, Booth 1971, Bolton et al. 1966, Armstrong & head et al. 1992, Kistler Skovgaard et al. 2001, O’Donnell et al. 2009,
Res. Bull. Agric. Exp. Stn Univ. Wis.: Summerell et al. 2010 Armstrong 1974, Kraft & Haglund et al. 1998, Katan 1999, Chakrabarti et al. 2011, Covey et al. 2014, Mirtalebi
11. 1928 1978, Haglund & Kraft 1979, Katan & Di Primo 1999, & Banihashemi 2014, Hansen et al. 2015, Taylor et
Fusarium oxysporum f. 8 W.C. Snyder, Coddington et al. 1987, Whitehead al. 2016, Williams et al. 2016, Van Dam & Rep 2017
Zentralbl. Bakteriol., 2. Abt.: 374. 1935 et al. 1992, Grajal-Martin et al. 1993
Fusarium oxysporum var. pisi (C.J.J. Hall)
Raillo, Fungi of the genus Fusarium: 254.
1950
Fusarium oxysporum var. orthoceras
(Appel & Wollenw.) Bilaǐ, Fusarii: 282. 1955
psidii Prasad et al. 1952 Gordon 1965, Armstrong & Arm- Gupta 2012, Mishra et al. 2013a, b, c, 2014
strong 1968, 1981, Booth 1971,
Summerell et al. 2010
pyracanthae McRitchie 1973, Armstrong & Armstrong 1981 Armstrong & Armstrong 1968, 1981,
Summerell et al. 2010
querci Gordon 1965 Gordon 1965, Armstrong & Arm-
strong 1968, 1981, Booth 1971,
Summerell et al. 2010
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

10 Persoonia – Volume 43, 2019
quitoense Ochoa et al. 2004
radicis-capsici Lomas-Cano et al. 2014, 2016 Lomas-Cano et al. 2014
radicis-cucumerinum Vakalounakis 1996 Summerell et al. 2010 Katan 1999, Katan & Di Vakalounakis & Fragkiadakis 1999, Vakalounakis
Primo 1999, Vakalounakis et al. 2004, 2005, Lievens et al. 2007, Van Dam
& Fragkiadakis 1999, & Rep 2017
Vakalounakis et al. 2004,
2005, Tok & Kurt 2010
radicis-lupini Weimer 1944 Gordon 1965, Booth 1971,
Summerell et al. 2010
radicis-lycopersici Jarvis & Shoemaker 1978 Summerell et al. 2010 Puhalla 1985, Correll et al. Kim et al. 2001, Skovgaard et al. 2001, Balmas et al.
1987, Katan et al. 1991, 2005, Hirano & Arie 2006, 2009, Bogale et al. 2007,
Kistler et al. 1998, Katan Hibar et al. 2007, O’Donnell et al. 2009, Huang et al.
1999, Katan & Di Primo 2013, Poli et al. 2013, Covey et al. 2014, Mirtalebi &
1999, Rosewich et al. Banihashemi 2014, Bertoldo et al. 2015, Taylor
1999, Di Primo et al. 2001, et al. 2016
Balmas et al. 2005, Huang
et al. 2013
radicis-vanillae Koyyappurath et al. 2016 Koyyappurath et al. 2016
ranunculi Garibaldi & Gullino 1985
rapae Enya et al. 2008 Enya et al. 2008 Enya et al. 2008
raphani Kendrick & Snyder 1942a Gordon 1965, Armstrong & Bosland & Williams 1987, Kistler & Benny 1989, Kistler et al. 1991, Kim et al.
Armstrong 1968, 1981, Booth Kistler et al. 1998, Katan 2001, Bogale et al. 2007, Hirano & Arie 2009, O’Donnell
1971, Summerell et al. 2010 1999, Katan & Di Primo et al. 2009, Lin et al. 2010, Srinivasan et al. 2010,
1999 Poli et al. 2012, 2013, Covey et al. 2014, Bertoldo
et al. 2015, Taylor et al. 2016, Van Dam & Rep
2017, Kim et al. 2017
rauvolfiae Janardhanan et al. 1964 Gordon 1965, Armstrong & O’Donnell et al. 2009
Armstrong 1968, 1981, Booth
1971, Summerell et al. 2010
rhois Snyder et al. 1949 Gordon 1965, Armstrong & Mbofung et al. 2007
Armstrong 1968, 1981, Booth
1971, Summerell et al. 2010
ricini Gordon 1965 Fusarium orthoceras var. ricini Gordon 1965, Armstrong & Prasad et al. 2008, Reddy et al. 2012
Wollenw., Biologico 6: 148. 1940 Armstrong 1968, 1981, Booth
1971, Summerell et al. 2010
samaneae Wellman 1972
sansevieriae Gupta et al. 1982
sedi Raabe 1960 Gordon 1965, Armstrong &
Armstrong 1968, 1981, Booth
1971, Summerell et al. 2010
sesami Gordon 1965, Booth 1971 Fusarium vasinfectum var. sesami Gordon 1965, Armstrong & Basirnia & Banihashemi O’Donnell et al. 2009, Li et al. 2012,
Zaprom., Pflanzenschutz-Vers. Armstrong 1968, 1981, Booth 2005 Bennett et al. 2013
Sta. Taschkent: 36 pp. 1926 1971, Summerell et al. 2010
sesbaniae Gordon 1965, Booth 1971 Gordon 1965, Armstrong &
Armstrong 1968, 1981, Booth
1971, Summerell et al. 2010
spinaciae Hungerford 1923 Fusarium spinaciae Sherb., Snyder & Hansen 1940, Gordon Armstrong & Armstrong 1976 Kistler et al. 1998, Katan Baayen et al. 2000, Kim et al. 2001, Skovgaard et al.
Phytopathology 13: 209. 1923 1965, Armstrong & Armstrong 1999, Katan & Di Primo 2001, Kawabe et al. 2007, Mbofung et al. 2007, Hirano
Fusarium oxysporum f. spinaciae 1968, 1981, Booth 1971, 1999, Takehara et al. & Arie 2009, O’Donnell et al. 2009, Poli et al. 2012,
(Sherb.) W.C. Snyder & H.N. Hansen, Summerell et al. 2010 2003 2013, Bennett et al. 2013, Okubara et al. 2013,
Amer. J. Bot. 27: 66. 1940 Covey et al. 2014, Mirtalebi & Banihashemi
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

11
L. Lombard et al.: Epitypification of Fusarium oxysporum
spinaciae (cont.) Fusarium redolens f. spinaciae 2014, Bertoldo et al. 2015
(Sherb.) Subram., Hyphomycetes:
an account of Indian species,
except Cercosporae: 690. 1971
stachydis Gordon 1965 Gordon 1965, Armstrong &
Armstrong 1968, 1981, Booth
1971, Summerell et al. 2010
strigae Elzein & Kroschel 2006 Elzein et al. 2008, Zimmermann et al. 2015, 2016
tabernaemontanae Pande & Rao 1990
tanaceti Hirooka et al. 2008 Hirooka et al. 2008
tracheiphilum Wollenweber 1931, Snyder & Hansen 1940 Fusarium tracheiphilum E.F. Sm. 1899 Snyder & Hansen 1940, Gordon Armstrong & Armstrong 1950, 1980, Correll et al. 1987, Kistler Gherbawy 1999, Bao et al. 2002, Hirano & Arie 2009,
Fusarium bulbigenum var. tracheiphilum 1965, Armstrong & Armstrong Hare 1953, Swanson & Van Gundy et al. 1998, Katan 1999, O’Donnell et al. 2009, Lin et al. 2010, Troisi et al.
(E.F. Sm.) Wollenw., Z. Parasitenk. 1968, 1981, Booth 1971, 1985, Smith et al. 1999 Katan & Di Primo 1999, 2010, Bennett et al. 2013, Poli et al. 2013, Bertoldo
(Berlin) 3: 413. 1931 Summerell et al. 2010 Bao et al. 2002 et al. 2015, Koyyappurath et al. 2016
Fusarium oxysporum f. tracheiphilum
(E.F. Sm.) W.C. Snyder & H.N. Hansen,
Amer. J. Bot. 27: 66. 1940
trifolii Bilaǐ 1955 Fusarium trifolii Jacz., Jb. Pfl. krankh. Gordon 1965, Armstrong &
Russl. VII-VIII, Abt. 6. 1917 Armstrong 1968, 1981, Booth
Fusarium oxysporum var. trifolii (Jacz.) 1971, Summerell et al. 2010
Raillo, Fungi of the genus Fusarium: 255.
1950
tuberosi Snyder & Hansen 1940 Fusarium oxysporum var. solani Snyder & Hansen 1940, Gordon Molnár et al. 1990, Venter Gherbawy 1999, Lievens et al. 2009a,
Raillo, Fungi of the genus Fusarium: 1965, Armstrong & Armstrong et al. 1992, Kistler et al. O’Donnell et al. 2009
254. 1950 1968, 1981, Booth 1971, 1998, Katan 1999
Fusarium oxysporum var. solani Summerell et al. 2010
(Raillo) Bilaǐ, Fusarii: 281. 1955
tulipae Snyder & Hansen 1940 Gordon 1965, Armstrong & Katan 1999, Katan & Gherbawy 1999, Baayen et al. 2000, Kim et al. 2001,
Armstrong 1968, 1981, Booth Di Primo 1999 Skovgaard et al. 2001, Hirano & Arie 2009, O’Donnell
1971, Summerell et al. 2010 et al. 2009, Poli et al. 2013, Mirtalebi & Banihashemi
2014, Bertoldo et al. 2015, Pinaria et al. 2015, Swett
& Uchida 2015, Van Dam & Rep 2017
vanillae Tucker 1927 Fusarium batatas var. vanillae Gordon 1965, Armstrong & Katan & Di Primo 1999 O’Donnell et al. 2009, Chakrabarti et al. 2011,
Tucker, J. Agric. Res. 44: 1121. 1927 Armstrong 1968, 1981, Booth Adame-García et al. 2015, Pinaria et al. 2015,
1971, Summerell et al. 2010 Koyyappurath et al. 2016
vasconcella Ochoa et al. 2004
vasinfectum Atkinson 1892 Fusarium vasinfectum G.F. Atk., Snyder & Hansen 1940, Gordon Armstrong & Armstrong 1958a, 1960, Puhalla 1985, Correll et al. Assigbetse et al. 1994, Fernandez et al. 1994,
Bulletin of the Alabama Agricultural 1965, Armstrong & Armstrong 1978a, Ibrahim 1966, Kappelman 1987, Katan & Katan 1988, Crowhurst et al. 1995, Moricca et al. 1998, Skovgaard
Experiment Station: 28. 1892 1968, 1981, Booth 1971, 1983, Chen et al. 1985, Assigbetse Hadar et al. 1989, Correll et al. 2001, Smith et al. 2001, Abd-Elsalam et al. 2002,
Fusarium oxysporum f. vasinfectum Summerell et al. 2010 et al. 1994, Fernandez et al. 1994, 1991, Fernandez et al. 2004, 2006, Abo et al. 2005, Kim et al. 2005, 2017,
(G.F. Atk.) W.C. Snyder & H.N. Hansen, Nirenberg et al. 1994, Skovgaard et 1994, Davis et al. 1996, McFadden et al. 2006, Wang et al. 2006, 2010,
Amer. J. Bot. 27: 66. 1940 al. 2001, Kim et al. 2005, Holmes et al. Kistler et al. 1998, Katan Mbofung et al. 2007, Zambounis et al. 2007, Bennett
2009, Guo et al. 2015 1999, Katan & Di Primo et al. 2008, 2013, Holmes et al. 2009, O’Donnell et
1999, Abo et al. 2005, al. 2009, Elliot et al. 2010, Chakrabarti et al. 2011,
Wang et al. 2010 Egamberdiev et al. 2013, 2014, Da Silva et al. 2014,
Covey et al. 2014, Doan et al. 2014, Cianchetta et al.
2015, Guo et al. 2015, Pinaria et al. 2015, Crutcher
et al. 2016, Taylor et al. 2016, Van Dam & Rep 2017,
Ortiz et al. 2017
voandzeiae Armstrong et al. 1975 Armstrong & Armstrong 1981 O’Donnell et al. 2009
zingiberi Trujillo 1963 Pappalardo et al. 2009 Katan & Di Primo 1999 Crowhurst et al. 1995, O’Donnell et al. 2009,
Pappalardo et al. 2009, Chakrabarti et al. 2011,
Gupta et al. 2014, Czislowski et al. 2017
Table 1 (cont.)
formae speciales Description Synonym(s) Listed Race(s) VCG(s) Molecular studies

