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Fungal Taxonomy and Nomenclature

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Chapter 2: Fungal Taxonomy and Nomenclature
Andrew M. Borman
Public Health England National UK Mycology Reference Laboratory, Myrtle Road, Bristol BS2
8EL, UK

Running Title: Taxonomy and Nomenclature

*Corresponding Author Address:

UK National Mycology Reference Laboratory
Public Health England South-West Regional Laboratory
Myrtle Road,
Kingsdown,
Bristol BS2 8EL
Tel: 0117 342 5030
Email: Andy.Borman@uhBristol.nhs.uk

Introduction to Fungal Taxonomy and Nomenclature

Fungi can be uni- or multicellular. Unicellular organisms which divide by budding have
traditionally been referred to as yeasts, whereas multicellular fungi (moulds) exist as
filament-like hyphae which grow via apical extension to form a mycelial mat. However,
moulds and yeasts are not taxonomically valid divisions, and many medically important
endemic fungi exhibit thermal or nutritional dimorphism allowing them to interconvert
between morphologically distinct yeast (or spherule) and mould forms. The fungal kingdom
is vast and ancient. Of the estimated 1-10 million species (Cannon, 1997; Blackwell, 2011;
Hawksworth 2001), only approximately 1% have been formally described and named. Of the
small proportion of fungi that have been named, less than 1000 are commonly reported as
the agents of animal and human infections, with only a small proportion of these being
capable of eliciting disease in immunocompetent hosts (Guarro et al., 1999).

Classical, Phenotype-based Taxonomy

Fungal reproduction can be asexual (mitosis), resulting in the production of usually large
numbers of asexual spores (conidia) that are essentially identical to the parent and permit
spread to and colonisation of new geographic niches, or sexual (resulting from hyphal and
nuclear fusion followed by meiosis) (Alexopoulos et al., 1996). Historically, the identification
and hierarchical classification of filamentous fungi has been based on examination of their
microscopic morphology, and in particular their reproductive structures (Ainsworth et al.,
1973, Sutton, 1973; Kendrick, 1979; Weresub and Hennebert, 1979; Sutton, 1980;
Subramanian, 1983; Sugiyama, 1987; Hennebert, 1991; Reynolds and Taylor, 1993;
Hawksworth, 1994; Guarro et al., 1999). On the basis of the types of sexual structures that
fungi can be induced to form, the kingdom was divided into a limited number of distinct
phyla: the Ascomycota, Zygomycota, and Basidiomycota for fungi with recognised sexual
(teleomorph) forms (Figure 2.1, 2.2). The sexual spores of Ascomycota (ascospores) are
typically produced in sacs (asci) contained in large, thick-walled structures called ascocarps
(although some species produce single, solitary ascospores). Basidiomycota produce sexual
basidiospores borne on basidia which are often contained within macroscopic fruiting
bodies (basidioma) typical of mushrooms, toadstools, smuts and bracket fungi. Conversely,
fungi classified in the phylum Zygomycota produce single thick-walled sexual zygospores
which are often highly ornamented. However, with a few notable exceptions, most fungi are
heterothallic and will only undergo sexual reproduction when two independent, compatible
isolates are present (Figure 2.2), and often in unique environmental conditions. Thus a
fourth form-division, the Deuteromycota (or Fungi Imperfecti) was created to encompass all
those fungi for which a sexual form (teleomorph) had not been described or could not be
induced (Figure 2.2) (reviewed in Talbot, 1971; Kendrick 1981; Seifert 1993). A similar
classification into Ascomycota, Zygomycota, and Basidiomycota could be achieved on the
basis of examination of the anamorph (asexual) forms of the fungi, including the presence of
specific hyphal structures (septa, clamp connections) and the types of asexual spores
formed and their method of production (conidiogenesis) in culture in the laboratory (Figure
2.1) (Hughes 1953; . The hyphae of Zygomycota are hyaline (colourless) broad, ribbon-like
and possess few to no septa (pauci-septate), and asexual conidia (termed sporangiospores)
are typically produced inside large sac-like structures (sporangia). Conversely, the hyphae of
filamentous Ascomycota (many medically important Ascomycota are yeasts) are narrow,
highly septate and may be either hyaline or dematiaceous (brown). Ascomycota typically
produce large numbers of conidia in culture on appropriate media, although the precise
mechanism of conidiogenesis varies dramatically. Finally, the filamentous Basidiomycota,
whilst producing hyphae similar to those of the Ascomycota, rarely produce conidia in
culture, and most often present as rapidly-growing floccose (fluffy) white colonies.
However, upon careful examination, the hyphae can be distinguished from those of the
Ascomycota by the presence of clamp connections (Alexopolous et al., 1996; Figure 2.1),
and occasionally delicate lateral projections termed spicules. It should however be noted
that most medically important Basidiomycota have anamorph states that are yeasts rather
than hyphal.

