1 s2.0 S0166061619300090 Main

available online at www.studiesinmycology.org STUDIES IN MYCOLOGY 94: 125–298 (2019).
Foliar pathogens of eucalypts*
P.W. Crous1,2*, M.J. Wingfield2,3, R. Cheewangkoon4, A.J. Carnegie5,6, T.I. Burgess2,7, B.A. Summerell8, J. Edwards9,10,
P.W.J. Taylor11, and J.Z. Groenewald1
1
Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD, Utrecht, The Netherlands; 2Department of Genetics, Biochemistry and Microbiology, Forestry and
Agricultural Biotechnology Institute (FABI), University of Pretoria, Pretoria, 0002, South Africa; 3Forestry and Agricultural Biotechnology Institute (FABI), University of
Pretoria, Pretoria, 0002, South Africa; 4Department of Plant Pathology, Faculty of Agriculture, Chiang Mai University, Chiang Mai, 50200, Thailand; 5Forest Health &
Biosecurity, Forest Science, NSW Department of Primary Industries – Forestry, Level 12, 10 Valentine Ave, Parramatta, NSW, 2150, Australia; 6School of
Environment, Science and Engineering, Southern Cross University, Lismore, NSW, 2480, Australia; 7Environmental and Conservation Sciences, Murdoch University,
90 South Street, Murdoch, WA, 6150, Australia; 8Royal Botanic Gardens and Domain Trust, Mrs Macquaries Rd, Sydney, NSW, 2000, Australia; 9Agriculture
Victoria Research, Department of Jobs, Precincts and Regions, AgriBio Centre, 5 Ring Road, LaTrobe University, Bundoora, Victoria, 3083, Australia; 10School of
Applied Systems Biology, LaTrobe University, Bundoora, Victoria, 3083, Australia; 11Faculty of Veterinary and Agricultural Sciences, The University of Melbourne,
Parkville, VIC, Australia
*Correspondence: P.W. Crous, p.crous@wi.knaw.nl
Abstract: Species of eucalypts are commonly cultivated for solid wood and pulp products. The expansion of commercially managed eucalypt plantations has chiefly
been driven by their rapid growth and suitability for propagation across a very wide variety of sites and climatic conditions. Infection of foliar fungal pathogens of eucalypts
is resulting in increasingly negative impacts on commercial forest industries globally. To assist in evaluating this threat, the present study provides a global perspective on
foliar pathogens of eucalypts. We treat 110 different genera including species associated with foliar disease symptoms of these hosts. The vast majority of these fungi
have been grown in axenic culture, and subjected to DNA sequence analysis, resolving their phylogeny. During the course of this study several new genera and species
were encountered, and these are described. New genera include: Lembosiniella (L. eucalyptorum on E. dunnii, Australia), Neosonderhenia (N. eucalypti on E. costata,
Australia), Neothyriopsis (N. sphaerospora on E. camaldulensis, South Africa), Neotrichosphaeria (N. eucalypticola on E. deglupta, Australia), Nothotrimmatostroma
(N. bifarium on E. dalrympleana, Australia), Nowamyces (incl. Nowamycetaceae fam. nov., N. globulus on E. globulus, Australia), and Walkaminomyces (W. medusae on
E. alba, Australia). New species include (all from Australia): Disculoides fraxinoides on E. fraxinoides, Elsinoe piperitae on E. piperita, Fusculina regnans on E. regnans,
Marthamyces johnstonii on E. dunnii, Neofusicoccum corticosae on E. corticosa, Neotrimmatostroma dalrympleanae on E. dalrympleana, Nowamyces piperitae on
E. piperita, Phaeothyriolum dunnii on E. dunnii, Pseudophloeospora eucalyptigena on E. obliqua, Pseudophloeospora jollyi on Eucalyptus sp., Quambalaria tasmaniae on
Eucalyptus sp., Q. rugosae on E. rugosa, Sonderhenia radiata on E. radiata, Teratosphaeria pseudonubilosa on E. globulus and Thyrinula dunnii on E. dunnii. A new
name is also proposed for Heteroconium eucalypti as Thyrinula uruguayensis on E. dunnii, Uruguay. Although many of these genera and species are commonly
associated with disease problems, several appear to be opportunists developing on stressed or dying tissues. For the majority of these fungi, pathogenicity remains to be
determined. This represents an important goal for forest pathologists and biologists in the future. Consequently, this study will promote renewed interest in foliar
pathogens of eucalypts, leading to investigations that will provide an improved understanding of the biology of these fungi.
Key words: Corymbia, Eucalyptus, Foliar pathogen, New taxa, Taxonomy.

Taxonomic novelties: new family: Nowamycetaceae Crous; New genera: Lembosiniella Crous, Neosonderhenia Crous, Neothyriopsis Crous, Neotrichosphaeria Crous
Studies in Mycology
& Carnegie, Nothotrimmatostroma Crous, Nowamyces Crous, Walkaminomyces Crous & Carnegie; New species: Disculoides fraxinoides Crous, Elsinoe piperitae
Crous, Fusculina regnans Crous, Lembosiniella eucalyptorum Crous & Carnegie, Marthamyces johnstonii Crous & Carnegie, Neofusicoccum corticosae Crous &
Summerell, Neosonderhenia eucalypti Crous, Neotrimmatostroma dalrympleanae Crous, Nowamyces globulus Crous, Nowamyces piperitae Crous, Phaeothyriolum
dunnii Crous & Carnegie, Pseudophloeospora eucalyptigena Crous, Pseudophloeospora jollyi Crous, Quambalaria tasmaniae Crous, Quambalaria rugosae Crous,
Sonderhenia radiata Crous, Teratosphaeria pseudonubilosa G. Perez & Carnegie, Thyrinula dunnii Crous & Carnegie; New combinations: Allelochaeta brevilata
(H.J. Swart & D.A. Griffiths) Crous, Blastacervulus robbenensis (Crous et al.) Crous, Lembosiniella eucalypti (Sivan. & R.G. Shivas) Crous, Neofusicoccum
versiforme (Z.Q. Yuan et al.) Crous, Neosonderhenia foliorum (Cooke) Crous, Neothyriopsis sphaerospora (Marasas) Crous, Neotrichosphaeria eucalypticola (Sivan.
& R.G. Shivas) Crous & Carnegie, Nothotrimmatostroma bifarium (Gadgil & M.A. Dick) Crous, Nothotrimmatostroma eucalyptorum (Crous & Carnegie) Crous,
Phacidium innumerum (Massee) Crous, Phaeothyriolum amygdalinum (Cooke & Massee) Crous & Carnegie, Phaeothyriolum corymbiae (Crous) Crous,
Phaeothyriolum eucalyptorum (Crous & W.B. Kendr.) Crous, Pseudocercospora tumulosa (Carnegie & Beilharz) Carnegie & Crous, Teratosphaeria delegatensis
(R.F. Park & Keane) Crous, Thyrinula parasitica (Crous) Crous, Walkaminomyces medusae (Carnegie & G.S. Pegg) Crous & Carnegie; New name: Thyrinula
uruguayensis Crous for Heteroconium eucalypti Crous & M.J. Wingf.; Epitypes: Arnaudiella eucalyptorum Crous & W.B. Kendr., Lembosiopsis eucalyptina Petr. &
Syd., Leptostromella eucalypti Cooke & Massee, Microthyrium amygdalinum Cooke & Massee, Mycosphaerella marksii Carnegie & Keane, Sphaerella cryptica
Cooke, Stigmina robbenensis Crous et al., Thyriopsis sphaerospora Marasas, Trichosphaeria eucalypticola Sivan. & R.G. Shivas, Trimmatostroma bifarium Gadgil &
M.A. Dick.
Available online 8 August 2019; https://doi.org/10.1016/j.simyco.2019.08.001.
INTRODUCTION
*
Dedication: This paper is dedicated to Dr H.J. Swart (1922–1993). Harry
Swart was an extraordinary mycologist with an exceptional gift for detailed The tree genera Angophora, Corymbia and Eucalyptus (Myrta-
mycological observations. His series of papers on the “Australian leaf-inhabiting ceae), colloquially termed eucalypts, accommodate more than
fungi” published in the then Transactions of the British Mycological Society, 800 species, the majority endemic to Australia (Pryor & Johnson
inspired the first author to dedicate much of his career to collecting and culturing 1971, Boland et al. 1992, Ladiges et al. 2003, Thornhill et al.
these incredible foliicolous fungi (see Australas. Pl. Pathol. 23: 29–35. 1994).
Peer review under responsibility of Westerdijk Fungal Biodiversity Institute.

© 2019 Westerdijk Fungal Biodiversity Institute. Production and hosting by ELSEVIER B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
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CROUS ET AL.
2019). Many species of Corymbia and Eucalyptus are grown climates, were very seriously damaged by leaf blotch disease
globally for commercial purposes and as amenity trees. To caused by species originally treated in the genus Mycosphaerella
meet the world’s demand for paper and pulp, there has in recent (Browne 1968, Lundquist 1987, Lundquist & Purnell 1987).
years been a rapid expansion in areas under afforestation, Plantations of E. globulus in certain areas in South Africa in the
particularly in South-East Asia and in the Southern Hemisphere 1930’s were abandoned due to severe damage from Ter-
(Eldridge et al. 1994, Cotterill & Brolin 1997, Martin 2003, atosphaeria (= Mycosphaerella) nubilosa (Lundquist 1987). When
Qi 2003). Importantly eucalypts, which make up 74 % of these species began to be commercially planted in Australia from
forested land in Australia (ABARES 2017), are the key compo- the 1990’s they also experienced severe damage from T. nubilosa
nents of most ecosystems and provide food and habitat for a and also T. cryptica (Mohammed et al. 2003, Milgate et al. 2005,
myriad of animals as well as a range of micro-organisms Carnegie 2007a, Barber et al. 2008, Hunter et al. 2009, Andjic
including fungi. et al. 2019). As Eucalyptus spp. such as E. camaldulensis,
The genus Eucalyptus is one of the most remarkable genera E. grandis and E. urophylla, generally suited to warmer summer
of trees. It includes at least 700 species, almost all of which are rainfall areas of the world, were deployed in parts of the tropics
native to Australia (Ladiges et al. 2003, Thornhill et al. 2019). The and Southern Hemisphere, leaf diseases caused by species of
remaining species, including Eucalyptus deglupta and Calonectria (Cylindrocladium) were found to cause serious leaf
E. urophylla, are endemic to Indonesia, New Guinea and the blight problems (Sharma & Mohanan 1982, Sharma et al. 1985,
Philippines. All but one (Corymbia papuana) of the 113 species in Blum et al. 1992, Park et al. 2000, Crous 2002, Old et al.
Corymbia are also native to Australia. These trees occur across 2003). Subsequently, species of Teratosphaeria (Kirramyces)
a vast diversity of habitats including moist forests, open wood- emerged as significant diseases on these species in the tropics
lands, deserts, rocky outcrops and swamplands (Potts & and sub-tropics (Andjic et al. 2019).
Pederick 2000). Likewise, they occur in a wide range of cli- Where eucalypts have been planted for the first time in new
matic conditions with adaptations to high or moderate rainfall, areas, they have tended not to be affected by diseases,
drought, bushfire, high and low temperatures and even snow including those affecting their shoots and leaves. But, as is
during winter. true for diseases of all plants, this situation has changed over
In the last comprehensive review of fungi occurring on these time with new disease (and insect) problems accumulating.
hosts, Sankaran et al. (1995) recorded more than 500 species of These diseases are caused by two distinct categories of
leaf-infecting fungi on eucalypts. There have been many other pathogens. They include native pathogens that have moved
studies on the fungi occurring on eucalypts (Park et al. 2000). onto eucalypts (host jumps) and those that have been acci-
These date back to descriptions of “Sphaerella” nubilosa and “S.” dentally introduced from Australia into the areas where the
cryptica in Cooke’s Handbook of Australian Fungi from samples eucalypts are planted (Burgess & Wingfield 2017). Native
sent to England from Australia (Cooke 1892). Interest in these pathogens of the foliage include for example species of Cal-
fungi has primarily been linked to the fact that Eucalyptus spp. onectria that can occur naturally in soils (Lombard et al.
(and to a lesser degree Corymbia spp.) have been extensively 2010a,b,c, Alfenas et al. 2015). The larger number of non-
planted outside their native range for more than 100 years native pathogens are eucalypt-specific such as species of
(Turnbull 2000). During this time, they have reached a point of Readeriella and Teratosphaeria.
being amongst few tree genera most commonly propagated in Very little is known regarding the majority of fungi occurring on
plantations, with almost 20 M ha now planted globally (http://git- the foliage of eucalypts (Sankaran et al. 1995). A relatively small
forestry-blog.blogspot.com/2008/09/eucalyptus-global-map- number of species are recognised as primary pathogens. These
2008-cultivated.html). These plantations sustain some of the have mostly emerged where species of eucalypts are intensively
world’s most important timber industries, which produce solid managed in plantations (Andjic et al. 2019). Others occurring on
wood and pulp products. Consequently, the health of eucalypt the leaves of trees in native stands are either recognised as pri-
trees is important to the sustainability of major forest industries, mary pathogens either because they belong to groups of fungi
and fungi are amongst the most common causes of diseases of known to cause disease or due to the nature of the symptoms with
these trees. which they are associated. But it is important to realise that even
Fungal pathogens have been recognised as threats to both for those fungi considered to cause leaf diseases, the majority
native-grown and plantation-grown eucalypt trees for many years have not been subjected to pathogenicity tests and thus proven to
(Browne 1968, Dick & Gadgil 1983, Sharma et al. 1985, be pathogens based on Koch’s postulates.
Lundquist 1987, Crous et al. 1989, Ferreira 1989, Carnegie The areas of origin of most leaf pathogens found on species of
et al. 1994, Andjic et al. 2019). This threat has become more eucalypts outside the native range of these trees remains un-
obvious as eucalypt plantation areas have increased, particularly known. For example, the very serious leaf and shoot pathogen,
during the course of the last 30 years. This expansion has been Teratosphaeria destructans, was first discovered on Eucalyptus
driven by their rapid growth and suitability for propagation across planted as non-natives in north Sumatra (Wingfield et al. 1996).
a very wide variety of sites and climatic conditions. The The pathogen has spread to many other areas of South East Asia
expansion of eucalypt plantings can also be attributed to the and most recently to South Africa (Andjic et al. 2011, Greyling et al.
growth of the paper and dissolving pulp industries, where the 2016). Yet its highly host-specific nature and the fact that popu-
short fibres of Eucalyptus spp. are required. lation genetic studies have shown strong clonality in invaded
The negative impact of foliar pathogens of eucalypts dates ranges suggest that it is native to Eucalyptus somewhere in the
back to the early propagation of these trees in plantations outside native range of these trees (Burgess & Wingfield 2017). One
Australia. One of the first species to be extensively planted was suggestion has been that it is native to Timor but this has yet to be
E. globulus, suited particularly to areas of the world with Medi- proven scientifically (Andjic et al. 2019).
terranean climates (Florence 1986). Many plantations of these There are numerous instances of foliar fungi being originally
trees, and the closely related E. nitens, suited to cool temperate described from exotic eucalypts then later found in Australia (e.g.
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EUCALYPT FOLIAR PATHOGENS
Carnegie et al. 1997, Maxwell et al. 2000, 2003, Jackson et al. one species, Q. coyrecup that is an aggressive stem canker
2005, Crous et al. 2007c, Burgess & Wingfield 2017). Many of pathogen of Corymbia (Paap et al. 2008). The two best known
these are likely to be endemic to Australia but found overseas species on leaves and shoots are Q. eucalypti that causes leaf and
first due to intensive interest in exotic eucalypt plantations. There shoot diseases of Eucalyptus spp. and Q. pitereka that is specific to
are, however, still many species of, for example Teratosphaeria, leaves and shoots of Corymbia spp. Both species are now found in
that have not been found in Australia (Carnegie et al. 2007). With plantations outside the native range of Eucalyptus and Corymbia
more intensive fungal forays in Australia, many new species are with Q. eucalypti having the most extensive global distribution.
now being discovered (Crous et al. 2017a, b, 2018a,b,c, present Interestingly, Q. eucalypti was first found and described in South
study). Africa (Wingfield et al. 1993), then found in Australia more than a
Diseases of eucalypts are emerging via host shifts from decade later (Carnegie 2007a, Pegg et al. 2008).
native plants to these trees where they are planted as exotics There are likely numerous records of eucalypt leaf fungi on
(Slippers et al. 2005). There are now many instances where species previously treated as Eucalyptus that are now known to
stem canker pathogens in the Cryphonectriaceae have been actually represent species of Corymbia. Those collected from
shown to have undergone host shifts from native Myrtales hosts only identified to genus (“Eucalyptus sp.”) would thus now
(including Myrtaceae, Melastomataceae, etc) to eucalypt be indistinguishable as being from Corymbia, cf. those collected
(Wingfield 1999, Wingfield 2003). These pathogens, new to from distinct species (e.g. Eucalyptus maculata = Corymbia
eucalypts, can be very damaging and they now threaten the maculata). It is reasonable to presume that the fungi on Cor-
trees in other areas where they are propagated, as well as in the ymbia spp. are distinct from those on Eucalyptus, as is found with
native ranges of these trees. A key driver of native pathogens Q. pitereka and Q. eucalypti. In recent years, Corymbia spp.
moving to non-native eucalypts propagated in plantations ap- have been increasingly deployed outside their native range for
pears to be the presence of native trees in the Myrtales and thus plantation development. This would justify a much greater focus
those relatively closely related to eucalypts facilitating a host shift on making comparisons of pathogens and other fungi that occur
(Burgess & Wingfield 2017, Crous et al. 2016a). on leaves specifically of Corymbia spp. It is relevant to recognise
A classic example of a primary eucalypt leaf and shoot that some records in the present study for “Eucalyptus” are also
pathogen that is not native on these trees but has adapted to likely to include some for Corymbia, because the two tree genera
infect them is the rust fungus Austropuccinia psidii. This path- were collectively treated as Eucalyptus until relatively recently
ogen has a wide host range on many genera of the Myrtaceae in (Hill & Johnson 1995).
South America and was first recorded causing significant dam- Many of the fungi listed in this study have been described
age to Eucalyptus in Brazil (Ferreira 1983), where South African from symptomatic dead leaves found in the litter below trees.
seed sources of E. grandis were shown to be particularly sus- Most are probably saprobes that grow and sporulate once leaves
ceptible (Coutinho et al. 1998, Glen et al. 2007). More recently, begin to die. Here it is important to recognise that like other
A. psidii has appeared in new areas including Hawaii, Japan, plants, eucalypt leaves have a vast microbiome and that many of
China, Australia, South Africa, New Caledonia, Indonesia, the fungi that form part of this community are seen only when
Singapore and most recently in New Zealand (Carnegie & Pegg leaves die. For example, Kemler et al. (2013) used a meta-
2018). The same genotype of the rust occurs in most of these genomics approach to show that living tissues of Eucalyptus
areas and this is known as the pandemic genotype (Stewart et al. trees include a very large number of fungal taxa, the most
2017, Carnegie & Pegg 2018). This is different to the genotype of common of which occur in groups that include important path-
A. psidii that occurs in South Africa, which has not been found on ogens such as those in the Capnodiales (Dothideomycetes).
these trees in plantations but has been shown to infect Euca- The primary aim of this study was to reconsider all the fungi
lyptus and Corymbia in greenhouse pathogenicity studies (Roux that have been recorded on diseased eucalypt foliage. Species
et al. 2013, 2016). simply associated with eucalypts, and assumed to be saprobic or
Many fungi occurring on eucalypt foliage are well-known of minor importance, are excluded. Various new species are
opportunists or latent pathogens. The best-studied are species described and the taxonomy of previously described species and
in the Botryosphaeriaceae and more specifically Neofusicoccum. genera is re-evaluated. Species associated with leaf spots rep-
The Botryosphaeriaceae include 24 genera, the majority of which resenting 110 genera are listed. For the majority of these fungi,
occur on trees or woody shrubs (Yang et al. 2017). Most are nothing is known regarding their biology or ecological impor-
found on the bark of branches and stems where they also cause tance. There are several genera that include well recognised
dieback symptoms. A much smaller number of these fungi are primary pathogens, known from experimental evidence to be
known from leaves including those of Eucalyptus spp. Within able to infect unwounded healthy plant tissue. These include
Australia only 24 species, mostly Neofusicoccum species, are (among others) Austropuccinia and species of Calonectria,
found associated with eucalypts (Burgess et al. 2019). It is Coniella, Elsinoe, Pseudocercospora, Quambalaria and Ter-
reasonable to assume that, like other Botryosphaeriaceae, they atosphaeria. Species in various genera such as Alternaria,
infect healthy tissues without causing symptoms and that they Botrytis and Neofusicoccum are commonly associated with
appear when these tissues are subjected to stress disease problems and are most likely opportunists that develop
(Schoeneweiss 1980, Slippers & Wingfield, 2007). In the case of on stressed or dying tissues. Many others are putative patho-
leaves, this stress might simply be related to senescence. gens or have other important roles, but these have yet to be
Pathogenicity studies are required to better understand the role studied. This presents an important goal for fungal biologists in
of these fungi as foliage pathogens. the future. It is hoped that this study will promote renewed in-
An important group of eucalypt leaf pathogens reside in the terest in eucalypt leaf fungi, leading to investigations that will
genus Quambalaria (de Beer et al. 2006). These pathogens are provide a better understanding of these fascinating and arguably,
apparently specifically native to Australia and they include at least insufficiently appreciated fungi.
www.studiesinmycology.org 127
CROUS ET AL.
MATERIALS AND METHODS fifth character state and all characters were unordered and of
unequal weight. A heuristic search option with 100 random taxon
Isolates additions and tree bisection and reconnection (TBR) as the
branch-swapping algorithm was used. Branches of zero length
Isolates were obtained from the culture collection (CBS) of the were collapsed and all multiple, equally most parsimonious trees
Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands were saved. The robustness of the trees obtained was evaluated
(WI), and the working collection of Pedro Crous (CPC), housed by 1 000 bootstrap replications (Hillis & Bull 1993). Other mea-
at the Westerdijk Institute, or were freshly isolated sures calculated included tree length (TL), consistency index
(Supplementary Table S1). Single-conidial and ascospore cul- (CI), retention index (RI) and rescaled consistency index (RC).
tures were obtained using the techniques described previously All resulting trees were printed with Geneious v. 7.0.6 (http://
(Crous et al. 1991, Crous 1998). www.geneious.com, Kearse et al. 2012). All new sequences
generated in this study were deposited in NCBI’s GenBank
nucleotide database (www.ncbi.nlm.nih.gov) and the accession
DNA extraction, amplification and sequencing numbers are listed in Supplementary Table S1. The alignments
and respective phylogenetic trees were deposited in TreeBASE,
Fungal mycelium of strains (Supplementary Table S1) was
study number S24522.
harvested with a sterile scalpel and the genomic DNA was iso-
lated using the UltraClean Microbial DNA Isolation Kit (MoBio
Laboratories, Inc., Solana Beach, CA, USA) following the man- Morphology
ufacturers’ protocols. A variety of loci were amplified and
sequenced, depending on which loci are commonly used for a Slide preparations were mounted in lactic acid, Shear’s mounting
specific genus. The “Genera of phytopathogenic fungi: GOPHY” fluid or water, from colonies sporulating on 2 % malt extract agar
(Marin-Felix et al. 2017, 2019) series provided a good starting (MEA), 2 % potato-dextrose agar (PDA), oatmeal agar (OA),
point for references pertaining to commonly used loci and their MEA (Crous et al. 2019c), autoclaved pine needles on 2 % tap
amplification conditions. The resulting fragments were water agar (PNA) (Smith et al. 1996), or autoclaved banana
sequenced in both directions using the PCR primers and the leaves (BLA), and incubated at 25 °C under continuous near-
BigDye Terminator Cycle Sequencing Kit v. 3.1 (Applied Bio- ultraviolet light to promote sporulation. Reference strains and
systems Life Technologies, Carlsbad, CA, USA). DNA specimens are maintained in the CBS culture collection. Sections
sequencing amplicons were purified through Sephadex G-50 through conidiomata were made by hand. Observations were
Superfine columns (Sigma-Aldrich, St. Louis, MO) in MultiScreen made with a Nikon SMZ25 dissection-microscope, and with a
HV plates (Millipore, Billerica, MA). Purified sequence reactions Zeiss Axio Imager 2 light microscope using differential interfer-
were analysed on an Applied Biosystems 3730xl DNA Analyzer ence contrast (DIC) illumination and images recorded on a Nikon
(Life Technologies, Carlsbad, CA, USA). The DNA sequences DS-Ri2 camera with associated software. Colony characters and
generated were analysed and consensus sequences were pigment production were noted after 2–4 wk of growth on MEA,
computed using SeqMan v. 7.0 (Lasergene, Madison, WI, USA). PDA and OA (Crous et al. 2009b) incubated at 25 °C. Colony
colours (surface and reverse) were scored using the colour
charts of Rayner (1970). Taxonomic novelties were deposited in
Phylogenetic analysis MycoBank (www.MycoBank.org; Crous et al. 2004a).
Additional sequences were downloaded from the NCBI’s Gen-
Bank nucleotide database as needed (Supplementary Tables S1
and S2). The downloaded and generated sequences for each RESULTS
gene were aligned with the online version of MAFFT v. 7 (Katoh
& Standley 2013). The alignments were manually checked and DNA amplification and phylogenetic analyses
improved where necessary using MEGA v. 7 (Kumar et al. 2016)
and were concatenated using the same software. The identity of generated sequences was confirmed against the
The phylogenetic methods used in this study included a NCBI GenBank nucleotide database. Several novelties were
Bayesian analysis performed with MrBayes v. 3.2.6 (Ronquist identified through the blast searches, which are described below.
et al. 2012), and Parsimony and distance analyses performed For some novelties, the blast results are discussed, whereas for
with PAUP v. 4.0b10 (Swofford 2003). Missing sequences, when others a phylogenetic tree was generated. Those trees are
present, were treated as missing data in the analyses. MrMo- discussed in the species notes of the respective species and in
deltest v. 2.2 (Nylander 2004) was used to determine the best the legends belonging to the trees.
nucleotide substitution model settings for each data partition in
order to perform a model-optimised Bayesian phylogenetic Taxonomy
reconstruction. The Markov Chain Monte Carlo (MCMC) analysis
of four chains started in parallel from a random tree topology, the Allelochaeta Petr., Sydowia 9: 464. 1955. Fig. 1.
heat parameter was set at 0.25 and trees were saved every 10 Synonyms: Vermisporium H.J. Swart & M.A. Will., Trans. Brit.
generations until the average standard deviation of split fre- Mycol. Soc. 81: 491. 1983.
quencies reached 0.01 (stop value). Burn-in was set to 25 % Discostromopsis H.J. Swart, Trans. Brit. Mycol. Soc. 73: 217.
after which the likelihood values were stationary. The Neigh- 1979.
bour-Joining (NJ) phylogenies performed with PAUP executed
1 000 bootstrap replicates using the HKY85 model. In the Foliicolous. Conidiomata stromatic and acervular, erumpent,
Maximum Parsimony analyses, alignment gaps were treated as unilocular to plurilocular, glabrous, dark brown to black; basal
128
Fig. 1. Allelochaeta spp. A–D. Disease symptoms on eucalypts. E–H. Allelochaeta neocylindrospora (CPC 20115). E. Conidiomata on PNA. F, G. Conidiophores. H. Conidia.
I–L. Allelochaeta samuelii (CPC 28912). I. Conidiomata on OA. J, K. Conidiophores. L. Conidia. Scale bars = 10 μm. (from Crous et al. 2018a).
stroma of textura angularis; dehiscing via an irregular split in the unicellular, hyaline, smooth. Ascomata perithecial, immersed in
overlying host tissue. Conidiophores arising from the upper cells host tissue, solitary or aggregated in cultures, covered by a small
of the basal stroma or lining the cavity of the conidioma, reduced clypeus, obpyriform; wall composed of several layers of thin-
to conidiogenous cells or branched and septate, hyaline or pale walled, brown cells. Asci basal, intermingled among paraphy-
brown at base, smooth, invested in mucus. Conidiogenous cells ses that degenerate at maturity, long elliptical, unitunicate with
discrete or integrated, ampulliform, lageniform, cylindrical, sub- thickened apex and indistinct apical structures. Ascospores
cylindrical, mostly hyaline, or pale brown below, smooth, prolif- biseriate, hyaline, 3-septate, ellipsoid, straight to inaequilateral.
erating percurrently. Conidia fusoid, naviculate, subcylindrical or
acerose, straight or curved, euseptate, wall thin and with or Type species: Allelochaeta gaubae Petr. [= A. dilophospora
without slight constrictions at the septa, smooth; median cells (Cooke) Crous]
hyaline to medium brown; end cells hyaline, bearing mostly a Symptoms: Leaf spots round to angular, amphigenous, often
single appendage at each end, which could be branched; ap- vein-limited, with distinct dark brown to purple-brown margin, and
pendages continuous with the conidium body or cellular, not erumpent brown acervuli visible under the hand lens, frequently
separated from the conidium body by septa, branched or un- exuding a slimy, pale brown to creamy or pinkish conidial mass.
branched, branches filiform or attenuated and flexuous; basal
appendage excentric, narrowly cuneiform to spike-like, podiform Notes: Based on a limited number of isolates and LSU nrDNA
or cellular. Microconidia present in some species, acerose, sequence data, Barber et al. (2011) concluded that Vermisporium
CROUS ET AL.
should be reduced to synonymy under Seimatosporium. How- Synonym: Seimatosporium brevicentrum (H.J. Swart & M.A.
ever, additional isolates and gene loci have revealed that this is a Will.) P.A. Barber & Crous, Persoonia 27: 96. 2011.
generic complex, and that Vermisporium represents a separate
Diagnosis: Conidia narrowly cylindrical, straight, curved or sig-
genus, for which the oldest available name is Allelochaeta
moid, 3-septate, hyaline, smooth walled, hyaline or almost hy-
(Crous et al. 2018a). Allelochaeta appears to be a common foliar
aline in mass, not constricted at the septa, 39–70 × 2–3.5
pathogen of eucalypts, and many more undescribed species
(av. = 54.3 × 2.8) μm; apical cell cylindrical with an obtuse apex,
await to be named.
(13–)18–25(–31) (av. = 21.3) μm long; 2nd cell from apex cy-
Allelochaeta acuta (H.J. Swart & M.A. Will.) Crous, Fungal Syst. lindrical, 5–10(–12) (av. = 7.3) μm long; 3rd cell from apex cy-
Evol. 2: 282. 2018. lindrical, (4–)5–9(–10) (av. = 6.7) μm long; basal cell with a
Basionym: Vermisporium acutum H.J. Swart & M.A. Will., Trans. truncate base, (9–)13–25(–28) (av. = 19.1) μm long; basal
Brit. Mycol. Soc. 81: 495. 1983. appendage excentric, single, cuneiform to podiform with an
Synonym: Seimatosporium acutum (H.J. Swart & M.A. Will.) P.A. obtuse tip, 1–3(–3.5) (av. = 2.1) μm long.
Barber & Crous, Persoonia 27: 93. 2011.
Description and illustration: Swart & Williamson (1983).
Diagnosis: Conidia narrowly fusiform, straight or curved, (2–)
Typus: Australia, Victoria, Mt Macedon, on leaves of E. ovata,
3(–4)-septate, hyaline, orange in mass, slightly or not constricted
1978, M.A. Williamson (holotype AW 78.09 ex Herb. MELU).
at septa, (39–)45–61(–66) × 3–4.5(–5) μm; apical cell narrowly
conical, attenuated to an acute apex, (11–)13–22 μm long; Materials examined: Australia, Victoria, Whittlesea, on E. ovata, 11 Oct. 1999,
second cell from apex cylindrical to sub-cylindrical, (10–) P.A. Barber, PAB99.11; Mt. Burchell, Grampians National Park, on E. serraensis,
12 Aug. 2000, P.A. Barber, GR0.02. New Zealand, Wellington, Karori Cemetery,
11–17 μm long; third cell from apex cylindrical to sub-cylindrical, on Eucalyptus sp., 14 Nov. 1996, B.J. Rogan, NZFRI-M 3645; Catchpool Forest,
9–15 (av. = 11.7) μm long; basal cell with a truncate base, (7–) on E. fastigata, 16 Oct. 1997, B.J. Rogan, NZFRI-M 3756.
9–12(–13) μm long; basal appendage excentric, single, narrowly
Allelochaeta brevilata (H.J. Swart & D.A. Griffiths) Crous,
cuneiform, 2–7 μm long.
comb. nov. MycoBank MB832013.
Description and illustration: Crous et al. (2018a).
Basionym: Seimatosporium brevilatum H.J. Swart & D.A. Grif-
Typus: Australia, Victoria, Rutherglen, on Eucalyptus sp., 1903, fiths, Trans. Brit. Mycol. Soc. 62: 360. 1974.
G.H. Robinson (holotype VPRI 2156); Victoria, Woorndoo, Synonym: Sarcostroma brevilatum (H.J. Swart & D.A. Griffiths)
S37°53’29.3” E 142°47’51.1”, on E. viminalis, 21 Oct. 2009, P.W. Nag Raj, Coelomycetous Anamorphs with Appendage-bearing
Crous (epitype CBS H-23434, culture ex-epitype CBS Conidia (Ontario): 780. 1993.
144168 = CPC 17646).
Diagnosis: Acervuli originating within the epidermis and breaking
Allelochaeta biseptata (H.J. Swart & M.A. Will.) Crous, Fungal through the outer epidermal wall and cuticle to form an irregular
Syst. Evol. 2: 283. 2018. rupture in the centre of the lesion; conidia 4-celled, short fusi-
Basionym: Vermisporium biseptatum H.J. Swart & M.A. Will., form, 13–21 × 5–7 μm, two median cells brown, thick-walled,
Trans. Brit. Mycol. Soc. 81: 492. 1983. together 8.5–14 um long, and end cells paler with long, thin
Synonym: Seimatosporium biseptatum (H.J. Swart & M.A. Will.) hyaline basal (6–16 μm long) and apical (7–18 μm long)
P.A. Barber & Crous, Persoonia 27: 94. 2011. appendages.
Diagnosis: Conidia narrowly cylindrical to acerose, straight, Description and illustration: Swart & Griffiths (1974).
curved or sigmoid, 2-septate, hyaline, orange in mass, not Typus: Australia, New South Wales, Rutherford, on leaves of
constricted at the septa, septa faint, 45–62(–69) × (1.5–)2–2.5 E. globulus, 3 Mar. 1970, F. McKeowen (holotype DAR 19582).
(av. = 54.6 × 2.0) μm; apical cell subcylindrical, tapering to a blunt Not known from culture or DNA.
apex, (17–)20–27 (av. = 23.2) μm long; middle cell short cy-
lindrical, (6–)7–11(–13) (av. = 9.3) μm long; basal cell cylindrical Notes: This species was regarded as a synonym of Seimato-
with a truncate base, (17–)20–29 (av. = 23.5) μm long; basal sporium fusisporum by Nag Raj (1993). However, because
appendage excentric, single, podiform, 1.5–3 (av. = 2.2) μm distinct species in this genus are frequently morphologically
long. similar, these are retained as two separate taxa, based on the
features as discussed by Swart & Griffiths (1974).
Description and illustration: Swart & Williamson (1983).
Allelochaeta cylindrospora (H.J. Swart) Crous, Fungal Syst.
Typus: Australia, Victoria, Melbourne, Box Hill, on E. melliodora, Evol. 2: 284. 2018.
11 Oct. 1903, C. French Jr. (holotype VPRI 2168); New South Basionym: Seimatosporium cylindrosporum H.J. Swart, Trans.
Wales, Northern Tablelands, 7.5 km E of Nundle on road to Brit. Mycol. Soc. 78: 267. 1982.
Hanging Rock (c. 100 m E of Hanging Rock track turnoff), Synonym: Vermisporium cylindrosporum (H.J. Swart) Nag Raj, in
S31°28’31” E151°10’59”, alt. 1 090 m, on E. oresbia, 20 Jul. Nag Raj, Coelomycetous anamorphs with appendage-bearing
2006, A.E. Orme & R. Johnstone, 732739 (epitype CBS H- conidia (Ontario): 965. 1993.
20743, cultures ex-epitype CBS 131116 = CPC 13584, CPC
Diagnosis: Conidia falcate, fusoid, 3-septate, slightly constricted
13585, 13586).
at septa, median cells pale brown, end cells hyaline, smooth-
Allelochaeta brevicentra (H.J. Swart & M.A. Will.) Crous, walled, (28–)35–45(–50) × (3.5–)4(–5) μm; apical cell
Fungal Syst. Evol. 2: 284. 2018. narrowly conical, attenuating toward apex with tubular
Basionym: Vermisporium brevicentrum H.J. Swart & M.A. Will., appendage, 13–15(–19) μm long; second cell from apex cylin-
Trans. Brit. Mycol. Soc. 81: 493. 1983. drical to subcylindrical, (7–)8–10(–12) μm long; third cell from
130
apex cylindrical to subcylindrical, (7–)8–9 μm long; basal cell Basionym: Cryptostictis falcata B. Sutton, Mycol. Pap. 88: 25.
with narrowly truncate base, 6–7 μm long; basal appendage 1963.
excentric, single, tubular to cuneiform, (7–)8–12(–17) μm long. Synonyms: Seimatosporium falcatum (B. Sutton) Shoemaker,
Canad. J. Bot. 42: 416. 1964.
Description and illustration: Nag Raj (1993).
Vermisporium falcatum (B. Sutton) Nag Raj, in Nag Raj, Coe-
Typus: Australia, Victoria, Melton, on E. behriana, 19 Mar. 1977, lomycetous anamorphs with appendage-bearing conidia
I. Pascoe (isotype MELU 2002-5-3); Victoria, Djerriwarrh Creek, (Ontario): 969. 1993.
on E. behriana, 12 May 1972, H.J. Swart (epitype CBS H-23435,
Diagnosis: Conidia falcate, fusiform or sigmoid, 3(–4)-septate,
culture ex-epitype CBS 144169 = CPC 28302 = VPRI 15692).
rarely 5-septate, distinct, slightly to strongly constricted at the
Allelochaeta euabalongensis Crous, Fungal Syst. Evol. 2: 286. septa, guttulate or not guttulate, median cells brown to pale
2018. brown, apical and basal cells pale brown to almost hyaline to
hyaline, periclinal wall verruculose or minutely verruculose,
Diagnosis: Conidia subcylindrical to narrowly fusoid, hyaline,
slightly thicker in the median cells, pale brown to dark brown in
smooth, guttulate, flexuous, 3-septate, constricted at septa or
mass, (31–)34–51(–59) × 4–6 μm; apical cell sub-cylindrical to
not, with prominent taper in apical cell to flexuous appendage,
narrowly conic, pale brown to almost hyaline, upper half hyaline,
(55–)60–70(–75) × (3–)3.5(–4) μm; apical cell attenuating to-
attenuated into a discernible conical or tubular appendage up to
ward a long thin apical appendage with subobtuse apex, (17–)
25 μm, total length including the appendage, (9–)
20–26(–30) μm long; second cell from apex cylindrical to sub-
cylindrical, (12–)13–15(–17) μm long; third cell from apex cy-
cylindrical, brown to pale brown, (6–)7–12(–13) μm long; third
lindrical to subcylindrical, (10–)11–12(–13) μm long; basal cell
cell from apex cylindrical to sub-cylindrical, brown to pale brown,
cylindrical to narrowly obconic with narrowly truncate base, (10–)
(5–)7–12(–13) μm long; basal cell obconic with a truncate base
11–12 μm long; basal appendage excentric, single, cuneiform
and basal appendage, pale brown to almost hyaline, lower half
with subobtuse tip, (7–)10–13 μm long.
hyaline, 5–9(–10) μm long; basal appendage single, excentric,
Description and illustration: Crous et al. (2018a). plectronoid to tubular and flexuous, attenuated to a point, (3–)
4–14(–17) μm long.
Typus: Australia, New South Wales, Euabalong, on leaves of
Eucalyptus sp., 1999, unknown collector (holotype CBS H- Descriptions and illustrations: Nag Raj (1993), Crous et al.
23436, cultures ex-type CBS 112504 = CPC 3777, CBS (2018a).
112332 = CPC 3776).
Typus: Australia, Victoria, on Eucalyptus sp., 1963, collector
Allelochaeta eucalypti (McAlpine) Crous, Fungal Syst. Evol. 2: unknown (holotype IMI 59166); New South Wales, Central Ta-
287. 2018. blelands, ca. 200 m WSW of ‘Coomber’ homestead, on Coomber
Basionym: Cylindrosporium eucalypti McAlpine, Proc. Linn. Soc. property, ca. 8 km SW of Rylstone, S32°50’04” E149°56’13”, alt.
N.S.W. 28: 97. 1903. 600 ± 10 m, on E. alligatrix, 17 Aug. 2006, R. Johnstone & A.E.
Synonyms: Seimatosporium eucalypti (McAlpine) H.J. Swart, Orme, 734259 (epitype CBS H-20744, cultures ex-epitype CBS
Trans. Brit. Mycol. Soc. 78: 268. 1982. 131117 = CPC 13578, CPC 13579, 13580).
Vermisporium eucalypti (McAlpine) Nag Raj, in Nag Raj, Coe-
Allelochaeta flexuosa Crous, Fungal Syst. Evol. 2: 289. 2018.
lomycetous anamorphs with appendage-bearing conidia
(Ontario): 966. 1993. Diagnosis: Conidia acerose, straight to curved, 3-septate, hya-
line, slightly constricted at septa, (70–)75–80(–85) × (3.5–)
Diagnosis: Conidia falcate, fusoid to sigmoid, 3-septate, slightly
4 μm; apical cell long conical, attenuating toward apex, (25–)
constricted at septa, guttulate, median cells pale brown, smooth,
30–36 μm long; second cell from apex cylindrical to sub-
(43–)55–65(–70) × (4–)4.5–5(–6) μm; apical cell cuneiform,
tapering prominently to a tubular apical appendage, up to 20 μm
lindrical to subcylindrical, 15–16 μm long; basal cell with
long; apical cell including appendage (16–)20–26(–30) μm;
narrowly truncate base, (9–)10–12 μm long; basal appendage
second cell cylindrical, hyaline to pale brown, (10–)
excentric, single, tubular with acute apex, (12–)15–18(–25) μm
11–14(–16) μm long; third cell cylindrical, hyaline to pale brown,
long.
(10–)11–13 μm long; basal cell subcylindrical to elongate-
obconical, with a small truncate base, 8–10(–12) μm; basal Description and illustration: Crous et al. (2018a).
appendage single, excentric, tubular and flexuous, attenuating to
Typus: Australia, South Australia, Kangaroo Island, Ravine des
a rounded apex, 10–20 μm long.
Casours Walk, on E. rugosa, Dec. 2011, W. Quaedvlieg (holo-
Descriptions and illustrations: Nag Raj (1993), Crous et al. type CBS H-23440, culture ex-type CBS 144171 = CPC 20173).
(2018a).
Allelochaeta fusispora (H.J. Swart & D.A. Griffiths) Crous,
Typus: Australia, Victoria, Dandenong Creek, on E. melliodora, Fungal Syst. Evol. 2: 289. 2018.
16 Nov. 1902, C. French Jr. (holotype VPRI 5927a); Tasmania, Basionym: Seimatosporium fusisporum H.J. Swart & D.A. Grif-
Lake St. Claire, on E. delegatensis, Mar. 2011, C. Mohammed fiths, Trans. Brit. Mycol. Soc. 62: 360. 1974.
(epitype CBS H-23439, culture ex-epitype CBS 144170 = CPC
Diagnosis: Conidia fusoid, 3-septate, brown, central cells verru-
12458).
culose, constricted at septa, (18–)20–22(–26) × 5(–6) μm; basal
Allelochaeta falcata (B. Sutton) Crous, Fungal Syst. Evol. 2: cell obconical, with truncate base, hyaline, smooth, 3–4 μm long;
288. 2018. two central cells thick-walled, medium brown, verruculose,
CROUS ET AL.
12–17 μm long; apical cell short-conical, 3–4 μm long, with acute Allelochaeta neofalcata Crous, Fungal Syst. Evol. 2: 294. 2018.
apex extending into apical tubular appendage, 10–20 μm long;
Diagnosis: Conidia falcate, fusoid, 3-septate, pale brown, end
basal appendage excentric, tubular, 15–20 μm long.
cells subhyaline, finely roughened, constricted at septa, (46–)
Description and illustration: Crous et al. (2018a). 48–55(–60) × 4–5 μm; apical cell subcylindrical to narrowly
conical, subhyaline, attenuating toward conical or tubular
Typus: Australia, Victoria, near Lake Merrimu, on leaf of
appendage, 20–22(–23) μm long; second cell from apex cylin-
E. polyanthemos, 30 Aug. 1973, H.J. Swart (isotype of Sei-
drical to subcylindrical, subhyaline, (11–)12–13(–14) μm long;
matosporium fusisporum, culture CBS 810.73, specimen CBS H-
third cell from apex cylindrical to subcylindrical, (12–)
17996) = ATCC 26928 = IMI 163446 (culture and specimen),
13–14(–16) μm long; basal cell obconical with narrowly truncate
culture C 73.22.
base, subhyaline, (7–)8–9(–10) μm long; basal appendage
Allelochaeta minor Crous, Fungal Syst. Evol. 2: 291. 2018. excentric, single, tubular, flexuous, (10–)11–13(–15) μm long.
Diagnosis: Conidia acerose, straight to curved, 3-septate, hya- Description and illustration: Crous et al. (2018a).
line, slightly constricted at septa, (25–)29–32(–35) × (3–)
Typus: Australia, Western Australia, Gerby, on E. radiata, 7 Nov.
3.5(–4) μm; apical cell narrowly conical, attenuating toward
2014, P.W. Crous (holotype CBS H-23450, culture ex-type CBS
apex, (10–)12–13(–16) μm long; second cell from apex cylin-
144178 = CPC 25455).
drical to subcylindrical, 6–7 μm long; third cell from apex cylin-
drical to subcylindrical, (6–)7(–8) μm long; basal cell with Allelochaeta neoorbicularis Crous, Fungal Syst. Evol. 2: 294.
narrowly truncate base, 6–7 μm long; basal appendage 2018.
excentric, single, cuneiform, (4–)5–6(–7) μm long.
Diagnosis: Conidia acerose, curved, 3-septate, hyaline, smooth,
Description and illustration: Crous et al. (2018a). constricted at septa, (45–)48–52(–55) × (4–)4.5(–5) μm; apical
cell attenuating toward conical, acute apex, (15–)
Typus: New Zealand, Auckland, Warkworth, Kaipara coast road,
Eucalyptus sp., 2015, R. Thangavel (holotype CBS H-23446,
cultures ex-type CBS 144175 = CPC 29354 = MPI T15_06344A,
CPC 29353).
with narrowly truncate base, (7–)8–9 μm long; basal appendage
Allelochaeta neoacuta Crous, Fungal Syst. Evol. 2: 292. 2018. excentric, single, cuneiform, (5–)6–7(–8) μm long.
Diagnosis: Conidia fusoid, falcate, (2–)3(–5)-septate, hyaline, Description and illustration: Crous et al. (2018a).
orange in mass, slightly or not constricted at septa, (43–)
Typus: Australia, Tasmania, Rosenberg, Eucalyptus sp., Dec.
49–55(–65) × 3(–4) (av. = 50 × 3) μm; apical cell narrowly
2011, W. Quaedvlieg (holotype CBS H-23451, culture ex-type
conical, attenuated to an acute apex, (11–)16–19(–22) μm long;
CBS 144179 = CPC 20140).
second cell from apex cylindrical to sub-cylindrical, (11–)
12–14(–16) μm long; third cell from apex cylindrical to sub- Allelochaeta neowalkeri Crous, Fungal Syst. Evol. 2: 294.
cylindrical, (8–)10–12 μm long; basal cell with a truncate 2018.
base, 8–10(–11) μm long; basal appendage excentric, single,
Diagnosis: Conidia acerose, hyaline, smooth, slightly curved, 3-
cuneiform, (3–)6–8(–10) μm long.
septate, constricted at septa, (50–)55–60(–65) × (3–)
Description and illustration: Crous et al. (2018a). 3.5(–4) μm; apical cell narrowly conical, attenuating toward
apex, (16–)20–22 μm long; second cell from apex cylindrical to
Typus: South Africa, Mpumalanga, Sabie, Sabie Forest Station,
subcylindrical, (13–)15–16(–17) μm long; third cell from apex
on leaves of E. smithii, 28 Sep. 1989, P.W. Crous (holotype
cylindrical to subcylindrical, (12–)13–14(–15) μm long; basal
PREM 50457, isotype CBS H-23448, cultures ex-type CBS
cell with narrowly truncate base, 10–12 μm long; basal
115131 = CPC 156, CBS 110733 = CPC 157, CBS
appendage excentric, single, cuneiform, (4–)6–7 μm long.
110734 = CPC 158, CBS 114876 = CPC 159).
Allelochaeta neocylindrospora Crous, Fungal Syst. Evol. 2:
292. 2018. Typus: Australia, Tasmania, Lovershill, on E. regnans, Dec.
2011, W. Quaedvlieg (holotype CBS H-23453, culture ex-type
Diagnosis: Conidia narrowly fusoid, straight to curved, 3-septate,
CBS 144181 = CPC 20144).
hyaline, orange in mass, not to slightly constricted at septa,
(42–)50–55(–60) × (3–)3.5(–4) μm; apical cell narrowly Allelochaeta obliquae Crous, Fungal Syst. Evol. 2: 295. 2018.
conical, attenuating toward apex, (15–)20–22 μm long; second
Diagnosis: Conidia subcylindrical, hyaline, smooth, guttulate,
cell from apex cylindrical to subcylindrical, 10–13(–16) μm long;
slightly curved, 3-septate, constricted at septa or not, (43–)
third cell from apex cylindrical to subcylindrical, 10–12(–15) μm
45–50(–55) × (3–)4 μm; apical cell narrowly conical, attenuating
long; basal cell with narrowly truncate base, 8–9(–10) μm long;
toward a knob-like apex, (13–)15–17 μm long; second cell from
basal appendage excentric, single, narrowly cuneiform with
apex cylindrical to subcylindrical, (12–)13–14(–15) μm long;
subobtuse tip, (7–)9–10(–15) μm long.
third cell from apex cylindrical to subcylindrical, (10–)
Description and illustration: Crous et al. (2018a). 11–12(–14) μm long; basal cell with narrowly truncate base,
(9–)10–11 μm long; basal appendage excentric, single, cunei-
Typus: Australia, South Australia, Kangaroo Island, Ravine des
form with obtuse apex, 2–3 μm long.
Casours Walk, on E. rugosa, Dec. 2011, W. Quaedvlieg (holo-
type CBS H-23447, culture ex-type CBS 144176 = CPC 20115). Description and illustration: Crous et al. (2018a).
132
Typus: Australia, South Australia, Cape Jervis, on E. obliqua, 25 cell narrowly conical, attenuating toward apex with tubular
Nov. 2011, W. Quaedvlieg (holotype CBS H-23454, culture ex- appendage, (12–)13–14(–15) μm long; second cell from apex
type CBS 144182 = CPC 20191). cylindrical to subcylindrical, (8–)9–10 μm long; third cell from
apex cylindrical to subcylindrical, 8–9(–10) μm long; basal cell
Allelochaeta obtusa (H.J. Swart & M.A. Will.) Crous, Fungal
with narrowly truncate base, 8–9 μm long; basal appendage
Syst. Evol. 2: 296. 2018.
excentric, single, tubular to cuneiform with subobtuse apex,
Basionym: Vermisporium obtusum H.J. Swart & M.A. Will.,
3–4 μm long.
Trans. Brit. Mycol. Soc. 81: 499. 1983.
Synonym: Seimatosporium obtusum (H.J. Swart & M.A. Will.) Description and illustration: Crous et al. (2018a).
P.A. Barber & Crous, Persoonia 27: 107. 2011.
Typus: Australia, South Australia, Cape Jervis, on Eucalyptus
Diagnosis: Conidia ellipsoid-fusiform to subcylindrical, straight, sp., 25 Nov. 2011, W. Quaedvlieg (holotype CBS H-23457,
slightly curved or slightly sigmoid, 3(–4)-septate, hyaline, smooth- culture ex-type CBS 144184 = CPC 20189).
walled, hyaline in mass, not or slightly constricted at the septa,
Allelochaeta parafalcata Crous, Fungal Syst. Evol. 2: 298.
(49–)54–92(–103) × (2.5–)3–4(–4.5) (av. = 70.6 × 3.3) μm;
2018.
apical cell subcylindrical, slightly attenuated to an obtuse tip, (14–)
20–31 (av. = 21.8) μm; second cell from apex cylindrical, Diagnosis: Conidia subcylindrical to fusoid, straight to curved,
13–25(–27) (av. = 17.6) μm; third cell from apex cylindrical, (10–) subhyaline to pale brown, finely verruculose, 3(–4)-septate,
12–21(–25) (av. = 16.3) μm; basal cell subcylindrical with a slightly constricted at septa, (42–)45–55(–70) × (3–)
narrow, truncate base, (8–)11–20(–24) (av. = 15) μm; basal 3.5(–4) μm; apical cell narrowly conical, attenuating toward
appendage tubular, single, excentric, cuneiform to podiform, often apex, (11–)14–15(–17) μm long; second cell from apex cylin-
somewhat swollen in the middle, 2–9(–11) (av. = 5.1) μm long. drical to subcylindrical, (10–)11–12(–13) μm long; third cell from
apex cylindrical to subcylindrical, (10–)11–12 μm long; basal cell
Description and illustration: Barber et al. (2011).
with narrowly truncate base, (5–)6–7(–8) μm long; basal
Typus: Australia, Sherbrooke Forest, Kallista, on E. regnans, 10 appendage excentric, single, cuneiform, (3–)6–7(–8) μm long.
Aug. 1976, H.J. Swart (holotype DAR 43110).
Materials examined: Australia, Victoria, Toolangi, on E. regnans, 16 July 1999,
P.A. Barber, PAB99.13; Victoria, Gellibrand, Otway State Forest, on E. obliqua, 1 Typus: Australia, Western Australia, Wandoo National Park, on
May 2002, P.A. Barber, PAB02.31. Corymbia sp., 13 Jul. 2011, W. Gams (holotype CBS H-23458,
Allelochaeta orbicularis (Cooke) Crous, Fungal Syst. Evol. 2: culture ex-type CBS 144185 = CPC 19840).
296. 2018. Allelochaeta paraorbicularis Crous, Fungal Syst. Evol. 2: 300.
Basionym: Stagonospora orbicularis Cooke, Grevillea 20: 6. 2018.
1891.
Synonyms: Seimatosporium orbiculare (Cooke) P.A. Barber & Diagnosis: Conidia cylindrical to acerose, straight to curved, 3-
Crous, Persoonia 27: 109. 2011. septate, hyaline, slightly constricted at septa, (35–)
Vermisporium orbiculare (Cooke) H.J. Swart & M.A. Will., Trans. 42–47(–52) × (3–)3.5(–4) μm; apical cell narrowly conical,
Brit. Mycol. Soc. 81: 497. 1983. attenuating toward apex, (11–)12–13(–15) μm long; second cell
from apex cylindrical to subcylindrical, (6–)10–12(–14) μm long;
Diagnosis: Conidia solitary, ellipsoid-fusoid to subcylindrical, third cell from apex cylindrical to subcylindrical, (9–)
gently curved or sigmoid, guttulate, 3-septate, hyaline, smooth
10–12(–13) μm long; basal cell with narrowly truncate base,
(orange in mass on leaves and in culture), (40–)
(8–)9–10(–11) μm long; basal appendage excentric, single,
50–55(–60) × (3–)3.5–4 μm; apical cell subcylindrical, tapering
cuneiform to podiform, (3–)4–5(–6) μm long.
toward an acutely rounded apex, 15–18(–23) μm long; second
cell from apex cylindrical, (11–)12–14(–15) μm long; third cell Description and illustration: Crous et al. (2018a).
from apex cylindrical, (10–)11–13 μm long; basal cell sub-
Typus: New Zealand, Auckland, Warkworth, Kaipara coast road,
cylindrical with narrow, truncate base, (8–)10–11(–12) μm;
Eucalyptus sp., 2015, R. Thangavel (holotype CBS H-23452,
basal appendage tubular, single, excentric, cuneiform to podi- culture ex-type CBS 144180 = CPC 29356 = MPI T15_06344B).
form, tapering towards a subobtuse apex, (2–)3–5(–6) μm long.
Allelochaeta polycarpae Crous, Fungal Syst. Evol. 2: 300.
2018.
Typus: Australia: Victoria, on Eucalyptus sp., 24 May 1886,
Diagnosis: Conidia subcylindrical, hyaline, smooth, guttulate,
F.M.C. (holotype K(M) 104759); New South Wales, Australian
flexuous, 3-septate, constricted at septa or not, with prominent
Botanic Garden, Mount Annan, on Corymbia henryi, 3 Mar. 2006,
taper in apical cell, (57–)60–70(–80) × (3–)4 μm; apical cell
B.A. Summerell (epitype CBS H-20745, culture ex-epitype CBS
attenuating toward a long thin apical appendage with subobtuse
131118 = CPC 12935) (treated as Seimatosporium obtusum in apex, (25–)30–35 μm long; second cell from apex cylindrical to
Barber et al. 2011).
subcylindrical, (13–)15–17 μm long; third cell from apex cylin-
Allelochaeta paracylindrospora Crous, Fungal Syst. Evol. 2: drical to subcylindrical, (13–)15–17 μm long; basal cell cylin-
297. 2018. drical to narrowly obconic with narrowly truncate base, (9–)
10–15 μm long; basal appendage excentric, single, cuneiform
Diagnosis: Conidia cylindrical to acerose, straight to curved, 3-
with subobtuse tip, (12–)15–19 μm long.
septate, not or slightly constricted at septa, pale brown in
mass, smooth-walled, (34–)38–45(–50) × 3(–3.5) μm; apical Description and illustration: Crous et al. (2018a).
CROUS ET AL.
Typus: Australia, Western Australia, Cape Riche, Wellstead, on but are shorter. This species can easily be distinguished from its
E. polycarpa, 21 Sep. 2015, P.W. Crous (holotype CBS H- closest phylogenetic neighbours based on rpb2, tef1 and tub2;
23462, culture ex-type CBS 144188 = CPC 28916). on ITS it differs with 2 nts from A. pseudoacuta and A. fusispora.
Allelochaeta pseudoacuta Crous, Fungal Syst. Evol. 2: 301. Allelochaeta pseudowalkeri Crous, Fungal Syst. Evol. 2: 304.
2018. 2018.
Diagnosis: Conidia narrowly fusoid, straight to curved, 3(–4)- Diagnosis: Conidia narrowly fusoid, straight to curved, guttulate,
septate, hyaline, orange in mass, not to slightly constricted at hyaline, smooth, becoming pale brown with age, 3-septate,
septa, (50–)57–65(–70) × (3.5–)4(–4.5) μm; apical cell becoming slightly constricted at septa, (45–)48–55(–60) × (3–)
narrowly conical, attenuating toward apex, (15–)19–21(–24) μm 3.5–4 μm; apical cell tapering prominently to an acutely rounded
long; second cell from apex cylindrical to subcylindrical, (13–) apex, (18–)20–22(–25) μm long; second cell cylindrical, (10–)
15–16(–17) μm long; third cell from apex cylindrical to sub- 11–13(–14) μm long; third cell cylindrical, (8–)10–12(–13) μm
cylindrical, (11–)12–13(–14) μm long; basal cell with narrowly long; basal cell cylindrical with narrowly truncate base, (6–)
truncate base, (9–)11–12(–17) μm long; basal appendage 7–8 μm long; basal appendage excentric, narrowly cuneiform,
excentric, single, narrowly cuneiform, (2–)4–6(–8) μm long. (8–)10–12(–13) μm long, tapering toward an acutely rounded
apex.
Typus: Australia, Tasmania, Lovershill, Eucalyptus sp., 4 Jan.
2012, W. Quaedvlieg (holotype CBS H-23463, culture ex-type Typus: Australia, Queensland, on Eucalyptus sp., 12 Jul. 2009,
CBS 144189 = CPC 20130). P.W. Crous (holotype CBS H-23469, culture ex-type CBS
144195 = CPC 17043).
Allelochaeta pseudofalcata Crous, Fungal Syst. Evol. 2: 302.
2018. Allelochaeta samuelii (Hansf.) Crous, Fungal Syst. Evol. 2:
305. 2018.
Diagnosis: Conidia falcate, fusoid, 3-septate, pale brown, end Basionym: Cylindrosporium samuelii Hansf., Proc. Linn. Soc.
cells subhyaline, smooth, slightly constricted at septa, (40–) N.S.W. 81: 46. 1956.
45–50(–55) × (3–)3.5(–4) μm; apical cell subcylindrical to Synonyms: Seimatosporium samuelii (Hansf.) J. Walker & H.J.
narrowly conical, subhyaline, attenuating toward conical or Swart, Trans. Brit. Mycol. Soc. 90: 287. 1988.
tubular appendage, (16–)17–18 μm long; second cell from apex Vermisporium samuelii (Hansf.) J.A. Simpson & Grgur., Muelleria
cylindrical to subcylindrical, subhyaline, (10–)11–12 μm long; 9: 239. 1996.
third cell from apex cylindrical to subcylindrical, (11–)12–14 μm
long; basal cell obconical with narrowly truncate base, subhya- Diagnosis: Conidia subcylindrical, hyaline, smooth, 3-septate,
line, 8–9 μm long; basal appendage excentric, single, tubular, straight to curved, slightly constricted at septa, guttulate, (45–)
flexuous, (7–)9–10 μm long. 50–70(–75) × 3(–4) μm; apical cell subcylindrical, tapering to a
distinct knob-like apex, 18–27 μm long; median cells cylindrical,
Description and illustration: Crous et al. (2018a). 13–24 μm long; basal cell obconical with a small truncate base,
Typus: Australia, Queensland, Mt. Crosby, on Eucalyptus cre- 16–24 μm; basal appendage single, excentric, cuneiform to
bra, 6 Aug. 1973, J. Alcorn, J.L. 73-133a, deposited as “Sei- podiform, attenuating to an obtuse or truncate tip, 2–4 μm long.
matosporium falcatum” (holotype CBS H-23466, isotypes BRIP Descriptions and illustrations: Barber et al. (2011), Crous et al.
5731, IMI 179066, culture ex-type BRIP 5731 = CBS (2018a).
144192 = CPC 28308 = VPRI 15701).
Typus: Australia, South Australia, on Eucalyptus sp., Sep. 1924, G.
Allelochaeta pseudoobtusa Crous, Fungal Syst. Evol. 2: 302. Samuel (holotype ADW 3840); Western Australia, Cape Riche,
2018. Wellstead, on E. angulosa, 21 Sep. 2015, P.W. Crous (epitype CBS
H-23470, culture ex-epitype CBS 144196 = CPC 28912).
Diagnosis: Conidia subcylindrical to ellipsoid-fusoid, straight to
gently curved, 3-septate, hyaline, smooth, orange in mass, not to Allelochaeta sparsifoliae Crous, Fungal Syst. Evol. 2: 305.
slightly constricted at septa, (48–)54–60(–65) × (3–) 2018.
3.5(–4) μm; apical cell subcylindrical, tip subobtuse, (14–)
Diagnosis: Conidia subcylindrical to acerose, hyaline, smooth,
guttulate, slightly curved, 3-septate, constricted at septa or not,
(45–)48–57(–60) × (3.5–)4 μm; apical cell narrowly conical,
attenuating toward an apical appendage with blunt apex, (15–)
with narrowly truncate base, (10–)11–12(–13) μm long; basal
17–21 μm long; second cell from apex cylindrical to sub-
appendage excentric, single, cuneiform to podiform, (3–)
cylindrical, (15–)16–17 μm long; third cell from apex cylindrical
5–7(–10) μm long.
to subcylindrical, (12–)13–15 μm long; basal cell cylindrical to
Description and illustration: Crous et al. (2018a). narrowly obconic with narrowly truncate base, (9–)
10–12(–14) μm long; basal appendage excentric, single,
Typus: Australia, Tasmania, Tasman Peninsula, S43°11'29.7"
cuneiform to podiform, 5–6 μm long.
E147°51'00.7", on leaves of E. viminalis, 14 Oct. 2006, B.A.
Summerell (holotype CBS H-23467, culture ex-type CBS Description and illustration: Crous et al. (2018a).
144193 = CPC 13590).
Typus: Australia, New South Wales, Wyong, on E. sparsifolia (=
Notes: Conidia of A. pseudoobtusa resemble those of A. obtusa E. oblonga), 23 Sep. 2007, B.A. Summerell (holotype CBS H-
[(49–)54–92(–103) × (2.5–)3–4(–4.5) (av. = 70.6 × 3.3) μm], 23455, culture ex-type CBS 144183 = CPC 14529).
134
Allelochaeta verrucispora (Nag Raj) Crous, Fungal Syst. Evol. mononematous, simple or irregularly and loosely branched, pale
2: 306. 2018. brown or brown, solitary or in fascicles. Conidiogenous cells
Basionym: Vermisporium verrucisporum Nag Raj, in Nag Raj, integrated, terminal becoming intercalary, polytretic, sympodial,
Coelomycetous anamorphs with appendage-bearing conidia or sometimes monotretic, cicatrized. Conidia catenate or solitary,
(Ontario): 972. 1993. dry, ovoid, obovoid, cylindrical, narrowly ellipsoid or obclavate,
Synonym: Seimatosporium verrucisporum (Nag Raj) P.A. Barber beaked or non-beaked, pale or medium olivaceous-brown to
& Crous, Persoonia 27: 111. 2011. brown, smooth or verrucose, with transverse and with or without
oblique or longitudinal septa. Septa can be thick, dark and rigid
Diagnosis: Conidia falcate, fusiform or sigmoid, 3(–4)-septate,
and an internal cell-like structure can be formed. Species with
rarely 5-septate, distinct, slightly to strongly constricted at the
meristematic growth are known. Ascomata small, solitary to
septa, guttulate or not guttulate, median cells brown to pale brown,
clustered, erumpent to (nearly) superficial at maturity, globose to
apical and basal cells pale brown to almost hyaline to hyaline,
ovoid, dark brown, smooth, apically papillate, ostiolate. Papilla
periclinal wall verruculose or minutely verruculose, slightly thicker
short, blunt. Peridium thin. Hamathecium of cellular pseudopar-
in the median cells, pale brown to dark brown in mass, (20–)
aphyses. Asci few to many per ascoma, (4–6–)8-spored, basal,
33–58(–63) × (3.5–)4–6(–6.5) (av. = 48.0 × 5.0) μm; apical cell
bitunicate, fissitunicate, cylindrical to cylindrical-clavate, straight
subcylindrical to narrowly conic, pale brown to almost hyaline,
or somewhat curved, with a short, furcate pedicel. Ascospores
upper half hyaline, attenuated into a discernible conical or tubular
muriform, ellipsoid to fusoid, slightly constricted at septa, yellow-
appendage up to 24 μm, total length including the appendage,
brown, without guttules, smooth, 3–7 transverse septa, 1–2
(7–)11–24(–36) (av. = 18.7) μm long; second cell from apex
series of longitudinal septa through the two original central
cylindrical to subcylindrical, brown to pale brown, (4–)8–14(–15)
segments, end cells without septa, or with one longitudinal or
(av. = 10.2) μm long; third cell from apex cylindrical to sub-
oblique septum, or with a Y-shaped pair of septa (from
cylindrical, brown to pale brown, (5–)7–14(–15) (av. = 10.5) μm
Woudenberg et al. 2013).
long; basal cell obconic with a truncate base and basal
appendage, pale brown to almost hyaline, lower half hyaline, (4–) Type species: Alternaria alternata (Fr.) Keissl.
5–9(–10) (av. = 7.1) μm long; basal appendage single, excentric,
Diagnosis: Leaf spots small, or pale brown, irregular, with diffuse
plectronoid to tubular and flexuous, attenuated to a point, (2–)
margins. Conidiophores and conidia medium golden-brown.
4–15(–25) (av. = 11.3) μm long.
Conidiophores simple, straight or curved, 1–3-septate. Conidia
Descriptions and illustrations: Nag Raj (1993), Barber et al. in branched chains, ovoid, obclavate, obpyriform, rarely ellip-
(2011). soidal, with a conspicuous basal pore, with or without a short
conical or cylindrical apical beak not exceeding one third of the
Typus: Australia, Victoria, Miles Creek, on E. regnans, 1915, C. conidial length, 18–63 × 7–18 μm.
French Jr. (holotype ADW 1784, isotype VPRI 1932b).
Symptom: Leaf spots mostly seen under humid nursery condi-
Allelochaeta walkeri (H.J. Swart & M.A. Will.) Crous, Fungal tions, associated with leaf scorch and herbicide damage, mostly
Syst. Evol. 2: 306. 2018. secondary.
Basionym: Vermisporium walkeri H.J. Swart & M.A. Will., Trans.
Brit. Mycol. Soc. 81: 495. 1983. Notes: Several species of Alternaria have been reported from
Synonym: Seimatosporium walkeri (H.J. Swart & M.A. Will.) P.A. eucalypt leaves, most commonly A. tenuissima and A. alternata
Barber & Crous, Persoonia 27: 113. 2011. (Magnani 1964, Sharma et al. 1985). The introduction of mo-
lecular techniques has had a significant impact on the taxonomy
Diagnosis: Conidia subcylindrical to fusoid, falcate, hyaline, of Alternaria (Woudenberg et al. 2013, 2015), and further
smooth, 3(–4)-septate, slightly constricted at septa, guttulate, research is now required to confirm the identity of the species
(52–)55–62(–67) × 3(–3.5) μm; apical cell conical, tapering to involved on eucalypts and elucidate their role as pathogens of
an obtuse apex, 18–27 μm long; second cell cylindrical, this host.
10–15 μm long; third cell cylindrical, 10–13 μm long; basal cell
with a small truncate base, 10–15 μm; basal appendage single, Amycosphaerella Quaedvlieg & Crous, Persoonia 33: 22. 2014.
excentric, cuneiform to podiform, 6–12 μm long. Fig. 3.
Descriptions and illustrations: Barber et al. (2011), Crous et al. Foliicolous, plant pathogenic. Ascomata pseudothecial,
(2018a). amphigenous, solitary, black, subepidermal, globose, with central
apical ostioles, becoming papillate; walls of 2–3 layers of me-
Typus: Australia, Victoria, Hume Highway, N of Wallen, on dium brown textura angularis, subhymenium of 1–2 layers of
E. obliqua, 29 Aug. 1980, H.J. Swart (holotype DAR 43109); hyaline cells. Asci obovoid to broadly ellipsoidal, straight or
Victoria, Melbourne, ‘Lamatina’s Farm’, S38°24’26.2”, incurved, 8-spored. Ascospores bi- to triseriate, overlapping,
E144°55’9”, on Eucalyptus sp., 12 Oct. 2009, P.W. Crous (epi- hyaline, guttulate, straight, fusoid-ellipsoidal with obtuse ends,
type CBS H-20746, cultures ex-epitype CBS 131119 = CPC widest in middle of apical cells, medianly 1-septate, tapering
17644, CPC 17645). toward both ends, but more prominently toward base.
Alternaria Nees, Syst. Pilze (Würzburg): 72. 1816 (1816–1817). Type species: Amycosphaerella africana (Crous & M.J. Wingf.)
Fig. 2. Quaedvlieg & Crous
Colonies effuse, usually grey, dark blackish brown or black. Notes: The genus Amycosphaerella represents a
Mycelium immersed or partly superficial; hyphae hyaline, mycosphaerella-like genus that requires DNA data for generic
olivaceous-brown or brown. Stroma rarely formed. Setae circumscription. Of the species presently known, A. africana has
and hyphopodia absent. Conidiophores macronematous, had a confused history, having been described several times
CROUS ET AL.
Fig. 2. Alternaria spp. Alternaria sect. Alternata. A. Colony on SNA. B–P. Conidia and conidiophores. B, N. A. daucifolii. C, L, M. A. arborescens. D, H–J. A. alternata. O.
A. gaisen. F. A. limoniasperae. F, K. A. tenuissima. G, P. A. longipes. Scale bars = 10 μm. (from Woudenberg et al. 2013).
under different names based on confusion related to its mode of Synonyms: Teratosphaeria africana (Crous & M.J. Wingf.) Crous
ascospore germination. & U. Braun, Stud. Mycol. 58: 8. 2007.
Mycosphaerella aggregata Carnegie & Keane, Mycol. Res. 98:
Amycosphaerella africana (Crous & M.J. Wingf.) Quaedvlieg &
415. 1994. nom. illegit. (Art. 53.1). (non Mycosphaerella aggre-
Crous, Persoonia 33: 23. 2014. Fig. 3.
gata (Schwein.) J.A. Stev. 1918).
Basionym: Mycosphaerella africana Crous & M.J. Wingf.,
Mycosphaerella gregaria Carnegie & Keane, Mycol. Res. 101:
Mycologia 88: 450. 1996.
843. 1997. nom. inval. (Art. 41.5, Melbourne).
136
Fig. 3. Amycosphaerella spp. A, D. Disease symptoms of A. ellipsoidea. B, C. Disease symptoms of A. africana. E–H. Asci, ascospores, and germinating ascospores of
A. africana. Scale bars = 10 μm.
Phaeophleospora gregaria (Carnegie & Keane) Quaedvlieg & Diagnosis: Ascospores tri- to multiseriate, overlapping, colour-
Crous, Persoonia 33: 23. 2014. nom. inval. (Art. 39.1, less, guttulate, thin-walled, straight, fusoid-ellipsoidal with obtuse
Melbourne). base and subobtuse apex, widest in middle of apical cell,
Mycosphaerella ellipsoidea Crous & M.J. Wingf., Mycologia 88: medianly 1-septate, not constricted at septum, tapering toward
452. 1996. both ends, but with more prominent taper toward base (7–)
Mycosphaerella buckinghamiae Crous & Summerell, Australas. 8–9(–11) × 2.5–3 μm.
Pl. Pathol. 29: 272. 2000.
Description and illustration: Crous (1998).
Mycosphaerella aurantia A. Maxwell, Mycol. Res. 107: 353.
2003. Typus: Kenya, on leaf litter of E. grandis, May 1995, M.J.
Wingfield (holotype PREM 54402, cultures ex-type CBS
Diagnosis: Leaf spots amphigenous, subcircular, 2–10 mm
111001 = CMW 5147 = CPC 1084, CPC 1085, 1086).
diam, pale brown, surrounded by slightly raised borders. Asco-
spores straight or slightly curved, fusoid-ellipsoidal, widest just Anthostomella Sacc., Atti Soc. Veneto-Trent. Sci. Nat., Padova,
above the septa, 1-septate, not constricted at median septa, (8–) Ser. 4 4: 84. 1875.
10–11 × (2–)2.5–3 μm.
Ascomata immersed or semi-immersed, clypeate or not, dark
Description and illustration: Crous & Wingfield (1996), Carnegie brown, solitary to clustered, globose to ellipsoid or conical.
& Keane (1997). Central ostiole usually lined with periphyses. Peridium usually
Typus: South Africa, Western Cape Province, Stellenbosch, of thin-walled brown textura angularis. Paraphyses fila-
Stellenbosch Mountain, on E. viminalis, Oct. 1994, P.W. Crous mentous, hyaline, flexuous, unbranched, septate. Asci (4–)
(holotype of M. africana PREM 51917, cultures ex-type CBS 8-spored, cylindrical to broadly so, unitunicate, short-
116154 = CMW 4945 = CPC 794, CBS 116155 = CPC 795, CBS pedicellate, usually with J+ subapical ring. Ascospores
680.95 = CBS 116259 = CPC 796). 1(–2)-seriate, ellipsoid to inequilaterally ellipsoid, unicellular,
or with a larger dark brown cell and a hyaline basal dwarf cell,
Notes: Ascospores can be up to 15 μm in length, not always mostly smooth-walled, with gelatinous sheath, with or without
constricted at septa, and in some cases, germinate from their germ slit.
polar ends with germ tubes parallel to the long axis, remain hy-
aline, and develop lateral branches (Maxwell et al. 2003). Col- Type species: Anthostomella limitata Sacc.
lections of A. africana known to date show that this pathogen has Symptoms: Leaf spots medium brown, subcircular with a raised,
variable morphology and results in diverse disease symptoms. thin border and narrow, red-purple margin, up to 8 mm diam.
Amycosphaerella keniensis (Crous & T.A. Cout.) Videira & Note: Several additional species with wider host ranges are
Crous, Stud. Mycol. 87: 329. 2017. known from Eucalyptus (Lu & Hyde 2000), although they appear
Basionym: Mycosphaerella keniensis Crous & T.A. Cout., Mycol. to be saprobes.
Mem. 21: 74. 1998.
CROUS ET AL.
Anthostomella eucalypti H.Y. Yip, Mycol. Res. 93: 75. 1989. Diagnosis: Conidia (7–)8–10(–11) × (5–)6(–7) μm, broadly
ellipsoid to obovoid, apex obtusely rounded, aseptate, non-
Diagnosis: Ascospores 14.5–17.5 × 5.5–8 × 3–5 μm, inequi-
apiculate, medium brown, thick-walled, smooth, striations along
laterally ellipsoidal or ellipsoidal with one side flattened and
length of conidium body, with prominent central guttule. Basal
rounded.
appendage absent or hyaline, tubular, smooth, thin-walled,
Description and illustration: Yip (1989). devoid of cytoplasm, 0–2 μm long, 2 μm diam.
Typus: Australia, Victoria, Ivanhoe, Darebin Parklands, Darebin Description and illustration: Marin-Felix et al. (2019).
Creek, on E. camaldulensis, 22 Feb. 1987, H.-Y. Yip (holotype
Typus: Malaysia, Sabah, on E. pellita, May 2015, M.J. Wingfield
MELU 7877). Not known from culture.
(holotype CBS H-23081, culture ex-type CBS 142518 = CPC
Anthostomella eucalyptorum Crous & M.J. Wingf., Fungal 27550).
Planet: 1. 2006. Fig. 4.
Apoharknessia eucalyptorum Crous & M.J. Wingf., Persoonia
Diagnosis: Ascospores uniseriate, ellipsoid, smooth-walled, with 38: 271. 2017. Fig. 5.
a central guttule, consisting of a larger brown cell, (7–)
Diagnosis: Conidia (8–)9–10(–11) × (5–)6–6.5(–7) μm,
8–9(–10) × 4–5(–5.5) μm, and a smaller, hyaline, basal dwarf
obliquely gibbose, aseptate, brown, smooth, thick-walled, with
cell, 1.5–2 μm long and wide, lacking a mucus sheath and germ
prominent central guttule, lacking striations, with conical short
slit.
apiculus. Basal appendage (1.5–)2–3(–3.5) × 2–2.5 μm, hya-
Description and illustration: Crous et al. (2006a). line, tubular, smooth, thin-walled, devoid of cytoplasm, 0–2 μm
long, 2 μm diam.
Typus: Indonesia, Northern Sumatra, on Eucalyptus sp., Feb.
2004, M.J. Wingfield (holotype CBS H-19727, cultures ex-type Description and illustration: Crous et al. (2017b).
CBS 120036 = CPC 11023, CPC 11024, 11025).
Apoharknessia Crous & S.J. Lee, Stud. Mycol. 50: 239. 2004. (holotype CBS H-23082, culture ex-type CBS 142519 = CPC
Fig. 5. 27546).
Similar to Harknessia, but distinct in having a hyaline, apical Apoharknessia insueta (B. Sutton) Crous & S.J. Lee, Stud.
apiculus, and not forming fluffy aerial mycelium on oatmeal or Mycol. 50: 240. 2004. Fig. 5.
malt extract agar, but growing within the medium, and also Basionym: Harknessia insueta B. Sutton, Mycol. Pap. 123: 20.
sporulating on naked hyphae. 1971.