12 Persoonia – Volume 43, 2019
naming of the multiple cryptic species recognised in this spe-
cies complex, Fusarium isolates were collected from the type
locality in Berlin, Germany, and the type substrate, Solanum
tuberosum. Using molecular phylogenetic and morphological
tools, an epitype is designated for F. oxysporum in the present
study based on these collections.
MATERIALS AND METHODS
Isolates
Tubers of S. tuberosum (potato), displaying symptoms of dry
rot, were collected from several vegetable gardens in Berlin,
Germany. Potato tubers were placed individually in paper
bags, stored at 4 °C until transported to the laboratory for
further processing. After surface-sterilisation of the potato tu-
bers using a 10 % (v/v) sodium hypochlorite solution, pieces
of symptomatic tissue were removed from the leading edges
of the rot lesions and plated onto 2 % (w/v) potato dextrose
agar (PDA) amended with 100 µg/mL penicillin and 100 µg/
mL streptomycin, and peptone pentachloronitrobenzene agar
(PCNB; Nash & Snyder 1962) and incubated at 25 °C in the
dark. Axenic cultures were prepared on PDA from characteristic
Fusarium colonies. Additional strains, previously identified as
F. oxysporum, were obtained from the culture collection (CBS)
of the Westerdijk Fungal Biodiversity Institute (WFBI), Utrecht,
the Netherlands, and the working collection of Pedro W. Crous
(CPC) housed at WFBI (Table 2).
DNA isolation, PCR and sequencing
Total genomic DNA was extracted from isolates grown for 7 d on
PDA at 24 °C using a 12/12 h photoperiod using the Wizard®
Genomic DNA purification Kit (Promega Corporation, Madison,
WI, USA), according to the manufacturer’s instructions. Partial
gene sequences were determined for the β-tubulin (tub2),
calmodulin (cmdA), the intergenic spacer region of the rDNA
(IGS), RNA polymerase II second largest subunit (rpb2) and
translation elongation factor 1-alpha (tef1), using PCR protocols
described elsewhere (O’Donnell et al. 1998, 2007, 2009, 2010,
Lombard et al. 2015). Primer pairs T1/CYLTUB1R (O’Donnell
& Cigelnik 1997, Crous et al. 2004) for tub2, Cal228F/CAL2Rd
(Carbone & Kohn 1999, Groenewald et al. 2013) for cmdA,
iNL11/iCNS1 and the internal sequencing primers NLa/CNSa
(O’Donnell et al. 2009) for IGS, 5f2/7cr (Liu et al. 1999, Sung
et al. 2007) for rpb2, and EF1/EF2 (O’Donnell et al. 1998) for
tef1, were used for amplifications of the respective gene re-
gions. Integrity of the sequences was ensured by sequencing
the amplicons in both directions using the same primer pairs
as were used for amplification. Consensus sequences for
each locus were assembled in MEGA v. 7 (Kumar et al. 2016),
with the exception of the IGS locus, which was assembled in
Geneious R11 (Kearse et al. 2012). All sequences generated
in this study were deposited in GenBank (Table 1).
Phylogenetic analyses
Sequences of the individual loci were aligned using MAFFT
v. 7.110 (Katoh et al. 2017) and manually corrected where
necessary. The individual gene datasets were assessed for
incongruency prior to concatenation using a 70 % reciprocal
bootstrap criterion (Mason-Gamer & Kellogg 1996). Three in-
dependent phylogenetic algorithms, Maximum Parsimony (MP),
Maximum Likelihood (ML) and Bayesian inference (BI), were
employed for phylogenetic analyses. Phylogenetic analyses
were conducted for the individual loci and then as a multilocus
sequence dataset that included the cmdA, rpb2, tef1 and tub2
sequences.
For BI and ML, the best evolutionary models for each locus
were determined using MrModeltest (Nylander 2004) and in-
corporated into the analyses. MrBayes v. 3.2.1 (Ronquist &
Huelsenbeck 2003) was used for BI to generate phylogenetic
trees under optimal criteria for each locus. A Markov Chain
Monte Carlo (MCMC) algorithm of four chains was initiated in
parallel from a random tree topology with the heating parameter
set at 0.3. The MCMC analysis lasted until the average standard
deviation of split frequencies was below 0.01 with trees saved
every 1 000 generations. The first 25 % of saved trees were
discarded as the ‘burn-in’ phase and posterior probabilities (PP)
were determined from the remaining trees.
The ML analyses were performed using RAxML v. 8.2.9 (ran-
domised accelerated (sic) maximum likelihood for high per-
formance computing; Stamatakis 2014) through the CIPRES
website (http://www.phylo.org) to obtain another measure of
branch support. The robustness of the analysis was evalu-
ated by bootstrap support (BS) with the number of bootstrap
replicates automatically determined by the software. For MP,
analyses were done using PAUP (Phylogenetic Analysis Using
Parsimony, v. 4.0b10; Swofford 2003) with phylogenetic rela-
tionships estimated by heuristic searches with 1 000 random
addition sequences. Tree-bisection-reconnection was used,
with branch swapping option set on ‘best trees’ only. All charac-
ters were weighted equally and alignment gaps treated as fifth
state. Measures calculated for parsimony included tree length
(TL), consistency index (CI), retention index (RI) and rescaled
consistence index (RC). Bootstrap (BS) analyses (Hillis &
Bull 1993) were based on 1 000 replications. Alignments and
phylogenetic trees derived from this study were uploaded to
TreeBASE (www.treebase.org).
Genealogical concordance phylogenetic species
recognition (GCPSR)
In order to establish the recombination levels between the newly
proposed species in this study and their closest phylogenetic
relatives, pairwise homoplasy index (PHI) analyses were done
on the respective concatenated multilocus datasets (Bruen et al.
2006). The analyses were conducted as described by Quaed-
vlieg et al. (2014) using SplitsTree v. 4.14.4 (Huson & Bryant
2006). Therefore, a PHI value below 0.05 (fW < 0.05) would
indicate the presence of significant recombination in the dataset.
Split graphs were constructed for visualization of the relation-
ships between closely related species.
Morphological characterisation
All isolates were characterised following the protocols described
by Leslie & Summerell (2006) using potato dextrose agar (PDA;
recipe in Crous et al. 2009), synthetic nutrient-poor agar (SNA;
Nirenberg 1976) and carnation leaf agar (CLA; Fisher et al.
1982). Colony morphology, pigmentation, odour and growth
rates were evaluated on PDA after 3 and 7 d at 24 °C with
a 12/12 h cool fluorescent light/dark cycle as described by
Sandoval-Denis et al. (2018) and using the colour charts of
Rayner (1970). Micromorphological characters were examined
using water as mounting medium on a Zeiss Axioskop 2 plus
with Differential Interference Contrast (DIC) optics and a Nikon
AZ100 stereomicroscope both fitted with Nikon DS-Ri2 high
definition colour digital cameras to photo-document fungal
structures. Measurements were taken using the Nikon software
NIS-elements D v. 4.50 and the 95 % confidence levels were
determined for the conidial measurements with extremes given
in parentheses. For all other fungal structures examined, only the
extremes are presented. To facilitate the comparison of relevant
micro- and macroconidial features, composite photo plates were
assembled from separate photographs using PhotoShop CSS.

13
L. Lombard et al.: Epitypification of Fusarium oxysporum
Fusarium callistephi CBS 187.53T Callistephus chinensis callistephi The Netherlands MH484693 MH484784 MH484875 MH484966 MH485057
CBS 115423 Agathosma betulina South Africa MH484723 MH484814 MH484905 MH484996 MH485087
F. carminascens CBS 144739 = CPC 25792 Zea mays South Africa MH484752 MH484843 MH484934 MH485025 MH485116
CBS 144740 = CPC 25793 Z. mays South Africa MH484753 MH484844 MH484935 MH485026 MH485117
CBS 144741 = CPC 25795 Z. mays South Africa MH484754 MH484845 MH484936 MH485027 MH485118
CBS 144738 = CPC 25800T Z. mays South Africa MH484755 MH484846 MH484937 MH485028 MH485119
F. contaminatum CBS 111552 Pasteurized fruit juice The Netherlands MH484718 MH484809 MH484900 MH484991 MH485082
CBS 114899T Pasteurized chocolate milk Germany MH484719 MH484810 MH484901 MH484992 MH485083
CBS 117461 Tetra pack with milky nutrition The Netherlands MH484729 MH484820 MH484911 MH485002 MH485093
F. cugenangense CBS 620.72 = DSM 11271 = NRRL 36520 Crocus sp. gladioli Germany MH484697 MH484788 MH484879 MH484970 MH485061
CBS 130304 = BBA 69050 = NRRL 25433 Gossypium barbadense vasinfectum China MH484739 MH484830 MH484921 MH485012 MH485103
CBS 130308 = ATCC 26225 = NRRL 25387 Human toe nail New Zealand MH484738 MH484829 MH484920 MH485011 MH485102
CBS 131393 Vicia faba Australia MH484746 MH484837 MH484928 MH485019 MH485110
F. curvatum CBS 247.61 = BBA 8398 = DSM 62308 = NRRL 22545 Matthiola incana matthiolae Germany MH484694 MH484785 MH484876 MH484967 MH485058
CBS 238.94 = NRRL 26422 = PD 94/184T Beaucarnia sp. meniscoideum The Netherlands MH484711 MH484802 MH484893 MH484984 MH485075
CBS 141.95 = NRRL 36251 = PD 94/1518 Hedera helix The Netherlands MH484712 MH484803 MH484894 MH484985 MH485076
F. duoseptatum CBS 102026 = NRRL 36115 Musa sapientum cv. Pisang ambon cubense Malaysia MH484714 MH484805 MH484896 MH484987 MH485078
F. elaeidis CBS 217.49 = NRRL 36358 Elaeis sp. elaeidis Zaire MH484688 MH484779 MH484870 MH484961 MH485052
CBS 218.49 = NRRL 36359 Elaeis sp. elaeidis Zaire MH484689 MH484780 MH484871 MH484962 MH485053
CBS 255.52 = NRRL 36386 Elaeis guineensis elaeidis Unknown MH484692 MH484783 MH484874 MH484965 MH485056
F. fabacearum CBS 144742 = CPC 25801 Z. mays South Africa MH484756 MH484847 MH484938 MH485029 MH485120
CBS 144743 = CPC 25802T Glycine max South Africa MH484757 MH484848 MH484939 MH485030 MH485121
CBS 144744 = CPC 25803 G. max South Africa MH484758 MH484849 MH484940 MH485031 MH485122
F. foetens CBS 120665 Nicotiana tabacum Iran MH484736 MH484827 MH484918 MH485009 MH485100
F. glycines CBS 176.33 = NRRL 36286 Linum usitatissium lini Unknown MH484686 MH484777 MH484868 MH484959 MH485050
CBS 214.49 = NRRL 36356 Unknown Argentina MH484687 MH484778 MH484869 MH484960 MH485051
CBS 200.89 Ocimum basilicum basilici Italy MH484706 MH484797 MH484888 MH484979 MH485070
CBS 144745 = CPC 25804 G. max South Africa MH484759 MH484850 MH484941 MH485032 MH485123
CBS 144746 = CPC 25808T G. max South Africa MH484760 MH484851 MH484942 MH485033 MH485124
F. gossypinum CBS 116611 Gossypium hirsutum vasinfectum Ivory Coast MH484725 MH484816 MH484907 MH484998 MH485089
CBS 116612 G. hirsutum vasinfectum Ivory Coast MH484726 MH484817 MH484908 MH484999 MH485090
CBS 116613T G. hirsutum vasinfectum Ivory Coast MH484727 MH484818 MH484909 MH485000 MH485091
F. hoodiae CBS 132474T Hoodia gordonii hoodiae South Africa MH484747 MH484838 MH484929 MH485020 MH485111
CBS 132476 H. gordonii hoodiae South Africa MH484748 MH484839 MH484930 MH485021 MH485112
CBS 132477 H. gordonii hoodiae South Africa MH484749 MH484840 MH484931 MH485022 MH485113
F. languescens CBS 645.78 = NRRL 36531T Solanum lycopersicum lycopersici Morocco MH484698 MH484789 MH484880 MH484971 MH485062
CBS 646.78 = NRRL 36532 S. lycopersicum lycopersici Morocco MH484699 MH484790 MH484881 MH484972 MH485063
CBS 413.90 = ATCC 66046 = NRRL 36465 S. lycopersicum lycopersici Israel MH484708 MH484799 MH484890 MH484981 MH485072
CBS 300.91 = NRRL 36416 S. lycopersicum lycopersici The Netherlands MH484709 MH484800 MH484891 MH484982 MH485073
CBS 302.91 = NRRL 36419 S. lycopersicum lycopersici The Netherlands MH484710 MH484801 MH484892 MH484983 MH485074
CBS 872.95 = NRRL 36570 S. lycopersicum radicis-lycopersici Unknown MH484713 MH484804 MH484895 MH484986 MH485077
CBS 119796 = MRC 8437 Z. mays South Africa MH484735 MH484826 MH484917 MH485008 MH485099
F. libertatis CBS 144748 = CPC 25782 Aspalathus sp. South Africa MH484750 MH484841 MH484932 MH485023 MH485114
CBS 144747 = CPC 25788 Aspalathus sp. South Africa MH484751 MH484842 MH484933 MH485024 MH485115
CBS 144749 = CPC 28465T Rock surface South Africa MH484762 MH484853 MH484944 MH485035 MH485126
F. nirenbergiae CBS 129.24 Secale cereale Unknown MH484682 MH484773 MH484864 MH484955 MH485046
CBS 149.25 = NRRL 36261 Musa sp. cubense Unknown MH484683 MH484774 MH484865 MH484956 MH485047
Species Culture accession1 Host/ substrate Special form Origin GenBank accession
cmdA IGS rpb2 tef1 tub2
Table 2 Details of Fusarium strains included in the phylogenetic analyses.