Although there is a staggering diversity in conidial forms and mechanisms of production,

especially amongst the Ascomycota, different fungi have undergone either convergent or
divergent evolution, with the result that many genetically unrelated species have very
similar conidial characteristics, and other genetically similar species have vastly different
morphological appearances. To further complicate matters, since fungi are of interest to a
wide range of scientists (medical mycologists, geneticists, phytopathologists, the
biotechnology industry etc.) and the sexual and asexual forms of fungi often develop
independently of each other with little obvious morphological relatedness, a single genetic
entity may have been described and named independently several times (Weresub and
Pirozynski, 1979).

Molecular approaches to identification and taxonomy

Since it is well established that different fungal species have dramatically varied antifungal
susceptibility profiles (see for example Pfaller et al., 1998), the accurate identification of the
agents of human infections is essential for appropriate patient management. However,
many fungal isolates from deep and chronic infections do not produce conidia in the
laboratory, rendering their conventional identification impossible. The advent of rapid and
cost effective approaches for extracting and analysing fungal DNA have vastly facilitated
accurate identification of clinically important isolates, and have also permitted a greater
taxonomic understanding of the fungal kingdom. Molecular analyses involving PCR
amplification and sequencing of conserved regions of the fungal genome encoding the rRNA
genes have permitted pan-fungal approaches to fungal identification (reviewed in Borman
et al., 2008), and indeed the Internal Transcribed Spacer region 1 (ITS1) has been proposed
and accepted as an excellent candidate for fungal DNA barcoding (Balajee et al., 2009;
Schoch et al., 2012). More sophisticated approaches examining additional protein-coding
gene loci have permitted the development of multi-locus sequencing approaches (MLST)
and fungal strain typing (reviewed in Peterson 2012). However, a more immediate impact of
such approaches has been the radical and continuous revision of the fungal tree of life
based on a phylogenetic approach to species recognition (PSR) (see Hawksworth 2006 for
review) where DNA sequence similarity is used to define species boundaries.

On the basis of such approaches, the fungal kingdom is now known to comprise at least
seven phyla, (Glomeromycota, Blastocladiomycota, Chytridiomycota,
Neocallimastigomycota, and Microsporidia) with the retention of Ascomycota and
Basidiomycota which now constitute the sub-kingdom Dikarya (Figure 2.3; Hibbett et al.,
2007). Organisms previously ascribed to the Deuteromycota can now be sequenced and
their correct positions in the fungal kingdom ascertained. More dramatically, the phylum
Zygomycota has been disbanded, after molecular approaches demonstrated unequivocally
that it was polyphyletic. The fungi previously classified in Zygomycota are now spread
between the phylum Glomeromycota, and four sub-phyla incertae sedis (of uncertain
position): the Mucoromycotina, Entomophthoromycotina, Kickxellomycotina and
Zoopagomycotina, with most of the medically important members contained in the order
Mucorales within Mucoromycotina (Figure 2.3).