Type species: Apoharknessia insueta (B. Sutton) Crous & S.J. Diagnosis: Conidia 10–11(–12.5) × 7.5–9 μm, obliquely gib-
Lee bose, aseptate, brown, smooth, thick-walled, with prominent
central guttule, lacking striations, with conical short apiculus.
Symptoms: Associated with medium brown, subcircular leaf
Basal appendage (1.5–)2–3(–3.5) Х 2–2.5 μm, hyaline,
spots that can cover large portions of the leaf.
tubular, smooth, thin-walled, devoid of cytoplasm, 0–2 μm long,
Notes: Although originally assumed to be a pathogen of minor 2 μm diam.
importance, Apoharknessia spp. have more recently been
Description and illustration: Lee et al. (2004).
associated with prominent leaf spotting of eucalypts in South
America and Asia (Crous, unpubl. data). Typus: Mauritius, Les Urares, on E. robusta, 19 Jan. 1933, G.
Orian (holotype IMI 22697). Brazil, Amazonia, Jari, on E. pellita,
Apoharknessia eucalypti Crous & M.J. Wingf., Stud. Mycol. 92:
8 Jul. 1996, P.W. Crous (epitype CBS H-9913, culture ex-epitype
51. 2018 (2019).
CBS 111377 = CPC 1451).
Fig. 4. Anthostomella eucalyptorum (CPC 11023). A. Immersed ascomata. B, C. Asci and ascospores. D. Asexual morph. Scale bars: A = 200 μm, others = 10 μm.
138
Fig. 5. Apoharknessia spp. A. Disease symptoms of Apoharknessia eucalyptorum on Eucalyptus pellita. B, C. Conidiomata. B. Apoharknessia eucalyptorum (ex-type CBS
142519). C. Apoharknessia insueta (ex-type CBS 111377). D–G. Conidiogenous cells and conidia. D, E. Apoharknessia eucalyptorum (ex-type CBS 142519). F, G. Apo-
harknessia insueta (ex-type CBS 111377). H, I. Conidia. H. Apoharknessia eucalyptorum (ex-type CBS 142519). I. Apoharknessia insueta (ex-type CBS 111377). Scale bars:
C = 25 μm; others = 10 μm. (from Marin-Felix et al. 2019).
Aurantiosacculus Dyko & B. Sutton, Mycologia 71: 922. 1979. Notes: Aurantiosacculus represents a genus of foliar pathogens
Fig. 6. that appears to be restricted to Eucalyptus. Species are regarded
to be of minor importance.
Conidiomata amphigenous, eustromatic, subepidermal,
becoming erumpent, rupturing epidermis, appearing bright or- Aurantiosacculus acutatus Crous & Summerell, Persoonia 28:
ange; subglobose to flattened, with ostiole central, opening via 70. 2012. Fig. 6.
irregular flaps in upper layer of conidioma. Conidiophores sub-
cylindrical, septate, hyaline, smooth, lining the inner layer of cavity, Diagnosis: Conidia hyaline, smooth, aseptate, sigmoid, apex
unbranched or branched below. Conidiogenous cells lageniform subobtuse, base swollen, obtuse with central, thickened,
to subcylindrical, hyaline, smooth, integrated, determinate; apex somewhat refractive scar, 1–1.5 μm diam, at times with marginal
with minute periclinal thickening and collarette. Conidia hyaline, frill, (40–)50–57(–67) × (2–)2.5(–3) μm.
smooth, aseptate, sigmoid, apex obtuse to subobtuse, base
swollen, obtuse with central, thickened, somewhat refractive scar, Description and illustration: Crous et al. (2012b).
at times with marginal frill.
Typus: Australia, Tasmania, Crescent Bay, S43°11'13.9",
Type species: Aurantiosacculus eucalypti (Cooke & Massee) E147°50'50.7", on E. viminalis, 14 Oct. 2006, B.A. Summerell &
Dyko & B. Sutton P. Summerell (holotype CBS H-20933, culture ex-type CBS
132181 = CPC 13704).
Symptoms: Associated with amphigenous, brown, irregular leaf
spots that have erumpent, characteristic yellow-orange Aurantiosacculus eucalypti (Cooke & Massee) Dyko & B.
conidiomata. Sutton, Mycologia 71: 922. 1979. Fig. 6.
CROUS ET AL.
Fig. 6. Aurantiosacculus spp. A, B. Disease symptoms of A. eucalypti (A), and A. acutatus (B). C, D. Close-up of conidiomata of A. eucalypti (C), and A. acutatus (D). E–H.
Conidiogenous cells and conidia of A. eucalyptorum (CPC 13229). Scale bars = 10 μm.
Basionym: Protostegia eucalypti Cooke & Massee, Grevillea ellipsoidal with obtuse ends, medianly 1-septate, widest in mid-
16(no. 79): 75. 1888. dle of apical cell, not constricted at septum or only slightly so.
Diagnosis: Conidia hyaline, smooth, aseptate, sigmoid, apex Type species: Australosphaerella nootherensis (Carnegie)
subobtuse, base swollen, obtuse with central, thickened, Videira & Crous
somewhat refractive scar, 51–81 × 2–3 μm.
Note: Very little is presently known regarding the importance or
Description and illustration: Dyko et al. (1979). distribution of Australosphaerella nootherensis (Carnegie et al.
2011); it is known only from a single location.
Typus: Australia, Victoria, Melbourne, on E. incrassata, Reader
no. 24 (holotype K(M) 195742). Not known from culture. Australosphaerella nootherensis (Carnegie) Videira & Crous,
Stud. Mycol. 87: 370. 2017. Fig. 7.
Aurantiosacculus eucalyptorum Crous & C. Mohammed,
Basionym: Mycosphaerella nootherensis Carnegie, Austral. Pl.
Persoonia 28: 71. 2012. Fig. 6.
Pathol. 40: 377. 2011.
Diagnosis: Conidia hyaline, smooth, aseptate, sigmoid, apex
Diagnosis: Leaf spots amphigenous, marginal, irregular
subobtuse, base swollen, obtuse with central, thickened,
(5–15 mm diam) to commonly blighting, light brown to reddish
somewhat refractive scar (2 μm diam), at times with marginal frill,
brown with prominent dark brown border (1–2 mm), often with
(48–)55–60(–67) × (2.5–)3–3.5(–4) μm.
purple staining emanating from necrotic area. Ascospores tri- to
Description and illustration: Crous et al. (2012b). multiseriate, overlapping, hyaline, straight to rarely curved,
fusoid-ellipsoidal with obtuse ends, medianly 1-septate, widest in
Typus: Australia, Tasmania, on E. globulus, 31 Aug. 2006, C. middle of apical cell, not constricted at septum or only slightly so,
Mohammed & M. Glen (holotype CBS H-20934, culture ex-type tapering towards both ends but more prominently towards the
CBS 130826 = CPC 13229). basal end, apical cell occasionally asymmetrical, (9.5–)
Australosphaerella Videira & Crous, Stud. Mycol. 87: 370. 2017. 10.5–11.5(−13) × 3–3.5 μm.
Description and illustration: Carnegie et al. (2011).
Ascomata pseudothecial, black, slightly erumpent, globose. Asci
aparaphysate, fasciculate, bitunicate, subsessile, obclavate to Typus: Australia, Queensland, Noosa Heads, on C. intermedia,
ellipsoidal, straight to incurved, 8-spored. Ascospores multi- 11 Aug. 2008, A.J. Carnegie (holotype and ex-type culture BRIP
seriate, overlapping, hyaline, straight to rarely curved, fusoid- 52584a).
140
Fig. 7. Australosphaerella nootherensis (BRIP 52584a). A. Disease symptoms. B, C. Germinating ascospores. D, E. Ascospores. Scale bars = 10 μm.
Austroafricana Quaedvlieg & Crous, Persoonia 33: 25. 2014. Typus: Australia, New South Wales, South Grafton, Grafton City
Fig. 8. Council Landfill Plantation, S29°46'21", E152°54'38", on leaves
of C. henryi, 16 Feb. 2006, A.J. Carnegie (holotype CBS H-
Foliicolous, plant pathogenic. Ascomata pseudothecial, black,
19833, isotype DAR 78031, cultures ex-type CBS
subepidermal to erumpent, globose, with central apical ostiole;
120730 = CPC 13119, CPC 13120, occurring with lembosina-like
wall consisting of 2–3 layers of medium brown textura angularis.
sp.).
Asci aparaphysate, but with remains of hamathecium visible,
fasciculate, bitunicate, subsessile, obovoid to ellipsoidal, straight Austroafricana keanei (Carnegie & G.S. Pegg) Quaedvlieg &
to slightly curved, 8-spored. Ascospores tri- to multiseriate, Crous, Persoonia 33: 25. 2014.
overlapping, hyaline, guttulate, thick-walled, straight, fusoid- Basionym: Teratosphaeria keanei Carnegie & G.S. Pegg, Aus-
ellipsoidal with obtuse ends, medianly 1-septate, tapering to- tralas. Pl. Pathol. 40: 368. 2011.
wards both ends, but more prominently towards the lower end;
Diagnosis: Leaf spots amphigenous, commonly irregular, occa-
ascospores with or without persistent mucus sheath. Germi-
sionally circular, 1–6 mm diam, separate, rarely confluent, red-
nating ascospores become either verruculose, brown and dis-
dish brown becoming greyish brown on adaxial surface,
torted, or remain hyaline and undistorted.
yellowish brown on abaxial surface, with prominent reddish
Type species: Austroafricana associata (Crous & Carnegie) brown border. Pseudothecia hypophyllous. Ascospores fusoid-
Quaedvlieg & Crous ellipsoid, constricted at median septum, hyaline, widest in mid-
dle of apical cell, 9.5–10.5 × 2.5–3 μm.
Notes: Species of Austroafricana are found to co-colonise le-
sions of their hosts together with other ascomycetes and to have Description and illustration: Carnegie et al. (2011).
wide host ranges. Austroafricana parva and A. associata, for
Typus: Australia, Queensland, Kingaroy, Berry’s Plantation, on
example, are known as pathogens of Eucalyptus and Protea-
E. globulus × E. camaldulensis, 14 Feb. 2004, A.J. Carnegie
ceae (Crous et al. 2008, Quaedvlieg et al. 2014).
(holotype BRIP 52593b, culture ex-type CBS 130524).
Austroafricana associata (Crous & Carnegie) Quaedvlieg &
Austroafricana parva (R.F. Park & Keane) Quaedvlieg & Crous,
Crous, Persoonia 33: 25. 2014. Fig. 8.
Persoonia 33: 25. 2014. Fig. 8.
Basionym: Mycosphaerella associata Crous & Carnegie, Fungal
Basionym: Mycosphaerella parva R.F. Park & Keane, Trans. Brit.
Diversity 26: 159. 2007.
Mycol. Soc. 79: 99. 1982.
Synonym: Teratosphaeria associata (Crous & Carnegie) Crous &
Synonyms: Teratosphaeria parva (R.F. Park & Keane) Crous &
U. Braun, Stud. Mycol. 58: 9. 2007.
U. Braun, Stud. Mycol. 58: 10. 2007.
Diagnosis: Leaf spots amphigenous, irregular to sub-circular, Mycosphaerella grandis Carnegie & Keane, Mycol. Res. 98: 414.
4–6 mm diam, medium brown, with a thin, raised, dark brown 1994.
border on the adaxial surface; dark brown with patches of grey
Diagnosis: Leaf spots amphigenous, confined to the margin of
due to the lifting cuticle on the abaxial surface, displaying
the leaf extending from the tip almost back to the petiole, often
numerous small cracks within the lesion tissue. Ascomata pre-
less than 1.5 cm wide, grey to yellow-brown often with a red-
dominantly hypophyllous, black, subepidermal to erumpent.
brown margin (less than 0.5 mm wide) on the adaxial surface,
Ascospores thick-walled, straight, fusoid-ellipsoidal with obtuse
occasionally confluent with lesions of Thyrinula eucalypti. As-
ends, widest in middle of apical cell, medianly 1-septate, con-
cospores multiseriate, overlapping, hyaline, guttulate, thick-
stricted at the septum, (12–)13–16(–17) × (3.5–)4–5(–6) μm;
walled, straight, ellipsoidal with obtuse ends, widest in middle
ascospores with persistent mucoid sheath.
of the apical cell, medianly 1-septate, constricted at septum,
Description and illustration: Crous et al. (2007a). tapering toward both ends, (7–)8–9(–10) × (2–)2.5(–3) μm.
CROUS ET AL.
Fig. 8. Austroafricana spp. A–D. Disease symptoms of A. parva. E–J. A. associata (CBS H-19833). E. Leaf spot with ascomata. F. Asci with ascospores. G–I. Germinating
ascospores. J. Ascospores. Scale bars = 10 μm. (from Crous et al. 2007a).
Description and illustration: Park & Keane (1982a). tonsure. Telia without sterile elements, partly developed within
uredinia; teliospores puccinioid, with pedicel, 2-celled, with a
Typus: Australia, Victoria, Nowa Nowa, on E. globulus, Jul.
vertical, median septum, ellipsoidal to ovoid, wall smooth, dark
1981, R.F. Park (holotype IMI 263258 (published as 263358));
yellow to brown, germ-pores one per cell, indistinct, formed by an
Victoria, Otway Ranges, (near Gellibrand), S38°34’6.3”,
area of thinned wall. Metabasidia 4-celled, producing four ba-
E143°32’22.5”, alt. 175 m, on E. globulus, Sep. 2005, I. Smith
sidiospores (from Beenken 2017).
(epitype CBS H-20098, cultures ex-epitype CBS 122892 = CPC
12421, CPC 12422, 12423). Type species: Austropuccinia psidii (G. Winter) Beenken
Austropuccinia Beenken, Phytotaxa 297: 55. 2017.
Notes: Austropuccinia psidii is native to South America where it
Spermogonia and aecia unknown. Uredia and telia on leaves, has caused severe disease in Eucalyptus plantations (Ferreira
buds and fruits of Myrtaceae. Uredinia without sterile elements; 1983) and guava (Psidium gaujava) plantations (Ribeiro &
urediniospores pedicellate, globose to ovoid, walls hyaline to Pommer 2004). It spread to Jamaica and Florida with subse-
light-yellow, ornamentation echinulate, sometimes with basal quent epidemics in all-spice (Pimenta dioica) (Smith 1935) and
142
exotic Melaleuca quinquqnervia (Rayachhetry et al. 1997). It has on some plant hosts. Urediniospores ellipsoidal or ovoid, pyr-
subsequently spread to Hawaii, Japan, China, Australia, New iform, yellowish brown, 17–21 × 13–17 μm; wall 1.5–2 μm
Caledonia, South Africa Singapore, Indonesia and New Zealand thick, finely echinulate with a distinctive smooth patch in the
(Carnegie & Pegg 2018). In Australia, it has established on over lower half, germ pore absent or inconspicuous. Telia on fruit,
350 plant species in 57 genera of Myrtaceae (Carnegie & Pegg leaves or stems, up to 0.5 mm diam., abaxial, erumpent, pul-
2018) and represents one of the more serious global quarantine vinate, yellow to yellowish brown. Teliospores cylindrical or
threats for the cultivation of eucalypts and other Myrtaceae. ellipsoidal, apex rounded, pale yellowish brown,
Globally, A. psidii is known from 525 species across 70 genera 23–50 × 14–28 μm; wall 1–2 μm thick, smooth, 2-celled,
(Soewarto et al. 2019). Several native species in Australia have remnant of pedicel remains attached up to 15 μm long (from
now been listed as critically endangered due to repeated infec- Shivas et al. 2014).
tion leading to tree mortality (Carnegie & Pegg 2018).
Descriptions and illustrations: Carnegie et al. (2010), Shivas
Austropuccinia psidii (G. Winter) Beenken, Phytotaxa 297: 55. et al. (2014), Beenken (2017).
2017. Fig. 9.
Typus: Brazil, S~ao Francisco do Sul, on Psidium guajava (=
Basionym: Puccinia psidii G. Winter, Hedwigia 23: 172. 1884.
Psidium pomiferum), Apr. 1884, E. Ule #14, (lectotype BR-MYC
Diagnosis: Symptoms range from minor leaf spots to severe 80409,93 (ex Museum Botanicum Berolinense); isolectotype
foliage and stem blight, as well as infection on flowers and fruit DAR 29790 (microscope slide)); Araquari, on P. guajava, 6 Feb.
Fig. 9. Austropuccinia psidii. A–G. Disease symptoms on leaves and petioles. H, I, L. Urediniospores. J, K. Urediniospores and teliospores. Scale bars = 10 μm.
CROUS ET AL.
2013, A.C. Alfenas & P.S. Machado (epitype VIC 42496; iso- Bagadiella koalae Crous et al., Persoonia 26: 127. 2011.
epitype BRIP 61521). See Machado et al. (2015).
Diagnosis: Conidiophores subcylindrical, brown, smooth, straight
Bagadiella Cheew. & Crous, Persoonia 23: 59. 2009. to gently curved, 1–3-septate, 15–30 × 3–4 μm. Conidiogenous
cells terminal, integrated, pale to medium brown, smooth,
Mycelium immersed, becoming superficial when incubated
7–15 × 2–3 μm; apex with flared collarette, 1–2 × 2–3 μm.
in moist chambers, pale to medium brown, consisting of
Conidia hyaline, smooth, curved, with bluntly rounded apex and
septate, branched, smooth. Chlamydospores absent. Caespituli
truncate base, (15–)17–20 × 1.5–2 μm.
suprastomatal, pale brown, compact, arising from pseudopar-
enchymatal tissue in the substomatal cavity, forming a rosette Description and illustration: Crous et al. (2011a).
with a central, basal point of attachment, giving rise to co-
Typus: Australia, Victoria, Otway Ranges, Kennett River, Great
nidiophores with a slimy conidial mass, up to 110 μm high
Ocean Road, on E. globulus, 18 Oct. 2009, P.W. Crous, I.G.
and 130 μm diam. Conidiophores micronematous, arranged
Pascoe, I.J. Porter & J. Edwards (holotype CBS H-20583,
in a rosette, cylindrical, mostly dichotomously branched,
culture ex-type CBS 129523 = CPC 17682).
slightly thick-walled, medium to pale grey-brown, straight or
slightly flexuous. Conidiogenous cells terminal, monophialidic,
Bagadiella lunata Cheew. & Crous, Persoonia 23: 60. 2009.
branched, subcylindrical to lageniform, at times constricted at
Fig. 10.
base of conidiogenous cell, tapering toward the apex, pale
brown, paler toward the apex, with a terminal, narrow, pale
Diagnosis: Conidiogenous cells predominantly terminal, mono-
olivaceous, vase-shaped, flaring collarette, constricted beneath
phialidic, branched, subcylindrical to lageniform, (8.5–)
the collarette, thickened and slightly darkened at the con-
11–13(–15) × 2.5–3.3 μm. Conidia borne in slimy heads, lunate,
idiogenous regions. Conidia borne in slimy heads, lunate,
curved, with a rounded apex, tapering toward a subtruncate
curved, apex rounded, with slight taper towards the subtruncate
base, hyaline, (15–)16–18(–22) × (1.3–)1.5(–1.7) μm.
base, hyaline. Ascomata immersed, globose, not visible on the
surface, with periphysate ostiolar channel; wall of 3–4 layers of Description and illustration: Cheewangkoon et al. (2009).
medium brown textura angularis. Asci 8-spored, cylindrical,
Typus: Australia, Tasmania, Mount Wellington Park S42°55’0”,
unitunicate, apical apparatus reacting in Melzer’s reagent.
E147°15’0” on E. delegatensis, 10 Oct. 2006, coll. B.A. Sum-
Paraphyses hyaline, smooth, cellular, intermingled between
merell, isol. P.W. Crous (holotype CBS H-20281, cultures ex-
asci, constricted at septa, unbranched. Ascospores uniseriate,
type CBS 124762 = CPC 13655, CPC 13656).
aseptate, guttulate, ellipsoid, ends acutely rounded, smooth,
hyaline. Bagadiella victoriae Crous et al., Persoonia 26: 125. 2011.
Type species: Bagadiella lunata Cheew. & Crous Diagnosis: Conidiophores subcylindrical, brown, smooth, straight
to gently curved, 1–3-septate, 30–50 × 2–3 μm. Conidiogenous
Symptoms: Leaf spots subcircular, inconspicuous or pale
cells terminal, integrated, pale to medium brown, smooth,
yellow, becoming more prominent once leaves are incu-
10–15 × 1.5–2 μm; apex with flared collarette, 1–2 × 2–3 μm.
bated in moist chambers due to prominent olivaceous to
Conidia hyaline, smooth, curved, with bluntly rounded apex and
grey sporulation on lesion. Species of Bagadiella are endo-
truncate base, (15–)17–22(–25) × (1–)1.5 μm.
phytes, with symptoms becoming visible on older senescent
leaves. Description and illustration: Crous et al. (2011a).
Notes: Bagadiella is a genus of foliar pathogens of minor Typus: Australia, Victoria, Main Ridge, 244 Shands Road,
importance. It appears to be an endophyte, which becomes Sunny Ridge Strawberry Farm, S38°24'3.1" E144°59'36.9" on
prominent (with chlorotic lesions and abundant pale yellow- Eucalyptus sp., 12 Oct. 2009, P.W. Crous, I.J. Porter & J.
brown conidiophore fascicles) once leaves are incubated in Edwards (holotype CBS H-20582, culture ex-type CBS
moist chambers. 129522 = CPC 17688).
Bagadiella eucalypti Crous, Persoonia 39: 431. 2017. Blastacervulus H.J. Swart, Trans. Brit. Mycol. Soc. 90: 289.
1988. emend.
Diagnosis: Conidiogenous cells integrated, as terminal ends on
hyphae, pale brown, smooth, subcylindrical, 10–17 × 2 μm. Conidiomata acervular to sporodochial, brown, subcuticular with
Conidia solitary, hyaline, smooth, subcylindrical, apex obtuse, a single layer of brown epidermal cells. Conidiophores reduced to
base truncate, strongly curved, (12–)14–17(–21) × (1.5–)2 μm. conidiogenous cells. Conidiogenous cells enteroblastic, phialidic,
Ascomata immersed, globose, not visible on the surface, ampulliform, determinate, discrete, hyaline, smooth, with minute
150–250 μm diam. Asci 8-spored, cylindrical, unitunicate, apical periclinal thickening. Conidia in a dry powdery mass, globose to
apparatus reacting in Melzer’s reagent, 110–130 × 10–13 μm. ellipsoidal to subcylindrical, medium brown, thick-walled, verru-
Paraphyses hyaline, smooth, cellular, intermingled between asci. culose, guttulate, aseptate to multiseptate, frequently in chains, at
Ascospores uniseriate, aseptate, guttulate, ellipsoid, ends times with a thin mucoid sheath, base truncate, unthickened.
acutely rounded, smooth, hyaline, (13–)15–16(–18) × (6.5–)
Type species: Blastacervulus eucalypti H.J. Swart
7 μm.
Symptoms: Leaf lesions prominent on leaf tips, amphigenous,
subcircular to irregular, discrete to influent, up to 2 mm diam,
Typus: Australia, Victoria, Nowa Nowa, on E. globulus, 30 Nov. medium brown at the middle, darker at the border, with a red-
2016, P.W. Crous (holotype CBS H-23310, culture ex-type CBS purple margin, with amphigenous, dark conidiomata at the
143439 = CPC 32619). margin, surrounded by indistinct border, not vein-limited.
144
Fig. 10. Bagadiella lunata. A. Caespituli on leaf. B–G. Rosettes of conidiophores giving rise to falcate, aseptate conidia. Scale bars = 10 μm. (from Cheewangkoon et al. 2009).
Note: Species of Blastacervulus are associated with dark brown, thick-walled, verruculose, guttulate, frequently in chains of
almost corky lesions, similar to those associated with Alysidiella two, at times with a thin mucoid sheath, base truncate,
and Thyrinula (see below), except that conidiomata are unthickened, 2–3 μm diam, apex obtuse, aseptate conidia,
frequently arranged in concentric circles or radiating rows. 5–6 × 4–5 μm, 1-septate (7–)8–9(–10) × 4–5 μm, 2-septate
(10–)12–14(–17) × 4–5 μm.
Blastacervulus eucalypti H.J. Swart, Trans. Brit. Mycol. Soc.
90: 289. 1988. Fig. 11. Description and illustration: Crous et al. (2016b).
Diagnosis: Conidiomata acervular, dark brown, single, 5–15 per Typus: Australia, Western Australia, Kalgan, Gull Rock National
lesion, frequently in radiating rows or circles. Conidia pale to Park, S35°00.520 E118°02.329, on E. adesmophloia, 22 Sep.
medium brown, aseptate, 5–7 × 5–8 μm, mostly subglobose to 2015, P.W. Crous (holotype CBS H-22862, culture ex-type CBS
broadly ovoid, slightly obtuse to truncate at the base, thick- 142065 = CPC 29450).
walled, 1–1.5 μm, forming branched chains of acropetal con-
Blastacervulus robbenensis (Crous et al.) Crous, comb. nov.
idia; ramoconidia with up to three hila, 1 μm wide.
MycoBank MB832015.
Description and illustration: Swart (1988), Cheewangkoon et al. Basionym: Stigmina robbenensis Crous et al., Mycol. Res. 99:
(2009), Giraldo et al. (2017). 651. 1995.
Synonyms: Heteroconium kleinzeense Crous & Z.A. Pretorius,
Typus: Australia, Victoria, Calder Highway near Woodend, on
as “kleinziense”, Fungal Diversity 25: 28. 2007.
E. obliqua, 13 Oct. 1979, H.J. Swart (holotype DAR 58998);
Alysidiella kleinziense (Crous & Z.A. Pretor.) Cheew. & Crous,
New South Wales, Mullion Creek, S33°06'48", E149°08'45", on
Mycol. Progr. 11: 63. 2012.
E. robertsonii subsp. hemisphaerica, 26 Jan. 2007, coll. B.A.
Alysidiella suttonii Cheew. & Crous, Mycol. Progr. 11: 63. 2012.
Summerell, isol. P.W. Crous (epitype CBS H-20278, culture ex-
epitype CBS 124759 = CPC 13956). Diagnosis: Leaf spots predominantly hypophyllous, frequently
not extending through the lamina, dark brown, with a whitish
Blastacervulus eucalyptorum Crous, Persoonia 37: 293. 2016.
centre and chlorotic margin, irregularly circular, separate,
Diagnosis: Conidiomata acervular, dark brown. Conidia in a 2–5 mm diam. Conidiophores 1–3-septate, 20–40 × 7–8 μm.
dry powdery mass, globose to ellipsoidal, medium brown, Conidiogenous cells proliferating percurrently, 7–10 × 7–8 μm.
CROUS ET AL.
Fig. 11. Blastacervulus eucalypti (CBS 124759). A–D. Leaf spots with characteristic black, acervular conidiomata. E. Vertical section though conidioma. F, G. Conidia. H.
Colony on MEA. I. Hyphae forming conidia in culture. Scale bars: B–D = 300 μm, E = 100 μm, all others = 10 μm. (from Giraldo et al. 2017).
Conidia subcylindrical to ellipsoid, (1–)3–5(–9)-distoseptate, sp., Apr. 2005, Z.A. Pretorius (holotype of Heteroconium
10–60(–72) × (6–)7–10 μm. kleinziense CBS H-19767, cultures ex-type CBS 120138 = CPC
12174, CPC 12175, 12176); Western Cape Province, Robben
Descriptions and illustrations: Crous et al. (1995, 2007b),
Island, on Eucalyptus sp., Sep. 1993, C.L. Lennox (holotype of
Cheewangkoon et al. (2012).
Stigmina robbenensis PREM 51730, isotype IMI 359478);
Typus: Cyprus, Larnaca, on Eucalyptus sp., 28 Mar. 2007, A. Robben Island, on Eucalyptus sp., May 2015, P.W. Crous, HPC
van Iperen (holotype of Alysidiella suttonii CBS H-20305, cul- 381 (epitype of Stigmina robbenensis designated here CBS H-
tures ex-type CBS 124780 = CPC 13957, CPC 13958, 13959). 24044, MBT388146, single conidial cultures ex-epitype CBS
South Africa, Northern Cape Province, Kleinzee, on Eucalyptus 145898 = CPC 27661, CPC 27659–27660, 27662).
146
Notes: Fresh collections of this pathogen were recently collected swollen towards base, probably spermatial in function (from
from Robben Island, South Africa, making it possible to resolve Hyde & Cannon 1999).
issues pertaining to the taxonomy and apparent restricted
Type species: Brobdingnagia nigeriensis (Sivan. & Okpala) K.D.
distribution of B. robbenensis. The latter pathogen was previ-
Hyde & P.F. Cannon
ously only known from Robben Island, and assumed to be ab-
sent from other regions where eucalypts are grown. Based on Notes: Very little is known regarding the biology B. eucalypticola,
these results, it is possible to confirm that B. robbenensis also and it will also need to be recollected to clarify its phylogenetic
occurs along the coastline of South Africa, where it was incor- position. It appears to be of minor importance as a pathogen.
rectly assumed to represent a distinct species, A. kleinziense,
Brobdingnagia eucalypticola Sivan. & R.G. Shivas, Fungal
and in Cyprus, were it was known as A. suttonii. See the
Diversity 11: 145. 2002. Fig. 14.
phylogenetic tree presented in Fig. 12 for relationships between
the species. Diagnosis: Leaf spots 1–2.5 mm diam, roughly circular “tar
spots”, the affected leaf tissue brown and necrotic with a purplish
Botrytis P. Micheli ex Pers., Neues Mag. Bot. 1: 120. 1794.
black to brown border at the edge of the area. Ascomata epi-
Colonies effuse, often grey, powdery, with visible brown co- genous to hologenous, developing below the cuticle or the
nidiophores and glistening heads of grey conidia. Sclerotia epidermis. Clypeus composed of epidermal and some palisade
frequently formed both on natural substrata and in culture. Co- tissue filled with fungal cells. Paraphyses numerous, simple or
nidiophores macronematous, mononematous, straight or flex- branched, hyaline, filamentous, septate. Asci broadly cylindrical
uous, smooth, brown, branched, often dichotomously or to cylindrical-clavate, unitunicate, non-amyloid, 75 138 ×
trichotomously, with branches mostly restricted to the apical re- 10–22 μm. Ascospores allantoid, hyaline, thin-walled, aseptate,
gion forming a stipe and a rather open head; branches often smooth to roughened, 38–115 × 7–11 μm.
markedly swollen at their ends to form hyaline conidiogenous
Description and illustration: Sivanesan & Shivas (2002a).
ampullae that collapse with age. Conidiogenous cells integrated,
terminal on branches, polyblastic, determinate, inflated, clavate Typus: Australia, Queensland, Brisbane, Gumdale, on Euca-
or subspherical, denticulate. Conidia solitary, simple, hyaline or lyptus sp., 4 Feb. 1986, M. Cox (holotype BRIP 15042). Not
pale brown, smooth, predominantly aseptate, ellipsoidal, known from culture.
obovoid, spherical or subspherical (adapted from Ellis 1971).
Caliciopsis Peck, Ann. Rep. N.Y. St. Mus. nat. Hist. 33: 32. 1883
Type species: Botrytis cinerea Pers. (1880).
Note: Botrytis cinerea causes a disease of plants including eu- Ascomata solitary, brown, superficial, stalked, ostiolate (con-
calypts known as grey mould. The disease is mainly problematic sisting of a cylindrical stalk and swollen upper part), centrum
in nurseries associated with high humidity and cool tempera- containing thin-walled, pseudoparenchymatous, hyaline tissue.
tures. It is occasionally found in young overstocked plantations Asci are bitunicate, 8-spored, apically rounded, pedicellate,
where the microclimate enables infection to occur. saccate, smooth, lacking a discharge mechanism. Ascospores
are globose, smooth to minutely verrucose, brown, aseptate
Botrytis cinerea Pers., Ann. Bot. (Usteri) 1: 32. 1794. Fig. 13.
(adapted from Benny et al. 1985). The asexual morph is phoma-
Diagnosis: Grey mould, leaf blight, dieback, diffuse brown leaf like (see C. eucalypti for details).
spots (along margins and leaf tips), and leaf blight. Co-
Type species: Caliciopsis pinea Peck
nidiophores erect, stipe straight, subhyaline to brown, septate,
branched towards the apex, branches lateral, alternate, at a wide Note: Nothing is known regarding the ecology or importance of
angle to the axis, successively developed from the base to the Caliciopsis eucalypti, and it is assumed to be of minor impor-
apex, branching again alternately, forming at each end a tance (but see Hypsotheca below).
globose, swollen conidiogenous cell bearing conidia on pedicels.
Caliciopsis eucalypti Crous, Persoonia 37: 295. 2016. Fig. 15.
Conidia obovoid to ellipsoid, synchronously formed, with a
slightly protuberant hilum, 8–16 × 6–12 μm, subhyaline to pale Diagnosis: Associated with medium brown, subcircular leaf spots
brown, smooth. Sclerotial morph developing in culture. up to 5 mm diam. Conidiomata solitary, erumpent, globose,
brown. Conidiophores lining the inner cavity, at times reduced to
Descriptions and illustrations: Mittal et al. (1987), Brown & Wylie
conidiogenous cells, ampulliform to doliiform, hyaline, smooth,
(1991), Park et al. (2000).
4–7 × 3–4 μm, or elongated, branched, 1–4-septate, with ter-
Brobdingnagia K.D. Hyde & P.F. Cannon, Mycol. Pap. 175: 47. minal, and lateral conidiogenous loci, 10–40 × 2–3 μm. Pa-
1999. raphyses intermingled among conidiophores. Conidia solitary,
hyaline, smooth, bean-shaped, ends obtuse, mostly slightly
Stromata broadly ellipsoid, matt black, shallowly domed, 1–4-
curved, 3–4 × 1.5 μm.
loculate, with shallow crateriform ostioles. Ascomata oblate,
walls hyaline, morphologically separate from stromatic tissue. Description and illustration: Crous et al. (2016b).
Interascal tissue present, of thin-walled, filiform paraphyses. Asci
Typus: Australia, Western Australia, Albany, Stirling Range
ovoid to broadly saccate or clavate, thin-walled, apparently
National Park, Mt Hassell walk, on E. marginata, 23 Sep. 2015,
evanescent, 8-spored. Ascospores hyaline, thin-walled, asep-
P.W. Crous (holotype CBS H-22863, culture ex-type CBS
tate, smooth, without sheath. Asexual morph with locules
142066 = CPC 28872).
completely lined by layer of conidiogenous cells formed on
cluster of short conidiophore, cylindrical, usually tapering to- Calonectria De Not., Comm. Soc. crittog. Ital. 2(fasc. 3): 477.
wards apex, proliferating percurrently. Conidia filiform, often 1867. Figs 16, 17.
CROUS ET AL.
Fig. 12. The first of 45 equally most parsimonious trees obtained from a phylogenetic analysis of the Blastacervulus/Thyrinula combined ITS, LSU, chs, second part of tef1 and
tub2 sequence alignment (54 strains including the outgroup; 2 919 characters analysed: 1 881 constant, 542 variable and parsimony-uninformative and 496 parsimony-
148
Fig. 13. Botrytis cinerea. A. Colony on SNA. B–D. Conidiophores. E. Conidia. Scale bars = 10 μm.
Synonym: Cylindrocladium Morgan, Bot. Gaz. 17: 191. 1892. dark brown-red, angular cells, merging with a erumpent stroma,
cells of the outer wall layer continuing into the pseudoparen-
Ascomata perithecial, solitary or in groups, globose to sub- chymatous cells of the erumpent stroma. Asci 8-spored, clavate,
globose to ovoid, yellow to orange to red or red-brown to brown, tapering to a long thin stalk. Ascospores aggregated in the upper
turning darker red to red-brown in KOH, rough-walled; perithecial third of the ascus, hyaline, smooth, fusoid with rounded ends,
apex consisting of flattened, thick-walled hyphal elements with straight to sinuous, unconstricted, or constricted at the septa.
rounded tips forming a palisade, discontinuous with warty wall, Megaconidiophores if present, borne on the agar surface or
gradually becoming thinner towards the ostiolar canal, and immersed in the agar; stipe extensions mostly absent; co-
merging with outer periphyses; perithecial base consisting of nidiophores unbranched, terminating in 1–3 phialides, or
informative). The tree was rooted to Leptosphaeria doliolum (GenBank JF740205.1, GQ387576.1, JF740162.1, GU349069.1, JF740144.1, respectively) and the scale bar
indicates the number of changes. Thickened branches represent those present in the strict consensus parsimony tree. Bootstrap support values higher than 49 % are shown at
the nodes and novelties are highlighted with bold text. Species names are indicated to the right of the tree. A superscript T denotes strains with a type status and ET those with
ex-epitype status. Tree statistics: TL = 2 585 steps, CI = 0.564, RI = 0.715, RC = 0.403. The analysis was performed using PAUP* v. 4.0b10 (Swofford 2003).
CROUS ET AL.
Fig. 14. Brobdingnagia eucalypticola (BRIP 15042). A. Leaf spots. B. Close-up of ascomata. C. Vertical section through ascoma. D–F. Asci and ascospores. Scale
bars = 10 μm.
sometimes with a single subterminal phialide; phialides straight aseptate, each terminal branch producing 1–6 phialides; phia-
to curved, cylindrical, seemingly producing a single conidium; lides cylindrical to allantoid, straight to curved, or doliiform to
periclinal thickening and an inconspicuous, divergent collarette reniform, hyaline, aseptate, apex with minute periclinal thickening
rarely visible. Megaconidia hyaline, smooth, frequently remaining and inconspicuous divergent collarette. Macroconidia cylindrical,
attached to the phialide, multi-septate, widest in the middle, bent rounded at both ends, straight or curved, widest at the base,
or curved, with a truncated base and rounded apical cell. Mac- middle, or first basal septum, 1- to multi-septate, lacking visible
roconidiophores consist of a stipe, a penicillate arrangement of abscission scars, held in parallel cylindrical clusters by colour-
fertile branches, a stipe extension, and a terminal vesicle; stipe less slime. Microconidiophores consist of a stipe and a penicillate
septate, hyaline or slightly pigmented at the base, smooth or or subverticillate arrangement of fertile branches. Primary
finely verruculose; stipe extensions septate, straight to flexuous, branches 0–1-septate, subcylindrical; secondary branches 0–1-
mostly thin-walled, terminating in a thin-walled vesicle of char- septate, terminating in 1–4 phialides; phialides cylindrical,
acteristic shape. Conidiogenous apparatus with 0–1-septate straight to slightly curved, apex with minute periclinal thickening
primary branches; up to eight additional branches, mostly and marginal frill. Microconidia cylindrical, straight to curved,
Fig. 15. Caliciopsis eucalypti (CPC 28872). A. Colony sporulating on MEA. B. Conidioma. C–E. Conidiogenous cells. F. Conidia. Scale bars = 10 μm.
150
Fig. 16. Disease symptoms associated with Calonectria. A, B. Defoliated and damaged trees in plantations. C, E. Cutting rot. D, F. Leaf spots. G. Stem canker with ascomata
developing. H. Stem canker with asexual (Cylindrocladium) sporulation.
rounded at apex, flattened at base, 1(–3)-septate, held in 1979). Defoliation caused by CLB decreases timber volume due
asymmetrical clusters by colourless slime (adapted from Crous to the reduced photosynthetic area (Berger et al. 2007, Alfenas
2002). et al. 2009). It also promotes weed growth due to light in the
understory (Alfenas et al. 2009).
Type species: Calonectria pyrochroa (Desm.) Sacc.
The most effective method to control this disease in the field is
Symptoms: Damping off, root rot, discrete pale brown leaf spots, by planting resistant species and genotypes (Alfenas et al. 2009).
leaf blight, shoot blight, stem cankers and root disease (Crous This can be difficult, because several different species are
2002). Calonectria pteridis, which is an important pathogen in associated with CLB even in the same area. Furthermore, there
Brazil, causes small (1–7 mm diam), round or elongated leaf is a movement of Calonectria species from natural forests to
spots, that are initially pale grey, becoming pale brown. Other commercial forest nurseries, and again from nurseries to com-
species such as those in the C. candelabra complex, and mercial plantations, which further complicates disease control
C. ovata, cause large, pale brown leaf botches (see Alfenas et al. (Alfenas et al. 2015).
2013).
Calonectria aciculata J.Q. Li et al., IMA Fungus 8: 273. 2017.
Notes: Calonectria leaf blight (CLB) is an important leaf and
Diagnosis: Calonectria aciculata can be distinguished from the
shoot disease of Eucalyptus wherever these trees are grown in
phylogenetically closely related Ca. colhounii, Ca. honghensis,
plantations (Alfenas et al. 2009, 2013). Although many more
and Ca. monticola in the longer macroconidia.
species have been associated with infections of Eucalyptus roots
and stems, only those confirmed to cause CLB are treated here. Description and illustration: Li et al. (2017).
On most Eucalyptus species, CLB is characterised by small,
Typus: China, Yunnan Province, Puer Region, Jinggu County,
circular or elongated pale grey to pale brown spots that extend
WeiYuan Town, on leaves of an E. urophylla × E. grandis hybrid
throughout the leaf blade, resulting in leaf drop and in some
clone, 16 Nov. 2014, S.F. Chen & J.Q. Li (holotype PREM 61941,
cases severe defoliation (Alfenas & Ferreira 1979, Alfenas et al.
culture ex-type CBS 142883 = CERC 5342 = CMW 47645).
CROUS ET AL.
Fig. 17. Calonectria spp. A–H. Sexual morphs. A–D. Perithecia. A. Calonectria asiatica (ex-type CBS 114073). B. Calonectria braziliensis (ex-type CBS 230.51 × CBS
114257). C. Calonectria fujianensis (ex-type CBS 127201). D. Section through perithecium of Calonectria asiatica (ex-type CBS 114073). E–F. Asci. E. Calonectria crousiana
(ex-type CBS 127198). F. Calonectria asiatica (ex-type CBS 114073). G–H. Ascospores. G. Calonectria fujianensis (ex-type CBS 127201). H. Calonectria acicola (ex-type CBS
114813). I–AB. Asexual morphs. I–L. Macroconidiophores. I. Calonectria malesiana (ex-type CBS 112752). J. Calonectria macroconidialis (ex-type CBS 114880). K. Calo-
nectria spathulata (ex-type CBS 555.92). L. Calonectria ovata (CBS 111307). M–O. Conidiogenous apparatus. M. Calonectria brachiatica (ex-type CBS 123700). N. Calonectria
ecuadoriae (ex-type CBS 111406). O. Calonectria hurae (CBS 114551). P. Microconidiophore of Calonectria reteaudii (ex-type CBS 112144). Q. Megaconidia of Calonectria
hurae (CBS 114551). R, S. Macroconidia. R. Calonectria angustata (ex-type CBS 109065). S. Calonectria chinensis (ex-type CBS 114827). T. Microconidia of Calonectria
pteridis (ex-type CBS 111793). U–AB. Terminal vesicles of stipe extensions. U. Calonectria brassicae (ex-type CBS 111869). V. Calonectria rumohrae (CBS 109062). W.
Calonectria cylindrospora (CBS 119670). X. Calonectria hongkongensis (ex-type CBS 114828). Y. Calonectria chinensis (ex-type CBS 114827). Z. Calonectria humicola (ex-type
CBS 125251). AA. Calonectria mexicana (ex-type CBS 110918). AB. Calonectria spathulata (ex-type CBS 555.92). Scale bars: A–C = 500 μm; D–F = 100 μm; G, H, M–P,
R–AB = 10 μm; I–L, Q = 20 μm. (from Marin-Felix et al. 2017).
152
Calonectria baviensis N.Q. Pham, Mycologia 111: 90. 2019. Diagnosis: Perithecia solitary or in groups of up to five, orange,
becoming red-brown with age. Ascospores fusoid with rounded
Diagnosis: Calonectria baviensis can be distinguished from its
ends, straight to slightly curved, (1–)3-septate, (56–)
closely related species Ca. acacicola, Ca. microconidialis and
58–69(–76) × (5–)6.5–7.5(–8) μm (av. = 64 × 7 μm). Vesicles
Ca. pentaseptata by the size of macroconidia [(82–)
clavate, (4 –)4.5–5(–6) μm diam. Macroconidia (59–)
87.5–104.5(–120) × (5–)6–7(–8) μm, 5-septate], and the ability
61–67(–75) × (4–)4.5–5.5(–6) μm (av. = 64 × 5 μm), (1–)3-
to produce microconidiophores and microconidia.
septate.
Description and illustration: Pham et al. (2019).
Description and illustration: Chen et al. (2011).
Typus: Vietnam, Hanoi, Ba Vi, from leaf of E. urophylla, Nov.
Typus: China, FuJian Province, on leaves of E. grandis, Aug.
2013, N.Q. Pham & T.Q. Pham (holotype PREM 62111, culture
2007, M.J. Wingfield (holotype PREM 60453, culture ex-type
ex-type CBS 143563 = CMW 47410).
CBS 127198 = CMW 27249).
Calonectria brasiliensis (Bat. & Cif.) L. Lombard et al., Stud.
Calonectria eucalypti L. Lombard et al., Stud. Mycol. 66: 47.
Mycol. 66: 19. 2010. Fig. 17.
2010.
Basionym: Cylindrocladium scoparium var. brasiliense Bat. &
Cif., Bol. Secr. Agric. (Pernambuco) 18(3–4): 188. 1952 [1951]. Diagnosis: Perithecia solitary or in groups, yellow to orange,
Synonym: Cylindrocladium brasiliensis (Bat. & Cif.) Peerally, becoming brown with age. Ascospores fusoid with rounded ends,
“braziliensis” CMI Descriptions of Pathogenic Fungi and Bacteria straight to slightly curved, (1–)3-septate, not or slightly con-
427. 1974. stricted at the septum, (25–)30–36(–56) × (3–)5–6(–8) μm
(av. = 33 × 6 μm). Vesicles broadly clavate, 4–6 μm diam.
Diagnosis: Vesicles ellipsoidal to obpyriform vesicle, 7–11 μm
Macroconidia cylindrical, rounded at both ends, straight, (66–)
diam. Macroconidia cylindrical, rounded at both ends, straight,
69–75(–80) × (5–)6 μm (av. = 72 × 6 μm), 3-septate.
(35–)36–40(–41) × 3–5 μm (av. = 38 × 3.5 μm), 1-septate.
Description and illustration: Lombard et al. (2010b).
Description and illustration: Lombard et al. (2010a).
Typus: Indonesia, Sumatra Utara, Aek Nauli, on leaf of
Typus: Brazil, Ceara State, on Eucalyptus sp., Sep. 1948, T.R.
E. grandis, May 2005, M.J. Wingfield (holotype PREM 60298,
Ciferri (holotype IMI 299576, culture ex-type CBS
culture ex-type CBS 125275 = CMW 18444).
230.51 = CMW 23671 = IMI 299576).
Calonectria eucalypticola R.F. Alfenas et al., Stud. Mycol. 80:
Calonectria candelabrum (Viegas) Rossman et al., Stud.
105. 2015.
Mycol. 80: 210. 2015.
Basionym: Cylindrocladium candelabrum Viegas, Bragantia 6: Diagnosis: Vesicles ellipsoidal to obpyriform, 5–7 μm diam.
370. 1946. Macroconidia straight to slightly curved, (43–)
Synonyms: Calonectria scoparia Ribeiro & Matsuoka, In: Ribeiro, 49–52(–55) × 3–5 μm (av. = 50 × 4 μm), 1-septate.
M.Sc. Thesis, Heterotalismo em C. scoparium Morgan: 28. 1978
Description and illustration: Alfenas et al. (2015).
(nom. inval., Art. 29).
Calonectria scoparia Peerally, Mycotaxon 40: 341. 1991. Typus: Brazil, Minas Gerais state, Santa Barbara, from stem of
Eucalyptus seedling, Dec. 2010, A.C. Alfenas (holotype CBS H-
Diagnosis: Perithecia solitary or in groups, red-brown. Asco-
21359, culture ex-type CBS 134847 = LPF124).
spores fusoid with rounded ends, slightly curved, 1-septate,
not or slightly constricted at the septum, (40–) Calonectria foliicola L. Lombard et al., Stud. Mycol. 80: 167.
45–50(–60) × 5–6 μm (av. = 48 × 5.5 μm). Vesicles ellipsoidal to 2015.
narrowly obpyriform, (5–)6–7(–8) μm diam. Macroconidia (45–)
Diagnosis: Vesicles obpyriform to ellipsoidal, 6–13 μm diam.
58–68(–80) × 4–5(–6) μm (av. = 60 × 4.5 μm), 1-septate.
Macroconidia (41–)44–50(–52) × (3–)4–5(–6) μm (av.
Description and illustration: Crous (2002). 47 × 5 μm), 1-septate.
Typus: Brazil, Bahia: Picad~ao, Conceiç~ao de Barra, on Description and illustration: Lombard et al. (2015).
E. grandis, Apr. 1992, A.C. Alfenas & F.A. Ferreira (neotype of
Typus: China, Guangxi Province, from E. urophylla × E. grandis
Ca. scoparia PREM 51045); Copener, Eucalyptus sp., A.C.
clone leaf, Mar. 2009, X. Zhou & G. Zhao (holotype CBS H-
Alfenas (neotype of Cy. candelabrum PREM 51044).
21472, cultures ex-type CBS 136641 = CERC 1728 = CMW
Calonectria cerciana L. Lombard et al., Persoonia 24: 7. 2010. 31393 = CPC 23491, CERC 1729 = CMW 31394 = CPC 23492,
CERC 1730 = CMW 31395).
Diagnosis: Vesicles fusoid to obpyriform, 8–13 μm diam.
Macroconidia (37–)41–46(–49) × 5–6 μm (av. = 44 × 5 μm), Calonectria fujianensis S.F. Chen et al., Persoonia 26: 8. 2011.
1-septate.
Diagnosis: Perithecia solitary or in groups of up to four, bright
Description and illustration: Lombard et al. (2010c). yellow, becoming orange with age. Ascospores fusoid with rounded
ends, (1–)3-septate, not or slightly constricted at the septum, (38–)
Typus: China, Guangdong Province, CERC nursery, on stems of
49–62(–72) × (5–)6–7.5(–8) μm (av. = 55.5 × 6.8 μm). Vesicles
E. urophylla × E. grandis hybrid cutting, Nov. 2007, M.J. Wing-
clavate, (3–)3.5–4.5(–5) μm diam. Macroconidia (48–)
field & X.D. Zhou (holotype PREM 60241, culture ex-type CBS
50–55(–60) × (2.5–)3.5–4.5(–5) μm (av. = 52.5 × 4 μm), (1–)3-
123693 = CMW 25309).
septate.
Calonectria crousiana S.F. Chen et al., Persoonia 26: 6. 2011.
Fig. 17. Description and illustration: Chen et al. (2011).
CROUS ET AL.
Typus: China, FuJian Province, on leaves of E. grandis, Aug. macroconidiale PREM 51035, culture ex-type CBS
2007, M.J. Wingfield (holotype PREM 60460, culture ex-type 114880 = CPC 307); ibid., E. grandis cuttings, Mar. 1990, P.W.
CBS 127201 = CMW 27257). Crous (holotype of Ca. macroconidialis PREM 51036).
Calonectria indusiata (Seaver) Crous, Taxonomy and Pathol- Calonectria matogrossensis R.A. Fernandes et al., Persoonia
ogy of Cylindrocladium (Calonectria) and Allied Genera (St 42: 393. 2019.
Paul): 94. 2002.
Diagnosis: Vesicles ellipsoid to obpyriform, 4–9 μm diam.
Basionym: Nectria indusiata Seaver, Mycologia 20: 58. 1928.
Macroconidia (42–)47–50 × (3.5–)4–5 μm (av. 47 × 4 μm), 1-
Synonym: Cylindrocladium theae (Petch) Subram., In: Sub-
septate.
ramanian, Hyphomycetes, an account of Indian species, except
Cercosporae: 731. 1971. Description and illustration: Crous et al. (2019a).
Diagnosis: Perithecia solitary or in groups, orange to red, Typus: Brazil, Mato Grosso, Primavera do Leste, on E. urophylla
becoming red-brown. Ascospores fusoid with rounded ends, clone I144, 2015, R.A. Alfenas (holotype UB24025).
straight to slightly curved, (1–)3-septate, not or slightly con-
Calonectria microconidialis L. Lombard et al., Stud. Mycol. 80:
stricted at the septum, (35–)40–55(–70) × (4–)6–7(–8) μm
175. 2015.
(av. = 53 × 7 μm). Vesicles narrowly clavate, (3–)4(–6) μm diam.
Macroconidia (65–)70–88(–96) × 5–6(–7) μm Diagnosis: Vesicles narrowly clavate, 3–7 μm diam. Macro-
(av. = 81 × 6 μm), (1–)3-septate. conidia (69–)78–98(–113) × 7–9(–10) μm (av. 88 × 8 μm),
4–6(–7)-septate.
Material examined: Ecuador, soil, Jun. 1997, M.J. Wingfield, culture CBS
Description and illustration: Lombard et al. (2015).
111399 = CPC 1620. Typus: China, Guangdong Province, Zhanjiang, CERC nursery,
Calonectria leguminum (Rehm) Crous, Taxonomy and Pa- on E. urophylla × E. grandis clone seedling leaf, Mar. 2009, G.
thology of Cylindrocladium (Calonectria) and Allied Genera (St Zhao (holotype CBS H-21473, cultures ex-type CBS
Paul): 107. 2002. 136638 = CERC 1822 = CMW 31487, CBS 136640 = CERC
Basionym: Nectria leguminum Rehm, Hedwigia 39: 221. 1900. 1827 = CMW 31492).
Synonym: Calonectria quinqueseptata Figueiredo & Namek.,
Calonectria multiseptata Crous & M.J. Wingf., Mycol. Res. 102:
Archos Inst. biol., S. Paulo 34: 93. 1967.
530. 1998.
Diagnosis: Perithecia solitary or in groups, orange to red-brown. Synonym: Cylindrocladium multiseptatum Crous & M.J. Wingf.,
Ascospores fusoid with rounded ends, slightly curved, sigmoid, Mycol. Res. 102: 530. 1998.
(1–)3-septate, not or slightly constricted at the septum, (30–)
50–80(–100) × (4–)5–7(–8) μm (av. = 70 × 6.5 μm). Vesicles Diagnosis: Perithecia solitary or in groups, orange, becoming
narrowly clavate, 2–3 μm diam. Macroconidia (45–) orange-red with age. Ascospores fusoid with rounded ends,
55–65(–70) × (4–)5(–6) μm (av. = 60 × 5 μm), (1–)3–5(–6)- straight to slightly curved, (1–)3–6(–9)-septate, not constricted
septate. at the septa, (45–)65–75(–110) × (5–)6–7(–8) μm
(av. = 70 × 6.5 μm). Megaconidia 6–10-septate, widest in the
middle, bent at right angles like a boomerang or curved, with a
Typus: Brazil, S~ao Paulo, Araruama farm, Jacupiranga, leaf truncate base and rounded apical cell, (120–)
spots on Annona squamosa, 1965, M.B. Figueiredo (holotype of 150–170(–220) × 8–9(–10) μm. Microvesicles clavate. Micro-
Ca. quinqueseptata NY 68-3); ibid., leaves of Eucalyptus sp., conidia 20–65 × 2.5–3.5 μm, 1–3-septate. Macroconidia
Nov. 1979, M.B. Figueiredo & T. Namekata (paratype NY 68-4). unknown.
Calonectria macroconidialis (Crous et al.) Crous, Canad. J.
Description and illustration: Crous et al. (1998).
Bot. 77: 1818. 1999.
Synonyms: Calonectria colhounii Peerally var. macroconidialis Typus: Indonesia, Northern Sumatra, leaves and stems of
Crous et al. Mycotaxon 46: 222. 1993. E. grandis, Jan. 1997, M.J. Wingfield (holotype of Ca. multi-
Cylindrocladium colhounii Peerally var. macroconidiale “macro- septata PREM 55343, holotype of Cy. multiseptatum PREM
conidialis” Crous et al., Mycotaxon 46: 222. 1993. 55344, culture ex-type CBS 112682 = CPC 1589).
Cylindrocladium macroconidiale (Crous et al.) Crous, Canad. J.
Calonectria ovata D. Victor & Crous, Syst. Appl. Microbiol. 20:
Bot. 77: 1818. 1999.
282. 1997. Fig. 17.
Diagnosis: Perithecia solitary or in groups, dull yellow. Asco- Synonym: Cylindrocladium ovatum El-Gholl et al., Canad. J. Bot.
spores straight to slightly curved, (1–)3-septate, not or slightly 71: 469. 1993.
constricted at the septa, (30–)50–65(–75) × (4–)5–7(–8) μm
Diagnosis: Perithecia solitary or in groups, orange, turning red-
(av. = 55 × 6 μm). Vesicles clavate, 3–5 μm diam. Macroconidia
brown with age. Ascospores fusoid with rounded ends, slightly
(68–)85–95(–112) × (5–)6–7(–8) μm (av. = 90 × 6.5 μm), (1–)
curved, sigmoid, 1–3(–7)-septate, not constricted at the septum,
3(–4)-septate.
(35–)55–70(–90) × (4–)5–6 μm (av. = 60 × 5.5 μm). Vesicles
Descriptions and illustrations: Crous et al. (1993b), Crous et al. ovate, 8–14 μm diam. Macroconidia straight or curved, (50–)
(1999). 65–80(–110) × 4–5(–6) μm (av. = 70 × 5 μm), 1(–3)-septate
(up to 7-septate have been observed in old cultures).
Typus: South Africa, Mpumalanga, Sabie, Frankfort nursery,
Eucalyptus cuttings, 16 Apr. 1991, P.W. Crous (holotype of Cy. Description and illustration: Victor et al. (1997).
154
Typus: Brazil, Para, Monte Dourado, E. urophylla, Mar. 1990, culture ex-type CBS 134824 = LPF367); ibid., culture CBS
N.E. El-Gholl (holotype of Cy. ovatum FLAS F55638, cultures 134822 = LPF365.
ex-type CBS 111299 = CPC 1437 = UFV 91, CBS 111307 = CPC
Calonectria pseudocolhounii S.F. Chen et al., Persoonia 26: 7.
1436 = UFV 90); Amazonas, on Eucalyptus sp., A.C. Alfenas,
2011.
UFV 55 (culture of asexual morph), heterothallic mating with
UNB 1026, Amazonas, on Eucalyptus sp., J.C. Dianese (culture Diagnosis: Perithecia solitary or in groups of up to four, bright
of asexual morph), 1996, P.W. Crous (holotype of Ca. ovata yellow, becoming orange with age. Ascospores fusoid with
PREM 51726). rounded ends, straight to slightly curved, (1–)3-septate, not or
slightly constricted at the septum, (44–)50–62(–74) × (5–)
Calonectria pauciramosa C.L. Schoch & Crous, Mycologia 91:
6–7(–8) μm (av. = 56 × 6.5 μm). Vesicles clavate, (3.5–)
289. 1999.
4–5(–6) μm diam. Macroconidia (49–)55–65(–74) × (3.5–)
Synonym: Cylindrocladium pauciramosum C.L. Schoch & Crous,
4–5(–5.5) μm (av. = 60 × 4.5 μm), (1–)3-septate.
Mycologia 91: 289. 1999.
Description and illustration: Chen et al. (2011).
Diagnosis: Perithecia solitary or in groups, orange to red-brown.
Ascospores fusoid with rounded ends, slightly curved, 1-septate, Typus: China, FuJian Province, on leaves of E. dunnii, Aug.
not or slightly constricted at the septum, (30–) 2007, M.J. Wingfield (holotype PREM 60456, culture ex-type
33–38(–40) × 6–7(–8) μm (av. = 35 × 6.5 μm). Vesicles CBS 127195 = CMW 27209).
obpyriform to ellipsoidal, (5–)7–9(–11) μm diam. Macroconidia
Calonectria pseudometrosideri R.F. Alfenas et al., Stud.
cylindrical, rounded at both ends, straight, (30–)
Mycol. 80: 118. 2015.
45–55(–60) × (3.5–)4–5 μm (av. = 50 × 4.5 μm), 1-septate.
Diagnosis: Vesicles ellipsoidal to obpyriform, 5–7 μm diam.
Description and illustration: Schoch et al. (1999).
Macroconidia straight to slightly curved, (40–)
Typus: Brazil × South Africa. Brazil, Bahia, Nursery, on Euca- 49–52(–60) × (3–)4.5(–5) μm (av. = 51 × 4.5 μm), 1-septate.
lyptus sp., Jul. 1990, A.C. Alfenas, South Africa, Western Cape,
Knysna, soil, Nov. 1994, P.W. Crous, heterothallic mating of STE-
U 1670 × STE-U 971 (holotype of Cy. pauciramosum PREM Typus: Brazil, Alagoas state, Maceio, from soil collected in
55752), Apr. 1997, C.L. Schoch (holotype of Ca. pauciramosa Eucalyptus plantation, Apr. 2011, M.M. Coutinho (holotype CBS
PREM 55754). 134845, preserved as metabolically inactive culture, culture ex-
type CBS 134845 = LPF210).
Calonectria pentaseptata L. Lombard et al., Persoonia 29: 157.
2012. Calonectria pseudoreteaudii L. Lombard et al., Persoonia 24:
8. 2010.
Diagnosis: Vesicles narrowly clavate, 2–6 μm diam.
Macroconidia (75–)87–109(–115) × (5–)6–8(–10) μm Diagnosis: Vesicles narrowly clavate, 3–5 μm diam. Macro-
(av. = 98 × 7 μm), 5(–8)-septate. conidia (88–)96–112(–119) × 7–9(–10) μm (av. = 104 × 8 μm),
5–8-septate.
Description and illustration: Lombard et al. (2010c).
Typus: Vietnam, Bavi, Hanoi, on Eucalyptus hybrid, Sep. 2011,
P.Q. Thu (holotype CBS H-21062, culture ex-type CBS Typus: China, Guangdong Province, CERC nursery, on stems of
133349 = BD 32-39). E. urophylla × E. grandis hybrid cutting, Nov. 2007, M.J. Wing-
field & X.D. Zhou (holotype PREM 60290, culture ex-type CBS
Calonectria propaginicola R.F. Alfenas et al., Stud. Mycol. 80:
123694 = CMW 25310).
115. 2015.
Calonectria pseudoscoparia L. Lombard et al., Stud. Mycol.
Diagnosis: Vesicles ellipsoidal, obpyriform to sphaer-
66: 53. 2010.
opedunculate, 5– 12 μm diam. Macroconidia straight to slightly
curved, (40–)48–51(–55) × 3–5 μm (av. = 49 × 4 μm), 1- Diagnosis: Vesicles obpyriform to ellipsoidal, 6–10 μm diam.
septate. Macroconidia cylindrical, rounded at both ends, straight, (41–)
45–51(–52) × 3–5 μm (av. = 48 × 4 μm), 1-septate.
Description and illustration: Lombard et al. (2010b).
Typus: Brazil, Para state, Santana, from Eucalyptus seedling,
Apr. 2011, A.C. Alfenas (holotype CBS H-21366, cultures ex- Typus: Ecuador, Pichincha Province, Las Golondrinas, Buenos
type CBS 134815 = LPF220); ibid., culture CBS Aires Nursery, from E. grandis cutting, Dec. 2004, M.J. Wingfield
134816 = LPF222. (holotype PREM 60305, culture ex-type CBS 125257 = CMW
15218).
Calonectria pseudocerciana R.F. Alfenas et al., Stud. Mycol.
80: 117. 2015. Calonectria pteridis Crous et al., Mycotaxon 46: 228. 1993.
Fig. 17.
Diagnosis: Vesicles obpyriform to sphaeropedunculate, 7–12 μm
Synonym: Cylindrocladium pteridis F.A. Wolf, J. Elisha Mitchell
diam. Macroconidia straight to slightly curved, (35–)
scient. Soc. 42: 59. 1926.
43–46(–55) × 3–5 μm (av. = 45 × 4 μm), 1-septate.
Diagnosis: Perithecia solitary or in groups, red-brown. Asco-
spores fusoid with rounded ends, slightly curved, sigmoid, 1(–3)-
Typus: Brazil, Para state, Santana, from stem of Eucalyptus septate, not constricted at the septum, (30–)45–60(–75) × (4–)
seedling, Apr. 2011, A.C. Alfenas (holotype CBS H-21366, 5–6(–7) μm (av. = 52 × 6 μm). Vesicles clavate to narrowly
CROUS ET AL.
ellipsoidal, 4–5(–6) μm diam. Macroconidia cylindrical, rounded Calonectria seminaria L. Lombard et al., Stud. Mycol. 80: 179.
at both ends, straight or curved, (50–)70–100(–130) × (4–) 2015.
5–6 μm (av. = 82 × 5.5 μm), 1(–3)-septate.
Diagnosis: Vesicles obpyriform to ellipsoid, 6–11 μm diam.
Description and illustration: Crous et al. (1993b). Macroconidia cylindrical, rounded at both ends, straight, (42–)
45–49(–52) × 3.5–4.5(–7) μm (av. 47 × 5 μm), 1-septate.
Typus: USA × Brazil. USA, Arachniodes adiantiformis, collection
date unknown, F. Schickedanz, ATCC 34395 (culture of asexual Description and illustration: Lombard et al. (2015).
morph), Brazil, E. grandis, collection date unknown, A.C. Alfe-
Typus: China, Guangdong Province, Zhanjiang, CERC nursery,
nas, PPRI 4180 (culture of asexual morph), 1992, P.W. Crous
on E. urophylla × E. grandis clone seedling leaf, Mar. 2009, G.
(holotype of Ca. pteridis PREM 51033).
Zhao (holotype CBS H-21475, culture ex-type CBS
Calonectria queenslandica L. Lombard et al., Persoonia 24: 8. 136632 = CERC 1785 = CMW 31450 = CPC 23488).
2010.
Calonectria spathulata El-Gholl et al., Mycotaxon 26: 159.
Diagnosis: Vesicles narrowly clavate, 3–4 μm diam. Macro- 1986. Fig. 17.
conidia (61–)65–73(–78) × (4–)5–6(–7) μm (av. = 69 × 6 μm), Synonym: Cylindrocladium spathulatum El-Gholl et al., Myco-
4–6-septate. taxon 26: 159. 1986.
Description and illustration: Lombard et al. (2010c).
Diagnosis: Perithecia solitary or in groups, orange, becoming
Typus: Australia, Queensland, Lannercost, on leaves of orange-red to red-brown with age. Ascospores fusoid with
E. urophylla, 15 Apr. 1991, B. Brown (holotype PREM 60243, rounded ends, straight to slightly curved, (1–)3-septate, not or
culture ex-type CBS 112146 = CMW 30604 = CPC slightly constricted at the septa, (38–)45–55(–60) × (4.5–)
3213 = DFRI00147). 5–6(–7) μm (av. = 50 × 5.5 μm). Vesicles ellipsoid to obpyriform
or clavate, 6–7(–10) μm diam. Macroconidia (48–)
Calonectria reteaudii (Bugn.) C. Booth, Mycol. Pap. 104: 41.
75–90(–100) × (4–)5–6 μm (av. = 80 × 6 μm), (1–)3(–6)-
1966. Fig. 17.
septate.
Basionym: Neonectria reteaudii Bugn. as “reteaudi”, Encycl.
Mycol. 11: 189. 1939.
Descriptions and illustrations: El-Gholl et al. (1986), Crous
Synonyms: Cylindrocladium reteaudii (Bugn.) Boesew., Trans.
(2002).
Brit. Mycol. Soc. 78: 554. 1982.
Cylindrocarpon reteaudii Bugn., Encycl. Mycol. 11: 189. 1939. Typus: Brazil, Santa Catarina, on living leaves of E. viminalis,
Cylindrocladium quinqueseptatum Boedijn & Reitsma, Rein- 1983, N.E. El-Gholl (holotypes of Ca. spathulata and Cy. spa-
wardtia 1: 59. 1950. thulatum FTCC 1001 and FLAS F54257a & b, respectively).
Diagnosis: Perithecia solitary or in groups, orange to red-brown. Calonectria sulawesiensis L. Lombard et al., Stud. Mycol. 66:
Ascospores fusoid with rounded ends, slightly curved, sigmoid, 53. 2010.
(1–)3(–5)-septate, not or slightly constricted at the septum,
Diagnosis: Vesicles broadly clavate to ellipsoidal, 5–7 μm diam.
(50–)65–85(–100) × (4–)5–6(–7) μm (av. = 70 × 5.5 μm).
Macroconidia cylindrical, rounded at both ends, straight, (41–)
Vesicles clavate, (3–)5(–6) μm diam. Macroconidia (50–)
45–51(–54) × (3–)4(–6) μm (av. = 48 × 4 μm), 1-septate.
75–95(–120) × (5–)6–7 μm (av. = 84 × 6.5 μm), (1–)5(–6)-
septate. Description and illustration: Lombard et al. (2010b).
Descriptions and illustrations: Crous (2002), Lombard et al. Typus: Indonesia, Sulawesi, on leaf of Eucalyptus sp., Jul. 2003,
(2010b). M.J. Wingfield (holotype PREM 60300, culture ex-type CBS
125277 = CMW 14878).
Typus: Vietnam, South Annam, leaves of Smithia bequaertii,
unknown collection date, F. Bugnicourt (holotype IMI 55922, Calonectria terrae-reginae L. Lombard et al., Persoonia 24: 9.
dried ex-type culture of Ca. reteaudii, depauperate); Binh Phuoc 2010.
Province, Chon Thanh, living leaves of E. camaldulensis, Oct.
Diagnosis: Vesicles narrowly clavate, 3–5 μm diam. Macro-
1998, M.J. Dudzinski & P.Q. Thu (neotypes of respectively Ca.
conidia 60–83(–87) × (4–)5–7(–8) μm (av. = 76 × 6 μm), 4–6-
reteaudii (PREM 57211) and Cy. reteaudii (PREM 57212), cul-
septate.
tures ex-type CBS 112144 = CMW 30984 = CPC 3201 (+) and
CPC 3204 (-)). Description and illustration: Lombard et al. (2010c).
Calonectria robigophila R.F. Alfenas et al., Stud. Mycol. 80: Typus: Australia, Queensland, Cardwell, Meunga, on leaves of
123. 2015. E. urophylla, 11 Apr. 1997, C. Hanwood (holotype PREM 60239,
culture ex-type CBS 112151 = CMW 30601 = CPC
Diagnosis: Vesicles acicular to clavate, 4– 5 μm diam. Macro-
3202 = DFRI00150).
conidia (45–)49–52(–60) × 3–5 μm (av. = 50 × 4 μm), 1-
septate. Calonectria terrestris L. Lombard et al., Stud. Mycol. 80: 182.
2015.
Diagnosis: Vesicles obpyriform to pyriform to broadly clavate,
Typus: Brazil, Maranh~ao state, Açailândia, on leaves of Euca-
5–12 μm diam. Macroconidia (33–)36–40(–41) × (3–)4–5 μm
lyptus sp., May 2011, R.F. Alfenas (holotype CBS H-21361,
(av. 38.5 × 4.5 μm), 1-septate.
culture ex-type CBS 134652 = LPF192); ibid., cultures LPF190,
CBS 134653 = LPF193. Description and illustration: Lombard et al. (2015).
156
Typus: China, Guangdong Province, from soil collected in a cells lageniform to cylindrical, sympodial, small denticles or
Eucalyptus plantation, Aug. 2009, X. Mou & R. Chang (holotype scars, terminal and lateral, subhyaline. Conidia falcate, cylindri-
CBS H-21478, cultures ex-type CBS 136642 = CERC cal or fusiform, 0–1-sepate, hyaline, smooth-walled. Chla-
1856 = CMW 35180, CBS 136643 = CERC 1868 = CMW mydospores not observed. Sexual morph unknown.
35364 = CPC 23493, CBS 136644 = CERC 1870 = CMW
Type species: Castanediella acaciae Crous et al.
35366 = CPC 23494, CBS 136645 = CERC 1854 = CMW
35178 = CPC 23496, CBS 136651 = CERC 1937 = CMW Symptoms: Leaf spots amphigenous when present, irregular,
37974 = CPC 23516). pale brown. The genus is endophytic, and symptoms are mostly
expressed on older senescent leaves.
Calonectria tetraramosa L. Lombard et al., Stud. Mycol. 80:
183. 2015.
Notes: Species of Castanediella have been isolated from
Diagnosis: Vesicles obpyriform, 4–10 μm diam. Macroconidia symptomatic Eucalyptus leaves but little is known about their role
(45–)46.5–49.5(–51) × (4–)4.5–5.5(–6) μm (av. 48 × 5 μm), 1- as pathogens. At present they are considered to be of minor
septate. importance.
Description and illustration: Lombard et al. (2015). Castanediella communis Crous & M.J. Wingf., Persoonia 37:
297. 2016. Fig. 18.
Typus: China, Guangdong Province, Zhanjiang, CERC nursery,
on E. urophylla × E. grandis clone seedling leaf, Mar. 2009, G. Diagnosis: Conidia solitary, hyaline, smooth, falcate with sub-
Zhao (holotype CBS H-21477, living culture ex-type CBS obtuse ends, guttulate, (13–)17–20(–22) × (2–)2.5(–3) μm.
136635 = CERC 1809 = CMW 31474 = CPC 23489).
Description and illustration: Crous et al. (2016b).
Calonectria variabilis Crous et al., Syst. Appl. Microbiol. 16:
272. 1993.
Synonym: Cylindrocladium variabile Crous et al., Syst. Appl.
27631).
Microbiol. 16: 272. 1993.
Castanediella eucalypti Crous et al., Persoonia 34: 189. 2015.
Diagnosis: Perithecia solitary or in groups of 2–3, red-brown.
Ascospores fusoid with rounded ends, straight to slightly Diagnosis: Conidia solitary, hyaline, smooth, falcate, slightly
curved, 1(–3)-septate, not or slightly constricted at the septa, curved, widest in middle with subobtuse ends, (15–)
(34–)38–50(–60) × 4–5(–6) μm (av. = 42 × 5 μm). Vesicles 18–21(–23) × 2–3 μm.
sphaeropedunculate to ovoid or ellipsoid to clavate, 6–11 μm
diam. Macroconidia straight or curved, (48–)
68–77(–85) × 4–5(–6) μm (av. = 73 × 5 μm), (1–)3(–4)- Typus: Malaysia, Sabah, on E. pellita, May 2014, M.J. Wingfield
septate, widest in middle of the lower conidium cell. (holotype CBS H-22226, cultures ex-type CBS 139897 = CPC
24746, CPC 24747).
Castanediella eucalypticola Crous & M.J. Wingf., Persoonia
Typus: Brazil, Para, near Belem, on Schefflera (= Didymopanax)
36: 351. 2016.
morototoni, 1990, F.C. de Albuquerque (holotype of Ca. varia-
bilis PREM 51039, holotype of Cy. variabile PREM 51041, Diagnosis: Conidia solitary, hyaline, smooth, falcate, straight to
culture ex-type of Cy. variabile AR 2675 = CBS 339.92 = CPC curved, widest in the middle, apex subobtusely rounded, base
2436 = IFO 32466 = IMI 354513 = PPRI 4174). truncate, 0.5 μm diam, (15–)20–26(–30) × (2.5–)3 μm.
Calonectria zuluensis L. Lombard et al., Stud. Mycol. 66: 25. Description and illustration: Crous et al. (2016c).
2010.
Typus: France, La Reunion, on E. robusta, 9 Mar. 2015, P.W.
Diagnosis: Perithecia solitary or in groups, orange to red, Crous & M.J. Wingfield (holotype CBS H-22604, culture ex-type
becoming red-brown with age. Ascospores fusoid with rounded CBS 141317 = CPC 26539).
ends, straight to slightly curved, 1-septate, not or slightly con-
Castanediella malaysiana Hern.-Restr. et al., Sydowia 68: 207.
stricted at the septum, (26–)29–34(–38) × 4–5 μm
2016.
(av. = 32 × 4 μm). Vesicles ellipsoid to obpyriform vesicles,
6–10 μm diam. Macroconidia cylindrical, rounded at both ends, Diagnosis: Conidia fusoid, curved, hyaline, smooth, 0–1-septate,
straight, (31–)34–38(–40) × 3–5 μm (av. = 36 × 4 μm), 1- apex acuminate, and base acuminate or slightly flattened,
septate. 18–30 × 2–3 μm.
Description and illustration: Lombard et al. (2010a). Description and illustration: Hernandez-Restrepo et al. (2016).
Typus: South Africa, KwaZulu-Natal, Kwambonambi, on Typus: Malaysia: Sabah, on E. brassiana, May 2014, M.J.
E. grandis clonal cutting, Feb. 2001, L. Lombard (holotype Wingfield (holotype CBS H-22659, culture ex-type CBS
PREM 60292, culture ex-type CBS 125268 = CMW 9188). 141509 = CPC 24918).
Castanediella M. Hern.-Restr. et al., Persoonia 36: 187. 2015. Cercospora Fresen., Beitr. Mykol. 3: 91. 1863.
Mycelium immersed and superficial, hyphae branched, septate, Having mycosphaerella-like sexual morphs. Mostly plant patho-
hyaline and brown, smooth-walled. Conidiomata if present genic, symptomless or usually causing distinct lesions. Mycelium
sporodochium- like. Conidiophores branched, pale brown to internal, rarely external; hyphae branched, septate, hyaline or
brown at the base and subhyaline at the apex. Conidiogenous almost so to usually pigmented, thin-walled, smooth, rarely faintly
CROUS ET AL.
Fig. 18. Castanediella communis (CPC 27631). A–C. Symptomatic leaves from which the fungus was isolated. D. Colony sporulating on SNA. E–G. Conidiophores with
conidiogenous cells. H. Conidia. Scale bars = 10 μm.
rough-walled. Stromata lacking to well-developed, substomatal, could be a secondary pathogen, as it was found to co-occur on
intra-epidermal to deeply immersed, mostly pigmented, lesions with Pseudocercospora paraguayensis. Conidia solitary,
composed of textura angulata or globosa. Conidiophores mon- hyaline, thick-walled, smooth, acicular, apex obtuse, base sub-
onematous, macronematous, solitary or fasciculate, in small to truncate to rounded, sometimes with a lateral branch near the
large fascicles, rarely in sporodochial conidiomata, emerging base, 3–7-septate, (30–)40–60(–80) × (2–)3–3.5 μm.
through stomata or erumpent, very rarely arising from superficial
hyphae, erect, continuous to multi-septate, hyaline or almost so
(subgen. Hyalocercospora) to mostly pigmented, pale olivaceous
to dark brown (subgen. Cercospora), wall smooth to somewhat
rough, thin to moderately thick; conidiogenous cells integrated,
terminal or intercalary, sometimes conidiophores aseptate, i.e.
reduced to conidiogenous cells, monoblastic, determinate to
usually polyblastic, proliferation sympodial, rarely percurrent,
conidiogenous loci (scars) conspicuous, thickened and
darkened-refractive, planate with minute central pore. Conidia
solitary, rarely in short chains (mainly under high humidity),
mostly scolecosporous, obclavate-cylindrical, acicular, filiform
and multi-euseptate, rarely amero to phragmosporous, broadly
ellipsoid-ovoid to broadly obclavate-cylindrical, but always hya-
line or subhyaline (with a pale greenish tinge), thin-walled,
smooth or almost so, hila thickened and darkened, conidial
secession schizolytic (Braun et al. 2013).
Type species: Cercospora apii Fresen.
Notes: Cercospora eucalyptorum appears to be a minor or
opportunistic foliar pathogen of Eucalyptus. The genus Pseu-
docercospora is much more important and prevalent on this host
than Cercospora.