14 Persoonia – Volume 43, 2019
F. nirenbergiae (cont.) CBS 181.32 = NRRL 36303 S. tuberosum USA MH484685 MH484776 MH484867 MH484958 MH485049
CBS 758.68 = NRRL 36546 S. lycopersicum lycopersici The Netherlands MH484695 MH484786 MH484877 MH484968 MH485059
CBS 744.79 = BBA 62355 = NRRL 22549 Passiflora edulis passiflorae Brazil MH484700 MH484791 MH484882 MH484973 MH485064
CBS 127.81 = BBA 63924 = NRRL 36229 Chrysanthemum sp. chrysanthemi USA MH484701 MH484792 MH484883 MH484974 MH485065
CBS 129.81 = BBA 63926 = NRRL 22539 Chrysanthemum sp. chrysanthemi USA MH484703 MH484794 MH484885 MH484976 MH485067
CBS 196.87 = NRRL 26219 Bouvardia longiflora bouvardiae Italy MH484704 MH484795 MH484886 MH484977 MH485068
CBS 840.88T Dianthus caryophyllus dianthi The Netherlands MH484705 MH484796 MH484887 MH484978 MH485069
CBS 115416 = CPC 5307 Agathosma betulina South Africa MH484720 MH484811 MH484902 MH484993 MH485084
CBS 115417 = CPC 5306 A. betulina South Africa MH484721 MH484812 MH484903 MH484994 MH485085
CBS 115419 = CPC 5308 A. betulina South Africa MH484722 MH484813 MH484904 MH484995 MH485086
CBS 115424 = CPC 5312 A. betulina South Africa MH484724 MH484815 MH484906 MH484997 MH485088
CBS 123062 = GJS 91-17 Tulip roots USA MH484737 MH484828 MH484919 MH485010 MH485101
CBS 130300 = NRRL 26368 Amputated human toe USA MH484743 MH484834 MH484925 MH485016 MH485107
CBS 130301 = NRRL 26374 Human leg ulcer USA MH484744 MH484835 MH484926 MH485017 MH485108
CBS 130303 S. lycopersicum radicis-lycopersici USA MH484741 MH484832 MH484923 MH485014 MH485105
CPC 30807 South Africa MH484768 MH484859 MH484950 MH485041 MH485132
F. odoratissimum CBS 794.70 = BBA 11103 = NRRL 22550 Albizzia julibrissin perniciosum Iran MH484696 MH484787 MH484878 MH484969 MH485060
CBS 102030 M. sapientum cv. Pisang mas cubense Malaysia MH484716 MH484807 MH484898 MH484989 MH485080
CBS 130310 = NRRL 25603 Musa sp. cubense Australia MH484740 MH484831 MH484922 MH485013 MH485104
F. oxysporum CBS 221.49 = IHEM 4508 = NRRL 22546 Camellia sinensis medicaginis South East Asia MH484690 MH484781 MH484872 MH484963 MH485054
CBS 144134ET S. tuberosum Germany MH484771 MH484862 MH484953 MH485044 MH485135
CBS 144135 S. tuberosum Germany MH484772 MH484863 MH484954 MH485045 MH485136
CPC 25822 Protea sp. South Africa MH484761 MH484852 MH484943 MH485034 MH485125
F. pharetrum CBS 144750 = CPC 30822 Aliodendron dichotomum South Africa MH484769 MH484860 MH484951 MH485042 MH485133
CBS 144751 = CPC 30824T A. dichotomum South Africa MH484770 MH484861 MH484952 MH485043 MH485134
F. trachichlamydosporum CBS 102028 = NRRL 36117 M. sapientum cv. Pisang awak legor cubense Malaysia MH484715 MH484806 MH484897 MH484988 MH485079
F. triseptatum CBS 258.50 = NRRL 36389T Ipomoea batatas batatas USA MH484691 MH484782 MH484873 MH484964 MH485055
CBS 116619 G. hirsutum vasinfectum Ivory Coast MH484728 MH484819 MH484910 MH485001 MH485092
CBS 119665 Sago starch Papua New Guinea MH484734 MH484825 MH484916 MH485007 MH485098
CBS 130302 = NRRL 26360 = FRC 755 Human eye USA MH484742 MH484833 MH484924 MH485015 MH485106
F. udum CBS 177.31 Digitaria eriantha South Africa MH484684 MH484775 MH484866 MH484957 MH485048
F. veterinarium CBS 109898 = NRRL 36153T Shark peritoneum The Netherlands MH484717 MH484808 MH484899 MH484990 MH485081
CBS 117787 Swab sample near filling apparatus The Netherlands MH484730 MH484821 MH484912 MH485003 MH485094
CBS 117790 Swab sample near filling apparatus The Netherlands MH484731 MH484822 MH484913 MH485004 MH485095
CBS 117791 Pasteurized milk-based product The Netherlands MH484732 MH484823 MH484914 MH485005 MH485096
CBS 117792 Pasteurized milk-based product The Netherlands MH484733 MH484824 MH484915 MH485006 MH485097
NRRL 54984 Mouse mucosa USA MH484763 MH484854 MH484945 MH485036 MH485127
NRRL 54996 Little blue penguin foot USA MH484764 MH484855 MH484946 MH485037 MH485128
NRRL 62542 Unknown animal faeces USA MH484765 MH484856 MH484947 MH485038 MH485129
NRRL 62545 Endoscope of veterinary clinic USA MH484766 MH484857 MH484948 MH485039 MH485130
NRRL 62547 Canine stomach USA MH484767 MH484858 MH484949 MH485040 MH485131
Fusarium sp. CBS 128.81 = BBA 63925 = NRRL 36233 Chrysanthemum sp. chrysanthemi USA MH484702 MH484793 MH484884 MH484975 MH485066
CBS 680.89 = NRRL 26221 Cucumis sativus cucurbitacearum The Netherlands MH484707 MH484798 MH484889 MH484980 MH485071
CBS 130323 Human nail Australia MH484745 MH484836 MH484927 MH485018 MH485109
1 ATCC: American Type Culture Collection, USA; BBA: Biologische Bundesanstalt für Land- und Forstwirtschaft, Berlin-Dahlem, Germany; CBS: Westerdijk Fungal Biodiverity Institute (WIFB), Utrecht, The Netherlands; CPC: Collection of P.W. Crous; DSM: Deutsche Sammlung von
Mikroorganismen und Zellkulturen GmbH, Braunschweig, Germany; FRC: Fusarium Research Center, Penn State University, Pennsylvania; GJS: Collection of Gary J. Samuels; IHEM: Institute of Hygiene and Epidemiology-Mycology Laboratory, Brussels, Belgium; MRC: National
Research Institute for Nutritional Diseases, Tygerberg, South Africa; NRRL: Agricultural Research Service Culture Collection, USA; PD: Collection of the Dutch National Plant Protection Organization, Wageningen, The Netherlands. T Ex-type culture; ETEpitype.
Species Culture accession1 Host/ substrate Special form Origin GenBank accession
cmdA IGS rpb2 tef1 tub2
Table 2 (cont.)

15
L. Lombard et al.: Epitypification of Fusarium oxysporum
RESULTS
Isolates
A total of 23 fusarium-like isolates were obtained from the
symptomatic tissues of the potato tubers. Of these, six isolates
displayed typical F. oxysporum-like phenotypes, of which two
(CBS 144134 and CBS 144135) were selected for further study.
Phylogenetic analyses
Approximately 500–650 bases were determined for cmdA,
tef1 and tub2, 880 bases for rpb2 and 2 650 bases for IGS.
Sequence comparisons of the IGS, rpb2 and tef1 gene regions
generated in this study, against those in the Fusarium-ID (http://
isolate.fusariumdb.org/blast.php) and Fusarium-MLST (http://
www.westerdijkinstitute.nl/fusarium/) databases revealed that
all isolates included in this study belonged to the FOSC. The
congruency analysis revealed no conflict between the cmdA,
rpb2, tef1 and tub2 sequence datasets and were therefore
combined. However, the IGS sequence dataset revealed major
conflict with several included taxa resolving into single lineages
due to the large number of ambiguous regions in this gene
region. Therefore, the IGS sequences were excluded from
further analyses.
For the BI and ML analyses, a K80 model for cmdA, an HKY+
G+I model for rpb2, an HKY+G for tef1 and SYM+I+G model for
tub2 were selected and incorporated into the analyses. The ML
tree topology confirmed the tree topologies obtained from the BI
and MP analyses, and therefore, only the ML tree is presented.
The combined four loci sequence dataset included 89 ingroup
taxa with F. foetens (CBS 120665) and F. udum (CBS 177.31)
as outgroup taxa. The dataset consisted of 2 679 characters
including gaps. Of these characters, 2 291 were constant, 211
parsimony-uninformative and 177 parsimony-informative. The
BI lasted for 1.2 M generations, and the consensus tree and
posterior probabilities (PP) were calculated from 8 814 trees
left after 2 937 were discarded as the ‘burn-in’ phase. The MP
analysis yielded 1 000 trees (TL = 574; CI = 0.747; RI = 0.858;
RC = 0.641) and a single best ML tree with -InL = 7353.014512
(Fig. 1).
In the phylogenetic tree (Fig. 1) the ingroup taxa resolved into
eight clades (I–VIII). Of these, Clades I, II, IV and VI represent
single well- (ML & MP-BS ≥ 75– 95 %; PP ≥ 0.95–0.98) to
highly (ML & MP-BS ≥ 96 %; PP ≥ 0.99–1.0) supported clades,
whereas Clades III, V, VII and VIII displayed substantial sub-
structure. Clade III included eight well- to highly supported sub-
clades as well as two single lineages. Sequence comparisons of
the rpb2 and tef1 sequences with those generated by Maryani
et al. (2019) revealed that both single lineages represented
F. duo septatum (CBS 102026) and F. tradichlamydosporum
(CBS 102028), respectively. Similarly, the subclade that include
isolates CBS 620.72, CBS 130304, CBS 130308 and CBS
131393 represent F. cugenangense. Both Clades V and VIII
resolved two subclades in each, and Clade VII included three
subclades. The phylogenetic relationships between Clades
I–VIII and their underlying subclades are further discussed in
the notes in the Taxonomy section.
The PHI tests revealed that no evidence of recombination
(fW = 0.43; Fig. 2a) was detected between each Clade (I–VIII)
and their underlining subclades. Similarly, the genealogical
exclusivity of the subclades in Clades III (fW = 0.43; Fig. 2b)
and VII (fW = 1.0; Fig. 2d), as well as between Clades IV–VIII
(fW = 0.06; Fig. 2c) was also confirmed. The basal subclade in
Clade VIII (fW = 0.031; Fig. 2c), however, showed significant
evidence for recombination among all isolates included.
Taxonomy
In this section we provide a new (emended) description of
F. oxysporum and designate an epitype for this species. The
following species are also recognised as new within the FOSC,
based on phylogenetic inference and morphological compari-
sons. Isolates CBS 128.81, CBS 680.89 and CBS 130323 in
Clade III are not treated further as these were sterile.
Fusarium callistephi L. Lombard & Crous, sp. nov. — Myco-
Bank MB826833; Fig. 3
Etymology. Name refers to the plant genus Callistephus from which this
fungus was isolated.
Typus. NetherlaNds, Oostenbrink, from Callistephus chinensis, 28 Feb.
1953, collector unknown (holotype CBS H-23608 designated here, culture
ex-type CBS 187.53).
Conidiophores carried on the aerial mycelium 60–110 µm
tall, unbranched or sparingly branched, bearing terminal or
intercalarily monophialides, often reduced to single phialides;
aerial phialides subulate to subcylindrical, smooth- and thin-
walled, 2–23 × 3 –4 µm, periclinal thickening inconspicuous
or absent; aerial conidia forming small false heads on the tips
of the phialides, hyaline, ellipsoidal to falcate, smooth- and
thin-walled, 0–1-septate; 0-septate conidia: (6–)7–11(–14) ×
2–3 µm (av. 9 × 3 µm); 1-septate conidia: (13–)14–18 (–20) ×
2–4 µm (av. 16 × 3 µm). Sporodochia pale luteous to pale rosy
vinaceous, formed abundantly on carnation leaves. Conidio-
phores in sporodochia verticillately branched and densely
packed, consisting of a short, smooth- and thin-walled stipe,
4–7 × 2–4 µm, bearing apical whorls of 2–3 monophialides or
rarely as single lateral monophialides; sporodochial phialides
subulate to subcylindrical, 9–13 × 3– 4 µm, smooth- and thin-
walled, sometimes showing a reduced and flared collarette.
Sporodochial conidia falcate, curved dorsiventrally with almost
parallel sides tapering slightly towards both ends, with a blunt
to papillate, curved apical cell and a blunt to foot-like basal cell,
3–4(–5)-septate, hyaline, smooth- and thin-walled; 3-septate
conidia: (28–)33 –39(–40) × 3–5 µm (av. 36 × 4 µm); 4-septate
conidia: (30–)35 –41(– 42) × 3 –5 µm (av. 38 × 4 µm); 5-septate
conidia: 36–44 (–47) × 4– 5 µm (av. 40 × 5 µm). Chlamydo-
spores not observed.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 2.9–4.2 mm /d at 24 °C. Colony surface
white to pale vinaceous, floccose with abundant aerial myce-
lium; colony margins irregular, lobate, serrate or filiform. Odour
absent. Reverse colourless, lacking diffusible pigment. On SNA,
hyphae hyaline, smooth-walled, lacking chlamydospores, aerial
mycelium sparse with moderate sporulation on the medium
surface. On CLA, aerial mycelium sparse with abundant pale
luteous to pale rosy vinaceous sporodochia forming on the
carnation leaves.
Additional material examined. south africa, Western Cape Province,
Piketberg, from Agathosma betulina, 2001, K. Lubbe, CBS 115423 = CPC
5311.
Notes — Fusarium callistephi formed a highly-supported
subclade in Clade III, closely related to F. cugenangense,
F. elaeidis and the untreated Fusarium clade. This species (co-
nidia 3–4 (–5)-septate) can be distinguished from F. cugen ang-
ense (conidia 3–6-septate; Maryani et al. 2019) and F. elaeidis
((1–)3 – 5-septate) based on septation of their macroconidia.
Additionally, F. cugenangense produces up to 3-septate micro-
conidia, a feature not seen in either F. callistephi or F. elaeidis.
Fusarium elaeidis readily formed polyphialidic conidiogenous
cells on the aerial mycelium, not seen in F. callistephi.

16 Persoonia – Volume 43, 2019
CBS 132477 (hoodiae)
CBS 144739
CBS 116619 (vasinfectum)
CBS 144743
CBS 128.81 (chrysanthemi)
CBS 221.49 (medicaginis)
CBS 187.53 (callistephi)
CBS 200.89 (basilici)
CBS 132476 (hoodiae)
Fusarium udum CBS 177.31
CPC 25822
CBS 102028 (cubense)
CBS 144742
CBS 144747
CBS 131393
CBS 130302
CBS 130323
CBS 102030 (cubense)
CBS 214.49 (dianthi)
CBS 680.89 (cucurbitacearum)
CBS 144134
CBS 255.52 (elaeidis)
CBS 115423
CBS 102026 (cubense)
CBS 144746
CBS 116611 (vasinfectum)
CBS 218.49 (elaeidis)
CBS 144749
CBS 144135
CBS 176.33 (lini)
CBS 144738
CBS 258.50 (batatas)
CBS 116613 (vasinfectum)
CBS 130308
CBS 217.49 (elaeidis)
CBS 144745
CBS 620.72 (gladioli)
CBS 119665
CBS 144741
CBS 130310 (cubense)
CBS 144740
CBS 130304 (vasinfectum)
CBS 132474 (hoodiae)
CBS 794.70 (perniciosum)
CBS 144748
CBS 116612 (vasinfectum)
Fusarium foetens CBS 120665
CBS 144744
100/50/1.0
80/85/1.0
-/-/0.99
-/-/0.99
97/96/1.0
78/66/0.95
-/-/0.99
99/78/0.99
-/-/1.0
66/-/-
89/66/1.0
84/71/1.0
100/50/1.0
-/-/0.96
95/96/1.0
100/100/1.0
96/96/1.0
79/-/1.0
78/68/1.0
63/61/-
100/99/1.0
52/54/-
84/-/1.0
77/73/1.0
99/98/1.0
100/99/1.0
x3
x3
x10
63/-/-
x3
Fusarium odoratissimum
Fusarium libertatis
Fusarium tradichlamydosporum
Fusarium duoseptatum
Fusarium fabacearum
Fusarium carminascens
Fusarium glycines
Fusarium gossypinum
Fusarium sp.
Fusarium callistephi
Fusarium elaeidis
Fusarium cugenangense
Fusarium hoodiae
Fusarium triseptatum
Fusarium oxysporum
I
II
III
IV
V
Fig. 1 The ML consensus tree inferred from the combined cmdA, rpb2, tef1 and tub2 sequence alignment. Thickened branches indicate branches present
in the ML, MP and Bayesian consensus trees. Support values (ML & MP bootstrap and posterior probability values) are indicated at the branches. The scale
bar indicates 0.02 expected changes per site. Clade numbers are provided on the right of the tree and these are used for reference in the treatment of the
species. The tree is rooted to F. foetens (CBS 120665) and F. udum (CBS 177.31). Epi- and ex-type strains are indicated in bold.