The Ascomycota are now divided into 3 sub-phyla with at least 14 Classes, and 60 orders
(Figure 2.4). The sub-phylum Taphrinomycotina to date contains only one medically
important fungal genus, Pneumocystis, which is formally located within the fungal kingdom.
The sub-phylum Saccharomycotina contains a single medically-important order, the
Saccharomycetales, which encompasses most pathogenic ascomycetous yeasts. The
remainder of the medically important ascomycete genera fall into the sub-phylum
Pezizomycotina, and are divided between at least 14 orders, including the Capnodiales
(Cladosporium and related genera); the Pleosporales (Alternaria, Bipolaris Curvularia,
Exserohilum, Ulocladium and many of the agents of dark-grain eumycetoma); the
Chaetothyriales (Cladophialophora, Exophiala, Fonsecaea, Phialophora, Ramichloridium and
Rhinocladiella); the Eurotiales (Aspergillus, Penicillium, Paecilomyces, Rasamsonia,
Talaromyces, Thermoascus); the Onygenales (the dermatophytes [Trichophyton,
Microsporum, Epidermophyton and fungi with Arthroderma teleomorphs], the thermally
dimorphic fungi with Ajellomyces teleomorphs [Blastomyces, Coccidioides, Emmonsia,
Histoplasma, Paracoccidioides], Chrysosporium, Lacazia, Myceliophthora and Nanniziopsis);
the Hypocreales (Acremonium and allied genera, Fusarium and allied genera,
Purpureocillium and Stachybotrys); the Microascales (Lomentospora, Scedosporium and
Scopulariopsis); the Sordariales (Chaetomium, Madurella, Phialemonium); the Dothideales
(Aureobasidium); Patellariales (Rhytidhysteron); the Choniochaetales (Lecythophora); the
Diaporthales (Phaeoacremonium); the Ophiostomatales (Sporothrix); and the
Calosphaeriales (Pleurostomophora). Even with this greatly revised taxonomy, some
medically important ascomycete genera (Neoscytalidium, Geomyces, Pseudogymnoascus)
remain incertae sedis, pending molecular analyses of more of the fungal kingdom (Figure
2.4).
Finally, the phylum Basidiomycota contains 3 sub-phyla (Pucciniomycotina,
Ustilaginomycotina and Agaricomycotina) and at least 46 orders (Figure 2.5). To date only
around a dozen basidiomycete genera have been formally associated with human infections
(Figure 2.5), and these are restricted to 5 orders. The Sporidiales (Pucciniomycotina)
contains the basidiomycete yeast genera Rhodotorula and Sporobolomyces, Trichosporon
and Cryptococcus are classified in the Tremellales (Agaricomycotina), Bjerkandera, Coprinus,
Irpex, Hormographiella, Perenniporia, Schizophyllum and Sporotrichum reside within the
Agaricales (Agaricomycotina), and Tilletiopsis is classified within the Georgefischeriales
(Ustilaginomycotina). A further genus of medical interest, Malassezia, is also a member of
the sub-phylum Ustilaginomycotina, but its exact position remains to be determined (Figure
2.5).

A further repercussion of widespread employment of molecular approaches to fungal

identification and classification is the discovery of cryptic species (species which can only be
distinguished by DNA sequencing) within many medically important morpho-species
(reviewed in Hawksworth 2006). It is now accepted that Aspergillus fumigatus is a species
complex that encompasses in excess of 40 cryptic species (reviewed in Johnson and Borman,
2009), and a similar number of cryptic species have been discovered in the Fusarium solani
species complex (see for example Short et al., 2013). In some cases, certain cryptic species
have been reported to have altered antifungal susceptibility profiles (Balajee et al., 2005;
Borman et al., 2008), whereas in others discrimination to species level within a species
complex appears to be less medically important (Borman et al., 2013). Nevertheless,
regardless of the medical relevance, the demonstration of cryptic species in most medically
important genera has further added to the complexity of fungal taxonomy and
nomenclature (see . ‘Impact of molecular approaches on fungal nomenclature’)