Cercospora eucalyptorum Crous, Mycol. Mem. 21: 33. 1998.
Fig. 19.
Diagnosis: Cercospora spp. are associated with brown, sub-
Fig. 19. Cercospora eucalyptorum (PREM 55301). Conidiophores and conidia.
circular to angular leaf spots. In the case of C. eucalyptorum, it
Scale bars = 10 μm. (from Crous 1998).
158
Description and illustration: Crous (1998). guttulate, ellipsoid, straight to allantoid, apex obtuse, base with
flattened hilum, 0.5 μm diam, (4 –)6–7.5( –10) × (2–)
Typus: Paraguay, Pirapo, Itapua, on Eucalyptus sp., 4 Dec.
2.5( –3) μm.
1983, T. Kobayashi, (holotype PREM 55301, derived from ho-
lotype of Pseudocercospora paraguayensis, TFM: FPH 5626b). Description and illustration: Crous et al. (2015b).
Not known from culture.
Typus: Colombia, Suiza, on E. urophylla × E. grandis, Jul. 2014,
Chrysofolia Crous & M.J. Wingf., Persoonia 34: 207. 2015. M.J. Wingfield (holotype CBS H-22238, cultures ex-type CBS
139909 = CPC 24986, CPC 24987).
Conidiomata pycnidial, separate to aggregated, exuding a yellow
slimy conidial mass; pycnidia globose, base immersed, green- Clypeophysalospora H.J. Swart, Trans. Brit. Mycol. Soc. 76:
brown in lactic acid, but bright yellow in Shear's; wall of 3–6 93. 1981.
layers of brown textura angularis; neck long, with a pale yellow
Ascomata perithecial, scattered, immersed, single or rarely
zone just above the agar surface, which then turns yellow-brown
in pairs, covered by a dark clypeus, with a wall consisting
while the apical part is pale yellow to almost subhyaline, termi-
of darkened, flattened, thin-walled cells and with a narrow
nating in an obtusely rounded apex with central ostiole surrounded
periphysate ostiole. Asci paraphysate, cylindrical or slightly
by loose hyphal elements. Conidiophores reduced to con-
swollen, unitunicate, with an amyloid apical ring and a pulvi-
idiogenous cells. Conidiogenous cells lining the inner cavity of
llus. Ascospores mostly uniseriate or irregularly arranged,
base, hyaline, smooth, ampulliform, tapering abruptly towards
hyaline, ellipsoid, with mucous outer wall layer. Asexual morph
neck, with several apical percurrent proliferations. Paraphyses
unknown.
interspersed among conidiogenous cells, hyaline, smooth, cylin-
drical, septate. Conidia solitary, hyaline, smooth, guttulate, ellip- Type species: Clypeophysalospora latitans (Sacc.) H.J. Swart
soid, straight to allantoid, apex obtuse, base with flattened hilum.
Notes: Clypeophysalospora latitans is a common fungus on
Type species: Chrysofolia colombiana Crous et al. Eucalyptus leaves. It is usually found sporulating on leaf litter, but
is also commonly associated with leaf spots, and assumed to be
Note: Since its original description based on infections in Colombia,
a minor pathogen. It appears more prominent on certain eucalypt
this pathogen has also been detected on leaves of Syzygium
species than others.
jambos in La Reunion (Hernandez-Restrepo et al. 2016).
Clypeophysalospora latitans (Sacc.) H.J. Swart, Trans. Brit.
Chrysofolia colombiana Crous et al., Persoonia 34: 207. 2015.
Mycol. Soc. 76: 95. 1981. Fig. 21.
Fig. 20.
Basionym: Physalospora latitans Sacc. Flora. Myc. Lusitan. 67.
Diagnosis: Associated with medium brown leaf spots; easily 1893.
recognisable by having erumpent conidiomata with bright Synonym: Amerostege latitans (Sacc.) Theiss., Ann. Mycol. 14:
yellow-brown necks. Conidia solitary, hyaline, smooth, 411. 1917.
Fig. 20. Chrysofolia colombiana (CPC 24986). A. Colony sporulating on OA. B–E. Conidiomata with long necks with aggregated hyphal elements. F. Conidiogenous cells. G.
Conidia. Scale bars = 10 μm.
CROUS ET AL.
Fig. 21. Clypeophysalospora latitans (CPC 14930). A. Disease symptoms. B, C. Typical erumpent, black ascomata. D–F. Vertical sections through ascomata. G–J. Asci with
ascospores. K. Ascoma on PNA. L. Ascoma in culture. M. Hamathecial tissue. Scale bars: D–F = 350 μm, all others = 10 μm.
Diagnosis: Leaf spots amphigenous, pale brown, large, sub- 6–9 μm, in vitro (11–)13–19(–20) × 5–7 μm, hyaline, sur-
circular, lacking prominent margins. The genus is clearly endo- rounded by a thin mucilaginous sheath.
phytic, but also pathogenic. Clypeus composed of dark brown
Description and illustration: Giraldo et al. (2017).
compressed host cells and fungal hyphae. Paraphyses up to
6.4 μm diam, hypha-like, flexuose, septate, numerous, hyaline Typus: Portugal, Algarve region, Faro, on Eucalyptus sp., 24
and embedded in a gelatinous matrix. Asci 8-spored, cylindrical, Jan. 2008, P.W. Crous (epitype of Clypeophysalospora latitans
unitunicate, thin-walled, with a J+ and an amyloid apical CBS H-20289, cultures ex-epitype CBS 141463 = CPC 14930,
ring, in vivo 115–141 × 11–16 μm, in vitro (107–) CPC 14931, 14932); Beira Litoral, Coímbra, (lectotype of
108–117(–118) × 9–10 μm. Ascospores uniseriate, unicellular, Physalospora latitans PAD 2747).
ellipsoidal, slightly curved, smooth-walled, in vivo 13–22 ×
Clypeosphaerella Guatimosim et al., Persoonia 37: 121. 2016.
160
Ascomata pseudothecial, epiphyllous, solitary, subcuticular to Synonyms: Coccomycella Höhn., Ann. Mycol. 15(5): 323. 1917.
erumpent, globose, walls of 2–3 layers of brown to dark brown Malenconia Bat. & H. Maia, Saccardoa 1: 98. 1960.
textura angularis, ostiole central. Asci bitunicate, aparaphysate,
Ascomata immersed in host epidermal tissue; hymenium
fasciculate, subsessile, 8-spored, obpyriform to ovoid, hyaline,
developing within pseudoparenchymatous stromatic tissue,
smooth. Ascospores inordinate, overlapping, fusoid, straight, 1-
becoming exposed after covering stromatic tissue develops
septate, slightly constricted at the septum, biguttulate, hyaline,
several radiate slits; paraphyses swollen at apices; ascus apex
thin-walled, smooth. Ascospores germinating at both ends,
non-amyloid; ascospores typically filiform, 0–1-septate, with
remaining hyaline, germ tubes following the main axis of the
gelatinous sheath (from Johnston 1986).
spore (from Guatimosim et al. 2016).
Type species: Clypeosphaerella quasiparkii (Cheew. et al.) Type species: Coccomyces coronatus (Schumach.) De Not.
Guatimosim et al. Notes: Johnston (1986) originally described C. globosus from
Notes: Causing leaf spots of Eucalyptus in Thailand. Very little is leaves of Nestegis lanceolata in New Zealand, and also recorded it
known about its pathology and ecology. as a wide host range species on Metrosideros fulgens and
Weinmannia racemosa. Dick (1990) reported Coccomyces glo-
Clypeosphaerella quasiparkii (Cheew. et al.) Guatimosim bosus from four species of Eucalyptus in New Zealand, while
et al., Persoonia 37: 121. 2016. Fig. 22. Johnston (2000) listed it on numerous species of Eucalyptus. The
Basionym: Mycosphaerella quasiparkii Cheew. et al., Persoonia ability of C. globosus to occur on a wide host range is supported
21: 85. 2008. by its DNA phylogeny (P. Johnston, unpubl. data). Johnston (2000)
Diagnosis: Leaf spots amphigenous, round to irregular, separate, also noted that the first record of Rhytismatales for Australia,
becoming confluent, medium brown on adaxial surface, pale namely Coccomyces delta occurring on Eucalyptus (Cooke 1892),
brown on abaxial surface, surrounded by a brown, raised border. could represent an older name for C. globosus. This species is not
Ascospores bi- to multiseriate, overlapping, hyaline, guttulate, a primary pathogen, appears to be an endophyte, and is found
thin-walled, straight to slightly curved, ellipsoidal to obovoid with sporulating on older weakened leaves, or on dead leaf tissue
obtuse ends, widest in the middle of the apical cell, medianly linked to lesions of other primary pathogens.
1-septate, not constricted at the septum, tapering toward Coccomyces globosus P.R. Johnst., New Zealand J. Bot. 24:
both ends, with a thin mucilaginous sheath, (9–)10–11(–12.5) × 99. 1986. Fig. 23.
(2.5–)3–3.5(–4.5) μm.
Leaf spots bleached, subcircular, 2–6 mm diam, with thin dark
Description and illustration: Cheewangkoon et al. (2008). brown zone line. Ascomata immersed, amphigenous, triangular
Typus: Thailand, Burirum, on Eucalyptus sp., Jul. 2007, P. in outline, with dark zone line, opening via three radiate flaps;
Suwannawong (holotype CBS H-20132, cultures ex-type CBS up to 600 μm diam; hymenium yellow, when dry becoming
123243 = CPC 15409); ibid., cultures CPC 15433, 15434. brown with age, with mucoid layer covering asci and paraph-
yses. Paraphyses 1.5–2 μm diam, septate, clavate, 2–3 μm
Coccomyces De Not., G. bot. ital. 2(7–8): 38. 1847. diam at apex, rarely branched, extending 10–25 μm
Fig. 22. Clypeosphaerella quasiparkii (CBS H-20132). A. Leaf spot. B. Section through an ascoma. C. Colony on MEA. D, E. Asci. F. Ascospores. G, H. Germinating as-
cospores. Scale bars: A = 1 mm; B = 20 μm; all others = 10 μm.
CROUS ET AL.
above asci, apex with mucoid sheath. Asci cylindrical, Ceuthosira Petr., Ann. Mycol. 22: 265. 1924.
97–125 × 5–6 μm, apex subobtuse, 8-spored with foot cell. Parafabraea Chen Chen et al., Fungal Biol. 120: 1291. 2016.
Ascospores (50–)55–65( –85) × 1.5(–2) μm, hyaline, smooth,
For synonymy see Nag Raj (1978), Sutton (1980), Duan et al. (2007).
slightly curved, with basal taper, apical mucoid cap only
observed in water mounts. Conidiomata pycnidial, erumpent on Mycelium immersed, consisting of branched, septate, hyaline to
OA, globose, brown, 90–150 μm diam, with central ostiole; wall pale brown hyphae. Ascomata apothecial, sessile to subsessile,
of 2–3 layers of brown textura angularis. Conidiophores lining short-stalked, gregarious or confluent, clustering on a basal
the inner cavity, hyaline, smooth, subcylindrical, branched, stroma, partly immersed, medium to dark brown. Disc turbinate,
1–3-septate, 15–30 × 3–5 μm. Conidiogenous cells terminal pale brown. Seta-like structures surrounding apothecia, rigid,
and intercalary, subcylindrical, 10–18 × 2–3 μm, proliferating pale brown, septate, cylindrical, straight or slightly curved,
inconspicuously percurrently at apex with delayed conidial slightly enlarged at truncate apex. Basal stroma subimmersed,
secession. Conidia solitary, aseptate, hyaline, smooth, sub- composed of irregular, pale to medium brown cells. Asci
cylindrical, straight, guttulate, ends bluntly rounded, (3–) inoperculate, clavate to cylindrical-clavate, apex rounded, short-
4–5( –6) × 1.5( –2) μm. pedicellate, base truncate, hyaline to pale brown, 8-spored.
Ascospores inequilateral, fusoid to ellipsoid, ends rounded,
Culture characteristics: Colonies flat, spreading, with sparse
straight or slightly curved, aseptate, thin-walled, hyaline, guttu-
aerial mycelium and smooth, lobate margins, reaching 30 mm
late. Paraphyses numerous, cylindrical, slender, wider at base,
diam after 2 wk at 25 °C. On MEA surface amber, reverse
septate, apex round, hyaline to pale brown. Conidiomata pyc-
fulvous. On PDA surface pale luteous, reverse amber. On OA
nidial, separate, globose or flattened at the base, black,
surface luteous.
immersed, unilocular; wall composed of brown, thick-walled
Typus: New Zealand, Auckland, Waitakere Ranges, Walker textura angularis; lower wall often thicker than upper region.
Bush Track, on fallen leaves of Nestegis lanceolata, 4 Aug. 1982, Ostiole single, circular, not papillate. Paraphyses hyaline, septate
P.R. Johnston, G.J. Samuels & E.H.C. McKenzie (holotype PDD at the base, intermingled among conidiophores, cylindrical or
44643). long clavate, collapsing at maturity. Conidiophores pale brown at
Material examined: Australia, Victoria, Tantawanglo State Forest, Tin Mine Road,
the base, hyaline above, branched, septate, smooth, thin-walled,
on Eucalyptus leaf litter, 28 Nov. 2016, P.W. Crous, HPC 1824 (CBS H-24022, formed from the inner cells of the pycnidial wall, sometimes
culture CBS 143429 = CPC 32711). confined to the base, or in short chains. Conidiogenous cells
phialidic, determinate, discrete and ampulliform to lageniform, or
Coleophoma Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Kl.,
integrated and subcylindrical, hyaline, smooth, with prominent
Abt. 1, 116: 637. 1907. Fig. 24.
periclinal thickening and collarette minute. Conidia hyaline,
Synonyms: Basilocula Bubak, Ann. Mycol. 12: 210. 1914.
aseptate, cylindrical, apex obtuse, base acute, thin-walled,
Rhabdostromellina Höhn., Ann. Mycol. 15: 303. 1917.
smooth, guttulate, straight (adapted from Sutton 1980, and
Bactropycnis Höhn., Hedwigia 62: 65. 1920.
Cheewangkoon et al. 2009).
Rhabdostromina Died., Ann. Mycol. 19: 297. 1921.
Xenodomus Petr., Ann. Mycol. 20: 206. 1922. Type species: Coleophoma crateriformis (Durieu & Mont.) Höhn.
Fig. 23. Coccomyces globosus (CPC 32711). A. Ascomata with radiate flaps. B–D. Asci and paraphyses. E. Ascospore. F. Conidiomata in culture. G, H. Conidiogenous cells.
I. Conidia. Scale bars = 10 μm.
162
Fig. 24. Coleophoma spp. A, B. C. eucalypti (VPRI 20394). A. Disease symptoms. B. Conidiomata on leaf spot. F. Conidia. C–E. C. eucalyptorum (CBS H-20770). Con-
idiogenous cells and paraphyses (typical for genus). Scale bars = 10 μm.
Symptoms: Associated with amphigenous brown leaf spots that Typus: Australia, Yapilika, Melville Island, Northern Territory, on
can vary from small spots to large botches, without prominent E. pellita, 18 May 1994, Z.Q Yuan (holotype VPRI 20394). Not
margins. known from culture.
Notes: Coleophoma eucalyptorum was found to co-occur with Coleophoma eucalypticola Crous, Fungal Biol. 120: 1404.
C. coptospermatis on the same leaf spots on Coptosperma lit- 2016.
torale in South Africa, suggesting that it has a wider host range Basionym: Neofabraea eucalypti Cheew. & Crous, Persoonia 23:
wider than only Eucalyptus (Crous & Groenewald 2016). 67. 2009.
Coleophoma represents a genus of foliar pathogens that can Synonym: Parafabraea eucalypti (Cheew. & Crous) Chen Chen
cause prominent leaf spots on a range of hosts, including et al., Fungal Biol. 120: 1318. 2015 (2016).
Eucalyptus. It is however not regarded as an important pathogen Diagnosis: Asci clavate to cylindrical-clavate, apex rounded,
on this host. short pedicellate, base truncate, hyaline to very pale brown, 8-
Coleophoma caliginosa (Cheew. et al.) Crous, Fungal Biol. spored, ascospores discharging through apical pore, (35–)
120: 1401. 2016. 40–45(–52) × 10–12 μm. Ascospores fusoid to ellipsoid,
Basionym: Cryptosporiopsis caliginosa Cheew. et al., Fungal aseptate, hyaline, ends rounded, unequal, straight or slightly
Diversity 44: 93. 2010. curved, thin-walled, guttulate, 10–14 × 4–6 μm.
Synonym: Parafabraea caliginosa (Cheew. et al.) Chen Chen Description and illustration: Cheewangkoon et al. (2009).
et al., Fungal Biol. 120: 1318. 2015 (2016).
Typus: Australia, Victoria, Otway, on E. globulus, 15 Feb.
Diagnosis: Leaf spots amphigenous, subcircular to irregular, 2007, coll. I. Smith, isol. P.W. Crous (holotype CBS H-20285,
medium brown. Conidia elongate ellipsoidal, mostly straight, cultures ex-type CBS 124810 = CPC 13755, CPC 13756,
broadly obtuse at the apex, tapering abruptly to a slightly pro- 13757).
truding basal scar, aseptate, hyaline, thick-walled, minutely
guttulate, (8.5–)15–17(–19) × (3.5–)4.5–5.5 μm. Coleophoma eucalyptorum Crous & Summerell, Persoonia 27:
137. 2011. Fig. 24.
Description and illustration: Cheewangkoon et al. (2010).
Diagnosis: Conidia hyaline, smooth, guttulate, cylindrical, apex
Typus: Australia, New South Wales, Northern Tablelands, Mt obtuse, base with flattened, truncate locus, (10–)
Mackenzie Nature Reserve (290504S; 1515805E), on 11–12(–14) × (2–)2.5 μm.
E. caliginosa, 1 Feb. 2007, B.A. Summerell (holotype CBS H-
20301, cultures ex-type CBS 124806 = CPC 14048, CPC 14049, Description and illustration: Crous et al. (2011b).
14050). Typus: Australia, New South Wales, Blue Mountains, Kurrajong
Heights, on E. piperita, 16 Nov. 2010, B.A. Summerell (holotype
Coleophoma eucalypti Z.Q. Yuan, Aust. Syst. Bot. 9: 342.
CBS H-20770, culture ex-type CBS 131314 = CPC 19299).
1996. Fig. 24.
Colletotrichum Corda, in Sturm, Deutschl. Fl., 3 Abt. (Pilze
Diagnosis: Conidia cylindrical, apex obtuse, base with flattened,
Deutschl.) 3(12): 41, tab. 21. 1831.
truncate locus, 7–11 × 1.5–2 μm.
Syn.: Glomerella Spauld. & H. Schrenk, Science, N.Y. 17: 751.
Description and illustration: Yuan (1996). 1903.
CROUS ET AL.
164
Ascomata perithecial, solitary or gregarious, globose to sub- entire or sometimes slightly irregularly lobate margin, ovate,
globose, dark brown to black, lined with paraphyses, ostiole globose or ampulliform, brown to medium brown.
periphysate. Peridium pale to medium brown flattened cells of Materials examined: Brazil, Minas Gerais, Viçosa, Forestry Nursery, UFV, on
textura angularis. Asci 8-spored, unitunicate, cylindrical to sub- Eucalyptus or Corymbia, 17 Mar. 2011, A.C. Alfenas, culture CBS 130237.
fusoid, short pedicellate, with an inamyloid, refractive ring at the Indonesia, on Eucalyptus sp., 2012, M.J. Wingfield, culture CBS 133193.
apex. Ascospores uni- to biseriate, aseptate, hyaline, smooth-
Colletotrichum indonesiense Damm et al., Stud. Mycol. 73: 71.
walled, cylindrical, oval, fusiform or rhomboid, straight or
2012.
curved, one end ± acute and one ended round or both ends
rounded. Conidiomata acervular, conidiophores and setae Diagnosis: Conidia hyaline, smooth-walled, aseptate, straight,
formed on cushions of pale to medium brown, roundish to cylindrical to fusiform with both ends acute, (10.5–)
angular cells, in culture, ranging from sporodochia-like aggre- 13–17.5(–19) × (3–)3.5–4 μm, av. ± SD = 15.4 ± 2.2 × 3.7 ±
gations of conidiophores directly on hyphae to closed con- 0.2 μm, L/W ratio = 4.1.
idiomata that open by rupture. Setae often present, straight, pale
Description and illustration: Damm et al. (2012a).
to dark brown, smooth-walled, verruculose to verrucose, 1–8-
septate, base cylindrical, conical or slightly inflated, tip ± Typus: Indonesia, Sumatra, Tele, from leaf spots developing
rounded to ± acute. Conidiophores hyaline to pale brown, simple after herbicide treatment of Eucalyptus sp., 1 Jan. 2008, M.J.
or septate, branched or unbranched, smooth-walled, sometimes Wingfield (holotype CBS H-20798, culture ex-type CBS
verruculose. Conidiogenous cells enteroblastic, hyaline to pale 127551 = CPC 14986).
brown, smooth-walled, discrete, cylindrical, ellipsoidal, doliiform
Notes: This species belongs to the C. acutatum species complex
or ampulliform, collarette usually distinct, periclinal thickening
and was isolated from leaf spots developing after herbicide
visible to conspicuous, sometimes extending to form new con-
treatment of Eucalyptus sp.; this is to date the only report (Damm
idiogenous loci (percurrent) or surrounded by a gelatinous
et al. 2012a).
coating. Conidia hyaline, smooth-walled, aseptate, cylindrical,
clavate, fusiform, sometimes ellipsoidal to ovoid, straight or Colletotrichum karstii Y.L. Yang et al., Cryptog. Mycol. 32: 241.
curved, apex rounded to acute, sometimes with a filiform 2011. Fig. 25.
appendage, base rounded to truncate, sometimes with a prom- Diagnosis: Conidia 12.5–19.5 × 6–8.5 μm (av. 15.9 ± 1.4 × 6.8 ±
inent hilum. Appressoria single or in small groups, pale to dark
0.5), smooth-walled, hyaline, straight, cylindrical, truncate at the
brown, with a globose, elliptical, clavate, navicular or irregular
base, obtuse at the apex.
outline and an entire, undulate or lobate edge.
Description and illustration (of the holotype on Vanda sp. and on
Type species: Colletotrichum lineola Corda Diospyros australis, respectively): Yang et al. (2011) and Damm
Symptoms: Anthracnose disease symptoms: defined, often et al. (2012b).
sunken necrotic spots on leaves, stems, flowers or fruits. Addi- Material examined: South Africa, Free State, Frankfort, on E. grandis, 1 Mar.
tionally, crown and stem rots, ripe rot, seedling blights and brown 1990, P.W. Crous, culture CBS 127535 = CPC 193.
blotch are caused by species of this genus.
Notes: The C. karstii strains from E. grandis in South Africa that
Notes: Most of the reports of Colletotrichum from Eucalyptus in are included in Damm et al. (2012b) were initially identified as
Farr & Rossman (2017) and other sources refer to C. boninense by Lubbe et al. (2004) or as C. gloeosporioides by
C. gloeosporioides. According to these sources, this species P.W. Crous (information from CBS strain database) and later
causes leaf spots, twig dieback or lesions on bark and wood in reidentified by Damm et al. (2012b) as C. karstii, which resides in
Australia, Bangladesh, Brazil, Myanmar, South Africa and the the C. boninense species complex. It occurs on many host
USA. Before its epitypification (Cannon et al. 2008) and the plants, is very common and occurs in geographically diverse
review of the C. gloeosporioides complex (Weir et al. 2012), the areas (Damm et al. 2012b).
species was often incorrectly identified, and reports could refer
Colletotrichum theobromicola Delacr., Bull. Soc. Mycol. Fr. 21:
to various species in diverse species complexes (Hyde et al.
191. 1905.
2009 and see above). Apart from C. indonesiense, C. karstii
and C. theobromicola there are ITS sequences of four Diagnosis: Conidia subcylindrical to clavate, often with broadly
Colletotrichum strains from Eucalyptus available in GenBank. rounded ends, straight, sometimes developing a median septum
These strains as well as two strains from the CBS collection in age. Appressoria irregular, often somewhat lobed, (5–)
(U. Damm, unpubl. data) belong to one or more species 6–10(–14) × (4–)5–6(–8) μm; terminal or lateral, dark brown.
from the "Musae clade" in the C. gloeosporioides complex;
Description and illustration (of the neotype on Theobroma ca-
they are consequently neither C. gloeosporioides s. str. nor
cao): Rojas et al. (2010).
C. theobromicola.
Material examined: Brazil, on E. urophylla × E. grandis, no collection data
Colletotrichum gloeosporioides s. lat. available, strain CMM 3559.
Diagnosis: Conidia cylindrical, (10–)15–20(–25) × (3–)4–6 μm, Notes: Colletotrichum theobromicola causes leaf spot and stem
apex obtuse, base subacute, aseptate, guttulate, hyaline, girdling in nurseries of E. urophylla × E. grandis (“urograndis”) in
smooth, forming septum before germination. Appressoria with
Fig. 25. Colletotrichum karstii (CBS 127597). A–B. Conidiomata. C. Tip of seta. D. Basis of seta. E–I. Conidiophores. J–N. Appressoria. O–P. Conidia. Q–R. Ascomata. S.
Outer surface of peridium. T. Peridium in cross section. U. Ascospores. V–X. Asci. Y. Paraphyses. A, C–F, O. from Anthriscus stem. B, G–N, P–Y. from SNA. A–B, Q. DM,
C–P, R–Y. DIC, Scale bars: A = 200 μm, B, Q = 100 μm, R = 50 μm, E, S = 10 μm. Scale bar of E applies to C–P. Scale bar of S applies to S–Y. (from Damm et al. 2012b).
CROUS ET AL.
the Brazilian states of Para and Minas Gerais (Rodrigues et al. lacking. Ascospores ellipsoid, aseptate, hyaline, at times
2014). becoming pale brown at maturity, smooth, with or without
mucoid caps. Conidiomata pycnidial, immersed to semi-
Coma Nag Raj & W.B. Kendr., Canad. J. Bot. 50: 614. 1972.
immersed, unilocular, glabrous, ostiolate. Ostiole central, cir-
Synonym: Ascocoma H.J. Swart, Trans. Brit. Mycol. Soc. 87:
cular or oval, often situated in a conical or rostrate neck. Con-
606. 1987 (1986).
idiomata wall brown to dark brown or black wall of thin, pale
Conidiomata stromatic, acervular, immersed to erumpent, uni- brown textura angularis on exterior, and hyaline, thin-walled,
locular, brown to black, walls of textura angularis; dehiscence by textura prismatica in the inner layers except at base, which
irregular splits in apical wall. Conidiophores reduced to con- has a convex, pulvinate tissue of hyaline textura angularis giving
idiogenous cells, sparsely septate, branched, cylindrical, hyaline, rise to conidiophores or conidiogenous cells. Conidiophores
smooth, encased in mucus, discrete, cylindrical to doliiform. mostly reduced to conidiogenous cells, occasionally septate and
Conidia euseptate, with cylindrical to ovoid upper cell, 3–4 times branched at base, invested in mucus. Conidiogenous cells
longer than lower cell, pale brown, minutely verruculose, con- discrete, cylindrical, subcylindrical, obclavate or lageniform,
stricted at septum, bearing appendages. Appendages tubular, hyaline, smooth-walled, proliferating percurrently, or with visible
unbranched, flexuous; apical appendage single; lateral ap- periclinal thickening. Conidia ellipsoid, globose, napiform, fusi-
pendages 1–3, inserted at different loci below septum between form or naviculate with a truncate base and an obtuse to apic-
the two cells. Stromata subcuticular, forming multilocular asco- ulate apex, unicellular, thin- or thick-walled, smooth, olivaceous
mata that open irregularly. Asci clavate to cylindrical, apex brown to brown, sometimes with a longitudinal germ-slit, with or
rounded, unitunicate, eight-spored. Ascospores fusoid with without a mucoid appendage extending from apex to base on
rounded ends, aseptate or unevenly 1-septate, hyaline. Pa- one side; basal hilum with or without short tubular basal
raphyses numerous, extending above asci, unbranched, septate, appendage. Spermatophores formed in same conidioma, hya-
with swollen apices. line, smooth, 1-septate with several apical conidiogenous cells,
or reduced to conidiogenous cells. Spermatogenous cells cells
Type species: Coma circularis (Cooke & Massee) Nag Raj &
hyaline, smooth, lageniform to subcylindrical, with visible apical
W.B. Kendr.
periclinal thickening. Spermatia hyaline, smooth, red-shaped
Note: Although Coma circularis is a biotrophic foliar path- with rounded ends.
ogen of several eucalypt species, it appears to be of limited
Type species: Coniella fragariae (Oudem.) B. Sutton 1977 (syn.
importance.
Coniella pulchella Höhn. 1918)
Coma circularis (Cooke & Massee) Nag Raj & W.B. Kendr.,
Symptoms: Leaf spots amphigenous, prominent, medium to pale
Canad. J. Bot. 50: 614. 1972. Fig. 26.
brown, with or without prominent margins, ranging from small
Basionym: Pestalozziella circularis Cooke & Massee, Grevillea
specks with irregularly arranged conidiomata to large conspic-
18(no. 88): 80. 1890.
uous blotches with concentric circles of conidiomata
Diagnosis: Produces subcuticular stromata with heavily melanised (C. eucalyptorum).
upper layers; 1–10 mm diam, red-brown with black centre. This
Notes: Species of Coniella are commonly isolated as endophytes
fungus is a typical biotroph (Swart 1986a). Conidia 1-septate,
on a wide range of plants. However, they are also important foliar
24–34 × 5.5–9.5 μm with appendages; upper cell cylindrical to
pathogens of Eucalyptus (e.g. C. eucalyptorum, C. wangiensis),
ovoid, pale brown, minutely verruculose; lower cell almost hyaline,
causing a range of leaf disease symptoms, with some species
5.5–13 × 3.5–5 μm; appendages tubular, unbranched, flexuous;
being more important than others. Coniella eucalyptorum can
apical appendage single, 16.5–24.5 × 1.5–3 μm; lateral ap-
cause significant disease to E. dunnii plantations in sub-tropical
pendages 1–3, 14.5–28.5 μm long.
Australia following periods of high rainfall (Carnegie 2007b).
Descriptions and illustrations: Swart (1986a), Beilharz & Pascoe Several species tend to be collected more on leaf litter than from
(2005). diseased trees.
Typus: Australia, Victoria Alps, on Eucalyptus sp., Mrs Martin Coniella africana L.V. Alvarez & Crous, Stud. Mycol. 85: 12.
480 (holotype K). Not known from culture. 2016.
Coniella Höhn., Ber. Deutsch. Bot. Ges. 36: 316. 1918. Fig. 27. Diagnosis: Conidia hyaline to pale yellowish when mature, cy-
Synonyms: Schizoparme Shear, Mycologia 15: 120. 1923. lindrical, sometimes bent to naviculate, apex acute to nearly
Baeumleria Petr. & Syd., Beih. Reprium nov. Spec. Regni veg. rounded, base truncate, smooth-walled, multi-guttulate, germ slit
42: 268. 1927. absent (14.5–)15–20.5(–21) × (2.5–)3(–3.5) μm (l : w = 5.6).
Pilidiella Petr. & Syd., Beih. Reprium nov. Spec. Regni veg. 42:
Description and illustrations: Alvarez et al. (2016).
462. 1927.
Anthasthoopa Subram. & K. Ramakr., Proc. Indian Acad. Sci., Typus: South Africa, Mpumalanga, Barberton, on E. nitens leaf
Sect. B 43: 173. 1956. litter, 11 May 1990, P.W. Crous (holotype CBS H-22706, iso-
Cyclodomella Mathur et al., Sydowia 13: 144. 1959. type PREM 51098, culture ex-type CBS 114133 = CPC 405).
Embolidium Bat., Broteria, N.S. 33(3–4): 194. 1964 non Sacc.
Coniella destruens (M.E. Barr & Hodges) L.V. Alvarez & Crous,
1978.
Stud. Mycol. 85: 13. 2016.
Pathogens, saprobes. Ascomata brown to black, collapsed Basionym: Gnomoniella destruens M.E. Barr & Hodges, Myco-
collabent, erumpent, becoming superficial, globose, papillate, logia 79: 782. 1987.
with central periphysate ostiole. Asci clavate to subcylindrical, Synonyms: Schizoparme destruens (M.E. Barr & Hodges)
with distinct apical ring, floating free at maturity. Paraphyses Samuels et al., Mycotaxon 46: 470. 1993.
166
Fig. 26. Coma circularis (VPRI 17238). A–D. Disease symptoms. E, F. Vertical section through ascomata. G, H. Asci. I. Ascospores. J, K. Conidiogenous cells giving rise to
conidia. L, M. Conidia. N, O. Spermatogenous cells. P. Spermatia. Scale bars: E, F = 150 μm, all others = 10 μm.
Pilidiella destruens Crous & M.J. Wingf., Mycol. Res. 108: 299. an acutely rounded apex and subtruncate base with minute scar,
2004. pale to medium brown, granular, wall of medium thickness, (10–)
12–13(–15) × (3–)4–5(–6) μm (l : w = 2.7).
Diagnosis: Ascospores ellipsoidal, hyaline, thick-walled, gran-
ular, with terminal mucous caps, (9–)11–13 × (4.5–)5–6 μm. Description and illustration: Samuels et al. (1993), van Niekerk
Conidia long, fusoid-ellipsoidal, widest in the middle, tapering to et al. (2004).
CROUS ET AL.
Fig. 27. Disease symptoms associated with Coniella spp. A, B, D–G. C. eucalyptorum. C. C. wangiensis. H–L. C. eucalyptigena (CBS 139893). H. Ascomata forming on OA. I.
Ostiolar area. J, K. Asci. L. Ascospores. M–Q. C. eucalyptorum (CBS 112640). M. Conidiomata forming on OA. N. Transverse section through a conidioma. O, P. Con-
idiogenous cells giving rise to conidia. Q. Conidia. Scale bars: H, M = 250 μm, N = 150 μm, others = 10 μm. (from Alvarez et al. 2016).
Typus: USA, Hawaii, on twigs of E. grandis, Oct. 2000, M.J. Coniella eucalypticola Nag Raj, Canad. J. Bot. 54: 1370. 1976.
Wingfield (holotype of Pilidiella destruens CBS H-6945, holo-
Diagnosis: Conidia resembling C. paracastaneicola, but conidia
type of Gnomoniella destruens NY, isotype BPI 596643).
smaller, 19–29 × 2.5–3.5 μm.
168
Description and illustration: Nag Raj (1976). slits absent, (8–)8.5–11(–11.5) × (3–)3.5–4.5(–5) μm (l :
w = 2.5), with mucoid appendage alongside conidium.
Typus: India, Bangalore, Hebbal, on dead leaves of Eucalyptus
sp. lying on the ground, 25 Nov. 1966, T.R. Nag Raj (holotype Description and illustration: Alvarez et al. (2016).
DAOM 150596). Not known from culture.
Typus: Malaysia, on C. torelliana, 2009, S.S. Lee (holotype
Coniella eucalyptigena (Crous & M.J. Wingf.) L.V. Alvarez & CBS H-22711, culture ex-type CBS 141598 = CPC 16659).
Crous, Stud. Mycol. 85: 14. 2016. Fig. 27.
Coniella minima B. Sutton & Thaung, Nova Hedwigia 26: 10.
Basionym: Pilidiella eucalyptigena Crous & M.J. Wingf., Per-
1975.
soonia 34: 179. 2015.
Diagnosis: Conidia pale brown, ovate to broadly ventricose,
Diagnosis: Ascospores ellipsoidal, hyaline, thin-walled, granular,
6.5–7.5 × 3.5–4.5 μm.
with terminal mucoid caps or lateral appendages up to 5 μm
diam, or ascospore entirely encased in sheath; sheath dis- Description and illustration: Sutton (1975).
appearing with age, and ascospores becoming pale brown and
Typus: Myanmar, Burma, Katha, Nandayan Plantation, on
surface appearing roughened (possibly remnants of sheath),
E. camaldulensis, 15 Nov. 1973, Mya Thaung (holotype IMI
(10–)12–13(–14) × (4–)5–6 μm (l : w = 2.2).
179300). Not known from culture.
Descriptions and illustrations: Crous et al. (2015b), Alvarez et al.
Coniella paracastaneicola L.V. Alvarez & Crous, Stud. Mycol.
(2016).
85: 24. 2016.
Typus: Malaysia, Sabah, on E. brassiana, May 2014, M.J.
Diagnosis: Conidia hyaline, becoming pale olivaceous with age,
Wingfield (holotype CBS H-22222, cultures ex-type CBS
smooth, solitary, granular to guttulate, fusoid to naviculate, apex
139893 = CPC 24793, CPC 24794).
obtuse, base truncate, (21–)25–28(–31) × (3–)4(–5) μm (l :
Coniella eucalyptorum (Crous & M. J. Wingf.) L.V. Alvarez & w = 6.5), with mucoid appendage along side of conidium.
Crous, Stud. Mycol. 85: 15. 2016. Fig. 27. Developing conidia and conidiophores are frequently encased in
Basionym: Pilidiella eucalyptorum Crous & M. J. Wingf., Mycol. a mucoid sheath.
Res. 108: 296. 2004.
Description and illustration: Alvarez et al. (2016).
Diagnosis: Leaf spots large, irregular, pale brown with diffuse
margins, frequently secondary, associated with primary patho- Typus: Australia, Victoria, Toolangi State Forest, S37°33’25.3”
gens or with insect or wind damage. Conidia medium to dark red- E145°31’55.9”, on leaves of Eucalyptus sp., 9 Nov. 2014, P.W.
brown, broadly ellipsoidal or limoniform, widest in the middle, Crous, J. Edwards & P.W.J. Taylor (holotype CBS H-22702,
tapering to an acutely rounded apex and a subtruncate base, culture ex-type CBS 141292 = CPC 20146).
multiguttulate, with a longitudinal germ slit, wall of medium Coniella wangiensis (Crous & Summerell) L.V. Alvarez &
thickness as in C. fragariae, but basal mucoid appendage less Crous, Stud. Mycol. 85: 29. 2016. Fig. 27.
common than in C. fragariae, (9–)10–12(–14) × (6–)7–8 μm (l : Basionym: Pilidiella wangiensis Crous & Summerell, Persoonia
w = 1.6). 28: 177. 2012.
Description and illustration: Alvarez et al. (2016). Diagnosis: Conidia broadly ellipsoidal to globose, apiculate,
Typus: Australia, Queensland, Lannercost, plantation, on granular with central guttule, hyaline when immature, becoming
E. grandis × E. tereticornis hybrid, 10 Aug. 1999, P.Q. Thu & R.J. medium brown, frequently with minute basal cellular appendage,
Gibbs (holotype CBS H-6946, culture ex-type CBS hyaline, cylindrical, 1–2 μm long; conidia at times flattened along
112640 = CPC 3904 = DFR 100185). one side, or collapsing with age; apex tapering to an apiculus,
1–2 μm diam, base tapering to a truncate hilum, 1–1.5 μm diam,
Coniella fusiformis L.V. Alvarez & Crous, Stud. Mycol. 85: 16. (9–)10–11(–13) × (7–)8–9(–10) μm (l : w = 1.2).
2016.
Diagnosis: Conidia medium to dark red-brown, broadly ellipsoidal
or limoniform, widest in the middle, tapering to an acutely Typus: Australia, Northern Territory, Wangi Falls, Litchfield Na-
rounded apex and a subtruncate base, multiguttulate, with a tional Park, on Eucalyptus sp., 24 Apr. 2011, P.W. Crous & B.A.
longitudinal germ slit, wall of medium thickness as in Summerell (holotype CBS H-20969, culture ex-type CBS
C. fragariae, but basal mucoid appendage less common than in 132530 = CPC 19397).
C. fragariae, (9–)10–12(–14) × (6–)7–8 μm (l : w = 1.6).
Coniothyrium Corda, Icon. fung. (Prague) 4: 38. 1840.
Conidiomata pycnidial, solitary to gregarious, scattered,
Typus: Indonesia, on Eucalyptus sp., 2011, M.J. Wingfield immersed, globose, unilocular, pale to dark brown, with central,
(holotype CBS H-22713, culture ex-type CBS 141596 = CPC papillate ostiole; conidiomatal wall composed of thick-walled,
19722). brown textura angularis. Conidiophores reduced to con-
idiogenous cells. Conidiogenous cells doliiform or ampulliform,
Coniella malaysiana L.V. Alvarez & Crous, Stud. Mycol. 85: 21.
hyaline, smooth, lining inner cavity, with percurrent proliferation.
2016.
Conidia ellipsoid to broadly clavate or subcylindrical, apex
Diagnosis: Conidia hyaline to pale brown, fusoid to ellipsoid, obtuse, base truncate, 0–1-septate, constricted at the septum,
inequilateral, apex acutely rounded, widest in the middle, brown, thin-walled, smooth to verruculose (Sutton 1980, de
tapering to a truncate base, yellowish brown, thick-walled, germ Gruyter et al. 2013).
CROUS ET AL.
Type species: Coniothyrium palmarum Corda Typus: Brazil, S~ao Paulo, Suzano, on E. saligna, Jun. 1995, P.W.
Crous & A.C. Alfenas (holotype PREM 55302, cultures ex-type
Coniothyrium eucalypticola B. Sutton, Mycol. Pap. 123: 34.
CBS 111306 = CMW 14907 = CPC 1455, CBS 111318 = CPC
1971.
1457).
Diagnosis: Leaf spots amphigenous, subcircular to irregular,
Coremiopassalora leptophlebae (Crous et al.) U. Braun et al.,
3–5 mm diam, pale brown, appearing darker due to exuding
Stud. Mycol. 87: 316. 2017. Fig. 28.
spore masses; border indistinct, margin absent. Conidia asep-
Basionym: Passalora leptophlebae Crous et al. (as “lepto-
tate, ovoid, thick-walled, verruculose, medium brown with a
phlebiae”), Persoonia 26: 131. 2011.
distinct basal marginal frill, (8.5–)9–11(–12) × (6–)7–8(–9) μm.
Diagnosis: Leaf spots amphigenous, subcircular to irregular or
Descriptions and illustrations: Sutton (1971a), Crous (1998).
angular, 1–6 mm diam, confined by leaf veins, medium brown,
Typus: Australia, Victoria, Kiata Lowan Sanctuary, Kiata, on with raised border and red-purple margin, becoming confluent;
E. leptophylla, Apr. 1957, I.D. Williams (holotype IMI 71347). sporulation amphigenous. Conidia solitary, rarely in branched
Not known from culture. chains, pale brown, smooth, guttulate, subcylindrical to narrowly
obclavate, base obconically truncate, apex subobtuse, (15–)
Note: This appears to be a species of Readeriella, but fresh
18–22(–27) × 3(–3.5) μm, 1–3-septate; hila thickened, dark-
collections are required to resolve its taxonomy.
ened, refractive, 1 μm diam.
Coniothyrium kallangurense B. Sutton & Alcorn, Nova Hed-
Description and illustrations: Crous et al. (2011a).
wigia 26: 11. 1975.
Typus: Brazil, Minas Gerais, Viçosa, University Forestry Nurs-
Diagnosis: Conidia ovoid to broadly ellipsoid or pyriform, con-
ery, on E. leptophleba, 23 Aug. 2010, P.W. Crous, A.C. Alfenas,
stricted at truncate base, aseptate, medium brown, finely ver-
R. Alfenas & O.L. Pereira (holotype CBS H-20585, culture ex-
ruculose, (4–)5–7(–10) × (2–)3–5(–7) μm.
type CBS 129524 = CPC 18480).
Descriptions and illustrations: Sutton (1975), Crous (1998).
Cylindrocladiella Boesew., Canad. J. Bot. 60: 2289. 1982.
Typus: Australia, Queensland, Kallangur, on E. microcorys, 22 Fig. 29.
Dec. 1971, J.L. Alcorn (holotype IMI 164061). Not known from Synonym: Nectricladiella Crous & C.L. Schoch, Stud. Mycol. 45:
culture. 54. 2000.
Notes: In the absence of cultures, the taxonomy of Coniothyrium Ascomata perithecial, superficial, solitary, basal stroma absent, globose
kallangurense will remain unresolved. However, the synasexual to obpyriform, collapsing laterally when dry, smooth, with several min-
morph has globose, brown, muriformly septate conidia observed ute, brown setae arising from the perithecial wall surface, red, KOH+;
in culture (Crous 1998), suggesting that this is not a member of ostiole consisting of clavate cells, lined with inconspicuous periphyses;
Coniothyrium s. str. perithecial wall consisting of a single region of 3–4 cell layers of textura
angularis, which become hyaline and slightly flattened towards the
Coremiopassalora U. Braun et al., Stud. Mycol. 87: 316. 2017.
centre. Asci unitunicate, 8-spored, cylindrical, sessile, thin-walled, with a
Differs from the genus Passalora by synnematous conidiophores flattened apex, and a refractive apical apparatus. Ascospores uni-
and catenate, hyaline to pale olivaceous conidia with distinct, seriate, overlapping, hyaline, ellipsoid to fusoid with obtuse ends,
slightly thickened and not darkened loci. smooth, 1-septate. Conidiophores monomorphic, penicillate, or dimor-
phic, penicillate and subverticillate, mononematous, hyaline; penicillate
Type species: Coremiopassalora eucalypti (Crous & Alfenas) U.
conidiophores comprising a stipe, a penicillate arrangement of fertile
Braun et al. (≡ Mycovellosiella eucalypti Crous & Alfenas)
branches, a stipe extension, and a terminal vesicle; subverticillate co-
Notes: The genus Coremiopassalora includes two species that nidiophores comprising a stipe, and one or two series of phialides; stipe
are morphologically similar to taxa in Passalora s.lat., but distinct septate, hyaline, smooth; stipe extensions aseptate, straight, thick-
from the type, Passalora bacilligera. walled, with one basal septum, terminating in a thin-walled vesicle of
characteristic shape. Conidiogenous apparatus with primary branches
Coremiopassalora eucalypti (Crous & Alfenas) U. Braun et al.,
aseptate or 1-septate; secondary branches aseptate, each terminal
Stud. Mycol. 87: 316. 2017.
branch producing 2–4 phialides; phialides cylindrical, straight, or dolii-
Basionym: Mycovellosiella eucalypti Crous & Alfenas, Mycol.
form to reniform to cymbiform, hyaline, aseptate, apex with minute
Mem. 21: 105. 1998.
periclinal thickening and collarette. Conidia cylindrical, rounded at both
Synonym: Passalora eucalypti (Crous & Alfenas) Crous & U.
ends, straight, (0–)1(–3)-septate, frequently slightly flattened at the
Braun, in Crous & Braun, CBS Biodiversity Ser. 1: 452. 2003.
base, held in asymmetrical clusters by colourless slime. Chlamydo-
Diagnosis: Leaf spots amphigenous, irregular to subcircular, grey spores brown, thick-walled, more frequently arranged in chains than
to pale brown in the center, becoming medium brown toward a clusters (from Crous 2002).
red-brown border, 1–15 mm diam. Conidia catenulate, pale
Type species: Cylindrocladiella parva (P.J. Anderson) Boesew.
olivaceous, smooth, thin-walled, subcylindrical to narrowly
obclavate, apex obtuse to truncate, base truncate, occurring in Symptoms: Brown water-soaked leaf spots and cutting rot.
branched or unbranched chains, 1(–3)-septate, (14–)
Note: Cylindrocladiella is mostly a problem in nurseries and is not
20–30(–40) × (1.5–)2–2.5 μm in vivo, (0–)1(–8)-septate, (10–)
found causing leaf spot disease in plantations.
15–30(–100) μm in vitro; hila thickened, darkened, refractive.
Cylindrocladiella lageniformis Crous, M.J. Wingf. & Alfenas,
Description and illustration: Crous (1998). Mycol. Res. 97: 441. 1993.
170
Fig. 28. Coremiopassalora leptophlebae (CBS H-20585). A–C. Disease symptoms. D–G. Conidiophores. H. Conidia. Scale bars = 10 μm.
Diagnosis: Conidiophores monomorphic. Vesicles lageniform to Davisoniella H.J. Swart, Trans. Brit. Mycol. Soc. 90: 289. 1988.
ovoid. Conidia (9–)11.5(–15) × (1.5–)2 μm.
Conidiomata stromatic, abaxial, solitary to gregarious, subepi-
Description and illustration: Crous & Wingfield (1993). dermal, erumpent, multilocular at maturity. Conidiomata wall
composed of brown-walled cells of textura angularis. Co-
Typus: Brazil, Espirito Santo, Aracruz nursery, on Eucalyptus
nidiophores reduced to conidiogenous cells. Conidiogenous cells
cuttings, A.C. Alfenas (holotype PREM 50927, culture ex-type
holoblastic, percurrent proliferation, flask-shaped. Conidia oval,
CBS 340.92 (wrongly cited as CBS 338.92 in the original
apex rounded, base truncate, with marginal frill, brown, verru-
publication) = PPRI 4449 = UFV 115).
culose (adapted from Swart 1988).
Cylindrocladiella lanceolata L. Lombard & Crous, Mycol. Progr.
Type species: Davisoniella eucalypti H.J. Swart
11: 851. 2012. Fig. 29.
Notes: The genus Davisoniella was established by Swart (1988)
Diagnosis: Conidiophores monomorphic. Vesicles lanceolate,
for what he perceived as a more stromatic form of Coniothyrium.
5–7 μm diam. Conidia (13–)15–17(–20) × 2–3 μm (av. 16 ×
Crous et al. (2006d) linked the fungus to its sexual morph, which
3 μm).
at the time was still referred to as “Mycosphaerella”. Morpho-
Description and illustration: Lombard et al. (2012). logically, Davisoniella resembles the Colletogloeopsis (Crous &
Wingfield 1997) asexual morph found with species of Ter-
Typus: Australia, Queensland, Brisbane, from soil, 18 Jul. 2009,
atosphaeria. Although Davisoniella is clearly a member of Ter-
P.W. Crous (holotype CBS H-20602, cultures ex-type CBS
atosphaeriaceae, its not clear if the genus should be treated as a
129566 = CPC 17567, CBS 129565 = CPC 17566).
synonym of Teratosphaeria, because DNA data are currently not
Cylindrocladiella pseudohawaiiensis L. Lombard & Crous, available to confirm this synonymy.
Mycol. Progr. 11: 857. 2012. Fig. 29.
Davisoniella eucalypti H.J. Swart, Trans. Brit. Mycol. Soc. 90:
Diagnosis: Conidiophores dimorphic. Vesicles clavate to ellip- 289. 1988. Fig. 30.
soidal, 6–8 μm diam. Conidia (11–)12–14(–15) × 2–4 μm (av. Synonym: Mycosphaerella davisoniella Crous, Stud. Mycol. 55:
13 × 3 μm). 111. 2006.
Description and illustration: Lombard et al. (2012). Diagnosis: Leaf spots amphigenous, subcircular to irregular,
Typus: Brazil, Sao Paulo, Aracruz nursery, on Eucalyptus 1–7 mm diam, discrete to confluent, medium brown, surrounded
cuttings, 1992, A.C. Alfenas (holotype CBS H-20607, culture by raised, red-purple margin. Conidiogenous cells subcylindrical
ex-type CBS 210.94 = IMI 361579 = PPRI 44500 = UFV to ampulliform or doliiform, 5–15 × 3–4 μm, medium brown,
1250). verruculose, proliferating several times percurrently near the
CROUS ET AL.
Fig. 29. Cylindrocladiella spp. A–D. Conidiophores of Cylindrocladiella spp. on infected alfalfa seeds A–B. Cylindrocladiella spp. sporulating on the seed coat of Medicago
sativa. C. Root rot of M. sativa seedling. D. Cylindrocladiella spp. on carnation leaf. E–AA. Asexual morph. E–I. Penicillate conidiophores. E. C. longistipitata (ex-type CBS
116075). F. C. kurandica (ex-type CBS 129577). G. C. lanceolata (ex-type CBS 129566). H. C. pseudoparva (ex-type CBS 129560). I. C. nederlandica (ex-type CBS 152.91).
J–N. Penicillate conidiogenous apparatus. J. C. hawaiiensis (ex-type CBS 129569). K. C. australiensis (ex-type CBS 129567). L. C. natalensis (ex-type CBS 114943). M.
C. cymbiformis (ex-type CBS 129553). N. C. ellipsoidea (ex-type CBS 129573). O–S. Subverticillate conidiophores. O. C. australiensis (ex-type CBS 129567). P.
C. longiphialidica (ex-type CBS 129557). Q. C. pseudohawaiiensis (ex-type CBS 210.94). R, S. C. natalensis (ex-type CBS 114943). T–Y. Terminal vesicles of stipe extensions.
T. C. hawaiiensis (ex-type CBS 129569). U. C. stellenboschensis (ex-type CBS 110668). V. C. cymbiformis (ex-type CBS 129553). W. C. variabilis (ex-type CBS 129561). X.
C. lanceolata (ex-type CBS 129566). Y. C. kurandica (ex-type CBS 129577). Z, AA. Conidia. Z. C. natalensis (ex-type CBS 114943). AA. C. brevistipitata (ex-type CBS
142783). Scale bars: E–I, O = 50 μm; J–N, P–AA = 10 μm (from Marin-Felix et al. 2019).
apex. Conidia solitary, brown, aseptate, verruculose, thick- from the brown basal rim of the conidium, (8–)
walled, oval with an obtuse apex and a truncate to subtruncate 10–12(–14) × 4.5–)5–6(–6.5) μm (av. 11 × 5.5 μm).
base with a prominent basal frill, which can extend up to 2 μm
Descriptions and illustrations: Swart (1988), Crous et al. (2006d).
172
Typus: Australia, Western Australia, Darling Ranges, Mund- Basionym: Mycosphaerella endophytica Crous & H. Sm. ter,
limup Block, on E. marginata, 24 Nov. 1981, F. Tay (holotype of Mycol. Mem. 21: 54. 1998.
D. eucalypti and M. davisoniellae DAR 58999). Not known from Synonyms: Pseudocercosporella endophytica Crous & H. Sm.
culture. ter, Mycol. Mem. 21: 55. 1998.
Mycosphaerella pseudoendophytica Crous & G. Hunter, Stud.
Devonomyces Videira & Crous, Stud. Mycol. 87: 353. 2017.
Mycol. 55: 118. 2006.
Ascomata pseudothecial, amphigenous, subepidermal, be-
Diagnosis: Leaf spots amphigenous, irregular, 5–20 mm diam,
coming erumpent, subglobose to globose, with apical, papillate
medium brown, centre frequently grey, border slightly raised,
ostiole; walls of 2–3 layers of medium brown textura angularis,
dark brown; coloured margin absent. Ascospores bi- to triseriate,
subhymenium of 1–2 layers of hyaline cells. Asci fasciculate,
overlapping, colourless, guttulate, thin-walled, straight, fusoid-
bitunicate, cylindrical to narrowly obovoid, straight or slightly
ellipsoidal with obtuse ends, widest in middle of apical cell,
incurved, 8-spored. Ascospores bi- to triseriate, overlapping,
medianly 1-septate, not constricted at septum, tapering toward
hyaline, guttulate, thin-walled, straight, fusoid-ellipsoidal with
both ends, but with more prominent taper toward base, (8–)
obtuse ends, medianly 1-septate. Mycelium internal, consisting
9–10(–11) × (2–)2.5–3 μm. Conidia solitary, colourless,
of septate, branched, hyaline to brown, smooth to verruculose
smooth, narrowly obclavate, 0–8-septate, irregularly curved,
hyphae. Caespituli sporodochial, situated on a brown stroma
rarely straight, apex obtuse, base long obconic-truncate, lateral
consisting of verruculose, brown, globose cells and hyphal ele-
branches common, secondary conidia forming on most mature
ments. Conidiophores rarely pigmented and verruculose in lower
primary conidia; conidia aggregated in slimy masses (13–)
part, mostly hyaline and smooth throughout, thick-walled, cylin-
25–45(–50) × 1.5–2(–2.5) μm.
drical, straight to irregularly curved, septate. Conidiogenous cells
terminal, hyaline, smooth, unbranched, straight or slightly curved, Description and illustration: Crous (1998).
proliferating sympodially. Conidia solitary, hyaline, smooth,
narrowly obclavate, septate, irregularly curved, rarely straight, Typus: South Africa, Western Cape Province, Stellenbosch,
apex obtuse, base long obconic-truncate, lateral branches Devon Valley, on Eucalyptus sp., Jun. 1995, P.W. Crous (holo-
common, secondary conidia forming on most mature primary type of M. endophytica PREM 54398, culture ex-type CBS
conidia; conidia aggregated in slimy masses (from Videira et al. 114662 = CPC 1193); KwaZulu-Natal, Enon, Richmond, on
2017). E. nitens, 3 May 2000, G. Hunter (holotype of
M. pseudoendophytica CBS H-19694, culture ex-type CBS
Type species: Devonomyces endophyticus (Crous & H. Sm. ter)
113288 = CMW 9098).
Videira & Crous
Note: Devonomyces forms an asexual morph with masses of Disculoides Crous et al., Persoonia 28: 71. 2012. Fig. 32.
slimy, hyaline conidia, undergoing microcyclic conidiation.
Conidiomata brown to black, amphigenous, subepidermal,
Devonomyces endophyticus (Crous & H. Sm. ter) Videira & acervular, opening by irregular rupture; wall of 2–3 layers of brown
Crous, Stud. Mycol. 87: 353. 2017. Fig. 31. textura angularis. Macroconidiophores 1–2-septate or reduced to
Fig. 30. Davisoniella eucalypti (DAR 58999). Asci with ascospores, conidiogenous Fig. 31. Devonomyces endophyticus (PREM 54398). Asci with ascospores,
cells with conidia, and conidiogenous cells with conidia of potential synasexual germinating ascospores, and conidiogenous cells with conidia undergoing micro-
morph. Scale bars = 10 μm. (from Crous et al. 2006d). cyclic conidiation. Scale bars = 10 μm. (from Crous et al. 2004b).
CROUS ET AL.
Fig. 32. Disculoides spp. A–H. D. eucalypti (CPC 17648). A, B. Disease symptoms. C–G. Conidiogenous cells giving rise to conidia. H. Conidia. I–L. D. fraxinoides (CPC
32420). I. Disease symptoms. J. Conidiogenous cells giving rise to conidia. K. Macro- and microconidia. L. Microconidia. Scale bars = 10 μm.
conidiogenous cells. Macroconidiogenous cells lining the inner Notes: Disculoides was recently established for a genus of foliar
cavity of conidioma, hyaline to olivaceous, smooth, subcylindrical pathogens on Eucalyptus. All species presently known are
to ampulliform, tapering to a long, thin neck, proliferating several associated with prominent leaf spot symptoms. Disculoides
times percurrently near apex, with flaring collarettes. Macroconidia calophyllae is reduced to synonymy with D. corymbiae based on
olivaceous, smooth, guttulate, ellipsoid to fusoid to somewhat multigene sequence data. Although the species treated can be
obclavate, straight to curved, apex subobtuse, base truncate, with distinguished based on their conidial dimensions, there is
prominent marginal frill, up to 1 μm long. Microconidiophores considerable overlap among them, and identifications are best
hyaline, smooth, ampulliform, 0–2-septate, branched below, or confirmed based on DNA sequence data.
reduced to conidiogenous cells, proliferating percurrently near
Disculoides corymbiae Crous, Persoonia 37: 313. 2016.
apex. Microconidia solitary, aseptate, hyaline, smooth, guttulate.
Synonym: Disculoides calophyllae Crous, Persoonia 38: 263. 2017.
Type species: Disculoides eucalypti Crous et al.
Diagnosis: Conidia hyaline, smooth, thick-walled, guttulate,
Symptoms: Associated with large, prominent, amphigenous ellipsoid to fusoid, straight to curved, (10–)12–14(–15) × (3.5–)
brown leaf spots, at times with a red-purple margin, 5–15 mm 4(–4.5) μm; apex subobtuse, base truncate, 1–1.5 μm diam,
diam. with minute marginal frill.
174
Description and illustration: Crous et al. (2016b). (on leaves of E. kingsmillii) having macroconidia that are ellipsoid
to fusoid, straight to curved, and larger in size (17–)
Typus: Australia, Western Australia, Denmark, Mount Lindesay
20–26(–30) × (6–)7(–8) μm.
Walk Trail, on C. calophylla, 19 Sep. 2015, P.W. Crous (holotype
CBS H-22872, culture ex-type CBS 142076 = CPC 28864); near Elsinoe Racib., Parasit. Alg. Pilze Java’s (Jakarta) 1: 14. 1900.
Kojonup, on C. calophylla, 18 Sept. 2015, P.W. Crous (holotype Fig. 34.
of Disculoides calophyllae CBS H-23125, culture ex-type CBS Synonym: Sphaceloma de Bary, Ann. Oenol. 4: 165. 1874.
142080 = CPC 29246). Additional synonyms in MycoBank.
Disculoides eucalypti Crous et al., Persoonia 28: 71. 2012. Ascostromata solitary, aggregated, or gregarious, wart-like, or as
Fig. 32. small distinctively coloured elevations, or pulvinate, immersed to
Synonym: Disculoides eucalyptorum Crous et al., Persoonia 28: semi-immersed, globose to subglobose, white, pale yellow or
73. 2012. brown, soft, multi-loculate, locules scattered in upper part of
ascostromata, cells of ascostromata comprising pseudoparen-
Diagnosis: Conidia olivaceous, smooth, guttulate, ellipsoid to
chymatous cells of textura globulosa to angularis. Locules with
fusoid to somewhat obclavate, straight to curved, (13–)
few to numerous asci inside each locule, ostiolate. Ostiole
14–16(–17) × (4–)5–6(–6.5) μm in vivo, (16–)
minute, periphyses absent. Asci 8-spored, bitunicate, fissituni-
17–20(–23) × (5–)6–6.5(–7) μm in vitro.
cate, saccate to globose, with a minute pedicel, and ocular
Description and illustration: Crous et al. (2012b). chamber. Ascospores irregularly arranged, oblong or fusiform
with slightly acutely rounded ends, with 2–3 transverse septa,
Typus: Australia, Victoria, Melbourne, S38°24'3.1"
hyaline, smooth-walled, lacking a sheath. Sphaceloma asexual
E144°59'36.9", on Eucalyptus sp., 12 Oct. 2009, P.W. Crous, J.
morph: Acervuli or sporodochia subepidermal, pseudoparen-
Edwards, I.J. Porter & I.G. Pascoe (holotype of D. eucalypti
chymatous. Conidiophores hyaline to pale-brown, polyphialidic.
CBS H-20935, culture ex-type CBS 132183 = CPC 17650);
Conidiogenous cells formed directly from the upper cells of the
Victoria, Woorndoo, S37°53'29.3" E142°47'51.1", on
pseudoparenchyma, mono- to polyphialidic, integrated or
E. viminalis, 17 Oct. 2009, P.W. Crous, J. Edwards, I.J. Porter &
discrete, determinate, hyaline to pale brown, without visible
I.G. Pascoe (holotype of D. eucalyptorum CBS H-20936, culture
periclinal thickening. Conidia hyaline, smooth, aseptate, ellip-
ex-type CBS 132184 = CPC 17648).
soidal, guttulate.
Disculoides fraxinoides Crous, sp. nov. MycoBank
Symptoms: Plant pathogenic, causing scab, leaf spot and
MB832016. Fig. 32.
anthracnose disease. Leaf spots are irregular, brown to red-
Etymology: Name refers to Eucalyptus fraxinoides, the host purple, with diffuse margins, and usually with white, flaky
species from which this fungus was isolated. centre, due to the raised, pealing off cuticle.
Associated with leaf spots. Conidiomata black, amphigenous, Notes: There are several species of Elsinoe that cause scab
subepidermal, acervular, opening by irregular rupture, disease on Eucalyptus foliage, although some are only known
200– 300 μm diam on banana leaf agar, 300 –600 μm diam on based on their asexual morphs. The disease can be prominent
OA; wall of 6 – 10 layers of brown textura angularis. Macro- on some eucalypt species, but it is generally regarded to be of
conidiophores reduced to conidiogenous cells or 1 – 2-septate, little importance. The genus Elsinoe was recently revised by Fan
10 – 15 × 4– 6 μm. Macroconidiogenous cells terminal and et al. (2017), who also provided a key to species occurring on
intercalary, hyaline, smooth, subcylindrical to ampulliform, Eucalyptus.
tapering to a long thin neck, 10 –15 × 3.5– 4 μm, proliferating
Elsinoe eucalypti Hansf. Proc. Linn. Soc. New South Wales 79:
percurrently at apex, with minute flaring collarette. Macro-
98. 1954.
conidia hyaline (pale brown in old cultures), aseptate, smooth,
thick-walled, guttulate, ellipsoid, straight to curved, (16– ) Diagnosis: Ascospores 20–28 × 7–8 μm.
19 – 21( –26) × (4 –)6 –7( – 8) μm; apex obtuse, base truncate,
Description: Hansford (1954).
2 μm diam, with minute marginal frill. Microconidiophores
hyaline, smooth, ampulliform, 0 – 2-septate, branched Typus: Australia, South Australia, Kangaroo Island, Rocky
below, or reduced to conidiogenous cells, 8 – 15 × 3 – 5 μm, River, on Eucalyptus sp., Feb. 1920, T. Osborn (holotype WARI
proliferating percurrently near apex. Microconidia solitary, 2010) (Hansford 1954). Not known from culture.
aseptate, hyaline, smooth, guttulate, (5 –)6 – 8( – 10) × (2– )
Elsinoe eucalypticola Cheew. & Crous, Persoonia 23: 64.
3( –3.5) μm.
2009. Fig. 34.
Culture characteristics: Colonies flat, spreading, with sparse
Diagnosis: Ascospores hyaline to pale brown, broadly ellipsoid
aerial mycelium and even, lobate margins, covering dish after 2
with rounded ends, with more prominent taper towards the base,
wk at 25 °C. On MEA surface dirty white, reverse luteous. On
with 4-transverse septa, and 0–3 vertical septa, and sometimes
PDA surface and reverse pale luteous. On OA surface pale
with oblique septa; mostly slightly constricted at the median
luteous.
septum, (16–)17–18(–20) × (6.5–)7–8 μm.
Typus: Australia, New South Wales, Merimbula, on
E. fraxinoides, 28 Nov. 2016, P.W. Crous, HPC 1808 (holotype
CBS H-24023, culture ex-type CBS 143430 = CPC 32420). Typus: Australia, Queensland, Cairns, Kuranda Kennedy
Highway, on Eucalyptus sp., 26 Sep. 2006, P.W. Crous (holo-
Note: There is considerable morphological variation within the
type CBS H-20283, cultures ex-type CBS 124765 = CPC 13318,
clade that includes this species (Fig. 33), with isolate CPC 29454
CPC 13319, 13320).
CROUS ET AL.
Elsinoe eucalyptigena Crous, Persoonia 37: 391. 2016. Conidiomata pycnidial, globose, pale to medium brown,
100–200 μm diam. Conidiophores lining the inner cavity, hyaline,
Diagnosis: Ascospores hyaline to pale brown, broadly ellipsoid
smooth, subcylindrical, 1–2-septate, straight to flexuous,
with rounded ends, with more prominent taper towards the base,
branched below, 15–30 × 3–4 μm. Conidiogenous cells hyaline,
with 4-transverse septa, and 0–3 vertical septa, and sometimes
smooth, terminal and intercalary, subcylindrical, straight to
with oblique septa; mostly slightly constricted at the median
slightly curved, 8–15 × 3–3.5 μm; proliferating inconspicuously
septum, (16–)17–18(–20) × (6.5–)7–8 μm.
percurrently at apex. Conidia solitary, aseptate, hyaline, smooth,
Description and illustration: Crous et al. (2016b). guttulate, subcylindrical, straight, apex obtuse, base truncate,
12–13 × 3(–3.5) μm.
Typus: Australia, Western Australia, Perth, King’s Park
Botanic Gardens, on E. kingsmillii, 27 Sep. 2015, P.W. Crous Culture characteristics: Colonies erumpent, with sparse aerial
(holotype CBS H-22913, culture ex-type CBS 142127 = CPC mycelium and uneven folded surface, with lobate margin,
29529). reaching 5 mm diam after 2 wk at 25 °C. On MEA, PDA and OA
surface and reverse scarlet, with diffuse scarlet pigment on PDA
Elsinoe eucalyptorum Crous & Summerell, Fungal Diversity 23:
and OA.
332. 2006.
Typus: Australia, New South Wales, Fitzroy Falls, Morton Na-
Diagnosis: Ascospores hyaline, smooth, thin-walled, broadly
tional Park, on E. piperita, 26 Nov. 2016, P.W. Crous, HPC 1747
ellipsoidal with rounded ends, with 1(–3) transverse septa, and
1–2 vertical or oblique septa; constricted at median septum,
32705).
(11–)13–15 × (4–)5(–6) μm.
Notes: Based on a megablast search of NCBI’s GenBank
Description and illustration: Summerell et al. (2006).
nucleotide database, the closest hits using the ITS sequence
Typus: Australia, New South Wales, 0.9 km west of Pacific had highest similarity to Elsinoe preissianae (GenBank
Highway on Middle Brother Road, ca. 11 km south of Kew, North KY173406.1; Identities = 750/771 (97 %), 3 gaps (0 %)),
Coast NSW, 31 42 38 S 152 42 20 E, alt. 40 m; on E. propinqua, Elsinoe lepagei (GenBank MH856598.1; Identities = 455/507
Feb. 2006, B.A. Summerell (holotype CBS H-19746, culture ex- (90 %), 11 gaps (2 %)), and Elsinoe theae (GenBank
type CBS 120084 = CPC 13052). NR_148174.1; Identities = 447/500 (89 %), 9 gaps (1 %)).
Closest hits using the rpb2 sequence had highest similarity to
Elsinoe piperitae Crous, sp. nov. MycoBank MB832017. Fig. 34.
Elsinoe barleriicola (GenBank KX887083.1; Identities = 567/689
Etymology: Name refers to Eucalyptus piperita, the host species (82 %), 2 gaps (0 %)), Elsinoe diospyri (GenBank KX887093.1;
from which this fungus was collected. Identities = 557/692 (80 %), 8 gaps (1 %)), and Elsinoe
Fig. 33. The first of four equally most parsimonious trees obtained from a phylogenetic analysis of the Disculoides combined ITS, cmdA and tub2 sequence alignment (12
strains including the outgroup; 1 563 characters analysed: 1 103 constant, 384 variable and parsimony-uninformative and 76 parsimony-informative). The tree was rooted to
Harknessia hawaiiensis (GenBank AY720723.1, AY720786.1, AY720755.1, respectively) and the scale bar indicates the number of changes. Thickened branches represent
those present in the strict consensus parsimony tree. Bootstrap support values higher than 49 % are shown at the nodes (parsimony bootstrap support / NJ HKY85 distance
bootstrap support) and novelties are highlighted with bold text. Species names are indicated to the right of the tree. A superscript T denotes strains with a type status. Tree
statistics: TL = 507 steps, CI = 0.957, RI = 0.833, RC = 0.797. The analyses were performed using PAUP v. 4.0b10 (Swofford 2003).
176
Fig. 34. Elsinoe eucalypticola (CBS H-20283). A, B. Disease symptoms. C–E. Ascomata with asci. F–H. Asci. I–M. Ascospores. Scale bars = 10 μm (from Cheewangkoon
et al. 2009). N–S. E. piperitae (CPC 32705). N. Colony on OA. O–R. Conidiogenous cells. S. Conidia. Scale bars = 10 μm.
leucospermi (GenBank KX887123.1; Identities = 551/686 Description and illustration: Crous et al. (2016b).
(80 %), no gaps). Closest hits using the tef1 sequence had
highest similarity to Elsinoe australis (GenBank JX993826.1;
National Park, Central lookout, on E. preissiana, 23 Sep. 2015,
Identities = 176/197 (89 %), 2 gaps (1 %)), and Elsinoe bidentis
(as Sphaceloma bidentis; GenBank KF421125.1; Identi-
142129 = CPC 29787).
ties = 295/377 (78 %), 28 gaps (7 %)).
Elsinoe salignae Crous & Carnegie, Persoonia 42: 333.
Elsinoe preissianae Crous, Persoonia 37: 393. 2016.
2019.
Diagnosis: Conidiogenous cells integrated, terminal, poly-
Diagnosis: Conidiogenous cells integrated, subcylindrical, hya-
phialidic, hyaline, smooth, ampulliform to subcylindrical or
line, smooth-walled, mono- to polyphialidic, 8–12 × 3–4 μm.
doliiform, 5–10 × 3–4 μm. Conidia hyaline, smooth, aseptate,
Conidia solitary, aggregating in mucoid mass, aseptate, hyaline,
granular, narrowly ellipsoid to subcylindrical, apex obtuse, base
smooth-walled, guttulate, subcylindrical to ellipsoid, apex obtuse,
truncate, 1–1.5 μm diam, (10–)11–12(–13) Х (3–)
base truncate, (4.5–)5–6(–6.5) × (2–)2.5 μm.
3.5( –4) μm.
CROUS ET AL.
Description and illustration: Crous et al. (2019a). Diagnosis: Leaf spots amphigenous, medium brown, subcircular.
Conidiomata scattered to gregarious, superficial, sporodochial,
Typus: Australia, New South Wales, Bulladelah, Bulladelah
cupulate, orange-yellow. Conidiophores arising from a stroma,
State Forest, S32°23’34.8” E152°15’11.9”, on E. saligna, 16 Apr.
septate, branched, pale orange-brown, 30–60 × 2.5–3.5 μm;
2016, A.J. Carnegie, HPC 2415 (holotype CBS H-23948, culture
forming a penicillate conidiogenous apparatus. Conidiogenous
ex-type CPC 35713 = CBS 145552).
cells integrated, terminal, lageniform to cylindrical, in whorls,
Elsinoe tectificae (Cheew. & Crous) Fan & Crous, Stud. Mycol. orange-brown, finely roughened to warty, thin-walled, with apical
87: 33. 2017. percurrent proliferations, 7–16 × 2.5–4 μm, apex 1–1.5 μm
Basionym: Sphaceloma tectificae Cheew. & Crous, Persoonia diam. Conidia (3–)4(–4.5) × 2 μm, aseptate, hyaline, smooth,
23: 79: 2009. thin-walled, ellipsoid, apex subobtuse, base truncate, 1 μm diam,
widest in upper third, lacking appendages.
Diagnosis: Conidiogenous cells phialidic, hyaline, smooth, un-
branched, occurring as lateral tips on hyphae, sometime with Description and illustration: Crous et al. (2016b).
one basal supporting cell, obclavate to cylindrical, tapering sharply
Typus: Malaysia, Kota Kinabalu, on leaf spots of E. pellita, 30
to a truncate apex, (4–)8–10 × 2.5–3(–5) μm. Conidia acroge-
May 2015, M.J. Wingfield (holotype CBS H-22871, cultures ex-
nous, ellipsoid to short cylindrical or obovoid, aseptate,
type CBS 142074 = CPC 28764, CBS 142075 = CPC 28748).
4–4.5 × 2–2.5 μm, hyaline, minutely guttulate, hila slightly rounded
to subtruncate; germinating conidia become slightly elongated and Eucasphaeria Crous, Fungal Diversity 25: 21. 2007.
swollen, thicker walled, guttulate, but remain hyaline. Morphologically similar to Plectosphaera, but lacking a clypeus,
and having unitunicate asci with an apical discharge mecha-
Description and illustration: Fan et al. (2017).
nism, producing an ascochytopsis-like asexual morph in
Typus: Australia, Northern Territory, road to Robin Falls, culture.
S14°10’20", E131°07’15", on E. tectifica, 23 Sep. 2007, coll. B.A.
Type species: Eucasphaeria capensis Crous
Summerell, isol. P.W. Crous (holotype CBS H-20296, cultures
ex-type CBS 124777 = CPC 14594, CPC 14595, 14596). Symptoms: Leaf spots irregular to subcircular, amphigenous,
medium brown, with or without dark brown margin.
Eucalyptostroma Crous & M.J. Wingf., Persoonia 37: 311. 2016.
Notes: Because the original description of Eucasphaeria
Conidiomata scattered to gregarious, superficial, sporodochial,
capensis noted the fact that the fungus also occurred on litter
cupulate, orange-yellow; basal stroma of textura intricata. Setae
(Crous et al. 2007b), its potential role as a foliar pathogen
absent. Conidiophores arising from a stroma, septate, branched,
remained unclear. However, with the recent description of
pale orange-brown; forming a penicillate conidiogenous appa-
Eucasphaeria rustici from leaf spots of two different species of
ratus. Conidiogenous cells integrated, terminal, lageniform to cy-
eucalypts in Australia (Crous et al. 2016b), it seems probable
lindrical, in whorls, orange-brown, finely roughened to warty, thin-
that this could be a genus of foliar pathogens, although of minor
walled, with apical percurrent proliferations, 7–16 × 2.5–4 μm,
importance.
apex 1–1.5 μm diam. Conidia (3–)4(–4.5) × 2 μm, aseptate,
hyaline, smooth, thin-walled, ellipsoid, apex subobtuse, base Eucasphaeria capensis Crous, Fungal Diversity 25: 21. 2007.
truncate, 1 μm diam, widest in upper third, lacking appendages. Fig. 36.
Type species: Eucalyptostroma eucalypti Crous & M.J. Wingf. Diagnosis: Ascospores hyaline, guttulate, fusoid-ellipsoidal,
mostly curved, 1-septate, not constricted at median septum,
Note: Although the genus Eucalyptostroma (Chaetosphaer-
widest at septum, tapering towards both subobtuse ends, mul-
iaceae) was found associated with leaf spots on Eucalyptus,
tiseriate, (17–)19–25(–28) × (3–)3.5(–4) μm. Conidiomata
nothing is known about its biology or ability to cause disease.
subepidermal, opening by irregular ruptures, acervuloid, up to
Eucalyptostroma eucalypti Crous & M.J. Wingf., Persoonia 37: 150 μm diam; wall consisting of 5–6 layers of brown cells of
311. 2016. Fig. 35. textura angularis, becoming hyaline towards inner conidiogenous
Fig. 35. Eucalyptostroma eucalypti (CPC 28764). A. Colony sporulating on MEA. B–D. Conidiophores with conidiogenous cells. E. Conidia. Scale bars = 10 μm (from Crous
et al. 2016a).
178
Fig. 36. Eucasphaeria capensis (CBS H-19764, CBS 120027). A–C. Sporulation on PDA. D. Ostiolar region. E. Periphyses. F, G. Asci. H–K. Conidia and conidiogenous cells.
Scale bars = 10 μm (from Crous et al. 2007b).
region. Conidia hyaline, minutely guttulate, predominantly Diagnosis: Conidia hyaline, guttulate, falcate, widest in middle,
falcate, widest in middle, aseptate, base subtruncate, apex 0–1-septate, base subtruncate, apex subobtusely rounded,
subobtusely rounded, (16–)20–22(–27) × (2–)2.5(–3) μm on (12–)17–22(–25) × (2.5–)3(–4) μm, covered in mucus.
host; in culture sporulating by means of inconspicuous spor-
odochia, conidia 0–2-septate, (22–)25–30(–40) × 3(–4) μm;
conidia covered in mucus. Typus: Australia, Western Australia, Perth, King’s Park Botanic
Gardens, on leaves of E. creta, 27 Sep. 2015, M.J. Wingfield
Typus: South Africa, Western Cape Province, on N7 from 28946).