17
L. Lombard et al.: Epitypification of Fusarium oxysporum
0.02
CBS 646.78 (lycopersici)
CBS 129.24
CBS 117790
CBS 302.91 (lycopersici)
CBS 758.68 (lycopersici)
CBS 117461
CBS 141.95
CBS 114899
CBS 238.94 (meniscoideum)
CBS 123062
CBS 144751
CBS 130303 (radicis-lycopersici)
CBS 115416
CBS 840.88 (dianthi)
CBS 117787
CBS 127.81 (chrysanthemi)
CBS 300.91 (lycopersici)
CBS 115424
CBS 130300
NRRL 62542
CBS 109898
CBS 144750
NRRL 62545
NRRL 62547
CBS 645.78 (lycopersici)
CPC 30807
CBS 117791
CBS 247.61 (matthiolae)
CBS 181.32
CBS 129.81 (chrysanthemi)
CBS 111552
CBS 130301
CBS 119796
CBS 872.95 (radicis-lycopersici)
CBS 196.87 (bouvardiae)
CBS 413.90 (lycopersici)
CBS 115419
CBS 149.25 (cubense)
CBS 115417
NRRL 54996
NRRL 54984
CBS 117792
CBS 744.79 (passiflorae)
100/98/1.0
87/86/1.0
93/81/1.0
72/52/-
73/67/0.99
99/94/1.0
93/75/0.98
98/86/1.0
84/71/0.95
Fusarium languescens
Fusarium pharetrum
Fusarium contaminatum
Fusarium veterinarium
Fusarium curvatum
Fusarium nirenbergiae
VI
VII
VIII
Fig. 1 (cont.)

18 Persoonia – Volume 43, 2019
Fig. 2 Splitgraphs showing the results of the pairwise homoplasy index (PHI) test of newly described taxa using both LogDet transformation and splits decomposition. PHI test results (fW) < 0.05 indicate significant recombination
within the dataset. a. Representatives of all phylogenetic species resolved in this study; b. phylogenetic species in Clade III; c. phylogenetic species in Clades IV– VIII; d. phylogenetic species in Clade VII; e. isolates representing
F. nirenbergiae.
Fusarium udum CBS 177.31
Fusarium foetens CBS 120665
CBS 646.78
CBS 645.78
CBS 144751
CBS 144750
CBS 111552
CBS 117787
CBS 141.95
CBS 238.94
CBS 840.88
CBS 115416
CBS 144135
CBS 144134
CBS 144749
CBS 144747
Fusarium tradichlamydosporum CBS 102028
Fusarium duoseptatum CBS 102026
CBS 144746
CBS 200.89
CBS 144739
CBS 144738
CBS 144743
CBS 144744
CBS 116611
CBS 116612
CBS 115423
CBS 187.53
CBS 131393
CBS 128.81
CBS 680.89
CBS 258.50
CBS 116619
CBS 114899
CBS 109898
CBS 132474
CBS 132476
CBS 218.49
CBS 217.49
CBS 130304
Fusarium triseptatum
Fusarium sp.
Fusarium cugenangense
Fusarium elaeidis
Fusarium callistephi
Fusarium gossypinum
Fusarium fabacearum
Fusarium carminascens
Fusarium glycines
Fusarium libertatis
Fusarium hoodiae
Fusarium oxysporum
Fusarium nirenbergiae
Fusarium curvatum
Fusarium veterinarium
Fusarium phraretrum
Fusarium contaminatum
Fusarium languescens
CBS 217.49
CBS 187.53
CBS 115423
CBS 116612
CBS 116611
CBS 144738
CBS 144643
CBS 200.89
CBS144746
CBS 102028 Fusarium tradichlamydosporum
CBS 102026 Fusarium duoseptatum
CBS 680.89
CBS 128.81
CBS 131393
CBS 218.49
CBS 144739
CBS 144744
CBS 130304
0.001
Fusarium sp.
Fusarium cugenangense
Fusarium elaeidis
Fusarium callistephi
Fusarium gossypinum
Fusarium carminascens
Fusarium fabacearum
Fusarium glycines
CBS 258.50
CBS 144134
CBS 144135
CBS 132476
CBS 144751
CBS 144750
CBS 141.95
CBS 238.94
CBS 840.88
CBS 115416
CBS 116619
CBS 111552
CBS 645.78
CBS 132474
Fusarium oxysporum
Fusarium triseptatum
Fusarium nirenbergii
Fusarium curvatum
CBS 117787
CBS 109898
Fusarium veterinatum
Fusarium hoodiae
CBS 646.78
Fusarium languescens
Fusarium phraretrum
CBS 114899
Fusarium contaminatum
CBS 109898
CBS 144750
CBS 144751
CBS 111552
Fusarium phraretrum
CBS 114899
Fusarium contaminatum
CBS 117787
Fusarium veterinarium
CBS 129.24
CBS 115424
CPC 30807
CBS 130301
CBS 149.25
CBS 181.32
CBS 123062
CBS 115416
CBS 115417
CBS 758.68
CBS 840.88
CBS 130300
CBS 744.79
CBS 129.81
CBS 130303
CBS 196.87
CBS 115419
CBS 127.81
1.0E-4
W= 0.031
e
1.0E-4
W= 1.0
d
0,001
W= 0.43
a
W= 0.43
b
0.001
W= 0.06
c

19
L. Lombard et al.: Epitypification of Fusarium oxysporum
Fusarium carminascens L. Lombard, Crous & Lampr., sp.
nov. — MycoBank MB826835; Fig. 4
Etymology. Name refers to the almost carmine exudates this fungus
produces in its aerial mycelium when grown on PDA.
Typus. south africa, KwaZulu-Natal Province, from Zea mays, 2008,
S.C. Lamprecht (holotype CBS H-23609 designated here, culture ex-type
CBS 144738 = CPC 25800).
Conidiophores carried on the aerial mycelium 35 –55 µm tall,
unbranched or sparingly branched, bearing terminal or interca-
larily phialides, often reduced to single phialides; aerial phialides
mono- and polyphialidic, subulate to subcylindrical, smooth- and
thin-walled, 8–18 × 3–4 µm, periclinal thickening inconspicu-
ous or absent; aerial conidia forming small false heads on the
tips of the phialides, hyaline, ellipsoidal to falcate, smooth- and
thin-walled, 0–1-septate; 0-septate conidia: (5–)7–11(–12) ×
2–3(–4) µm (av. 9 × 3 µm); 1-septate conidia: (12–)13–15(–18)
× 2– 4 µm (av. 14 × 3 µm). Sporodochia bright orange, formed
abundantly on carnation leaves. Conidiophores in sporodochia
verticillately branched and densely packed, consisting of a
short, smooth- and thin-walled stipe, 4–9 × 2–4 µm, bearing
apical whorls of 2–3 monophialides or rarely as single lateral
monophialides; sporodochial phialides subulate to subcylin-
drical, 5–13 × 2 –4 µm, smooth- and thin-walled, sometimes
showing a reduced and flared collarette. Sporodochial conidia
falcate, curved dorsiventrally with almost parallel sides tapering
slightly towards both ends, with a blunt to papillate, curved api-
cal cell and a blunt to foot-like basal cell, (2–)3 –4(–5)-septate,
hyaline, smooth- and thin-walled; 2-septate conidia: 16–19 ×
3–4 µm (av. 18 × 3 µm); 3-septate conidia: (21–)26–36(–40) ×
3–5 µm (av. 31 × 4 µm); 4-septate conidia: (31–)33–43(–44) ×
4–5 µm (av. 38 × 4 µm); 5-septate conidia: 45– 51 × 4 µm (av.
48 × 4 µm). Chlamydospores globose to subglobose, formed
terminally, 4–8 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.1–4.0 mm/d at 24 °C. Colony surface
vinaceous purple to livid purple, floccose with abundant aerial
mycelium which produce an almost carmine exudate; colony
margins irregular, lobate, serrate or filiform. Odour absent.
Reverse dark livid to livid purple, lacking diffusible pigment.
On SNA, hyphae hyaline, smooth-walled, with abundant chla-
mydospores, aerial mycelium sparse with abundant sporulation
on the medium surface. On CLA, aerial mycelium sparse with
abundant bright orange sporodochia forming on the carnation
leaves.
Additional materials examined. south africa, KwaZulu-Natal Province,
from Zea mays, 2008, S.C. Lamprecht, CBS 144739 = CPC 25792, CBS
144740 = CPC 25793, CBS 144741 = CPC 25795.
Notes — Fusarium carminascens formed a well-supported
subclade in Clade III, closely related to F. fabacearum and
F. glycines. This species produced an almost carmine coloured
exudate in its aerial mycelium, a feature not observed in any of
the other strains studied here. Furthermore, F. carminascens
produces polyphialidic conidiogenous cells on its aerial myce-
lium, not observed in F. fabacearum or F. glycines.
Fig. 3 Fusarium callistephi (ex-type culture CBS 187.53). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light;
b. reverse of colony on PDA; c. conidiophores on surface of carnation leaf; d. sporodochia on carnation leaves; e– i. conidiophores and phialides on aerial my-
celium; j–k. sporodochia and sporodochial conidiophores; l. aerial conidia (microconidia); m. sporodochial conidia (macroconidia). — Scale bars: e– m = 10 µm.

20 Persoonia – Volume 43, 2019
Fusarium contaminatum L. Lombard & Crous, sp. nov. — Myco-
Bank MB826836; Fig. 5
Etymology. Name refers to the fact that this fungus was isolated from
contaminated food products.
Typus. GermaNy, Schluchtern, from pasteurized chocolate milk, Apr. 2004,
J. Houbraken (holotype CBS H-23610 designated here, culture ex-type CBS
114899).
Conidiophores carried on the aerial mycelium 15 –85 µm tall,
unbranched or branched, bearing a single terminal or a whorl
of 2–4 monophialides or intercalarily monophialides, often
reduced to single phia lides; aerial phialides subulate to sub-
cylindrical, smooth- and thin-walled, 7–22 × 2–5 µm, periclinal
thickening inconspicuous or absent; aerial conidia forming small
false heads on the tips of the phialides, hyaline, ellipsoidal
to falcate, smooth- and thin-walled, 0–1-septate; 0-septate
conidia: 5–9 (–11) × 2–4 µm (av. 7 × 3 µm); 1-septate conidia:
(9–)10–14 (–17) × 2–4 µm (av. 12 × 3 µm). Sporodochia bright
orange, formed sparsely on carnation leaves. Conidiophores in
sporodochia verticillately branched and densely packed, con-
sisting of a short, smooth- and thin-walled stipe, 7–13 × 4 µm,
bearing apical whorls of 2–3 monophialides or rarely as single
lateral monophialides; sporodochial phialides subulate to sub-
cylindrical, 4–9 × 2–3 µm, smooth- and thin-walled, sometimes
Fig. 4 Fusarium carminascens (ex-type culture CBS 144738). a –b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e– f. sporodochia on carnation leaves; g– j. conidiophores and phialides on
aerial mycelium; g– h. monophialides; i–j. polyphialides; k –l. chlamydospores; m– p. sporodochia and sporodochial conidiophores; o–p. phialides of sporo-
dochial conidiophores; q. aerial conidia (microconidia); r. sporodochial conidia (macroconidia). — Scale bars: g– r = 10 µm.

21
L. Lombard et al.: Epitypification of Fusarium oxysporum
showing a reduced and flared collarette. Sporodochial conidia
falcate, curved dorsiventrally with almost parallel sides tapering
slightly towards both ends, with a blunt to papillate, curved api-
cal cell and a blunt to foot-like basal cell, (2–)3-septate, hyaline,
smooth- and thin-walled; 2-septate conidia: (14–)15–17 × 3–4
µm (av. 16 × 3 µm); 3-septate conidia: (18 –)20– 26(–28) × 3–5
µm (av. 23 × 4 µm). Chlamydospores not observed.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.1–4.5 mm/d at 24 °C. Colony surface
white to pale vinaceous, floccose with abundant aerial myce-
lium; colony margins irregular, lobate, serrate or filiform. Odour
absent. Reverse rosy vinaceous, lacking diffusible pigment. On
SNA, hyphae hyaline, smooth-walled, lacking chlamydospores,
aerial mycelium sparse with abundant sporulation on the me-
dium surface. On CLA, aerial mycelium sparse with abundant
orange sporodochia forming on the carnation leaves.
Additional materials examined. NetherlaNds, from pasteurized fruit juice,
date and collector unknown, CBS 111552; from tetra pack with milky nutrition,
2005, collector unknown, CBS 117461.
Notes — Fusarium contaminatum formed a highly-supported
subclade in Clade VII, closely related to F. pharetrum and
F. veterinarium. This species produces small, 2 – 3-septate
macroconidia, whereas F. pharetrum produces much larger,
3(– 4)-septate macroconidia and F. veterinarium produces
slightly smaller, 1–(2–)3-septate macroconidia. None of these
three species produced any chlamydospores on SNA.
Fusarium curvatum L. Lombard & Crous, sp. nov. — Myco-
Bank MB826837; Fig. 6
Etymology. Name refers to the strongly curved sporodochial conidia pro-
duced by this fungus.
Typus. NetherlaNds, from Beaucarnia sp., 1994, J.W. Veenbaas-Rijks (holo-
type CBS H-23611 designated here, culture ex-type CBS 238.94 = NRRL
26422 = PD 94/184).
Conidiophores carried on the aerial mycelium 25 –56 µm tall,
unbranched or sparingly branched, bearing terminal or interca-
Fig. 5 Fusarium contaminatum (ex-type culture CBS 114899). a– b. Colony on PDA; a. Surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c –d. conidiophores on surface of carnation leaf; e–f. sporodochia on carnation leaves; g– k. conidiophores and phialides
on aerial mycelium; l. false head carried on phialide on aerial mycelium; m– p. sporodochia and sporodochial conidiophores; q. aerial conidia (microconidia);
r. sporodochial conidia (macroconidia). — Scale bars: g– l, q– r = 10 µm; m–p = 20 µm.