Impact of molecular approaches on fungal nomenclature

Fungal nomenclature is governed by strict rules established by the International Code of
Nomenclature (ICN) for algae, plants and Fungi. Under the ICN dictates, a name for a fungal
species is only valid if it is in Latin binomial form, has been submitted to MycoBank, and has
a living culture (usually of the type strain) permanently preserved in an accepted Culture
Collection. Fungal name changes have always been permitted, and historically concerned
the most commonly encountered fungal species which had been described and named
independently many times by different authorities, only for those isolates later to be
demonstrated to be identical and thus the names synonymous (Figure 2.6). Under such
circumstances, the earliest valid name was usually retained. Molecular approaches to fungal
identification have to some extent facilitated this process, as fungi which are genetically
indistinguishable should in theory be synonymous. However, there are caveats to this
approach, especially when genomic regions that are fairly conserved are employed to
establish phylogenetic relationships with fungi that have diverged more recently. For
example, initial approaches to examine dermatophyte phylogeny using conserved loci were
unable to distinguish Trichophyton tonsurans, an anthropophilic agent of tinea capitis in
humans, and T. equinum, a zoophilic cause of horse ringworm (reviewed in Borman and
Summerbell, 2015). Further studies using additional loci did however demonstrate that the
two are distinct genetically as well as morphologically and clinically. Similar discussions
continue regarding T. rubrum, the principal cause of tinea pedis and tinea unguinum in
developed countries, and T. soudanense, an agent almost exclusively of tinea capitis that
originated on the African sub-continent (Borman dermatophytes). Thus, fungal taxonomy
and nomenclature remains something of a battleground between the “lumpers” (who
propose to synonymise fungal species with quite distinct geographical niches, different
morphological appearances and even spectra of clinical presentations) and the “splitters”
(who use very precise genetic loci to demonstrate innumerable cryptic species within
morpho-species that are morphologically and clinically indistinguishable).

A second acceptable reason for fungal name changes is when studies demonstrate that a
species should not remain in its current genus. Effectively, when a new fungal genus is
erected, it is done so around a type species, with additional different species being
accommodated in the genus over time. Molecular approaches have demonstrated that in
many cases, these additional species are quite unrelated to the type species, and have
clearly undergone considerable convergent phenotypic evolution. For example, Absidia
corymbifera, which had been retained as a valid name for nearly a century was shown to be
genetically distinct from Absidia repens, the type of the genus, and was briefly renamed
Mycocladus corymbifer, before attaining its final genetically compatible nomenclatural
resting place as Lichtheimia corymbifera, a name by which it had been briefly known from
1903 (Figure 2.6). In such cases, wherever possible, the species epithet is retained (after
adjusting according to the rules of Latin grammar), in an attempt to minimise nomenclatural
confusion. Other examples include Scytalidium dimidiatum (now Neoscytalidium
dimidiatum) which is genetically distant from the type species S. lignicola, and Paecilomyces
lilacinus (now Purpureocillium lilacinum) which is not even in the same fungal class as P.
variotii, the type species of the genus (Figure 2.4).

For other common fungi, a more cautious approach has been proposed. Since the type
species of Aspergillus is A. glaucus, on the basis of phylogenetic approaches most other
Aspergillus species should therefore be removed from the genus, and renamed with their
teleomorph names, which according to convention should take precedence over anamorph
names. This would result in at least 9 new teleomorph genera to encompass the other
former members of Aspergillus (reviewed in Samson et al., 2014). Similarly, the type species
of Fusarium is F. sambucinum, which has a Gibberella teleomorph. Thus, all those current
Fusarium species which have teleomorphs other than Gibberella should be removed,
including Fusarium solani (teleomorph Haematonectria haematococca) and Fusarium
dimerum. However, several large working groups, containing many of the scientists who
originally demonstrated the polyphyletic natures of these two genera have proposed that
the status quo be maintained in the face of phylogenetic evidence, to “preserve established
research connections” in the diverse communities interested in Fusarium (Geiser et al.,
2013) and “maintain the prevailing, broad concept of Aspergillus” (Samson et al., 2014).
Although at odds with much molecular phylogenetic evidence, this nomenclatural
obfuscation, which is supported by the International Commission of Penicillium and
Aspergillus (ICPA), will certainly reduce confusion in medical mycology, where accepted
nomenclatural changes have to be filtered down to clinicians. A non-exhaustive list of
accepted and likely future nomenclatural changes to common medically important fungi can
be found in Table 2.1, although this is likely subject to considerable change.