Malmesbury towards Cape Town, on living leaves and leaf litter
Euteratosphaeria Quaedvl. & Crous, Persoonia 33: 25. 2014.
of Eucalyptus sp., Jan. 2006, P.W. Crous (holotype CBS
H-19764, cultures ex-type CBS 120028 = CPC 12796, CPC Ascomata pseudothecial, solitary, black, immersed becoming
12797). erumpent, globose; ostiole apical, central; wall of 2–3 layers of
medium brown textura angularis. Asci aparaphysate, fasciculate,
Eucasphaeria rustici Crous & T.I. Burgess, Persoonia 37: 317.
bitunicate, subsessile, obovoid to narrowly ellipsoid, straight or
2016.
CROUS ET AL.
slightly incurved, 8-spored. Ascospores tri- to multiseriate, Notes: In a review of the Passalora complex, Crous & Braun
overlapping, hyaline, guttulate, thin-walled, straight, ellipsoid with (2003) included genera such as Mycovellosiella, Phaeoramu-
obtuse ends, medianly 1-septate, tapering towards both ends, laria and Fulvia as synonyms of Passalora. Videira et al. (2017)
but more prominently towards the lower end. also showed that these genera were phylogenetically distinct.
Type species: Euteratosphaeria verrucosiafricana (Crous & M.J. Exopassalora zambiae (Crous & T.A. Cout.) Videira & Crous,
Wingf.) Quaedvl. & Crous Stud. Mycol. 87: 380. 2017. Fig. 38.
Basionym: Passalora zambiae Crous & T.A. Cout., Stud. Mycol.
Note: Presently only known from Australia and Indonesia, and
50: 209. 2004.
not regarded a significant pathogen.
Euteratosphaeria verrucosiafricana (Crous & M.J. Wingf.)
diam, medium brown, surrounded by a raised, brown border.
Quaedvl. & Crous, Persoonia 33: 25. 2014. Fig. 37.
Conidia catenulate, chains simple or branched, medium brown,
Basionym: Mycosphaerella verrucosiafricana Crous & M.J.
smooth, narrowly ellipsoidal, tapering to subtruncate, with flat-
Wingf., Stud. Mycol. 55: 125. 2006.
tened ends, straight or slightly curved, 0(–2)-septate,
Diagnosis: Leaf spots amphigenous, irregular to sub-circular, 10–20 × 2–3 μm in vitro.
5–15 mm diam, pale brown to grey, surrounded by a raised,
dark brown border, and a thin, red-purple margin. Ascospores tri-
to multiseriate, overlapping, hyaline, guttulate, thin-walled, Typus: Zambia, on E. globulus, 21 Aug. 1995, T. Coutinho
straight, ellipsoid with obtuse ends, medianly 1-septate, widest (holotype CBS H-9895; cultures ex-type CBS 112971 = CPC
in the middle of the apical cell, constricted at the septum, 1227, CBS 112970 = CPC 1228).
tapering towards both ends, but more prominently towards the
Fairmaniella Petr. & Syd., Beih. Reprium nov. Spec. Regni veg.
lower end, (7–)8–9(–10) × 3(–3.5) μm in vivo.
42: 481. 1927. (1926).
Descriptions and illustrations: Crous et al. (2006d), Quaedvlieg
et al. (2014).
Typus: Indonesia, Northern Sumatra, on Eucalyptus sp., Feb.
2004, M.J. Wingfield (holotype CBS H-19705, culture ex-type
CBS 118496 = CPC 11167).
Exopassalora Videira & Crous, Stud. Mycol. 87: 380. 2017.
Mycelium composed of brown hyphae, smooth to rough, irreg-
ularly branched, septate, with dark brown chlamydospore-like
hyphal swellings. Conidiophores arising from the mycelium,
medium brown, smooth, simple or branched, straight to curved.
Conidiogenous cells terminal and intercalary, subcylindrical, pale
to medium brown, smooth, proliferating sympodially, con-
idiogenous loci conspicuous, darkened, refractive. Conidia
catenate, in simple or branched chains, medium brown, smooth,
narrowly ellipsoidal, tapering to subtruncate, straight or slightly
curved, hila slightly thickened and darkened.
Type species: Exopassalora zambiae (Crous & T.A.Cout.)
Videira & Crous Fig. 38. Exopassalora zambiae (CBS 112971). Conidia, conidiogenous cells and
germinating ascospores. Scale bar = 10 μm (from Crous et al. 2004b).
Fig. 37. Euteratosphaeria verrucosiafricana (CPC 11167). A. Leaf spot. B, C. Asci. D, E. Ascospores. F, G. Germinating ascospores. Scale bars = 10 μm (from Crous et al.
2006d).
180
Conidiomata acervular, subepidermal, dark brown, solitary; wall Fusculina Crous & Summerell, Fungal Diversity 23: 334. 2006.
composed of thick-walled, medium brown textura epidermoidea
Fusculina is intermediary between Fusicoccum and Disculina,
to prismatica; conidioma with brown hyphal threads, verruculose,
having pycnidial conidiomata, and hyaline, ellipsoid conidia that
septate, 3–4 μm diam, giving rise to conidia in a sympodial
arise from percurrently proliferating conidiogenous cells.
fashion, holoblastic. Conidia ellipsoid to ovate, base obtuse to
truncate, apex obtuse, aseptate, pale brown, thick-walled, Type species: Fusculina eucalypti Crous & Summerell
punctulate.
Notes: Very little is known regarding the biology of Fusculina or
Type species: Fairmaniella leprosa (Fairm.) Petr. & Syd. whether this is a pathogen. A third species, F. eucalyptorum
(Crous et al. 2018b) is not treated here, as it was collected as an
Notes: Many records reporting Fairmaniella leprosa from coun-
endophyte, and has not been associated with any foliar disease
tries such as South Africa (Crous et al. 1989a) and Chile
symptoms.
(Wingfield et al. 1995), actually refer to Blastacervulus eucalypti.
The interpretation of F. leprosa was confused for many years, Fusculina eucalypti Crous & Summerell, Fungal Diversity 23:
and Sutton (1980) treated mixed collections as representative of 334. 2006.
F. leprosa, giving an incorrect interpretation of its conidiogenesis.
Diagnosis: Leaf spots amphigenous, circular, medium brown with
The question was resolved by Swart (1988) who introduced the
a raised border, 2–6 mm diam. Conidiomata amphigenous, black
genus Blastacervulus. Originally, he assumed that these col-
on leaves, pycnidial. Primary conidiogenous cells hyaline,
lections also represented F. leprosa (H.J. Swart, pers. comm.).
smooth, ampulliform to subcylindrical, proliferating several times
Fairmaniella is not known from culture, and its relationship to
percurrently near apex, 5–15 × 4–8 μm, giving rise to sub-
Blastacervulus remains unresolved.
cylindrical primary conidia. Primary conidia hyaline, granular,
Fairmaniella leprosa (Fairm.) Petr. & Syd., Beih. Reprium nov. smooth, straight to somewhat irregular, subcylindrical, apex
Spec. Regni veg. 42: 481. 1927. 1926. obtuse, base truncate, (16–)18–20(–22) × 3–4 μm.
Basionym: Coniothyrium leprosum Fairm., in Millspaugh & Nut-
tall, Publications of the Field Museum of Natural History, Botany
Series 5: 328. 1923. Typus: Australia, New South Wales, Wollemi National Park, on
Synonym: Melanconium eucalypticola Hansf., Proc. Linn. Soc. Eucalyptus sp., Jan. 2006, B. Summerell (holotype CBS H-
New South Wales 81: 47. 1956. 19747, culture ex-type CBS 120083 = CPC 12946).
Diagnosis: Leaf spots dark brown, varying from angular to cir- Fusculina regnans Crous, sp. nov. MycoBank MB832018.
cular, and acervular pustules also vary in diameter and height, Fig. 39.
depending on the host species (Swart 1988). Conidiomata
Etymology: Name refers to Eucalyptus regnans, the host species
acervular, producing long, flexuous multiseptate, brown co-
from which it was collected.
nidiophores with phialidic conidiogenous cells, each forming a
fertile locus beneath a septum. Conidia pale brown, aseptate,
Conidiomata immersed, becoming erumpent, black, occurring
thick-walled, ellipsoid to ovoid, apex obtuse, base truncate,
in leaf spots of Allelochaeta. Sporulating on OA: conidiomata
5–7.5 × 3–4 m.
eustromatic, opening via irregular split, grey-black, occurring
Description and illustration: Swart (1988). in clusters, 150 –300 μm diam; wall of 6 –8 layers of brown
textura angularis. Conidiophores lining the inner cavity, hy-
Typus: Australia, South Australia, Adelaide, on E. fasciculosa,
aline, smooth, 0 – 2-septate, subcylindrical, 15 – 25 × 5 –7 μm.
1924, G. Samuel (holotype of Melanconium eucalypticola WARI
Conidiogenous cells terminal, integrated, subcylindrical to
3846, isotype IMI 57994). Not known from culture. USA, Cali-
ampulliform, hyaline, smooth, with numerous prominent
fornia, fruits of cultivated, 3 Jun. 1920, collector unknown (ho-
percurrent proliferations in apical region, 5– 20 × 5 –7 μm.
lotype of Coniothyrium leprosum Nuttall 408).
Paraphyses intermingled among conidiophores, hyaline,
Fig. 39. Fusculina regnans (CPC 32732). A. Disease symptoms on F. regnans. B. Conidiomata on OA. C–F. Conidiogenous cells. G. Conidia. Scale bar = 10 μm.
CROUS ET AL.
smooth, septate, 25 – 40 × 4– 5 μm. Conidia solitary, asep- unilocular to convoluted and multilocular, brown; walls composed
tate, hyaline, smooth, guttulate, subcylindrical, apex obtuse, of thin-walled, pale brown to brown textura angularis. Ostiolar
base truncate to bluntly rounded, (14 – )17 –20( – 23) × (3.5– ) opening central, circular, wide, surrounded by brown furfura-
4 –5( –6) μm. ceous cells. Conidiophores lining the inner cavity, or limited to a
basal layer in some species; usually reduced to conidiogenous
Culture characteristics: Colonies erumpent, spreading, surface
cells, rarely septate and branched; commonly invested in mucus.
folded, with sparse aerial mycelium and even, lobate margins,
Conidiogenous cells discrete, ampulliform, lageniform, sub-
reaching 7 mm diam after 2 wk at 25 °C. On MEA surface dirty
cylindrical to cylindrical, hyaline, smooth, giving rise to macro-
white, reverse buff. On PDA surface olivaceous grey, reverse
conidia, and in some cases also microconidia in the same
smoke grey. On OA surface dirty white.
conidioma, proliferating one to several times percurrently;
Typus: Australia, Victoria, Kinglake National Park, on secession rhexolytic. Macroconidia consisting of a conidium
E. regnans, 1 Dec. 2016, P.W. Crous, HPC 1885 (holotype CBS body and a basal appendage, delimited by a septum; conidium
H-24025, culture ex-type CBS 143428 = CPC 32732). body unicellular, of various shapes, thick-walled, smooth, brown,
with or without light and dark coloured longitudinal bands, oc-
Notes: Phylogenetically, this species is related to F. eucalypti casionally longitudinally striate, guttulate; basal appendage
(Fig. 40). The two ITS sequences are 96 % (697/723, including 3 cellular, cylindrical to subcylindrical, hyaline, fIexuous, thin-
gaps) similar. walled and devoid of contents; apical appendage mostly lack-
Golovinomyces (U. Braun) V.P. Heluta, Biol. Zh. Armenii 41: ing, when present elongated, attenuated; in some species the
357. 1988. conidium body and basal appendage are invested in a thin layer
Basionym: Erysiphe sect. Golovinomyces U. Braun, Feddes of mucus. Microconidia oval to ellipsoid, aseptate, hyaline,
Repert. 88: 659. 1978 (1977). smooth. Ascomata perithecial, single or aggregated, immersed,
disc furfuraceous brown, neck emergent to depressed; wall of
Asexual morph: Euoidium - mycelium external; hyphal 3–5 layers of brown textura angularis. Asci unitunicate, cylin-
appressoria usually nipple-shaped; conidia in chains with drical to clavate, hyaline, smooth, 8-spored, with apical appa-
sinuate outline, without fibrosin bodies. Sexual morph: chas- ratus. Paraphyses hyaline, septate, interspersed among asci.
mothecia with mycelium-like appendages and two to numerous Ascospores aseptate, uni- to biseriate, ellipsoid to fusoid, hya-
usually 2-spored asci (for full description see Braun & Cook line, thick-walled, guttulate, smooth.
2012).
Type species: Harknessia eucalypti Cooke
Type species: Golovinomyces cichoracearum (DC.) V.P. Heluta
Symptoms: Small leaf spots, vein limited or not, irregular to
Notes: This powdery mildew pathogen has a wide host range subcircular, medium brown (with chlorotic margin on some eu-
other than Eucalyptus spp. It is less common on Eucalyptus than calypts), mostly occurring along leaf margins, with conidiomata
the well-known powdery mildew Podosphaera aphanis. occurring on leaves and petioles.
Golovinomyces orontii (Castagne) V.P. Heluta, Ukr. bot. Zh. Notes: Most species of Harknessia appear to be endophytes,
45: 63. 1988. Fig. 41. and Crous & Rogers (2001) concluded that the majority are
Basionym: Erysiphe orontii Castagne, Suppl. Cat. Pl. Mars.: 52. secondary or weak pathogens. Numerous species are known to
1851. occur on Eucalyptus (Sutton & Pascoe 1989b, Crous et al.
Diagnosis: White mycelial growth on leaves and stems. Co- 1993d, 2007b, 2012c, 2016b, Nag Raj 1993, Lee et al. 2004,
nidiophores erect, foot-cells straight, often curved in basal half. Summerell et al. 2006), but only a few are routinely associated
Conidia ellipsoid-ovoid to doliiform-subcylindrical, 25–40 × (10–) with leaf spots, such as H. eucalyptorum and H. hawaiiensis.
15–23(–25) μm. Generally, species in the genus are regarded to be of little
economical importance (Park et al. 2000).
Description, illustration and specimens: Braun & Cook (2012).
Harknessia eucalyptorum Crous et al., Mycologia 85: 109.
Harknessia Cooke, Grevillea 9(no. 51): 85. 1881. Figs 42, 43. 1993. Fig. 43.
Mycelium internal, branched, septate, hyaline to pale brown. Synonym: Wuestneia eucalyptorum Crous et al., Mycologia 85:
Conidiomata eustromatic to pycnidial, immersed, globose, 112. 1993.
Fig. 40. The single most parsimonious tree obtained from a phylogenetic analysis of the Fusculina ITS sequence alignment (four strains including the outgroup; 690 characters
analysed: 393 constant, 284 variable and parsimony-uninformative and 13 parsimony-informative). The tree was rooted to Delitschia confertaspora (GenBank MF459003.1) and
the scale bar indicates the number of changes. Bootstrap support values higher than 49 % are shown at the nodes and novelties are highlighted with bold text. Species names
are indicated to the right of the tree and GenBank accession numbers between round parentheses. A superscript T denotes strains with a type status. Tree statistics: TL = 339
steps, CI = 0.985, RI = 0.615, RC = 0.606. The analysis was performed using PAUP v. 4.0b10 (Swofford 2003).
182
Fig. 41. Golovinomyces orontii. A–E. Disease symptoms. F, G. Conidial chains. H. Conidia. Scale bar = 10 μm.
Diagnosis: Macroconidia holoblastic, broadly ventricose with Typus: Australia, Western Australia, Perth, King's Park Botanic
central guttule. aseptate. dark brown, astriate, apex obtuse Gardens, on E. platyphylla, 26 Sep. 2015, M.J. Wingfield (ho-
to bluntly apiculate, base truncate, 16 – 29 × 9 – 24 μm lotype CBS H-23105, culture ex-type CBS 142542 = CPC
(av. 22 × 11 μm), basal appendage 3 – 16 μm long. Asco- 28862).
spores aseptate, uni- or biseriate, ellipsoidal, with obtuse
Hyalozasmidium U. Braun et al., Stud. Mycol. 87: 375. 2017.
ends, hyaline, thick-walled, guttulate, smooth, 13 – 28 ×
8 – 13 μm. Mycelium composed of subhyaline to pale brown hyphae,
smooth, branched and septate, producing large swollen propa-
Description and illustration: Crous et al. (1993d).
gules that occur terminaly or laterally on hyphal strands. Co-
Typus: South Africa, Western Cape Province, Stellenbosch, nidiophores medium to dark brown, unbranched, smooth to
Stellenbosch mountain, on E. andrewsii, 20 Dec. 1989, P.W. verruculose, becoming constricted at septa, eventually dis-
Crous (holotype PREM 50830, isotypes DAOM 211794, IMI articulating, with each conidiophore giving rise to a single
338270b, culture ex-type CBS 111115 = CPC 85). conidium. Conidiogenous cells apical and intercalary, mono- or
polyblastic, straight, proliferating sympodially, with conidiogenous
Harknessia hawaiiensis F. Stevens & E. Young, Bull. Bernice P.
loci unthickened or slightly thickened, located at shoulders and
Bishop Museum 19: 136. 1925.
apex. Conidia hyaline, thick-walled, subcylindrical, with multiple
Diagnosis: Conidia globose, 9–13.5 × 8–11 μm (av. 11 × 9 μm), transverse septa, developing irregular swellings which can form
with striations; basal appendage 1–8 μm long. branches with obtuse ends, body granular, basal cell tapering
prominently towards the conidiophore. Differs from the genus
Zasmidium, by bearing hyaline conidia.
Typus: USA, Hawaii, Oahu, Waipio, on E. robusta, 1 Jul. 1919,
Type species: Hyalozasmidium aerohyalinosporum (Crous &
H.L. Lyon, Lyon 124 (holotype IMI 147757).
Summerell) Videira & Crous (≡ Zasmidium aerohyalinosporum
Harknessia platyphyllae Crous, Persoonia 38: 313. 2017. Crous & Summerell)
Diagnosis: Conidia (16–)17–19(–21) × (11–)12–13(–15) μm Note: Very little is known regarding the biology or pathogenicity
(av. 18 × 12.5 μm) in vitro, broadly ellipsoid, apex acutely of species in this genus.
rounded, aseptate, apiculate, pale yellow-brown, thick-walled,
Hyalozasmidium aerohyalinosporum (Crous & Summerell)
smooth, striations along length of conidium, multi-guttulate. Basal
Videira & Crous, Stud. Mycol. 87: 375. 2017. Fig. 44.
appendage (4–)6–8(–20) × 2–2.5 μm.
Basionym: Zasmidium aerohyalinosporum Crous & Summerell,
Description and illustration: Crous et al. (2017b). Persoonia 23: 144. 2009.
CROUS ET AL.
Fig. 42. Leaf spot disease symptoms associated with Harknessia spp. on different Eucalyptus hosts. A. H. fusiformis (CPC 13649). B. H. hawaiiensis (15003). C, D.
H. rhabdosphaera (CPC 13593 and CPC 12847). E. H. globispora (CPC 14924). F. H. eucalyptorum (CPC 12697) (from Crous et al. 2012c).
Synonym: Paramycosphaerella aerohyalinosporum (Crous & Hypsotheca Ellis & Everh., J. Mycol. 1: 128. 1885.
Summerell) Guatimosim et al., Persoonia 37: 124. 2016. Synonyms: Capnodiella (Sacc.) Sacc. & D. Sacc., Syll. fung.
(Abellini) 17: 621. 1905. [based on Capnodium maximum]
Diagnosis: Leaf spots amphigenous, irregular grey-brown, sur-
Sorica Giesenh., Ber. dt. bot. Ges. 22: 195. 1904. [based on
rounded by a dark brown to red-purple margin; spots varying
Sorica dusenii]
from specks to spots, up to 7 mm diam; hypophyllous surface
well colonised by black, submerged spermatogonia. Conidia Ascomata separate or loosely grouped, not arising from a visible
hyaline, thick-walled, subcylindrical, with 3–30 transverse septa, stroma, dark brown to black, ventricose, straight or curved,
cells 5–15 μm long, developing irregular swellings which can elongate with a submedian to suprabasal swollen ascigerous
form branches, 4–7 μm wide, 40–150 μm long, apex and lateral locule. Ascomatal wall of textura porrecta to textura intricata. Asci
branches with obtuse ends; body granular, basal cell tapering 8-spored, elongating at maturity and extending up the ascoma
prominently towards the conidiophore. Typical brown, verrucu- neck to the apex before deliquescing to release ascospores at or
lose, obclavate conidia with thickened scars rarely observed. below the ostiole; discharged ascospores accumulating in a dry
reddish brown mass at the ostiole. Ascospores golden brown,
thick-walled, smooth, depressed globose to subellipsoid. Pyc-
Typus: Australia, Northern Territory, Road to Robin Falls, nidial and hyphomycetous morphs produced. Pycnidial con-
S13°31'01.3", E131°16'22.5", 126 m, on leaves of E. tectifica, 23 idiomata solitary, dark brown to black, globose or depressed
Sep. 2007, coll. B.A. Summerell, isol. P.W. Crous (holotype of globose, or short stipitate, with a prominent papillate ostiole, wall
Zasmidium aerohyalinosporum CBS H-20274, cultures ex-type of textura angulata to textura intricata. Conidiophores hyaline,
CBS 125011 = CPC 14636, CPC 14637). arising from the inner cells of the pycnidial wall, simple
184
Fig. 43. Harknessia spp. A–F. H. eucalyptorum (CPC 12697). A. Ascoma with short neck, oozing ascospores. B. Vertical section through conidioma. C, D. Asci. E. Ascal tip. F.
Ascospores. G, H. Conidiomata oozing conidia. I, J. Conidiogenous cells giving rise to conidia of H. pseudohawaiiensis (CPC 17380). K. Conidiogenous cells giving rise to
conidia of H. kleinzeeina (CPC 16277). L. Conidiogenous cells giving rise to conidia of H. gibbosa (CPC 12473). M. Conidia of H. australiensis (CPC 13596). N. Conidia of
H. pseudohawaiiensis (CPC 17379). O. Conidia of H. eucalyptorum (CPC 12697). P, Q. Conidia of H. kleinzeeina (CPC 16277). Scale bars: B = 170 μm, all others = 10 μm.
CROUS ET AL.
Fig. 44. Hyalozasmidium aerohyalinosporum (CPC 14636). A. Leaf spot with black spermatogonia (small black dots). B–E. Germinating ascospores on MEA. F–Q. Con-
idiogenous cells giving rise to conidia (arrow denotes point of attachment). Scale bars = 10 μm (from Crous et al. 2009b).
186
ampulliform or elongate, septate. Conidiogenous cells phialidic erumpent, eustromatic, acervular to sporodochial, composed of
with an inconspicuous collarette. Conidia hyaline, asymmetrical, thick-walled, dark to reddish brown textura angularis. Co-
oblong to allantoid or fusoid, aseptate, smooth. Hyphomycetous nidiophores dark to reddish brown, coarsely verrucose, cylin-
morph with mucoid heads of conidia scattered on short lateral drical, unbranched, septate, formed from the upper cells of the
phialides, phaeoacremonium-like, sub-hyaline to pale brown, conidiomata. Conidiogenous cells integrated, dark to reddish
smooth or rough. Conidiogenous cells lageniform, the collarettes brown, coarsely verrucose to tuberculate, cylindrical, with several
usually inconspicuous or flared (phialophora-like). Conidia percurrent enteroblastic proliferations. Conidia holoblastic, dark
aseptate, ellipsoid-ovoid, smooth. to reddish brown, coarsely verrucose to tuberculate, with 0-
several eusepta, straight to curved, obtuse or acute at apex,
Type species: Hypsotheca subcorticalis [Basionym: Sphaer-
truncate at base, cylindrical to fusiform. Conidiogenesis: a suc-
onaema subcorticale, perithecia occurring inside the bark of
cession of conidia is formed by holoblastic conidial ontogeny,
Quercus, New Jersey, USA, type at K]
delimitation by a transverse septum, schizolytic secession,
Notes: Species of Hypsotheca are well-known plant pathogens replacement wall building apex leading to enteroblastic percur-
and cause serious canker diseases on species of Eucalyptus rent conidiogenous cell proliferation followed by holoblastic
(Pascoe et al. 2018, Crous et al. 2019b). Further collections are conidial ontogeny, with successive conidia seceding at pro-
required however to elucidate the role of Hypsotheca as foliar gressively higher levels (Sutton & Crous 1997).
pathogens, but recent collections suggest that species in the
Type species: Lecanostictopsis kamatii (Ullasa) B. Sutton &
genus are more common than previously assumed.
Crous
Hypsotheca eucalyptorum Crous & Carnegie, Persoonia 42:
Notes. The genus Lecanostictopsis appears to have co-evolved
367. 2019.
with species of Syzygium (Sutton & Crous 1997). A single
Diagnosis: Conidiomata sparsely formed in culture. Dominant species, L. eucalypti, is known from Eucalyptus in India. It is,
morph hyphomycetous. Conidiophores erect on superficial hy- however, possible that this host was originally incorrectly iden-
phae, 0–1-septate, unbranched, subcylindrical, straight to flex- tified, and represents a species of Syzygium, although it was not
uous, brown, verruculose, 5–20 × 1.5–2.5 μm. Conidiogenous possible to resolve this question based on the specimen
cells terminal, pale brown, verruculose, subcylindrical, phialidic examined. Lecanostictopsis is allied to the Mycosphaerellaceae
with flared collarette, 2–3 μm diam, 5–15 × 1.5–2.5 μm. Conidia (Crous, unpubl. data).
aseptate, solitary, hyaline, smooth, guttulate, subcylindrical with
Lecanostictopsis eucalypti Crous, Mycol. Mem. 21: 38. 1998.
obtuse ends, (3–)3.5–4(–4.5) × 1.5(–2) μm.
Fig. 46.
Diagnosis: Leaf spots amphigenous, dark brown, sometimes with
Typus: Australia, New South Wales, Kyogle, Boorabee State a grey center, angular, elongated, 2–20 mm in length, 2–8 mm
Forest, McCorquodale plantation, S28°36’46.5” E153°3’2.7”, on in width, confined by leaf veins, border raised, margin absent, or
E. grandis × camaldulensis clone, 20 Apr. 2016, A.J. Carnegie, thin and red when present. Conidia holoblastic, brown, coarsely
HPC 2431 (holotype CBS H-23966, culture ex-type CPC verruculose to tuberculate, 0–3-septate, cylindrical to fusiform,
35734 = CBS 145576). straight to curved, apex obtuse to bluntly rounded, base truncate,
(11–)22–25(–35) × 4–5(–6) μm.
Hypsotheca pleomorpha (Patricia McGee & I. Pascoe) Crous,
Fungal Syst. Evol. 3: 87. 2019. Fig. 45. Description and illustration: Crous (1998).
Basionym: Caliciopsis pleomorpha Patricia McGee & I. Pascoe,
Typus: India, Karnataka, Devarayanadurga, on Eucalyptus sp., 7
Fungal Syst. Evol. 2: 50. 2018.
Mar. 1968, K.A. Lucy-Channamma (holotype BPI 436205). Not
Diagnosis: Conidiomata pycnidial, globose, ostiolate, brown, known from culture.
50–200 μm diam, separate (on PNA), or aggregated in a brown
Lembosiniella Crous, gen. nov. MycoBank MB832019.
stroma (on PDA, MEA). Conidiophores arising from inner layer,
hyaline, smooth, subcylindrical, branched, 1–4-septate, Etymology: Name reflects its similarity to the genus Lembosina.
5–20 × 3–4 μm. Conidiogenous cells subcylindrical to doliiform,
Phytopathogenic. Leaf spots amphigenous, dark brown to black,
hyaline, smooth, terminal and intercalary, phialidic with prominent
superficial, irregularly rounded. Hysterothecia linear to rarely Y-
periclinal thickening, 3–6 × 2–4 μm. Conidia solitary, aseptate,
shaped, superficial, amphigenous, black, opening by a central lon-
hyaline, smooth, granular, fusoid-ellipsoid, mostly somewhat
gitudinal slit. Pseudoparaphyses filiform, hyaline, simple or
curved, apex obtuse, tapered towards base, truncate, 0.5 μm
branched, septate. Asci broadly ellipsoidal to obclavate, bitunicate, 8-
diam, (3–)4–5(–6) × 1.5(–2) μm.
spored, sessile to short-stalked. Ascospores fusoid, hyaline to brown,
Typus: Australia, Victoria, Flinders, Orcadia Park, on medianly 1-septate, constricted, smooth to echinulate, guttulate.
E. cladocalyx ‘Nana’, Nov. 1991, L. Wesley (holotype VPRI
17721). Type species: Lembosiniella eucalyptorum Crous
Material examined: Australia, New South Wales, Fitzroy Falls, Morton National Notes: Lembosiniella is ecologically different from Lembosina,
Park, on E. piperita, 2014, P.W. Crous, HPC 1762, culture CBS 144636 = CPC which appears to be a saprobe on Rhododendron (Crous
32144. unpubl. data), whereas Lembosiniella causes prominent
round, brown leaf spots on Eucalyptus. The ascal endotunica
Lecanostictopsis B. Sutton & Crous, Mycol. Res. 101: 215. 1997.
is multi-layered in Lembosina, but this is not the case in
Mycelium immersed, intercellular, branched, septate, dark to Lembosiniella. Although the 1-septate fusoid-ellipsoid asco-
reddish brown. Conidiomata epidermal to subepidermal, spores are brown, and smooth to verruculose in both genera,
CROUS ET AL.
Fig. 45. Hypsotheca pleomorpha (CPC 32144). A–D. Ascomata on host, oozing asci and ascospores. E. Asci. F. Ascospores. G, H. Pycnidial conidioma sporulating in culture.
I. Conidioma with ostiole. J–M. Conidiogenous cells giving rise to conidia. N. Conidia. O–Q. Conidiogenous cells. R. Conidia. Scale bars: A–C = 120 μm, all others = 10 μm.
188
30–35(–37) × 8–9(–10) μm. Germinating ascospores dark

brown, distorting, verruculose, covered in mucoid layer, germi-
nating with 1–2 brown, verruculose germ tubes that are promi-
nently constricted at septa; ascospores 5–12 μm diam, with germ
tubes parallel or perpendicular to the long axis of the spore.
Culture characteristics: Colonies erumpent, spreading, with
sparse aerial mycelium and smooth margin, reaching 5 mm diam
after 2 wk at 25 °C. On MEA surface greenish black, reverse
olivaceous grey; on PDA surface greenish black, reverse oliva-
ceous grey with diffuse yellow pigment in agar; on OA surface
greyish sepia.
Typus: Australia, New South Wales, Tooloom, Beury State
Forest plantation, Tin Hut Forestry Rd, S28°35’35.0”,
E152°21’44.9”, on E. dunnii, 20 Jan. 2016, A.J. Carnegie, HPC
1443 (holotype CBS H-24028, culture ex-type CBS
144603 = CPC 31816).
Notes: Lembosina eucalypti is reminiscent of Thyrinula eucalypti

(= Aulographina eucalypti), except that the ascospores are hy-
aline in the latter genus. The type species of Lembosina occurs
Fig. 46. Lecanostictopsis eucalypti (BPI 436205). Aggregated conidiophores giving as a saprobe on twigs of Rhododendron in the Netherlands, and
rise to warty, brown conidia. Scale bars = 10 μm (from Crous 1998). is not congeneric with Lembosiniella eucalypti, which has hys-
terothecia opening with a longitudinal slit, and is pathogenic to
those of Lembosina contain a basal mucoid appendage (plug), Eucalyptus. Lembosiniella eucalyptorum [ascospores (26–)
which is absent in Lembosiniella. Lembosiniella eucalyptorum 30–35(–37) × 8–9(–10) μm] is similar to L. eucalypti (asco-
is common but not damaging on mature and senescing leaves spores 20–32 × 5.5–7.5 μm), although ascospores of
of E. dunnii and C. variegata in plantations in subtropical L. eucalyptorum are longer and wider (Sivanesan & Shivas
Australia. 2002c). The ITS sequence of Lembosiniella eucalyptorum is
not close to any sequence in GenBank, whereas the LSU
Lembosiniella eucalypti (Sivan. & R.G. Shivas) Crous, comb.
sequence is 95 % similar to those of species of Elsinoe, sug-
nov. MycoBank MB832020. Fig. 47.
gesting an affinity with Myrangiales.
Basionym: Lembosina eucalypti Sivan. & R.G. Shivas, Fungal
Diversity 11: 167. 2002. Macrohilum H.J. Swart, Trans. Brit. Mycol. Soc. 90: 288. 1988.
Diagnosis: Leaf spots amphigenous, dark brown to black, su-
perficial, irregular to subcircular, up to 8 mm diam. Hysterothecia Conidiomata immersed, becoming erumpent, medium brown,
linear to rarely Y-shaped, superficial, amphigenous, black, globose. Conidiogenous cells lining the inner cavity, pale brown,
300–450 × μm long, 75–115 μm high, opening by a central cylindrical, proliferating percurrently near the apex. Conidia sol-
longitudinal slit. Ascospores fusiform, hyaline to brown, 1-septate itary, medium to dark brown, ovoid, smooth, guttulate, developing
in the middle, constricted, smooth to echinulate, guttulate, a single supra-median septum, thick-walled, frequently con-
conglobate, 20–32 × 5.5–7.5 μm. stricted at the septum, apex obtuse, base truncate with a visible
scar.
Description and illustration: Sivanesan & Shivas (2002c).
Type species: Macrohilum eucalypti H.J. Swart
Typus: Australia, Queensland, Coen, on Eucalyptus sp., 18 Jul. Notes: Until recently, this monotypic genus has been obscure,
1999, R.G. Shivas & M. Gunther (holotype BRIP 25821). Not and little is known about its biology or pathogenicity (Crous et al.
known from culture. 2015a). It is generally regarded as a minor pathogen of
Lembosiniella eucalyptorum Crous & Carnegie, sp. nov. Eucalyptus.
MycoBank MB832021. Fig. 47. Macrohilum eucalypti H.J. Swart, Trans. Brit. Mycol. Soc. 90:
Etymology: Name refers to Eucalyptus, the host genus from 288. 1988. Fig. 48.
which this fungus was collected. Diagnosis: Leaf spots amphigenous, medium brown, subcircular,
Ascomata predominantly hypophyllous, black, superficial, hys- associated with leaf margins. Conidia solitary, medium to dark
terothecial, 150–350 μm diam, with feathery margin, opening by brown, ovoid, smooth, guttulate, developing a single, dark
longitudinal or Y-shaped split, margin with cells of textura epi- brown, supra-median septum, thickwalled, frequently constricted
dermoidea. Asci intermingled among pseudoparaphyses, hyaline, at the septum, apex obtuse, base truncate and protruding, with
septate, hypha-like, 3–4 μm diam, with obtuse ends. Asci obovoid a visible scar, 2–3 μm wide, (15–)17–19(–20) × (8–)
to ellipsoid, bitunicate, 8-spored, with apical chamber, 2–3 μm 10–12(–13) μm.
diam, 40–60 × 21–30 μm. Ascospores multiseriate in asci, fusoid-
Descriptions and illustrations: Swart (1988), Crous et al. (2015a).
ellipsoid, straight, ends obtuse, prominently constricted at
median septum, thick-walled, with 1–2 large guttules per cell, Typus: Australia, Victoria, Panton Hill, on living leaves of
hyaline, smooth, becoming brown and verruculose, (26–) E. polyanthemos, 27 Mar. 1971, H.J. Swart (holotype DAR
CROUS ET AL.
Fig. 47. Lembosiniella eucalypti (BRIP 25821). A. Disease symptoms. B. Surface view of ascomata. C. Vertical section through hysterothecium. D. Ascus. E. Ascospores.
Scale bars = 10 μm. F–M. L. eucalyptorum (CPC 31816). F, G. Disease symptoms. H. Hysterothecium. I. Colony on MEA. J, K. Asci. L. Ascospores. M. Germinating as-
cospores. Scale bars: H = 300 μm, all others = 10 μm.
59000); Northern Territories, Darwin, Kurrajong Heights, on thickened and darkened, microcyclic conidiation observed in
E. piperita, Apr. 2011, P.W. Crous (epitype CBS H-22279; cul- culture. Spermatogonia forming on OA. Spermatia cylindrical
ture ex-epitype CBS 140063 = CPC 19421). with obtuse ends, smooth, hyaline.
Madagascaromyces U. Braun et al., Stud. Mycol. 87: 376. Type species: Madagascaromyces intermedius (Crous & M.J.
2017. Wingf.) Videira & Crous (≡ Passalora intermedia Crous & M.J.
Wingf.)
Mycelium composed of pale to medium brown hyphae,
septate, branched, smooth, 2 – 3 μm. Conidiophores solitary, Notes: The genus Madagascaromyces is known only from
medium brown, smooth, subcylindrical, simple or branched, Madagascar and was probably introduced along with Eucalyptus
straight to variously curved or geniculate-sinuous. Con- material from Asia. Nothing is known about the biology of species
idiogenous cells terminal and intercalary, proliferating sym- in the genus.
podially, with one or multiple conidiogenous loci that are
thickened and darkened. Conidia solitary, pale brown, smooth, Madagascaromyces intermedius (Crous & M.J. Wingf.) Videira
guttulate, subcylindrical when small, narrowly obclavate when & Crous, Stud. Mycol. 87: 376. 2017. Fig. 49.
larger, apex subobtuse, base long obconically subtruncate, Basionym: Passalora intermedia Crous & M.J. Wingf., Persoonia
straight to slightly curved, 1- or multiseptate, with hila 22: 88. 2009.
190
Fig. 48. Macrohilum eucalypti (CPC 19421). A. Conidiomata on PNA. B. Conidiomata on OA. C–E. Conidiogenous cells. F. Conidia. Scale bars: A = 200 μm, B = 300 μm, all
others = 10 μm (from Crous et al. 2015a).
Fig. 49. Madagascaromyces intermedius (CPC 15745). A. Spermatogonium forming on OA. B. Spermatia. C–F. Conidiophores giving rise to conidia. G, H. Conidia. Scale
bars = 10 μm (from Crous et al. 2009b).
Diagnosis: Conidia solitary, pale brown, smooth, guttulate, sub- slightly curved, 1–8-septate, (35–)50–75(–100) × (2.5–)3 μm;
cylindrical when small, becoming narrowly obclavate when larger, hila thickened and darkened, 1–1.5 μm wide; microcyclic con-
apex subobtuse, base long obconically subtruncate, straight to idiation observed in culture.
CROUS ET AL.
Description and illustration: Crous et al. (2009b). Asci 70–90 × 6–7 μm, cylindrical to subsaccate, widest point in
lower third, base foot-like cell, apex rostrate, 8-spored, apex with
Typus: Madagascar, Morondavo, on E. calmadulensis, Aug. flat-tipped top, 1.5 μm diam. Ascospores (52–)
2007, M.J. Wingfield (holotype CBS H-20197, culture ex-type 62–65(–70) × 1.5–2 μm, tapering towards each end, straight to
CBS 124154 = CPC 15745). slightly curved, subcylindrical, medianly 1-septate, apex and
Marthamyces Minter, Mycotaxon 87: 50. 2003. base with mucoid caps, globose, 2–3 μm diam. Conidiomata
forming on OA: separate, pycnidial, globose, brown,
Ascomata apothecial, angular, immersed, opening with irregular 250–300 μm diam, opening by irregular rupture; wall of 3–6
flaps, upper layer poorly defined, with crystalline material layers of brown textura angularis. Conidiophores arising from
covering asci. Asci cylindrical, unitunicate, opening by central inner layer, hyaline, smooth, subcylindrical, branched, 1–8-
pore, 8-spored, with ascospores in fascicle. Ascospores hyaline, septate, 15–40 × 2.5–3 um. Conidiogenous cells sub-
filiform, thin-walled, smooth, transversely septate. Paraphyses cylindrical with apical taper, straight to flexuous, terminal and
slender, thin-walled, smooth, septate, often branched at the tips. intercalary, 9–12 × 1.5–2 μm, phialidic with periclinal thickening.
Type species: Marthamyces emarginatus (Cooke & Massee) Conidia aseptate, solitary, hyaline, smooth, subcylindrical,
Minter slightly curved, widest in middle, apex subobtuse, base truncate,
(11–)12–15(–17) × 1 μm.
Notes: The taxonomy of Marthamyces emarginatus (Martha-
mycetaceae) on Eucalyptus required clarification based on DNA Culture characteristics: Colonies flat, spreading, with moderate
data. Although this taxon has been reported from New Zealand aerial mycelium and smooth, lobate margin, reaching 40 mm
on Meterosideros, the specimens from New Zealand and diam after 2 wk at 25 °C. On MEA, PSA and OA surface and
Australia seem to represent different species (P. Johnston, pers. reverse dirty white to saffron.
comm.). Although ascomata are usually common on leaf litter,
they are also frequently associated with bleached (dirty grey- Typus: Australia, New South Wales, Billys Creek, Clouds Creek
white) lesions, suggesting they are of minor importance. Mar- State Forest, S30°6’17.3” E152°37’23.2, on E. dunnii, 21 Apr.
thamyces johnstonii is distinguished by having characteristic flat- 2018, A.J. Carnegie, HPC 2425 (holotype CBS H-24029, culture
tipped asci, and broad, clavate paraphyses. This species has ex-type CBS 145892 = CPC 35761).
also been identified from New Zealand (isolate AU97_7) (P. Notes: Based on a megablast search of NCBI’s GenBank
Johnston unpubl. data). nucleotide database, the closest hits using the ITS sequence had
Marthamyces emarginatus (Cooke & Massee) Minter, Myco- highest similarity to “Marthamyces sp. voucher PDD 81847”
taxon 87: 51. 2003. Fig. 50. (GenBank MK599212.1; Identities = 718/724 (99 %), no gaps),
Basionym: Stictis emarginata Cooke & Massee, in Cooke, Marthamyces emarginatus (GenBank MK599210.1; Identi-
Grevillea 18: 7. 1889. ties = 701/728 (96 %), 6 gaps (0 %)), and Marthamyces des-
Synonyms: Propolis emarginata (Cooke & Massee) Sherwood, moschoeni (GenBank KJ606679.1; Identities = 701/729 (96 %),
Mycotaxon 5: 323. 1977. 7 gaps (0 %)). Closest hits using the LSU sequence are “Mar-
Coccomyces martinae Hansf., Proc. Linn. Soc. N.S.W. 81: 40. thamyces sp. voucher PDD 81847” (GenBank MK599204.1;
1956. Identities = 958/958 (100 %), no gaps), Marthamyces emargi-
natus (GenBank MK599203.1; Identities = 953/959 (99 %), 1 gap
Diagnosis: Usually saprobic (endophytic, sporulating on dead (0 %)), and Coccomyces proteae (GenBank JN712515.1; Iden-
tissue), but occasionally associated with small, circular to irreg- tities = 953/959 (99 %), 1 gap (0 %)).
ular, bleached lesions; widespread in occurrence. Ascospores
55–70 × 1.5 μm, hyaline, filiform, thin-walled, smooth, trans- Microthyrium Desm., Annls Sci. Nat., Bot., ser. 2 15: 137. 1841.
versely septate. Paraphyses slender, thin-walled, 1.5–2 μm Synonyms: Calothyriella Höhn., Ber. dt. bot. Ges. 35: 251. 1917.
diam, smooth, often branched at the tips. Microthyrina Bat., Publç~oes Inst. Micol. Recife 260: 51. 1960.
Phragmothyrium Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Description: Sherwood (1977), Minter (2003). Kl., Abt. 1 121: 347 (9 repr.). 1912.
Typus: Australia, Victoria, on Eucalyptus sp., Mrs Martin 439
Mycelium superficial, abundant, comprising almost colourless,
(holotype of Stictis emarginata in K – not seen).
branched, septate hyphae, with subcuticular haustoria. Thyro-
Marthamyces johnstonii Crous & Carnegie, sp. nov. Myco- thecia circular, solitary or gregarious, superficial, membrana-
Bank MB832022. Fig. 50. ceous or slightly carbonaceous, pale brown to brown, with a
poorly developed basal layer, easily separated from host surface,
Etymology: Named in honour of the New Zealand mycologist,
with a prominent darker central ostiole; in section lenticular.
Peter R. Johnston, who revised Marthamyces and related
Upper wall of textura epidermoidea, radiating outwardly in par-
genera, bringing clarity to the Leotiomycetes.
allel rows from the darker central ostiole. Peridium comprising a
Ascomata on leaf litter, not associated with leaf spots; ascomata single stratum of cells, outer cells of small heavily pigmented
150–300 μm diam, round in outline, immersed without margin, cells of textura epidermoidea, inner cells pale brown textura
concolourous, becoming raised at maturity, opening by radiate angularis. Hamathecium comprising asci inclined from the base
slits, flaps folding back to expose crystalline hymenium; surface and rim towards the central ostiole. Pseudoparaphyses not
with scattered white crystals; walls of pale brown cells, becoming observed. Asci numerous, 8-spored, bitunicate, fissitunicate,
hyaline towards inner region. Paraphyses 3–4 μm diam near narrowly obclavate to long fusiform, with small knob-like pedicel
base, branched at base or not, becoming clavate towards thick- or pedicel lacking, ocular chamber not apparent, not staining blue
walled apex, 3–6 μm diam, 5–7-septate, extending above asci. in IKI. Ascospores overlapping 2–3-seriate, fusoid to ellipsoidal,
192
Fig. 50. Marthamyces emarginatus (VPRI 22068). A, B. Disease symptoms. C. Leaf spot with ascomata. D. Vertical section through ascoma. E–K. Asci with ascospores.
Marthamyces johnstonii (CPC 35761). L. Ascomata on leaf tissue. M, N. Asci with ascospores. O. Ascospores. P. Conidiomata on OA. Q, R. Conidiophores giving rise to
conidia. S. Conidia. Scale bars: C, L, P = 300 μm, D = 150 μm, all others = 10 μm.
CROUS ET AL.
hyaline, 1-septate, fine appendages sometimes present, smooth- aparaphysate, fasciculate, bitunicate, subsessile, obovoid,
walled. straight to slightly curved, 8-spored. Ascospores multiseriate,
overlapping, hyaline, guttulate, thin-walled, straight to slightly
Type species: Microthyrium microscopicum Desm.
curved, fusoid-ellipsoidal with obtuse ends, widest in middle of
Notes: Although obscure, a similar fungus was found to be a apical cell, medianly 1-septate.
pathogen of older leaves of E. globulus plantations in Victoria,
Australia (Barber 1998). Little information is available and this Type species: Mycodiella eucalypti Crous
report can only be confirmed when fresh collections become Notes: Mycodiella can be distinguished from other
available. Swart (1986c) noted that M. eucalypticola (sporulates mycosphaerella-like genera only based on DNA data, as the
on dead leaves) resembles Phaeothyriolum microthyrioides species are morphologically very similar, and lack asexual
(sporocarps on living leaves), which is a foliar pathogen of morphs. Other than being found on Eucalyptus foliage, little is
Eucalyptus (see below). known about the biology of Mycodiella.
Microthyrium eucalypticola Speg., Anal. Mus. nac. B. Aires, Mycodiella eucalypti Crous, Persoonia 37: 337. 2016. Fig. 52.
Ser. 3 17: 426. 1908.
Diagnosis: Leaf spots amphigenous, irregular, pale brown, with
Diagnosis: Leaf spots circular, chlorotic, up to 30 mm diam. raised dark brown border. Ascospores multiseriate, overlapping,
Thyrothecia form on older or fallen leaves and are arranged in hyaline, guttulate, thin-walled, straight to slightly curved, fusoid-
concentric circles on chlorotic zones; assumed to be weakly ellipsoidal with obtuse ends, widest in middle of apical cell,
pathogenic, appearing as very small black spots. Thyrothecia medianly 1-septate, not constricted at the septum, tapering to-
superficial, flat, conical with circular ostiole and irregular margin, wards both ends, but more prominently towards lower end, (11–)
lacking superficial mycelium. Ascospores 2-celled, hyaline, 12–13(–15) × (2.5–)3(–3.5) μm.
fusoid, straight to curved, constricted at septum, with gelatinous
sheath, 12–14 × 3 μm (ascospore dimensions from Spegazzini Description and illustration: Crous et al. (2016b).
1909; specimen not seen). Typus: Australia, Western Australia, Porongurup, Porongurup
Description: Adapted from Park et al. (2000). National Park, S34°41'18.6" E117°55'56", on E. diversicolor, 24
Sep. 2015, P.W. Crous (holotype CBS H-22885, cultures ex-
Typus: Argentina, Buenos Aires, La Plata, on E. globulus, May type CBS 142097 = CPC 29226, CBS 142098 = CPC 29458).
1906, collector unknown (holotype LPS 1429 – not seen).
Mycodiella sumatrensis (Crous & M.J. Wingf.) Crous, Per-
Mulderomyces Crous et al., Persoonia 36: 359. 2016. soonia 37: 337. 2016.
Conidiomata pycnidial, solitary, pale brown, erumpent, globose, Basionym: Mycosphaerella sumatrensis Crous & M.J. Wingf.,
with central ostiole; wall of 6–8 layers of subhyaline to pale brown Stud. Mycol. 55: 124. 2006.
textura angularis. Conidiophores lining the inner cavity, hyaline, Diagnosis: Leaf spots amphigenous, irregular to subcircular,
smooth, subcylindrical, septate, branched. Conidiogenous cells 2–10 mm diam, pale brown with a dark brown, raised border,
hyaline, smooth, subcylindrical, terminal and lateral; proliferating and thin, red-purple margin. Ascospores multiseriate, over-
sympodially, scars inconspicuous. Conidia cylindrical, hyaline, lapping, hyaline, guttulate, thin-walled, straight,
smooth, guttulate, straight with subobtuse ends, 2–6-septate, fusoid–ellipsoidal with obtuse ends, medianly 1-septate, widest
prominently constricted at septa (cells linked by a narrow in middle of apical cell, not constricted at the septum, tapering
isthmus), with mature conidia breaking into phragmospores. towards both ends, but more prominently towards the lower end,
Type species: Mulderomyces natalis Crous et al. (12–)13–15(–16) × (3–)4 μm, in vivo.
Note: Nothing is known regarding the biology of Mulderomyces Description and illustration: Crous et al. (2006d).
natalis, which appears to be a minor foliar pathogen of Typus: Indonesia, Northern Sumatra, on Eucalyptus sp., Feb.
Eucalyptus. 2004, M.J. Wingfield (holotype CBS H-19704, culture ex-type
Mulderomyces natalis Crous et al., Persoonia 36: 359. 2016. CBS 118499 = CPC 11171).
Fig. 51. €
Mycosphaerella Johanson, Ofvers. K. Svensk. Vetensk.-Akad.
Diagnosis: Associated with large, pale brown, amphigenous leaf Förhandl. 41(no. 9): 163. 1884 (1884–1885).
spots. Conidia cylindrical, hyaline, smooth, guttulate, straight with = Ramularia Unger (see below)
subobtuse ends, 2–6-septate, prominently constricted at septa
(cells linked by a narrow isthmus), with mature conidia breaking Notes: When Crous (1998) revised this group of foliar patho-
into phragmospores, (22–)50–75(–90) × (2–)3 μm. gens on Eucalyptus, they were all treated under the aggregate
genus “Mycosphaerella”. However, Mycosphaerella has since
Description and illustration: Crous et al. (2016c). been reduced to synonymy under Ramularia (Rossman et al.
Typus: Australia, Victoria, Melbourne, Moonee Ponds Creek, on 2015, Videira et al. 2015a, b, 2016), and many species origi-
Eucalyptus sp., 2 Nov. 2014, P.W. Crous, J. Edwards & P.W.J. nally named in Mycosphaerella have been relocated else-
Taylor (holotype CBS H-22609, culture ex-type CBS where, e.g. the Dissoconiaceae or Teratosphaeriaceae
141296 = CPC 25519). (Quaedvlieg et al. 2014). The species listed below lack cul-
tures or DNA data, and thus cannot safely be allocated to
Mycodiella Crous, Persoonia 37: 337. 2016. another genus. Therefore, pending fresh collections needed to
Ascomata pseudothecial, brown, erumpent, globose; wall con- resolve their taxonomy, they are still listed here under their
sisting of 2–3 layers of medium brown textura angularis. Asci original names.
194
Fig. 51. Mulderomyces natalis (CPC 25519). A, B. Disease symptoms. C. Conidiomata sporulating on MEA. D–F. Conidiogenous cells giving rise to conidia. G, H. Conidia.
Scale bars = 10 μm.
Fig. 52. Mycodiella eucalypti (CPC 29226). A. Leaf spot. B, C. Asci. D. Ascospores. E. Germinating ascospores. Scale bars = 10 μm.
“Mycosphaerella” longibasalis Crous & M.J. Wingf., Mycol. Note: Although no cultures could be obtained for this pathogen, it
Mem. 21: 79. 1998. is presumed to be a species of Teratosphaeria based on the size
of its ascospores, and general morphological characteristics.
diam, pale brown, surrounded by a raised, medium brown “Mycosphaerella” eucalypti (Wakef.) Hansf., Proc. Linn. Soc.
border; coloured margin absent. Ascospores multiseriate, over- N.S.W. 82: 218. 1957.
lapping, colourless, guttulate, thin-walled, straight to slightly Basionym: Hypospila eucalypti Wakef., Bull. Misc. Inf., Kew(4):
curved, fusoid-ellipsoidal, base obtuse, apex subobtuse, widest 190. 1912.
above middle of apical cell, unequally 1-septate, not constricted
Diagnosis: Leaf spots amphigenous, but not extending
at septum, tapering toward both ends, but with slightly more so
through the leaf lamina, circular, raised, appearing warty,
toward base (22–)23–25(–30) × 3.5–4(–5) μm, apical cell
medium brown, 3 –4 mm diam; border irregular, margin absent,
9–11 μm long, basal cell 13–15 μm long.
or thin, red-brown and inconspicuous. Ascospores multi-
Description and illustration: Crous (1998). seriate, overlapping, hyaline, appearing olivaceous in some
asci, guttulate, thick-walled, straight to curved, obovoid to
Typus: Colombia, La Selva, on E. grandis, Jun. 1994, M.J.
subcylindrical with obtuse ends, widest near apex, medianly
Wingfield (holotype PREM 54403).
1-septate, becoming constricted at septum, tapering toward
Additional material examined: Colombia, Sinai, on E. grandis, Jun. 1995, M.J. both ends, but more prominently toward base, (36– )
Wingfield, specimen PREM 54404. 38 –52( – 60) × 6( – 7) μm.
CROUS ET AL.
Description and illustration: Crous (1998). Note: Mycosphaerelloides is currently a monotypic genus known
from Madeira and the Netherlands and appears to not be specific
Typus: Australia, Queensland, Brisbane, on Eucalyptus sp.,
to Eucalyptus.
1912, F.M. Bailey (holotype K). Not known from culture.
Additional material examined: Australia, Queensland, Brisbane, on Eucalyptus
Mycosphaerelloides madeirae (Crous & Denman) Videira &
sp., 31 Jan. 1913, C.T. White, BRIP 125a. Crous, Stud. Mycol. 83: 100. 2016.
Basionym: Mycosphaerella madeirae Crous & Denman, Stud.
Notes: Mycosphaerella eucalypti is morphologically similar to Mycol. 50: 204. 2004.
several other large-spored species of Teratosphaeria, and would
Synonym: Paramycosphaerella madeirae (Crous & Denman)
better be accommodated in the latter genus. However, fresh
Guatimosim et al., Persoonia 37: 127. 2016, as “madeirensis”.
collections are required to resolve its phylogeny.
Diagnosis: Leaf spots amphigenous, subcircular, 2–15 mm diam,
Mycosphaerelloides Videira & Crous, Stud. Mycol. 83: 99.
medium brown, surrounded by a slightly raised, red-purple border.
2016.
Ascospores 3- to multiseptate, overlapping, hyaline, guttulate, thin-
Leaf spots amphigenous, subcircular, 2–15 mm diam, medium walled, straight to slightly curved, fusoid-ellipsoid with subobtuse
brown, surrounded by a slightly raised, red-purple border. Asco- ends, apex frequently acutely rounded, medianly 1-septate, widest
mata pseudothecial, predominantly epiphyllous, single, black, in the middle of the apical cell, not constricted at the septum,
immersed, becoming erumpent, globose, up to 120 μm diam; tapering towards both ends, but more prominently towards the
apical ostiole 10–15 μm diam; wall of 2–3 layers of medium brown lower end, (9–)10–13(–15) × 2.5–3(–3.5) μm in vivo.
textura angularis. Asci aparaphysate, fasciculate, bitunicate, sub-
sessile, obovoid to narrowly ellipsoid, straight or slightly incurved, 8-
spored, 30–50 × 8–12 μm. Ascospores 3- to multiseriate, over- Typus: Portugal, Madeira, Party Farm, on E. globulus, Apr.
lapping, hyaline, guttulate, thin-walled, straight to slightly curved, 2000, S. Denman (holotype CBS H-9898, cultures ex-type CBS
fusoid-ellipsoid with subobtuse ends, apex frequently acutely 112895 = CMW 14458 = CPC 3745, CBS 112301 = CPC 3747).
rounded, medianly 1-septate, widest in the middle of the apical cell,
Neoceratosperma Crous & Cheew., Persoonia 32: 255. 2014.
not constricted at the septum, tapering towards both ends, but
more prominently towards the lower end, (9–) Mycelium consisting of branched, septate, brown, verruculose,
10–13(–15) × 2.5–3(–3.5) μm in vivo. Mycelium internal and hyphae that turn warty with age. Conidiophores reduced to
external, consisting of smooth, branched, septate, pale to medium conidiogenous cells, or septate, erect, brown, verruculose, un-
brown, 3–6 μm wide hyphae; external mycelium extensive on branched, subcylindrical, dark brown and smooth at the base.
abaxial leaf surface. Conidiomata fasciculate, hypophyllous, me- Conidiogenous cells subcylindrical, brown, verruculose, but
dium brown, up to 90 μm wide and 150 μm high. Conidiophores conidiogenous apical area smooth, forming a short rachis that
arising from superficial mycelium, or aggregated in loose fascicles proliferates sympodially; loci somewhat thickened and darkened.
arising from the upper cells of a brown stroma up to 80 μm wide Conidia solitary, rarely in unbranched chains, subcylindrical,
and 90 μm high; conidiophores pale to medium brown, smooth, medium brown, becoming dark brown, verruculose, becoming
unbranched or branched, 1–5-septate, subcylindrical, straight to warty, distoseptate, less obvious when older (dark brown, warty),
variously curved, 15–45 × 2.5–4 μm; conidiogenous cells terminal straight to irregularly curved; apex obtuse, base truncate, but
or lateral, unbranched, subcylindrical, pale brown, smooth, prolif- scars somewhat thickened and darkened.
erating sympodially, or 1–4 times percurrently near apex,
Type species: Neoceratosperma eucalypti Crous & Cheew.
7–15 × 2.5–3 μm; conidiogenous loci inconspicuous. Conidia
solitary, pale brown, smooth, subcylindrical, but tapering from a Notes: Neoceratosperma is a zasmidium-like genus associated
subtruncate base towards a subobtuse apex, 3–6- or multiseptate, with leaf spots of different host plants, including Eucalyptus. It
35–85 × 2.5–4 μm, hila neither thickened nor darkened-refractive. appears to be of minor importance as a pathogen.
Type species: Mycosphaerelloides madeirae (Crous & Denman) Neoceratosperma eucalypti Crous & Cheew., Persoonia 32:
Videira & Crous (≡ Mycosphaerella madeirae) 257. 2014. Fig. 53.
Fig. 53. Neoceratosperma eucalypti (CPC 23465). A. Conidiophores in culture. B. Conidiogenous cell giving rise to conidia. C–E. Conidia. Scale bars = 10 μm.
196
Diagnosis: Leaf spots spreading inward from the leaf margins, Etymology: Name reflects the host species it was collected from,
irregular, medium brown, surrounded by thick, dark brown E. corticosa.
border. Conidia solitary, rarely in unbranched chains, sub-
Diagnosis: Conidia hyaline, smooth, thin-walled, granular,
cylindrical, medium brown, becoming dark brown, verruculose,
ellipsoidal, apex obtuse, base subtruncate, somewhat flattened,
becoming warty, 1–7-disto-septate, less obvious when older
at times with minute marginal frill, (8 –)15–17( –18) × (7–)8 μm.
(dark brown, warty), straight to irregularly curved,
In vitro producing brown Dichomera conidiomata up to
40–150(–200) × 3–4 μm; apex obtuse, base truncate, but scars
200 μm diam. Conidia globose, obpyriform or obovoid,
somewhat thickened and darkened, 1–1.5 μm diam.
brown at maturity, smooth, thin-walled, with 1–2 transverse and
Description and illustration: Crous et al. (2014b). 1–3 vertical or oblique septa, (10–)13–14( –15) × (9–)
10(–11) μm.
Typus: Thailand, Chiang Mai, on Eucalyptus sp., Sep. 2013, R.
Cheewangkoon (holotype CBS H-21712, culture ex-type CBS Description and illustration: Summerell et al. (2006).
137998 = CPC 23465).
Typus: Australia, New South Wales, Wollemi National Park, on
Neoceratosperma yunnanensis (Barber et al.) Guatimosim E. corticosa, 3.5 km E of final turnoff to Dunn’s Swamp (ca. 5 km
et al., Persoonia 37: 123. 2016. E of Currant Mountain Gap), Central Tablelands NSW,
Basionym: Mycosphaerella yunnanensis Barber et al., Fungal S32°51’28”, E150°14’11”, alt. 740 m, Feb. 2006, B. Summerell
Diversity 24: 150. 2007. (holotype designated here CBS H-19745, cultures ex-type CBS
Synonym: Xenomycosphaerella yunnanensis (Barber et al.) 120081 = CPC 12925, CPC 12926, 12927).
Quaedvl. & Crous, Persoonia 33: 24. 2014.
Notes: Phillips et al. (2013) regarded this species as a synonym
Diagnosis: Leaf spots spreading inward from the leaf margins, of N. vitifusiforme, but lacked tef1 sequence data for a detailed
irregular, up to 14 mm wide and 140 mm long, medium brown, comparison. These data were generated by Yang et al. (2017),
surrounded by thick, dark brown border. Ascospores bi- to showing them closely related, but distinct. In the original publi-
triseriate, overlapping, hyaline, non-guttulate, thick-walled, cation (Summerell et al. 2006), the holotype specimen was not
straight to slightly curved, fusoid-ellipsoidal with obtuse ends, indicated as such, rendering the species invalid (Art 40.6, Mel-
medianly or unequally 1-septate, widest at the middle or near bourne code). It is validated here.
the apex of the apical cell, not or slightly constricted at the
Neofusicoccum eucalyptorum (Crous et al.) Crous et al., Stud.
septum, tapering towards both ends but more prominently to-
Mycol. 55: 248. 2006.
wards lower end, (9.5 –)10–12.5 × (2 –)2.5–3(–3.5) μm
Basionym: Fusicoccum eucalyptorum Crous et al., Mycologia 93:
(av. = 11.2 × 2.7 μm).
280. 2001.
Synonyms: Phoma australis Cooke, Grevillea 15: 17. 1886.
Description and illustration: Burgess et al. (2007).
Idiocercus australis (Cooke) H.J. Swart, Trans. Brit. Mycol. Soc.
Typus: China, Yunnan, Lancang, on E. urophylla, May 2005, B. 90: 283. 1988.
Dell (holotype MURU 407, culture ex-type CBS 119975 = CMW Botryosphaeria eucalyptorum Crous et al., Mycologia 93: 280.
23443). 2001.
Neofusicoccum Crous et al., Stud. Mycol. 55: 247. 2006. Diagnosis: Ascospores irregularly biseriate, hyaline, aseptate,
granular, becoming pale brown with age, (20–)23–26(–28) × (7–)
Resembling species of Fusicoccum, but distinct in forming a
8–9(–11) μm, prominently inequilateral when young, less so when
dichomera-like synasexual morph with brown, globose to pyri-
mature, fusoid, widest in the middle, base obtuse, apex obtuse to
form conidia.
subobtuse. Conidia hyaline, granular, ovoid to slightly clavate, apex
Type species: Neofusicoccum parvum (Pennycook & Samuels) obtuse, tapering towards a subtruncate or bluntly rounded base,
Crous et al. sometimes with a minute marginal frill visible on younger conidia,
(20–)22–25(–28) × (6–)7–8(–9) μm in vivo, (18–)
Symptoms: Leaf spots brown, amphigenous, irregular, raised, at
20–23(–25) × 7–8(–12) μm in vitro.
times with a thin, red-purple to chlorotic margin.
Description and illustration: Smith et al. (2001).
Notes: Phillips et al. (2013) showed that isolates identified by
Barber et al. (2005) were conspecific with N. vitifusiforme. The Typus: South Africa, Mpumalanga, Sabie, on E. grandis, 1995,
epithet “eucalypti” predates that of “vitifusiforme”. However, as H. Smith (holotype PREM 56604); representative culture CBS
most species of Neofusicoccum form Dichomera synasexual 115791 = CMW10125.
morphs, and the fact that the isolate from Barber et al. (2005)
Neofusicoccum ribis (Slippers et al.) Crous et al., Stud. Mycol.
was not ex-type, we will refrain from introducing a new
55: 249. 2006.
combination.
Basionym: Fusicoccum ribis Slippers et al., Mycologia 96: 96.
Species of Neofusicoccum most commonly occur as endo-
2004.
phytes in Eucalyptus leaves and twigs, causing stem cankers
Synonym: Botryosphaeria ribis Grossenb. & Duggar, Tech. Bull.
and dieback. However, under certain conditions, some of them
N.Y. Agric. Exp. St. 18: 128. 1911.
are associated with leaf spots.
Neofusicoccum corticosae Crous & Summerell, sp. nov. Diagnosis: Ascospores fusoid to ellipsoid, often round at the
MycoBank MB832023. Fig. 54. ends then broadly ellipsoidal, hyaline, unicellular, smooth
Synonym: Neofusicoccum corticosae Crous & Summerell, with granular contents, biseriate in the ascus, (14–)
Fungal Diversity 23: 337. 2006. nom. inval. Art 40.6. 18–23(–27) × 6–8(–10) μm (av. 20.5 × 7.1 μm), L/W = 2.9.
CROUS ET AL.
Fig. 54. Neofusicoccum spp. A–G. N. corticosae (CPC 12925). A. Leaf spot. B, C. Conidiogenous cells. D. Conidia. E. Conidiogenous cells giving rise to dichomera-like
conidia. F, G. Dichomera-like conidia. H–J. N. vitifusiforme (CBS H-7756). H. Sporulation on PNA. I. Conidiogenous cells. J. Conidia. Scale bars = 10 μm.
Conidia subglobose, obpyriform or rarely obovoid to broadly longitudinal septa, becoming brown when mature with 1–4
fusiform or fusiform, apex subobtuse to obtuse, base truncate to transverse septa, 0–3 longitudinal septa, and 0–4 oblique
bluntly rounded. Subglobose, obpyriform conidia (7–) septa. Broadly fusiform to fusiform conidia (12–)
8–13.5(–17) × (6.5–)7–9.5(–10.5) μm, hyaline to pale brown 13.5–22.5(–24) × (5–)5.5–8 μm, brown with 2–7 transverse
when immature with one transverse septum and 0–2 septa, and 0–2 oblique septa.
198
Description and illustration: Phillips et al. (2013). Conidiomata acervular or pycnidial, subglobose, globose, clavate,
solitary or aggregated, dark brown to black, immersed to erum-
Typus: Of asexual morph; USA, New York, Ithaca, on Ribes
pent, unilocular or irregularly plurilocular; exuding dark brown to
sp., 2000, G. Hudler, (holotype of asexual morph PREM
black conidia in a slimy, globose mass. Conidiophores indistinct,
57368, ex-type culture CBS 115475 = CMW 7772); USA, New
often reduced to conidiogenous cells. Conidiogenous cells
York, Geneva, on Ribes vulgare, 1911, J.G. Grossenbacher &
discrete, cylindrical, ampulliform to lageniform, hyaline, smooth,
B.M. Duggar (lectotype of sexual morph CUP-A (F.Col.
thin-walled; conidiogenesis initially holoblastic, becoming percur-
3408)).
rent to produce additional conidia at slightly higher levels. Conidia
fusoid, ellipsoid to subcylindrical, straight to slightly curved, 4-
Neofusicoccum versiforme (Z.Q. Yuan et al.) Crous, comb.
septate; basal cell conic to subcylindrical, with a truncate base,
nov. MycoBank MB832024.
hyaline or pale brown to olivaceous, thin and rugose to smooth-
Basionym: Dichomera versiformis Z.Q. Yuan et al., Nova Hed-
walled; three median cells doliiform, wall rugose to verruculose,
wigia 70: 140. 2000.
versicoloured, septa darker than the rest of the cell; apical cell
Diagnosis: Conidia in vitro variable, subglobose, obovoid or hyaline, conic to cylindrical, thin- and smooth-walled; with tubular
obpyriform to ellipsoidal or broadly fusiform, apex subobtuse to apical appendages, one to many, filiform or attenuated, flexuous,
obtuse, base truncate to bluntly rounded. Subglobose, obovoid branched or unbranched; basal appendage single, tubular, un-
or obpyriform conidia 8–17(–19) × 6.5–9.5 μm branched, centric (from Maharachchikumbura et al. 2014).
(av. = 13.6 × 9.3 μm), hyaline to pale brown when immature
Type species: Neopestalotiopsis protearum (Crous & L. Swart)
becoming brown when mature, with 0–3 transverse septa, 0–1
Maharachch. et al.
longitudinal and 0–1 oblique septae. Ellipsoidal and broadly
fusiform conidia 16–22(–25) × (5–)5.5–6.5(–8) μm Note: Based on a recent taxonomy presented for pestalotiopsis-
(av. = 19.3 × 5.9 μm), hyaline to pale brown when immature with like fungi (Maharachchikumbura et al. 2014), previous reports all
0–4 transverse septa and no longitudinal or oblique septa need to be reconfirmed.
becoming brown and muriform when mature with 4–5 transverse
Neopestalotiopsis eucalypticola Maharachch. et al., Stud.
septa and 0–2 oblique septa.
Mycol. 79: 138. 2014. Fig. 55.
Description and illustration: Barber et al. (2005).
Diagnosis: Water-soaked or pale brown leaf spots. Conidia
Typus: Australia, Victoria: Nareen, on E. camaldulensis, 7 Sep. fusoid, ellipsoid, straight to slightly curved, 4-septate, (22–)
1999, P.A. Barber (epitype VPRI 31989, ex-epitype culture 23–30(–31) × (9–)7.5–9(–9.5) μm; basal cell conic to obconic
WAC12403); Tasmania, Smithton, on E. nitens, 27 Aug.1998, with a truncate base, hyaline, rugose and thin-walled, 5–7 μm
Z.Q. Yuan & T. Wardlaw (holotype VPRI 22038a). long; three median cells doliiform, (15.5–)16–19.5(–20) μm
long, wall rugose, versicoloured, septa darker than the rest of the
Note: Slippers et al. (2013) regarded this species as a member of
cell (second cell from the base pale brown, 5–7 μm long; third
Neofusicoccum but did not formally introduce a new combination.
cell darker brown, 4.5–7.5 μm long; fourth cell darker brown,
Neofusicoccum vitifusiforme (Van Niekerk & Crous) Crous 5–7 μm long); apical cell 4.5–7.5 μm long, hyaline, cylindrical to
et al., Stud. Mycol. 55: 249. 2006. Fig. 54. subcylindrical, rugose and thin-walled; with 1–2 tubular apical
Basionym: Fusicoccum vitifusiforme Van Niekerk & Crous, appendages, arising as an extension of the apical cell, un-
Mycologia 96: 793. 2004. branched, attenuated, flexuous, (20–)32–55(–66) μm long;
Synasexual morph: Dichomera eucalypti (G. Winter) B. Sutton, basal appendage single, tubular, unbranched, centric, 6–11 μm
Mycol. Pap. 138: 182. 1975. long.
Basionym: Camarosporium eucalypti G. Winter, Revue Mycol.,
Description and illustration: Maharachchikumbura et al. (2014).
Toulouse 8 (32): 212. 1886.
Typus: Unknown country, on E. globulus, Jun. 1937, H.W.
Diagnosis: Conidia hyaline, granular, fusoid to ellipsoid, widest in
Wollenweber (holotype CBS H-15658, ex-type culture CBS
the upper third with an obtuse apex and flattened, subtruncate
264.37 = BBA 5300).
base, (18–)19–21(–22) × (4.5–)5.5–6.5(–8) μm in vitro, L/W
ratio = 3.3. Dichomera synasexual morph: Conidia subglobose, Neophysalospora Crous & M.J. Wingf., Persoonia 33: 247.
obpyriform or obovoid, apex obtuse, base truncate to bluntly 2014.
rounded, (9–)9.5–13(–14.5) × (6.5–)8–10.5(–11) μm, hyaline
Ascomata globose, solitary, brown, immersed, with central
to pale brown when immature with 0–3 transverse septa, 0–2
ostiole; wall of 2–3 layers of brown textura angularis. Asci cy-
longitudinal septa, and 0–2 oblique septa, becoming brown
lindrical, hyaline, stipitate, unitunicate with apical mechanism
when mature with 1–3 transverse septa, 0–3 longitudinal septa,
staining in Melzer’s reagent, ascospores uniseriate but over-
and 0–2 oblique septa.
lapping, with 8 ascospores per ascus. Paraphyses intermingled
Descriptions and illustrations: Barber et al. (2005), Phillips et al. among asci, hyaline, septate, branched. Ascospores hyaline,
(2013). smooth, guttulate, fusoid-ellipsoid, with acutely rounded ends.
Conidiomata globose, solitary to aggregated, brown, with central
Typus: South Africa, Western Cape Province, Stellenbosch, on
ostiole; wall of 2–3 layers of brown textura angularis. Co-
Vitis vinifera, 2002, J.M. van Niekerk (holotype of Fusicoccum
nidiophores lining the inner conidiomatal wall, subcylindrical,
vitifusiforme CBS H-7756, culture ex-type CBS 110887 = CPC
straight to curved, branched, septate, or reduced to con-
5252).
idiogenous cells, hyaline to pale brown, smooth or finely verru-
Neopestalotiopsis Maharachch. et al., Stud. Mycol. 79: 135. culose. Conidiogenous cells ampulliform to subcylindrical,
2014. hyaline, smooth, terminal or intercalary, phialidic with flared
CROUS ET AL.
Fig. 55. Neopestalotiopsis eucalypticola (CBS 264.37) A. Conidiomata sporulating on PNA. B. Conidiomata on PDA. C–E. Conidiogenous cells. F–K. Conidia. Scale
bars = 10 μm (from Maharachchikumbura et al. 2014).
collarette at the apex. Conidia solitary, hyaline, smooth, sub- Description and illustration: Crous et al. (2014c).
cylindrical, curved, with obtuse apex and truncate base, thick-
Typus: Mozambique, on C. henryi, 1 Feb. 2014, M.J. Wingfield
walled.
Type species: Neophysalospora eucalypti Crous & M.J. Wingf. 24209).
Notes: Neophysalospora is reminiscent of Clypeophysalospora, Neosonderhenia Crous, gen. nov. MycoBank MB832025.
and the two genera are easily confused. The fungus has been
collected from leaf litter but has also been associated with cutting Etymology: Name reflects its similarity to the genus Sonderhenia.
rot and prominent leaf spot symptoms. Leaf spots amphigenous, pale brown, circular, 2–3 mm diam,
Neophysalospora eucalypti Crous & M.J. Wingf., Persoonia red-purple margin. Ascomata pseudothecial, immersed, with
33: 247. 2014. Fig. 56. central ostiole; wall of 3–6 layers of brown textura angularis. Asci
8-spored, bitunicate, sessile, fasciculate with small ocular
Diagnosis: Associated with brown leaf spots in plantations chamber. Ascospores hyaline, smooth, guttulate, median
and cutting rot in nurseries. Ascospores hyaline, smooth, septate. Conidiomata globose, immersed, subepidermal with
guttulate, fusoid-ellipsoid, widest in middle, tapering to acutely central ostiole, brown; wall of 3–6 layers of brown textura
rounded ends, (13– )15 –17( – 19) × (4– )4.5( – 5) μm. Conidia angularis. Conidiophores reduced to inconspicuous con-
solitary, hyaline, smooth, subcylindrical, curved, with obtuse idiogenous cells lining the cavity, doliiform, pale brown, prolifer-
apex and truncate base, thick-walled, (13 –)14– 15( –16) × ating percurrently. Paraphyses hyaline, subcylindrical, sparsely
1.5 μm. septate. Conidia solitary, medium brown, verrcuculose,
200
Fig. 56. Neophysalospora eucalypti (CPC 24209). A. Leaf spots. B. Conidiomata on SNA. C. Conidiogenous cells. D. Conidia. E. Asci with ascospores. F. Germinating
ascospores. Scale bars = 10 μm.
distoseptate with central pore at each septum, apex subobtuse, Although all species of Neosonderhenia have conidiomatal pa-
base truncate. raphyses, this feature is not definitive, as some species of
Sonderhenia (e.g. S. radiata) also exibit this feature. These
Type species: Neosonderhenia eucalypti Crous
genera are thus best distinguished based on their DNA
Notes: Neosonderhenia has pycnidial conidiomata, distoseptate phylogeny.
conidia with a central pore, brown, percurrently proliferating
Neosonderhenia eucalypti Crous, sp. nov. MycoBank
conidiogenous cells, and a teratosphaeria-like sexual morph.
MB832026. Fig. 57.
Fig. 57. Neosonderhenia spp. A–D. N. eucalypti (CBS 145081). A. Leaf spot. B–D. Conidiogenous cells and conidia. E–H. N. foliorum [K(M) 251545]. E. Ascus with as-
cospores. F. Transverse section through conidioma. G. Conidiogenous cells. H. Conidia. Scale bars: F = 250 μm, all others = 10 μm.
CROUS ET AL.
Etymology: Name refers to Eucalyptus, the host genus from Leaf spots amphigenous, pale brown, circular, 2–3 mm diam,
which this fungus was isolated. with immersed, slightly erumpent, black, separate sporocarps
representing conidiomata or ascomata. Ascomata pseudothecial,
Leaf spots dark brown, amphigenous, subcircular, 1–3 mm diam,
100–250 μm diam, with central ostiole; wall of 3–6 layers of
with prominent red-purple margin. Conidiomata separate, pycnid-
brown textura angularis. Asci 8-spored, bitunicate, sessile,
ial, brown, globose, 200–350 μm diam with central ostiole; wall of
fasciculate with small ocular chamber, 1–2 μm diam,
6–8 layers of brown textura angularis. Conidiophores reduced to
50–80 × 19–22 μm. Ascospores multiseriate, hyaline, smooth,
conidiogenous cells lining the inner cavity, hyaline to pale brown,
guttulate, obovoid, widest in middle of apical cell, slightly con-
smooth to verruculose, 5–12 × 5–7 μm, proliferating percurrently
stricted at median septum, straight to curved, (26–)
at apex. Paraphyses hyaline, hypha-like, septate, 3–4 μm diam,
27–28(–29) × (5–)6(–7) μm. Conidiomata globose, immersed,
up to 60 μm long, intermixed between conidiophores. Conidia
subepidermal with central ostiole, brown, 150–250 μm diam; wall
solitary, brown, verruculose, fusoid-ellipsoid, 3-distoseptate with
of 3–6 layers of brown textura angularis. Conidiophores reduced
central septal pore in each septum, apex subobtuse, base trun-
to inconspicuous conidiogenous cells lining the cavity, doliiform,
cate, 3–4 μm diam, (27–)32–36(–40) × (8–)9(–10) μm.
pale brown, proliferating percurrently, 4–9 × 4–5 μm. Paraphyses
Culture characteristics: Colonies erumpent, spreading, surface sparse, hyaline, subcylindrical, sparsely septate, up to 30 μm
folded, with moderate aerial mycelium and smooth, lobate long, 2–3 μm diam. Conidia solitary, medium brown, verrcucu-
margin, reaching 10 mm diam after 2 wk at 25 °C. On MEA, PDA lose, 3-distoseptate with central pore at each septum, apex
and OA surface dirty white, reverse ochreous. subobtuse, base truncate, (15–)18–20 × (8–)9–10(–11) μm.