22 Persoonia – Volume 43, 2019
larily phialides, often reduced to single phialides or as phialidic
pegs; aerial phialides mono- and polyphialidic, subulate to sub-
cylindrical, smooth- and thin-walled, 3–30 × 2–5 µm, periclinal
thickening inconspicuous or absent; aerial conidia forming small
false heads on the tips of the phialides, hyaline, ellipsoidal
to falcate, smooth- and thin-walled, 0–1-septate; 0-septate
conidia: (4–)5 –9(–11) × 2–4 µm (av. 7 × 3 µm); 1-septate
conidia: (10–)11–13 × 2 –4 µm (av. 12 × 3 µm). Sporodochia
orange, formed abundantly on carnation leaves. Conidiophores
in sporodochia verticillately branched and densely packed,
consisting of a short, smooth- and thin-walled stipe, 8–10 × 2–4
µm, bearing apical whorls of 2–3 monophialides or rarely as
single lateral monophialides; sporodochial phialides subulate to
subcylindrical, 8–22 × 2– 4 µm, smooth- and thin-walled, some-
times showing a reduced and flared collarette. Sporodochial
conidia falcate, strongly curved or curved dorsiventrally with
almost parallel sides tapering slightly towards both ends, with
a blunt to papillate, curved apical cell and a blunt to foot-like
basal cell, (2–)3 –5-septate, hyaline, smooth- and thin-walled;
2-septate conidia: (15–)16– 22(– 23) × 3 –4 µm (av. 19 × 3 µm);
Fig. 6 Fusarium curvatum (ex-type culture CBS 238.94). a–b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light;
b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e–f. sporodochia on carnation leaves; g –i. conidiophores, monophialides and
polyphialides (arrows) on aerial mycelium; j. phialidic pegs on aerial mycelium; k– o. sporodochia and sporodochial conidiophores; p. aerial conidia (microco-
nidia); q. sporodochial conidia (macroconidia). — Scale bars: g– i, n = 20 µm; j, o–q = 10 µm, k– m = 50 µm.

23
L. Lombard et al.: Epitypification of Fusarium oxysporum
3-septate conidia: (18–)27– 39(–41) × 3– 5 µm (av. 33 × 4 µm);
4-septate conidia: (34–)37– 43(–46) × 3– 5 µm (av. 40 × 4 µm);
5-septate conidia: (30–)38 –46(–51) × 3–5 µm (av. 42 × 4 µm).
Chlamydospores not observed.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.1–4.5 mm/d at 24 °C. Colony surface
pale vinaceous to rosy vinaceous, floccose with abundant aerial
mycelium; colony margins irregular, lobate, serrate or filiform.
Odour absent. Reverse pale vinaceous, lacking diffusible pig-
ment. On SNA, hyphae hyaline, smooth-walled, lacking chlamy-
do spores, aerial mycelium sparse with abundant sporulation
on the medium surface. On CLA, aerial mycelium sparse with
abundant orange sporodochia forming on the carnation leaves.
Additional materials examined. GermaNy, Berlin-Dahlem, from Matthiola
incana, Feb. 1957, W. Gerlach, CBS 247.61 = BBA 8398 = DSM 62308 =
NRRL 22545. – NetherlaNds, from Hedera helix, 1994, J.W. Veenbaas-Rijks,
CBS 141.95 = NRRL 36251 = PD 94/1518.
Notes — Fusarium curvatum formed a highly-supported sub-
clade in Clade VIII, closely related to F. nirenbergiae. This
species produces strongly curved 3-septate macroconidia and
aerial polyphialidic conidiogenous cells, distinguishing it from
F. nirenbergiae. Additionally, F. curvatum failed to produce any
chlamydospores on SNA, whereas F. nirenbergiae produced
abundant chlamydospores.
Fusarium elaeidis L. Lombard & Crous, sp. nov. — MycoBank
MB826838; Fig. 7
Etymology. Name refers to the host plant genus Elaeis, from which this
fungus was first isolated.
Typus. Zaire, from Elaeis sp., 1949, T. Gogoi (holotype CBS H-23612
designated here, culture ex-type CBS 217.49 = NRRL 36358).
Conidiophores carried on the aerial mycelium 25 –65 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily phialides, often reduced to single phialides or as phia-
lidic pegs; aerial phialides mono- and polyphialidic, subulate
to subcylindrical, smooth- and thin-walled, 3–14 × 3– 4 µm,
periclinal thickening inconspicuous or absent; aerial conidia
forming small false heads on the tips of the phialides, hyaline,
ellipsoidal to falcate, smooth- and thin-walled, 0–1-septate;
0-septate conidia: 6–10(–13) × 2–3 µm (av. 8 × 3 µm); 1-sep-
tate conidia: (9–)11–15(–17) × 2–4(–5) µm (av. 13 × 3 µm).
Sporodochia pale rosy vinaceous to orange, formed abundantly
on carnation leaves. Conidiophores in sporodochia verticillately
Fig. 7 Fusarium elaeidis (ex-type culture CBS 217.49). a –b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light;
b. reverse of colony on PDA; c –d. conidiophores on surface of carnation leaf; e–f. sporodochia on carnation leaves; g. false head carried on a phialidic peg
on aerial mycelium; h. phialidic peg; i– j. conidiophores and phialides on aerial mycelium; j. polyphialide; k –l. sporodochia and sporodochial conidiophores;
m. aerial conidia (microconidia); n. sporodochial conidia (macroconidia). — Scale bars: g– n = 10 µm.

24 Persoonia – Volume 43, 2019
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 3–9 × 2 –3 µm, bearing apical whorls of
2–3 monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 8–12 × 2–4
µm, smooth- and thin-walled, sometimes showing a reduced
and flared collarette. Sporodochial conidia falcate, curved dor-
siventrally with almost parallel sides tapering slightly towards
both ends, with a blunt to papillate, curved apical cell and a blunt
to foot-like basal cell, (1–)3 – 5-septate, hyaline, smooth- and
thin-walled; 1-septate conidia: (14–)15– 25(– 32) × 2 –4 µm (av.
20 × 3 µm); 2-septate conidia: (17–)19 –25 × 3 – 4 µm (av. 22
× 4 µm); 3-septate conidia: (22–) 30–40(– 46) × (2–)3–4 µm
(av. 35 × 4 µm); 4-septate conidia: (34 –)36– 40(–43) × 3 – 5 µm
(av. 38 × 4 µm); 5-septate conidia: (36 –)37–43(–50) × 3– 5 µm
(av. 40 × 4 µm). Chlamydospores not observed.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 2.6–3.4 mm /d at 24 °C. Colony surface
pale rosy vinaceous grey, floccose with abundant aerial my-
celium; colony margins irregular, lobate, serrate or filiform.
Odour absent. Reverse pale rosy vinaceous, lacking diffusible
pigment. On SNA, hyphae hyaline, smooth-walled, lacking chla-
mydospores, aerial mycelium sparse with abundant sporulation
on the medium surface. On CLA, aerial mycelium sparse with
abundant pale rosy vinaceous to orange sporodochia forming
on the carnation leaves.
Additional materials examined. Zaire, from Elaeis sp., 1949, T. Gogoi,
CBS 218.49 = NRRL 36359. – uNkNowN locality, from Elaeis guineensis,
1952, J. Fraselle, CBS 255.52 = NRRL 36386.
Notes — Fusarium elaeidis formed a highly-supported sub-
clade in Clade III, closely related to F. callistephi, F. cugenan-
gense and the untreated Fusarium clade. See notes under
F. callistephi for distinguishing morphological features.
Fusarium fabacearum L. Lombard, Crous & Lampr., sp. nov.
— MycoBank MB826839; Fig. 8
Etymology. Name refers to the plant family, Fabaceae, which includes
the plant host Glycine max from which this fungus was first isolated.
Typus. south africa, North West Province, from Glycine max, 2010, S.C.
Lamprecht (holotype CBS H-23613 designated here, culture ex-type CBS
144743 = CPC 25802).
Conidiophores carried on the aerial mycelium 25 –50 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
11–15 × 3– 4 µm, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: (4–) 5– 9(–13) × 2–3 µm (av. 7 ×
Fig. 8 Fusarium fabacearum (ex-type culture CBS 144743). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c. conidiophores on surface of carnation leaf; d. sporodochia on carnation leaves; e. false head carried on a phialide on
aerial mycelium; f– h. conidiophores and phialides on aerial mycelium; i–k. sporodochia and sporodochial conidiophores; l. chlamydospore; m. aerial conidia
(microconidia); n. sporodochial conidia (macroconidia). — Scale bars: e– h, k–n = 10 µm; i– j = 50 µm.

25
L. Lombard et al.: Epitypification of Fusarium oxysporum
3 µm); 1-septate conidia: (12–)13 –15(–16) × 3– 4 µm (av. 14 ×
3 µm). Sporodochia pale luteous to orange, formed abundantly
on carnation leaves. Conidiophores in sporodochia verticillately
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 4–7 × 3 µm, bearing apical whorls of 2 – 3
monophialides or rarely as single lateral monophialides; sporo-
dochial phialides subulate to subcylindrical, 7–10 × 2 – 4 µm,
smooth- and thin-walled, sometimes showing a reduced and
flared collarette. Sporodochial conidia falcate, curved dorsiven-
trally with almost parallel sides tapering slightly towards both
ends, with a blunt to papillate, curved apical cell and a blunt to
foot-like basal cell, (1–)3 –4 (–5)-septate, hyaline, smooth- and
thin-walled; 1-septate conidia: (15–)16– 24(– 25) × 3 –4 µm (av.
20 × 3 µm); 3-septate conidia: (24 –)27–33(–36) × (2–)3– 5 µm
(av. 30 × 4 µm); 4-septate conidia: (32 –)33– 37(– 40) × 3 – 5 µm
(av. 35 × 4 µm); 5-septate conidia: (35–)38–44 × 3–4 µm (av.
41 × 4 µm). Chlamydospores globose to subglobose, formed
terminally, 5–8 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.0–4.4 mm /d at 24 °C. Colony surface
pale vinaceous grey to vinaceous grey, floccose with abundant
aerial mycelium; colony margins irregular, lobate, serrate or
filiform. Odour absent. Reverse pale vinaceous grey, lacking
diffusible pigment. On SNA, hyphae hyaline, smooth-walled,
with abundant chlamydospores, aerial mycelium sparse with
abundant sporulation on the medium surface. On CLA, aerial
mycelium sparse with abundant pale luteous to orange sporo-
dochia forming on the carnation leaves.
Additional materials examined. south africa, North West Province, from
Glycine max, 2010, S.C. Lamprecht, CBS 144744 = CPC 25803; from Zea
mays, 2008, C.M. Bezuidenhout, CBS 144742 = CPC 25801.
Notes — Fusarium fabacearum formed a highly-supported
subclade in Clade III, closely related to F. carminascens and
F. glycines. See notes under F. carminascens for distinguishing
morphological features.
Fusarium glycines L. Lombard, Crous & Lampr., sp. nov. —
MycoBank MB826840; Fig. 9
Etymology. Name refers to the plant genus Glycine from which this fungus
was isolated.
Typus. south africa, North West Province, from Glycine max, 2010, S.C.
Lamprecht (holotype CBS H-23614 designated here, culture ex-type CBS
144746 = CPC 25808).
Fig. 9 Fusarium glycines (ex-type culture CBS 144746). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light; b.
reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e– f. sporodochia on carnation leaves; g– i. conidiophores and phialides on aerial
mycelium; j –k. sporodochia and sporodochial conidiophores; l. chlamydospore; m. aerial conidia (microconidia); n. sporodochial conidia (macroconidia). — Scale
bars: g– i, l–n = 10 µm; j– k = 50 µm.

26 Persoonia – Volume 43, 2019
Conidiophores carried on the aerial mycelium 5– 45 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
15–25 × 2–4 µm, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: 7–11(–13) × 3– 4 µm (av. 9 × 3
µm); 1-septate conidia: (13 –)14–16(–18) × 3– 4 µm (av. 15 ×
3 µm). Sporodochia bright orange, formed abundantly on car-
nation leaves. Conidiophores in sporodochia verticillately branch-
ed and densely packed, consisting of a short, smooth- and thin-
walled stipe, 4–9 × 2– 4 µm, bearing apical whorls of 2–3
monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 12–14 ×
2–5 µm, smooth- and thin-walled, sometimes showing a
reduced and flared collarette. Sporodochial conidia falcate,
curved dorsiventrally with almost parallel sides tapering slightly
towards both ends, with a blunt to papillate, curved apical cell
and a blunt to foot-like basal cell, (1–)3 – 5-septate, hyaline,
smooth- and thin-walled; 1-septate conidia: 20–25 × 3– 4 µm
(av. 23 × 3 µm); 3-septate conidia: 37–43(–48) × 4–5 µm (av.
38 × 4 µm); 4-septate conidia: 44 – 46(–51) × 4– 5 µm (av. 42 ×
4 µm); 5-septate conidia: 43– 49(–52) × 4–5 µm (av. 46 × 4 µm).
Chlamydospores globose to subglobose, formed terminally,
4–8 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.0–4.4 mm /d at 24 °C. Colony surface
vinaceous, floccose with abundant aerial mycelium; colony
margins irregular, lobate, serrate or filiform. Odour absent.
Reverse vinaceous, lacking diffusible pigment. On SNA, hyphae
hyaline, smooth-walled, with abundant chlamydospores, aerial
mycelium sparse with abundant sporulation on the medium
surface. On CLA, aerial mycelium sparse with abundant bright
orange sporodochia forming on the carnation leaves.
Additional materials examined. arGeNtiNa, substrate unknown, date un-
known, C.J.M. Carrera, CBS 214.49 = NRRL 36356 = LCF F-245. – italy,
from Ocimum basilicum, 1989, G. Tamiette & A. Matta, CBS 200.89. – south
africa, North West Province, from Glycine max, 2010, S.C. Lamprecht, CBS
144745 = CPC 25804. – uNkNowN locality, from Linum usitatissium, 1933,
E.C. Stakman, CBS 176.33 = NRRL 36286.
Notes — Fusarium glycines formed a highly-supported sub-
clade in Clade III, closely related to F. carminascens and F. fa-
bacearum. See notes under F. carminascens for distinguishing
morphological features.
Fusarium gossypinum L. Lombard & Crous, sp. nov. — Myco-
Bank MB826841; Fig. 10
Etymology. Name refers to the plant genus Gossypium from which this
fungus was isolated.
Typus. ivory coast, Bouaké, wilted Gossypium hirsutum, Sept. 1995,
K. Abo (holotype CBS H-23615 designated here, culture ex-type CBS 116613).
Conidiophores carried on the aerial mycelium 35–75 µm tall, un-
branched or sparingly branched, bearing terminal or intercalarily
monophialides, often reduced to single phialides; aerial phial-
ides subulate to subcylindrical, smooth- and thin-walled, 3–30
× 2–4 µm, periclinal thickening inconspicuous or absent. Micro-
conidia forming small false heads on the tips of the phia lides,
hyaline, ellipsoidal to falcate, smooth- and thin-walled, 0–1-sep-
tate; 0-septate conidia: (5–)6 –8(–11) × 2–4 µm (av. 7 × 3 µm);
1-septate conidia: (11–)12–14(–15) × 2–4 µm (av. 15 × 3 µm).
Macroconidia also formed by phialides on aerial mycelium,
falcate, curved dorsiventrally with almost parallel sides taper-
ing slightly towards both ends, with a blunt to papillate, curved
Fig. 10 Fusarium gossypinum (ex-type culture CBS 116613). a –b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e. false head carried on a phialide on aerial mycelium; f– h. conidiophores
and phialides on aerial mycelium; i. aerial conidia (microconidia); j. sporodochial conidia (macroconidia). — Scale bars: e = 20 µm; f– j = 10 µm.