One fungus = One name

The existence of both anamorph and teleomorph states that share little morphological Comment [AM1]: This can just be
deleted
relatedness for many medically important fungi, and which have often been described
separately many years apart has resulted in many fungal species having several names. In
2011, the “Amsterdam Declaration” (Hawksworth et al., 2011) proposed a substantial
modification of Article 59 of the code of nomenclature that would enforce the
abandonment of dual nomenclature in fungi. From Jan 2013, each fungus must retain a
single name, with the decisions on which names should be retained being taken or at least
ratified by the Nomenclature Committee for Fungi (Hawksworth 2011). The pre-existing
convention, by which the teleomorph name should always take precedence, could certainly
lead to considerable confusion in the medical mycology community, which has widely
ignored the convention previously as it was usually the anamorph state that was isolated in
the laboratory. To somewhat soften the blow to medical mycologists, additional changes to
the code included “Follow the principal of priority of publication when selecting the generic
name to adopt” which seemingly ends the previous precedence of teleomorph over
anamorph, with the caveat “except where the younger generic name is far better known”.
To further lesson unnecessary, and transient nomenclatural instability, working groups and
committees established under the auspices of the International Commission on the
Taxonomy of Fungi (ICTF) and the Nomenclature Committee for Fungi (NCF) will propose
lists of retained (protected) and rejected names for key genera, with only definitive changes
being ratified. Whilst this might seem like a satisfactory fix, it still has its adversaries (Gams
et al., 2011), who argue that if this applies at both the genus and species level, then
numerous new combinations or specific conservation proposals would be required, and that
many important teleomorph genera would also be abolished. There is also the question of
who should and how to decide whether an anamorph or teleomorph name is more popular.
Suggestions that have been explored to date include the use of internet search engines to
examine the number of “hits” against a given fungal name (ICPA; Geiser et al., 2013).
Whatever the final decisions, it is inevitable that fungal nomenclature is heading for stormy
waters, which hopefully in the end will lead to a more stable and scientifically robust
taxonomic understanding of the fungal kingdom.

Disclaimers
To the best of the author’s knowledge, the taxonomic and nomenclatural affiliations
discussed in this chapter were accurate at the time of writing. Any political views expressed
in gest are the author’s own and do not reflect the position of Public Health England.