Typus: Australia, New South Wales, Mildura, Mungo National Typus: Australia, Victoria, Box Hill, on Eucalyptus sp. (Stringy
Park, on E. costata, 28 Aug. 2015, B.A. Summerell, HPC 2229 bark), Mrs Martin (holotype K(M) 251545). Not known from
(holotype CBS H-24041, culture ex-type CBS 145081 = CPC culture.
34405); idem., CBS 145082 = CPC 34395.
Note: Little is known regarding this species, except that it has
Notes: Other than Neosonderhenia foliorum, N. eucalypti should also been reported on E. sieberi in East Gippsland (Park et al.
also be compared to Sonderhenia, which presently includes two 2000).
species, S. eucalyptorum (conidia 40–48 × 5–6 μm; Hansford
Neothyriopsis Crous, gen. nov. MycoBank MB832028.
1954) and S. eucalypticola (conidia 20–26 × 9–11 μm; Swart
& Walker 1988), both associated with small, circular leaf spots Etymology: Name reflects its similarity to the genus Thyriopsis.
on Eucalyptus. Swart & Walker (1988) treated Hendersonia
Ascostromata scutellate to conical, suborbicular, irregularly
fraserae (conidia 23–28 × 6–9 μm; Hansford 1954) as synonym
tuberculate, subcuticular, black, producing coralloid, hyaline
of Hendersonia eucalypticola, which was placed in a new genus
haustoria in the epidermal cells, guard cells, cells surrounding
as Sonderhenia eucalypticola.
the stomatal cavity and the first layer of mesophyll; uni- to
Neosonderhenia foliorum (Cooke) Crous, comb. nov. Myco- multilocular, opening by irregular apical fissures exposing the
Bank MB832027. Fig. 57. ascospores; upper wall stromatic, composed of several layers of
Basionym: Stilbospora foliorum Cooke, Grevillea 20(no. 93): 6. dark brown flattened, thick-walled cells; basal wall well devel-
1891. oped, composed of two layers of cells, the outer layer composed
Fig. 58. Neothyriopsis sphaerospora (epitype, CBS H-24031). A. Disease symptoms. B. Vertical section through ascoma. C–G. Asci and ascospores. Scale bars = 10 μm.
202
of pale brown, relatively thin-walled cells, which extend laterally Mycelium superficial, composed of pale brown, smooth, septate,
for some distance beyond the base of the upper wall as an branched, thick hyphae. Ascomata perithecial, globose to sub-
intracuticular, unicellular layer of radially elongated, hyaline cells, globose with a flattened base and central ostiole, superficial,
and an inner layer. Asci parallel on the basal wall between hy- amphigenous, dark brown, setose, scattered to a few closely
aline, deliquescent pseudoparaphyses, broadly clavate, wall grouped. Setae numerous, pale brown, septate, thick-walled,
thickened, especially at the apex, bitunicate, deliquescing before smooth, giving a star-like appearance to the ascoma. Paraphy-
spore maturation. Ascospores hyaline when immature, becoming ses hyaline, septate, deliquescing early. Asci cylindrical to
dark brown, thick-walled, subglobose to globose, finely verru- cylindric-clavate to narrowly ellipsoid, shortstalked, unitunicate,
cose, medianly 1-septate (adapted from Marasas 1966). 8-spored, with an amyloid apical structure staining blue in
Meltzer’s reagent. Ascospores narrowly ellipsoid to fusoid, hy-
Type species: Neothyriopsis sphaerospora (Marasas) Crous
aline, aseptate, smooth, guttulate when young, mostly straight to
Notes: This pathogen is known to occur in Australia, Brazil, Chile slightly curved, obliquely uniseriate to biseriate inside ascus.
and South Africa (Park et al. 2000). All attempts to cultivate it
have proven unsuccessful, as ascospores germinate (on MEA, Type species: Neotrichosphaeria eucalypticola (Sivan. & R.G.
PDA), but die shortly thereafter. Shivas) Crous & Carnegie
As biotrophic pathogen, Neothyriopsis sphaerospora is Notes: Very little is known regarding the biology of this pathogen,
distinct from the genus Thyriopsis, which occurs on needles of which was described from symptomatic leaves of E. deglupta
Pinus spp. Thyriopsis has thyrothecia that open by linear fis- and C. torelliana in Queensland, Australia (Sivanesan & Shivas
sures, sometimes Y-shaped, asci are bitunicate, 8-spored, and 2002b). Because of the lack of DNA data for the type species,
contain ascospores that are ellipsoidal, 1-septate, with cells of T. pilosa, Reblova & Gams (2016) recommended accepting
roughly equal size, rounded at the ends, highly constricted at the Trichosphaeria as the only member of the Trichosphaeriaceae,
septa, hyaline to yellowish brown (von Arx & Müller 1975), which until it has been recollected and sequenced. Neotrichosphaeria
clearly distinguish it from N. sphaerospora. The ascostromata of is distinguished from Trichosphaeria (Reblova & Gams 2016) in
Neothyriopsis are more similar to those of Thyriopsis, having that it lacks a periphysate ostiole, and has numerous, very long
upper walls composed of several layers of stromatic cells, and a and flexuous setae, paraphyses that dissolve during maturation,
basal wall of two cell layers. In addition to the difference in as- asci with a visible discharge mechanism, and ascospores that
cospores, Neothyriopsis also has well developed haustoria, are hyaline and aseptate.
which are absent in Thyriopsis.
Neotrichosphaeria eucalypticola (Sivan. & R.G. Shivas) Crous
Neothyriopsis sphaerospora (Marasas) Crous, comb. nov. & Carnegie, comb. nov. MycoBank MB832031. Fig. 59.
MycoBank MB832029. Fig. 58. Basionym: Trichosphaeria eucalypticola Sivan. & R.G. Shivas,
Basionym: Thyriopsis sphaerospora Marasas, Bothalia 9: 206. Fungal Diversity 9: 172. 2002.
1966.
Leaf spots amphigenous, subcircular, brown, with irregular mar-
Diagnosis: Associated with corky lesions on leaves of Eucalyptus gins. Mycelium superficial, composed of pale brown, smooth,
species. The ascostromata are raised, brown to black in colour, septate, branched, up to 5 μm thick hyphae. Ascomata perithecial,
and occur singly or in clusters on both leaf surfaces. These globose to subglobose with a flattened base and central ostiole,
structures are usually found on healthy leaf tissue, and subse- superficial, amphigenous, dark brown, setose, scattered to a few
quently form small, discrete, corky, necrotic lesions, extending closely grouped, 170–250 μm wide, 150–230 μm high. Setae
through the leaf lamina. This fungus infects both juvenile and numerous, pale brown, septate, thick-walled, smooth, up to
mature foliage, and severe infection leads to premature defoli- 600 μm long and 7 μm wide, giving a star-like appearance to the
ation. Ascospores are dark brown, thick-walled, subglobose to ascoma. Peridium up to 26 μm thick is composed of polygonal to
globose, finely verruculose, medianly 1-septate, angular, dark brown, thick-walled cells towards the outerside, and
10–15 × 10–12.5 μm. subhyaline to hyaline, thin-walled, compressed cells towards the
Description and illustration: Marasas (1966). interior. Paraphyses hyaline, septate, deliquescing early. Asci cy-
lindrical to cylindric-clavate to narrowly ellipsoid, shortstalked,
Typus: South Africa, Gauteng, Nylstroom, on E. camaldulensis, unitunicate, 8-spored, 70–95 × 9.5–11 μm, with an amyloid apical
Aug. 1963, W.F.O. Marasas (holotype PREM 42659); Western structure staining blue in Meltzer’s reagent. Ascospores narrowly
Cape Province, Stellenbosch, Stellenbosch Mountain, on ellipsoid to fusoid, hyaline, aseptate, smooth, guttulate when
Eucalyptus sp., 2003, P.W. Crous (epitype designated here CBS young, mostly straight to slightly curved, obliquely uniseriate to
H-24031, MBT388155). biseriate inside ascus, 16–23 × 4.5–6 μm (adapted from
Note: The ITS sequence obtained directly from fungal material Sivanesan & Shivas 2002b).
removed from the leaf surface showed some similarity to the Typus: Australia, Queensland, Babinda, on E. deglupta, 6 Jul.
genus Blastacervulus, e.g. with Blastacervulus eucalypti (as 2000, M.H. Ivory (holotype BRIP 27808); New South Wales,
Heteroconium eucalypti; GenBank DQ885893.1, 707/821 (86 %), Kew, Burrawan State Forest, on E. microcorys, 14 Apr. 2018,
55 gaps (6 %)). A.J. Carnegie, HPC 2416 (epitype designated here CBS H-
Neotrichosphaeria Crous & Carnegie, gen. nov. MycoBank 24045, MBT388156, culture ex-epitype CBS 145891 = CPC
MB832030. 35777).
Etymology: Name reflects its similarity to the genus Notes: Based on a megablast search of NCBI’s GenBank
Trichosphaeria. nucleotide database, the closest hits using the ITS sequence had
CROUS ET AL.
Fig. 59. Neotrichosphaeria eucalypticola (CBS H-24045). A, B. Disease symptoms (BRIP 27808). C. Ascomata on leaf midrib. D. Ascoma with setae. E–H. Asci. I. Pa-
raphyses. J, K. Setae. Scale bars = 10 μm.
highest similarity to Iodosphaeria sp. DSM-2019a voucher MFLU ascomata), 8-spored, obovoid to broadly ellipsoidal, straight to
19-0719 (GenBank MK737501.1; Identities = 505/575 (88 %), 23 slightly incurved. Ascospores tri- to multiseriate, fusoid-ellipsoidal
gaps (4 %)), and Iodosphaeria tongrenensis (as Iodosphaeria sp. with obtuse ends, hyaline, smooth, pale brown and verruculose
QL-2015 voucher MFLU 15-0393, GenBank KR095282.1; in old asci, becoming 3-septate, not constricted at median
Identities = 444/505 (88 %), 23 gaps (4 %)). Closest hits using septum, thick-walled, guttulate, widest in the middle of the apical
the LSU sequence are Iodosphaeria tongrenensis (as Iodos- cell, with persistent mucous sheath. Conidiomata sporodochial,
phaeria sp. QL-2015 voucher MFLU 15-0393, GenBank at times concentrically arranged, dark brown to black, dry,
KR095283.1; Identities = 766/780 (98 %), 1 gap (0 %)), Iodos- powdery, confined to the lesions. Conidiophores branched at
phaeria sp. DSM-2019a voucher MFLU 19-0719 (GenBank base, pale brown, smooth, loosely aggregated, pale brown.
MK722172.1; Identities = 764/780 (98 %), 1 gap (0 %)), and Conidiogenous cells terminal, cylindrical to doliiform, holothallic,
Pseudosporidesmium knawiae (GenBank MH874823.1; Identi- pale brown. Conidia formed in basipetal chains, smooth, medium
ties = 741/782 (95 %), 7 gaps (0 %)). brown, 4-celled, consisting of two basal cells with truncate lateral
sides (adhesion scars present when catenulate), each giving rise
Neotrimmatostroma Quaedvl. & Crous, Persoonia 33: 27. to a secondary globose apical cell, that may extend and develop
2014. two additional septa; septa dark brown and thick-walled between
the primary and secondary cells.
Ascomata pseudothecial, separate, dark brown, subepidermal,
becoming erumpent, globose; ostiole apical, central, frequently Symptoms: Leaf spots separate, coalescing with age, medium
opening by irregular rupture; wall of 2–3 layers of dark brown, brown, subcircular with indistinct margins, 2–7 mm diam,
thick-walled textura angularis. Asci fasciculate, bitunicate, apar- amphigenous, with prominent olivaceous brown sporodochia,
aphysate (remains of the hamathecium observed in some giving lesions a sooty appearance.
204
Type species: Neotrimmatostroma excentricum (B. Sutton & Neotrimmatostroma excentricum (B. Sutton & Ganap.)
Ganap.) Quaedvl. & Crous Quaedvl. & Crous, Persoonia 33: 27. 2014. Fig. 62.
Basionym: Trimmatostroma excentricum B. Sutton & Ganap.,
Notes: Neotrimmatostroma is a common genus of phytopatho-
New Zealand J. Bot. 16: 529. 1978.
genic fungi observed on eucalypt leaves. The disease can be
Synonyms: Catenulostroma excentricum (B. Sutton & Ganap.)
quite serious on some eucalypt species, but is generally
Crous & U. Braun, Stud. Mycol. 58: 10. 2007.
considered to be of minor importance in commercial forestry
Mycosphaerella excentrica Crous & Carnegie, Fungal Diversity
plantations.
26: 164. 2007.
Neotrimmatostroma dalrympleanae Crous, sp. nov. Myco- Teratosphaeria excentrica (Crous & Carnegie) Crous & U. Braun,
Bank MB832032. Fig. 60. Stud. Mycol. 58: 10. 2007.
Diagnosis: Conidia in basipetal chains, smooth, pale brown, 4-
Etymology: Name refers to Eucalyptus dalrympleana, the host
celled, consisting of an upper and a lower truncate ended pri-
species from which this fungus was collected.
mary cell separated by a thick, dark brown transverse septum.
Mycelium immersed, consisting of pale brown, septate, Each cell so delimited produces a smaller, single, lateral, globose
branched, 2–3 μm diam hyphae. Conidiomata sporodochial, secondary cell on the same side of the conidium. The conidial
chiefly epiphyllous, concentrically arranged, dark brown, dry wall opposite the lateral cells is considerably thicker than the rest
powdery, discrete, up to 400 μm diam. Conidiophores micro- of the conidium. The two primary cells combined are
nematous, branched, septate, medium brown, smooth, densely 9–11 × 3–4 μm, and secondary cells are 2.5–4.5 μm.
aggregated, with differential thickening of periclinal wall, one side
Descriptions and illustrations: Sutton & Ganapathi (1978), Crous
thinner than the other, up to 30 μm tall, 3–4 μm diam. Con-
et al. (2007a).
idiogenous cells holothallic, integrated, terminal, doliiform to
subcylindrical, 5–10 × 2.5–3 μm. Conidia in sparsely branched Typus: Australia, New South Wales, Kempsey, Mackenzie
chains, smooth, pale brown, 4-celled, consisting of upper and Creek Road, Byrne Plantation, S30°53'15" E152°27'47", on
lower cells with truncate ends, separated by a thick, dark brown E. agglomerata, 13 Apr. 2005, G. Price (holotype of sexual
transverse septum, each primary cell with a smaller, lateral, morph CBS H-19829, isotype DAR 78033, culture ex-type CBS
globose secondary cell on either side of the primary septum. The 121102 = CPC 13092)
two primary cells together are (7–)8–9(–10) μm diam, the
Neotrimmatostroma paraexcentricum Crous et al., Persoonia
secondary cells 5–6 μm diam.
36: 383. 2016. Fig. 63.
Diagnosis: Conidia in sparsely branched chains, smooth, pale
sparse aerial mycelium and feathery margin, reaching 4 cm after
brown, 4-celled, consisting of upper and lower cells with truncate
2 wk at 25 °C. On MEA, PDA and OA surface olivaceous grey.
ends, separated by a thick, dark brown transverse septum, each
Typus: Australia, New South Wales, South East National Forest primary cell with a smaller, lateral, globose secondary cell on
Park, on E. dalrympleana, 28 Nov. 2016, P.W. Crous, HPC 1815 either side of the primary septum. The two primary cells together
(holotype CBS H-24033, culture ex-type CBS 144609 = CPC are 9–11 μm diam, the secondary cells 4–5 μm diam.
32605).
Description and illustration: Crous et al. (2016c).
Notes: Neotrimmatostroma dalrympleanae is phylogenetically
distinct from other closely related species (LSU phylogeny Typus: Australia, Victoria, Phillip Island, Oswin Roberts
presented in Fig. 61). In addition, the ITS sequence is most Reserve, on Eucalyptus sp., 8 Nov. 2014, P.W. Crous, J.
similar to Neotrimmatostroma paraexcentricum (GenBank Edwards & I.G. Pascoe (holotype CBS H-22621, culture ex-type
NR_145124.1; 516/533 (97 %), 4 gaps (0 %)). CBS 141325 = CPC 25594).
Fig. 60. Neotrimmatostroma dalrympleanae (CPC 32605). A–E. Conidiogenous cells giving rise to conidia. F. Conidia. Scale bars = 10 μm.
CROUS ET AL.
Elsinoe jasminicola CBS 212.63 (KX886997.1)

Fumagospora capnodioides CBS 131.34 (EU019269.1)
1 Scorias leucadendri CBS 131318 (JQ044456.1) Capnodiaceae
Scorias spongiosa CBS 325.33 (GU214696.1)
Graphiopsis chlorocephala CPC 11969 (EU009458.2)
1
Verrucocladosporium dirinae CBS 112794 (EU040244.1)
0.95 1
Toxicocladosporium leucadendri CBS 131317 (JQ044455.1)
Cladosporium cycadicola CBS 137970 (KJ869179.1) Cladosporiaceae
1
1 Cladosporium perangustum CPC 18228 (JF499855.1)
Cladosporium allicinum CPC 5101 (GU214408.1)
Cladosporium phaenocomae CBS 128769 (JF499857.1)
1 Chaetothyrina artocarpi MFLUCC 15-1082 (MF614834.1)
1 Phaeothecoidiella illinoisensis CBS 125223 (GU117901.1) Phaeothecoidiellaceae
Phaeothecoidiella missouriensis CBS 125222 (MH874962.1)
CPC 32400
1
CPC 32901 Nowamyces piperitae, sp. nov.
1 CPC 32946
CPC 33553 Nowamycetaceae, fam. nov.
CPC 32722
0.90 Nowamyces globulus gen. et sp. nov.
0.99 CPC 32894
0.92
CPC 32724
1 Zymoseptoria passerinii CBS 120382 (JQ739843.2)
Zymoseptoria tritici CPC 18116 (KX287264.1)
0.90
Zymoseptoria brevis CPC 18102 (KF252148.1)
1 Lecanosticta acicola CBS 133791 (NG_042762.1)
1 Lecanosticta pini CBS 871.95 (GQ852598.1)
Phaeophleospora hymenocallidis CBS 139911 (NG_058168.1)
Phaeophleospora eucalypticola CPC 26523 (KX228318.1)
0.99 Phaeophleospora pteridivora CPC 24683 (NG_059578.1)
Phaeophleospora stramenti CBS 118909 (KF901942.1)
1 0.99
Phaeophleospora eugeniicola CPC 2558 (FJ493209.2)
Mycosphaerelloides madeirae CBS 112895 (DQ204756.1)
0.94
Phaeophleospora concentrica CPC 3615 (FJ493205.1)
Virosphaerella irregularis CBS 123242 (MH874810.1)
1 Zasmidium musigenum CBS 365.36 (EU041858.1) Mycosphaerellaceae
Zasmidium cellare CBS 146.36 (NG_057791.1)
0.98 Zasmidium scaevolicola CBS 127009 (KF251789.1)
0.99 0.95 Sonderhenia eucalypticola CBS 112502 (KF902019.1)
0.87 Sonderhenia eucalypticola CMW 20333 (DQ267574.1)
0.88 Sonderhenia eucalyptorum CBS 120220 (DQ923536.1)
1 Sonderhenia sp. CPC 17710
Pseudophaeophleospora stonei CBS 120830 (KF901847.1)
CPC 34395
Neosonderhenia eucalypti, gen. et sp. nov.
CBS 145081
Pseudophaeophleospora atkinsonii ICMP 17862 (GU214463.1)
1
Pallidocercospora ventilago CPC 21817 (KF777229.1)
0.99 Pallidocercospora crystallina ZJUM 2 (KP895884.1)
Pallidocercospora irregulariramosa CPC 1362 (GU214441.1)
0.01
Fig. 61. Consensus phylogram (50 % majority rule) of 182 252 trees resulting from a Bayesian analysis of the Capnodiales LSU sequence alignment (90 strains including the
outgroup; 779 aligned positions; 258 unique site patterns). The tree was rooted to Elsinoe jasminicola (GenBank KX886997.1) and the scale bar indicates the expected changes
per site. Bayesian posterior probabilities (PP) >0.84 are shown at the nodes and novelties are highlighted with bold text. Family names are indicated to the right of the tree and
GenBank accession numbers between round parentheses. Thickened branches represent those present in the strict consensus parsimony tree. The analyses were performed
using MrBayes v. 3.2.6 (Ronquist et al. 2012) and PAUP v. 4.0b10 (Swofford 2003). The parsimony analysis (only strict consensus branches showed) yielded the maximum of
1 000 equally most parsimonious trees and had the following statistics: 529 constant, 59 variable and parsimony-uninformative and 193 parsimony-informative characters;
TL = 890 steps, CI = 0.415, RI = 0.842, RC = 0.349.
Nothotrimmatostroma Crous, gen. nov. MycoBank MB832033. Conidiomata sporodochial, dark brown to black, dry, powdery,
confined to the lesions. Conidiophores branched at base, pale
Etymology: Referring to its morphological similarity with brown, smooth, loosely aggregated, pale brown. Conidiogenous
Trimmatostroma. cells terminal, cylindrical to doliiform, holothallic, pale brown.
206
1 Austroafricana parva CBS 110503 (KT186510.1)

Austroafricana associata CBS 112224 (GU301874.1)
1 Devriesia staurophora CBS 375.81 (KT186509.1)
Devriesia tardicrescens CBS 128770 (JF499860.1)
Phaeothecoidea intermedia CBS 124994 (NG_057841.1)
1 Phaeothecoidea eucalypti CPC 13010 (EU019280.1)
1
Phaeothecoidea minutispora CBS 124995 (GQ852629.1)
1
Nothotrimmatostroma bifarium, comb. nov. CBS 143488
0.97 CPC 17586 (JF951174.1)
Nothotrimmatostroma eucalyptorum, comb. nov.
CBS 129578 (MH876908.1)
Suberoteratosphaeria xenosuberosa CBS 134747 (KF901879.1)
1
Suberoteratosphaeria suberosa CPC 11032 (GQ852718.1)
0.93
Suberoteratosphaeria pseudosuberosa CBS 118911 (KF902144.1)
Neotrimmatostroma dalrympleanae, sp. nov. CPC 32605
1 Neotrimmatostroma paraexcentricum CBS 141325 (NG_058243.1)
1 Neotrimmatostroma excentricum CBS 121102 (KF901840.1)
1 Penidiella aggregata CBS 128772 (JF499862.1)
Euteratosphaeria verrucosiafricana CBS 118497 (KF901992.1)
Teratosphaeria fibrillosa CPC 1876 (GU214506.1)
Teratosphaeria alboconidia CPC 14597 (FJ493221.1)
Teratosphaeria pseudocryptica CBS 118503 (KF442547.1)
Teratosphaeriaceae
Teratosphaeria miniata CBS 125006 (KF442570.1)
Teratosphaeria zuluensis 157B2 (JQ732966.1)
Teratosphaeria gauchensis CBS 120303 (EU019290.1)
Teratosphaeria stellenboschiana CBS 124989 (GQ852715.2)
Penidiella columbiana CBS 486.80 (EU019274.2)
Teratosphaeriopsis pseudafricana CBS 111171 (KF902085.1)
Teratosphaericola pseudoafricana CBS 111168 (KF902045.1)
0.99
Pseudoteratosphaeria ohnowa CPC 1005 (GU214511.1)
Pseudoteratosphaeria secundaria CBS 115608 (EU019306.2)
1 Pseudoteratosphaeria flexuosa CBS 111048 (KF901978.1)
Pseudoteratosphaeria africana CBS 144595 (MK442558.1)
1
Pseudoteratosphaeria perpendicularis CBS 118367 (KF901972.1)
Pseudoteratosphaeria stramenticola CBS 118506 (KF901948.1)
Myrtapenidiella pleurocarpae CBS 142531 (KY979822.1)
Myrtapenidiella eucalypti CBS 123246 (MH874813.1)
Myrtapenidiella balenae CBS 142527 (KY979816.1)
1
Myrtapenidiella tenuiramis CBS 124993 (GQ852626.1)
Myrtapenidiella eucalyptorum CBS 139920 (MH878664.1)
Myrtapenidiella eucalyptigena CBS 142526 (KY979815.1)
Myrtapenidiella sporadicae CPC 29200 (KY173513.1)
0.85
Myrtapenidiella corymbia CBS 124769 (MH874924.1)
0.01
Fig. 61. (Continued).
Conidia formed in basipetal chains, smooth, medium brown, 4- of minor importance. Morphologically there is no obvious differ-
celled, consisting of two basal cells with truncate lateral sides ence between these two genera, and they are best separated
(adhesion scars present when catenulate), each giving rise to a based on their DNA phylogeny (LSU phylogeny presented in
secondary globose apical cell, that may extend and develop Fig. 61).
additional septa; septa dark brown and thick-walled between the
primary and secondary cells. Nothotrimmatostroma bifarium (Gadgil & M.A. Dick) Crous,
comb. nov. MycoBank MB832034. Fig. 64.
Symptoms: Leaf spots separate, coalescing with age, medium Basionym: Trimmatostroma bifarium Gadgil & M.A. Dick, New
brown, subcircular with indistinct margins, amphigenous, with Zealand J. Bot. 21: 49. 1983.
prominent olivaceous brown sporodochia, giving lesions a sooty Synonym: Neotrimmatostroma bifarium (Gadgil & M.A. Dick)
appearance (similar to spots of Neotrimmatostroma). Quaedvl. & Crous, Persoonia 33: 27. 2014.
Type species: Nothotrimmatostroma bifarium (Gadgil & M.A.
Diagnosis: Conidia formed in two basipetal chains, brown, con-
Dick) Crous
sisting of two parallel, laterally fused rows, with a common thick-
Notes: Species of Nothotrimmatostroma are less commonly ened, transverse base and obtuse apices, 12–24 × 6–14 μm,
observed than those of Neotrimmatostroma, and appear to be 6–10-celled when mature (with 3–5 cells per row).
CROUS ET AL.
Fig. 62. Neotrimmatostroma excentricum (CPC 13092). A. Leaf spot with ascomata. B, C. Asci. D. Ascospores. E. Colony on OA. F, G. Conidia. Scale bars = 10 μm.
Fig. 63. Neotrimmatostroma paraexcentricum (CPC 25594). A, B. Leaf spots with conidiomata. C. Close-up of leaf spot. D–F. Conidiogenous cells giving rise to conidia. G.
Conidia. Scale bars = 10 μm.
208
Fig. 64. Nothotrimmatostroma bifarium (CPC 32833). A. Leaf spot with conidiomata. B–F. Conidiogenous cells giving rise to conidia. G. Conidia. Scale bars = 10 μm.
Descriptions and illustrations: Gadgil & Dick (1983), Park et al. basal cell that is globose, thin-walled, pale brown,
(2000). 4–5 × 2–2.5 μm.
Description based on CPC 32833: Leaf spots corky, medium Description and illustration: Crous et al. (2011a).
brown, irregular, 2–10 mm diam. Sporodochia hypophyllous,
Typus: Australia, New South Wales, Ebor, S30°14'21"
dark brown, in concentric clusters, 150–220 μm diam. Co-
E152°31'55", on E. laevopinea, 28 Jul. 2009, A.J. Carnegie
nidiophores micronematous, branched, septate, medium brown,
smooth, densely aggregated, up to 50 μm tall, 3–4 μm diam.
17586).
Conidiogenous cells holothallic, integrated, terminal, doliiform to
subcylindrical, 9–11 × 3–4 μm. Conidia in branched chains, Nowamycetaceae Crous, fam. nov. MycoBank MB832036.
smooth, medium brown, 2–7-celled, consisting of two parallel
Etymology: Name refers to the genus Nowamyces.
rows unequal in length, fused at common thickened base, apices
obtuse, (15–)17–20(–22) × (8–)9(–10) m, 5–6-celled in vivo, Ascomata pseudothecial, immersed, substomatal, black,
(13–)15–19(–25) × (7–)8–9(–10) μm, (2–)5–6(–7)-celled globose, with apical ostiole; wall of 2–4 layers of medium brown
in vitro. textura angularis. Asci fasciculate, bitunicate, obovoid to broadly
ellipsoidal, straight to incurved, 8-spored. Ascospores multi-
seriate, overlapping, hyaline, guttulate, thin-walled, straight to
sparse aerial mycelium and even, lobate margins, reaching
slightly curved, fusoid-ellipsoidal with obtuse ends, 1-septate.
5 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface
olivaceous grey, reverse iron-grey. Nowamyces Crous, gen. nov. MycoBank MB832037.
Typus: Australia, New South Wales, Merimbula, on Etymology: Name refers to Nowa Nowa, Australia, where this
E. dalrympleana, 28 Nov. 2016, P.W. Crous, HPC 1815 (epitype fungus was collected.
designated here CBS H-24032, MBT388158, culture ex-epitype
CBS 143488 = CPC 32833). New Zealand, Kinleith, on Presumed phytopathogenic, associated with leaf spots of Ter-
E. regnans, Sep. 1981, D.J. Rawcliffe (holotype NZFRI 5, atosphaeria nubilosa. Ascomata pseudothecial, predominantly
isotype PDD 42845). hypophyllous, immersed, substomatal, black, globose, with api-
cal ostiole; wall of 2–4 layers of medium brown textura angularis.
Nothotrimmatostroma eucalyptorum (Crous & Carnegie) Asci fasciculate, bitunicate, obovoid to broadly ellipsoidal,
Crous, comb. nov. MycoBank MB832035. straight to incurved, 8-spored. Ascospores multiseriate, over-
Basionym: Catenulostroma eucalyptorum Crous & Carnegie, lapping, hyaline, guttulate, thin-walled, straight to slightly curved,
Persoonia 26: 149. 2011. fusoid-ellipsoidal with obtuse ends, widest in middle of the apical
Diagnosis: Conidia catenulate, smooth, pale brown, 4-celled, cell, medianly 1-septate, prominently constricted at septum,
upper two primary cells 7–9 × 2–4 μm, with truncate ends where tapering toward both ends. Ascospores become distorted, brown
attached, 1.5–2 μm diam, cells separated from each other by a and verruculose upon germination. No asexual morph produced
broad, dark brown area; each primary cell giving rise to a smaller in culture.
CROUS ET AL.
Type species: Nowamyces globulus Crous Typus: Australia, Victoria, Nowa Nowa, on E. globulus, 30 Nov.
2016, P.W. Crous, HPC 1875 (holotype CBS H-24034, cultures
Notes: Species of Nowamyces are teratosphaeria-like in
ex-type CBS 144598 = CPC 32722, CBS 144599 = CPC 32724,
morphology, and can only be reliably identified based on DNA
CBS 144600 = CPC 32726, CBS 144601 = CPC 32894).
sequence (LSU phylogeny presented in Fig. 61). Nowamyces
piperitae was associated with prominent leaf spots, suggesting Notes: Nowamyces globulus was collected in an attempt to find
that the genus is plant pathogenic. authentic material of Austroafricana parva, which was first iso-
lated from Nowa Nowa on E. globulus, where it occurred in older
Nowamyces globulus Crous, sp. nov. MycoBank MB832038.
lesions of T. nubilosa. However, the present collection proved to
Fig. 65.
be distinct, in that the ascospores are larger, (11–)
Etymology: Name refers to E. globulus, the host species from 13–16(–18) × (4–)4.5(–5) μm, those of Austroafricana parva
which this fungus was collected. being (7–)8–9(–10) × (2–)2.5(–3) μm (Crous 1998), and those
of its suggested synonym, Mycosphaerella grandis, being
Leaf spots amphigenous, subcircular, 4–15 mm diam, pale (10.5–)12.3(–14.5) × (3–)3.8(–4.5) μm (Carnegie & Keane
brown, with raised dark brown border; occurring on older leaf 1994). Furthermore, colonies proved to be homothallic in
spots of Teratosphaeria nubilosa. Ascomata pseudothecial, culture.
predominantly hypophyllous, immersed, substomatal, black,
globose, (40–)90–120 μm diam; apical ostiole 10 μm diam; wall Nowamyces piperitae Crous, sp. nov. MycoBank MB832039.
of 2–4 layers of medium brown textura angularis. Asci fascicu- Fig. 65.
late, bitunicate, obovoid to broadly ellipsoidal, straight to Etymology: Name refers to Eucalyptus piperita from which the
incurved, 8-spored, 28–35(–40) × (8–)9–10(–13) μm. Asco- fungus was collected.
spores multiseriate, overlapping, hyaline, guttulate, thin-walled,
straight to slightly curved, fusoid-ellipsoidal with obtuse ends, Leaf spots amphigenous, irregular, 3–30 mm diam, medium
widest in middle of the apical cell, medianly 1-septate, promi- brown, with raised red-brown border. Ascomata pseudothecial,
nently constricted at septum, tapering toward both ends, (11–) amphigenous, dark brown, immersed, inconspicuous, globose,
13–16(–18) × (4–)4.5(–5) μm. Ascospores initially germinating 80–120 μm diam; wall of 3–4 layers of brown textura angularis.
at right angles to the long axis, but eventually both germ tubes Asci aparaphysate, fasciculate, obovoid with ocular chamber,
grow parallel to the long axis, spores become distorted, brown, straight to slightly curved, 8-spored, 30–45 × 10–12 μm. As-
verruculose, (4–)5(–6) μm diam. cospores tri- to multiseriate, overlapping, hyaline, guttulate,
thin-walled, straight to slightly curved, obovoid with obtuse
Culture characteristics: Colonies erumpent, spreading, with ends, widest near apex of apical cell, medianly 1-septate,
sparse aerial mycelium and smooth, lobate margin, reaching not constricted at septum, tapering towards both ends,
5 mm diam after 2 wk at 25 °C. On MEA, PDA and OA surface but more prominently towards lower end, (13–)
and reverse olivaceous grey. Cultures are homothallic, and 15–16( –18) × 4(–4.5) μm. Ascospores germinating with germ
produce the sexual morph on OA. tubes parallel to the long axis of the spore, becoming
Fig. 65. Nowamyces spp. A–D. N. globulus (CPC 32722). A, B. Asci. C. Ascospores. D. Germinating ascospores. E–J. N. piperitae (CPC 32400). E. Disease symptoms. F, G.
Asci. H. Ascospores. I, J. Germinating ascospores. Scale bars = 10 μm.
210
constricted and somewhat distorted at medium septum, 4 μm Clypeus amphigenous, gregarious, rounded-pulvinate, black and
diam; germinating spores have a prominent mucoid sheath shiny. Ascomata perithecial, solitary, immersed, subglobose with
visible on the agar surface. papillate ostiole. Paraphyses sparse, filiform. Asci fusoid,
attenuated towards rounded apices, without a cap, substipitate,
8-spored. Ascospores filiform, parallel in ascus, as long as
sparse aerial mycelium and smooth, lobate margins, reaching
ascus, hyaline, pluriguttulate, becoming multiseptate (from
10 mm diam after 1 mo at 25 °C. On MEA, PDA and OA surface
Hanlin et al. 1992).
olivaceous grey, reverse iron-grey.
Type species: Ophiodothella atromaculans (Henn.) Höhn.
Typus: Australia, New South Wales, Fitzroy Falls, on E. piperita,
28 Nov. 2016, P.W. Crous, HPC 1757 (holotype CBS H-24035, Note: Very little is known regarding the biology of this fungus.
Ophiodothella longispora H.J. Swart, Trans. Brit. Mycol. Soc.
Notes: Nowamyces globulus and N. piperitae are phyloge- 79: 567. 1982. Fig. 67.
netically related but some intraspecific variation is found in
Diagnosis: Infected tissue swelling, with discrete black perithecial
each of the species (Fig. 66). We have refrained naming
ascomata embedded in the tissue. Ascospores parallel, asep-
cryptic species until a broader geographic sampling can be
tate, hyaline, 150–200 × 4–6 μm, with a gelatinous outer wall
made.
layer, somewhat curved, with rounded ends.
Ophiodothella (Henn.) Höhn., Sber. Akad. Wiss. Wien, Math.-
Description and illustration: Swart (1982).
naturw. Kl., Abt. 1 119: 940 [64 repr.]. 1910.
Fig. 66. The first of seven equally most parsimonious trees obtained from a phylogenetic analysis of the Nowamyces ITS sequence alignment (19 strains including the
outgroup; 465 characters analysed: 344 constant, 95 variable and parsimony-uninformative and 26 parsimony-informative). The tree was rooted to Ramularia endophylla
(GenBank AY490763.1) and the scale bar indicates the number of changes. Thickened branches represent those present in the strict consensus parsimony tree. Bootstrap
support values higher than 49 % are shown at the nodes (parsimony bootstrap support / NJ HKY85 distance bootstrap support) and novelties are highlighted with bold text.
Species names are indicated to the right of the tree. A superscript T denotes strains with a type status. Tree statistics: TL = 148 steps, CI = 0.899, RI = 0.899, RC = 0.808. The
analyses were performed using PAUP v. 4.0b10 (Swofford 2003).
CROUS ET AL.
Fig. 67. Ophiodothella longispora (DAR 35028). A, D. Disease symptoms. B. Vertical section through ascoma. C–G. Asci with ascospores. H. Ascospores. Scale bars: B,
D = 600 μm, C, E–G = 25 μm, H = 10 μm.
Typus: Australia, Victoria, Kinglake, on E. goniocalyx, Sep. Description and illustration: Swart (1982).
1980, V. Beilharz (holotype DAR 35028). Not known from
Typus: Australia, South Australia, Noarlunga Gorge, on
culture.
E. rostrata, May 1924, G. Samuel (holotype K(M) 176511,
Pachysacca Syd., Annls Mycol. 28: 435. 1930. Fig. 68. isotype ADW 3822). For additional specimens see Swart (1982).
Ascostromata black, immersed between epidermis and mesophyll,
solitary or scattered, multiloculate; peridium composed of pale Pachysacca pusilla H.J. Swart, Trans. Brit. Mycol. Soc. 79: 268.
brown to brown cells of textura angularis. Hamathecium lacking 1982.
pseudoparaphyses. Asci 8-spored, bitunicate, fissitunicate, broadly
Diagnosis: Asci develop in locules stroma formed between
obovoid at base, apical part subcylindrical, short pedicellate.
epidermis and mesophyll. Ascospores hyaline, cylindrical, 4-
Ascospores multiseriate, hyaline, smooth-walled, transversely
celled, 38–46 × 4–5 μm.
septate, narrowly subcylindrical, with broadly rounded ends.
Description and illustration: Swart (1982).
Type species: Pachysacca eucalypti Syd.
Typus: Australia, Victoria, Mr Dandenong, on E. regnans, Sept.
Symptoms: Ascomata develop between the epidermis and 1979, G. Marks (holotype DAR 37766). Not known from culture.
mesophyll, inducing a swelling of the leaf. Leaf spots are Pachysacca samuelii (Hansf.) H.J. Swart, Trans. Brit. Mycol.
amphigenous, with a less well-developed stroma developing on Soc. 79: 267. 1982. Fig. 68.
the opposite side of the leaf; the fungus forms spreading dendritic Basionym: Pseudosphaeria samuelii Hansf., Proc. Linn. Soc.
or large circular stromata. N.S.W. 79: 118. 1954.
Notes: The fungus has a restricted distribution within a stand, Diagnosis: Asci develop in locules, stroma formed between
usually being confined to a few trees in a forest. Fungal matu- epidermis and mesophyll. Ascospores hyaline, elongate cylin-
ration depends on rainfall, and new infections occur during the drical, 4-celled, 60–65 × 6–7 μm.
cooler months (Park et al. 2000). Attempts to culture species of
Pachysacca have thus far proven unsuccessful. Description and illustration: Swart (1982).
Typus: Australia, South Australia, Cape Jervis peninsula, on
Pachysacca eucalypti Syd., Annls Mycol. 28: 435. 1930. E. obliqua, Jan. 1924, G. Samuel (lectotype ADW 2069).
Fig. 68. For additional specimens see Swart (1982). Not known from
Diagnosis: Asci develop in locules stroma formed between culture.
epidermis and mesophyll. Ascospores hyaline, cylindrical, 3-
celled, 90–120 × 7 μm. Pallidocercospora Crous, Stud. Mycol. 75: 73. 2012. Fig. 69.
212
Fig. 68. Pachysacca spp. A, B, D. Disease symptoms of P. eucalypti (BRIP 27863). C. Disease symptoms of Pachysacca sp. E. Disease symptoms of P. samuelii (BPRIP
27863a). F, G. Spermatogonium and spermatogenous cells of P. eucalypti. H, I, N–Q. Vertical section through ascomata, asci and ascospores of P. samuelii. J–M. Asci and
ascospores of P. eucalypti. Scale bars = 10 μm.
CROUS ET AL.
Fig. 69. Disease symptoms of Pallidocercospora spp. A, B. P. crystallina (PREM 51922). C–E. P. heimii (PREM 51749). F–N. P. crystallina. F–H. Vertical section through
ascomata. I–L. Asci. M. Conidiogenous cells on hyphae. N. Ascospores. Scale bars = 10 μm.
Ascomata single, black, immersed, globose, glabrous; wall cell. Conidia solitary, straight to irregularly curved, guttulate,
of 3 –4 layers of medium brown textura angularis. Asci pale olivaceous to olivaceous-brown, subcylindrical to
fasciculate, bitunicate, aparaphysate, subsessile, 8-spored, narrowly obclavate, multiseptate; hila neither thickened nor
ellipsoid to obclavate or cylindrical, straight or curved, darkened.
numerous. Ascospores 2 –multi-seriate, oblique, overlapping,
Type species: Pallidocercospora heimii (Crous) Crous
straight ellipsoidal to obovoid, colourless, smooth, 1-septate.
Mycelium predominantly immersed, consisting of olivaceous- Notes: Pallidocercospora is characterised by having pale oliva-
brown hyphae, smooth, branched, septate, 2 – 4 μm diam. ceous, smooth conidia (Crous et al. 2004b), and is known to form
Conidiophores in vivo fasciculate, or occurring singly on red crystals when cultivated on agar (on WA, SNA, PDA, MEA),
superficial mycelium as lateral projections, unbranched or which distinguishes it from Pseudocercospora.
branched, septate, cylindrical, straight to geniculate-sinuous,
olivaceous-brown. Conidiogenous cells integrated, terminal, Pallidocercospora crystallina (Crous & M.J. Wingf.) Crous &
cylindrical, straight to geniculate-sinuous, olivaceous-brown, M.J. Wingf., Stud. Mycol. 75: 74. 2012. Fig. 69.
proliferating sympodially or percurrently; conidiogenous loci Basionym: Pseudocercospora crystallina Crous & M.J. Wingf.,
unthickened, not darker than the surrounding conidiogenous Mycologia 88: 451. 1996.
214
Synonym: Mycosphaerella crystallina Crous & M.J. Wingf., olivaceous to pale brown, finely verruculose, guttulate, narrowly
Mycologia 88: 451. 1996. obclavate with a rounded to subobtuse apex and long obconic-
truncate to truncate base, 4- to multiseptate, widest near the
first basal septum or in the middle of the basal cell, (25–)
diam, coalescing to form larger blotches, light brown, surrounded
40–90(–150) × (2–)2.5–3(–3.5) μm in vitro.
by raised, dark brown borders on the adaxial surfaces, whitish-
brown on the lower surfaces, surrounded by raised, con- Description and illustration: Crous & Wingfield (1996).
colourous borders on the abaxial surfaces. Ascospores bi- to
triseriate, overlapping, colourless, guttulate, thin-walled, straight, Typus: Indonesia, N. Sumatra, Lake Toba area, on Eucalyptus
rarely curved, obovoid, with obtuse basal and bluntly obtuse sp., Mar. 1996, M.J. Wingfield (holotype of sexual morph PREM
apical cells, widest near apex, medianly 1-septate, not con- 54966; holotype of asexual morph PREM 54967; cultures ex-
stricted at septum, tapering toward both ends, but with more type CBS 111190 = CMW 3046 = CPC 1312, CPC 1311).
prominent taper toward the base, (11–)
12–14(–15) × 3–3.5(–4) μm. Conidia solitary, sinuous, smooth, Pallidocercospora irregulariramosa (Crous & M.J. Wingf.)
olivaceous, narrowly obclavate with a subobtuse apex and long Crous & M.J. Wingf., Stud. Mycol. 75: 74. 2012.
obconic-truncate base, widest in the middle of the basal cell, Basionym: Pseudocercospora irregulariramosa Crous & M.J.
guttulate, multiseptate, 50–200 × 2–3 μm. Wingf., Canad. J. Bot. 75: 785. 1997.
Synonym: Mycosphaerella irregulariramosa Crous & M.J. Wingf.,
Description and illustration: Crous & Wingfield (1996). Canad. J. Bot. 75: 785. 1997.
Typus: South Africa, Kwazula-Natal Province, Umvoti, on Diagnosis: Leaf spots amphigenous, subcircular, 3–15 mm
E. bicostata, Oct. 1994, M.J. Wingfield (holotypes PREM 51922, diam, grey to pale brown, surrounded by a slightly raised border
sexual morph; PREM 51923, asexual morph, cultures ex-type and dark brown margin. Ascospores multiseriate, overlapping,
CBS 681.95 = CMW 3033 = CPC 802, CPC 800, 801). hyaline, guttulate, thin-walled, straight, fusoid-ellipsoidal with
Pallidocercospora heimii (Crous) Crous, Stud. Mycol. 75: 74. obtuse ends, widest in middle of apical cells, medianly 1-septate,
2012. Fig. 69. not constricted at septum, tapering toward both ends, but with
Basionym: Pseudocercospora heimii Crous, S. African For. J. slightly more so toward base (7–)8–10 × (1.5–)2–2.5 μm.
172: 4. 1995. Conidia solitary, pale to medium brown, verruculose, guttulate,
Synonyms: Mycosphaerella heimii Crous, S. African For. J. 172: thick-walled, subcylindrical with subobtuse apex and truncate
2. 1995. base, multiseptate, irregular in width on host material (not in
Mycosphaerella heimii Bouriquet, Encycl. Mycol. 12: 418. 1946. culture), variously curved, (35–)45–75(–85) × 2.5–3 μm in vivo,
nom. nud. 70–200 × 1.5–2 μm in vitro.
Diagnosis: Leaf spots discrete, amphigenous, medium brown, Description and illustration: Crous & Wingfield (1996).
elongated, irregular, 5–15 mm diam, surrounded by a promi- Typus: South Africa, Northern Province, Tzaneen, on
nently raised, brown margin; also associated with tip blight E. saligna, Mar. 1996, M.J. Wingfield (holotype of sexual morph
symptoms. Ascospores 2–3-seriate, oblique, overlapping, PREM 54964; holotype of asexual morph PREM 54965; cul-
straight ellipsoidal, obtuse at each end, hyaline, smooth, 1- tures ex-type CBS 114774 = CPC 1360, CBS 111211 = CPC
septate, prominently guttulate, not constricted at median 1362, CPC 1361).
septum, widest in the middle of upper cell, tapering more
prominently toward the base, (8–)9–11(–12) × 2–2.5(–3) μm. Paramycosphaerella Crous & Jol. Roux, Persoonia 31: 245.
Conidia solitary, irregularly curved, guttulate, olivaceous brown, 2013.
narrowly obclavate with subacute apex, obconic-truncate base Ascomata erumpent, amphigenous, brown, globose, with central
and unthickened hilum, 55–300 × 2.5–3 μm, multiseptate. ostiole; wall of 2–3 layers of brown textura angularis. Asci
Description and illustration: Crous & Swart (1995). fasciculate, bitunicate with apical chamber, 8-spored, sub-
cylindrical to narrowly ellipsoid. Ascospores tri- to multiseriate,
Typus: Madagascar, Moramanga, on Eucalyptus sp., Apr. 1994, thin-walled, guttulate, not to very slightly constricted at septum,
P.W. Crous (PREM 51749, holotype of sexual morph; PREM obovoid, remaining hyaline.
51748, holotype of asexual morph, cultures ex-type CBS
110682 = CMW 4942 = CPC 760, CPC 761). Type species: Paramycosphaerella brachystegia Crous & Jol.
Roux
Pallidocercospora heimioides (Crous & M.J. Wingf.) Crous &
M.J. Wingf., Stud. Mycol. 75: 74. 2012. Note: Paramycosphaerella marksii is a common foliar pathogen
Basionym: Pseudocercospora heimioides Crous & M.J. Wingf., on eucalypts, easily recognised by large leaf spots that have a
Canad. J. Bot. 75: 787. 1997. thin red-purple margin, and ascospores that are fusoid-ellipsoidal
Synonym: Mycosphaerella heimioides Crous & M.J. Wingf., with an asymmetrical apical cell.
Canad. J. Bot. 75: 787. 1997. Paramycosphaerella intermedia (M.A. Dick & K. Dobbie)
Diagnosis: Leaf spots not observed. Ascospores multiseriate, Quaedvlieg & Crous, Persoonia 33: 23. 2014.
overlapping, hyaline, guttulate, thin-walled, straight to slightly Basionym: Mycosphaerella intermedia M.A. Dick & K. Dobbie,
curved, fusoid-ellipsoidal with obtuse ends, widest in middle of New Zealand J. Bot. 39: 272. 2001.
apical cells, medianly 1-septate, not constricted at septum, Diagnosis: Leaf spots amphigenous, up to 10 mm diam,
tapering toward both ends, but slightly so toward base, (7.5–) becoming confluent, pale brown on abaxial surface, rust-brown
8–10(–11) × (2–)2.5–3 μm. Conidia solitary, terminal, with dark brown margin on adaxial surface with red-purple
CROUS ET AL.
zone. Ascospores hyaline, straight to curved, apices obtuse, MBT388177, culture ex-epitype CBS 110920 = CPC 935); Vic-
medianly 1-septate, widest in middle of apical cell, toria, Nowa Nowa, on E. botryoides, 11 Nov. 1990, A.J. Carnegie
12–16 × 2–4 μm. (holotype IMI 353731).
Descriptions and illustrations: Dick & Dobbie (2001), Quaedvlieg Notes: Disease symptoms associated with species of Para-
et al. (2014). mycosphaerella can be severe, although defoliation has not been
observed as a result of infection, as in the case of
Typus: New Zealand, Bay of Plenty, Rotoehu Forest, Kohekohe
Teratosphaeria.
Road, on E. saligna, 30 Jun. 1998, L. Renney (holotype NZFRI-
M 3831, culture ex-type NZFS 301.10 = CBS 114356 = CMW Parapallidocercospora Videira et al., Stud. Mycol. 87: 310.
7163 = CPC 10902 = NZFS 301 K/1). 2017.
Paramycosphaerella marksii (Carnegie & Keane) Quaedvlieg Leaf spots amphigenous, irregular to subcircular. Ascomata
& Crous, Persoonia 33: 23. 2014. Fig. 70. pseudothecial, predominantly hypophyllous, black, subglobose
Basionym: Mycosphaerella marksii Carnegie & Keane, Mycol. to globose, with apical ostiole, walls of 2–3 layers of medium
Res. 98: 414. 1994. brown textura angularis. Asci fasciculate, bitunicate, subsessile,
cylindrical to narrowly ellipsoidal, straight or slightly incurved.
Diagnosis: Leaf spots amphigenous, subcircular to irregular, Ascospores bi- to multiseriate, overlapping, hyaline, guttulate,
3–20 mm diam, pale brown, surrounded by raised, medium thin-walled, straight to slightly curved, fusoid-ellipsoidal, obovoid,
brown borders, and frequently with red-purple margins. Asco- medianly 1-septate, not constricted at septum or only slightly
spores bi- to multiseriate, overlapping, colourless, guttulate, thin- constricted, tapering toward both ends but more prominently
walled, straight to slightly curved, fusoid-ellipsoidal with an toward the base. Spermogonia intermixed with the ascomata or
obtuse basal and asymmetrical apical cell, widest in middle of the with the asexual morph, hyaline and rod-shaped. Mycelium in-
apical cell, medianly 1-septate, not constricted at septum, ternal and external, hyphae light brown, septate, branched,
tapering toward both ends, but with more so toward base, (11–) smooth. Conidiophores arising from superficial mycelium, from
12–14(–16) × 2–2.5(–3) μm. the upper cells of a brown stroma; conidiophores light brown,
Descriptions and illustrations: Carnegie & Keane (1994), smooth, aseptate or septate, subcylindrical, straight to variously
Quaedvlieg et al. (2014). curved, unbranched. Conidiogenous cells terminal, unbranched,
light brown, smooth, tapering to flat-tipped apical loci, prolifer-
Typus: Australia, Victoria, Briagolong, on E. globulus, 14 Oct. ating sympodially, rarely percurrently near apex. Conidia solitary,
1994, A.J. Carnegie (PREM 51932, epitype designated here, light brown, smooth to finely verruculose, septate, guttulate,
Fig. 70. Paramycosphaerella marksii (CPC 14655). A. Disease symptoms (note typical red margin). B. Ascomata forming in culture. C. Ascomatal periphyses. D–F. Asci. G.
Ascospores. Scale bars = 10 μm.
216
narrowly obclavate or subcylindrical, tapering towards the base, Parapallidocercospora thailandica (Crous et al.) Videira &
straight to curved (from Videira et al. 2017). Crous, Stud. Mycol. 87: 310. 2017.
Basionym: Mycosphaerella thailandica Crous et al., Stud. Mycol.
Type species: Parapallidocercospora colombiensis (Crous et al.)
50: 465. 2004.
Videira & Crous
Synonyms: Pseudocercospora thailandica Crous et al., Stud.
Notes: Morphologically, these taxa appear typical members of Mycol. 50: 465. 2004.
Pseudocercospora s. str. and are difficult to identify in the Pallidocercospora thailandica (Crous et al.) Phook. et al., Fungal
absence of DNA sequence data. Little is known about their Diversity 80: 21. 2016.
biology, but the fact that Parapallidocercospora thailandica oc-
Diagnosis: Ascospores tri- to multiseriate, overlapping, hyaline,
curs on more than one host plant genus in Thailand suggests
guttulate, thin-walled, fusoid-ellipsoidal medianly 1-septate,
that it deserves further study.
widest in middle of the apical cell, slightly constricted at the
Parapallidocercospora colombiensis (Crous & M.J. Wingf.) septum, (9–)10–11(–12) × (2–)2.5–3 μm. Conidia solitary, pale
Videira & Crous, Stud. Mycol. 87: 310. 2017. Fig. 71. brown, smooth, guttulate, narrowly obclavate to subcylindrical,
Basionym: Pseudocercospora colombiensis Crous & M.J. apex subobtuse, base long obconically subtruncate, straight to
Wingf., Mycol. Mem. 21: 42. 1998. curved, 3–6-septate, (25–)30–45(–60) × 2–2.5(–3) μm; hila
Synonym: Mycosphaerella colombiensis Crous & M.J. Wingf., inconspicuous.
Mycol. Mem. 21: 41. 1998.
Diagnosis: Leaf spots amphigenous, irregular to subcircular,
1–15 mm diam, pale brown, with raised dark brown borders; Typus: Thailand, Chachoengsao Prov., Sanamchaikhet, on
coloured margin absent. Ascospores bi- to triseriate, over- Acacia mangium, 28 May 2003, K. Pongpanich (holotype CBS
lapping, hyaline, guttulate, thin-walled, straight to slightly curved, H-9875, of both M. thailandica and P. thailandica, cultures ex-
obovoid with an obtuse base and rounded apex, widest near type CBS 116367 = CPC 10547, CPC 10548, 10549).
apex, medianly 1-septate, not constricted at septum, tapering Parapenidiella Crous & Summerell, Persoonia 29: 185. 2012.
toward both ends, but more prominently toward base (11–)
12–14(–15) × 3–3.5(–4) μm. Conidia solitary, light brown, Mycelium consisting of branched, septate, smooth subhyaline to
smooth to finely verruculose, guttulate, narrowly obclavate or pale brown hyphae. Conidiophores macronematous, occasion-
subcylindrical, apex obtuse, base long obconic-truncate or cy- ally micronematous; macronematous conidiophores arising from
lindrical and truncate, straight to curved, 1–5-septate, (25–) superficial mycelium, solitary, erect, pale brown, thin-walled,
30–45(–60) × 2.5–3(–3.5) μm. smooth to finely verruculose; terminally penicillate, unbranched
in terminal part; conidiogenous apparatus composed of a series
Description and illustration: Crous (1998). of conidiogenous cells and/or ramoconidia. Conidiogenous cells
Typus: Colombia, Pinal Farm, on E. urophylla, May 1995, M.J. integrated, terminal or intercalary, unbranched, pale brown,
Wingfield (holotype PREM 54397, cultures ex-type CBS smooth, tapering to a flattened or rounded apical region, mono-
110968 = CPC 1105, CPC 1104, 1106). or polyblastic, sympodial, giving rise to a single or several sets of
ramoconidia on different levels; with relatively few conidiogenous
loci, slightly thickened, slightly darkened. Conidia in branched
acropetal chains. Ramoconidia 0–1-septate, pale brown,
smooth, thin-walled, fusoid-ellipsoidal to subcylindrical. Conidia
subcylindrical, fusoid to ellipsoid-ovoid, aseptate, pale olivaceous
to pale brown, smooth, thin-walled, catenate; hila truncate,
slightly thickened, somewhat darkened.
Type species: Parapenidiella tasmaniensis (Crous & M.J. Wingf.)
Crous
Notes: Parapenidiella is distinguished from Penidiella by having
pale brown, unbranched, penicillate conidiophores, with oliva-
ceous to pale brown, long, branched conidial chains.
Parapenidiella pseudotasmaniensis (Crous) Crous, Persoonia
29: 185. 2012. Fig. 72.
Basionym: Penidiella pseudotasmaniensis Crous, Persoonia 23:
126. 2009.
Diagnosis: Leaf spots amphigenous, irregular to subcircular or
circular, 2–5 mm diam, medium brown, with a raised border and
thin, red-purple margin. Ascospores bi- to triseriate, overlapping,
hyaline, guttulate, thin-walled, straight, fusoid-ellipsoidal with
obtuse ends, widest in middle of apical cell, medianly 1-septate,
constricted at the septum, tapering towards both ends, but more
prominently towards the lower end, prominently guttulate,
Fig. 71. Parapallidocercospora colombiensis (PREM 54397). Asci, ascospores,
germinating ascospores, conidiogenous cells and conidia. Scale bars = 10 μm.
covered in mucilaginous sheath, which largely disappears at
(from Crous 1998). maturity, (8–)9(–10) × 3(–3.5) μm. Secondary ramoconidia
CROUS ET AL.
Fig. 72. Parapenidiella pseudotasmaniensis (CPC 12400). A. Disease symptoms. B. Leaf spot. C. Somewhat superficial ascomata. D. Broken ascoma with asci. E–G. Asci. H.
Germinating ascospore. I. Colony on PDA. J. Sporulation on pine needle. K–N. Conidiophores giving rise to catenulate conidia. Scale bars: C = 100 μm, all others = 10 μm.
218
subcylindrical, aseptate, pale brown, smooth, subcylindrical to Diagnosis: Leaf spots amphigenous, grey to pale brown, sub-
narrowly fusoid, 5–9 × 1.5–2 μm. Intercalary and terminal circular to angular, 1–4 mm diam, borders light brown, coloured
conidia aseptate, pale brown, smooth, subcylindrical to narrowly margins absent. Conidia solitary, olivaceous to pale brown,
fusoid, 5–9 × 1.5–2 μm; scars thickened and somewhat verruculose, thick-walled, obclavate, apex broadly rounded, base
darkened. obconic-truncate, straight to curved, 1–3(–7)-septate, (30–)
35–45(–55) × 4–5(–6) μm; hila thickened, darkened, refractive,
2 μm wide.
Typus: Australia, Victoria, on E. globulus, Sep. 2005, coll. I.W.
Smith, isol. P.W. Crous (holotype CBS H-20252, cultures ex-
type CPC 12400 = CBS 124991, CPC 12401, 12402). Typus: Australia, Queensland, Coopers, on E. morrisii, 29 May
1967, F.D. Hockings (holotype IMI 151656). Not known from
Parapenidiella tasmaniensis (Crous & M.J. Wingf.) Crous,
culture.
Persoonia 29: 185. 2012.
Basionym: Mycovellosiella tasmaniensis Crous & M.J. Wingf., Brux. 19: 300.
Pestalotiopsis Steyaert, Bull. Jard. bot. Etat
Mycol. Res. 102: 527. 1998. 1949. Fig. 74.
Synonyms: Passalora tasmaniensis (Crous & M.J. Wingf.) Crous
Conidiomata acervular, epidermal to subepidermal, separate or
& U. Braun, CBS Diversity Ser. (Utrecht) 1: 472. 2003.
confluent, formed of brown, thin-walled of textura angularis;
Mycosphaerella tasmaniensis Crous & M.J. Wingf., Mycol. Res.
dehiscence irregular. Conidiophores hyaline, branched and
102: 527. 1998.
septate at the base and above, cylindrical or lageniform, formed
Diagnosis: Leaf spots amphigenous, subcircular, 2–30 mm from the upper cells of the pseudoparenchyma. Conidiogenous
diam., grey to light brown, separate, but coalescing with age to
form large blotches, surrounded by a raised, medium brown
border and a diffuse, red-purple margin. Ascospores tri- to
multiseriate, overlapping, colourless, guttulate, thin-walled,
straight, fusoid-ellipsoidal with obtuse ends, widest in middle of
apical cell, medianly 1-septate, not constricted at septum,
tapering toward both apices, but with more prominent taper to-
wards lower end (10–)11–12(–13) × (2.5–)3(–4) μm. Conidia
catenulate, chains branched, olivaceous, smooth, eguttulate,
subcylindrical, narrowly ellipsoidal or fusiform, tapering toward
rounded ends with flattened, darkened, refractive, thickened
loci, straight to slightly curved, 0(–1)-septate, (4–)
8–12(–20) × 2–2.5 μm in vivo and in vitro.
Typus: Australia, Tasmania, on E. nitens, Nov. 1996, M.J.
Wingfield (holotypes PREM 55339 of sexual morph, PREM
55340 of asexual morph, cultures ex-type CBS 111687 = CMW
14780 = CPC 1555, CBS 114556 = CMW 14663 = CPC 1556,
CPC 1557).
Passalora Fr., Summa veg. Scand., Sectio Post. (Stockholm):
500. 1849.
Mycelium internal, consisting of hyaline, branched, septate hy-
phae. Stromata absent or small. Conidiophores emerging
through stomata, in fascicles, unbranched or branched, straight
to flexuous, at times with a single basal septum, usually up to 3-
septate, medium brown, somewhat swollen in the conidiogenous
region. Conidiogenous cells integrated, terminal, with flat,
somewhat thickened and darkened loci. Conidia solitary, oliva-
ceous to pale brown, thin-walled, smooth, straight or gently
curved, mostly didymosporous, constricted at septum, with
somewhat thickened, darkened and refractive hila (from Videira
et al. 2017).
Type species: Passalora bacilligera (Mont. & Fr.) Fresen.
Note: The phylogenetic relationships of this species cannot be
resolved without fresh collections, but it clearly represents a
genus distinct from Passalora s. str.
“Passalora” morrisii Crous, Mycol. Mem. 21: 108. 1998. Fig. 73. Passalora morrisii (IMI 151656). Conidiophores and conidia. Scale
Fig. 73. bar = 10 μm (from Crous 1998).
CROUS ET AL.
Fig. 74. Pestalotiopsis spp. A. Disease symptoms on Eucalyptus sp. in a nursery. B. Emerging ascomata neck. C–E. Asci with ascospores. F. Conidiomata sporulating on
PDA. G, H. Conidiogenous cells. I. Conidia. Scale bars = 10 μm.
cells holoblastic, annellidic, indeterminate, integrated, cylindrical, Pestalotiopsis colombiensis Maharachch. et al., Stud. Mycol.
hyaline, smooth, with several percurrent proliferations. Conidia 79: 158. 2014.
fusiform, straight or slightly curved, 4-euseptate; basal cell hy-
aline, truncate, with an endogenous, cellular, simple or rarely Diagnosis: Conidia ellipsoid, straight to slightly curved,
branched appendage; apical cell conic, hyaline, with 2 or more 4-septate, slightly constricted at septa, (19–)21–27(–28.5) ×
apical, simple or branched, spathulate or espathulate append- 5.5–7.5(–8) μm; basal cell conic to acute with truncate base,
ages; median cells brown, sometimes versicoloured, thicker- minutely verruculose and thinwalled, 5–7.5 μm long; three me-
walled, smooth or verruculose dian cells, (13–) 13.5–16.5(–17) μm long, doliiform, thick-
walled, verruculose, concolourous, brown (second cell from
Type species: Pestalotiopsis guepinii (Desm.) Steyaert
base 5–6.5 μm long; third cell 4.5–6 μm long; fourth cell
Symptoms: Causing necrotic leaf spots and blights. 5–6.5 μm long); apical cell cylindrical to subcylindrical, hyaline,
thin- and smooth-walled, 3.5–5 μm long; with 2–3 tubular apical
Notes: Pestalotiopsis spp. are generally regarded as endophytic
appendages (mostly 3), arising from the apical crest, un-
fungi of minor importance, although they can be associated with
branched, filiform, (11–)13–25(–28) μm; basal appendage sin-
severe infections in nurseries. It is commonly isolated as a
gle, tubular, unbranched, centric, 2–5 μm long.
saprophyte from eucalypt leaves damaged by other agents, such
as herbicide, drought, insects or other fungi. Description and illustration: Maharachchikumbura et al. (2014).
220
Typus: Colombia, from living leaves of E. urophylla × E. grandis, cylindrical-clavate to clavate, crozier present, usually 8- or 4-
2004, M.J. Wingfield (holotype CBS H-21764, culture ex-type spored; apical apparatus with a well-developed apical thick-
CBS 118553 = CPC 10969). ening, IKI + or −, Mlz−, but often blue after pretreatment with
KOH. Ascospores inequilateral, ovoid, ellipsoid, or fusoid,
Pestalotiopsis disseminata (Thüm.) Steyaert, Bulletin Jard. straight or curved, thin-walled, smooth, hyaline, filled with
Bot. l Etat Bruxelles 19: 319. 1949. numerous oil droplets, aseptate; mostly later 1–3(–7)-septate, or
Basionym: Pestalotia disseminata Thüm., Inst. Coimbra: no. 578. muriform, sometimes forming clavate to cylindrical, aseptate,
1879. hyaline, thin- and smooth-walled conidia from minute openings or
phialides produced during germination. Paraphyses filiform,
Diagnosis: Conidia broadly fusoid to fusoid-clavate, straight or
septate, simple or branched, hyaline, smooth-walled, apical cells
somewhat curved, 5-celled, upper cell conical to cylindrical,
often swollen. Conidiomata immersed or erumpent, eustromatic
hyaline, fairly thin-walled, apical setulae central, (2–)3(–4),
with a single or several fusing cavities formed on basal stroma,
rather stout, up to 1.2 μm wide, 11–20 μm long, with a blunt tip,
subglobose, irregularly pulvinate to conical or claviform;
three intermediate cells concolourous or the upper two inter-
sometimes stromatic acervular, almost plane to pustulate. Co-
mediate cells slightly darker, dull olivaceous-brown to vinaceous-
nidiophores simple or branched, smooth, hyaline, acrogenous or
brown, contents guttulate, walls smooth, slightly constricted at
acropleurogenous. Conidiogenous cells discrete or integrated,
the septa when mounted in water, and thickened up to 1 μm
determinate, phialidic, or indeterminate, proliferating percurrently,
especially in the upper two intermediate cells and in the septa,
cylindrical to narrowly ampulliform, giving rise to macro- and/or
basal cell hyaline, thin-walled, tapering into a filiform pedicel (2–)
microconidia. Macroconidia present or absent, ellipsoid, pyriform,
2.5–4.5(–5) μm long; conidium body (18–)
claviform or fusoid, rounded or somewhat pointed at apex,
20–24(–25) × 6.5–7(–8) μm.
rounded or attenuated and with a barely or distinctly protruding
Description and illustration: Crous et al. (2006c). scar at base, aseptate, thin-walled, hyaline, smooth, mostly filled
Material examined: New Zealand, North Island, Kerikeri, on living leaves of
with numerous oil droplets, becoming 1–3(–6)-septate, finally
E. botryoides, 17 Oct. 2003, M.A. Dick, cultures CBS 118552 = CPC 10950, CPC often muriform; sometimes forming microconidia from minute
10951). opening in wall or from phialides. Microconidia present or absent,
Pezicula Tul. & C. Tul., Sel. Fung. Carpol. 3: 182. 1865. cylindrical, apex rounded, base truncate, aseptate, hyaline,
smooth, thin-walled, contents granular. (Adapted from Verkley
Synonyms: Ocellaria (Tul. & C. Tul.) P. Karst., Bidr. K€ann. Finl.
1999, Chen et al. 2016).
Nat. Folk 19: 21, 242. 1871.
Stictis subgen. Ocellaria Tul. & C. Tul., Select. Fung. Carpol. 3: Type species: Pezicula carpinea (Pers.) Tul. & C. Tul. ex Fuckel
128. 1865.
Symptoms: Leaf spots amphigenous, subcircular to irregular,
Dermatella P. Karst., Bidr. K€ann. Finl. Nat. Folk 19: 16, 209.
medium brown.
1871.
Dermatina (Sacc.) Höhn., Sber. Akad. Wiss. Wien, Math.-naturw. Notes: Pezicula californiae clusters basal to other taxa in the
Kl., Abt. 1 118: 1512. 1909. genus Pezicula, and although we were originally of the opinion
Cryptosporiopsis Bubak & Kabat, Hedwigia 52: 360. 1912. that it represented a distinct genus, it was eventually retained in
Pachydiscula Höhn., Z. G€arungsphysiol. 5: 210. 1915. Pezicula, pending further collections (Chen et al. 2016). Pezicula
Lagynodella Petr., Annls Mycol. 20: 207. 1922. is a genus of well-known plant pathogens, but the species
Additional synonyms in Verkley (1999). presently known from eucalypts appear to be of relatively minor
importance.
Apothecia erumpent, sessile to short-stalked, solitary or in
clusters on a basal stroma. Disc circular to irregular, pruinose, Pezicula californiae (Cheew. et al.) P.R. Johnst., IMA Fungus 5:
pale whitish, yellow, orange, olivaceous, or orange-brown to dark 103. 2014. Fig. 75.
brown, 0.2–3 mm diam. Margin first entire, often with a slightly Basionym: Cryptosporiopsis californiae Cheew. et al., Fungal
raised rim, persistent, or irregularly torn. Asci inoperculate, Diversity 44: 91. 2010.
Fig. 75. Pezicula californiae (CPC 32352). A. Disease symptoms on leaves of E. globulus. B. Conidiomata sporulating on OA. C, D. Conidiogenous cells. E. Conidia. Scale
bars = 10 μm.
CROUS ET AL.
Diagnosis: Conidiomata pycnidial to acervular, superficial or Phacidiella eucalypti Crous, Fungal Diversity 25: 30. 2007.
partly immersed, medium to dark brown, with cream conidial Fig. 76.
masses. Conidia elongate ellipsoidal, straight or slightly
Diagnosis: Leaf spots amphigenous, medium brown, subcircular,
curved, some inaequilateral, apex obtuse or broadly acute,
with dark brown margin. Conidia subhyaline to pale brown,
tapering abruptly to a slightly protruding scar at the base,
smooth, subcylindrical to barrel-shaped, ends bluntly rounded,
1.5– 2 μm wide; aseptate, hyaline, thick-walled, with 5
occurring in disarticulating, unbranched, short chains, (4–)
30 guttules per conidium, (12.5 –)15– 18(– 27.5) × (4.2 –)
5–6(–7) × (2–)2.5 μm.
4.5– 5.2( –5.8) μm.
Typus: South Africa, Western Cape Province, Stellenbosch
Typus: USA, California, on Eucalyptus sp., Mar. 2009, S. Den-
Mountain, on Eucalyptus sp., 10 Jan. 2006, P.W. Crous (holo-
man (holotype CBS H-20302, cultures ex-type CBS
type CBS H-19768, cultures ex-type CBS 120255 = CPC 12745,
124819 = CPC 13819, CPC 13820, 13821).
CPC 12746, 12747).
Phacidiella P. Karst., Hedwigia 23: 85. 1884.
Phacidium Fr., Observ. Mycol. 1: 167. 1815; nom. cons. Fig. 77.
Conidiomata eustromatic, immersed, separate, rarely aggre- Synonyms: Phacidiostroma Höhn., Ber. dt. bot. Ges. 35: 420.
gated, dark brown, initially globose, later opening via irregular 1917.
splits, becoming cupulate; wall of pale brown, thin textura Ceuthospora Grev., Scot. Crypt. Flora 5: 253. 1827; nom. cons.
angularis, lateral walls of textura porrecta. Conidiophores hya-
Foliicolous or caulicolous. Ascomata amphigenous, scattered or
line, branched at base, septate, forming from the base as well as
gregarious, circular, immersed, becoming erumpent, rupturing host
laterally. Conidiogenous cells holothallic, integrated, hyaline,
tissue by irregular stellate splits, of dark brown pseudoparenchymatal
smooth, cylindrical. Conidia arthric, disarticulating from con-
cells of textura globulosa, inner layer with periphysoids, invested in
idiogenous cells, producing long branched chains, with youngest
mucilage. Hymenium of asci and paraphyses; basal stroma present
conidia at the base, hyaline, straight, smooth, ends truncate, but
or absent. Asci clavate, (4–)8-spored, with amyloid (staining blue in
apical conidium with subacute apex, long doliiform, aseptate
Melzer’s reagent) apical discharge mechanism. Ascospores asep-
(from Sutton 1980).
tate, ellipsoid to ellipsoid-fusoid, uni- to biseriate, hyaline, smooth,
Type species: Phacidiella salicina P. Karst. lacking mucoid appendages. Paraphyses septate, hyaline, smooth,
branched, anastomosing, invested in mucilage. Conidiomata pyc-
Note: Considered to be a minor pathogen. nidial, immersed, becoming erumpent, uni- to multilocular, brown,
Fig. 76. Phacidiella eucalypti (CBS H-19768, CBS 120038). A, B. Leaf spots with conidiomata. C. Colony on PDA. D–H. Conidia and conidiogenous cells. Scale bars = 10 μm
(from Crous et al. 2007b).
222
Fig. 77. Phacidium spp. A– H. Phacidium sp. A, D. Ascomata on leaf. B, C. Vertical section through ascoma. E– G. Asci. H. Ascospores. I – O. Phacidium
mollerianum (CBS 574.66). I. Conidiomata on autoclaved barley leaves. J – M. Conidiogenous cells. N, O. Conidia. Bars: A = 300 μm, all others = 10 μm (from Crous
et al. 2014a).
with ostiole; wall of textura angularis to textura globulosa. Co- Phacidium innumerum (Massee) Crous, comb. nov. Myco-
nidiophores branched or simple, septate, hyaline, smooth, invested in Bank MB832040.
mucilage. Conidiogenous cells phialidic, at times proliferating per- Basionym: Ceuthospora innumera Massee, Bull. Misc. Inf., Kew:
currently, subcylindrical to ampulliform, smooth, hyaline, invested in 182. 1899.
mucilage. Conidia subcylindrical, aseptate, hyaline, smooth, with
Diagnosis: Conidia are hyaline, cylindrical or slightly tapering,
irregular funnel-shaped apical mucilaginous appendage.
with rounded ends and a funnel-shaped, gelatinous apical
Type species: Phacidium lacerum Fr. appendage, 16–20 × 2.5–3 μm.
Symptoms: Associated with inconspicuous brown leaf spots. Description and illustration: Swart (1988).
This genus is a minor pathogen of Eucalyptus, and is usually
Typus: Australia, Tasmania, Rodway, on Eucalyptus sp.,
found sporulating on leaf litter, or dead leaf tips.
collection date and collector unknown (holotype K(M) 190756).
CROUS ET AL.
Notes: Conidiomata densely aggregated on one side of the leaf, Typus: Kenya, Maragua, Central Province, on E. grandis, 13 Jul.
with conidia 16–20 × 2.5–3 μm (Swart 1988). Conidiomata are 2013, J. Roux (holotype PREM 61155).
recognised under the dissecting microscope by having multiple,
Phaeophleospora Rangel, Arq. Mus. Nac., Rio de Janeiro 18:
somewhat papillate, black, slightly protruding, prominent ostioles.
162. 1916.
Phacidium mollerianum (Thüm.) Crous, IMA Fungus 5: 187.
Conidiomata pycnidial, aggregated or separate, becoming
2014. Fig. 76.
erumpent, lifting the epidermis; pycnidia black, subglobose,
Basionym: Sphaeropsis molleriana Thüm., Inst. Coimbra 27: 40.
unilocular; wall of brown textura epidermoidea in surface view,
1879.
and of textura angularis to textura intricata in vertical section,
Synonyms: Phoma molleriana (Thüm.) Sacc., Syll. fung. 3: 110.
base of 2–3 layers; ostiole irregular, central. Conidiophores
1884.
mostly reduced to conidiogenous cells, or multi-septate, sub-
Macrophoma molleriana (Thüm.) Berl. & Voglino, Syll. fung.,
cylindrical, branched or not, brown, verruculose. Conidiogenous
Addit. I: 314. 1886.
cells terminal, discrete, brown, verruculose, subcylindrical or
Ceuthospora molleriana (Thüm.) Petr., Annls Mycol. 23: 29.
doliiform, with several inconspicuous percurrent proliferations, or
1925.
at times with sympodial proliferation. Conidia solitary, exuded in
Diagnosis: Conidia hyaline, smooth, granular, subcylindrical, chirrus, subcylindrical to obclavate, apex obtuse, base obconi-
(9–)10–12(–13) × (2–)2.5 μm; apical mucoid appendage only cally truncate, thick-walled, euseptate, straight to irregularly
visible when mounted in water. curved, hyaline to subhyaline, smooth to verruculose, granular;
hila with a minute marginal frill. Spermatogenous cells devel-
oping in conidiomata before the development of conidia, hyaline,
Materials examined: Italy, Sardinia, Su Cologone, on Eucalyptus sp., 6 May 1971, ampulliform. Spermatia hyaline, smooth, rod-shaped.
W. Gams, CBS H-10285, 10286, culture CBS 365.72. The Netherlands, Baarn,
on Polygonatum odoratum, 8 Aug. 1966, H.A. van der Aa, culture CBS 574.66. Type species: Phaeophleospora eugeniae Rangel
Notes: Ceuthospora molleriana was originally described from
Eucalyptus leaves collected in Portugal. Conidia are hyaline, Notes: Phaeophleospora represents an important genus of foliar
smooth, granular, subcylindrical, (9–)10–12(–13) × (2–)2.5 μm; pathogens of Myrtaceae, especially Eugenia, and to some extent
apical mucoid appendage only visible when mounted in water also Eucalyptus. However, the generic concept has widened
(Crous et al. 2014a). considerably, and probably contains some taxa that will even-
tually be placed elsewhere, as they are morphologically quite
Phakopsora Dietel, Ber. dt. bot. Ges. 13: 333. 1895. distinct from the type.
Pycnidia and Aecia unknown. Uredinia subepidermal, erumpent, Phaeophleospora scytalidii (Crous & M.J. Wingf.) Quaedvlieg
surrounded by encircling, incurved paraphyses. Urediniospores & Crous, Persoonia 33: 23. 2014.
solitary, obovate-globoid to ellipsoid, pale yellow, with obscure Basionym: Mycosphaerella scytalidii Crous & M.J. Wingf., Stud.
germ pores. Telia subepidermal, non-erumpent, lenticular, black. Mycol. 55: 120. 2006.