27
L. Lombard et al.: Epitypification of Fusarium oxysporum
apical cell and a blunt to foot-like basal cell, (1–)3-septate, hya-
line, smooth- and thin-walled; 1-septate conidia: 16–18 × 3 µm
(av. 17 × 3 µm); 2-septate conidia: 21–23 × 3 –4 µm (av. 22 ×
3 µm); 3-septate conidia: (24–)27–35(– 38) × 3 –4 µm (av. 31
× 4 µm). Sporodochia absent. Chlamydospores not observed.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 1.6–2.8 mm /d at 24 °C. Colony surface
white to pale rosy vinaceous, floccose with abundant aerial
mycelium; colony margins irregular, lobate, serrate or filiform.
Odour absent. Reverse pale rosy vinaceous, lacking diffusible
pigment. On SNA, hyphae hyaline, smooth-walled, lacking chla-
mydospores, aerial mycelium sparse with abundant sporulation
on the medium surface. On CLA, aerial mycelium sparse lacking
sporodochia on the carnation leaves.
Additional materials examined. ivory coast, Bouaké, wilted Gossypium
hirsutum, Sept. 1995, K. Abo, CBS 116611 and CBS 116612.
Notes — Fusarium gossypinum formed a unique highly-
supported subclade in Clade III. This species failed to produce
any sporodochia on the carnation leaf pieces, but still produced
abundant 3-septate macroconidia on the aerial mycelium. Other
species included in Clade III, all readily produced sporodochia
on carnation leaves.
Fusarium hoodiae L. Lombard, Crous & Lampr., sp. nov. —
MycoBank MB826842; Fig. 11
Etymology. Name refers to the plant genus Hoodia from which this fungus
was isolated.
Typus. south africa, Northern Cape Province, Prieska, root of Hoodia
gordonii, 2002, O.A. Philippou (holotype CBS H-23616 designated here,
culture ex-type CBS 132474).
Conidiophores carried on the aerial mycelium 40 –60 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
15–24 × 2–3 µm, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: (5–) 6–10(–16) × 2– 4 µm (av.
8 × 3 µm); 1-septate conidia: (11–)12–16(–17) × 3–4 µm (av.
Fig. 11 Fusarium hoodiae (ex-type culture CBS 132474). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light;
b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e–f. sporodochia on carnation leaves; g– h. conidiophores and phialides on aerial
mycelium; i –k. sporodochia and sporodochial conidiophores; l. chlamydospore; m. aerial conidia (microconidia); n. sporodochial conidia (macroconidia). — Scale
bars: g– h, l–n = 10 µm; i = 50 µm; j– k = 20 µm.

28 Persoonia – Volume 43, 2019
14 × 3 µm). Sporodochia pale vinaceous to light orange, formed
abundantly on carnation leaves. Conidiophores in sporodochia
verticillately branched and densely packed, consisting of a
short, smooth- and thin-walled stipe, 7–11 × 3–5 µm, bearing
apical whorls of 2–3 monophialides or rarely as single lateral
monophialides; sporodochial phialides subulate to subcylindri-
cal, 7–13 × 2–5 µm, smooth- and thin-walled, sometimes show-
ing a reduced and flared collarette. Sporodochial conidia falcate,
curved dorsiventrally with almost parallel sides tapering slightly
towards both ends, with a blunt to papillate, curved apical cell
and a blunt to foot-like basal cell, (1–)3 (– 4)-septate, hyaline,
smooth- and thin-walled; 1-septate conidia: 20–33 × 3– 5 µm
(av. 25 × 4 µm); 3-septate conidia: (20 –)27–39 (–45) × 3– 5 µm
(av. 33 × 4 µm); 4-septate conidia: (35–)36–46(–51) × 4–5
µm (av. 41 × 5 µm). Chlamydospores globose to subglobose,
formed terminally, 4–11 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.1–4.5 mm/d at 24 °C. Colony surface
pale vinaceous grey to livid vinaceous, floccose with abundant
aerial mycelium; colony margins irregular, lobate, serrate or
filiform. Odour absent. Reverse livid purple to pale vinaceous
grey, lacking diffusible pigment. On SNA, hyphae hyaline,
smooth-walled, with abundant chlamydospores, aerial myce-
lium sparse with abundant sporulation on the medium surface.
On CLA, aerial mycelium sparse with abundant pale vinaceous
to light orange sporodochia forming on the carnation leaves.
Additional materials examined. south africa, Northern Cape Province,
Prieska, root of Hoodia gordonii, 2002, O.A. Philippou, CBS 132476, CBS
132477.
Notes — Fusarium hoodiae formed a weakly supported
clade constituting Clade IV in this phylogenetic study. All three
isolates studied here, produced pale vinaceous to pale orange
sporodochia on the carnation leaf pieces, unique for all the
isolates studied.
Fusarium languescens L. Lombard & Crous, sp. nov. — Myco-
Bank MB826843; Fig. 12
Etymology. Name refers to the wilting symptoms associated with infec-
tions of this fungus.
Typus. morocco, Solanum lycopersicum, date and collector unknown (holo-
type CBS H-23617 designated here, culture ex-type CBS 645.78 = NRRL
36531).
Conidiophores carried on the aerial mycelium 25 –30 µm
tall, unbranched or sparingly branched, bearing terminal or
intercalarily monophialides, often reduced to single phialides;
aerial phialides subulate to subcylindrical, smooth- and thin-
walled, 7–22 × 2–4 µm, periclinal thickening inconspicuous or
absent; aerial conidia forming small false heads on the tips of
the phialides, hyaline, ellipsoidal to falcate, smooth- and thin-
walled, 0–1-septate; 0-septate conidia: (4–) 5–9(–12) × 2–3
Fig. 12 Fusarium languescens (ex-type culture CBS 645.78). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c. conidiophores on surface of carnation leaf; d. sporodochia on carnation leaves; e– h. conidiophores and phialides on
aerial mycelium; i– k. sporodochia and sporodochial conidiophores; l. chlamydospore; m. aerial conidia (microconidia); n. sporodochial conidia (macroco-
nidia). — Scale bars: e– h, l–n = 10 µm; i– k = 20 µm.

29
L. Lombard et al.: Epitypification of Fusarium oxysporum
µm (av. 7 × 3 µm); 1-septate conidia: (9–)11–15 × 2–4 µm (av.
13 × 3 µm). Sporodochia light orange, formed abundantly on
carnation leaves. Conidiophores in sporodochia verticillately
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 5–10 × 3–4 µm, bearing apical whorls of
2–3 monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 10–14 × 2–4
µm, smooth- and thin-walled, sometimes showing a reduced
and flared collarette. Sporodochial conidia falcate, curved dor-
siventrally with almost parallel sides tapering slightly towards
both ends, with a blunt to papillate, curved apical cell and a
blunt to foot-like basal cell, 1–3(– 5)-septate, hyaline, smooth-
and thin-walled; 1-septate conidia: (15–)18– 23(– 30) × 3 –4 µm
(av. 20 × 3 µm); 2-septate conidia: (14–)16–22(–24) × 4 µm
(av. 19 × 3 µm); 3-septate conidia: (22 –)26–38(–47) × 3–5
µm (av. 32 × 4 µm); 5-septate conidia: 32–40 × 4 –5 µm (av.
36 × 5 µm). Chlamydospores globose to subglobose, formed
terminally, 6–9 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.1–4.5 mm/d at 24 °C. Colony surface
flesh to rosy vinaceous, floccose with abundant aerial myce-
lium; colony margins irregular, lobate, serrate or filiform. Odour
absent. Reverse pale luteous, lacking diffusible pigment. On
SNA, hyphae hyaline, smooth-walled, with abundant chlamydo-
spores, aerial mycelium sparse with abundant sporulation on
the medium surface. On CLA, aerial mycelium sparse with
abundant light orange sporodochia forming on the carnation
leaves.
Additional materials examined. israel, Bet Dagan, Solanum lycopersicum,
1986, R. Cohn, CBS 413.90 = ATCC 66046 = NRRL 36465. – morocco, Sola-
num lycopersicum, date and collector unknown, CBS 646.78 = NRRL 36532.
– NetherlaNds, Solanum lycopersicum, 1991, D.H. Elgersma, CBS 300.91 =
NRRL 36416, CBS 302.91 = NRRL 36419. – south africa, Zea mays, date and
collector unknown, CBS 119796 = MRC 8437. – uNkNowN locality, Solanum
lycopersicum, date and collector unknown, CBS 872.95 = NRRL 36570.
Notes — Fusarium languescens forms the highly-supported
Clade VI, which mostly includes strains associated with tomato
wilt. This species displays morphological overlap with several
species treated here. Therefore, phylogenetic inference is
needed to accurately identify this species.
Fusarium libertatis L. Lombard, Crous, sp. nov. — MycoBank
MB826844; Fig. 13
Etymology. Name refers to ‘freedom’. Fusarium libertatis was isolated from
the rock surfaces in the stone quarry on Robben Island where the prisoners
were forced to work. It is named in remembrance of all those who through the
centuries were incarcerated on the Island for their different political beliefs.
Typus. south africa, Western Cape Province, Robben Island, Van Rie-
beeck’s Quarry, from rock surfaces, May 2015, P.W. Crous (holotype CBS
H-23618 designated here, culture ex-type CBS 144749 = CPC 28465).
Conidiophores carried on the aerial mycelium 2– 30 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily phialides, often reduced to single phialides; aerial
phialides mono- and polyphialidic, subulate to subcylindrical,
smooth- and thin-walled, 8–13 × 2–4 µm, sometimes proliferat-
ing percurrently, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: (6–)7– 9(–11) × 2 – 4 µm (av.
8 × 3 µm); 1-septate conidia: (11–)12–14(–15) × 2–4 µm (av.
13 × 3 µm). Sporodochia bright orange, formed abundantly on
carnation leaves. Conidiophores in sporodochia verticillately
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 4–8 × 3 –4 µm, bearing apical whorls of
2–3 monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 6–12 × 2–4
µm, smooth- and thin-walled, sometimes showing a reduced
and flared collarette. Sporodochial conidia falcate, curved dor-
siventrally with almost parallel sides tapering slightly towards
both ends, with a blunt to papillate, curved apical cell and a
blunt to foot-like basal cell, 1–3-septate, hyaline, smooth- and
thin-walled; 1-septate conidia: (15–)17–21(–23) × 2–4 µm (av.
19 × 3 µm); 2-septate conidia: (18 –)20– 24(–25) × 2–3(–4) µm
(av. 22 × 4 µm); 3-septate conidia: (24–)30– 38(–40) × (2–)3–5
µm (av. 34 × 4 µm). Chlamydospores globose to subglobose,
formed terminally and intercalarily, carried singly, 5–9 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 2.3–4.4 mm /d at 24 °C. Colony surface
vinaceous, floccose with abundant aerial mycelium; colony
margins irregular, lobate, serrate or filiform. Odour absent.
Reverse vinaceous, lacking diffusible pigment. On SNA, hyphae
hyaline, smooth-walled, with abundant chlamydospores, aerial
mycelium sparse with abundant sporulation on the medium
surface. On CLA, aerial mycelium sparse with abundant bright
orange sporodochia forming on the carnation leaves.
Additional materials examined. south africa, Western Cape Province,
from Aspalathus sp., 2008, C.M. Bezuidenhout, CBS 144747 = CPC 25788,
CBS 144748 = CPC 25782.
Notes — Fusarium libertatis formed a unique well-supported
clade Clade (II). This species readily produced polyphialidic
conidio genous cells on its aerial mycelium and can be distin-
guished from the other species (F. carminascens, F. curvatum
and F. elaeidis) found to produce polyphialides by only produc-
ing up to 3-septate macroconidia, whereas the other polyphi-
alidic species produce up to 5-septate macroconidia.
Fusarium nirenbergiae L. Lombard & Crous, sp. nov. — Myco-
Bank MB826845; Fig. 14
Etymology. Named in honour of Prof. H.I. Nirenberg for her contribution
to our understanding of Fusarium taxonomy.
Typus.
NetherlaNds, Aalsmeer, from Dianthus caryophyllus, 1988, H. Rat-
tink (holotype CBS H-23619 designated here, culture ex-type CBS 840.88).
Conidiophores carried on the aerial mycelium 18 –50 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
8–24 × 2–4 µm, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: (5–) 6–10(–11) × 2– 4 µm (av.
8 × 3 µm); 1-septate conidia: (9–)10–14(–15) × 2–4 µm (av.
12 × 3 µm). Sporodochia bright orange, formed abundantly on
carnation leaves. Conidiophores in sporodochia verticillately
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 6–14 × 3–5 µm, bearing apical whorls of
2–3 monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 8–18 × 2–4
µm, smooth- and thin-walled, sometimes showing a reduced
and flared collarette. Sporodochial conidia falcate, curved dor-
siventrally with almost parallel sides tapering slightly towards
both ends, with a blunt to papillate, curved apical cell and a
blunt to foot-like basal cell, 1–5-septate, hyaline, smooth- and
thin-walled; 1-septate conidia: 15–29 (–34) × 3–4 µm (av. 22
× 4 µm); 2-septate conidia: (18 –)19 –31(–39) × 2– 4(–5) µm
(av. 25 × 3 µm); 3-septate conidia: (30 –)32– 40(–43) × 3 – 4 µm
(av. 36 × 4 µm); 4-septate conidia: (34 –)36– 44(–48) × 3 – 5 µm
(av. 40 × 4 µm); 5-septate conidia: (36 –)43– 59(–66) × 3 – 5 µm
(av. 51 × 4 µm). Chlamydospores globose to subglobose,
formed terminally, 4–6 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 2.9–4.2 mm /d at 24 °C. Colony surface