References
AINSWORTH, G.C., SPARROW, F.K. & SUSSMAN, A.S. (Eds.) 1973. The Fungi: An Advanced
Treatise, Volume IVA – A Taxonomic Review with Keys: Ascomycetes and Fungi
Imperfecti. Academic Press, London: 1-7.
ALEXOPOULOS, C.J., MIMS, C.W. AND BLACKWELL, M. 1996. Introductory Mycology, IV
edition.John Wiley & Sons, New York: 1-868.
BALAJEE, S.A., GRIBSKOV, J.L., HANLEY, E., NICKLE, D., & MARR, K.A. 2005. Aspergillus
lentulus sp. nov., a new sibling species of A. fumigatus. Eukaryot. Cell.4, 625-632.
BALAJEE, S.A., BORMAN, A.M., BRANDT, M.E., CANO, J., CUENCA-ESTRELLA, M., DANNAOUI,
E., GUARRO, J., HAASE,G., KIBBLER, C.C., MEYER, W., O’DONNELL, K., PETTI, C.A.,
RODRIGUEZ-TUDELA, J-L., SUTTON, D., VELEGRAKI, A., & WICKES, B.L. 2009.
Sequence-based identification of Aspergillus, Fusarium and the Mucorales in the
 clinical mycology laboratory: where are we and where should we go from here? J. Clin.
Microbiol. 47, 877-884.
BORMAN, A.M., LINTON, C.J., MILES, S-J., & JOHNSON, E.M. 2008 Molecular identification
of pathogenic fungi. J. Antimicrob. Chemother. 61, i7-i12.
BORMAN, A.M., PETCH, R., LINTON, C.J., PALMER, M.D., BRIDGE, P.D., & JOHNSON, E.M.
2008. Candida nivariensis, an emerging pathogenic fungus with multidrug resistance
to antifungal agents. J. Clin. Microbiol. 46, 933-938.
BORMAN, A.M., SZEKELY, A., LINTON, C.J., PALMER, M.D., BROWN, P., & JOHNSON, E.M.
2013. Epidemiology, antifungal susceptibility, and pathogenicity of Candida africana
isolates from the United Kingdom. J. Clin. Microbiol. 51, 967-972.
BORMAN, A.M. & SUMMERBELL, R.C. 2015. Trichophyton, Microsporum, Epidermophyton,
and Agents of Superficial Mycoses, p 2128-2152. In Jorgensen J, Pfaller M, Carroll K,
Funke G, Landry M, Richter S, Warnock D (eds), Manual of Clinical Microbiology, 11th
Edition. ASM Press, Washington, DC.
CANNON, P.F. 1997. Strategies for rapid assessment of fungal diversity. Biodiversity and
Conservation 6, 669-680.
GAMS, W., & JAKLITSCH, W. (& 77 signatories). 2011. A critical response to the “Amsterdam
Declaration”. Mycotaxon 116, 1-12.
GEISER, D.M., AOKI, T., BACON, C.W., BAKER, S.E., BHATTACHARYYA, M.K., BRANDT, M.E. et
al., 2013. One fungus, one name: defining the genus Fusarium in a scientifically robust
way that preserves longstanding use. Phytopathology. 103, 400-408.
GUARRO, J., GENE, J. & STCHIGEL, A.M. 1999. Developments in fungal taxonomy. Clin.
Microbiol. Rev. 12, 454-500.
HAWKSWORTH, D.L. (ed.) 1994. Ascomycete systematics: problems and perspectives in the
nineties. Plenum Press, New York.
HAWKSWORTH D.L. 2001. The magnitude of fungal diversity: The 1.5 million species
estimate revisited. Mycological Res. 105, 1422–1432.
HAWKSWORTH, D.L. 2006. Pandora's mycological box: molecular sequences vs. morphology
in understanding fungal relationships and biodiversity. Rev. Iberoam. Micol. 23, 127-
133.
HAWKSWORTH, D.L., CROUS, P.W., REDHEAD, S.A., REYNOLDS, D.R., SAMSON, R.A., SEIFERT
K.A., TAYLOR J.W., & WINGFIELD M.J. [& 69 signatories]. 2011. The Amsterdam
Declaration on Fungal Nomenclature. IMA Fungus 2, 105–112; Mycotaxon 116, 91–
500
HAWKSWORTH D.L. 2011. A new dawn for the naming of fungi: impacts of decisions made in
Melbourne in July 2011 on the future publication and regulation of fungal names.
MycoKeys 1, 7–20; IMA Fungus 2,155–162.
HENNEBERT, G.L. 1991. Art. 59 and the problem with pleoanamorphic fungi. Mycotaxon 40,
479-496.
HIBBETT D.M., BINDER M., BISCHOFF J. F., BLACKWELL M., CANNON P.F., ERIKSSON O.,
HUHNDORF S. et al. 2007. A higher-level phylogenetic classification of the Fungi.
Mycological Res. 111, 509–547.
HOFFMANN K., JPAWŁOWSKA J., WALTHER G., WRZOSEK M., DE HOOG G.S., BENNY G.L.,
KIRK P.M., & VOIGT K. 2013. The family structure of the Mucorales: a synoptic revision
based on comprehensive multigene-genealogies. Persoonia. 30, 57-76.
JOHNSON, E.M. & BORMAN, A.M. 2009. Identification of Aspergillus at the species level:
the importance of conventional methods; microscopy and culture. pp 55-74. In
Aspergillus and Aspergillosis. Ed. Alessandro Pasqualotto. Springer Press.
KENDRICK, B. (ed.) 1979. The Whole Fungus: The Sexual-Asexual Synthesis. Volume 1-2.
National Museums of Canada, Ottawa.
KENDRICK, B. 1981. The history of conidial fungi, Pages 3–18 in GT Cole and B Kendrick, eds.
Biology of Conidial Fungi. New York, Academic Press.
BLACKWELL, M. 2011. The Fungi: 1,2,3…5.1 million species?. Am. J. Botany 98; 426-438.
PETERSON, S.W. 2012. Aspergillus and Penicillium identification using DNA sequences:
barcode or MLST? Appl. Microbiol. Biotechnol. 95, 339-344.
PFALLER M.A., JONES R.N., MESSER S.A., EDMOND M.B., WENZEL R.P. 1998. National
surveillance of nosocomial blood stream infection due to species of Candida other
than Candida albicans: frequency of occurrence and antifungal susceptibility in the
SCOPE Program. Diagn. Microbiol. Infect. Dis. 31: 327-332.
REYNOLDS, D.R. & TAYLOR, J.W. (eds.) 1993. The fungal holomorph: Mitotic, meiotic and
pleomorphic speciation in fungal systematics. CAB International, Oxon.
SAMSON, R.A., VISAGIE, C.M., HOUBRAKEN, J., HONG, S.B., HUBKA, V., KLAASSEN, C.H.,
PERRONE, G., SEIFERT, K.A., SUSCA, A., TANNEY, J.B., VARGA, J., KOCSUBÉ, S., SZIGETI,
G., YAGUCHI,T., & FRISVAD, J.C. 2014. Phylogeny, identification and nomenclature of
the genus Aspergillus. Stud. Mycol. 78, 141-173.
SCHOCH, C.L., SEIFERT, K.A., HUHNDORF, S., ROBERT, V., SPOUGE, J.L., LEVESQUE, C.A.,
CHEN, W.; Fungal Barcoding Consortium; Fungal Barcoding Consortium Author List.
2012. Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA
barcode marker for Fungi. Proc. Natl. Acad. Sci. U S A. 109, 6241-6246.
SEIFERT, K.A. 1993. Integrating anamorphic fungi into the fungal system, Pages 79–85 in DR
Reynolds, and JW Taylor, eds. The Fungal Holomorph: mitotic, meiotic and
pleomorphic speciation in fungal systematics. Wallingford, UK, CAB International.
Short, D.P., O'Donnell, K., Thrane, U., Nielsen, K.F., Zhang, N., Juba, J.H., & Geiser, D.M.
2013. Phylogenetic relationships among members of the Fusarium solani species
complex in human infections and the descriptions of F. keratoplasticum sp. nov. and F.
petroliphilum stat. nov. Fungal. Genet. Biol. 53, 59-70.
SUBRAMANIAN, C.V. 1983. Hyphomycetes: taxonomy and biology. Academic Press, London.
SUGIYAMA, J. (ed.) 1987. Pleomorphic Fungi: The diversity and its taxonomic implications.
Kodansha Ltd., Tokyo: 1-325.
SUTTON, B.C. 1973. Coelomycetes. In: The fungi: An advanced treatise. Vol. IVA. A
taxonomic review with keys: ascomycetes and fungi imperfecti (eds. G.C. Ainsworth,
F.K. Sparrow and A.S. Sussman). Academic Press, New York: 513-582.
SUTTON, B.C. 1980. Coelomycetes - Fungi Imperfecti with pycnidia, acervuli and stromata.
Commonwealth Mycological Institute, Kew, Surrey, England.
TALBOT, P.H.B. 1971. Principles of Fungal Taxonomy. The Macmillan Press, London: 1- 274.
WERESUB, L.K. & HENNEBERT, G.L. 1979. Anamorph and teleomorph: terms for organs of
reproduction rather than karyological phases. Mycotaxon 8, 181-186.
WERESUB, L.K. & PIROZYNSKI, K.A. 1979. Pleomorphism of fungi as treated in the history of Comment [SCW2]: Please can I
mycology and nomenclature. In: Kendrick, B (ed) The Whole Fungus: The Sexual- confirm own copyright for the images and
that they haven’t been previously
Asexual Synthesis. Vol 1, 17-30. National Museums of Canada, Ottawa. published.
Comment [AM3]: Yes, I do “own”
Figure Legends copyright, and they have Not been
previously published.
 Comment [SCW4]:
Ideally all figures should be 10-x15cm at
Figure 2.1: Approaches to fungal identification and taxonomy. Common features of the 300dpi. The supplied images are all less
sexual cycles (left-hand side) and asexual cycles (right-hand side) of Basidiomycota, than 2x2cm at the correct dpi.