Teliospores aseptate, chestnut brown to golden brown; formed in
succession from basal hymenium. Diagnosis: Leaf spots amphigenous, irregular to sub-circular,
1–8 mm diam, grey to medium brown, with a raised, dark
Type species: Phakopsora punctiformis (Barclay & Dietel) Dietel brown border. Ascospores tri- to multiseriate, overlapping, hya-
line, guttulate, thin-walled, straight, fusoid–ellipsoidal with sub-
Notes: Austropuccinia psidii and P. myrtacearum are the only two obtuse ends, medianly 1-septate, widest in the middle of the
rust species confirmed from eucalypts. Phakopsora myrtacea- apical cell, constricted at the septum, tapering towards both
rum is widespread in southern and eastern Africa (Kenya, ends, but more prominently towards the lower end,
Mozambique, South Africa), where it occurs on E. cloeziana, 8–10 × (2.5–)3 μm in vivo.
E. grandis and E. nitens. It is considered an important quarantine
pathogen that may cause damage to eucalypt forestry and other Description and illustration: Crous et al. (2006d).
species of Myrtaceae (Maier et al. 2015). Typus: Colombia, Angela Maria, on E. urophylla, Jan. 2004, M.J.
Phakopsora myrtacearum McTaggart et al., Pl. Pathol. 65: 192. Wingfield (holotype CBS H-19696, culture ex-type CBS
2015. Fig. 78. 118493 = CPC 10998).
Diagnosis: Leaf spots amphigenous, small, brown, angular, Phaeoramularia Munt.-Cvetk., Lilloa 30: 182. 1960.
coalescing with age, with pale yellow to pale brown uredinia. Phytopathogenic, usually forming leaf spots, occasionally almost
Spermogonia, aecia and telia unknown. Uredinia amphigenous, symptomless. Mycelium internal, composed of subhyaline to
the majority hypophyllous, 0.1–0.5(–1) mm diam, sometimes on pigmented hyphae, septate, branched, smooth to rough. Stro-
vein-limited, raised lesions, approx. 0.3–1 mm, scattered or mata almost absent to well-developed, pigmented. Co-
aggregated in groups, sub-epidermal in origin, erumpent, sur- nidiophores macronematous, mononematous, in small to large
rounded by torn epidermis, pale brown, paraphyses absent. fascicles, rarely solitary, arising from internal hyphae or stromata,
Urediniospores obovoid, mostly pyriform to ellipsoid, (20–) emerging through stomata or erumpent through the cuticle, erect,
23–26(–30) × 14–17(–20) μm (av. 24.5 × 15.8 lm), walls straight, subcylindrical to flexuous, geniculate-sinuous, simple,
2–3 μm thick, sub-hyaline to pale brown; spore walls finely and rarely branched, continuous to septate, pale yellowish green,
densely echinulate, germ pores inconspicuous (from Maier et al. olivaceous to brown, smooth to rough, thin-walled. Con-
2015). idiogenous cells integrated, terminal, occasionally intercalary,
Description and illustration: Maier et al. (2015). sometimes conidiophores reduced to a single conidiogenous
224
Fig. 78. Phakopsora myrtacearum (PREM 61155). A, B. Uredinia and lesions on abaxial leaf surface. C, D. Urediniospores, equatorial and surface views, respectively. E.
Urediniospores under scanning electron microscope. Scale bars C–E = 10 μm (from Maier et al. 2015).
cell, polyblastic, proliferation sympodial, rarely percurrent, con- Phaeothyriolum Syd., Annls Mycol. 36: 305. 1938. Fig. 80.
idiogenous loci thickened and darkened. Conidia catenate, Synonym: Murramarangomyces corymbiae Crous, Persoonia
sometimes in branched chains, ellipsoid-ovoid, subcylindrical, 39: 385. 2017.
fusiform, continuous to euseptate, subhyaline to pigmented,
Leaf spots circular, indistinct, confined to one side of the leaf.
smooth to rough, ends obtuse, truncate or subacute; hila thick-
Mycelium interepidermal, brown, septate, branched, giving leaf
ened and darkened; conidial secession schizolytic (see Videira
spots a red-purple appearance without any superficial mycelium
et al. 2017).
being present; mycelium aggregating in substomatal cavity,
Type species: Phaeoramularia gomphrenicola (Speg.) Munt.- giving rise to suprastomatal ascomata, either on living leaves, or
Cvetk. on leaf litter. Ascomata thyrothecial, in radial orientation or
clusters, circular, brown, superficial, flattened, with lobed, smooth
Note: Fresh collections are required to resolve the phylogenetic
to irregular margin of textura epidermoidea or angularis. Asci
relationships of this species, which is probably not a member of
ellipsoid-obclavate, pedicellate, 8-spored, with small ocular
Phaeoramularia s. str.
chamber. Pseudoparaphyses hyaline, septate, anastomosing or
Phaeoramularia eucalyptorum Crous, Mycol. Mem. 21: 117. not, hypha-like, dissolving at maturity, or persistent. Ascospores
1998. Fig. 79. tri- to multiseriate, hyaline, 1-septate, guttulate, granular, slightly
constricted at median septum or not, fusoid-ellipsoid to obovoid,
Diagnosis: Leaf spots amphigenous, variable from small specks
encased in prominent mucoid sheath that is mostly absent at
to subcircular spots, occurring along the leaf margin, pale brown,
maturity, becoming brown once discharged onto agar. Mycelium
border raised, margin red-brown. Conidia catenulate, pale
in vitro consisting of hyaline, smooth hyphae, irregular, con-
olivaceous to subhyaline, smooth, thin-walled, guttulate, sub-
stricted at septa, forming brown, thick-walled cells that become
cylindrical to narrowly obclavate, apex obtuse, base narrow, long
fertile conidiogenous cells, aggregated in cauliflower-like clusters
obconic-truncate, occurring in unbranched chains, 1–6-septate,
of brown, doliiform to globose, brown conidiogenous cells,
(15–)25–50(–90) × 2–2.5(–3) μm; hila thickened, darkened,
phialidic, giving rise to solitary conidia. Conidia in slimy mass,
refractive.
aseptate, hyaline, smooth, bacilliform.
Type species: Phaeothyriolum eucalyptinum Syd.
Typus: Malaysia, Cameron highlands, Cameron nursery, on
Phaeothyriolum amygdalinum (Cooke & Massee) Crous &
E. saligna, May 1995, M.J. Wingfield (holotype PREM 54405).
Carnegie, comb. nov. MycoBank MB832041. Fig. 80.
CROUS ET AL.
16–18 × 6–7 μm (Hansford 1956, Swart 1986c). This species is

common in Corymbia plantations in eastern Australia (Carnegie
et al. 2008, as Phaeothyriolum microthyrioides), as well as native
Corymbia forest, but never associated with defoliation.
Phaeothyriolum corymbiae (Crous) Crous, comb. nov.
Mycobank MB832042. Fig. 80.
Basionym: Murramarangomyces corymbiae Crous, Persoonia
39: 385. 2017.
Leaf spots on living leaves, up to 20 mm diam, diffuse, circular,
border indistinct, confined to one side of the leaf. Ascomata
thyrothecial, suprastomatal, up to 180 μm diam, lacking super-
ficial hyphae, dark brown, scattered, flattened, with central
ostiole, star-shaped, 10–20 μm diam; margin irregular, lobed
textura angularis. Pseudoparaphyses hyaline, septate, hyphae-
like, anastomosing, 2–3 μm diam, dissolving at maturity. Asci
8-spored, bitunicate, broadly ellipsoid, stipitate, curved, apical
chamber weakly developed, 1 μm diam, (25–)
30–35(–38) × 12–14(–15) μm. Ascospores tri- to multiseriate,
medianly 1-septate, thick-walled, straight to slightly curved,
fusoid-ellipsoid, granular, guttulate, widest just above septum,
becoming constricted at septum, enclosed in mucoid sheath
extending over ends of spore, up to 5 μm diam, but dissolving at
maturity, (15–)16–17(–18) × 4(–4.5) μm; ascospores turn
brown at germination. Mycelium in vitro consisting of hyaline,
smooth, 2–5 μm diam hyphae, irregular, constricted at septa,
forming brown, thick-walled cells that become fertile con-
idiogenous cells, aggregated in cauliflower-like clusters of dolii-
form to globose, brown conidiogenous cells, 3–5 μm diam,
Fig. 79. Phaeoramularia eucalyptorum (PREM 54405). Conidiophores, con-
phialidic, giving rise to solitary conidia. Conidia in slimy mass,
idiogenous cells and conidia. Scale bar = 10 μm (from Crous 1998).
aseptate, hyaline, smooth, bacilliform, 3–5 × 1.5–2 μm.
Typus: Australia, New South Wales, close to Murramarang, on
Basionym: Microthyrium amygdalinum Cooke & Massee, Gre- C. maculata, 27 Nov. 2016, P.W. Crous (holotype of Murra-
villea 19(no. 92): 90. 1891. marangomyces corymbiae CBS H-23281, culture ex-type CBS
143434 = CPC 33000).
Leaf spots on living leaves, up to 20 mm diam, diffuse, circular,
border indistinct, confined to one side of the leaf. Ascomata
Additional material examined: Australia, New South Wales, Clouds Creek State
thyrothecial, suprastomatal, 100–160 μm diam, lacking su- Forest, on E. saligna, 21 Apr. 2018, A.J. Carnegie, HPC 2423 (sexual morph),
perficial hyphae, dark brown, scattered, flattened, with central culture CBS 145942 = CPC 35789.
ostiole, star-shaped, 15–30 μm diam; margin of even, lobed
Notes: Phaeothyriolum corymbiae resembles P. eucalyptinum
textura angularis. Pseudoparaphyses hyaline, septate, hyphae-
(asci 40–55 × 15–18 μm, ascospores 15–17 × 5–7 μm, with
like, anastomosing, 2–3 μm diam, dissolving at maturity. Asci
numerous pseudoparaphyses; Swart 1986c), but is distinct in
8-spored, bitunicate, broadly ellipsoid, stipitate, straight to
that it has smaller asci, narrower ascospores, and pseudopar-
curved, apical chamber weakly developed, 1–2 μm diam,
aphyses that dissolve at maturity. The asexual morph, Murra-
40–55 × (13–)15–18 μm. Ascospores tri- to multiseriate,
marangomyces corymbiae was associated with black
medianly 1-septate, broadly ellipsoid to obovoid, granular,
thyrothecia, although the sexual-asexual link could not be
guttulate, thick-walled, widest at septum or in middle of apical
confirmed at the time (Crous et al. 2017a). As we show here, it is
cell, not constricted at septum, enclosed in mucoid sheath
the asexual morph of Phaeothyriolum.
extending over ends of spore, up to 5 μm diam, (13–)
15–17(–18) × (6–)7 μm; ascospores turn brown at germina- Phaeothyriolum dunnii Crous & Carnegie, sp. nov. MycoBank
tion, and mature ascospores become constricted at septum, MB832043. Fig. 80.
and lack a sheath.
Etymology: Name reflects the host species from which it was
Typus: Australia, southeastern coast of South Australia, collected, Eucalyptus dunnii.
Spenser Gulf, on E. amygdalinus var. linearus (probably
Leaf spots developing on leaf litter, up to 15 mm diam, diffuse,
E. radiata), 1890, collector unknown (holotype K); New South
circular, border indistinct, confined to one side of the leaf.
Wales, Mallanganee, Sugar Loaf State Forest, on C. variegata,
Ascomata thyrothecial, suprastomatal, up to 250 μm diam,
17 Apr. 2018, A.J. Carnegie, HPC 2421 (CBS H-24030 epitype
lacking superficial hyphae, dark brown, scattered, flattened, with
designated here MBT388161, culture ex-epitype CPC 36084).
central ostiole, circular, 15–25 μm diam; margin of even, lobed
Notes: The present collection is a good fit for Microthyrium textura epidermoidea. Pseudoparaphyses hyaline, septate,
amygdalinum, with asci 50–60 × 15–20 μm, and ellipsoid to hypha-like, anastomosing, 1.5–2 μm diam, extending above
obovoid ascospores that become constricted at the septum, asci, persistent at maturity. Asci 8-spored, bitunicate,
226
Fig. 80. Phaeothyriolum spp. A–D. Disease symptoms. E–H. P. amygdalinum (HPC 2421). E. Thyrothecial ascomata. F, G. Asci. H. Ascospores. I–L. P. corymbiae (HPC
2423). I. Thyrothecial ascomata. J, K. Asci. L. Ascospores. M–P. P. dunnii (HPC 2433). M. Thyrothecial ascomata. N, O. Asci. P. Ascospores. Q–T. P. eucalyptorum (HPC
2269). Q. Thyrothecial ascomata. R, S. Asci. T. Ascospores. Scale bars = 10 μm.
CROUS ET AL.
subcylindrical to fusoid-ellipsoid, stipitate, curved, apical cham- PREM 51297); Western Cape Province, Stellenbosch, Stellen-
ber weakly developed, 1 μm diam, (38–)40–45(–55) × (9–) bosch Mountain, on Eucalyptus leaf litter, Sep. 1990, P.W. Crous
10–11(–12) μm. Ascospores tri- to multiseriate, medianly 1- (paratype PREM 51298); Western Cape Province, Stellenbosch,
septate, fusoid, granular, guttulate, widest at septum, not con- Stellenbosch Mountain, on Eucalyptus leaf litter, Jan. 2014, P.W.
stricted at septum, enclosed in mucoid sheath extending over Crous, HPC 2269 (epitype designated here CBS H-24021
ends of spore, up to 5 μm diam, (13–)18–20(–22) × (3.5–)4 ìm; MBT388163).
ascospores turn brown and verruculose at germination.
Notes: The original paper describing this fungus assumed that it
Typus: Australia, New South Wales, Tabulum, Keybarbin State was a saprobe, as thyrothecia were collected on leaf litter (Crous
Forest, on E. dunnii, 17 Apr. 2018, A.J. Carnegie, HPC 2433 & Kendrick 1994). The recent collection from the same site on
(holotype CBS H-24036, culture ex-type CBS 145916 = CPC symptomatic living leaves suggests that this is a foliar pathogen,
35847). which then also questions its placement in the genus Arnaudiella.
Furthermore, the link with its purported asexual morph, Glio-
Notes: Phaeothyriolum dunnii is described as a new species cladiopsis, has since been shown to be incorrect (Lombard et al.
based on the fact that although it resembles Microthyrium euca- 2015), as the latter genus resides in the Nectriaceae. Species of
lypticola morphologically (ascospores 12–14 × 3 μm; Swart Gliocladiopsis are common on leaf litter, and cultures of
1986c), it is distinct in having larger ascospores. The phyloge- P. eucalyptorum are extremely slow-growing, explaining why the
netic relationship between the Phaeothyriolum species known wrong fungus was eventually cultured, and assumed to be the
from sequence data is shown in Fig. 81. This species is common asexual morph. Phaeothyriolum eucalyptorum is morphologically
on adult foliage of E. dunnii in mature plantations in subtropical similar to P. microthyrioides, in that the upper reddish pigmen-
Australia, mostly found on fallen leaves. On the rare occasion that tation is visible on leaves long before and sporocarps appear.
one has access to live adult foliage (e.g. fallen tree or branch), the
beginnings of leaf spots (i.e. discolouration) can be found. Phaeothyriolum eucalyptorum differs from P. microthyrioides
in the textura epidermoidea cells of its upper thyrothecial wall,
Phaeothyriolum eucalyptorum (Crous & W.B. Kendr.) Crous, and ascospores that are not prominently constricted at the me-
comb. nov. MycoBank MB832044. Fig. 80. dian septum, although the morphology of P. microthyrioides can
Basionym: Arnaudiella eucalyptorum Crous & W.B. Kendr., be resolved only once the type has been located. Because this
Canad. J. Bot. 72: 60. 1994. taxon is associated with leaf spots it cannot be accommodated in
Arnaudiella and is better placed in Phaeothyriolum. It resembles
Forming leaf spots, up to 3 cm diam, diffuse, circular, with
Microthyrium eucalypticola in having thyrothecia that develop on
indistinct border, reddish brown, confined to one side of the leaf,
litter, having a lobed upper wall of textura epidermoidea, and
pigment due to subepidermal hyphae that extend 1 mm away
ascospores with a persistent sheath. It is primarily distinguished
from colonised substomatal cavities. No sporocarps visible, but
from M. eucalypticola by having larger ascospores.
developing within 7–10 d when incubated in damp chambers.
Ascomata thyrothecial, circular, suprastomatal, 90–300 μm Although it was not possible to culture this fungus, amplifi-
diam, lacking superficial hyphae, dark brown, scattered, flat- cation and sequencing of an ITS sequence from fungal material
tened, with star-shaped central ostiole, margin even, lobed; was possible to allow phylogenetic placement of this species
upper wall of textura epidermoidea. Pseudoparaphyses not (Fig. 81).
observed. Asci 8-spored, bitunicate, obpyriform to subcylindrical,
Phaeothyriolum microthyrioides (G. Winter) H.J. Swart,
short pedicellate, straight to slightly curved, with ocular chamber
absent, or weakly developed, 1 μm diam, 30–45 × 13–16 μm.
Basionym: Asterina microthyrioides G. Winter, Hedwigia 24: 23.
Ascospores tri- to multiseriate, medianly 1-septate, fusiform,
1885.
granular and guttulate, widest just above the septum, not to €
Synonyms: Seynesia microthyrioides (G. Winter) Theiss., Osterr.
slightly constricted at septum, (14–)17–18(–20) × (4–)
Bot. Zeitschr. 63: 125. 1913.
5–5.5(–6) μm, with persistent mucoid sheath extending over
Phaeothyriolum eucalyptinum Syd., Annls Mycol. 36: 305. 1938.
ends of the spore (up to 5 μm in lactic acid, 8 μm in water),
Micromicrothelia eucalyptina (Syd.) E. Müll., in Müller & von Arx,
ascospores turn brown and verruculose with age.
Beitr. Kryptfl. Schweiz 11(no. 2): 327. 1962.
Typus: South Africa, Limpopo Province, Goudriver Game
Diagnosis: Leaf spots vary per host. Spots are chlorotic,
Lodge, on Eucalyptus leaf litter, May 1991, P.W. Crous (holotype
and at times with a red margin, ranging from 5 mm
Fig. 81. The single most parsimonious trees obtained from a phylogenetic analysis of the Phaeothyriolum ITS sequence alignment (six strains including the outgroup; 585
characters analysed: 297 constant, 238 variable and parsimony-uninformative and 50 parsimony-informative). The tree was rooted to Melanconiella hyperopta (GenBank
NR_160258.1) and the scale bar indicates the number of changes. Bootstrap support values higher than 49 % are shown at the nodes and novelties are highlighted with bold
text. Species names are indicated to the right of the tree. A superscript T denotes strains with a type status and ET those with ex-epitype status. Tree statistics: TL = 364 steps,
CI = 0.962, RI = 0.745, RC = 0.717. The analysis was performed using PAUP v. 4.0b10 (Swofford 2003).
228
(E. polyanthemos) to 5 cm (E. globulus), at times with radial Note: Phloeosporella eucalypticola is characterised by having
orientation, or forming clusters; thyrothecia have irregular acervuli, short, polyblastic, lageniform conidiogenous cells,
ostioles. The disease is sometimes referred to as “greasy 3–4 × 4–8 μm, and hyaline, filiform, 1-septate conidia with a
spot” because of the appearance of the colonies as dark prominent taper to acutely rounded apices, and truncate bases,
circular areas on a lighter background. Asci are saccate, 35–170 × 2–4 μm.
bitunicate, 40 – 50 × 18 – 21 μm, and contain eight 1-septate,
hyaline ascospores, 19 × 5 μm. Phyllosticta Pers., Traite sur les Champignons Co-
mestibles (Paris): 55. 147. 1818.
Description and illustration: Swart (1986c).
Conidiomata and spermatogonia pycnidial, immersed, subepi-
Notes: The type specimen of Asterina microthyrioides (Australia, dermal to erumpent, unilocular, rarely multilocular, glabrous,
on leaves of E. pilularis var. acmenoides = E. acmenoides)
ostiolate, dark brown to black; ostiole circular to oval; pycnidial wall
could not be located, and the treatment of Phaeothyriolum
of thick-walled, dark brown textura angularis, with inner layers of
microthyrioides by Swart (1986c) is based on his interpretation
hyaline to pale brown, thin-walled textura prismatica to angularis.
of many different specimens. Although he correctly resolved
Conidiophores lining the cavity of the conidioma, reduced to
the presence of interepidermal hyphae, he missed the presence
conidiogenous cells, invested in mucus. Conidiogenous cells
of pseudoparaphyses (not present in all taxa), and the pres-
discrete, producing macroconidia and spermatia (also produced in
ence of a mucoid sheath in ascospores (absent in older
separate spermatogonia), ampulliform, lageniform, doliiform to
ascospores). subcylindrical, hyaline, smooth, proliferating several times per-
Phloeosporella Höhn., Annls Mycol. 22: 201. 1924. currently near the apex, invested in a mucoid layer. Spermatog-
enous cells ampulliform to lageniform or subcylindrical, hyaline
Mycelium immersed, branched, septate, hyaline. Conidiomata
smooth, phialidic. Conidia ellipsoid-fusoid to obovoid or ovoid,
acervular, subepidermal, ± circular, discrete, composed of hya-
rarely subcylindrical, aseptate, broadly rounded at the apex, often
line to pale brown, thin-walled textura angularis. Conidiogenous
tapering strongly toward the base, unicellular, hyaline, smooth-
cells holoblastic, sympodial, discrete, indeterminate, hyaline,
walled, guttulate to granular, often enclosed in a persistent
smooth, lageniform to cylindrical, with one to two broad, flat
mucilaginous sheath, and bearing an unbranched, tapering,
unthickened apical scars, formed from the upper pseudoparen- straight to curved, mucoid apical appendage. Spermatia hyaline,
chyma. Conidia hyaline, 2-euseptate, thin-walled, smooth, gut-
smooth, granular, subcylindrical or dumbbell-shaped, with rounded
tulate, straight, curved or irregular, tapered gradually to an
or blunt ends. Ascomata pseudothecial, separate to gregarious,
obtuse apex and abruptly to a truncate base.
globose to subglobose, brown to black, unilocular with a central
Type species: Phloeosporella ceanothi (Ellis & Everh.) Höhn. ostiole. Asci bitunicate, fissitunicate, clavate to subcylindrical, 8-
spored, fasciculate, stipitate, with an ocular chamber. Pseudo-
Phloeosporella eucalypticola H.Y. Yip, Australas. Pl. Path. 26:
paraphyses mostly absent at maturity, filamentous, branched,
26. 1997. Fig. 82.
septate when present. Ascospores bi- to triseriate, hyaline, gut-
Diagnosis: Leaf spots subcircular to irregular, 3–6 × 8–11 mm, tulate to granular, aseptate, ellipsoid, ellipsoid-fusoid to limoniform,
pale brown, separate, amphigenous. Conidia filiform, hyaline, 1- smooth-walled, usually with mucilaginous caps at ends, or sur-
septate, 74–82 × 2–2.5 μm; apical cells tapering gradually to an rounded by a mucilaginous sheath (from Wikee et al. 2013).
obtuse tip; basal cells ca. 40 % of the length of the conidium, with
Type species: Phyllosticta convallariae Pers., nom. inval. (=
a small truncate base. P. cruenta (Fr.) J. Kickx f.)
Description and illustration: Yip (1997).
Symptoms: Associated with amphigenous, small, brown leaf
Typus: Australia, Victoria, Errinundra, on E. radiata × E. dives, spots.
1971, D.H. Ashton (holotype BRIP 21999). No cultures are
Notes: Phyllosticta eucalyptorum, which was described from
available.
E. grandis leaves in Brazil (Crous et al. 1993c), was shown to
Fig. 82. Phloeosporella eucalypticola (BRIP 21999). A. Disease symptoms. B, C. Vertical section through conidiomata. D. Conidia. Scale bars = 10 μm.
CROUS ET AL.
be a synonym of P. capitalensis by Wikee et al. (2013). The 8 × 1 μm (Cooke 1891), suggesting this is a spermatial morph of
latter species is a common endophyte, although it is associated some ascomycete, probably a member of Teratosphaeriaceae,
with leaf spots on older leaves of different host genera under and not Piggotia s. str. The fungus treated by Thrower is prob-
favourable conditions. Other species known from Eucalyptus ably an asexual species of Diaporthales occurring on leaf litter,
include P. eucalyptina (conidia 18–20 × 5–6 μm; on and those specimens from Brazil represent members of
E. globulus, Tunisia, which could be the same as the uniden- Teratosphaeriaceae.
tified sterile isolate from Spain on E. globulus, CPC 11336),
Pilidium Kunze, in Kunze & Schmidt, Mykologische Hefte
P. eucalypti (on E. globulus in Portugal; = a spermatial morph of
(Leipzig) 2: 92. 1823. Fig. 83.
a Mycosphaerella sp.; Van der Aa & Vanev 2002), and
P. extensa (on Eucalyptus sp. in Caifornia; = a cryptosporiopsis- Conidiomata often of two kinds, most commonly encountered as
like coelomycete; Van der Aa & Vanev 2002). No serious dis- black pycnidia; also producing open, discoid, stalked sporodochia.
ease symptoms have been associated with Phyllosticta spp. on Pycnidia globose to oblong, sessile, dark brown to black at maturity,
Eucalyptus. pale brown when young, superficial, separate, uniloculate, smooth.
Pycnidial wall of two regions: outer region dark brown, inner region
Piggotia Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 7: 95.
hyaline. Opening by a stellate slit, rupturing irregularly, or lacking.
1851.
Conidiophores short, cylindrical. Conidiogenous cells entero-
Conidiomata acervular, subcuticular, applanate or irregularly blastic, phialidic, developing acropleurogenously. Conidia asep-
pulvinate with copious conidial masses, separate or frequently tate, hyaline, fusiform to falcate with ends slightly pointed,
coalescing to form large composite conidiomata, black, cuticle straight to curved, smooth (from Rossman et al. 2004).
dark brown, lower wall of pale to medium brown almost cuboid
Type species: Pilidium acerinum (Alb. & Schwein.) Kunze
textura prismatica. Ostiole absent, dehiscence by irregular
rupture of the cuticle. Conidiophores branched and septate only Notes: This pathogen is rather common on eucalypts, but is
near the base, pale brown, smooth, cylindrical, usually straight, mostly a problem in nurseries, and is seldomly seen in planta-
formed from the upper cells of the acervulus. Conidiogenous tions. It can be especially problematic in cutting nurseries, where
cells, annellidic, indeterminate, discrete, cylindrical, pale brown, it infects cuttings via the the leaf wounds made during the cutting
smooth or verruculose above, with one to two simple or ragged process.
percurrent proliferations. Conidia pale brown, thin-walled, asep-
Pilidium lythri (Desm.) Rossman, IMA Fungus 5: 105. 2014.
tate, smooth, cylindrical to almost cuneiform, apex obtuse, base
Basionym: Dacrymyces lythri Desm., Pl. Crypt. Nord France,
truncate (from Sutton 1980).
Edn 1: no. 1545. 1846.
Type species: Piggotia ulmi (Grey.) Keissl. (= Asteroma ulmi Synonyms: Hainesia lythri (Desm.) Höhn., Sber. Akad. Wiss.
Grey., P. astroidea (Berk.) Berk. & Broome) Wien, Math.-naturw. Kl., Abt. 1 115: 687 [39 of repr.] 1906.
Pezizella lythri (Desm.) Shear & B.O. Dodge, Mycologia 13(3):
Piggotia substellata Cooke, Grevillea 20(no. 93): 6. 1891.
149. 1921.
Diagnosis: Referred to as winter leaf spot disease, as the dis-
ease occurs mainly on E. regnans during the winter in Victoria. Diagnosis: Primarily a nursery pathogen, causing water soaked
Especially severe in wet winters. Visible as dull water-soaked brown leaf spots, and stem lesions, leading to seedling death
patches, often on leaf tips of the lower leaves. Pycnidial con- under conditions of high humidity. Conidiomata pycnidial and
idiomata formed on the leaf surface after the rupture of the sporodochial (discoid). Conidia aseptate, hyaline, fusoid to
cuticle; conidia hyaline, 1-celled, elliptical, 8–13 × 2–2.5 μm). falcate with ends slightly pointed, straight to curved, smooth,
5.5–8(–11) × 1.5–2(–2.5) μm.
Descriptions and illustrations: Macauley & Thrower (1966),
Ashton & Macauley (1972). Description and illustration: Rossman et al. (2004).
IMI 101643: Conidiomata pycnidial, globose, immersed, black Materials examined: Brazil, Minas Gerais, Viçosa, on E. globulus, May 1993,
with slightly papillate neck, 150–250 μm diam, mostly hypo- P.W. Crous, culture CPC 15142; Rio Grande do Sul, Guaiba, on E. globulus, May
1993, A.C. Alfenas, CPC 15154.
phyllous; wall of 6–10 layers of brown textura angularis. Co-
nidiophores reduced to conidiogenous cells lining inner cavity. Plectosphaera Theiss., Ann. Mycol. 14: 413. 1917 (1916).
Conidiogenous cells hyaline, smooth, ampulliform,
Von Arx & Müller (1954) regarded Plectosphaera as accom-
5–10 × 3–5 μm, with long thin neck with visible percurrent
modating non- or weakly stromatic Phyllachora species. Cannon
proliferation. Conidia aseptate, hyaline, smooth, thin-walled,
(1996) did not concur with this view, as the type species,
subcylindrical, straight, apex obtuse, inner plane flat, outer
P. bersamae, is a typical stromatic member of Phyllachora. The
plane convex, hilum truncate, situated on inner plane, 1 μm diam,
genus Plectosphaera thus requires revision, and its phylogenetic
(12–)13–14(–15) × (2.5–)3 μm.
affinity to other fungi remains unclear.
Materials examined: Australia, Victoria, Univ. of Melbourne, on E. regnans, 5
May 1964, IMI 101643 (Diaporthales?). Brazil, Sao Paulo, on E. paniculata, 16
Type species: Plectosphaera bersamae (Lingelsh.) Theiss.
Jan. 1973, I.A.S. Gibson, IMI 173095 (spermatial morph of Pallidocercospora Notes: Pascoe (1990) reported on the pseudoclypeus and
sp.); on E. saligna, 9 Jun. 1973, C. Hodges, IMI 176992 (spermatial morph of
Teratosphaeria sp.).
apparent bitunicate nature of its asci and speculated regarding its
correct classification. Summerell et al. (2006) showed that this
Notes: This fungus is common on leaf litter and in the soil fungus resides in the Amphisphaeriaceae.
associated with E. regnans and is splash-dispersed (Macauley &
Thrower 1966, Ashton & Macauley 1972). Spores were originally Plectosphaera eucalypti (Cooke & Massee) H.J. Swart, Trans.
described as cylindrical, straight with obtuse ends, hyaline, Brit. Mycol. Soc. 76: 91. 1981. Fig. 84.
230
Fig. 83. Pilidium spp. A. Sporodochial conidiomata forming on SNA. B. Leaf spot with immersed conidiomata. C. Vertical section through conidioma. D, E. Conidiogenous cells
giving rise to conidia. F. Conidia. Scale bars: B, C = 150 μm, all others = 10 μm.
Basionym: Trabutia eucalypti Cooke & Massee, Grevillea 17(no. Type species: Podosphaera myrtillina Kunze
82): 43. 1888.
Symptoms: White mycelium growth on leaves and stems.
Synonyms: Phyllachora eucalypti (Cooke & Massee) Theiss. &
Syd., Ann. Mycol. 13(5/6): 539. 1915. Notes: This pathogen has a wide host range other than on
Placostroma eucalypti (Cooke & Massee) Hansf., Proc. Linn. Eucalyptus spp. It is widely distributed on Eucalyptus, occurring
Soc. N.S.W. 81(1): 29. 1956. on all continents where eucalypts are planted. It is not consid-
ered a significant pathogen.
Diagnosis: Colonies developing on either leaf surface, but
prominent on only one leaf surface, ca. 2.5 – 50 mm diam, Podosphaera aphanis (Wallr.) U. Braun & S. Takam.,
roughly circular to ellipsoidal, reddish to reddish brown, often Schlechtendalia 4: 26. 2000. Fig. 85.
developing a convex rise in the leaf tissue, with numerous, Basionym: Alphitomorpha aphanis Wallr., Ann. Wetter.
small, 0.5 – 2 mm diam, black, circular domes, each uni- Gesellsch. Ges. Naturk. 4: 242. 1819.
loculate, ostiole distinct as a minute apical pore in each dome.
On the opposite leaf surface, visible sometimes as a region of Diagnosis: Conidiophores erect, up to 300 μm long, foot-cells
brownish, roughly circular, discoloured leaf tissue, or as a cylindrical, increasing somewhat in width towards the apex,
roundish, puckered, region in the leaf tissue. Ascospores ar- 30–160 × 8–13.5 μm. Conidia with fibrosin bodies, ellipsoid-
ranged uni-biseriately, often overlapping, 36 – 41 × ovoid, doliiform-limoniform, (20–)25–40(–45) × (10–)
11.5 – 14 μm, inclusive of thick, hyaline, epispore layer, fusiform 14–26 μm.
to narrowly ellipsoidal, sometimes ovoid, often inaequilateral, Description, illustration and specimens: Braun & Cook (2012).
hyaline, with a hyaline epispore layer 2.5 – 5 μm thick, often
thickest at poles. Pseudocercospora Speg., Anales Mus. Nac. Hist. Nat. Buenos
Aires, Ser. 3, 20: 437. 1910. Fig. 86.
Descriptions and illustrations: Swart (1981), Pascoe (1990).
Foliicolous, chiefly phytopathogenic, but also endophytic;
Material examined: Australia, Northern Territory, Alice Springs, Olive Pink Bot-
anic Gardens, on E. orbifolia, Oct. 2005, B. Summerell, CBS H-19751, cultures commonly associated with leaf spots, but also occurring on fruit.
CBS 120063 = CPC 12955, CPC 12956. Ascomata pseudothecial, single to aggregated, black, immersed
becoming erumpent, globose; apical papillate ostiole; wall of 3–4
Podosphaera Kunze, in Kunze & Schmidt, Mykologische Hefte
layers of medium brown textura angularis, base consisting of
(Leipzig) 2: 111. 1823.
2–3 layers of hyaline cells. Asci aparaphysate, fasciculate,
Asexual morph: Fibroidium - mycelium external; hyphal appres- bitunicate, subsessile, ovoid to obclavata to ellipsoid, straight or
soria indistinct to nipple-shaped; conidia in chains with crenate incurved, 8-spored. Ascospores 3- to multiseriate, overlapping,
outline, with fibrosin bodies. Sexual morph: chasmothecia with hyaline, guttulate, thin-walled, slightly curved, rarely straight,
dichotomously branched appendages (sect. Podosphaera) or fusoid-ellipsoidal with obtuse ends, 1(–3)-septate, tapering to-
mycelium-like unbranched or irregularly branched appendages ward both ends. Mycelium internal and external, consisting of
(sect. Sphaerotheca) and a single, (6–)8-spored ascus per smooth, septate, subhyaline to brown, branched hyphae. Stroma
ascoma (for detailed description see Braun & Cook 2012). absent to well-developed. Conidiophores in vivo arranged in
CROUS ET AL.
Fig. 84. Plectosphaera eucalypti (CPC 12955). A. Disease symptoms. B. Leaf spot with black immersed ascomata. C, D. Vertical section through ascomata. E–H. Asci with
ascospores. Scale bars: C, D =400 μm, all others = 10 μm.
loose to dense fascicles, sometimes forming distinct synnemata minute marginal frill, straight to curved, rarely sigmoid, smooth to
or sporodochia, emerging through stomata or erumpent through finely verruculose; hila usually unthickened, not darkened, at
the cuticle, often arising from substomatal or subcuticular to most somewhat refractive, occasionally slightly thickened along
intraepidermal stromata, or occurring singly on superficial hy- the rim, or rarely flat, and slightly thickened and darkened, but
phae, short to long, septate or continuous, i.e. conidiophores never pronounced (Crous et al. 2013).
may be reduced to conidiogenous cells, simple to branched and
Type species: Pseudocercospora vitis (Lev.) Speg.
straight to geniculate-sinuous, pale to dark brown, smooth to
finely verruculose. Conidiogenous cells integrated, terminal, Symptoms: Leaf spots amphigenous, angular to subcircular or
occasionally intercalary, polyblastic, sympodial, or monoblastic, irregular, pale to medium brown, frequently surrounded by a
proliferating percurrently via inconspicuous or darkened, irregular raised, dark brown border with a diffuse red-purple margin;
annellations, at times denticulate, pale to dark brown; scars single, discrete lesions may coalesce with age; small olivaceous
inconspicuous, or only thickened along the rim, or flat, and brown tufts of conidiophores are usually visible under the hand
slightly thickened and darkened, but never pronounced. Conidia lens. In some species, superficial mycelium may be more
solitary, rarely in simple chains, subhyaline, olivaceous, pale to common, in which case only fluffy olivaceous brown growth
dark brown, usually scolecosporous, i.e. obclavate-cylindrical, would be seen.
filiform, acicular, and transversely plurieuseptate, occasionally
Notes: Although some species of Pseudocercospora can cause
also with oblique to longitudinal septa, conidia rarely amero- to
prominent leaf spot symptoms (e.g. P. eucalyptorum), these are
phragmosporous, short subcylindrical or ellipsoid-ovoid, aseptate
mostly found on older foliage, and thus appear to be of little
or only with few septa, apex subacute to obtuse, base obconi-
importance on Eucalyptus. They are relatively common on
cally truncate to truncate, or bluntly rounded, with or without a
mature foliage, but not damaging, on a wide range of hosts in
232
Fig. 85. Podosphaera aphanis (BRIP 50406). A. Disease symptoms. B. Leaf spot with sporulation. C, D. Conidial chains. Scale bars = 10 μm.
Fig. 86. Pseudocercospora spp. A–D. Disease symptoms of P. eucalyptorum. E–I. P. chiangmaiensis (CPC 15412). E. Leaf spot. F, G. Conidiogenous cells. H, I.
Conidia. J–N. P. tereticornis (CPC 13299). J. Leaf spots. K, L. Conidiophores with conidiogenous cells with percurrent proliferation. M, N. Conidia with guttules. Scale
bars = 10 μm.
CROUS ET AL.
native stands and in eastern Australia. Disease symptoms also Diagnosis: Leaf spots amphigenous, subcircular to angular,
vary depending on the host species infected. 2–6 mm diam, pale to medium brown, surrounded by a slightly
raised, dark-brown border, becoming confluent with age, leading
Pseudocercospora acerosa U. Braun & M.A. Dick, N.Z. Jl For.
to leaf blight from the leaf tip. Conidia solitary, subcylindrical to
Sci. 32: 223. 2002.
narrowly obclavate, tapering toward the subobtuse apex; base
Diagnosis: Leaf spots amphigenous, subcircular to angular- obconic-subtruncate, (2–)3–5(–10)-septate, straight to slightly
irregular, 1–10 mm diam or confluent, pale to medium dark curved, pale to medium brown, smooth, thin-walled, guttulate,
brown, dingy greyish brown, margin indefinite or with a narrow (40–)50–60(–100) × (2–)2.5–3(–3.5) μm (up to 140 μm long in
darker border or marginal line, reddish brown to dark brown, moist chambers).
margin or sometimes entire spots somewhat raised. Conidia
solitary, 40–180 × (2.5–)3–5.5(–6) μm, short conidia sub-
cylindrical, rarely obclavate, longer conidia broadly acicular (i.e., Typus: Thailand, Chiang Mai, Doi Lor, on E. camaldulensis, Jun.
gradually attenuated from a truncate base towards the apex), 4- 2007, P. Suwannawong (holotype CBS H-20133, culture ex-type
to 15-septate, subhyaline, pale olivaceous to olivaceous-brown, CBS 123244 = CPC 15412); ibid., cultures CPC 15450, 15451.
smooth, wall thin to slightly thickened, apex obtuse in short
Pseudocercospora crispans G.C. Hunter & Crous, Stud.
conidia and obtuse to subacute in longer conidia, base truncate,
Mycol. 75: 82. 2012 (2013).
rarely slightly narrowed at the very base (short obconically
truncate), (2.5–)3–4(–5) μm wide, hila unthickened, non- Diagnosis: Leaf spots amphigenous, angular to irregular, pre-
pigmented. dominantly occurring next to or close to the mid-rib, 2–15 mm
diam, pale brown on the upper side of the leaf, and pale to darker
Description and illustration: Braun & Dick (2002).
brown on the bottom side of the lesion, surrounded by a raised,
Typus: New Zealand, Christchurch, Hagley Park (MC), on dark brown border with a diffuse red pigment emanating away from
E. baxteri, 26 Sep. 2001, P.M. Bradbury (holotype NZFRI-M the border; single, discrete lesions may coalesce to form larger
4580). Not known from culture. lesions. Conidia solitary, smooth, guttulate, curved to undulate, pale
brown, 3–9-septate, apex acute to subacute, base truncate, (40–)
Pseudocercospora basiramifera Crous, Mycol. Mem. 21: 120.
65–96(–102) × (2–)3(–4) μm; hila unthickened, not darkened.
1998.
Diagnosis: Leaf spots amphigenous, subcircular or angular,
2–10 mm diam, dark brown, border diffuse, margin red when Typus: South Africa, Western Cape Province, Knysna, on
present. Conidia solitary, pale olivaceous, smooth, thin-walled, Eucalyptus sp., Jan. 2008, P.W. Crous (holotype CBS H-20392,
guttulate, narrowly obclavate, apex obtuse, base long obconic- culture ex-type CBS 125999 = CPC 14883).
truncate, 3–10-septate, (35–)50–70(–80) × 2(–3) μm; hila
Pseudocercospora crousii U. Braun & M.A. Dick, N.Z. Jl For.
thickened along the rim as associated in species of
Sci. 32: 224. 2002.
Paracercospora.
Diagnosis: Leaf spots amphigenous, subcircular to angular-
irregular, sometimes oblong, 1–15 mm diam, sometimes
Typus: Thailand, on E. pellita, May 1995, M.J. Wingfield (ho- confluent and larger, brown, reddish brown, later greyish brown
lotype PREM 54406, cultures ex-type CBS 111072 = CPC 1266, to greyish white, with a darker narrow margin or marginal line,
CBS 114757 = CPC 1267). often somewhat raised, brown to purplish violet, finally often
blackish. Conidia solitary, obclavate or obclavatecylindrical, short
Pseudocercospora basitruncata Crous, Mycol. Mem. 21: 123.
conidia sometimes cylindrical, (20–)35–115(–140) × 3–5 μm, 4-
1998.
to 10-septate, subhyaline to pale olivaceous or olivaceous-
Diagnosis: Leaf spots amphigenous, variable, 1–20 mm diam, brown, smooth or almost so, wall thin to slightly thickened,
subcircular, irregular or angular, pale brown, becoming dark apex obtuse to subacute, base obconically truncate, (1–)
brown near the border; margin thin, red-brown; lesions light 1.5–2(–3) μm diam, hila unthickened, non-pigmented.
brown on lower surface without margins. Conidia solitary,
Description and illustration: Braun & Dick (2002).
olivaceous to pale brown, finely verruculose, thick-walled,
guttulate, straight or curved, subcylindrical, lower part cylindri- Typus: New Zealand, Whakarewarewa Forest (BP), on
cal, tapering above to an obtuse apex, base mostly truncate, E. regnans, 18 Sep. 2001, M. Dick (holotype NZFRI-M 4550).
rarely obconic-truncate, straight to geniculate-sinuous, mostly
Pseudocercospora cubae Crous, Mycol. Mem. 21: 125. 1998.
only slightly curved, 1–6(–11)-septate, (25–)45–70(–90) ×
2.5–3(–3.5) μm. Diagnosis: Leaf spots amphigenous, irregular, 2–5 mm diam,
pale brown, becoming medium brown toward a raised border,
margin thin to absent, red-purple. Conidia solitary, olivaceous,
Typus: Colombia, Astorga, 5 km North of Nemocon, Cundina- smooth to finely verruculose, thick-walled, guttulate, straight or
marca, alt. 2 615 m, on Eucalyptus sp., 5 Jan. 1942, J. Orjuela- curved, subcylindrical, tapering from a truncate to long obconic-
Navarrete (holotype BPI 436146, isotype Herbario de Fitopa- truncate base to an obtuse apex, 0–3-septate, (20–)
tologia Dept. de Agric. Bogota 00990). Not known from culture. 30–40(–50) × (2–)2.5–3 μm; hila 1.5–2 μm wide, inconspic-
uous, unthickened, not darkened.
Pseudocercospora chiangmaiensis Cheew. et al., Persoonia
21: 87. 2008. Fig. 86. Description and illustration: Crous (1998).
234
Typus: Cuba, on Eucalyptus sp., 26 Mar. 1966, R.V. Martinez Pseudocercospora eucalyptorum Crous et al., Mycol. Res. 93:
(holotype IMI 121688). Not known from culture. 394. 1989. Fig. 86.
Synonym: Pseudocercospora pseudoeucalyptorum Crous, Stud.
Pseudocercospora deglupta Crous, Mycol. Mem. 21: 127.
Mycol. 50: 210. 2004.
1998.
Diagnosis: Leaf spots amphigenous, subcircular to angular,
3–10 mm diam, pale to medium brown, surrounded by a raised,
diam, dark brown on the upper surface with a diffuse, irregular
brown border. Conidia solitary, pale brown, smooth, cylindrical,
border; dark to medium brown on lower surface; coloured margin
bases truncate, apices bluntly rounded, thick-walled with irreg-
absent. Conidia solitary, pale to medium brown, verruculose,
ular swellings, straight or curved, 1–7-septate, (25–)
thick-walled, guttulate, narrowly obclavate, apex obtuse, base
59–70(–90) × 2.5–3(–4) μm in vivo, 30–65 × 2.5–3 μm, 3–6-
obconic-subtruncate, 0–8-septate, (35–)45–70(–80) × (3–)
septate in vitro; hila inconspicuous.
3.5–4(–4.5) μm; hila 2 μm wide, inconspicuous, unthickened,
not darkened. Description and illustration: Crous et al. (1989b).
Description and illustration: Crous (1998). Typus: South Africa, Western Cape Province, Stellenbosch,
Stellenbosch Mountain, on E. nitens, Aug. 1988, P.W. Crous
Typus: Malaysia, on E. deglupta, 19 Jan. 1981, I.A.S. Gibson
(holotype PREM 49112, cultures ex-type CBS
(holotype IMI 255250). Not known from culture.
110777 = CMW5228 = CPC 16, CPC 11–15, 17).
Pseudocercospora denticulata Crous, Mycol. Mem. 21: 129.
Note: This species is common on adult foliage of a wide range of
1998.
Eucalyptus spp.
Diagnosis: Leaf spots amphigenous, varying from irregular
Pseudocercospora flavomarginata G.C. Hunter et al., Fungal
specks to larger, irregular, angular or subcircular lesions,
Diversity 22: 80. 2006.
confined by leaf veins, 1–8 mm diam, medium brown, with a
slightly raised border, margin mostly absent, but red-purple Diagnosis: Leaf spots distinct, scattered over leaves, amphige-
when present; larger, subcircular lesions are light brown in nous, circular to angular, 320 mm diam, pale to dark brown
the centre. Conidia solitary, olivaceous, finely verruculose, gut- becoming darker with age, definite chlorotic margin on abaxial and
tulate, obclavate to subcylindrical, straight or curved, apex adaxial leaf surfaces but more obvious on adaxial leaf surfaces.
obtuse, base obconic-truncate, 1–5-septate, (25–)40–50(–70) × Conidia solitary, straight or slightly curved, pale brown, smooth,
2–2.5(–3) μm; hila 1-1.5 μm wide, inconspicuous, unthickened, thick-walled, guttulate, acicular to obclavate, apex obtuse, base
not darkened. rounded to long obconic-truncate, 2–7-septate, (28–)
46–54(–90) × (2–)3(–4) μm; hila unthickened, not darkened.
Description and illustration: Hunter et al. (2006).
Typus: Dominican Republic, on Eucalyptus sp., 25 Apr. 1970,
R. Heliczer (holotype BPI 436203). Not known from culture. Typus: Thailand, Chang Gao Province near Pratchinburi, on
E. camaldulensis, 2004, M.J. Wingfield (holotype PREM 58952,
Pseudocercospora epispermogonia Crous & M.J. Wingf.,
cultures ex-type CBS 118841 = CMW 13586, CBS
Mycologia 88: 456. 1996.
118823 = CMW 13590, CBS 118824 = CMW 13594).
Diagnosis: Conidia solitary, narrowly obclavate with a subobtuse
apex and long obconic-subtruncate base, straight or curved, Pseudocercospora fori (G.C. Hunter et al.) G.C. Hunter et al.,
28–65 × 2–3 μm, pale olivaceous, smooth, guttulate, 1–7- Persoonia 23: 108. 2009.
septate; hilum inconspicuous, 1–2 μm wide. Basionym: Mycosphaerella fori G.C. Hunter et al., Mycol. Res.
108: 677. 2004.
Description and illustration: Crous & Wingfield (1996). Diagnosis: Leaf spots amphigenous, subcircular to irregular,
Typus: South Africa, Northern Province, Tzaneen, Magoe- 2–30 mm diam, becoming confluent along leaf margins, grey,
baskloof, on E. grandis × E. saligna hybrid, Oct. 1994, G. Kemp surrounded by a dark brown outer zone, and a thin red-purple
(holotype PREM 51936). Not known from culture. border, confluent with the leaf surface; abaxial surface medium
brown with a dark brown outer zone. Ascospores 3- to multi-
Notes: Conidia of Ps. epispermogonia failed to grow in culture. seriate, overlapping, hyaline, guttulate, thick-walled, straight to
The cultures associated with the type specimen, were generated slightly curved, narrowly ellipsoid with subobtuse apices, medi-
from single ascospores of Paramycosphaerella marksii. anly 1-septate, widest at unconstricted septum, tapering equally
Pseudocercospora eucalypticola S.K. Singh & Bhalla, as toward both ends, (12–)14–16(–20) × 3(–3.5) μm in vivo,
“kolanesis”, Indian Phytopath. 53: 401. 2000. 11–14 × 2.5–3.5 μm in vitro.
Diagnosis: Leaf spots amphigenous, angular, vein limited, Description and illustration: Hunter et al. (2004).
sometimes coalescing to form large blotches. Conidia straight to Typus: South Africa, Northern Province, Tzaneen, on
curved, smooth-walled, 2–4-septate, cylindrical, apex sub- E. grandis, 2000, G.C. Hunter (holotype PREM 57305, culture
obtuse, base obclavate, subhyaline, 30.5–90 × 2–3.5 μm. ex-type CBS 113285 = CMW 9095).
Description and illustration: Singh & Bhalla (2000). Pseudocercospora gracilis Crous & Alfenas, Mycologia 87:
Typus: India, Uttar Pradesh, Mirzapur, on Eucalyptus sp., Feb. 123. 1995.
1997, S.K. Singh (holotype HCIO 42554, isotype Herb. No. 05/ Synonym: Mycosphaerella gracilis Crous & Alfenas, Mycologia
97). Not known from culture. 87: 123. 1995.
CROUS ET AL.
Diagnosis: Leaf spots amphigenous, irregular, 2 – 6 mm Pseudocercospora natalensis Crous & T.A. Cout., Mycol.
diam, pale brown, surrounded by a raised, thin red to Mem. 21: 136. 1998.
brown border. Ascospores 3- to multiseriate, overlapping,
colourless, guttulate, thin-walled, slightly curved, rarely Diagnosis: Leaf spots amphigenous, angular to subcircular,
straight, narrowly ellipsoid with obtuse apices, 1-septate, 1–9 mm diam, pale to dark brown, confined by leaf veins,
widest at unconstricted septum, tapering toward both ends, frequently becoming darker toward the raised border, margin
(10 – )15 – 18( – 20) × (2 – )2.5 – 3 μm. Conidia solitary, cylin- absent or thin and red when present. Conidia solitary, olivaceous
drical, straight or curved, apex obtuse, base truncate, to pale brown, smooth to verruculose, thick-walled, guttulate,
35 – 100 × 2 – 3 μm, indistinctly 3 – 11-septate, guttulate, straight or curved, subcylindrical with a truncate base, tapering to
pale olivaceous, hilum unthickened. an obtuse apex, 4–8(–11)-septate, (30–)50–85(–110) × (2–)
2.5–3(–3.5) μm in vivo; hila 2–3.5 μm wide, inconspicuous,
Description and illustration: Crous & Alfenas (1995).
unthickened, not darkened; conidia (65–)85–100(–120) × (2–)
Typus: Indonesia, North of Sumatra, on E. urophylla, 22 Nov. 2.5–3 μm in vitro.
1993, A.C. Alfenas (holotype PREM 51718 of sexual morph;
PREM 51719 of asexual morph, cultures ex-type CBS
242.94 = CPC 729, CBS 243.94 = CMW 14455 = CPC 730). Typus: South Africa, Kwazulu-Natal Province, Kwambonambi,
on E. nitens, 22 Nov. 1995, T. Coutinho (holotype PREM 54412,
Pseudocercospora haiweiensis Crous & X. Zhou, Stud. Mycol.
cultures ex-type CBS 111069 = CMW 14777 = CPC 1263, CBS
75: 87. 2012 (2013).
111070 = CMW 14784 = CPC 1264, CBS 111071 = CPC 1265).
Diagnosis: Leaf spots amphigenous, irregular to subcircular or
Pseudocercospora norchiensis Crous, Fungal Diversity 26:
angular, 2–4 mm diam, brown, with raised border, and at times
172. 2007.
with a red-purple margin. Conidia solitary, brown, finely verru-
culose, guttulate, subcylindrical, apex obtuse, base obconically Diagnosis: Leaf spots amphigenous, irregular to subcircular,
subtruncate to truncate, straight to gently curved, 3(–5)-septate, 2–6 mm diam, medium brown with a raised border, and a thin
(25–)30–40(–45) × 3(–4) μm; hila unthickened, neither dark- red-purple margin. Conidia solitary, medium brown, smooth,
ened nor refractive, 1.5 μm wide. prominently guttulate, obclavate, apex subobtuse, base short
obconically truncate, straight to slightly curved, (0–)5–9(–12)-
septate, (50–)80–120(–140) × (5–)6(–7) μm; hila inconspic-
Typus: China, Hai Wei, on Eucalyptus sp. (APP 21), 3 Jun. 2007, uous, 2–3 μm wide.
X. Zhou (holotype CBS H-20866, culture ex-type CBS
131584 = CPC 14084).
Typus: Italy, Viterbo, Norchia, on Eucalyptus sp., Apr. 2005, W.
Pseudocercospora irregularis Crous, Mycol. Mem. 21: 134.
Gams (holotype CBS H-19841, culture ex-type CBS
1998.
120738 = CPC 13049).
Diagnosis: Leaf spots amphigenous, angular to irregular,
Pseudocercospora paraguayensis (Kobayashi) Crous, Myco-
1–7 mm diam, confined by leaf veins, pale brown, becoming
taxon 57: 270. 1996.
dark brown near a raised border; margin red when present.
Basionym: Cercospora paraguayensis Kobayashi, Trans. Mycol.
Conidia solitary, medium brown, becoming olivaceous near apex,
Soc. Japan 25: 263. 1984.
thick-walled, verruculose, guttulate, straight or slightly curved,
Synonym: Pseudocercospora eucalypti Goh & Hsieh, Cerco-
obclavate, apex rounded to obtuse, base obconic-truncate,
spora and similar fungi from Taiwan: 244. 1990.
conidium body with irregular swellings, 1–7-septate, (17–)
45–65(–90) × 3.5–4(–5) μm; hila 1.5–2 μm wide, inconspic- Diagnosis: Leaf spots amphigenous, angular, confined by leaf
uous, unthickened, not darkened. veins, pale to medium brown, 1–4 mm diam, coalescing with
age, border slightly raised, margin absent or thin, red when
present. Conidia solitary, smooth to finely verruculose, oliva-
Typus: Peru, Lima, on Eucalyptus sp., 2 Aug. 1928, E.V. Abbott ceous to light brown, straight to sinuous, thick-walled, guttulate,
(holotype BPI 436140). Not known from culture. narrowly obclavate, apex obtuse, base obconic-truncate, (1–)
3–5(–7)-septate, (28–)40–65(–75) × 2.5–3(–3.5) μm; hila
Pseudocercospora madagascariensis Crous & M.J. Wingf.,
1.5–2 μm wide, inconspicuous, unthickened, not darkened.
Persoonia 22: 88. 2009.
Diagnosis: Leaf spots amphigenous, subcircular to circular,
1–2 mm diam, medium brown with sporulation within and Typus: Paraguay, Pirapo, Itapua, on Eucalyptus sp., 4 Dec.
adjacent to lesion. Conidia solitary, pale brown, smooth, sub- 1983, T. Kobayashi (holotype TFM: FPH 5626).
cylindrical to narrowly obclavate, apex subobtuse, base long
Pseudocercospora robusta Crous & M.J. Wingf., Mycol. Mem.
obconically subtruncate to truncate, straight to curved, 1–3(–4)-
21: 141. 1998.
septate, (15–)30–45(–60) × 2(–2.5) μm; hila and scars
inconspicuous. Diagnosis: Leaf spots amphigenous, but more prominent on
upper surface, subcircular, 4–8 mm diam, pale brown with an
indistinct border. Conidia solitary, pale to medium brown, ver-
Typus: Madagascar, Morondavo, on E. camaldulensis, Aug. ruculose, thick-walled, guttulate, straight to sinuous, sub-
2007, M.J. Wingfield (holotype CBS H-20192, cultures ex-type cylindrical, apex bluntly rounded, base long obconic-truncate to
CBS 124155 = CPC 14621, CPC 14622). subtruncate, 3–9-septate, (45–)70–80(–110) × (2.5–)3–3.5 μm
236
in vivo; hila 2–3 μm wide, inconspicuous, unthickened, not Diagnosis: Leaf spots amphigenous, irregular to subcircular,
darkened; conidia (55–)70–80(–100) × (2–)2.5–3.5 μm in vitro. 1–5 mm diam, medium brown with a raised border and thin, red-
purple margin. Conidia solitary, medium brown, smooth, guttu-
late, subcylindrical, apex obtuse, at times conidia somewhat
Typus: Malaysia, Cameron Highlands, on E. robusta, May 1995, elongated, then apex subobtuse, widest at truncate base, straight
M.J. Wingfield (holotype PREM 54413, cultures ex-type CBS to curved, 1–6-septate, (30–)40–65(–85) × (4–)5(–6) μm; hila
111175 = CMW 5151 = CPC 1269, CPC 1270). inconspicuous, with minute marginal frill.
Pseudocercospora sphaerulinae Crous & M.J. Wingf., Sydo- Description and illustration: Crous et al. (2009b).
wia 55: 145. 2003.
Typus: Australia, Queensland, Cairns, 48 km from Mareeba,
Synonym: Mycosphaerella sphaerulinae Crous & M.J. Wingf.,
Eureka Creek, 17°17'13.2"S, 145°02'27.4"E, alt. 468 m, on
Sydowia 55: 145. 2003.
leaves of E. tereticornis, 27 Sep. 2006, leg. P.W. Crous & B.
Diagnosis: Leaf spots amphigenous, subcircular, coalescing to Summerell (holotype CBS H-20256, cultures ex-type CBS
form larger blotches, mostly along leaf margins, up to 30 mm 125214 = CPC 13299, CPC 13300, 13301, 13315–13317).
diam, more prominent on adaxial than abaxial leaf surface, pale
Note: This species is common on mature foliage of E. tereticornis
brown, surrounded by a dark brown, diffuse border on adaxial
in remnant stands in eastern Australia.
surface; spots pale brown with indistinct border on abaxial
surface. Ascospores tri- to multiseriate, overlapping, hyaline, Pseudocercospora tumulosa (Carnegie & Beilharz) Carnegie
guttulate, thin-walled, slightly curved, fusoid-ellipsoidal with & Crous, comb. nov. MycoBank MB832045.
obtuse ends, widest in middle of spore, 1-septate when Basionym: Mycosphaerella tumulosa Carnegie & Beilharz,
immature, 3-septate at maturity, not constricted at septa, Mycologia 99: 465. 2007.
tapering equally towards both ends, (20 –)25–30(–35) × (3–)
Diagnosis: Leaf spots amphigenous, subcircular to irregular,
3.5–4 μm. Conidia solitary, pale brown, smooth, guttulate,
2 –15 mm diam, single to confluent, beginning as irregular
subcylindrical, apex subobtuse, base truncate, straight to
chlorotic blotches, turning carmine-red before necrosis, slightly
slightly curved, 5–10-septate, 70–100 × 2–3 μm; hila
raised and corky, yellow-brown to red-brown on the abaxial
inconspicuous.
surface, medium brown to grey-brown on the adaxial surface,
Description and illustration: Crous et al. (2003). becoming grey-brown to grey with age, especially in center,
maturation of leaf spot often not even, with irregular margins
Typus: Chile, on E. nitens, May 2001, M.J. Wingfield (holotype
and red-brown border 1 mm wide, often with diffuse chlorotic
and red-brown to purple halo surrounding leaf spot. Asco-
Pseudocercospora subulata Z.Q. Yuan et al., Nova Hedwigia spores 2– 3-seriate, overlapping, narrowly clavate to fusoid-
71: 416. 2000. ellipsoidal, straight to slightly curved, hyaline, basal end
Synonym: Pseudocercospora pseudobasitruncata U. Braun & M. obtuse, apical end prominently tapered, apical cell occasionally
Dick, N.Z. Jl For. Sci. 32: 228. 2002. asymmetrical, widest at midpoint of apical cell, medianly 1-
septate, not constricted at septum or slightly so,
Diagnosis: Leaf spots amphigenous, subcircular to angular-
15 –19 × 4.5–5.5 μm. Conidia solitary, pale olivaceous to
irregular, 1–10 mm diam or confluent and larger, pale to dark
subhyaline, smooth to faintly verruculose, straight or slightly
brown or blackish brown, later pale greyish brown to dingy grey,
curved, narrowly obclavate or subulate to cylindrical, often
margin narrow, somewhat raised, reddish-purplish or pale to dark
variable in diameter, gradually attenuating to a subobtuse or
brown or with a raised marginal line and diffuse reddish-purplish
subacute apex and more abruptly to an obconically or narrowly
halo. Conidia solitary, most conidia broadly acicular (i.e., grad-
obclavate, subtruncate or truncate base, anastomosing, 3 –7-
ually attenuated from a truncate base towards the apex), some
septate, (31–)50–80( –95) × 3.5–4.5 μm.
shorter conidia obclavate-cylindrical, 40–100 × 4–7 μm, 3- to
10-septate, pale olivaceous-brown, apex obtuse, base truncate, Description and illustration: Carnegie et al. (2007).
occasionally short obconically truncate, 4–5 μm wide, hila
Typus: Australia, Queensland, Kingaroy, Coolabunia Planta-
unthickened, non-pigmented.
tion, on E. moluccana, 14 Feb. 2004, A.J. Carnegie (holotype
Descriptions and illustrations: Yuan et al. (2000), Braun & Dick DAR 77424, culture ex-type CBS 121158 = AC0213i = DAR
(2002). 77424; isotype NSWF 005313, culture ex-isotype NSWF
005313).
Typus: Australia, Tasmania, near Birralee, Pughs South, on
E. nitens, 11 Feb. 1999, D. de Little, NFP 11 (holotype of Note: Based on the blast results of the ITS sequence
Pseudocercospora subulata VPRI 22042). New Zealand, Kain- of the ex-type culture (GenBank NR_137549.1), a new
garoa Forest (TO), on E. nitens, 11 Jul. 1984, D.J. Slater-Hayes combination for this species in Pseudocercospora is
(holotype of Pseudocercospora pseudobasitruncata NZFRI-M warranted.
2068).
Pseudophaeophleospora C. Nakash. et al., Stud. Mycol. 87:
Material examined: New Zealand, North Island, KeriKeri, on leaves of 311. 2017.
E. botryoides, 17 Oct. 2003, M.A. Dick, culture CBS 118489 = CPC 10849.
Differs from the genus Phaeophleospora, in vitro, by forming
Pseudocercospora tereticornis Crous & Carnegie, Persoonia conidiophores arising from hyphae without stromata composed
23: 129. 2009. Fig. 86. from swollen hyphal cells. The conidiophores are often reduced
to conidiogenous cells, which are subhyaline to pale blackish
CROUS ET AL.
brown, proliferating percurrently, and conidia are concolourous, Description and illustration: Crous et al. (2007c).
pale brown, distoseptate.
Typus: Australia, Queensland, Cairns, Kuranda, Karoomba
Type species: Pseudophaeophleospora stonei (Crous) C. River Walk, S16°49’08.8”, E145°38’ 24.7”, on Eucalyptus sp., 19
Nakash. et al. Aug. 2006, P.W. Crous & J. Stone (holotype CBS H-19835,
cultures ex-type CBS 120830 = CPC 13330, CPC 13331,
Notes: The genus Pseudophaeophleospora is monotypic.
13332).
Pseudophaeophleospora is morphologically similar to Phaeo-
phleospora, and the two genera can be distinguished only by Pseudophloeospora Crous & R.G. Shivas, Persoonia 25: 141.
means of DNA sequence data. 2010. Fig. 88.
Pseudophaeophleospora stonei (Crous) C. Nakash., et al. Conidiomata amphigenous, pycnidial, globose, medium brown;
Stud. Mycol. 87: 311. 2017. Fig. 87. pycnidial wall consisting of 3–6 layers of brown textura angularis.
Basionym: Phaeophleospora stonei Crous, Fungal Diversity 26: Conidiophores lining the cavity, hyaline, smooth, reduced to
169. 2007. conidiogenous cells, or with 1–2 supporting cells, subcylindrical,
branched below or unbranched. Conidiogenous cells terminal or
Diagnosis: Leaf spots amphigenous, circular to subcircular, pale
lateral, hyaline, smooth, tapering to an acutely truncate apex;
brown with a raised, dark brown border, and thin, red-purple
proliferating inconspicuously percurrently at apex. Conidia hya-
margin, 1–4 mm diam. Conidia subcylindrical to narrowly
line, smooth, filiform, flexuous, subcylindrical, tapering to an
obclavate, widest at basal septum, tapering to a subtruncate,
acutely rounded apex and truncate base, not thickened nor
flattened hilum with minute marginal frill, and tapering in the
darkened, transversely euseptate.
apical cell to an obtuse apex; cellular content granular to not so
in vivo, conidia 3(–6)-euseptate (septa appear thicker in Shear’s Type species: Pseudophloeospora eucalypti Crous & R.G.
than in clear lactic acid, but never distoseptate); conidia guttulate Shivas
and darker brown in vitro, but similar in dimensions, (25–)
Notes: The genus Pseudophloeospora was established for a
30–33(–35) × (3.5–)4(–5) μm.
species associated with leaf spots of a Eucalyptus sp. collected
Fig. 87. Pseudophaeophleospora stonei (CBS 120830). A. Leaf spots. B. Leaf spot with exuding conidial cirrhus. C. Colony on OA. D, E. Conidiogenous cells. F. Conidia. Scale
bars = 10 μm.
238
Fig. 88. Pseudophloeospora spp. A–C. P. eucalyptorum (CPC 25600). A. Disease symptom. B. Conidiomata on OA. C. Conidiogenous cells with conidia. E. Disease
symptoms of P. eucalypti (CPC 17051). D, F–H. P. eucalyptigena (CPC 32532). D, F. Conidiogenous cells. G, H. Conidia. I–M. P. jollyi (CPC 17137). I–K. Conidiogenous cells.
L, M. Conidia. Scale bars = 10 μm.
in Queensland, Australia (Crous & Shivas 2010). Although it has Typus: Australia, Queensland, Brisbane, Jolley’s Lookout,
pycnidial conidiomata, and percurrently proliferating con- S27°23'59.8" E152°48'23.7", on Eucalyptus sp., 15 Jul. 2009,
idiogenous cells that give rise to scolecosporous, hyaline conidia, P.W. Crous & R.G. Shivas (holotype CBS H-20493, culture ex-
Pseudophloeospora (Orbiliales) is unrelated to Septoria s. str. type CBS 128212 = CPC 17051).
and Phloeospora s. str. (Capnodiales, Mycosphaerellaceae).
Pseudophloeospora eucalyptigena and P. jollyi need to be Pseudophloeospora eucalyptigena Crous, sp. nov. Myco-
compared to P. eucalypti (conidia 3-septate, (60–) Bank MB832046. Fig. 88.
65–75(–80) × (1.5–)2(–2.5) μm; Crous & Shivas 2010),
Etymology: Named after the host genus Eucalyptus from which it
and P. eucalyptorum (conidia 3-septate, (30–)
was isolated.
50–67(–75) × 2.5(–3) μm; Crous et al. 2016c). Although conidia
of P. eucalyptigena overlap with those of P. eucalypti in di- Conidiomata pycnidial on host, but more acervular on SNA, pale
mensions, they differ in that those of P. eucalyptigena frequently brown, up to 300 μm diam; wall of 3–4 layers of subhyaline textura
form an excentric basal appendage, lacking in the other two angularis. Conidiophores lining the inner cavity, hyaline, smooth,
species. Conidia of P. jollyi are similar to those of P. eucalyptorum, subcylindrical, branched, 1–6-septate, 20–35 × 3–4 μm. Con-
but differ in being shorter and narrower. Members of Pseudo- idiogenous cells terminal and intercalary, smooth, ampulliform,
phloeospora are considered to be minor pathogens of Eucalyptus, tapering to truncate apex, proliferating percurrently at apex,
though this single collection of P. eucalyptorum on Banksia, sug- 10–16 × 3–3.5 μm. Conidia hyaline, smooth, filiform, guttulate,
gests that it also occurs on Proteaceae. flexuous, subcylindrical, apex long subobtuse, base truncate,
1–1.5 μm, 3-septate. Several conidia form a basal excentric
Pseudophloeospora eucalypti Crous & R.G. Shivas, Persoo-
appendage (vermisporium-like), up to 10 μm long, 1.5–2 μm
nia 25: 141. 2010. Fig. 88.
diam, conidia (45–)65–77(–85) × 2–2.5 μm.
Diagnosis: Leaf spots amphigenous, irregular, pale brown, with
Culture characteristics: Colonies flat, spreading, with moderate
raised, thin, red-brown margins, 3–10 mm diam. Conidia hyaline,
aerial mycelium and feathery, lobate margins, reaching 30 mm
smooth, guttulate, filiform, flexuous, subcylindrical, widest in
diam after 2 wk at 25 °C. On MEA surface smoke grey, reverse
lower third, tapering to an acutely rounded apex and truncate
olivaceous grey. On PDA surface pale mouse grey, reverse pale
base, 0.5–1 μm wide, not thickened nor darkened, (60–)
olivaceous grey. On OA surface dirty white.
65–75(–80) × (1.5–)2(–2.5) μm, 3-septate.
Note: Phylogenetically, this species is related to, but distinct from
Description and illustration: Crous & Shivas (2010).
Pseudophloeospora eucalypti (Fig. 89).
CROUS ET AL.
Typus: Australia, New South Wales, Nullica State Forest, on lobate margins, reaching 20 mm diam after 2 wk at 25 °C. On
E. obliqua, 29 Nov. 2016, P.W. Crous, HPC 1842 (holotype CBS MEA surface isabelline, reverse hazel. On PDA surface and
H-24037, culture ex-type CBS 143441 = CPC 32532). reverse dirty white. On OA surface buff.
Pseudophloeospora eucalyptorum Crous et al., Persoonia 36: Typus: Australia, Queensland, Moreton Bay Region, Jolley’s
391. 2016. Fig. 88. Lookout, on Eucalyptus sp., 15 Jul. 2009, P.W. Crous (holotype
Diagnosis: Leaf spots amphigenous, angular to irregular, medium CBS H-24038, culture ex-type CBS 145603 = CPC 17137).
to dark brown, 2–7 μm diam with raised border. Conidia hyaline, Note: Phylogenetically, this species is distinct from all other
smooth, filiform, guttulate, flexuous, subcylindrical, widest in lower Pseudophloeospora species currently known from sequence
third, tapering to an acutely rounded apex, and truncate base, data (Fig. 89).
1.5 μm diam, 3-septate, (30–)50–67(–75) × 2.5(–3) μm.
Pseudoplagiostoma Cheew. et al., Fungal Diversity 44: 96.
Description and illustration: Crous et al. (2016c). 2010. Fig. 90.
Typus: Australia, Victoria, near Gurdies Winery, Gurdies-St. Ascomata perithecial, immersed in host tissue, oblique to hori-
Helier Road, The Gurdies, S38°22'49.8" E145°34'23.4", on zontal, depressed globose or elliptical, dark brown to black; beak
Eucalyptus sp., 7 Nov. 2014, P.W. Crous, J. Edwards & I.G. usually erumpent epiphyllously, eccentric to lateral; ostiole lined
Pascoe (holotype CBS H-22624, culture ex-type CBS with periphyses; peridium coriaceous, with sparse hyphae visible
141327 = CPC 25600). growing into the host tissue; stromatic tissue not formed. Asci
Pseudophloeospora jollyi Crous, sp. nov. MycoBank subcylindrical to long obovoid, lacking paraphyses, unitunicate,
MB832047. Fig. 88. with non-amyloid subapical ring, wedge-shaped, refractive, with
canal leading to the apex. Ascospores hyaline, ellipsoidal,
Etymology: Named after the former Brisbane mayor, William tapering towards rounded ends, usually straight, medianly 1-
Alfred Jolly, who visited the lookout around 1927. septate, wall smooth, with terminal, elongate, hyaline append-
ages. Conidiomata acervular to pycnidial, subcuticular to
Conidiomata pycnidial on host, but more acervular on SNA,
epidermal, wall composed of textura angularis. Conidiophores
brown, 250–350 μm diam; wall of 3–4 layers of subhyaline
absent. Conidiogenous cells cylindrical to ampulliform, prolifer-
textura angularis. Conidiophores lining the inner cavity,
ating enteroblastically with periclinal thickening and collarette, or
hyaline, smooth, subcylindrical, branched, 1–3-septate,
percurrently proliferating in the apical part. Conidia holoblastic,
10–20 × 2.5–3 μm. Conidiogenous cells terminal and interca-
ellipsoid, with obtuse apex and a flat protruding scar at the base,
lary, smooth, subcylindrical, tapering to truncate apex, prolifer-
0-septate.
ating percurrently at apex, at times also sympodial,
5–15 × 1.5–2 μm. Conidia hyaline, smooth, filiform, guttulate, Type species: Pseudoplagiostoma eucalypti Cheew. et al.
flexuous, subcylindrical, apex long subobtuse, base truncate, 3-
septate, (35–)60–60(–65) × (1.5–)2 μm. Symptoms: Leaf spots amphigenous, subcircular to irregular,
medium brown with blackish brown, reverse medium brown,
Culture characteristics: Colonies erumpent, spreading, with 3–20 mm diam, at times with a purple-brown margin, which can
sparse aerial mycelium, folded, cracked surface, and smooth, be dark brown in reverse.
Fig. 89. The single most parsimonious trees obtained from a phylogenetic analysis of the Pseudophloeospora ITS sequence alignment (eight strains including the outgroup;
623 characters analysed: 479 constant, 128 variable and parsimony-uninformative and 16 parsimony-informative). The tree was rooted to Discosia pseudoartocreas (GenBank
NR_132068.1) and the scale bar indicates the number of changes. Bootstrap support values higher than 49 % are shown at the nodes and novelties are highlighted with bold
text. Species names are indicated to the right of the tree. A superscript T denotes strains with a type status. Tree statistics: TL = 163 steps, CI = 0.969, RI = 0.828, RC = 0.802.
The analysis was performed using PAUP v. 4.0b10 (Swofford 2003).
240
Fig. 90. Pseudoplagiostoma spp. A–D. Disease symptoms. E–Y. P. eucalypti (CBS H-20303) E, F. Leaf spots. G, I. Ascomata. H. Ascomatal wall. I. Cross section though
ascomata. J. Ostiole. K–N. Asci. O. Ascospores. P. Conidioma. Q. Cross section though conidiomata. R–T. Conidia attached to conidiogenous cells with percurrent prolif-
eration. U. Conidia. V. Colony on MEA. W, X. Conidia and conidiogenous cells. Y. Microcyclic conidiation. Scale bars: E = 5 mm, F = 1 mm, G, I = 50 μm, H = 5 μm, J–N = 30 μm,
O, W–Y = 20 μm, P = 200 μm, Q = 70 μm, R–U = 15 μm (from Cheewangkoon et al. 2010).
CROUS ET AL.
Notes: Pseudoplagiostoma is one of the most common fungi of Ps. eucalypti CBS H-20303, cultures ex-type CBS
occurring on Eucalyptus leaves. In many cases it appears to be a 124807 = CPC 13341, CPC 13342, 13343).
primary pathogen, although it is also observed to co-occur with
Pseudoplagiostoma oldii Cheew. et al., Fungal Diversity 44:
other pathogens, presumably in a secondary role, and is also a
101. 2010.
common endophyte. More research is required to resolve its role
as pathogen. Diagnosis: Conidia holoblastic, hyaline, guttulate, smooth, thick-
walled, ellipsoidal, aseptate, slightly curved, apex obtuse, base
Pseudoplagiostoma corymbiae Crous & Summerell, Persoo-
tapering to a flat, protruding scar, (15–)17–20(–23) × (6–)
nia 28: 151. 2012.
7–8(–9) μm; on MEA, (11–)14–17(–20) × (6–)7–9(–11) μm.
Diagnosis: Leaf spots large, up to 3 cm diam, subcircular to
somewhat irregular, medium brown with thin red-brown border.
Conidia aseptate, hyaline, smooth, thick-walled, (1–2 μm diam),
Typus: Australia, Queensland, Lannercost, on E. camaldulensis,
guttulate, elongate ellipsoidal, straight, apex broadly obtuse,
6 Jan. 2007, K. Old (holotype CBS H-20300, cultures ex-type
tapering at base to a truncate hilum (1 μm diam), with minute
CBS 124808 = CMW 6675 = CPC 14155).
marginal frill, (14–)16–18(–19) × (7–)8–9(–10) μm.
Pseudoplagiostoma variabile Cheew. et al., Fungal Diversity
44: 102. 2010.
Typus: Australia, Northern Territory, Harrison Dam Conservation
Diagnosis: Conidia holoblastic, hyaline, guttulate, smooth, thin to
Area, S12°41.953' E131°24.008', on Corymbia sp., 25 Apr.
slightly thick-walled, ellipsoid, aseptate, slightly curved,
2011, P.W. Crous & B.A. Summerell (holotype CBS H-20957,
frequently constricted in the middle, apex obtuse, base tapering
to flat protruding scar, (12.5–)15.5–17.5(–23.5) × (5.5–)
Pseudoplagiostoma corymbiicola Crous, Persoonia 41: 259. 6.5–8(–9) μm; on MEA, (6.5–)15.5–17(–19) × (6.5–)
2018. 7.5–9(–10.5) μm.
Diagnosis: Conidia aseptate, hyaline, smooth, thick-walled, Description and illustration: Cheewangkoon et al. (2010).
(1–2 μm diam), guttulate, elongate ellipsoidal, straight, apex
Typus: Uruguay, on E. globulus, 5 Aug. 2002, M.J. Wingfield
broadly obtuse, tapering at base to a truncate hilum (1 μm diam),
(holotype CBS H-20304, cultures ex-type CBS 113067 = CPC
with minute marginal frill, (15–)16–17(–20) × (6–)7(–8) μm.
5320, CPC 5321).