30 Persoonia – Volume 43, 2019
pale vinaceous to vinaceous, floccose with abundant aerial
mycelium; colony margins irregular, lobate, serrate or filiform.
Odour absent. Reverse pale vinaceous grey to greyish lilac,
lacking diffusible pigment. On SNA, hyphae hyaline, smooth-
walled, with abundant chlamydospores, aerial mycelium sparse
with moderate sporulation on the medium surface. On CLA,
aerial mycelium sparse with abundant bright orange sporo-
dochia forming on the carnation leaves.
Additional materials examined. BraZil, from Passiflora edulis, 1968,
W. Gerlach, CBS 744.79 = BBA 62355 = NRRL 22549. – italy, Napoli, Castel-
lammare di Stabia, from Bouvardia longiflora, July 1986, B. Aloj, CBS 196.87
= NRRL 26219. – NetherlaNds, Berkel, from Solanum lycopersicum, 16 May
1968, G. Weststeijn, CBS 758.68 = NRRL 36546. – south africa, Western
Cape Province, Riebeeck-Wes, from Agathosma betulina, 2001, K. Lubbe,
CBS 115424 = CPC 5312; Stellenbosch, Elsenberg farm, from Agathosma
betulina, 2001, K. Lubbe, CBS 115416 = CPC 5307, CBS 115417 = CPC
5306, CBS 115419 = CPC 5308. – USA, California, from amputated human
toe, unknown date and collector, CBS 130300 = NRRL 26368; Florida, from
Solanum tuberosum, 1923, H.W. Wollenweber, CBS 181.32 = NRRL 36303;
from Chrysanthemum sp., date unknown, G.M. Armstrong & J.K. Armstrong,
CBS 127.81 = BBA 63924 = NRRL 36229; Florida, from Chrysanthemum sp.,
date unknown, A.W. Engelhard, CBS 129.81 = BBA 63926 = NRRL 22539;
Maryland, Beltsville, from tulip roots, 1991, R.L. Lumsden, CBS 123062 =
GJS 91-17; Florida, Immokalee, from Solanum lycopersicum, date unknown,
J. Swezey, CBS 130303; Texas, San Antonio, from human leg ulcer, date
and collector unknown, CBS 130301 = NRRL 26374. – uNkNowN locality,
from Secale cereale, date unknown, H.W. Wollenweber, CBS 129.24; from
Musa sp., date unknown, E.W. Mason, CBS 149.25 = NRRL 36261.
Fig. 13 Fusarium libertatis (ex-type culture CBS 144749). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light;
b. reverse of colony on PDA; c –e. conidiophores on surface of carnation leaf; g–k. conidiophores and phialides on aerial mycelium; g –h. monophialides;
i– k. polyphialides; l –n. sporodochia and sporodochial conidiophores; n. phialides of sporodochial conidiophores; o – p. chlamydospores; q. aerial conidia
(microconidia); r. sporodochial conidia (macroconidia). — Scale bars: c– r = 10 µm.

31
L. Lombard et al.: Epitypification of Fusarium oxysporum
Notes — Fusarium nirenbergiae formed a well-supported
subclade in Clade VIII, closely related to F. curvatum. See notes
under F. curvatum for distinguishing morphological features.
Fusarium oxysporum Schltdl., Fl. Berol. 2: 139. 1824 — Fig.
15
Synonyms. Fusarium bulbigenum Cooke & Massee, Grevillea 16: 49.
1887.
Fusarium vasinfectum G.F.Atk., Bull. Alabama Agric. Exper. Station 41:
19. 1892.
Fusarium dianthi Prill. & Delacr., Compt. Rend. Acad. Sci. 129: 744. 1899.
Fusarium lini Bolley, Proc. Ann. Meeting Soc. Prom. Agr. Sci. 21: 1– 4.
1902.
Fusarium orthoceras Appel & Wollenw., Arb. Kaiserl. Biol. Anst. Ld.- u.
Forstw. 8: 152. 1910.
Fusarium citrinum Wollenw., Maine Agric. Exp. Sta. Bull. 219: 256. 1913.
Fusarium angustum Sherb., Cornell Univ. Agric. Exp. Sta. Mem. 6: 203.
1915.
Fusarium lutulatum Sherb., Cornell Univ. Agric. Exp. Sta. Mem. 6: 209.
1915.
Fusarium bostrycoides Wollenw. & Reinking, Phytopathology 15: 166.
1925.
Diplosporium vaginae Nann., Atti Reale Accad. Fisiocrit. Siena sér. 4, 17:
491. 1926.
For additional synonyms see Index Fungorum and MycoBank.
Typification. GermaNy, Berlin, from rotten tuber of Solanum tuberosum,
1824, D.L.F. von Schlechtendal, HAL 1612 F, holotype in HAL; (epitype
designated here: GermaNy, Berlin, from rotten tuber of Solanum tuberosum,
17 Oct. 2017, L. Lombard, epitype CBS H-23620, MBT382397, culture ex-
epitype CBS 144134).
Conidiophores carried on the aerial mycelium 15–75 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
11–40 × 2 – 4 µm, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: (4–) 6–10(–11) × 2– 4 µm (av.
8 × 3 µm); 1-septate conidia: 13–15(–16) × 2 – 4 µm (av. 14
× 3 µm). Sporodochia bright orange, formed abundantly on
carnation leaves. Conidiophores in sporodochia verticillately
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 4–10 × 4–5 µm, bearing apical whorls of
2–3 monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 8–13 × 3–5
µm, smooth- and thin-walled, sometimes showing a reduced
and flared collarette. Sporodochial conidia falcate, curved dor-
siventrally with almost parallel sides tapering slightly towards
both ends, with a blunt to papillate, curved apical cell and a
blunt to foot-like basal cell, (1–)3(–5)-septate, hyaline, smooth-
and thin-walled; 1-septate conidia: (21–)22 –26 × 4–5 µm (av.
24 × 4 µm); 2-septate conidia: 20– 26(–27) × 4– 5 µm (av. 23
× 4 µm); 3-septate conidia: (22–)25 –29(– 31) × 4–5 µm (av.
27 × 4 µm); 4-septate conidia: (30–)31– 35 × 4 – 5 µm (av. 33
× 5 µm); 5-septate conidia: 35–38 × 5–6 µm (av. 37 × 5 µm).
Chlamydospores globose to subglobose, formed intercalarily
or terminally, 5–10 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.0–4.0 mm /d at 24 °C. Colony surface
pale vinaceous, floccose with abundant aerial mycelium; colony
margins irregular, lobate, serrate or filiform. Odour absent. Re-
verse vinaceous to rosy vinaceous, lacking diffusible pigment.
On SNA, hyphae hyaline, smooth-walled, producing abundant
chlamydospores, aerial mycelium sparse with abundant sporu-
lation on the medium surface. On CLA, aerial mycelium sparse
with abundant bright orange sporodochia forming on the carna-
tion leaves.
Fig. 14 Fusarium nirenbergiae (ex-type culture CBS 840.88). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c. conidiophores on surface of carnation leaf; d. sporodochia on carnation leaves; e. conidiophores and phialides on aerial
mycelium; f– g. sporodochia and sporodochial conidiophores; h. chlamydospore; i. aerial conidia (microconidia); j. sporodochial conidia (macroconidia). — Scale
bars: e, h– j = 10 µm; f–g = 50 µm.

32 Persoonia – Volume 43, 2019
Additional materials examined. GermaNy, from rotten tuber of Solanum
tuberosum, 17 Oct. 2017, L. Lombard, CBS 144135. – south africa, Western
Cape Province, from Protea sp., date unknown, C.M. Bezuidenhout, CPC
25822. – south east asia, from Camellia sinensis, 1949, F. Bugnicourt, CBS
221.49 = IHEM 4508 = NRRL 22546.
Notes — Fusarium oxysporum formed a well-supported
subclade in Clade V with F. triseptatum as closest relative.
Both species in Clade V displayed some morphological over-
lap. However, the 1-septate ((21–)22–26 × 4–5 µm (av. 24 ×
4 µm) and 2-septate (20–26(–27) × 4 – 5 µm (av. 23 × 4 µm)
macroconidia of F. oxysporum are larger than those of F. tri-
septatum ((18–)19– 23(–24) × 3 –4 µm (av. 20 × 3 µm) and
17–25(– 26) × 3 µm (av. 21 × 3 µm), respectively), whereas
the 3-septate ((25–)27– 39(–47) × 4–5 µm (av. 33 × 3 µm)),
4-septate ((31–)34 – 40(– 41) × 4– 5 µm (av. 37 × 4 µm)) and
5-septate ((33–48 (–49) × 4–5 µm (av. 40 × 4 µm)) macroco-
nidia of F. triseptatum are larger than those of F. oxysporum
((22–)25 –29(– 31) × 4– 5 µm (av. 27 × 4 µm), (30–)31– 35 ×
4–5 µm (av. 33 × 5 µm) and 35– 38 × 5– 6 µm (av. 37 × 5 µm),
respectively). Additionally, all isolates of F. oxysporum produced
abundant bright orange sporodochia on carnation leaf pieces,
not observed for any of the F. triseptatum isolates studied.
Fusarium pharetrum L. Lombard & Crous, sp. nov. — Myco-
Bank MB826846; Fig. 16
Etymology. Name refers to the practice of the Southern African indige-
nous San people of hollowing out the tubular branches of the host plant,
Aloidendron dichotomum, to form quivers (Latin pharetra) for their arrows.
Fig. 15 Fusarium oxysporum (ex-epitype culture CBS 144134). a–b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e –f. sporodochia on carnation leaves; g –j. conidiophores and phialides
on aerial mycelium; k– n. sporodochia and sporodochial conidiophores; o– p. chlamydospores; q. aerial conidia (microconidia); r. sporodochial conidia (macro-
conidia). — Scale bars: g– h, m–r = 10 µm; i– l = 50 µm.

33
L. Lombard et al.: Epitypification of Fusarium oxysporum
Typus. south africa, from Aliodendron dichotomum, 2000, F. van der
Walt & G.J. Marais (holotype CBS H-23621 designated here, culture ex-type
CBS 144751 = CPC 30824).
Conidiophores carried on the aerial mycelium 20–75 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
4–28 × 2–5 µm, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: 5– 9(–13) × 2– 3 µm (av. 7 × 3
µm); 1-septate conidia: (10–)12–16 (–18) × 2– 4 µm (av. 14 × 3
µm). Sporodochia rosy vinaceous to orange, formed abundantly
on carnation leaves. Conidiophores in sporodochia verticillately
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 5–10 × 3–5 µm, bearing apical whorls of
2–3 monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 7–13 × 3–4
µm, smooth- and thin-walled, sometimes showing a reduced
and flared collarette. Sporodochial conidia falcate, curved dor-
siventrally with almost parallel sides tapering slightly towards
both ends, with a blunt to papillate, curved apical cell and a
blunt to foot-like basal cell, 3(– 4)-septate, hyaline, smooth- and
thin-walled; 3-septate conidia: (22–)27– 35(–39) × 3–5 µm (av.
31 × 4 µm); 4-septate conidia: (34 –)36–40(–41) × 3–5 µm (av.
36 × 5 µm). Chlamydospores not observed.
Fig. 16 Fusarium pharetum (ex-type culture CBS 144751). a– b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light;
b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e –f. sporodochia on carnation leaves; g –h. false heads carried on a phialide
on aerial mycelium; i– l. conidiophores and phialides on aerial mycelium; m– p. sporodochia and sporodochial conidiophores; q. aerial conidia (microconidia);
r. sporodochial conidia (macroconidia). — Scale bars: g– l, q– r = 10 µm; m–p = 50 µm.

34 Persoonia – Volume 43, 2019
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.1–4.5 mm/d at 24 °C. Colony surface
rosy vinaceous, floccose with abundant aerial mycelium; colony
margins irregular, lobate, serrate or filiform. Odour absent.
Reverse rosy vinaceous, lacking diffusible pigment. On SNA,
hyphae hyaline, smooth-walled, lacking chlamydospores, aerial
mycelium sparse with abundant sporulation on the medium
surface. On CLA, aerial mycelium sparse with abundant rosy
vinaceous to orange sporodochia forming on the carnation
leaves.
Additional material examined. south africa, from Aliodendron dichoto-
mum, 2000, F. van der Walt & G.J. Marais, CBS 144750 = CPC 30822.
Notes — Fusarium pharetrum formed a well-supported sub-
clade in Clade VII, closely related to F. contaminatum and F. vete-
rinarium. See notes under F. contaminatum for distinguishing
morphological features.
Fusarium triseptatum L. Lombard & Crous, sp. nov. — Myco-
Bank MB826847; Fig. 17
Etymology. Name refers to the abundant 3-septate macroconidia pro-
duced by this fungus.
Typus. USA, locality unknown, from Ipomoea batatas, 1950, T.T. McClure
(holotype CBS H-23622 designated here, culture ex-type CBS 258.50 =
NRRL 36389).
Conidiophores carried on the aerial mycelium 5– 40 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
6–22 × 2–4 µm, periclinal thickening inconspicuous or absent.
Microconidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: (5–) 6–10(–13) × 1– 3 µm (av.
8 × 3 µm); 1-septate conidia: (12–)14–16(–18) × 2 –4 µm (av.
15 × 3 µm). Macroconidia also formed by phialides on aerial
mycelium, falcate, curved dorsiventrally with almost paral-
lel sides tapering slightly towards both ends, with a blunt to
papillate, curved apical cell and a blunt to foot-like basal cell,
(1–)3(–5)-septate, hyaline, smooth- and thin-walled; 1-septate
conidia: (18–)19– 23(– 24) × 3 –4 µm (av. 20 × 3 µm); 2-septate
conidia: 17–25(– 26) × 3 µm (av. 21 × 3 µm); 3-septate conidia:
(25–)27–39(– 47) × 4–5 µm (av. 33 × 3 µm); 4-septate conidia:
(31–)34 –40(– 41) × 4–5 µm (av. 37 × 4 µm); 5-septate conidia:
33–48 (–49) × 4 –5 µm (av. 40 × 4 µm). Sporodochia absent.
Chlamydospores globose to subglobose, formed terminally,
5–12 µm diam.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 2.2–3.4 mm /d at 24 °C. Colony surface
pale vinaceous grey to vinaceous grey, floccose with abundant
aerial mycelium; colony margins irregular, lobate, serrate or
filiform. Odour absent. Reverse pale vinaceous grey, lacking
diffusible pigment. On SNA, hyphae hyaline, smooth-walled,
Fig. 17 Fusarium triseptatum (ex-type culture CBS 258.50). a–b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white light;
b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e– f. false heads carried on a phialide on aerial mycelium; g–j. conidiophores
and phialides on aerial mycelium; k– l. chlamydospores; m. microconidia; n. macroconidia. — Scale bars: e, g–n = 10 µm; f = 20 µm.