Ascomycota and “Zygomycota”. We will require high resolution versions of

the 14 part figures to include the figure in
the book.
Figure 2.2: Traditional separation of the fungal Kingdom into three phyla (Ascomycota,
Comment [AM5]: I am sending
Basidiomycota, Zygomycota) and the form division Deuteromycota for those fungi without a individual images of the 14 parts. They are
known sexual stage. the best resolution I can provide.
Unfortunately, due to the file sizes, I will be
sending them individually or in small
Figure 2.3: Recent taxonomy of the fungal Kingdom, based on Hibbett et al., 2007. batches.

Organisms previously accommodated in the phylum “Zygomycota” are highlighted in the Comment [SCW6]: As each figure is
called out separately they should be
blue shaded lozenge. The positions of the more common, medically important mould labelled individually. I have split the figures
genera are shown. up and deleted the original title ‘Historical
and current taxonomy of the Fungal
Kingdom’.
Figure 2.4: Revised taxonomy of the Ascomycota. The positions of medically relevant genera Comment [SCW7]: Are Figure 2.4 and
is shown. 2.5 to be added at the call-out or is this
simply to cross reference the figures?

Figure 2.5: Revised taxonomy of the Basidiomycota. The position of medically relevant What is the point of the shaded box in the
figure (not clear/no key)?
genera is shown.
Comment [AM8]: They are simply to
cross reference and should not be called-
Figure 2.6: The taxonomic (A) and Nomenclatural (B) merry-go-round. (A) The principal out all at once.
taxonomic revisions of Lichtheimia corymbifera; dashed arrows indicate temporary changes, The point of the shaded lozenge is already
the solid arrow indicates the final (hopefully) taxonomic position (B) Some of the various explained in the legend.
nomenclatural synonyms of L. corymbifera over the past century (adapted from Index Comment [SCW9]: What do you mean
Fungorum). by based on? Have you adapted an existing
figure from the cited article? Please can
you send us the original publication to
Table 2.1: Accepted and likely future nomenclatural changes to human pathogenic fungi. check if permission will be needed.

The original name by which the species was described, intermediary (transitional) names Comment [AM10]: The figure is not an
adaptation of an existing one. It is drawn
and the currently accepted name are shown. Note that in many cases, the name commonly from the data presented in the Hibbett
employed by clinicians and mycologists is a transitional name and not the original name. In paper, as are many other similar ones that
have been published since by other
most cases the species epithet has been retained to limit nomenclatural confusion. authors
Comment [AM11]: THAT IS FINE
Comment [SCW12]: We will need to
split the image over several pages (most
likely split in three)
...
Comment [AM13]: A new 3 part,
editable figure is provided.
Comment [SCW14]: What do you
mean by adapted? Have you adapted an
existing figure from the cited source?
Please can you send us a link to the direct...
Comment [AM15]: No, this is an
original figure drawn by myself, from data
that exists on the Index Fungorum website.
Comment [SCW16]: Table to be
supplied.
Comment [AM17]: The table is now
supplied
Formatted: Space After: 10 pt, Line
spacing: Multiple 1.15 li
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