Pseudosydowia Thambug. & K.D. Hyde, Fungal Diversity 68:
Typus: Australia, New South Wales, Dyraaba, Dyraaba plan- 140. 2014.
tation, on C. variegata, 14 Mar. 2015, A.J. Carnegie, HPC 2027
Ascomata amphigenous, black, subepidermal, becoming erum-
(holotype CBS H-23753, culture ex-type CPC 33275 = CBS
pent, separate or aggregated in clusters, globose with apical
145052).
ostiole; wall consisting of several layers of dark brown textura
Pseudoplagiostoma eucalypti Cheew. et al., Fungal Diversity angularis. Asci 8-spored, bitunicate, fasciculate, broadly ellipsoid
44: 98. 2010. Fig. 90. to clavate, straight to slightly curved, subsessile with a well-
Basionym: Cryptosporiopsis eucalypti Sankaran & B. Sutton, developed ocular chamber. Ascospores overlapping, bi- to mul-
Mycol. Res. 99: 828. 1995. tiseriate, hyaline, (1–)3(–4)-septate at maturity, constricted at the
median septum, aguttulate, thin-walled, straight to slightly curved,
Diagnosis: Leaf spots amphigenous, subcircular to irregular, obovoid to ellipsoid with obtuse ends, tapering towards both ends,
medium brown with blackish brown, reverse medium brown, but more prominently towards the lower end. Coniothyrium-like
3–20 mm diam, surrounded by a purple-brown margin, which is synanamorph: Conidiomata pycnidial to acervular, dark brown,
dark brown in reverse. Ascospores ellipsoid, tapering to rounded semi-thick-walled. Conidiophores reduced to conidiogenous cells,
ends, widest at septum, hyaline, bi- to triseriate overlapping, annellidic, integrated, indeterminate, formed from the inner cells of
fasciculate, medianly 1-euseptate; not constricted at the the conidiomatal wall, hyaline to pale brown, smooth, slightly thick-
septum, with 1–2 large guttules in each cell, thin-walled, straight, walled, short-cylindrical to narrowly ampulliform, slightly tapered
(15–)17–19(–21) × (5–)6(–7) μm; with hyaline, cylindrical ap- toward the apex. Conidia aseptate, medium brown to olivaceous-
pendages at both polar ends at maturity, expanded at the base, brown, ellipsoid to ovoid, not guttulate, thin-walled.
tapering towards the apex, 5.5–7 μm wide, (8–)15–20(–30) μm
Type species: Pseudosydowia eucalypti (Verwoerd & du Plessis)
long. Conidiomata medium to dark brown, acervular to pycnidial.
Thambug. & K.D. Hyde
Conidia holoblastic, hyaline, guttulate, smooth, thick-walled,
ellipsoid, aseptate, slightly curved, frequently slightly narrow at Notes: Sydowia eucalypti was originally described by Verwoerd
the middle, with obtuse apex; base tapering to flat protruding & du Plessis (1931) as Sphaerulina eucalypti from Eucalyptus
scar, (15–)17–19(–23) × (6.5–)7–8 (–8.5) μm; on MEA, (14–) leaves collected in Tulbach, South Africa, where it was recol-
16–19(–22) × (6–)7–9(–11) μm. lected by Crous et al. (2003). Although the sexual morph is rarely
observed, the selenophoma-like asexual morph is commonly
encountered as pycnidia or acervuli on eucalypt leaves in lots
Typus: USA, Hawaii, Kauai, on E. grandis, 23 May 1978, C.S. of countries where this crop is grown. It is frequently associ-
Hodges (holotype of Cryptosporiopsis eucalypti IMI 237416f). ated with pale brown leaf spots, forming a yeast-like growth in
Venezuela, on E. urophylla, Oct. 2006, M.J. Wingfield (holotype culture. This fungus is pleomorphic, forming selenophoma- and
242
coniothyrium-like morphs on leaves. It does not appear to be of Synonyms: Sydowia eucalypti (Verwoerd & du Plessis) Crous,
major importance. Sydowia 55: 143. 2003.
Selenophoma eucalypti Crous et al., Mycol. Res. 99: 648. 1995.
Pseudosydowia eucalypti (Verwoerd & du Plessis) Thambug.
& K.D. Hyde, Fungal Diversity 68: 140. 2014. Fig. 91. Diagnosis: Leaf spots amphigenous, subcircular, becoming
Basionym: Sphaerulina eucalypti Verwoerd & du Plessis, S. Afr. confluent, covering large areas of the leaf, pale brown, sur-
J. Sci. 28: 296. 1931. rounded by a narrow, raised, dark brown margin. Coniothyrium-
Fig. 91. Pseudosydowia eucalypti (CPC 5247). A–D. Disease symptoms. E. Conidiomata giving rise to spore masses on host leaf. F. conidia of Selenophoma morph. G.
Colony on MEA. H. Colony on pine needle agar. I, J. Conidia of coniothyrium-like synasexual morph. K. Conidiogenous cell. L–N. Asci and ascospores. Scale bars: E,
G = 10 mm; F, I = 20 μm; H = 100 μm; L–N = 40 μm (from Cheewangkoon et al. 2009).
CROUS ET AL.
like synanamorph on PNA: Conidiomata pycnidial to acervular, Synonym: Teratosphaeria flexuosa (Crous & M.J. Wingf.) Crous
dark brown, semi-thick-walled, up to 120 μm diam and 200 μm & U. Braun, Stud. Mycol. 58: 10. 2007.
high. Conidiophores reduced to conidiogenous cells, annellidic,
Diagnosis: Leaf spots occurring on juvenile foliage, being
integrated, indeterminate, formed from the inner cells of the
colonised by M. suberosa on older foliage, amphigenous, sub-
conidiomatal wall, hyaline to pale brown, smooth, slightly thick-
circular, 1–8 mm diam, pale to medium brown, border incon-
walled, short-cylindrical to narrowly ampulliform, slightly
spicuous or slightly raised, coloured margin absent; lesions pale
tapered toward the apex, (2.5–)3.5–5.5 × 1.8–3.3 μm. Conidia
brown and indistinct on lower surface. Ascospores tri- to multi-
aseptate, medium brown to olivaceous-brown, ellipsoid to ovoid,
seriate, overlapping, colourless, guttulate, thin-walled, straight to
not guttulate, thin-walled, 6–8(–10) × (2.3–)3–5.5 μm. Asco-
slightly curved, fusoid-ellipsoidal with an obtuse basal and sub-
spores bi- to multiseriate, overlapping, hyaline, non-guttulate,
obtuse apical end, widest in middle of apical cell, medianly 1-
thin-walled, straight to slightly curved, obovoid to ellipsoid with
septate, not constricted at septum, tapering toward both ends,
obtuse ends, widest at apical septum, or middle of apical cell if 1-
but more prominently toward base, (9–)10–12(–13) × (2–)
septate, (1–)3(–4)-septate at maturity, constricted at median
2.5–3 μm.
septum, tapering towards both ends, but more prominently to-
wards lower end, 10–19 × 5–7 μm. Selenophoma morph: Descriptions and illustrations: Crous (1998), Crous et al. (2007a).
Conidiomata pycnidial, amphigenous, subepidermal, con-
colourous with the lesion, with a thin-walled basal layer con- Typus: Colombia, La Selva, on E. globulus, May 1995, M.J.
sisting of 2–3 layers of walled textura angularis, up to 120 μm Wingfield (holotype PREM 54401, cultures ex-type CBS
diam; dehiscence by irregular rupture of the upper wall; 111012 = CMW 5224 = CPC 1109, CPC 1107, 1108).
becoming stromatic on leaf litter, wall developing 3–6 layers of
thick-walled textura angularis, black, separate or aggregated, Pseudoteratosphaeria gamsii (Crous) Quaedvl. & Crous,
intermingled between pseudothecia, up to 200 μm diam. Conidia Persoonia 33: 29. 2014.
hyaline, smooth, fusoid with an acutely rounded apex and sub- Basionym: Mycosphaerella gamsii Crous, Stud. Mycol. 55: 113.
truncate base, straight to falcate, 8–13(–15) × 2–4(–5) μm 2006.
in vivo. Synonym: Teratosphaeria gamsii (Crous) Crous & U. Braun,
Stud. Mycol. 58: 10. 2007.
Descriptions and illustrations: Verwoerd & du Plessis (1931),
Crous et al. (1995, 2003), Cheewangkoon et al. (2009). Diagnosis: Leaf spots amphigenous, irregular, 1–20 mm diam,
medium brown, with a raised, dark brown border. Ascospores tri-
Typus: South Africa, Western Cape Province, Tulbach, on to multiseriate, overlapping, hyaline, guttulate, thin-walled,
Eucalyptus sp., collection date unknown, B.J. Dippenaar (ho- straight, fusoid–ellipsoidal, medianly 1-septate, widest in the
lotype PREM 46423, ex Herb. L. Verwoerd, 472). middle of the apical cell, constricted at the septum, tapering
Pseudoteratosphaeria Quaedvl. & Crous, Persoonia 33: 29. towards both ends, but more prominently towards the lower end,
2014. (8–)9–10 × (2–)3 μm in vivo.
Ascomata pseudothecial, epiphyllous, single, black, subepider- Description and illustration: Crous et al. (2006d).
mal, globose; ostiole central, apical; wall of 2–3 layers of me- Typus: India, Palampur, on Eucalyptus sp., Mar. 2004, W. Gams
dium brown textura angularis. Asci aparaphysate, fasciculate, & M. Arzanlou (holotype CBS H-19690, cultures ex-type CBS
bitunicate, subsessile, obovoid to broadly ellipsoid, slightly 118495 = CPC 11138, CPC 11139, 11140).
incurved, 8-spored. Ascospores multiseriate, overlapping, hya-
line, guttulate, thinwalled, straight, fusoid-ellipsoidal, ellipsoidal or Pseudoteratosphaeria ohnowa (Crous & M.J. Wingf.) Quaedvl.
obovoid with obtuse ends, medianly 1-septate, widest in the & Crous, Persoonia 33: 29. 2014.
middle of the apical cell, constricted at the septum, tapering Basionym: Mycosphaerella ohnowa Crous & M.J. Wingf., Stud.
towards both ends, but more prominently towards the lower end. Mycol. 50: 206. 2004.
Synonym: Teratosphaeria ohnowa (Crous & M.J. Wingf.) Crous
Type species: Pseudoteratosphaeria perpendicularis (Crous & & U. Braun, Stud. Mycol. 58: 10. 2007.
M.J. Wingf.) Quaedvl. & Crous
Symptoms: Leaf spots amphigenous, subcircular to irregular, 2–10 mm diam, medium brown, with a raised border which is
medium brown, with or without a dark brown or red-purple red-brown on the adaxial surface, and medium brown on the
margin. abaxial surface. Ascospores 2–3-seriate, overlapping, hyaline,
Notes: Pseudoteratosphaeria is very similar to Teratosphaeria guttulate, thick-walled, straight, obovoid with obtuse ends,
and is best distinguished from that genus based on DNA medianly to unequally 1-septate, widest near the apex of the
phylogeny. Although the genus includes several species that apical cell, not constricted at the septum, tapering towards both
appear to be primary pathogens, some also co-occur with other ends, but more prominently towards lower end, (10–)
taxa, suggesting they could be secondary. Research is required 12–14(–15) × (3–)3–4 μm in vivo.
to clarify their biology and their possible role as plant
pathogens. Description and illustration: Crous et al. (2004b).
Pseudoteratosphaeria flexuosa (Crous & M.J. Wingf.) Typus: South Africa, Mpumalanga, Hazy View, on E. grandis, 27
Quaedvl. & Crous, Persoonia 33: 29. 2014. Mar. 1995, M.J. Wingfield (holotype PREM 51912, cultures ex-
Basionym: Mycosphaerella flexuosa Crous & M.J. Wingf., Mycol. type CBS 112896 = CPC 1004, CBS 112973 = CPC 1005, CBS
Mem. 21: 58. 1998. 110949 = CPC 1006).
244
Pseudoteratosphaeria perpendicularis (Crous & M.J. Wingf.) Basionym: Mycosphaerella secundaria Crous & Alfenas, Stud.
Quaedvl. & Crous, Persoonia 33: 29. 2014. Mycol. 55: 122. 2006.
Basionym: Mycosphaerella perpendicularis Crous & M.J. Wingf., Synonym: Teratosphaeria secundaria (Crous & Alfenas) Crous &
Stud. Mycol. 55: 113. 2006. U. Braun, Stud. Mycol. 58: 11. 2007.
Synonym: Teratosphaeria perpendicularis (Crous & M.J. Wingf.)
Diagnosis: Ascospores tri- to multiseriate, overlapping, hyaline,
guttulate, thin-walled, straight, ellipsoidal with subobtuse ends,
Diagnosis: Leaf spots amphigenous, irregular to sub-circular, medianly 1-septate, widest close to the apex of the apical cell,
5–15 mm diam, medium brown, frequently with an orange- constricted at the septum, tapering towards both ends, but more
red discolouration in the central part; border raised, dark prominently towards the lower end, 8–10 × 2.5–3 μm in vivo.
brown. Ascospores multiseriate, overlapping, hyaline, guttulate,
thin-walled, straight, fusoid–ellipsoidal with obtuse ends,
medianly 1-septate, widest in the middle of the apical cell, Typus: Brazil, Bahia, Teixeira de Freitas, on Eucalyptus sp., 8
constricted at the septum, tapering towards both ends, but more Jun. 2004, A.C. Alfenas (holotype CBS H-19697, culture ex-
prominently towards the lower end, (8–)9–10(–12) × (2.5–) type CBS 118507 = CPC 11551, CPC 11552, 11553).
3 μm in vivo.
Pseudoteratosphaeria stramenticola (Crous & Alfenas)
Description and illustration: Crous et al. (2006d). Quaedvl. & Crous, Persoonia 33: 29. 2014. Fig. 92.
Basionym: Mycosphaerella stramenticola Crous & Alfenas, Stud.
Typus: Colombia, Suiza, on E. grandis × E. urophylla, Jan.
Mycol. 55: 123. 2006.
2004, M.J. Wingfield (holotype CBS H-19691, cultures ex-type
Synonyms: Teratosphaeria stramenticola (Crous & Alfenas)
CBS 118367 = CPC 10983, CPC 10984, 10985).
Pseudoteratosphaeria secundaria (Crous & Alfenas) Quaedvl. Mycosphaerella parkiiaffinis Crous & M.J. Wingf., Fungal Di-
& Crous, Persoonia 33: 29. 2014. versity 26: 168. 2007.
Fig. 92. Pseudoteratosphaeria stramenticola (CBS H-19823). A. Leaf spot. B. Colony on MEA. C, D. Asci. E. Ascospores. F–H. Germinating ascospores. Scale bar = 10 μm
(from Crous et al. 2007c).
CROUS ET AL.
Teratosphaeria parkiiaffinis (Crous & M.J. Wingf.) Crous & U. 19699, cultures ex-type CBS 118506 = CPC 11438, CPC 11439,
Braun, Stud. Mycol. 58: 10. 2007. 11440).
Diagnosis: Leaf spots amphigenous, irregular to subcircular, Pseudozasmidium Videira & Crous, Stud. Mycol. 87: 378. 2017.
6–30 mm diam, pale to medium brown, with a thin, raised, dark Fig. 93.
brown border, and a red-purple margin. Ascospores tri- to mul-
Pseudothecia amphigenous, aggregated, black, immersed and
tiseriate, overlapping, hyaline, guttulate, thin-walled, straight,
becoming erumpent, wall of 2–3 layers of medium brown textura
fusoid-ellipsoidal with obtuse ends, widest in middle of apical cell,
angularis. Asci aparaphysate, fasciculate, bitunicate, subsessile,
medianly 1-septate, constricted at the septum, tapering towards
narrowly ellipsoid or obclavate to cylindrical, straight or slightly
both ends, but more prominently towards the lower end, (8–)
incurved, 8-spored. Ascospores bi- to triseriate, overlapping,
9–10 × 3(–3.5) μm.
hyaline, straight to slightly curved, ellipsoid or fusoid-ellipsoid,
Description and illustration: Crous et al. (2007c). with obtuse ends, medianly 1-septate, not constricted to
slightly constricted at the septum, symmetrical cells or widest at
Typus: Brazil, Bahia, Eunapolis, on Eucalyptus sp., 23 May the middle of the apical cell, tapering towards both ends or more
2004, A.C. Alfenas (holotype of M. stramenticola, CBS H- prominently towards lower end. Ascospore germination parallel
Fig. 93. Pseudozasmidium spp. A, B. Disease symptoms of P. parkii on Eucalyptus grandis (PREM 50668). C. Close-up of leaf spots with pseudothecia. D–L. P. nabiacense
(CBS H-20273). D. Leaf spot. E–H. Conidiophores developing in culture. I–L. Conidia. Scale bars = 10 μm.
246
to perpendicular to the long axis of the spore. Mycelium internal 35–45(–55) × (2–)3–3.5 μm; hila and scars thickened, dark-
and external, internal hyphae branched, septate, smooth and ened and refractive, 1–1.5 μm wide.
hyaline, external hyphae verruculose and pale to medium brown,
terminal hyphal ends may develop clusters of globose, multi-
celled chlamydospore-like structures. Conidiophores pale to Typus: Australia, New South Wales, Nabiac, on Eucalyptus sp.
medium brown, smooth to verruculose, erect, subcylindrical, (red gum), 30 Nov. 2005, coll. A.J. Carnegie, isol. P.W. Crous
straight or curved, branched or unbranched, repeatedly genic- (holotype CBS H-20273, cultures ex-type CBS 125010 = CPC
ulate, septate, sometimes reduced to conidiogenous cells. 12748, CPC 12749, 12750).
Conidiogenous cells terminal, smooth to verruculose, pale brown
Pseudozasmidium parkii (Crous & Alfenas) Videira & Crous,
to brown, proliferating sympodially, sometimes repeatedly
Stud. Mycol. 87: 379. 2017. Fig. 93.
geniculate, with conidiogenous loci thickened and darkened-
Basionym: Stenella parkii Crous & Alfenas, Mycologia 87: 121.
refractive. Conidia single, pale brown to olivaceous brown,
1995.
smooth to verruculose, obclavate, narrowly obclavate to sub-
Synonyms: Zasmidium parkii (Crous & Alfenas) Crous & U.
cylindrical, obtuse apex and obconically truncate base, straight
Braun, Schlechtendalia 20: 102. 2010.
or curved, 1- to multiseptate, hila thickened and darkened-
Paramycosphaerella parkii (Crous & Alfenas) Guatimosim et al.,
refractive.
Persoonia 37: 127. 2016.
Type species: Pseudozasmidium parkii (Crous & Alfenas) Mycosphaerella parkii Crous et al., Mycol. Res. 97: 582. 1993.
Videira & Crous (≡ Stenella parkii Crous & Alfenas) Diagnosis: Leaf spots amphigenous, varying from small to large,
Notes: Species of Pseudozasmidium are known to occur on round or slightly irregular spots, light brown in colour, sur-
eucalypts in South America and Asia. Nothing is known about rounded by a thin red to brown line, forming a raised margin in
their relative importance, although Z. parkii was associated with most cases. Ascospores 2–3-seriate or irregularly arranged,
prominent leaf spotting on Eucalyptus spp. in Brazil (Crous et al. oblique, overlapping, straight ellipsoidal, obtuse at each end,
1993c, Crous & Alfenas 1995). colourless, smooth, 1-septate, eguttulate, not constricted at
septum, widest in middle of upper cell, tapering more promi-
Pseudozasmidium eucalypti (Crous & Summerell) Videira & nently toward base, (8.5–)9–13(–15) × (2–)2.5–3(–3.5) μm
Crous, Stud. Mycol. 87: 378. 2017. in vivo, (6.5–)8–11(–9.5) × (2–)2.5–3 μm in vitro. Conidia
Basionym: Stenella eucalypti Crous & Summerell, Fungal Di- olivaceous brown, verruculose, single, obclavate, apex obtuse,
versity 26: 177. 2007. base long obconic-truncate, straight or curved, indistinctly 1–8-
Synonym: Zasmidium eucalypti (Crous & Summerell) Crous & U. septate, 25–200 × 2–2.5 μm; hila slightly thickened and
Braun, Schlechtendalia 20: 101. 2010. darkened.
Diagnosis: Leaf spots amphigenous, irregular to angular specks, Descriptions and illustrations: Crous et al. (1993c), Crous &
1–3 mm diam, pale brown with dark brown, with raised, dark Alfenas (1995).
brown spots inside lesions, presumably due to insect damage;
Typus: Brazil, Aracruz, Florestal nursery, on E. grandis, 24 Feb.
borders raised, margins absent to red-purple, but the latter may
1990, M.J. Wingfield (holotype of Mycosphaerella parkii PREM
be due to co-colonisation of a Pseudocercospora sp. Conidia
50668, ex-type culture CBS 387.92 = CMW 14775 = CPC 353);
solitary, pale brown, finely verruculose, guttulate, subcylindrical
S~ao Paulo, on E. saligna, Apr. 1993, P.W. Crous (holotype of
to narrowly obclavate, apex subobtuse, base long obconically
Stenella parkii PREM 51713).
subtruncate to obconically subtruncate, straight to slightly
curved, (0–)1–3(–5)-septate, (10–)20–35(–60) × (2–) Pseudozasmidium vietnamense (Barber & T.I. Burgess)
3–4(–6) μm; hila thickened, darkened, refractive, 1.5–2 μm Videira & Crous, Stud. Mycol. 87: 379. 2017.
wide. Basionym: Mycosphaerella vietnamensis Barber & T. I. Burgess,
Fungal Diversity 24: 148. 2007.
Synonym: Paramycosphaerella vietnamensis (Barber & T.I.
Typus: Australia, Queensland, Cairns, Eureka Creek, 48 km Burgess) Guatimosim et al., Persoonia 37: 128. 2016.
from Mareeba, S17°11’13.2”, E145°02’27.4”, alt. 468 m, on
Diagnosis: Leaf spots amphigenous, sub-circular to circular,
E. tereticornis, 26 Aug. 2006, P.W. Crous (holotype CBS H-
vein-limited, 2–5 mm diam, medium brown, surrounded by a
19830, culture ex-type CBS 121101 = CPC 13302).
thin, raised, brown border and occasionally a spreading pink
Pseudozasmidium nabiacense (Crous & Carnegie) Videira & margin. Ascospores bi- to triseriate, overlapping, hyaline, gut-
Crous, Stud. Mycol. 87: 379. 2017. Fig. 93. tulate, thick-walled, straight to slightly curved, fusoid-ellipsoidal
Basionym: Zasmidium nabiacense Crous & Carnegie, Persoonia with obtuse ends, medianly 1-septate, widest at the middle of
23: 142. 2009. the apical cell, slightly constricted at the septum, tapering to-
Synonym: Paramycosphaerella nabiacensis (Crous & Carnegie) wards both ends but more prominently towards lower end,
Guatimosim et al., Persoonia 37: 127. 2016. 10–12 × 3–4 μm.
Description and illustration: Burgess et al. (2007).
Diagnosis: Leaf spots amphigenous, irregular to angular,
3–15 mm diam, medium brown with a concolourous, raised Typus: Vietnam, South East Forestry Institute nursery, on
border. Conidia solitary, medium brown, finely verruculose, gut- E. grandis hybrid, 6 Jul. 2004, coll. T.I. Burgess, isol. P.A. Barber
tulate, narrowly obclavate, apex obtuse, base long obconically (holotype MURU 411, ex-type culture CBS 119974 = CMW
subtruncate, straight to slightly curved, 3–5-septate, (20–) 23441 = MUCC 66 = VTN1).
CROUS ET AL.
Quambalaria J.A. Simpson, Australas. Mycol. 19: 60. 2000. thickened pale scars, flat or on small geniculations or on
Fig. 94. small denticles. Conidia holoblastic, hyaline, aseptate, smooth,
Synonym: Fugomyces Sigler, Manual Clin. Microbiol. 2: 1753. variable in shape, clavate to elongated-clavate, cylindrical,
2003. fusiform or narrowly pyriform, or obovoid, single, with a slightly
refractive non-thickened scar at one or both ends, germination
Pustules arising from stromatic layer, subcuticular or in the
by one or two hyphae. Secondary conidia arising from primary
stomatal cavity, white. Conidiophores erumpent, single or
conidia (= ramoconidia), either by budding or from very
caespitose, hyaline, simple, rarely branched, sparsely septate,
short conidiogenous cells with scars and sympodial
with sympodial proliferation and small slightly refractive non-
Fig. 94. Quambalaria spp. A–C. Disease symptoms of Q. pitereka on Eucalyptus sp. D–I. Conidiophores, conidiogenous cells, primary and secondary conidia of Q. pusilla
(CPC 14499). J–M. Conidiophores, conidiogenous cells, primary and secondary conidia of Q. tasmaniae (CPC 25464). N–Q. Conidiophores, conidiogenous cells, primary and
secondary conidia of Q. rugosae (CPC 20162). Scale bars = 10 μm
248
proliferation, forming short simple or branched chains (from Diagnosis: Primary conidia obovoid, subcylindrical, pyriform,
Simpson 2000). subclavate or somewhat irregularly shaped,
5–20 × 2.5–5(–6.5) μm, aseptate, hyaline, smooth or almost so,
Type species: Quambalaria pitereka (J. Walker & Bertus) J.A.
ends obtuse, occasionally with a single or rarely several sec-
Simpson
ondary conidia, apical or subapical, shape and size agreeing with
Symptoms: Lesions develop on immature leaves and stems, those of the primary conidia; hila conspicuous, unthickened, not
forming discrete chlorotic spots with necrotic centres that darkened, but somewhat refractive.
develop into large, sporulating lesions. Under favourable con-
Description and illustration: Braun (1998).
ditions the disease can cause stem girdling on seedlings, and
leaf and shoot blight on ministumps. Quambalaria pitereka Typus: Australia, New South Wales, W. Pennant Hill, Forestry
causes leaf and shoot blight on Corymbia spp., regardless of Nursery, on leaves and shoots of E. eximia, 3 Apr. 1970, A.L.
conditions. Bertus & J. Walker (holotype DAR 19773).
Quambalaria spp. can cause shoot blight, which is a major Note: Quambalaria pitereka is a major disease in Corymbia
disease affecting spotted gum (Corymbia spp.). Several species plantations in eastern Australia, causing leaf and shoot blight,
are associated with leaf spots, shoot blight and tip death, which results in branch death and multi-stemmed trees
resulting in loss of growth, apical dominance, affecting tree form, (Carnegie 2007b, Pegg et al. 2009).
and tree death in severe cases.
Quambalaria pusilla (U. Braun & Crous) J.A. Simpson, Aus-
Quambalaria cyanescens (de Hoog & G.A. de Vries) Z.W. de tralas. Mycol. 19: 61. 2000. Fig. 94.
Beer et al., Stud. Mycol. 55: 295. 2006. Basionym: Sporothrix pusilla U. Braun & Crous, in Braun,
Basionym: Sporothrix cyanescens de Hoog & G.A. de Vries, Monogr. Cercosporella, Ramularia Allied Genera (Phytopath.
Antonie van Leeuwenhoek 39: 515. 1973. Hyphom.) 2: 418. 1998.
Synonyms: Cerinosterus cyanescens (de Hoog & G.A. de Vries) Synonym: Quambalaria simpsonii Cheew. & Crous, Persoonia
R.T. Moore, Stud. Mycol. 30: 216. 1987. 23: 77. 2009.
Fugomyces cyanescens (de Hoog & G.A. de Vries) Sigler, In
Diagnosis: Primary conidia aseptate, hyaline, smooth, thin-
Murray et al., Manual of Clinical Microbiology 2: 1763. 2003.
walled, fusiform or ellipsoid, with tapered base, (4.6–)
Diagnosis: Primary conidia hyaline, smooth-walled or finely 5.5–8.5(–10.4) × (2.5–)3(–3.5) μm, usually giving rise to one or
verrucose, obovoidal, 3–4 μm long, somewhat larger several obovoid to fusoid secondary conidia, (2.5–)
(3.5–6.5 μm long) when bearing secondary conidia. 3–4.5(–6.2) × (1.7–)2–2.5(–2.9) μm, sometimes giving rise to
1–4 obovoid, tertiary conidia, (2–)2.5(–3.4) × (1.3–)
Description and illustration: de Hoog & de Vries (1973).
1.5–1.8(–2.3) μm.
Typus: Netherlands, Groningen, from skin of man, 18 Oct 1959,
Descriptions and illustrations: Braun (1998), Cheewangkoon
T.F. Visser (holotype CBS H-7779, culture ex-type CBS
et al. (2009).
357.73 = CMW 5583 =MUCL 19329).
Typus: Australia, Northern Territory, Edith Falls S14°05'20",
Note: Quambalaria cyanescens is not considered a primary
E132°05'12"E, on E. tintinnans, 1 Jan. 2007, coll. B.A. Sum-
pathogen of eucalypts.
merell, isol. P.W. Crous (holotype of Q. simpsonii CBS H-20291,
Quambalaria eucalypti (M.J. Wingf. et al.) J.A. Simpson, culture ex-type CPC 14499 = CBS 124772, CPC 14500, 14501).
Australas. Mycol. 19: 61. 2000. Thailand, on E. camaldulensis, Oct. 1995, M.J. Wingfield (ho-
Basionym: Sporothrix eucalypti M.J. Wingf. et al., Mycopatho- lotype of Q. pusilla in HAL).
logia 123: 160. 1993.
Note: The synonymy of Q. pusilla with Q. simpsonii is based on
Diagnosis: Primary conidia hyaline, smooth-walled, fusoid, DNA sequences derived from the dried fungarium specimen of
aseptate, 6–12 × 2.5–4 ìm, secondary conidia obovoid to Q. pusilla (W. de Beer, in prep.).
ellipsoid, 3–5 × 1.5–2.5 μm.
Quambalaria rugosae Crous, sp. nov. MycoBank MB832048.
Description and illustration: Wingfield et al. (1993). Fig. 94.
Typus: South Africa, KwaZulu-Natal, Kwambonambi, on Etymology: Name refers to Eucalyptus rugosa, the host species
E. grandis leaves, 19 May 1987, M.J. Wingfield (holotype PREM from which this fungus was isolated.
51089, culture ex-type CBS 118844 = CMW 1101); KwaZulu-
Mycelium consisting of hyaline, smooth, branched, septate,
Natal, Kwambonambi, on E. grandis leaves, 2001, L. Lombard
1.5–2 μm diam hyphae. Conidiophores solitary, erect, sub-
(epitype PREM 58939, culture ex-epitype CBS 119680 = CMW
cylindrical, flexuous, branched, multiseptate, or reduced to con-
11678).
idiogenous cells on hyphae, hyaline, smooth, up to 150 μm long,
Quambalaria pitereka (J. Walker & Bertus) J.A. Simpson, with fertile lateral branches. Conidiogenous cells integrated, ter-
Australas. Mycol. 19: 60. 2000. Fig. 94. minal and intercalary, 5–25 × 1.5–2 μm, hyaline, smooth, sub-
Basionym: Ramularia pitereka J. Walker & Bertus, Proc. Linn. cylindrical, apex slightly swollen due to aggregated subdenticulate
Soc. N.S.W. 96: 108. 1971. conidiogenous loci; scars slightly refractive, not thickened, trun-
Synonym: Sporothrix pitereka (J. Walker & Bertus) U. Braun, In: cate, 0.5 μm diam. Conidia aseptate, hyaline, smooth, thin-walled,
Braun, Monogr. Cercosporella, Ramularia Allied Genera (Phy- aggregated in clusters; primary (ramo-) conidia fusoid-ellipsoid,
topath. Hyphom.) 2: 416. 1998. prominently tapered at base, (6–)7–8(–10) × (2–)2.5(–3) μm;
CROUS ET AL.
secondary and tertiary conidia aseptate, hyaline, smooth, Pseudovularia Speg., Anales Mus. Nac. Buenos Aires, Ser. 3,
obovoid, (3–)3.5–4(–5) × (1.5–)2(–2.5) μm; hila slightly refrac- 20: 418. 1910.
tive, not thickened nor darkened, 0.5 μm diam. For additional synonyms see Braun (1998).
Culture characteristics: Colonies flat, spreading, with folded Mostly phytopathogenic (leaf spots, chlorosis or necrosis),
surface and sparse aerial mycelium, smooth, lobate margin, sometimes saprobic or mycophylic. Conidiophores individual or
reaching 20 mm diam on MEA, PDA and OA; surface powdery, synnematous, sometimes forming small to sporodochial caes-
white; reverse luteous (MEA), pale luteous (PDA). pituli, emerging through stomata or through the cuticle, straight,
subcylindrical to geniculate-sinuous, continuous or septate, hy-
Typus: Australia, Kangaroo Island, on E. rugosa, Dec. 2011, W.
aline or in some species with a faintly reddish tinge, occasionally
Quaedvlieg (holotype CBS H-24040, culture ex-type CBS
branched, thin-walled, usually smooth but rarely rough. Con-
145601 = CPC 20162).
idiogenous cells integrated, terminal, polyblastic, sympodially
Note: Phylogenetically, this species is distinct from other elongating, straight to geniculate-sinuous, conidiogenous loci
Quambalaria species currently known from sequence data, being conspicuously thickened, darkened and refractive, coronate
closest related to Q. pitereka (Fig. 95). (cladosporoid). Conidia consistently solitary or in simple or
branched chains, solitary conidia 0–1-septate, catenate conidia
Quambalaria tasmaniae Crous, sp. nov. MycoBank aseptate to multiseptate (mostly 1–4 eusepta), hyaline, in a few
MB832049. Fig. 94. species with a faintly reddish tinge, usually ellipsoid-ovoid, cy-
lindrical-fusiform, rarely filiform, occasionally constricted at the
Etymology: Name refers to Tasmania, the island from which this septa, thin-walled, smooth to verruculose-echinulate, hila distinct,
fungus was collected. slightly to conspicuously thickened, darkened, refractive; conidial
secession schizolytic (Videira et al. 2016).
Associated with leaf spots. Mycelium consisting of hyaline,
smooth, branched, septate, 1.5–2 μm diam hyphae. Co- Type species: Ramularia pusilla Unger
nidiophores solitary, erect, subcylindrical, straight to geniculate- Note: Species of Ramularia are generally regarded to be of minor
sinuous, reduced to conidiogenous cells on hyphae, or up to 3- importance as foliar pathogens of Eucalyptus.
septate, hyaline, smooth, 5–50 × 1.5–2 μm. Conidiogenous
cells integrated, terminal and intercalary, 5–15 × 1.5–2 μm, hy- Ramularia eucalypti Crous, Fungal Diversity 26: 174. 2007.
aline, smooth, subcylindrical, apex slightly swollen due to aggre- Fig. 96.
gated subdenticulate conidiogenous loci; scars slightly refractive, Diagnosis: Leaf spots amphigenous, irregular, subcircular or
not thickened, truncate, 0.5–1 μm diam. Conidia aseptate, hyaline, angular, confined by leaf veins, medium brown with a thin, red-
smooth, thin-walled, arranged in clusters; primary (ramo-) brown border, specks 1–2 mm diam, or larger spots and
conidia fusoid-ellipsoid, prominently tapered at base, (5.5–) blotches up to 4 cm diam. Conidia catenulate in branched chains,
6–7(–8) × (1.5–)2(–3) μm; secondary (rarely tertiary) conidia smooth, hyaline; ramoconidia subcylindrical to fusoid-ellipsoidal,
aseptate, hyaline, smooth, obovoid, (2–)3–3.5(–5) × 1.5(–2) μm; 0–1-septate, (10–)12– 15(–18) × (2.5–)3(–4) μm; secondary
hila slightly refractive, not thickened nor darkened, 0.5 μm diam. conidia fusoid-ellipsoidal, occurring in branched chains of up to
Culture characteristics: Colonies flat, spreading, with folded 15 μm long, (5–)6–7(–8) × 3(–3.5) μm; hila darkened, thick-
surface and sparse aerial mycelium, smooth, lobate margin, ened, refractive, up to 1 μm wide.
reaching 30 mm diam on MEA, PDA and OA; surface powdery, Descriptions and illustrations: Crous et al. (2007c), Videira et al.
white; reverse luteous to ochreous (MEA), pale luteous (PDA). (2015a).
Typus: Australia, Tasmania, Royal Tasmanian Botanical Gar- Typus: Italy, Norcia, on C. grandifolia, 10 May 2006, W. Gams
den, on Eucalyptus sp., 12 Nov. 2014, P.W. Crous, HPC 121 (holotype CBS H-19832, culture ex-type CBS 120726 = CPC
(holotype CBS H-24039, culture ex-type CBS 145602 = CPC 13043).
25464).
Ramularia glennii Videira & Crous, Persoonia 33: 57. 2014.
Additional material examined: Australia, Tasmania, next to highway, on Euca-
lyptus sp., 12 Nov. 2014, P.W. Crous, HPC 120, culture CBS 145598 = CPC Diagnosis: Ramoconidia hyaline, smooth to finely verruculose,
25462. subcylindrical to clavate or oval, 0–1-septate, hyaline, (6–)
Note: Phylogenetically, this species is distinct from other 9–11(–15) × (2–)3(–4) μm. Intercalary conidia hyaline, smooth
Quambalaria species currently known from sequence data, being to finely verruculose, aseptate, fusiform or oval, (5–)
closest related to Q. eucalypti (Fig. 95). 6.5–8(–12) × (2–)2.5(–3) μm, in branched chains of up to
seven. Terminal conidia, hyaline, smooth to finely verruculose,
Ramularia Unger, Exanth. Pflanzen (Wien): 169. 1833. emend. aseptate, obovoid, (3–)5–5.5(–8) × (1.5–)2(–3) μm; hila
U. Braun (nom. cons.) thickened, darkened, refractive, 0.5–1 μm diam.
Synonyms: Didymaria Corda, Icon. fung. (Prague) 5: 9. 1842.
Septocylindrium Bonord. ex Sacc., Michelia 2: 15. 1880. Description and illustration: Videira et al. (2015a).
Acrotheca Fuckel, Jahrb. Vereins Naturk. Herzogth. Nassau 15: Typus: Netherlands, Rotterdam, Maasstad Ziekenhuis (Clara),
43. 1860. on human bronchial alveolar lavage, 2011, unknown collector,
Phacellium Bonord., in Rabenh., Fungi Eur. Exs., Edn. 2, ser. 2: dep. A. van Duin (holotype CBS H-21617, culture ex-type CBS
no. 288. 1860. 129441).
Ovularia Sacc., Michelia 2: 17. 1880.
Ophiocladium Cav., Z. Pflanzenkrankh. 3: 26. 1893. Readeriella Syd. & P. Syd., Ann. Mycol. 6: 484. 1908. Fig. 97.
250
Fig. 95. The first of 384 equally most parsimonious trees obtained from a phylogenetic analysis of the Quambalaria ITS sequence alignment (105 strains including the
outgroup; 581 characters analysed: 317 constant, 189 variable and parsimony-uninformative and 75 parsimony-informative). The tree was rooted to Burgoa verzuoliana
(GenBank NR_145334.1) and the scale bar indicates the number of changes. Thickened branches represent those present in the strict consensus parsimony tree. Bootstrap
support values higher than 49 % are shown at the nodes and novelties are highlighted with bold text. Species names are indicated to the right of the tree and GenBank
accession numbers between round parentheses. A superscript T denotes strains with a type status and ET those with ex-epitype status. Tree statistics: TL = 364 steps,
CI = 0.860, RI = 0.973, RC = 0.837. The analysis was performed using PAUP v. 4.0b10 (Swofford 2003).
Synonyms: Nothostrasseria Nag Raj, Canad. J. Bot. 61: 23. broadly ellipsoid, straight to slightly curved, 8-spored. Asco-
1983. spores bi- to triseriate, overlapping, hyaline, guttulate, thin-
Cibiessia Crous, Fungal Diversity 26: 151. 2007. walled, straight, fusoid-ellipsoidal with obtuse ends, medianly
1-septate, not to slightly constricted at the septum; encased in a
Ascomata pseudothecial, amphigenous, black, subepidermal,
mucus sheath. Conidiomata pycnidial, brown, globose; wall
globose; wall consisting of 2–3 layers of medium brown textura
consisting of 2–3 layers of brown textura angularis.
angularis. Asci aparaphysate, fasciculate, bitunicate, subsessile,
CROUS ET AL.
Fig. 95. (Continued).
Conidiophores 0–1-septate, pale brown, finely verruculose, limoniform, at times tapering to a tubular basal appendage
ampulliform to doliiform. Conidiogenous cells pale brown, finely (present in some species), thick-walled, with darker pigmentation
verruculose, ampulliform to doliiform, proliferating several times in the lateral projections (present in some species), but with more
percurrently near apex, mono- or polyphialidic. Conidia solitary, prominent constriction between the projections and the base.
medium brown, aseptate, smooth, granular, base truncate, with Cibiessia synasexual morph: Hyphae pale brown, smooth,
three apical, lateral, obtuse projections, deltoid, or ellipsoid to 3–5 μm diam, disarticulating at septa to form short, pale brown,
252
cylindrical conidia with obtusely rounded to subtruncate ends; Readeriella dendritica (Crous & Summerell) Crous & Sum-
conidia 0(–1)-septate, becoming medium brown, verruculose, merell, Persoonia 23: 111. 2009. Fig. 97.
ellipsoidal to subglobose or globose, with dehiscence scars Basionym: Mycosphaerella dendritica Crous & Summerell,
clearly visible on conidial body. Fungal Diversity 26: 161. 2007.
Synonyms: Teratosphaeria dendritica (Crous & Summerell)
Type species: Readeriella mirabilis Syd. & P. Syd.
Symptoms: Leaf spots amphigenous, irregular to subcircular, Spilomyces dendriticus Hansf., Proc. Linn. Soc. New South
2–8 mm diam, pale to grey-brown, with raised borders and thin, Wales 81: 32. 1956.
dark brown margins. Nothostrasseria dendritica (Hansf.) Nag Raj, Canad. J. Bot. 61:
25. 1983.
Notes: Species of Readeriella are very common on eucalypt
foliage and appear to also be endophytic. Although some Diagnosis: Leaf spots amphigenous, irregular to subcircular,
species are associated with prominent leaf spot disease symp- 2–8 mm diam, pale to grey-brown, with raised borders and thin,
toms, the majority are regarded as pathogens of minor dark brown margins. Ascospores bi- to triseriate, overlapping,
importance. hyaline, guttulate, thin-walled, straight, fusoid-ellipsoidal with
obtuse ends, widest just above the septum, medianly 1-septate,
Readeriella angustia Crous & Summerell, Persoonia 23: 132.
not to slightly constricted at the septum, tapering towards both
2009.
ends, but more prominently towards the lower end, (11–)
12–13(–15) × 3–3.5(–4.5) μm; encased in a mucus sheath.
Diagnosis: Leaf spots amphigenous, raised, dark brown with a
Conidia consisting of an ellipsoid body with obtuse apex, tapering
raised border and thin red-brown margin, subcircular to angular,
to a tubular basal appendage; body medium brown, verruculose,
1–3 mm diam. Conidia narrowly ellipsoid to subcylindrical with
6–12 × 3–4 μm; tubular appendage separated from the
rounded ends, pale brown, smooth to finely verruculose,
conidium body by a septum, unbranched, hyaline, smooth,
3–5 × 1.5–2 μm. Cibiessia morph: cylindrical conidia with
4–15 × 1–1.5 μm.
obtusely rounded to subtruncate ends; aseptate conidia
2–4 × 2–3 μm, 1(–3)-septate conidia 5–10 × 2–4.5 μm. Description and illustration: Crous et al. (2007c).
Description and illustration: Crous et al. (2009b). Typus: Australia, New South Wales, Rylstone, S32°39'31",
E150°12'30", on E. deanei, Feb. 2006, coll. B.A. Summerell, isol.
Typus: Australia, Tasmania, Mount Wellington, S42°55'0",
P.W. Crous (holotype of Mycosphaerella dendritica CBS H-19772,
E147°15'0", on E. delegatensis, 14 Oct. 2006, coll. B.A. Sum-
cultures ex-type CBS 120032 = CPC 12709, CPC 12710, 12711).
merell (holotype CBS H-20259, cultures ex-type CBS
124997 = CPC 13608, CPC 13609, 13610, CBS 124998 = CPC Readeriella dimorphospora (Crous & C. Mohammed) Crous,
13618, CPC 13619, 13620). Persoonia 23: 111. 2009.
Readeriella callista (Syd.) Crous & Summerell, Persoonia 23: Basionym: Cibiessia dimorphospora Crous & C. Mohammed,
111. 2009. Fig. 97. Fungal Diversity 26: 151. 2007.
Basionym: Coniothyrium callistum Syd., Ann. Mycol. 35: 358. Diagnosis: Readeriella morph: conidia narrowly ellipsoid to
1937. subcylindrical with rounded ends, pale brown, smooth to finely
Synonym: Microsphaeropsis callista (Syd.) B. Sutton, Mycol. verruculose, 4.5–7 × 2–2.5 μm. Cibiessia morph: cylindrical
Pap. 123: 35. 1971. conidia with obtusely rounded to subtruncate ends; aseptate
Diagnosis: Leaf spots brown, raised spots, 5–10 mm diam. conidia 5–7 × 2–3 μm, 1(–3)-septate conidia 5–9 × 2–3 μm;
Conidia 6–8.5 × 3.5–5.5 μm, 1-celled, brown, smooth, thick- conidia developing further, becoming medium brown, predomi-
walled, with a central guttule, elliptical, tapering to truncate base. nantly aseptate, verruculose, ellipsoidal to subglobose or
globose, 5–7 μm diam, with dehiscence scars clearly visible on
Description and illustration: Crous et al. (2009b). conidial body; inner layer of the dehiscence scar extends past
Typus: Australia, New South Wales, Bulli, on E. haemastoma, the outer layer.
Aug. 1935, F. Fraser (holotype IMI 21230); New South Wales, Description and illustration: Crous et al. (2007c).
Woodford, S33°43'30", E150°29'25", on E. sclerophylla, Oct.
2006, coll. B.A. Summerell, isol. P.W. Crous (epitype CBS H- Typus: Australia, Tasmania, on E. nitens, Oct. 2005, C.
20246, cultures ex-type CBS 124986 = CPC 13615, CPC 13616, Mohammed (holotype CBS H-19762, cultures ex-type CBS
13617). 120034 = CPC 12636, CPC 12637, 12638).
Readeriella deanei Quaedvl. et al., Persoonia 33: 30. 2014. Readeriella ellipsoidea Crous, Persoonia 38: 283. 2017.
Diagnosis: Conidia solitary, aseptate, ellipsoid to limoniform, Diagnosis: Conidia solitary, narrowly ellipsoid, apex obtuse,
tapering towards a bluntly rounded, subobtuse, thickened apex, tapering to a narrowly truncate base, 1 μm diam, yellow brown in
base subtruncate and thickened, hyaline becoming medium to mass, finely roughened, (4–)5(–6) × (2–)2.5 μm.
golden brown, finely verruculose, (8–)9–10(–11) × 4(–5) μm. Description and illustration: Crous et al. (2017b).
Description and illustration: Quaedvlieg et al. (2014). Typus: Australia, Western Australia, Albany, Stirling Range
Typus: Australia, New South Wales, Wollemi National Park, on National Park, Bluff Knoll, S34°22'3.8" E118°14'313", on Euca-
E. deanei, 9 Feb. 2006, B. Summerell (holotype CBS H-21136, lyptus sp., 23 Sep. 2015, P.W. Crous (holotype CBS H-23088,
culture ex-type CBS 134746 = CPC 12715). culture ex-type CBS 142525 = CPC 29153).
CROUS ET AL.
Readeriella eucalypti (Gonz. Frag.) Crous, Fungal Diversity 23: 21135, cultures ex-type CBS 134745 = CPC 12727, CPC 12728,
342. 2006. Fig. 97. 12729).
Basionym: Coniothyrium olivaceum var. eucalypti Gonz. Frag.,
Readeriella menaiensis Crous & Summerell, Persoonia 23:
Bolm Soc. broteriana, Coimbra, ser. 2 2: 59. 1924.
135. 2009.
Synonym: Microsphaeropsis eucalypti (Gonz. Frag.) B. Sutton,
Mycol. Pap. 123: 37 (1971) Diagnosis: Leaf spots amphigenous, irregular to subcircular,
medium brown, frequently grey-brown in center, border raised,
Diagnosis: Associated with brown, subcircular leaf spots
dark brown, 2–5 mm diam. Conidia solitary, medium brown,
on living leaves. Conidia globose, subglobose to pyriform,
aseptate, smooth, granular, base truncate, with three apical,
base truncate, thick-walled, dark-brown, smooth-walled,
lateral, obtuse projections, deltoid, thick-walled, with darker
6 –7 × 5 –6 μm; in older cultures conidia are frequently more
pigmentation in the lateral projections, 9–10(–11) μm long, (7–)
pyriform in shape (onset of germination?), and also vary from
8–9 μm wide at apex.
being smooth to rough.
Typus: Australia, New South Wales, Menai, Central Coast,
Typus: Portugal, Taboaco, on E. globulus, Nov. 1922, leg. J.
S34°00'38", E151°00'57", on E. oblonga, 23 Sep. 2007, coll.
Macedo Pinto (holotype MA). Spain, on E. globulus, Apr. 2004,
B.A. Summerell, isol. P.W. Crous (holotype CBS H-20265,
M.J. Wingfield (epitype CBS H-19741, cultures ex-epitype CBS
120079 = CPC 11184, CPC 11185, 11186).
14449).
Readeriella eucalyptigena Crous & Summerell, Persoonia 23: Readeriella minutispora (Crous & Carnegie) Crous & Carnegie,
133. 2009. Fig. 97. Persoonia 23: 111. 2009.
Diagnosis: Leaf spots amphigenous, irregular to angular, pale Basionym: Cibiessia minutispora Crous & Carnegie, Fungal Di-
to medium brown, with a raised, dark brown border, 2 –5 mm versity 26: 153. 2007.
diam. Conidia solitary, medium brown, aseptate, smooth, Diagnosis: Cibiessia morph: conidia cylindrical with obtusely
granular, base truncate, with three apical, lateral, obtuse pro- rounded to subtruncate ends; aseptate conidia 4–6 × 2–3 μm,
jections, deltoid, thick-walled, with darker pigmentation in the 1(–2)-septate conidia 6–10 × 2–3 μm; conidia developing further,
lateral projections, but with more prominent constriction be- becoming medium brown, predominantly aseptate, verruculose,
tween the projections and the base, (7–)8–9( – 10) μm long, ellipsoidal to subglobose or globose, with dehiscence scars clearly
(7–)8( –9) μm wide at apex. Microconidia ellipsoidal with visible on conidial body. Readeriella synasexual morph not seen.
subtruncate bases and obtuse apices, hyaline, smooth,
3– 5 × 2–3.5 μm. Description and illustration: Crous et al. (2007c).
Description and illustration: Crous et al. (2009b). Typus: Australia, New South Wales, South Grafton, Grafton City
Council Landfill Plantation, S29°46’21”, E152°54’38”, on
Typus: Australia, New South Wales, Southern Tablelands, on C. henryi, 16 Feb. 2006, A.J. Carnegie (holotype CBS H-19839,
E. dives, Apr. 2006, coll. B.A. Summerell, isol. P.W. Crous isotype DAR 78030, cultures ex-type CBS 120894 = CPC
(holotype CBS H-20261, cultures ex-type CBS 124999 = CPC 13071, CPC 13072, 13073).
13026, CPC 13027, 13028).
Readeriella mirabiliaffinis Quaedvl. et al., Persoonia 33: 31.
Readeriella lehmannii Crous, Persoonia 37: 397. 2016. 2014.
Diagnosis: Conidia solitary, medium brown, aseptate, granular, Diagnosis: Conidia solitary, medium brown, aseptate, smooth,
verruculose, thick-walled, ellipsoid to clavate, apex obtuse, base granular, base truncate, with three apical, lateral, obtuse pro-
truncate, 1 μm diam, apex and base frequently slightly thickened, jections, deltoid, thick-walled, with darker pigmentation in the
(5–)6(–7) × (2.5–)3(–3.5) μm, lacking a sheath. lateral projections, but with more prominent constriction between
Description and illustration: Crous et al. (2016b). the projections and the base, (9–)10–11(–12) μm long, (8–)
9–10(–11) μm wide at apex.
National Park, Central lookout, on E. lehmannii, 23 Sep. 2015, Description and illustration: Quaedvlieg et al. (2014).
P.W. Crous (holotype CBS H-22916, culture ex-type CBS Typus: Australia, Tasmania, Tasman Peninsula, Brown Moun-
142131 = CPC 28935). tain walk, S43°11'13.9", E147°50'50.7", on E. delegatensis, 14
Readeriella limoniforma Quaedvl. et al., Persoonia 33: 31. Oct. 2006, P. Summerell & B. Summerell (holotype CBS H-
2014. 21134, culture ex-type CBS 134744 = CPC 13611).
Diagnosis: Conidia solitary, aseptate, ellipsoid to limoniform, Readeriella mirabilis Syd. & P. Syd., Annls Mycol. 6: 484. 1908.
tapering towards a bluntly rounded, subobtuse, thickened apex,
base subtruncate, hyaline becoming medium to golden brown, Diagnosis: Leaf spots amphigenous, irregular to subcircular,
finely verruculose, (6–)7–8(–10) × 3(–4) μm. medium brown, border raised, dark brown, 2–8 mm diam.
Conidia solitary, medium brown, aseptate, smooth, granular, base
Description and illustration: Quaedvlieg et al. (2014).
truncate, with three apical, lateral, obtuse projections, deltoid,
Typus: Australia, New South Wales, Wollemi National Park, on thick-walled, with darker pigmentation in the lateral projections,
Eucalyptus sp., 9 Feb. 2006, B. Summerell (holotype CBS H- (7–)9–10(–11) μm long, (7–)8–9(–10) μm wide at apex.
254
Description and illustration: Crous et al. (2009b). not observed, but epiphyllous remnants intermixed with those of
Paramycosphaerella marksii. Conidia holoblastic, solitary,
Typus: Australia, Victoria, on E. capitellata, Jun. 1907, F.M. aseptate, pale to medium brown, finely verruculose, base sub-
Reader (holotype in S); Victoria, on E. globulus, Sep. 2005, coll. truncate, apex flattened with three lateral, obtuse projections,
I.W. Smith, isol. P.W. Crous (epitype CBS H-20262, cultures ex- deltoid, whole conidia 3–5 μm long and wide.
epitype CBS 125000 = CPC 12370, CPC 12371, 12372).
Readeriella nontingens (Crous & Summerell) Crous & Sum-
merell, Persoonia 23: 111. 2009. Typus: New Zealand, North Island, Kerikeri, on E. botryoides,
Basionym: Cibiessia nontingens Crous & Summerell, Fungal 17 Oct. 2003, M.A. Dick (holotype CBS H-9892, culture ex-
Diversity 26: 154. 2007. type CBS 114357 = CMW 14913 = CPC 10895 = NZFRI-M
5098).
Diagnosis: Leaf spots amphigenous, irregular to subcircular;
spots variable from small specks (1 mm diam) to larger spots Readeriella patrickii Crous & Summerell, Persoonia 23: 111.
(7 mm diam), or coalescing to form larger blotches, medium 2009. Fig. 97.
brown, with a raised border and thin, red-purple margin. Diagnosis: Leaf spots amphigenous, subcircular to irregular, pale
Ascospores tri- to multiseriate, overlapping, hyaline, guttulate, to medium brown, with a raised, dark brown border, up to 5 mm
thin-walled, straight, fusoid-ellipsoidal with obtuse ends, diam. Conidia solitary, medium brown, aseptate, granular, finely
widest just above the septum, medianly 1-septate, constricted verruculose, thick-walled, ellipsoid to obclavate, widest below the
at the septum, tapering towards both ends, but more promi- obtuse apex, base subtruncate to truncate, 1 μm wide, with
nently towards the lower end, (9– )10 –11 × 2 –3( – 3.5) μm; inconspicuous marginal frill, (6–)7–8(–9) × (2.5–)3(–3.5) μm;
several ascospores showed remnants of a mucus sheath. covered in a persistent mucilaginous sheath.
Readeriella synasexual morph: Conidia narrowly ellipsoid
to subcylindrical with rounded ends, pale brown, smooth, Description and illustration: Crous et al. (2009b).
4 – 6 × 2 – 3 μm. Typus: Australia, Tasmania, Tasman Peninsula, S43°11'29.7",
Description and illustration: Crous et al. (2007c). E147°51'00.7", on E. amygdalina, 14 Oct. 2006, coll. P. Sum-
merell & B.A. Summerell, isol. P.W. Crous (holotype CBS H-
Typus: Australia, New South Wales, McWilliam Drive, Douglas 20248, cultures ex-type CBS 124987 = CPC 13602, CPC 13603,
Park S34°11’0", E150°43’0", on E. tereticornis, Jul. 2006, B. 13604).
Summerell (holotype CBS H-19840, cultures ex-type CBS
120725 = CPC 13217, CPC 13218, 13219). Readeriella pseudocallista Crous & Summerell, Persoonia 23:
133. 2009. Fig. 97.
Readeriella novaezelandiae Stud. Mycol. 50: 211. 2004.
Diagnosis: Leaf spots amphigenous, circular to irregular, medium
Diagnosis: Leaf spots irregular to subcircular, medium brown to brown, with a raised, brown border, and thin red-purple margin,
red-brown, margins raised, 2–15 mm diam. Intact pseudothecia up to 6 mm diam. Conidia solitary, medium brown, aseptate,
Fig. 96. Ramularia eucalypti (CBS H-19832). A, B. Leaf spots. C. Conidiophores. D–G. Conidia in chains. Scale bars: C = 10 μm, D = 6 μm (from Crous et al. 2007c).
CROUS ET AL.
Fig. 97. Readeriella spp. A, B. General disease symptoms (BRIP 29864). C, D. Asci of R. dendritica. E. Typical Readeriella colony on MEA. F, G. Vertical sections through
conidiomata. H, I. Conidiogenous cells and conidia of R. pseudocallista. J, K. Conidiogenous cells and conidia of R. eucalyptigena. L. Conidia of Cibiessia synasexual morph. M,
N. Conidia of Nothostrasseria synasexual morph of R. dendritica. O, P. Conidiogenous cells and conidia of R. eucalypti. Q, R. Conidiogenous cells and conidia of R. menaiensis.
S. Conidia of R. callista (CPC 13615). T. Conidia of R. patrickii (CPC 13602). Scale bars = 10 μm.
256
finely verruculose, ellipsoidal, tapering towards a subobtuse lamina, with pycnidia oozing black spore masses onto the leaf
apex and subtruncate or truncate base (1 μm wide), (5–) surface. Conidia consisting of an ellipsoid body with obtuse apex,
7–8 × (4.5–)5–5.5 μm. tapering to a tubular basal appendage; body medium brown,
verruculose, (7–)8–10(–11) × (2.5–)3–3.5(–4) μm; tubular
appendage separated from the conidium body by a septum,
Typus: Australia, New South Wales, Central Coast, S33°05’01", unbranched, hyaline, smooth, (5–)6–7(–8) × 1–1.5 μm.
E151°07’39", on E. prominula, Oct. 2006, coll. B.A. Summerell,
isol. P.W. Crous (holotype CBS H-20263, cultures ex-type CBS
125001 = CPC 13599, CPC 13600, 13601). Typus: Australia, Tasmania, Mount Wellington Park, S42°55'0",
E147°15'0", on E. delegatensis, 14 Oct. 2006, coll. B.A. Sum-
Readeriella readeriellophora Crous & Mansilla, Stud. Mycol.
merell, isol. P.W. Crous (holotype CBS H-20264, cultures ex-
50: 207. 2004.
type CBS 125002 = CPC 13631, CPC 13632).
Synonyms: Mycosphaerella readeriellophora Crous & Mansilla,
Stud. Mycol. 50: 207. 2004. Rehmiodothis Theiss. & Syd., Annls Mycol. 12: 192. 1914.
Teratosphaeria readeriellophora (Crous & Mansilla) Crous & U.
Ascomata superficial, black, thyrothecial with central ostiole. Asci
Braun, Stud. Mycol. 58: 11. 2007.
fusoid-ellipsoid, 8-spored, intermingled among paraphyses. As-
Diagnosis: Leaf spots amphigenous, subcircular, 4–6 mm diam, cospores hyaline, smooth-walled, unequally 1-septate, with
grey to medium brown, with a raised, red-brown border. Asco- smaller basal cell.
spores 2–3-seriate, overlapping, hyaline, guttulate, thick-walled,
Type species: Rehmiodothis osbeckiae (Berk. & Broome)
straight, obovoid with obtuse ends, unequally 1-septate, widest in
Theiss. & Syd.
the middle of the apical cell, not to slightly constricted at the
septum, tapering towards both ends, but more prominently to- Symptoms: Tar spot fungi, with immersed stroma beneath
wards the lower end, (11–)13–14(–16) × (3.5–)4–4.5 μm epidermis, having only an ostiole breaking though the surface.
in vivo. Conidia holoblastic, solitary, ellipsoidal to limoniform,
Notes: Species of Rehmiodothis are rare, and not well-known on
tapering towards a bluntly rounded, subobtuse, thickened apex,
Eucalyptus. Swart (1987) treated the two species cited below, but
base subtruncate, initially hyaline, becoming yellow- to green-
as these have not been recollected since, and are not known from
brown, and finally dark brown, aseptate, finely verruculose,
culture, they will not be further treated here. Species of Rehmio-
(5–)6–7(–9) × (3–)4(–4.5) μm; inconspicuous marginal frill
dothis are considered to be minor pathogens.
present.
Rehmiodothis eucalypti (Cooke & Massee) H.J. Swart, Trans.
Brit. Mycol. Soc. 89: 487. 1987.
Typus: Spain, Pontevedra, Lourizan, Areeiro, on E. globulus, Basionym: Phyllachora eucalypti Cooke & Massee, in Cooke,
2003, J.P. Mansilla (holotype CBS H-9901, culture ex-type CBS Grevillea 16(no. 77): 5. 1887.
114240 = CMW 14233 = CPC 10375). Synonym: Montagnella eucalypti (Cooke & Massee) Sacc., Syll.
fung. (Abellini) 9: 1048. 1891.
Readeriella tasmanica Crous & Summerell, Persoonia 23: 133.
2009. Diagnosis: Different from R. inaequalis in having smaller stro-
Diagnosis: Leaf spots predominantly hypophyllous, irregular to mata, smaller and more crowded ascocarps, and larger asco-
subcircular, medium brown, 1–4 mm diam, raised above the leaf spores, 25–39 × 8–13 μm.
Fig. 98. Rehmiodothis inaequalis (VPRI 15214). A. Disease symptoms. B. Vertical section through ascomata. C. Close up of leaf spot. D, E. Asci. F. Ascospores. Scale
bars = 10 μm.
CROUS ET AL.
Description and illustration: Swart (1987). Seiridium Nees, Syst. Pilze (Würzburg): 22. 1816 (1816–1817).
Typus: Australia, Victoria, Gippsland, Bunyip, on Eucalyptus Ascomata perithecial, immersed to semi-erumpent, depressed,
(stringy bark) litter, Mar. 1887, F. Campbell (holotype K(M) 49489). globose to pyriform, scattered or confluent; peridium dark brown,
Additional specimens in Swart (1987). Not known from culture. pseudoparenchymatous. Ostioles central, slightly papillate,
black, periphysate. Paraphyses hyaline, smooth, filiform. Asci
Rehmiodothis inaequalis (Cooke) H.J. Swart, Trans. Brit. cylindrical, 8-spored, unitunicate, thin-walled, stipitate, with an
Mycol. Soc. 89: 487. 1987. Fig. 98. apical amyloid ring. Ascospores cylindrical-oblong, euseptate,
Basionym: Dothidella inaequalis Cooke, Grevillea 20(no. 93): 5. septa often thicker than the wall, yellow- to dark brown, guttu-
1891. late. Conidiomata acervuloid to pycnidioid, semi-immersed to
Synonyms: Phyllachora inaequalis (Cooke) Cooke, Handb. erumpent, uni- to plurilocular, brown or black, glabrous,
Austral. fungi: 298. 1892. dehiscing by irregular splits in the upper wall. Conidiophores
Placostroma inaequale (Cooke) Theiss. & Syd., Annls Mycol. lining the cavity of the conidioma, septate and sparsely
13(3/4): 407. 1915. branched at the base, or reduced to conidiogenous cells,
Diagnosis: A typical "tar spot" fungus, with flattened dome- hyaline, smooth. Conidiogenous cells discrete, integrated,
shaped glossy black stromata, and with ascocarp ostioles ampulliform to lageniform or subcylindrical, hyaline, smooth,
emerging through the leaf surface as slightly protruding gelati- proliferating percurrently at the apex. Conidia fusiform, dis-
nous structures. Ascostromata with sparse ostioles emerging toseptate (septal pores present or not), end cells hyaline, me-
through leaf surface as slightly protruding gelatinous structures; dian cells dark brown to brown, wall thick, smooth or with
ascospores 2-celled, hyaline, with basal cell less than half the striations, constricted at septa or not; apical cell with a single,
length of the apical cell, 22–29 × 6–7 μm. cellular, unbranched or branched, appendage; basal cell with or
without a centric, unbranched or sometimes branched
Description and illustration: Swart (1987). appendage.
Typus: Australia, Victoria, on Eucalyptus litter, collection date Type species: Seiridium marginatum Nees
unknown, J.M. Martin 824 (holotype K(M) 49520). Additional
specimens in Swart (1987). Not known from culture. Symptoms: Small, brown, irregular, amphigenous leaf spots.
Fig. 99. Seiridium kartense (CBS 142629). A. Sporodochia on SNA immersed in agar. B. Sporodochium on SNA erumpent from agar. C. Conidia. D, E. Conidiophores. F.
Polyblastic conidiogenesis. G. Conidiogenous cell proliferating percurrently showing multiple collarettes. Scale bars: A, B = 100 μm; C–G = 10 μm (from Bonthond et al. 2018).
258
Note: The genus Seiridium appears to be of minor importance as transversely distoseptate, not constricted, verruculose, apex
a foliar pathogen of Eucalyptus. obtuse, base truncate with marginal frill.
Seiridium eucalypti Nag Raj, Coelomycetous Anamorphs with Type species: Sonderhenia eucalyptorum (Hansf.) H.J. Swart &
Appendage-bearing Conidia (Ontario): 862. 1993. J. Walker
Diagnosis: Conidia lunate to falcate, straight to curved, 5-septate, Notes: Sonderhenia has pycnidial conidiomata, distoseptate
not striate, bearing two appendages, euseptate with visible pores, conidia with a central pore, brown, percurrently proliferating
(23.5–)27.5–33(–36) × (8–)9–10.5(–11) μm, av. ± SD = 30.3 ± conidiogenous cells, and a teratosphaeria-like sexual morph.
2.7 × 9.6 ± 0.7 μm (n = 32); basal cell obconic with a truncate Species of Sonderhenia are morphologically distinguished based
base, hyaline, walls smooth, 4–6 μm; four median cells, smooth, on their conidial morphology (shape, taper, and dimensions).
cylindrical to doliiform, brown to dark brown, and septa darker than Although Sonderhenia is quite common on eucalypts in Australia,
the rest of the cells (second cell from base 4.5–8 μm long; third New Zealand, South America and Europe, it appears to be of
cell 4–6.5 μm long; fourth cell 3.5–6 μm long; fifth cell 4.5–7 μm minor importance. Some species (e.g. S. eucalytorum) can oc-
long); apical cell conical, hyaline, smooth, 2.5–6 μm long; apical casionally be severe on sapplings in regenerating forests in
appendage single, centric, 8.5–15.5 μm long; basal appendage, Australia, for example on E. sieberi and E. fraxinoides.
single, cylindrical, centric and excentric, 6.5–10 μm long.
Sonderhenia eucalypticola (A.R. Davis) H.J. Swart & J.
Descriptions and illustrations: Nag Raj (1993), Bonthond et al.
Walker, Trans. Brit. Mycol. Soc. 90: 640. 1988. Fig. 100.
(2018).
Basionym: Hendersonia eucalypticola A.R. Davis, Pomona Coll.
Typus: Australia, South Australia, Adelaide, Mt. Lofty Summit, J. econ. Bot. 2: 251. 1912.
on Eucalyptus sp., 16 Oct. 1979, B. Kendrick (holotype DAOM Synonyms: Hendersonia fraserae Hansf., Proc. Linn. Soc.
215255); Tasmania, on E. delegatensis, 13 Sep. 1996, Z.Q. Yuan N.S.W. 79: 135. 1954.
(epitype CBS H-23145, culture ex-epitype CBS 343.97 = ECF Mycosphaerella walkeri R.F. Park & Keane, Trans. Brit. Mycol.
149). Soc. 83: 104. 1984.
Seiridium kartense Bonthond et al., Persoonia 40: 109. 2017 Diagnosis: Leaf spots amphigenous, round to confluent and
(2018). Fig. 99. irregular, 0.5–2 mm diam, grey, margin purple when young,
becoming dark red to brown and raised with age. Ascospores
Diagnosis: Conidia lunate to falcate, curved, 5-septate, not stri- 2–3-seriate, colourless, guttulate, straight or slightly curved,
ate, bearing two appendages, euseptate with visible pores, fusiform, 1-septate, widest just above median septum, slightly
(23.5–)27.5–30.5(–32) × (6.5–)7.5–8.5(–9) μm, av. ± constricted at septum, 19–27 × 4–6 μm. Conidia ellipsoid to
SD = 28.5 ± 1.8 × 8.0 ± 0.4 μm (n = 38); basal cell obconic with a ovoid, brown, 3-distoseptate, not constricted at septa, verrucu-
truncate base, hyaline, walls smooth, 4–5.5 μm; four median lose, apex obtuse, base truncate with marginal frill,
cells, smooth, cylindrical to doliiform, dark brown (second cell 19–31 × 6–12 μm. Swart & Walker (1988) treated Hendersonia
from base 5.5–6.5 μm long; third cell 5–5.5 μm long; fourth cell fraserae (conidia 23–28 × 6–9 μm; Hansford 1954) as synonym
4.5–5 μm long; fifth cell 4.5–5.5 μm long); apical cell conical, of Hendersonia eucalypticola.
hyaline, smooth, 3–4 μm long; appendages cylindrical, attenu-
Descriptions and illustrations: Swart & Walker (1988), Park &
ated, unbranded, slightly spathulate; apical appendage single,
Keane (1984), Crous (1998).
centric, 6.5–8.5 μm long; basal appendage consistently present,
single, cylindrical, excentric, 6–8.5 μm long.
Typus: Australia, Victoria, Skipton, on E. globulus, 8 Dec. 1982,
Description and illustration: Bonthond et al. (2018). R.F. Park, (holotype of M. walkeri DAR 45724, isotype of
M. walkeri IMI 280473). USA, California, Claremont, on
Typus: Australia, South Australia, Kangaroo Island, on
E. globulus, 20 Dec. 1911, A.R. Davis, POM 249135, F. columb.
E. cladocalyx, 15 Dec. 2012, W. Quaedvlieg (holotype CBS H-
3813 (lectotype of H. eucalypticola, slide as DAR 55638;
duplicate of F. columb. 3813 also as DAR 55846).
Sonderhenia H.J. Swart & J. Walker, Trans. Brit. Mycol. Soc. 90:
Sonderhenia eucalyptorum (Hansf.) H.J. Swart & J. Walker,
640. 1988. Fig. 100.
Trans. Brit. Mycol. Soc. 90: 640. 1988. Fig. 100.
Ascomata pseudothecial, amphigenous, on one side of each Basionym: Hendersonia eucalyptorum Hansf., Proc. Linn. Soc.
lesion, often 1–3, intermingled with pycnidia, immersed, black, N. S. W. 79: 135. 1954.
punctiform, globose to subglobose; apical ostiole substomatal; Synonym: Mycosphaerella swartii R.F. Park & Keane, Trans. Brit.
wall olive-brown, of 3–4 layers of textura angularis. Asci Mycol. Soc. 83: 99. 1984.
fasciculate, bitunicate, subsessile, 8-spored, ovoid to obclavate,
Diagnosis: Leaf spots amphigenous, round to confluent and
straight to incurved. Ascospores 2–3-seriate, hyaline, guttulate,
irregular, 0.5–2 mm diam, grey, surrounded by a purple margin
straight or slightly curved, fusiform, 1-septate. Conidiomata
when young, which becomes dark red to brown and raised with
pycnidial, amphigenous, subepidermal with central non-
age. Ascospores 2–3-seriate, colourless, guttulate, straight or
projecting ostiole, scattered, black, globose; wall of 2–3 layers
slightly curved, fusiform, 1-septate, widest just above median
of brown textura angularis. Paraphyses present or absent, hy-
septum, slightly constricted at septum, 20–27 × 4–6 μm. Conidia
aline, septate, hypha-like, intermixed between conidiogenous
ellipsoid to cylindrical, straight or bent, brown, 3-distoseptate, not
cells. Conidiogenous cells minute, olivaceous, proliferating
constricted, verruculose, apex obtuse, base truncate with mar-
enteroblastically and percurrently, lining the pycnidium. Conidia
ginal frill, 25–49 × 5–10 μm (conidia originally cited as
ellipsoid to cylindrical or ovoid, straight or bent, brown,
40–48 × 5–6 μm; Hansford 1954).
CROUS ET AL.
Fig. 100. Sonderhenia spp. A–D. Disease symptoms. E–H. S. eucalypticola. E. Vertical section through conidioma. F, G. Conidiogenous cells giving rise to conidia. H. Conidia.
I–K. S. eucalyptorum. I. Vertical section through conidioma. J. Conidiogenous cells giving rise to conidia. K. Conidia. L–O. S. radiata (CBS 145600). L–N. Conidiogenous cells.
O. Conidia. Scale bars = 10 μm.
Descriptions and illustrations: Swart (1988), Park & Keane Leaf spots circular to subcircular, 1–5 mm diam, amphigenous,
(1984), Crous (1998), Summerell et al. (2006). brown with red margin. Conidiomata separate, pycnidial, brown,
180–250 μm diam, with central ostiole; wall of 6–8 layers of brown
Typus: Australia, Mt. Gambier, on E. leucoxylon, 9 Dec. 1982,
textura angularis. Conidiophores reduced to conidiogenous cells
R.F. Park (holotype of M. swartii DAR 45719, isotype of
lining the inner cavity, hyaline to pale brown, smooth to verruculose,
M. swartii IMI 280474); South Australia, Clare Valley, on
5–10 × 4–7(–10) μm, proliferating percurrently at apex. Paraph-
E. leucoxylon, Aug. 1922, T. Osborn (holotype of
yses hyaline, septate, hypha-like, 2–3 μm diam, up to 80 μm long,
H. eucalyptorum K(M) 137253).
intermixed between conidiogenous cells. Conidia solitary, (25–)
Sonderhenia radiata Crous, sp. nov. MycoBank MB832050. 27–30(–34) × (6–)6.5–7(–8) μm, medium brown, verruculose,
Fig. 100. straight to slightly curved, 3-distoseptate, with central pore in
septum, fusoid-ellipsoid to subcylindrical, apex subobtusely
Etymology: Name refers to Eucalyptus radiata, the host species
tapered, base truncate, 2–3 μm diam, with minute marginal frill.
from which this fungus was isolated.
260
Culture characteristics: Colonies erumpent, spreading with fol- Conidiophores reduced to conidiogenous cells or 1 –2 sup-
ded surface and moderate aerial mycelium, smooth, lobate porting cells, hyaline, subcylindrical, lining the inner layer of
margin, reaching 20 mm diam after 2 wk at 24 °C. On MEA conidioma. Conidiogenous cells hyaline, smooth, sub-
surface dirty white, reverse umber; on PDA surface dirty white, cylindrical to ampulliform; proliferating sympodially but also
reverse pale luteous, on OA surface dirty white. percurrently near apex. Conidia hyaline, smooth, guttulate,
filiform, apex subobtuse, base long obconically truncate,
Typus: Australia, Victoria, Cathedral Ranges State Park, start of
transversely euseptate.
Jawbone track, S37°24’, E145°45’, on E. radiata, 4 Feb. 1986,
I.G. Pascoe (holotype CBS H-24042, culture ex-type CBS Type species: Sphaerulina myriadea (DC.) Sacc.
145600 = CPC 18218 = VPRI 13393a).
Notes: In the past, Sphaerulina was separated from Mycos-
Additional material examined: Australia, New South Wales, on leaves of phaerella on the basis of ascospore septation (Crous et al.
E. piperita, 26 Nov. 2016, P.W. Crous, HPC 1747, culture CPC 32134. 2003). Sphaerulina was shown to have septoria-like asexual
Notes: Phylogenetically, this species is distinct from other Son- morphs (Quaedvlieg et al. 2013, Verkley et al. 2013).
derhenia species currently known from sequence data, being
Sphaerulina cercidis (Fr.) Quaedvl., G.J.M. Verkley & Crous,
closest related to Sonderhenia eucalypticola (Fig. 101). Some Stud. Mycol. 75: 345. 2013. Fig. 102.
intraspecific variation exists for this species, but more isolates
Basionym: Septoria cercidis Fr., in Leveille, Annls Sci. Nat., Bot.,
are needed to determine whether this is indicative of two species.
ser. 3 9: 251. 1848.
Morphologically, it is a rather distinct species of Sonderhenia, as
Synonym: Septoria provencialis Crous, Stud. Mycol. 55: 127.
conidiomatal paraphyses were observed in culture, but are ab-
2006.
sent on S. eucalypticola and S. eucalyptorum.
Diagnosis: Leaf spots amphigenous, dark brown, angular,
Sphaerulina Sacc., Michelia 1(no. 4): 399. 1878.
confined by leaf veins, 1–6 mm diam, becoming confluent with
Ascomata pseudothecial, immersed, subepidermal, erumpent age. Conidia solitary in vivo, finely guttulate, subcylindrical to
single to clustered, globose, papillate. Ostiole central, with narrowly obclavate, with subobtuse apex, and obconically sub-
hyaline periphyses; wall of textura angularis, composed of 2 – 4 truncate base, variously curved to irregular, mostly widest in the
layers of brown cells. Hamathecium dissolving at maturity. Asci middle of the basal cell, tapering towards the apex, (1–)2(–3)-
bitunicate, fissitunicate, clustered, cylindrical to obclavate, septate, (12–)30–40(–45) × 2.5–3(–4) μm.
rounded at apex, with or without a shallow apical chamber,
short-stipitate or sessile, with 8 bi- to triseriate ascospores. Descriptions and illustrations: Quaedvlieg et al. (2013), Verkley
Ascospores subcylindrical to fusiform, rounded at ends, slightly et al. (2013).
tapered, straight or slightly curved, 1 – 3-septate, with a primary Typus: France, Provence, Cheval Blanc camping site, on
septum nearly median, hyaline, smooth, without sheath or
Eucalyptus sp., 29 Jul. 2005, P.W. Crous (holotype of
appendages. Conidiomata pycnidial, separate but frequently
S. provencialis CBS H-19701, cultures ex-type CBS
aggregated, with or without brown stromatic tissue in a stroma;
118910 = CPC 12226, CPC 12227, 12228).
globose, black, exuding a creamy conidial mass via a central
ostiole; wall of 3 – 12 layers of dark brown, textura angularis. Staninwardia B. Sutton, Trans. Brit. Mycol. Soc. 57: 540. 1971.
Fig. 101. The first of 11 equally most parsimonious trees obtained from a phylogenetic analysis of the Sonderhenia combined ITS, act, cmdA and tub2 sequence alignment (15
strains including the outgroup; 1 686 characters analysed: 1 254 constant, 325 variable and parsimony-uninformative and 107 parsimony-informative). The tree was rooted to
Cladosporium allicinum (GenBank DQ780399.1, EF679489.1, EF679565.1, EF101451.1, respectively) and the scale bar indicates the number of changes. Thickened branches
represent those present in the strict consensus parsimony tree. Bootstrap support values higher than 49 % are shown at the nodes (parsimony bootstrap support / NJ HKY85
distance bootstrap support) and novelties are highlighted with bold text. Species names are indicated to the right of the tree. A superscript T denotes strains with a type status.