35
L. Lombard et al.: Epitypification of Fusarium oxysporum
with abundant chlamydospores, aerial mycelium sparse with
abundant sporulation on the medium surface. On CLA, aerial
mycelium sparse lacking sporodochia on the carnation leaves.
Additional materials examined. ivory coast, Béoumi, wilted Gossypium
hirsutum, Oct. 1996, K. Abo, CBS 116619. – PaPua New GuiNea, Suki village,
from sago starch, 2005, A. Greenhill, CBS 119665. – USA, Tennessee, from
human eye, collector and date unknown, CBS 130302 = NRRL 26360 = FRC
755.
Notes — Fusarium triseptatum formed a highly-supported
subclade in Clade V, closely related to F. oxysporum. See notes
under F. oxysporum for distinguishing morphological features.
Fusarium veterinarium L. Lombard & Crous, sp. nov. — Myco-
Bank MB826849; Fig. 18
Etymology. Name refers to the fact that this fungus was isolated mostly
from veterinary samples.
Typus. NetherlaNds, from shark peritoneum, date unknown, C. Hoek (holo-
type CBS H-23623 designated here, culture ex-type CBS 109898 = NRRL
36153).
Conidiophores carried on the aerial mycelium 12 –90 µm tall,
unbranched or sparingly branched, bearing terminal or inter-
calarily monophialides, often reduced to single phialides; aerial
phialides subulate to subcylindrical, smooth- and thin-walled,
8–24 × 2–4 µm, periclinal thickening inconspicuous or absent;
aerial conidia forming small false heads on the tips of the phia-
lides, hyaline, ellipsoidal to falcate, smooth- and thin-walled,
0–1-septate; 0-septate conidia: (4–) 6–8(–11) × 2–4 µm (av.
7 × 3 µm); 1-septate conidia: (9–)10–14(–15) × 2–4 µm (av.
12 × 3 µm). Sporodochia bright orange, formed abundantly on
carnation leaves. Conidiophores in sporodochia verticillately
branched and densely packed, consisting of a short, smooth-
and thin-walled stipe, 8–13 × 3–4 µm, bearing apical whorls of
2–3 monophialides or rarely as single lateral monophialides;
sporodochial phialides subulate to subcylindrical, 10–15 × 2–4
µm, smooth- and thin-walled, sometimes showing a reduced
and flared collarette. Sporodochial conidia falcate, curved dor-
siventrally with almost parallel sides tapering slightly towards
both ends, with a blunt to papillate, curved apical cell and a blunt
to foot-like basal cell, 1–(2 –) 3-septate, hyaline, smooth- and
Fig. 18 Fusarium veterinarium (ex-type culture CBS 109898). a –b. Colony on PDA; a. surface of colony on PDA after 7 d at 24 °C under continuous white
light; b. reverse of colony on PDA; c– d. conidiophores on surface of carnation leaf; e –f. sporodochia on carnation leaves; g –i. conidiophores and phialides
on aerial mycelium; j– l. sporodochia and sporodochial conidiophores; m. aerial conidia (microconidia); n. sporodochial conidia (macroconidia). — Scale bars:
g– n = 10 µm.

36 Persoonia – Volume 43, 2019
thin-walled; 1-septate conidia: (12–)15–19(–20) × 3–4 µm (av.
17 × 3 µm); 2-septate conidia: (16–)17–21(– 24) × 3–4 µm (av.
19 × 3 µm); 3-septate conidia: (19 –)20–24(–27) × 3–4 µm (av.
22 × 3 µm). Chlamydospores not observed.
Culture characteristics — Colonies on PDA with an average
radial growth rate of 3.1–4.5 mm/d at 24 °C. Colony surface
pale vinaceous grey, floccose with moderate aerial mycelium
appearing wet; colony margins irregular, lobate, serrate or
filiform. Odour absent. Reverse pale vinaceous, lacking diffusi-
ble pigment. On SNA, hyphae hyaline, smooth-walled, lacking
chlamydospores, aerial mycelium sparse with abundant sporu-
lation on the medium surface. On CLA, aerial mycelium sparse
with abundant orange sporodochia forming on the carnation
leaves.
Additional materials examined. NetherlaNds, from swab sample near
filling apparatus, Apr. 2005, J. Houbraken, CBS 117787, CBS 117790; from
pasteurized milk-based product, Apr. 2005, J. Houbraken, CBS 117791,
CBS 117792. – USA, California, from endoscope of veterinary clinic, date
and collector unknown, NRRL 62545; from canine stomach, date and col-
lector unknown, NRRL 62547; Massachusetts, from mouse mucosa, date
and collector unknown, NRRL 54984; from little blue penguin foot, date and
collector unknown, NRRL 54996; Texas, from unknown animal faeces, date
and collector unknown, NRRL 62542.
Notes — Fusarium veterinarium formed a highly-supported
subclade in Clade VII, closely related to F. contaminatum and
F. pharetrum. See notes under F. contaminatum for distinguish-
ing morphological features.
DISCUSSION
Fusarium taxonomy and the underlying phylogenetic backbone
on which it is based, is undergoing continuous revision. In
modern day fungal taxonomy, phylogenetic inference plays a
vital role to resolve the identity of cryptic species due to the
paucity of morphological features. However, a key component
of a robust phylogeny is the availability of living ex-type mate-
rial to serve as basic reference point or ‘phylogenetic anchor’
on which comparative taxonomy can be based (Booth 1975).
Epi- and/or neotypification provides a vital means where upon
stability can be enforced into a chaotic classification system as
being applied to F. oxysporum today.
Snyder & Hansen’s (1940) treatment of the section Elegans to
represent only F. oxysporum, has resulted in a much too broad
definition of this species. Based on this, the current morphologi-
cal characters used to define F. oxysporum include aseptate
microconidia forming false heads on short monophialides,
commonly 3-septate macroconidia formed on monophialides
or branched conidiophores in sporodochia, and chlamydo-
spores that are either formed abundantly and quickly or slowly
with some strains not forming them at all (Leslie & Summerell
2006, Fourie et al. 2011). In this study, all isolates were found
to produce not only aseptate microconidia, but abundant 1-sep-
tate microconidia, all of which were carried on false heads.
Several species were also found to form polyphialides (e.g.,
F. carminascens, F. curvatum, F. elaeidis and F. libertatis),
a characteristic not associated with F. oxysporum morpho-
logy (Gerlach & Nirenberg 1982, Nelson et al. 1983, Leslie
& Summerell 2006). Additionally, the majority of the species
introduced here produced 4- to 5-septate macroconidia in the
same abundance as the 3-septate macroconidia. Gerlach &
Nirenberg (1982) also indicated the presence of 7-septate
macroconidia, but these were not observed in this study given
the media and growth conditions we employed. The ex-epitype
strain of F. oxysporum designated here, agrees well with the
morphological characteristics described by Wollenweber &
Reinking (1935), Booth (1971), Gerlach & Nirenberg (1982) and
Nelson et al. (1983). This strain produced abundant aseptate
and 1-septate microconidia on monophialides only, abundant
3-septate macroconidia with much fewer 1-, 2-, 4- and 5-septate
macroconidia on its sporodochia, and smooth-walled globose
chlamydospores carried intercalarily and/ or terminally. Although
this strain was isolated from a potato tuber displaying symptoms
of dry rot, the ability of this strain to induce these symptoms
requires further investigation. Comparisons of the 15 novel
Fusarium taxa introduced here, revealed subtle morphological
distinctions between the species.
Fusarium carminascens, F. curvatum, F. elaeidis and F. libertatis
readily formed polyphialides on the aerial mycelium, a feature
not known for F. oxysporum (Wollenweber & Reinking 1935,
Booth 1971, Gerlach & Nirenberg 1982, Nelson et al. 1983,
Leslie & Summerell 2006). These four species are further
distinguished from each other by the degree of septation and
curvature of their macroconidia. Both F. carminascens and
F. liber tatis readily formed chlamydospores in culture, whereas
no chlamydospores were observed for F. curvatum and F. elaei-
dis. Furthermore, all strains of F. carminascens produced an
almost carmine red exudate on the aerial mycelium on PDA, not
observed for any other strains studied here. The strong curva-
ture of the macroconidia of F. curvatum is also a unique feature.
The remaining 11 novel species introduced here can be dis-
tinguished based on the degree of septation and dimensions
of the macroconidia and the formation of chlamydospores in
culture. Of these, F. contaminatum, F. gossypinum, F. hoodiae,
F. languescens, F. pharetrum, F. triseptatum and F. veterinarium
displayed some morphological overlap with the ex-epitype strain
of F. oxysporum. However, F. contaminatum, F. gossypinum,
F. pharetrum and F. veterinarium did not form chlamydospores
in culture. These four species are easily distinguished based
on macroconidial dimensions with F. contaminatum and F. vete-
rinarium producing the smallest macroconidia. Fusarium hoo-
diae, F. languescens and F. triseptatum readily formed chla-
mydospores in culture and can be distinguished from each
other and F. oxysporum based on their sporodochia. All
strains of F. triseptatum failed to produce any sporodochia on
the carnation leaf pieces, whereas F. hoodiae formed distinct
pale vinaceous to pale orange sporodochia compared to the
only pale orange sporodochia of F. languescens. Fusarium
callistephi, F. fabacearum, F. glycines and F. nirenbergiae are
easily distinguished from each other and F. oxysporum by the
degree of macroconidial septation and dimensions. However,
these subtle morphological differences need to be supported
by phylogenetic inference to accurately discriminate between
these novel species introduced in the FOSC in this study.
Individual analyses of the partial sequences of the four gene re-
gions (cmdA, rpb2, tef1 and tub2) included in this study (results
not shown) revealed that the tef1 gene region provided the best
resolution to discriminate the novel species introduced here.
The rpb2 gene region also provided good resolution, but with
lower statistical support, whereas the cmdA and tub2 provided
little to no support. However, the addition of the latter two gene
regions to either or both the rpb2 and tef1 greatly increased the
statistical support of each Clade (I–VIII) and their underlining
subclades. Genealogical concordance phylogenetic species
recognition analyses also indicated that there was no evidence
of recombination detected between any of the Clades and
subclades resolved in this study. Analysis of the IGS gene re-
gion (results not shown) provided contradictory tree topologies
and support values, with several strains in Clades III, VII and
VIII forming single lineages. Although O’Donnell et al. (2015)
advocates the use of rpb1, rpb2 and tef1 for sequence-based
identification of Fusarium species, attempts to generate rpb1
sequence data in this study failed for the majority of strains
included in this study.

37
L. Lombard et al.: Epitypification of Fusarium oxysporum
Previous studies of FOSC revealed a high phylogenetic di-
versity within this complex, resolving three (O’Donnell et al.
1998, Brankovics et al. 2017), four (O’Donnell et al. 2004) and
five (Laurence et al. 2012) phylogenetic clades, respectively.
Comparisons of all these clades with those resolved in this
study, revealed that Clade I in this study correlates well with
Clade 1 resolved by O’Donnell et al. (1998, 2004), Laurence
et al. (2012) and Brankovics et al. (2017). Similarly, Clade VIII
in this study matched with Clade 3 of each of these studies.
Clade III correlated with Clade 2 resolved by O’Donnell et al.
(2004) and Brankovics et al. (2017), and Clade V correlated
with clades 4 and 5 of Laurence et al. (2012), and Clade 4 of
O’Donnell et al. (2004). Clades II, IV, VI and VII resolved in
this study did not match any of the clades resolved in these
previous studies.
Comparisons of the origin of the strains studied here revealed
some correlation within most of the Clades (and subclades). All
veterinarian strains included in this study clustered together with
some strains originating from equipment used in food process-
ing in a highly-supported subclade representing F. veterinarium.
Similarly, three strains collected from contaminated dairy pro-
ducts and fruit juice clustered together in the highly-supported
(sub)clade representing F. contaminatum. The majority of the
isolates collected from tomato (Solanum lycopersicum) also
cluster together in a clade representing F. languescens, with a
few clustering in the F. nirenbergiae (sub)clade. In contrast to
these few highlighted examples, all medically related strains
clustered in various well- to highly supported clades, represent-
ing F. cugenangense, F. nirenbergiae, F. triseptatum and the
untreated Fusarium clade. The highest host/substrate diversity
was found in the F. nirenbergiae (sub)clade which included
several special forms in addition to the medically related strains.
The application of the special form and pathotype classification
system can only be successfully applied if the species bounda-
ries are well established (Woudenberg et al. 2015), which is
clearly not the case within the FOSC. For the FOSC, special
forms are defined by the accessory chromosome obtained
via horizontal gene transfer, and the pathotype on the type of
virulence genes carried by this chromosome and should not be
confused with the species boundaries within the FOSC. There-
fore, epitypification of F. oxysporum in this study has resulted
in the recognition of 21 phylogenetic species of which 15 are
provided with names here. Although this study includes only
a small subset of strains belonging to the FOSC, the inclusion
of more isolates will provide a much better perspective on the
cryptic diversity within this important species complex, allowing
additional species to be recognised. Furthermore, it is hoped
that with the epitypification of F. oxysporum, the confusing and
sometimes complicated subspecific classification systems that
have been applied to the FOSC in the past will become obsolete
and be replaced by a more stable and convenient species-level
classification system. We believe that such a system will allow
for better communication between Fusarium researchers in the
medical, environmental and phytopathological fields.
Acknowledgements The authors thank the technical staff, A. van Iperen,
D. Vos-Kleyn and Y. Vlug for their valuable assistance with cultures.
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