Tree statistics: TL = 508 steps, CI = 0.951, RI = 0.872, RC = 0.830. The analyses were performed using PAUP v. 4.0b10 (Swofford 2003).
CROUS ET AL.
Fig. 102. Sphaerulina cercidis (CBS 118910). A. Leaf spot. B. Section through conidioma. C. Conidiogenous cells. D. Conidia. E. Colony on OA. Scale bars = 10 μm.
Conidiomata amphigenous, black, erumpent acervuli; basal Diagnosis: Leaf spots amphigenous, dark brown to black, circular,
stroma consisting of dense, aggregated hyaline cells giving rise 2–5 mm diam, rarely extending through the leaf lamina, margins
to conidiophores. Conidiophores hyaline, cylindrical, densely diffuse, slightly chlorotic. Conidia aggregated in mucus, fusoid-
aggregated, covered in mucus, 0–2-septate, unbranched or ellipsoidal with rounded apex and truncate base, brown, verru-
branched below. Conidiogenous cells integrated, terminal, hya- culose, predominantly 1-septate, but each cell can develop an
line, smooth, covered in mucus, proliferating several times per- additional septum with age (though the first is observed in the
currently near apex. Conidia aggregated in mucus, fusoid- apical cell), constricted at septa, (8–)10–14(–17) × (3–)
ellipsoidal with rounded apex and truncate base, brown, verru- 4(–5) μm, basal cell shorter and narrower (4–)5–7 × 3–4 μm
culose, predominantly 1-septate, but each cell can develop an than the apical cell (4–)5–8 × (3–)4–5 μm; conidia remain
additional septum with age, constricted at septa; mucilaginous covered in a mucilaginous sheath (up to 4 μm wide), and also form
sheath persistent. short conidial chains. In culture conidial chains are more promi-
nent, and conidia can become up to 20 μm long and 7 μm wide.
Type species: Staninwardia breviuscula B. Sutton
Symptoms: Leaf spots amphigenous, dark brown to black, cir-
cular, 2–5 mm diam, rarely extending through the leaf lamina, Typus: Australia, New South Wales, 1.4 km along Lakes Way
margins diffuse, slightly chlorotic. from intersection with Green Point Drive, then ca. 500 m NW
from Lakes Way along dirt access road, ca. 10 km S of Forster,
Notes: The genus Staninwardia is known from two species,
Booti Booti National Park, North Coast NSW, S32°15’45”,
namely S. suttonii from Australia, and S. breviuscula reported
E152°31’33”, alt. 5 m, on leaves of E. robusta, Mar. 2006, B.
from India, Mauritius and Sri Lanka (Sutton 1980). Staninwardia
Summerell (holotype CBS H-19750, culture ex-type CBS
is characterised by having acervuli on corky brown leaf spots,
120061 = CPC 13055, CPC 13056, 13057).
with brown, catenulate conidia covered in a mucilaginous sheath.
Stigmina Sacc., Michelia 2(no. 6): 22. 1880.
Staninwardia breviuscula B. Sutton, Trans. Brit. Mycol. Soc.
57: 541. 1971. = Pseudocercospora Speg. (see above)
Diagnosis: Causing small circular lesions on leaves, and lentic- Type species: Stigmina platani (Fuckel) Sacc.
ular to elongated, often confluent, lesions (up to 10 mm long) on Symptoms: Leaf spots amphigenous, circular to subcircular, pale
the stems and petioles of an unidentified eucalypt species in to medium brown, becoming red-purple for some taxa, with or
Mauritius. Numerous epiphyllous acervuli located in epidermis; without raised border.
conidia pale brown, thick-walled, verruculose, 2-celled and often
constricted at the median septum, doliiform to clavate, Notes: The genus Stigmina (based on S. platani) is a synonym of
7–10.5 × 3.5–5 μm, formed in chains of up to 3, intercalary Pseudocercospora (Braun & Crous 2006, Crous et al. 2006b,
conidia truncate at both ends with the remains of a basal frill, 2013). However, previously several cercosporoid taxa were
apical conidia truncate only at the base, each conidial chain indiscriminently placed in Stigmina, because they had rough,
enveloped in a mucilaginous sheath about 2 μm thick which brown, percurrently proliferating conidiogenous cells, and disto-
persists on mature detached conidia. or muriformly septate conidia. These taxa are clearly not
congeneric with Pseudocercospora s. str., but in the absence of
Description and illustration: Sutton (1971b). molecular data, it is also not possible to determine their generic
Typus: Mauritius, Nouvelle Decouverte, on Eucalyptus sp., Apr. affinities. For the present, they are retained in “Stigmina”.
1969, G.M. Lallmahomed (holotype IMI 139473). Not known “Stigmina” eucalypticola B. Sutton & Pascoe, Mycol. Res. 92:
from culture. 215. 1989.
Staninwardia suttonii Crous & Summerell, Fungal Diversity 23: Synonym: Peltosoma eucalypti Hansf., Proc. Linn. Soc. N.S.W.
343. 2006. Fig. 103. 81: 44. 1956.
262
Fig. 103. Staninwardia suttonii (CBS H-19750). A, B. Leaf spots with conidiomata. C–G. Conidiophores giving rise to pigmented, septate conidia. H. Colony on MEA. I, J.
Conidiogenous cells giving rise to conidia in vitro (note persistent mucilaginous sheath). Scale bars: A, B = 400 μm, C = 10 μm, H = 4 mm, I = 10 μm (from Summerell et al.
2006).
CROUS ET AL.
Diagnosis: Leaf spots amphigenous, subcircular, 1–7 mm diam, Suberoteratosphaeria pseudosuberosa (Crous & M.J. Wingf.)
pale to medium brown, becoming red-purple; border indistinct, Quaedvlieg & Crous, Persoonia 33: 31. 2014. Fig. 104.
coloured margin absent. Conidia holoblastic, verruculose, medium Basionym: Mycosphaerella pseudosuberosa Crous & M.J.
brown, broadly ellipsoid, apex obtuse, base truncate with marginal Wingf., Stud. Mycol. 55: 118. 2006.
frill, transversely (1–)3-distoseptate, and occasionally with a lon- Synonym: Teratosphaeria pseudosuberosa (Crous & M.J.
gitudinal distoseptum, (13–)15–18(–20) × (6–)7–8 μm. Wingf.) Crous & U. Braun, Stud. Mycol. 58: 11. 2007.
Description and illustration: Sutton & Pascoe (1989a). Diagnosis: Leaf spots amphigenous, associated with brown,
Typus: Australia, South Australia, Pinnaroo, on E. oleosa, Sep. corky spots on leaf petioles. Ascospores tri- to multiseriate,
1924, G. Samuel, WARI 384 (holotype IMI 57990). Not known overlapping, hyaline, guttulate, thick-walled, straight to slightly
from culture. curved, fusoid-ellipsoidal with obtuse ends, medianly 1-septate,
widest at the middle of the apical cell, constricted at the septum,
“Stigmina” eucalyptorum Crous & T. Coutinho, Mycol. Mem. tapering towards both ends, but more prominently towards the
21: 144. 1998. lower end, (11–)12–14(–15) × (3–)3.5(–4) μm in vivo;
Diagnosis: Leaf spots amphigenous, circular, separate, 1–3 mm frequently surrounded by an irregular mucous sheath.
diam, on young and mature foliage, pale to medium brown, Description and illustration: Crous et al. (2006d).
becoming dark brown at the raised border, with a diffuse, reddish
margin. Conidia holoblastic, apical, solitary, brown, (8–) Typus: Uruguay, on leaves and petioles of Eucalyptus sp., Apr.
9–11(–15) × (3.5–)4–5(–6) μm, roughly verruculose with lon- 2005, M.J. Wingfield (holotype CBS H-19695, culture ex-type
gitudinal striations, transversely 0–2-distoseptate, ellipsoidal to CBS 118911 = CPC 12085).
fusiform, apex obtuse, base subtruncate with a marginal frill. Suberoteratosphaeria suberosa (Crous et al.) Quaedvlieg &
Description and illustration: Crous (1998). Crous, Persoonia 33: 31. 2014. Fig. 104.
Basionym: Mycosphaerella suberosa Crous et al., Mycologia 85:
Typus: Tanzania, on E. maidenii, Jun. 1995, T. Coutinho (ho- 707. 1993.
lotype PREM 54414). Not known from culture. Synonym: Teratosphaeria suberosa (Crous et al.) Crous & U.
“Stigmina” hansfordii B. Sutton & Pascoe, Mycol. Res. 92: Braun, Stud. Mycol. 58: 11. 2007.
216. 1989. Diagnosis: Leaf spots amphigenous, 0.5–1.5 cm diam, medium
Diagnosis: Leaf spots amphigenous, circular to irregular, to dark brown, corky with irregular margins, frequently sur-
2–15 mm diam, discrete, becoming confluent, pale brown, with a rounded by a red-purple border on the upper, smoother lesion
purple margin. Conidia holoblastic, lateral, verruculose, medium surface. Ascospores 2–3-seriate or irregularly arranged, oblique,
brown, medianly 1-distoseptate, occasionally with one oblique overlapping, straight ellipsoidal, obtuse at each end, hyaline to
distoseptum, ellipsoid to fusoid, apex obtuse to subobtuse, base pale brown, smooth, 1-septate, guttulate, slightly constricted at
truncate with a marginal frill, (10–)12–15(–17.5) × (5–) the median septum, widest in middle of upper cell, tapering more
6–7(–8) μm; conidia produced in a pale brown gelatinous matrix. toward one end, (10–)12–16(–17) × (3–)3.5–5(–6) μm.
Description and illustration: Sutton & Pascoe (1989a). Description and illustration: Crous (1998).
Typus: Australia, South Australia, Ooldea, on E. pyriformis, Aug. Typus: Brazil, Espírito Santo, Santa Catarina, on E. dunnii, Aug.
1922, G. Samuel (holotype ADW 2011, isotype IMI 320192). 1992, M.J. Wingfield (holotype PREM 51082, culture ex-type
Not known from culture. CBS 436.92 = CMW5226 = CPC 515).
Suberoteratosphaeria Quaedvl. & Crous, Persoonia 33: 31. Suberoteratosphaeria xenosuberosa Quaedvlieg et al., Per-
2014. Fig. 104. soonia 33: 31. 2014.
Foliicolous and caulicolous, plant pathogenic. Ascomata pseu- Diagnosis: Leaf spots similar to those reported for S. suberosa,
dothecial, solitary or aggregated, black, superficial to subepi- corky, and erumpent. Ascospores bi- to triseriate, hyaline, ellip-
dermal, globose, glabrous; ostiole apical, central, papillate, lined soidal with rounded ends, medianly 1-septate, constricted at the
with periphyses; wall of 3–4 layers of medium brown textura septum, straight to slightly curved, guttulate, thick-walled, widest
angularis, subhymenium of 3–5 layers of hyaline cells. Asci in middle of apical cell, (10–)11–13(–15) × (4–)4.5(–5) μm;
fasciculate, bitunicate, aparaphysate, subsessile, 8-spored, ascospores brown and verruculose at germination.
ellipsoid to obclavate, straight or curved. Ascospores bi- to tri-
seriate or irregularly arranged, oblique, overlapping, straight Description and illustration: Quaedvlieg et al. (2014).
ellipsoidal, obtuse at each end, hyaline to pale brown, smooth, 1- Typus: Australia, Queensland, Coolabunia Plantation, Kingaroy,
septate, guttulate, with or without mucoid sheath. Germinating on E. moluccana, 14 Feb. 2004, A.J. Carnegie (holotype CBS
ascospores become brown and verruculose. H-21138, culture ex-type CBS 134747 = CPC 13093 = NSWF
Type species: Suberoteratosphaeria suberosa (Crous et al.) 005175).
Quaedvlieg & Crous Teratosphaeria Syd. & P. Syd., Annls Mycol. 10: 39. 1912. Figs
Notes: Suberoteratosphaeria forms corky spots on leaves and 105–107.
petioles. It is primarily distinguished based on its characteristic
leaf spots, and secondly by its germinating ascospores that Ascomata separate, immersed; at times linked in densely
become brown, verruculose, and germinate by two or multiple branched stromatic tissue, forming fibrils, subglobose to globose,
germ tubes. black, uniloculate, thick-walled, with well developed ostiolar
264
Fig. 104. Suberoteratosphaeria spp. A–D. Disease symptoms. E, F. S. suberosa (PREM 51082). E. Vertical section through pseudothecium. F. Germinating ascospores. G–K.
S. pseudosuberosa (CBS H-19695). G. Asci with ascospores. H. Germinating ascospores. I. Colony on PDA. J, K. Conidia. Scale bars = 10 μm.
periphyses, somewhat pseudoparaphysoidal in immature asco- ellipsoid to flexuous, obclavate, smooth to verrucose, thin- to
mata. Peridium thick-walled, comprised of 2–3 layers of brown thick-walled, brown, mostly with basal dehiscence scars.
textura angularis; inner cells of flattened, hyaline cells. Hama-
Type species: Teratosphaeria fibrillosa Syd. & P. Syd.
thecium mostly absent, when present consisting of subcylindrical,
branched, septate pseudoparaphyses, or reduced to hyaline Symptoms: Associated with discrete leaf spots that could
periphysoids lining the ostiole. Asci 8-spored, bitunicate, sessile, develop into blights, and defoliation. Symptoms mostly restricted
clavate, apex rounded or tapered, short pedicellate with apical to leaves, but can also occur on petioles and twigs, and some
chamber. Ascospores overlapping, fusoid to cylindrical, 1-septate, species can cause cankers.
slightly constricted at the septum or not, curved to straight,
Notes: Some species of Teratosphaeria cause distinctive dis-
rounded at both ends, upper cell slightly obtuse, thick-walled,
ease symptoms. Teratosphaeria cryptica commonly infects
hyaline, at times becoming brown and somewhat verruculose,
young expanding leaves, often causing the leaf lamina to
at times with non-persistent mucoid sheath. Asexual morph
become crinkled and contorted, being commonly referred to as
hyphomycetous or coelomycetous. Hyphae dark brown, septate,
“crinkle leaf disease” (Marks et al. 1982, Park & Keane 1982b).
branched, mostly immersed. Conidiomata acervular to sporodo-
Teratosphaeria nubilosa forms characteristic leaf blotches (or
chial or pycnidial, solitary, brown, with pseudoparenchymatal wall
blight). Leaf blotches are a result of several individual lesions
of textura angularis. Conidiogenous cells integrated, terminal or
coalescing to form larger spreading blotches over the leaf
reduced to conidiogenous cells, proliferating percurrently and
surface. Species of Teratosphaeria with Kirramyces asexual
sympodially. Conidia aseptate to septate, solitary or in chains,
morphs (T. destructans, T. epicoccoides, T. eucalypti, T.
CROUS ET AL.
Fig. 105. Disease symptoms of Teratosphaeria spp. A. T. cryptica on E. globulus. B. T. verrucosa spots on Eucalyptus sp. C. T. eucalypti on Eucalyptus sp. D. T. nubilosa on
E. globulus. E. T. pseudoeucalypti on Eucalyptus sp. F. T. fimbriata on Eucalyptus sp. G. T. destructans on Eucalyptus sp. H. T. epicoccoides on Eucalyptus sp.
novaehollandiae, T. pseudoeucalypti, T. tiwiana and T. viscida) Diagnosis: Leaf spots amphigenous, irregular, 3–6 mm diam,
are associated with severe blight diseases on of Eucalyptus grey, with a thin, raised, brown border. Conidia solitary, pale
leaves, buds and shoots (Andjic et al. 2007a,b,c, 2010a,b, brown to brown, aseptate, guttulate, verruculose, ellipsoidal to
2019). Symptoms are similar and include chlorotic spots that subcylindrical, apex obtuse, base truncate or bluntly rounded,
become brown to purple with diffuse borders, and red-brown (4–)5–6(–7) × (2–)2.5–3 μm.
margins, or necrotic lesions delimited by veins, and the pres-
ence of black conidial masses on the leaf surface (Andjic et al.
2016). Teratosphaeria epicoccoides has a host range of more Typus: Australia, Northern Territory, ENE Pine Creek,
than 200 eucalypt species and is found in all areas where S13°40’49", E131°59’04.9", on E. miniata, 23 Sep. 2007, coll.
eucalypts are grown, but rarely causing significant disease B.A. Summerell, isol. P.W. Crous (holotype CBS H-20266,
(Andjic et al. 2019). cultures ex-type CBS 125004 = CPC 14598, CPC 14597,
14599).
The ability of some species in the Teratosphaeria complex to
infect several different hosts (“pogo-stick” hypothesis; Crous & Teratosphaeria alcornii Crous, Persoonia 23: 114. 2009.
Groenewald 2005), does not seem to apply to Teratosphaeria Synonyms: Stigmina eucalypti Alcorn, Trans. Brit. Mycol. Soc.
s. str., where species appear to be quite host specific, at least at 60: 151. 1973.
generic level. Several of the species discussed by Hunter et al. Batcheloromyces eucalypti (Alcorn) Crous & U. Braun, Stud.
(2011) have since been reallocated to other genera. The most Mycol. 58: 12. 2007.
damaging eucalypt foliar pathogens appear to be species of
Diagnosis: Conidiomata sporodochial, amphigenous, emerging
Teratosphaeria, including T. cryptica, T. nubilosa, T. destructans,
through stomata, up to 150 μm diam. Conidia medium to dark
T. pseudoeucalypti and T. viscida (Wingfield et al. 1996, Crous
brown, verrucose, broadly ellipsoid to subcylindrical, apex
1998, Andjic et al. 2007a,b,c, 2010a, 2016, Hunter et al. 2009,
obtuse, base truncate, (1–)3-distoseptate, (20–)
2011).
25–31(–36) × 7–10(–12) μm, frequently constricted at septa;
Teratosphaeria species can be spread to new areas to initiate
marginal frill present.
new epidemics in native and exotic environments (Hunter et al.
2011). This appears to happen primarily via infected seed or Descriptions and illustrations: Alcorn (1973), Crous (1998),
asymptomatic nursery stock (Maxwell et al. 2003, Jackson et al. Crous et al. (2007a).
2008, Andjic et al. 2011).
Typus: Australia, Queensland, Brisbane, on E. tessellaris, 19
Teratosphaeria alboconidia Crous & Summerell, Persoonia 23: Nov. 1969, A. Skoien (holotype of Stigmina eucalypti IMI
136. 2009. 161747, culture ex-type CBS 313.76 = CPC 3632 = IMI 161747).
266
Fig. 106. Asexual structures of Teratosphaeria spp. A. Immersed conidiomata with conidial cirrhi of T. epicoccoides. B. Conidia masses of T. corymbiae. C. Vertical section
through conidioma of T. epicoccoides. D. Conidiogenous cells of T. viscida. E, F. Conidiogenous cells of T. epicoccoides. G. Conidia of T. australiensis. H. Conidia of
T. corymbiae. I. Conidia of T. viscida. J. Conidia of T. epicoccoides. Scale bars = 10 μm.
Teratosphaeria angophorae (Andjic et al.) Andjic et al., Per- gradually tapering toward apex, truncate to subtruncate at base
soonia 23: 114. 2009. with marginal frill, (9.5–)11–14(–16) × (2–)2.5–3.5(–4)
Basionym: Kirramyces angophorae Andjic et al., Mycol. Res. (av. = 12.5 × 3 μm).
111: 1193. 2007.
Description and illustration: Andjic et al. (2010b).
Diagnosis: Leaf spots amphigenous, circular to irregular, Typus: Australia, Queensland, Rosedale, on E. grandis, 2007,
2–8 mm diam, single to confluent, red–brown with prominent G. Whyte (holotype MURU440, culture ex-type CBS
purple border. Conidia single, aseptate to 1–3 euseptate, sub- 125243 = MUCC668).
hyaline to pale brown, verruculose, fusoid, subcylindrical to
ellipsoidal, straight to slightly curved; base truncate with a mar- Teratosphaeria australiensis (B. Sutton) Crous, Persoonia 23:
ginal frill, apex sub-obtuse to obtuse, (4.5–)9–15(–19) × (1.5–) 114. 2009. Fig. 106.
2.5–4(–4.5) μm (av. 12 × 3.5 m). Basionym: Leptomelanconium australiense B. Sutton, Nova
Hedwigia 25: 163. 1974.
Description and illustration: Andjic et al. (2007a). Diagnosis: Leaf spots irregular medium brown blotches with red-
Typus: Australia, New South Wales, Greenwich, Lane Cove purple margins, and prominently erumpent conidiomata (very
Bushland, on Angophora floribunda, 27 Feb. 2005, A.J. Carnegie, characteristic). Conidia aseptate, medium brown, ovoid,
MURU 426 (holotype DAR 77452, culture ex-type DAR 77452). 7–11 × 4.5–6.5 μm.
Teratosphaeria aurantia Whyte & Andjic, Fungal Diversity 43: Descriptions and illustrations: Crous et al. (2009b), Taylor et al.
30. 2010. (2012).
Diagnosis: Leaf spots epiphyllous and hypophyllous, extending Typus: Australia, Western Australia, York, on C. ficifolia,
through leaf lamina, pale brown, conspicuously circular, collection date unknown, H.L. Harvey (holotype IMI 159079a);
0.5–5 mm diam with corky-brown margins. Conidia ellipsoidal, Western Australia, Perth, Murdoch University Campus, on
0–1-septate, subhyaline to medium brown, smooth, guttulate, C. ficifolia, Jul. 2009, V. Andjic (epitype MURU 453; culture ex-
CROUS ET AL.
Fig. 107. Teratosphaeria molleriana (CBS H-19826). A. Leaf spot (arrows indicate subepidermal ascomata). B. Sporulation on MEA. C, D. Conidia and conidiogenous cells. E,
F. Asci. G. Ascospores. H. Germinating ascospores. Scale bar = 10 μm.
epitype CBS 125244 = MUCC 731); Western Australia, Perth, Diagnosis: Leaf spots pale brown, irregular, amphigenous, up to
Murdoch University Campus, on C. calophylla, Aug. 2009, G.E. 7 mm diam; associated with wasp damage. Conidia pale brown,
St.J. Hardy (paratype MURU 446; culture ex-paratype CBS verruculose, frequently bi-guttulate, characteristically narrowly
124580 = MUCC 695). ellipsoidal, apex subobtuse, base subtruncate, predominantly
straight, with inconspicuous, minute marginal frill, (8–)
Teratosphaeria biformis Whyte & Andjic, Fungal Diversity 43:
9–10(–12) × 3(–4) μm.
30. 2010.
Diagnosis: Leaf spots epiphyllous and hypophyllous, pale brown,
conspicuously circular, 0.5–5 mm diam, extending through leaf Typus: Australia, New South Wales, on E. blakelyi, 13.5 km
lamina. Conidia holoblastic, subhyaline but becoming melanised along Glen Davis road from Capertee, Central Tablelands
when mature, aseptate, ovoid, thick-walled, truncate at base with S33°08’13", E150°04’46", alt. 554 m, Mar. 2006, B. Summerell
a minute marginal frill, (6–)7–10(–11) × (2.5–)3–4(–4) (holotype CBS H-19743, cultures ex-type CBS 120089 = CPC
(av. = 8.5 × 3.5 μm). 12837, CPC 12838, 12839).
Description and illustration: Andjic et al. (2010b). Teratosphaeria brunneotingens (Crous & Summerell) Crous &
Typus: Australia, Queensland, Rosedale, on E. globulus, 2007, Summerell, Persoonia 23: 114. 2009.
G. Whyte (holotype MURU 438, culture ex-type CBS Basionym: Readeriella brunneotingens Crous & Summerell,
124578 = MUCC 693). Stud. Mycol. 58: 26. 2007.
Teratosphaeria blakelyi (Crous & Summerell) Crous & Sum- Diagnosis: Leaf spots amphigenous, irregular specks up to 3 mm
merell, Persoonia 23: 114. 2009. diam, medium brown with a thin, raised, concolourous border.
Basionym: Colletogloeopsis blakelyi Crous & Summerell, Fungal Conidia brown, smooth to finely verruculose, ellipsoidal to sub-
Diversity 23: 328. 2006. cylindrical, apex obtuse to subobtuse, tapering to a subtruncate
Synonym: Readeriella blakelyi (Crous & Summerell) Crous & U. or truncate base (1–1.5 μm wide) with inconspicuous, minute
Braun, Stud. Mycol. 58: 26. 2007. marginal frill, (5–)6–7(–8) × 2–3(–3.5) μm in vitro, becoming 1-
268
septate; in older cultures becoming swollen, and up to 2-septate, E150°36’19", alt. 255 m, Mar. 2006, B. Summerell (holotype
15 μm long and 5 μm wide. CBS H-19744, cultures ex-type CBS 120087 = CPC 12940, CPC
12941, 12942). The CPC numbers were wrongly cited as CPC
12840–12842 in the original publication.
Typus: Australia, Queensland, Cairns, Eureka Creek, 48 km
Teratosphaeria coolabuniensis G.S. Pegg & Carnegie, Aus-
from Mareeba, S17°11'13.2”, E145°02'27.4”, alt. 468 m, on
tralas. Pl. Path. 40: 373. 2011.
E. tereticornis, 26 Aug. 2006, P.W. Crous (holotype CBS H-
19838, culture ex-type CBS 120747 = CPC 13303). Diagnosis: Leaf spots amphigenous, mostly subcircular, some-
Teratosphaeria calophyllae Taylor et al., as “calophylla”, Mycol. times irregular or angular, often confined by larger leaf veins,
Progr. 11: 163. 2012. 5–12 mm diam, separate, becoming confluent, reddish brown on
adaxial, reddish brown to greyish brown on abaxial, with a
Diagnosis: Leaf spots amphigenous, confluent, irregular and slightly raised, narrow, dark brown border on the upper leaf
angular, vein limited spreading inwards from the margins 54 mm, surface. Ascospores bi- to multiseriate, obovoid, with obtuse
pale greyish brown in colour, lesion margins becoming dark red ends, medianly 1-septate, not constricted, hyaline, widest in
to brown and slightly raised. Ascospores hyaline, widest at middle of apical cell, tapering towards basal end, smooth, gut-
middle of apical cell, straight to slightly curved, septate, over- tulate, 8–12 × 2–3 μm.
lapping, constricted at septum, ends obtuse narrowing towards
the base (8–)8.5–11.5(–13) × 2.5–4(–4.5) μm. Description and illustration: Carnegie et al. (2011).
Description and illustration: Taylor et al. (2012). Typus: Australia, Queensland, Coolabunia, Coolabunia Plantation,
on C. torelliana × C. variegata, 14 Feb. 2004, G.S. Pegg & A.J.
Typus: Australia, Western Australia, Perth, Kings Park Botanical Carnegie (holotype BRIP 52585a, culture ex-type BRIP 52585a).
Gardens on C. calophylla, Aug. 2007, K. Taylor (holotype
MURU 457; culture ex-type CBS 124584); Perth, Kings Park Teratosphaeria corymbiae (Carnegie et al.) Carnegie et al.,
Botanical Gardens, on C. calophylla, Aug. 2007, K. Taylor Persoonia 23: 115. 2009. Fig. 106.
(isotype MURU 460; culture ex-isotype MUCC 701). Basionym: Kirramyces corymbiae Carnegie, Andjic & P.A.
Barber, Mycol. Res. 111: 1193. 2007.
Teratosphaeria complicata Crous & Summerell, Persoonia 23:
136. 2009. Diagnosis: Leaf spots amphigenous, subcircular to irregular,
single to confluent, 1–10 mm diam, yellow-brown with thin
Diagnosis: Leaf spots amphigenous, circular, 10–20 mm diam, green-brown to red-purple margin. Conidia holoblastic, fusiform,
brown, with a raised, thin, red-purple border. Conidia sub- straight to mostly curved, occasionally sigmoidal, apex sub-
cylindrical to ellipsoid, straight to curved, brown, verruculose, obtuse, base truncate, not prominently guttulate, pale brown,
almost warty, with longitudinal striations, apex obtuse, base aseptate, (14–)17–23(–24) × 3.5–5 m.
bluntly rounded or truncate, with a flaring marginal frill; wall thick,
0–3-septate, eventually disarticulating with age into aseptate Description and illustration: Carnegie et al. (2007).
conidia; 3-septate conidia up to 30 μm long, 2-septate up to Typus: Australia, New South Wales, Mandalong (native forest),
20 μm long, 1-septate up to 13 μm long, aseptate conidia, (6–) on C. maculata, 15 Jan. 2003, A.J. Carnegie (holotype DAR
7–8 × (4.5–)5–6 μm in vivo. 77445d, culture ex-type DAR 77445).
Description and illustration: Crous et al. (2007b). Note: This species is common but not damaging on Corymbia in
Typus: Australia, Northern Territory, ENE Pine Creek, plantations and native forests in eastern Australia.
S13°40’49", E131°59’04.9", on E. miniata, 23 Sep. 2007, coll. Teratosphaeria crispata Carnegie, Australas. Pl. Path. 40: 376.
B.A. Summerell, isol. P.W. Crous (holotype CBS H-20267, 2011.
14537). Diagnosis: Leaf spots amphigenous, irregular to angular, often
bordered by veins, mostly 1–5 mm diam, single to confluent, light
Teratosphaeria considenianae (Crous & Summerell) Crous & brown to greyish brown with purplish brown to dark brown raised
Summerell, Persoonia 23: 115. 2009. border. Ascospores tri- to multiseriate, overlapping, hyaline, straight
Basionym: Colletogloeopsis considenianae Crous & Summerell, to slightly curved, fusoid-ellipsoidal with obtuse ends, medianly 1-
Fungal Diversity 23: 329. 2006. septate, widest in middle of apical cell, slightly constricted at
Synonym: Readeriella considenianae (Crous & Summerell) septum, tapering towards both ends, but more prominently towards
Crous & U. Braun, Stud. Mycol. 58: 26. 2007. the basal end, (10–)12–14(−15) × 3.5–4 μm.
Diagnosis: Leaf spots amphigenous, circular, medium brown,
1–4 mm diam, surrounded by a prominent red-purple margin.
Conidia medium brown, verruculose, ellipsoidal, apex obtuse, base Typus: Australia, New South Wales, Tumbarumba, Mannus
subtruncate to truncate, straight to slightly curved, State Forest, on E. bridgesiana, 23 May 2007, A.J. Carnegie
with inconspicuous, minute marginal frill, (6–)7–9(–10) × 3(–4) μm. (holotype DAR 77453b, culture ex-type CBS
Description and illustration: Summerell et al. (2006). 130523 = DAR77453).
Typus: Australia, New South Wales, Blaxland, on Teratosphaeria cryptica (Cooke) Crous & U. Braun, Stud.
E. consideniana, in Blaxland War Memorial Park, opposite Mycol. 58: 10. 2007. Fig. 105.
Blaxland Public School, intersection of Wilson Way and Great Basionym: Sphaerella cryptica Cooke, Grevillea 20(no. 93): 5.
Western Highway, Central Coast NSW, S33°44’14", 1891.
CROUS ET AL.
Synonyms: Mycosphaerella cryptica (Cooke) Hansf., Proc. Linn. Basionym: Kirramyces destructans M.J. Wingf. & Crous, S. Afr.
Soc. N.S.W. 81: 35. 1956. J. Bot. 62: 325. 1996.
Colletogloeum nubilosum Ganap. & Corbin, Trans. Brit. Mycol. Synonyms: Phaeophleospora destructans (M.J. Wingf. & Crous)
Soc. 72: 237. 1979. Crous et al., S. Afr. J. Bot. 63: 113. 1997.
Colletogloeopsis nubilosum (Ganap. & Corbin) Crous & M.J. Readeriella destructans (M.J. Wingf. & Crous) Crous & U. Braun,
Wingf., Canad. J. Bot. 75: 668. 1997. Stud. Mycol. 58: 26. 2007.
Diagnosis: Leaf spots circular to irregular, discrete to confluent, Diagnosis: Leaf spots irregular to subcircular, 10–20 mm diam,
light brown to red-brown, variable in size, becoming dark grey with or blighting large areas of the lamina, pale brown on both sur-
age and pseudothecial formation, frequently with a prominent faces, border diffuse, margin red-brown when present. Symp-
purple, raised margin; spots frequently coalesce, and cause a toms vary from shoot blight to large subcircular leaf spots or leaf
distortion of the leaf lamina. Ascospores 2–3- or multiseriate, blight. Conidia holoblastic, solitary, dry, exuding in long cirrhi,
colourless, ellipsoidal with rounded ends, 1-septate, constricted at forming black masses on the leaf surface; cylindrical, apex
median septum, 9–17.5 × 2–5.5 μm. Conidia single, subhyaline, obtuse, or tapering slightly from uppermost septum to a subacute
smooth-walled, cylindrical, straight or curved, apex obtuse, base apex as in T. eucalypti; base truncate, 1.5–2 μm wide, minute
truncate with a marginal frill, (8.5–)10–15(–18) × 4–5(–6) μm. marginal frill mostly absent; variously curved, rarely straight,
thick-walled, pale brown, verruculose, guttulate, (1–)3-
Descriptions and illustrations: Park & Keane (1982a), Crous
euseptate, (30–)50–65(–70) × 2.5(–3) μm.
(1998).
Description and illustration: Wingfield et al. (1996).
Typus: Australia, Victoria, Melbourne, on Eucalyptus sp., Mrs.
Martin 753 (holotype of S. cryptica in K); Victoria, Otway Typus: Indonesia, Northern Sumatra, Lake Toba area, Aek
Ranges, on E. globulus, 15 Feb. 2007, I.W. Smith (epitype Nauli, on E. grandis, Feb. 1996, M.J. Wingfield (holotype of
designated here CBS H-24043, MBT388169, cultures ex-epitype Kirramyces destructans PREM 54416, cultures ex-type CBS
CBS 145895 = CPC 13839, CBS 145896 = CPC 13842). New 111369 = CMW 5219 = CPC 1366, CPC 1367, 1368; paratype of
Zealand, Auckland, Auckland University, on E. gigantea, Apr. Kirramyces destructans PREM 54689); Sumatra, Sabisa, on
1977, A. Nathan Ganapathi (holotype of Colletogloeum nubi- E. grandis, Feb. 1996, M.J. Wingfield (paratype of Kirramyces
losum PDD 37677). destructans PREM 54688); Taratung, on E. grandis, Feb. 1996,
M.J. Wingfield (paratype of Kirramyces destructans PREM
Teratosphaeria delegatensis (R.F. Park & Keane) Crous,
54690); Northern Sumatra, Habinsarin, on E. grandis, Feb. 1996,
comb. nov. MycoBank MB832051.
M.J. Wingfield (paratype of Kirramyces destructans PREM
Basionym: Mycosphaerella delegatensis R.F. Park & Keane,
54691).
Synonyms: Stagonospora delegatensis R.F. Park & Keane, Teratosphaeria dimorpha (Crous & Carnegie) Crous & Sum-
Trans. Brit. Mycol. Soc. 83: 95. 1984. merell, Persoonia 22: 42. 2009.
Phaeophleospora delegatensis (R.F. Park & Keane) Crous, Basionym: Colletogloeopsis dimorpha Crous & Carnegie, Fungal
Mycol. Mem. 21: 51. 1998. Diversity 23: 331. 2006.
Kirramyces delegatensis (R.F. Park & Keane) Andjic, Mycol. Synonym: Readeriella dimorpha (Crous & Carnegie) Crous & U.
Res. 111: 1191. 2007. Braun, Stud. Mycol. 58: 26. 2007.
Diagnosis: Leaf spots amphigenous, yellow-green, becoming Diagnosis: Leaf spots amphigenous, medium to dark brown,
dark brown, and finally grey with raised margins, round to irregular to angular, with a raised border, 2–5 mm diam. Conidia
confluent, 5–10 mm diam. Ascospores multiseriate, hyaline, (7–)9–11(–13) × (3–)4(–5) μm, medium brown, finely verru-
straight, fusiform to clavate, with the widest point in the upper half culose, guttulate, ellipsoidal to fusiform, straight, apex sub-
of the apical cell, 1-septate, without a constriction at the median obtuse, widest in middle if fusiform, or in lower third of conidium if
septum, 16–25 × 3–5 μm. Conidia cylindrical, straight or curved, ellipsoidal, tapering towards a subtruncate base, 1–1.5 μm wide;
colourless, smooth, apex obtuse, base truncate, unequally 1- with age some conidia become median septate, usually at the
septate, 21–51 × 3–5 μm. onset of microcyclic conidiation.
Descriptions and illustrations: Park & Keane (1984). Description and illustration: Summerell et al. (2006).
Typus: Australia, Victoria, Narbethong, on E. delegatensis, 30 Typus: Australia, New South Wales, Rosewood, Carabost State
Jun. 1982, R.F. Park (holotype of M. delegatensis DAR 45718, Forest, on Eucalyptus sp., Jan. 2006, A. Carnegie (holotype
isotype of M. delegatensis DAR 280471; isotype of CBS H-19739, isotype DAR 77443, culture ex-type CBS
K. delegatensis DAR 280472). Not known from culture. 120086 = CPC 12919).
Notes: A single sequence in GenBank under the name Mycos- Teratosphaeria dunnii Crous & Carnegie, Persoonia 42: 327.
phaerella delegatensis (GenBank AB435070.1 based on IMI 2019.
362252 and derived from E. camaldulensis in Ethiopia) blasts Diagnosis: Conidiomata pycnidial, solitary, brown, globose,
with species of Septoria and the identification of this sequence is 90–200 μm diam. Conidiophores lining the inner cavity, sub-
therefore doubtful. Furthermore, this pathogen is also not known cylindrical, pale brown, 1–2-septate, branched or not,
from outside Australia. Surveys to recollect this species by the 7–20 × 2.5–4 μm. Conidiogenous cells subcylindrical to dolii-
authors have thus far been unsuccessful. form, medium brown, verruculose, proliferating percurrently at
Teratosphaeria destructans (M.J. Wingf. & Crous) M.J. Wingf. apex, 5–8 × 3.5–4 μm. Conidia solitary, aseptate, thick-walled,
& Crous, Persoonia 23: 115. 2009. Fig. 105. guttulate, golden brown, verruculose, subcylindrical to fusoid-
270
ellipsoid, apex subobtuse, base truncate, 1.5–2 μm diam with Synonyms: Pseudocercospora eucalypti (Cooke & Massee) Y.L.
minute marginal frill, (6–)8–9(–11) × (2.5–)3(–3.5) μm. Guo & X.J. Liu, Mycosystema 2: 234. 1989.
Kirramyces eucalypti (Cooke & Massee) J. Walker et al., Mycol.
Res. 96: 920. 1992.
Typus: Australia, New South Wales, Yabbra State Forest, Boomi Phaeophleospora eucalypti (Cooke & Massee) Crous et al., S.
Creek plantation, on E. dunnii, 19 Apr. 2016, A.J. Carnegie, HPC Afr. J. Bot. 63: 113. 1997.
2430 (holotype CBS H-23945, culture ex-type CPC Septoria normae Heatler, Studies on Septoria n. sp., causing a
35653 = CBS 145548). leaf blotch of Eucalyptus dalrympleana Maid. 1961. M.Sc.
Thesis, Univ. of Sydney, Sydney, Australia.
Teratosphaeria epicoccoides (Cooke & Massee) Rossman &
Septoria pulcherrima Gadgil & M.A. Dick, N.Z. Jl Bot. 21: 49.
W.C. Allen, IMA Fungus 6: 519. 2015. Fig. 106.
1983.
Basionym: Cercospora epicoccoides Cooke & Massee, Grevillea
Stagonospora pulcherrima (Gadgil & M.A. Dick) H.J. Swart,
19: 91. 1891.
Synonyms: Kirramyces epicoccoides (Cooke & Massee) J.
Readeriella pulcherrima (Gadgil & M.A. Dick) Crous & U. Braun,
Walker et al., Mycol. Res. 96 919. 1992.
Stud. Mycol. 58: 26. 2007.
Phaeophleospora epicoccoides (Cooke & Massee) Crous et al.,
S. Afr. J. Bot. 63: 113. 1997. Diagnosis: Leaf spots amphigenous, subcircular to irregular,
Readeriella epicoccoides (Cooke & Massee) Crous & U. Braun, medium brown with a slightly raised border and wide, red-purple
Stud. Mycol. 58: 11. 2007. margin, 2–10 mm diam. Conidia holoblastic, solitary, sub-
Hendersonia grandispora McAlpine, Proc. R. Soc. N.S.W. 28: 99. cylindrical to narrowly obclavate, tapering toward an subobtuse
1903. apex, base truncate to long obconic-truncate, straight to flex-
Phaeoseptoria eucalypti Hansf., Proc. Linn. Soc. N.S.W. 82: 225. uous, pale brown, finely verruculose, not prominently guttulate,
1957. (0–)1(–2)-euseptate, (25–)35–50(–55) × (2–)3–4 μm; incon-
Phaeoseptoria luzonensis Tak. Kobay., Trans. Mycol. Soc. Japan spicuous basal marginal frill present.
19: 377. 1978.
Descriptions and illustrations: Walker et al. (1992), Crous (1998).
Mycosphaerella suttonii Crous & M.J. Wingf., as “suttoniae”,
Canad. J. Bot. 75: 783. 1997. Typus: Australia, Victoria, Oakleigh, on Eucalyptus sp., before
Teratosphaeria suttonii (Crous & M.J. Wingf.) Crous & U. Braun, 1889, Mrs Martin No. 436 (holotype of Cercospora eucalypti
Stud. Mycol. 58: 11. 2007. K(M) 39487).
Mycosphaerella obscuris P.A. Barber & T.I. Burgess, Fungal
Teratosphaeria fimbriata (Crous & Summerell) Crous & U.
Diversity 24: 146. 2007.
Braun, Stud. Mycol. 58: 10. 2007. Fig. 105.
Teratosphaeria obscuris (P.A. Barber & T.I. Burgess) P.A. Barber
Basionym: Mycosphaerella fimbriata Crous & Summerell, Fungal
& T.I. Burgess, Persoonia 23: 115. 2009.
Diversity 26: 166. 2007.
Diagnosis: Leaf spots primarily epiphyllous to hypophyllous,
Diagnosis: Leaf spots amphigenous, irregular to circular, 5–15 mm
angular to irregular, purple with an indistinct border, 0.5–3 mm
diam, medium to dark brown, with radiating superficial mycelium,
diam, but brown spots up to 7(–25) mm diam with red-purple
spreading from ascomata that are predominantly in the middle of
margins are frequently observed in the field. Ascospores multi-
the lesion; hyphae red-brown, 5–8 μm wide, thick-walled, verru-
seriate, overlapping, colourless, guttulate, thin-walled, straight to
culose, aggregating in hyphal strands (also in vitro), with
curved, obovoid with obtuse ends, widest near apex, medianly 1-
chlamydospore-like cells, up to 15 μm diam, aggregating in clusters;
septate, not constricted at septum, tapering more prominently
forming spermatogonia in the outer region of the lesion (also formed
toward base (10–)11–12(–13) × (2.5–)3–3.5 μm. Conidia
in vitro). Ascospores multiseriate, overlapping, hyaline, guttulate,
holoblastic, apical, solitary, medium brown, (15–)
thin-walled, straight to slightly curved, obovoid, with obtuse ends,
30–35(–55) × (4–)5–6 μm, verruculose, 0–5-transversely
widest near the apex of the apical cell, mostly medianly 1-septate,
euseptate, guttulate, straight to variously curved, obclavate to
constricted at the septum; larger ascospores tend to be unequally 1-
subcylindrical, apex rounded to obtuse, base truncate to obconic-
septate, with the upper cell being up to 13 μm long, and the bottom
truncate with a marginal frill; primary conidia frequently forming
cell up to 17 μm long, tapering towards both ends, but more
lateral branches or secondary conidia via microcyclic conidiation.
prominently towards the lower end, (18–)22–17(–30) × (6–)
Descriptions and illustrations: Walker et al. (1992), Crous & 7(–8) μm; ascospores frequently with a persistent mucous sheath.
Wingfield (1997), Crous (1998).
Typus: Indonesia, Northern Sumatra, Lake Toba area, on
Typus: Australia, Queensland, Cairns, S16°56’23.3”,
Eucalyptus sp., Mar. 1996, M.J. Wingfield (holotype of
E145°32’34.6”, on leaves of Corymbia sp., 26 Aug. 2006, P.W.
M. suttoniae PREM 54963a, cultures ex-type CMW 5348 = CPC
Crous (holotype CBS H-19828, cultures ex-type CBS
1345, CPC 1346, 1347); idem., PREM 54963b of asexual morph,
120736 = CPC 13324, CPC 13325, 13326).
derived from single-ascospore cultures on MEA. Australia,
Victoria, on Eucalyptus sp., before 1891, Mrs Martin No. 600 Teratosphaeria foliensis Andjic & S. Jacks., Fungal Diversity
(holotype of asexual morph K(M) 39488). 43: 32. 2010.
Diagnosis: Leaf spots raised, pale brown, circular up to 9 mm
Teratosphaeria eucalypti (Cooke & Massee) Crous, Persoonia
diam; border medium to dark brown, raised with a purple margin.
23: 115. 2009. Fig. 105.
Conidia solitary, hyaline to subhyaline, guttulate, aseptate,
Basionym: Cercospora eucalypti Cooke & Massee, in Cooke,
smooth, ellipsoidal to obovoid, base truncate to subtruncate with
Grevillea 18(no. 85): 7. 1889.
CROUS ET AL.
marginal frill, apex obtuse, (8–)8.5–11(–12.5) × (2.7–) Diagnosis: Leaf spots amphigenous, circular, 1.5–5 mm diam,
3–3.5(–5) (av. = 10.5 × 3.5 m). pale brown, becoming medium brown toward the raised
border; margin thin, red-purple. Conidia holoblastic, solitary, cy-
Description and illustration: Andjic et al. (2010b).
lindrical, apex obtuse, base truncate, 2–4 μm wide, with a mar-
Typus: Australia, New South Wales, Commins plantation, ginal frill; straight to curved, thick-walled, medium brown,
S34°350 100 , E146°90 5600 , on E. globulus, May 2005, S. Collins & verruculose, guttulate, (1–)3(–4)-euseptate, (35–)40–48(–50) ×
S. Jackson (holotype MURU 442, culture ex-type CBS 5–6(–7) μm.
124581 = MUCC 670).
Descriptions and illustrations: Walker et al. (1992), Crous (1998).
Teratosphaeria henryi Crous & Carnegie, Persoonia 42: 315.
Typus: Australia, New South Wales, Wiseman's Ferry, on
2019.
E. eximia, Jul. 1949, L.R. Fraser (holotype DAR 3832). Not
Diagnosis: Conidiomata pycnidial, solitary, brown, 90–120 μm known from culture.
diam. Conidiophores reduced to conidiogenous cells lining
Teratosphaeria majorizuluensis Crous & Summerell, Persoo-
cavity. Conidiogenous cells brown, verruculose, subcylindrical
nia 23: 137. 2009.
with slight apical taper, proliferating percurrently at apex,
6–12 × 3–4 μm. Conidia solitary, brown, verruculose, aseptate, Diagnosis: Leaf spots amphigenous, irregular blotches up to
granular, fusoid, apex subobtuse, base truncate, 2 μm diam, with 3 cm diam, medium brown with a thin, raised, dark brown border,
minute marginal frill, (7–)8–10(–11) × (2.5–)3(–4) μm. and red-purple margin. Conidia brown, verruculose, ellipsoidal to
subcylindrical, apex obtuse to subobtuse, tapering to a sub-
truncate or truncate base (1–2 μm wide) with inconspicuous,
Typus: Australia, New South Wales, Tallawandi plantation, minute marginal frill, (4–)5–6(–7) × (1.5–)2(–2.5) μm in vivo;
South Grafton, on C. henryi, 17 Apr. 2016, A.J. Carnegie, HPC 6–9 × 2.5–4 μm in vitro.
2417 (holotype CBS H-23939, culture ex-type CPC
35715 = CBS 145539).
Typus: Australia, New South Wales, Middle Head, Sydney
Teratosphaeria hortaea Crous & M.J. Wingf., Persoonia 22: 89.
Harbour National Park, S33°49’51", E151°15’31", on
2009.
E. botryoides, Feb. 2006, coll. B.A. Summerell, isol. P.W. Crous
Diagnosis: Conidia ellipsoid, pale to medium brown, apex (holotype CBS H-19773, cultures ex-type CBS 120040 = CPC
obtusely rounded, widest in middle, tapering towards a sub- 12712, CPC 12713, 12714).
truncate base, 1 μm wide, (4–)5–6(–7) × (2–)2.5(–3) μm. On
Teratosphaeria mexicana (Crous) Crous & U. Braun, Stud.
MEA conidia become 1-septate, and frequently undergo micro-
Mycol. 58: 10. 2007.
cyclic conidiation (percurrently), and in general are darker brown,
Basionym: Mycosphaerella mexicana Crous, Mycol. Mem. 21:
up to 15 μm long, 5 μm wide, with minute marginal frill, and
81. 1998.
subtruncate to truncate base.
Diagnosis: Leaf spots amphigenous, round to subcircular,
1–8 mm diam, pale brown in the centre, becoming medium
Typus: Madagascar, Morondavo, on E. camaldulensis, Aug. brown toward the raised border, surrounded by a dark brown to
2007, M.J. Wingfield (holotype CBS H-20194, culture ex-type black margin (the leaves are leathery and brown, and the mar-
CBS 124156 = CPC 15716 = A8). gins may have been red-purple when collected). Ascospores bi-
to triseriate, overlapping, colourless to olivaceous, guttulate,
Teratosphaeria juvenalis Crous & M.J. Wingf., Persoonia 22:
smooth to verruculose, thick-walled, straight to slightly curved,
44. 2009.
fusoid-ellipsoidal with obtuse ends, widest in middle of the apical
Diagnosis: Leaf spots raised, medium brown, circular, up to 7 mm cell, medianly 1-septate, not to slightly constricted at septum,
diam; border medium to dark brown, raised, with a red-purple tapering toward both ends, but more so toward base (15–)
margin. Conidia ellipsoidal to subcylindrical, apex subobtuse, 17–22(–24) × (5–)5.5–6(–7) μm.
base truncate to subtruncate, (1–)2–3(–4) μm wide, generally
widest at the median, thin-walled, guttulate, verruculose, (10–)
11–13(–15) × (4–)5(–6) μm; basal marginal frill present. Typus: Mexico, Piedras Negras, on Eucalyptus sp., 26 Jan.
1986, D. Sailors (holotype BPI 602163).
Teratosphaeria micromaculata Whyte & Andjic, Fungal Di-
Typus: South Africa, Western Cape Province, Stellenbosch,
versity 43: 32. 2010.
Stellenbosch Mountain, on E. cladocalyx, Apr. 1988, P.W. Crous
(holotype CBS H-20180, culture ex-type CBS 110906 = CMW Diagnosis: Leaf spots epiphyllous and hypophyllous, dark brown
13347 = CPC 40). circular lesion 0.5–2 mm diam, with a raised purple margin
followed by a pale brown margin, extending through leaf lamina.
Teratosphaeria lilianiae (J. Walker et al.) Crous & Andjic,
Conidia ellipsoidal to ovoid, aseptate, thick-walled, hyaline when
Persoonia 23: 115. 2009.
produced but becoming melanised, truncate to subtruncate at
Basionym: Kirramyces lilianiae J. Walker, B. Sutton & Pascoe,
base with marginal frill, (5–)5–7(–7.5) × (2–)2–3(–3.5)
Mycol. Res. 96: 921. 1992.
(av. = 6 × 2.5 μm).
Synonym: Phaeophleospora lilianiae (J. Walker et al.) Crous
et al., S. Afr. J. Bot. 63: 115. 1997. Description and illustration: Andjic et al. (2010b).
272
Typus: Australia, Queensland, Boonah, on E. globulus, 2007, G. N40°00’39”, W8°36’2.3”, alt. 77 m, on Eucalyptus sp., 13 Oct.
Whyte (holotype MURU 437, cultures ex-type CBS 2006, P.W. Crous & A.J.L. Phillips (epitype of S. molleriana CBS
124582 = MUCC 647, MUCC 648). H-19826, cultures ex-epitype CBS 120746 = CPC 13398, CPC
13399, 13400).
Teratosphaeria miniata Crous & Summerell, Persoonia 23: 139.
2009. Teratosphaeria multiseptata (Carnegie) Carnegie, Persoonia
23: 115. 2009.
Diagnosis: Leaf spots amphigenous, irregular, 3–12 mm diam,
Basionym: Mycosphaerella multiseptata Carnegie, Mycologia 99:
medium brown, with a raised, brown border, and thin, red-purple
471. 2007.
margin. Conidia solitary, brown, aseptate, verruculose, ellipsoidal
to subcylindrical, apex obtuse to subobtuse, tapering to a sub- Diagnosis: Leaf spots on Angophora subvelutina: amphigenous,
truncate or truncate base (1–2 μm wide), with inconspicuous circular to irregular, single to confluent, 2–7 mm diam, yellow-
marginal frill, (5–)6–7(–8) × (2.5–)3(–3.5) μm; older cultures have brown becoming grey-brown on adaxial surface, yellow-brown
conidia that become swollen, broadly ellipsoid, 1-septate, up to to red-brown on abaxial surface, with prominent red-brown
10 μm long and 5 μm wide (more pronounced on MEA than on OA). border; on Angophora costata: amphigenous, subcircular with
irregular margins, bordered by veins, single to confluent,
3–12 mm diam, yellow-brown becoming grey brown on adaxial
Typus: Australia, Northern Territory, ENE Pine Creek, surface, yellow-brown to red-brown on abaxial surface, with
S13°40’49", E131°59’04.9", on E. miniata, 23 Sep. 2007, coll. prominent red-brown border, often with anthocyanin pigmenta-
B.A. Summerell, isol. P.W. Crous (holotype CBS H-20269, tion surrounding border. Ascospores 2–3-seriate, overlapping,
cultures ex-type CBS 125006 = CPC 14514, CPC 14515, hyaline, straight, rarely curved, narrowly obovoid to fusiform with
14516). obtuse ends, medianly or unequally 1-septate, widest in middle
of apical cell, mostly not constricted at septum, tapering to basal
Teratosphaeria molleriana (Thüm.) Crous & U. Braun, Stud.
end, (12.5–)13.5–17(–19) × (3–)3.5–4.5(–5) μm.
Mycol. 58: 10. 2007. Fig. 107.
Basionym: Sphaerella molleriana Thüm., Revista Inst. Sci. Lit. Description and illustration: Carnegie et al. (2007).
Coimbra 28: 31. 1881.
Typus: Australia, New South Wales, Whiporie, Pintexan Prop-
Synonyms: Mycosphaerella molleriana (Thüm) Lindau, Nat.
erty, native forest, on Angophora subvelutina, 18 Jan. 2005, A.J.
Pfanzenfam. 1: 424. 1897.
Carnegie (holotype DAR 77438, culture ex-type CBS
Colletogloeopsis molleriana Crous & M.J. Wingf., Canad. J. Bot.
121312 = DAR 77438).
75: 670. 1997.
Readeriella molleriana (Crous & M.J. Wingf.) Crous & U. Braun, Teratosphaeria novaehollandiae V. Andjic et al., IMA Fungus
Stud. Mycol. 58: 10. 2007. 7: 260. 2016.
Mycosphaerella vespa Carnegie & Keane, Mycol. Res. 102:
Diagnosis: Leaf spots circular to irregular, 2.5–35 mm diam,
1275. 1998.
single to confluent, medium brown to pale brown with red brown
Mycosphaerella ambiphylla A. Maxwell, Mycol. Res. 107: 354.
border on the top surface, pale brown at the bottom. Conidia
2003.
solitary, 1–3-septate, hyaline to subhyaline, slightly verruculose,
Teratosphaeria xenocryptica Crous & M.J. Wingf., Persoonia 23:
cylindrical, straight to slightly curved, thick-walled, base truncate
139. 2009.
sometimes with marginal frill, apex obtuse, (20–)
Diagnosis: Leaf spots amphigenous, subcircular to irregular, 35–40(–50) × (1.5–)2–3.5(–4) μm.
becoming confluent, 2–10 mm diam, pale brown, becoming
Description and illustration: Andjic et al. (2016).
darker brown toward the raised, medium brown border on the
upper surface; border concolourous on the lower surface; col- Typus: Australia, Western Australia, Kununurra, on
oured margin absent. Ascospores bi- to multiseriate, over- E. camaldulensis, Apr. 2012, A. Maxwell & V. Andjic (holotype
lapping, colourless, guttulate, thin-walled, straight to curved, BRIP 59486, culture ex-type AQISWA 201302 = BRIP 59486).
fusoid-ellipsoidal with obtuse ends, widest in middle of apical
Teratosphaeria nubilosa (Cooke) Crous & U. Braun, Stud.
cells, medianly 1-septate, not or only slightly constricted at
Mycol. 58: 10. 2007.
septum, tapering toward both ends, but more so toward base
Basionym: Sphaerella nubilosa Cooke, Grevillea 19(no. 91): 61.
(11–)12–14(–17) × (2.5–)3.5–4(–4.5) μm. Conidia single,
1891.
medium brown, verruculose, subcylindrical to ellipsoidal, straight
Synonyms: Mycosphaerella nubilosa (Cooke) Hansf., Proc. Linn.
to slightly curved, occasionally with a lateral protuberance that
Soc. N.S.W. 81: 36. 1956.
can develop into a secondary conidium; apex obtuse, base
Mycosphaerella juvenis Crous & M.J. Wingf., Mycologia 88: 453.
truncate with a marginal frill, (7–)9–12(–13) × (2.5–)
1996.
3–3.5(–4) μm; conidia rarely becoming 1-septate in culture.
Mycosphaerella quasicercospora Crous & T.A. Cout., Stud.
Descriptions and illustrations: Crous & Wingfield (1997), Crous Mycol. 55: 119. 2006.
(1998). Teratosphaeria quasicercospora (Crous & T.A. Cout.) Crous & U.
Typus: Portugal, Lusitania, on E. globulus, Jul. 1879, Fr. Moller
Diagnosis: Leaf spots amphigenous, varying from pin spots or
(holotype of Sphaerella molleriana in K); Abrantes, on
flecks to small, round or irregular spots, frequently circular to
E. globulus, Jul. 1995, S. McCrae (holotype of Colletogloeopsis
irregular, up to 15 mm diam, becoming confluent to form larger
molleriana PREM 54395, cultures ex-type CBS 111164 = CMW
blotches up to 3 cm diam on older leaves, pale brown, sur-
4940 = CPC 1214, CBS 111165 = CPC 1215); Lisbon,
rounded by a raised dark brown border, and a thin red-purple
CROUS ET AL.
diffuse margin. Ascospores bi- to triseriate, overlapping, hyaline, occasionally delineated by veins, often with purple pigmentation
non-guttulate, thin-walled, but the septum appearing thicker than surrounding part of the leaf spot. Commonly older (grey) lesions
the side walls, straight to slightly curved, obovoid with obtuse are located within younger (yellowish brown to reddish brown)
ends, medianly or unequally 1-septate, not or slightly constricted lesions. Ascospores multiseriate, overlapping, straight to curved,
at the septum, tapering towards both ends, but more prominently fusiform with obtuse ends, tapering slightly to basal end, mostly
towards the lower end, (8–)13–14(–16) × (2.5–)3–4(–4.5) μm unequally 1-septate (the basal cell longer), not constricted or only
in vivo; apical cell (4–)5–6 μm, basal cell (4–)7–9 μm long. slightly so, widest in upper half of apical cell, (32–)40–52 × (4–)
6–8 μm.
Descriptions and illustrations: Park & Keane (1982a), Crous
(1998), Crous et al. (2004b). Description and illustration: Carnegie et al. (2011).
Typus: Australia, Victoria, Melbourne, on Eucalyptus sp., Mrs. Typus: Australia, New South Wales, Guest Plantation, Ebor, on
Martin 584 (holotype of Sphaerella nubilosa in K); Victoria, E. dunnii, 15 Feb. 2005, G. Price (holotype DAR 77441, culture
Briagolong, on E. globulus, 16 Sep. 1994, A. Carnegie (epitype ex-type DAR 77441).
CBS H-9902, culture ex-epitype CBS 116005 = CPC 937).
Teratosphaeria profusa Crous & Carnegie, Persoonia 23: 139.
Tanzania, on E. maidenii, May 1995, T. Coutinho (holotype of
2009.
M. quasicercospora PREM 54971, culture ex-type CBS
111161 = CPC 1098). Diagnosis: Isolated from leaf spots together with several other
fungi. Conidia solitary, brown, verruculose, with two prominent
Note: New DNA sequence data generated of M. quasicercospora
guttules, ellipsoidal to subcylindrical, 0(–1)-septate, apex sub-
(CBS 111161; Crous et al. 2006d) showed that it is a synonym of
obtuse, tapering to a subtruncate or truncate base (1–2 μm
T. nubilosa.
wide), with inconspicuous marginal frill, (7–)8–10(–13) × (2.5–)
Teratosphaeria ovata (H.J. Swart) Crous & Summerell, Per- 3(–3.5) μm.
soonia 22: 44. 2009.
Basionym: Coniothyrium ovatum H.J. Swart, Trans. Brit. Mycol.
Soc. 86: 495. 1986. Typus: Australia, Bago State Forest, on E. nitens, 22 Dec. 2005,
Synonym: Coniothyrium parvum H.J. Swart, Trans. Brit. Mycol. coll. A.J. Carnegie, isol. P.W. Crous (holotype CBS H-20270,
Soc. 86: 495. 1986. culture ex-type CBS 125007 = CPC 12821).
Diagnosis: Leaf spots amphigenous, raised, medium brown, Teratosphaeria pseudocryptica (Crous) Crous & U. Braun,
circular, 1–2 mm diam; border medium to dark brown, raised; Stud. Mycol. 58: 11. 2007.
margin absent on E. dives, but thin and red-purple on Basionym: Mycosphaerella pseudocryptica Crous, Stud. Mycol.
E. melliodora. Conidia ellipsoidal, apex subobtuse, base trun- 55: 116. 2006.
cate, generally widest at or below the median, finely verruculose,
(6–)7–9(–11) × 3–3.5(–4) μm; basal marginal frill present.
0.5–2 mm diam, pale brown, with a raised, red-brown margin.
Descriptions and illustrations: Swart (1986b), Crous et al. (2009a). Ascospores multiseriate, overlapping, hyaline, granular, thin-
walled, straight, fusoid–ellipsoidal with obtuse ends, medianly
Typus: Australia, Victoria, Calder Highway, on E. dives, 17 May 1-septate, widest at the middle of the apical cell, constricted at
1983, H.J. Swart (holotype of C. ovatum DAR 49461). the septum, tapering towards both ends, but more prominently
towards the lower end, (11–)12–14(–15) × (3–)3.5(–4) μm,
Teratosphaeria pluritubularis (Crous & Mansilla) Crous & U.
in vivo. Conidia holoblastic, solitary, aseptate, fusoid with obtuse
to subobtuse apices and truncate bases, medium brown, finely
Basionym: Mycosphaerella pluritubularis Crous & Mansilla, Stud.
verruculose, (10–)12–14(–17) × (3.5–)4(–6) μm; inconspic-
Mycol. 55: 114. 2006.
uous basal marginal frill present.
Diagnosis: Leaf spots amphigenous, irregular to sub-circular,
5–15 mm diam, pale to medium brown, surrounded by a thin,
raised, dark brown border. Ascospores multiseriate, overlapping, Typus: New Zealand, Wellington Botanical Garden, on Euca-
hyaline, prominently guttulate, thin-walled, straight, obovoid with lyptus sp., Mar. 2004, J.A. Stalpers (holotype CBS H-19693,
subobtuse ends, medianly 1-septate, widest at the middle of the cultures ex-type CBS 118504 = CPC 11267, CPC 11268, 11269,
apical cell, constricted at the septum, tapering towards both sexual morph; CPC 11264–11266, asexual morph).
ends, but more prominently towards the lower end, (8–)
Teratosphaeria pseudoeucalypti Andjic & T.I. Burgess, Pl.
9–10(–11) × 3(–4) μm in vivo.
Path. 59: 908. 2010.
Diagnosis: Leaf spots subcircular to irregular, 2–15 mm diam,
Typus: Spain, on E. globulus, Nov. 2004, J.P. Mansilla (holotype single to confluent, often blighting on E. grandis hybrids, initially
CBS H-19692, culture ex-type CBS 118508 = CPC 11697). pale green, turning chlorotic before becoming necrotic, pale to
medium brown with red-purple margin on the upper and lower
Teratosphaeria praelongispora Carnegie, Australas. Pl. Path.
surface. Conidia solitary, 0–3-septate, subhyaline to pale brown,
40: 378. 2011.
slightly verruculose, cylindrical, straight to variously curved,
Diagnosis: Leaf spots amphigenous, sub-circular to irregular, thick-walled, base truncate sometimes with marginal frill, apex
2–10 mm diam., single to often confluent, yellowish brown to obtuse, (26–)31.5–40(–58) × (1.7–)2–2.5(–3.5) μm.
reddish brown becoming grey with age, with prominent raised
reddish brown to dark brown border (especially on E. dunnii), Description and illustration: Andjic et al. (2010a).
274
Typus: Australia, Queensland, Miriam Vale, on medium brown, finely verruculose, widest below the middle,
E. grandis × E. camaldulensis, Aug. 2005, G. Pegg (holotype (6.5–)7–9(–10) × (3–)3.5(–4) μm.
MURU 450, culture ex-type CBS 124577 = MUCC 607).
Teratosphaeria pseudonubilosa G. Perez & Carnegie, sp.
nov. MycoBank MB832052.
Mountain, on Eucalyptus sp., 4 Dec. 2004, P.W. Crous (holotype
Synonym: Teratosphaeria pseudonubilosa G. Perez & Carnegie,
Australas. Pl. Path. 43: 72. 2014. nom. inval. (Art. 42, Melbourne).
Teratosphaeria tinarooa Andjic & T.I. Burgess as “tinara”,
Etymology: Name reflects its similarity to T. nubilosa.
Fungal Diversity 43: 34. 2010.
Diagnosis: Leaf spots occurring on both juvenile and adult foliage
Diagnosis: Leaf spots amphigenous, irregular blotches, 1–4 mm
of E. globulus. Leaf lesions on juvenile foliage amphigenous,
diam, pale, brown with purple border. Conidia subhyaline
circular to sub-circular, up to 15 mm diam, yellow to brown in
becoming brown when mature, aseptate, guttulate, thick-walled,
colour, surrounded by a thick, raised, brown border. Single le-
obovoid to broadly ellipsoidal, base truncate to subtruncate with
sions frequently coalescing to form larger blotches across the
minute marginal frill, apex obtuse, (3.5–)6–7.5(–9.5) × (1.8–)
leaf surfaces. On adult foliage, lesions amphigenous, angular,
3–3.5(–4.5) (av. = 6.5 × 3.1 μm),
showing conspicuous brown border, more prominent than on
juvenile foliage. Ascospores 2–3 seriate, overlapping, hyaline, Description and illustration: Andjic et al. (2010b).
thin-walled, guttulate, straight, ellipsoidal, medially 1-septate,
widest at the middle of the apical cell, slightly constricted at the Typus: Australia, northern Queensland, Mareeba, Lake Tinaroo,
septum, tapering towards both ends but more prominently to- on Corymbia sp., Aug. 2006, T.I. Burgess (holotype MURU 445,
wards the basal end, 10.6–15.5 × 2.3–4.2 μm. cultures ex-type CBS 124583 = MUCC 666, MUCC 665 = TIN4).
Description and illustration: Perez et al. (2014). Teratosphaeria tiwiana Andjic et al., IMA Fungus 7: 260. 2016.
Typus: Australia, Victoria, Kinglake, S37°27’ E145°12’, on Diagnosis: Leaf spots circular to irregular, 3–20 mm diam, single
E. globulus in commercial plantation, Oct. 2008, G. Perez (holo- to confluent, medium brown to pale brown with red brown border
type PREM 60480, culture ex-type CBS 135621 = CMW 30745). on the top surface, pale brown at the bottom. Conidia solitary,
1–2-septate, predominantly with 1 septum, pale brown, slightly
Notes: The name T. pseudonubilosa is invalid, as it was pub- verruculose, cylindrical, straight to variously curved, thick-walled,
lished without any MycoBank number, which is a requirement for base truncate sometimes with marginal frill, apex obtuse, (26.5–)
valid publication (Art. 42, Melbourne). It is therefore validated 35–40(–44.5) × (2–)2.5–3(–3.5) μm.
here.
Description and illustration: Andjic et al. (2016).
Teratosphaeria rubida Taylor et al., as “rubidae”, Mycol. Progr.
11: 163. 2012. Typus: Australia, Northern Territory, Tiwi Island, on
E. grandis × E. urophylla hybrids, Aug. 2007, T.I. Burgess (ho-
Diagnosis: Leaf spots single to confluent circular to irregular, lotype BRIP 63496, culture ex-type CBS 141549 = BRIP 63496).
amphigenous, 3–15 mm diam, raised, corky, surrounded by dark
brown and carmine red margin. Conidia pale brown to medium Teratosphaeria toledana (Crous & Bills) Crous & U. Braun,
brown, aseptate, thick-walled, lightly verruculose, ellipsoidal, Stud. Mycol. 58: 11. 2007.
base sub-truncate and truncate apex obtuse, (11–) Basionym: Mycosphaerella toledana Crous & Bills, Stud. Mycol.
12.5–16 × (4.5–)5.5–6(–6.5) μm. 50: 208. 2004.
Synonyms: Phaeophleospora toledana Crous & Bills, Stud.
Description and illustration: Taylor et al. (2012). Mycol. 50: 208. 2004.
Typus: Australia, Western Australia, Yalgorup National Park, on Readeriella toledana (Crous & Bills) Crous & U. Braun, Stud.
C. calophylla, Aug. 2007, P.A. Barber (holotype MURU 443, Mycol. 58: 11. 2007.
culture ex-type CBS 124579 = MUCC 658); Western Australia, Kirramyces toledana (Crous & Bills) Andjic, Mycol. Res. 111:
Yalgorup National Park, on C. calophylla, Oct. 2008, P.A Barber 1192. 2007.
(isotype MURU458). Diagnosis: Leaf spots amphigenous, irregular to subcircular or
Teratosphaeria stellenboschiana (Crous) Crous, Persoonia angular, frequently confined by leaf veins, 3–6 mm diam, me-
23: 115. 2009. dium brown, with or without a red-purple border, spots aggre-
Basionym: Colletogloeopsis stellenboschiana Crous, Stud. gating with age, forming irregular blotches. Ascospores 2–3-
Mycol. 55: 110. 2006. seriate, overlapping, hyaline, guttulate, thin-walled, straight to
Synonyms: Readeriella stellenboschiana (Crous) Crous & U. slightly curved, fusoid–ellipsoid with subobtuse ends, medianly
Braun, Stud. Mycol. 58: 26. 2007. 1-septate, widest in the middle of the apical cell, not constricted
Kirramyces stellenboschiana (Crous) Andjic as “stellenbochi- at the septum, tapering towards both ends, but more prominently
ana”, Mycol. Res. 111: 1192. 2007. towards the lower end, (7–)8–10(–11) × 3(–3.5) μm. Conidia
fusoid with acutely rounded apices and truncate bases, medium
Diagnosis: Leaf spots amphigenous, circular to subcircular, brown, verruculose, aseptate, (8–)10–12(–14) × (2.5–)
0.5–3 mm diam, pale brown, with a raised border and red-purple 3–3.5(–4) μm; base with minute marginal frill.
margin. Conidia holoblastic, solitary, aseptate, ellipsoidal, with
subobtuse apex and subtruncate base with minute marginal frill, Description and illustration: Crous et al. (2004b).
CROUS ET AL.
Typus: Spain, Toledo, on Eucalyptus sp., May 2003, P.W. Crous cylindrical, straight to variously curved, thick-walled, base trun-
& G. Bills (holotype CBS 9896, culture ex-type CBS cate sometimes with marginal frill, apex obtuse, (30.5–)
113313 = CMW 14457). 47–60(–78.5) × (2–)2.5–3.5(–4) μm.
Teratosphaeria velox Crous & Summerell as “veloci”, Persoo- Description and illustration: Andjic et al. (2007c).
nia 22: 46. 2009.
Typus: Australia, Queensland, Mareeba, on E. grandis, Aug.
Diagnosis: Leaf spots amphigenous, subcircular, 1–5 mm diam, 2005, T.I. Burgess, G.E.St.J. Hardy, A.J. Carnegie & G. Pegg
pale brown with a thin, raised border and red-purple margin. (holotype BRIP 49804, culture ex-type CBS
Conidia solitary, brown, aseptate, verruculose, guttulate, ellip- 121157 = FNQ147 = MUCC 453).
soidal to subcylindrical, apex subobtuse, tapering to a sub-
Teratosphaericola Quaedvl. & Crous, Persoonia 33: 32. 2014.
truncate or truncate base (1–2 μm wide), with inconspicuous
marginal frill, (6–)8–10(–11) × (2.5–)3(–3.5) μm. Ascomata pseudothecial, solitary, black, immersed becoming
erumpent, globose; ostiole apical, central; wall of 2–3 cell layers
of medium brown textura angularis. Asci aparaphysate, fascic-
Typus: Australia, Northern Territory, ENE Pine Creek, ulate, bitunicate, subsessile, narrowly ellipsoid to subcylindrical,
S13°40’49.0’’, E131°57’04.9’’, on E. miniata, 23 Sep. 2007, B.A. slightly incurved, 8-spored. Ascospores tri- to multiseriate,
Summerell (holotype CBS H-20182, cultures ex-type CBS overlapping, hyaline to pale brown, guttulate, thin-walled, straight
124061 = CPC 14602, CPC 14600, 14601). to slightly curved, smooth to finely roughened, fusoid-ellipsoidal
with subobtuse ends, medianly 1-septate. Spermatogonia similar
Teratosphaeria verrucosa Crous & M.J. Wingf., Persoonia 22:
to the ascomata in morphology. Spermatia hyaline, smooth, rod-
46. 2009.
shaped with rounded ends.
Diagnosis: Leaf spots absent, sporulating with long black cirrhi
Type species: Teratosphaericola pseudoafricana (Crous & T.A.
from submerged pycnidia in apparently healthy, green tissue, or
Cout.) Quaedvl. & Crous
occurring in lesions that are amphigenous, raised, medium
brown, circular, up to 7 mm diam; border medium to dark brown, Notes: Teratosphaericola is best distinguished from Ter-
raised, with a red-purple margin, occurring in association with atosphaeria based on DNA sequence data. The genus is
T. juvenalis. Conidia ellipsoidal, apex subobtuse, base truncate to monotypic, and nothing is known about its biology.
subtruncate, generally widest at or below the median, thick-
Teratosphaericola pseudoafricana (Crous & T.A. Cout.)
walled, verrucose, (7–)8–10(–15) × (4–)5(–6) μm in vitro,
Quaedvl. & Crous, Persoonia 33: 32. 2014. Fig. 108.
(6–)7–9(–10) × (4–)5(–6) μm in vivo.
Basionym: Mycosphaerella pseudafricana Crous & T.A. Cout.,
Description and illustration: Crous et al. (2009a). Stud. Mycol. 55: 115. 2006.
Synonym: Teratosphaeria pseudafricana (Crous & T.A. Cout.)
Typus: South Africa, Western Cape Province, Stellenbosch,
Stellenbosch Mountain, on E. cladocalyx, Apr. 1988, P.W. Crous
(holotype CBS H-20183, culture ex-type CBS 113621 = CPC 42). Diagnosis: Leaf spots amphigenous, irregular to sub-circular,
2–7 mm diam, medium brown, surrounded by a thin, raised,
Teratosphaeria viscida (Andjic et al.) Andjic et al. as “viscidus”, concolourous border. Ascospores tri- to multiseriate, overlapping,
Persoonia 23: 115. 2009. Fig. 106. hyaline to pale brown, guttulate, thin-walled, straight to slightly
Basionym: Kirramyces viscidus Andjic et al., Australas. Pl. Path. curved, smooth to finely roughened, fusoid-ellipsoidal with sub-
36: 485. 2007. obtuse ends, medianly 1-septate, widest in the middle of the
apical cell, constricted at the septum, tapering towards both
Diagnosis: Leaf spots circular to irregular, 3–20 mm diam, single ends, but more prominently towards the lower end, (8–)
to confluent, pale to medium brown with red brown border on the 9–10(–11) × (2.5–)3 μm in vivo.
top surface, pale brown at the bottom. Conidia solitary, 0–3-
septate, subhyaline to pale brown, slightly verruculose, Description and illustration: Crous et al. (2006d).
Fig. 108. Teratosphaericola pseudoafricana (PREM 54973). A. Leaf spot. B–D. Asci with ascospores. E. Germinating ascospores. Scale bars = 10 μm.
276
Fig. 109. Thyrinula eucalypti. A–E. Disease symptoms. F, J. Vertical section through ascomata. G, H. Asci. I. Hysterothecia on lesion surface. K. Ascospores. L. Section
through pycnidium showing needle-like conidia. Scale bars = 10 μm.
Typus: Zambia, on E. globulus, Aug. 1995, T. Coutinho (holo- Ascomata black, superficial, hysterothecial, elongate, occa-
type PREM 54973, cultures ex-type CBS 114782 = CPC 1230, sionally Y-shaped, opening by a longitudinal slit; margin without
CPC 1229, 1231). restricted brown, superficial hyphae. Asci bitunicate with ocular
chamber, broadly ellipsoid to somewhat obovoid, containing 8
Thyrinula Petr. & Syd., Annls Mycol. 22: 373. 1924. Figs 109,
ascospores, intermixed among hyphal-like, septate, branched
110.
pseudoparaphyses. Ascospores 1-septate, constricted at
Synonym: Alysidiella Crous, Fungal Diversity 23: 325. 2006.
septum, ends obtusely rounded, widest just above septum,
Spermatogonia pycnidial, pimple-like, scutiform, black, super- guttulate, hyaline, smooth (becoming brown and verruculose
ficial, glabrous. Spermatophores reduced to spermatogenous with a mucoid sheath at germination (5–8 μm diam), with
cells, small, hyaline, phialidic. Spermatia solitary, hyaline, solitary germ tube growing characteristically down into agar
smooth, filiform-cylindrical, aseptate, appearing to have a medium).
spermatial-like function, as they do not germinate in culture.
Type species: Thyrinula eucalyptina Petr. & Syd.
Conidiomata sporodochial or as solitary, erect conidiophores on
lesions, consisting of brown, verrucose, thick-walled, branched, Symptoms: “Target spot or corky spot” leaf spots are roughly
septate, hyphae. Conidiogenous cells holoblastic, scars indis- circular, dark brown, corky, raised, necrotic, 1–6 mm diam;
tinct to thickened along the rim, not darkened nor refractive. distinctive in only developing part-way through the leaf lamina,
Setae and hyphopodia absent. Conidia dry, in branched or occurring with sporocarps on the lower or upper leaf surface;
simple acropetal chains, ellipsoidal to subcylindrical, medium margins indistinct or chlorotic to red-purple. Lesions also occur
brown, thick-walled, verruculose, aseptate to multiseptate. on leaf petioles, small branches and on stems. Initially black,
CROUS ET AL.
Fig. 110. Thyrinula spp. A–C. T. dunnii (CPC 12977). A. Leaf spot with sporocarps. B. Hysterothecium viewed from above. C. Asci. D–G. T. eucalypti (CPC 12986). D. Leaf spot with
sporocarps. E. Hysterothecium viewed from above. F. Asci. G. Spermatogonium. H–J. T. eucalyptina (CPC 35990). H. Leaf spot with sporocarps. I. Asci. J. Spermatogonium oozing
spermatia. K–M. T. eucalyptina ((K-M 262956). K. Leaf spot with sporocarps. L. Transverse section through hysterothecium. M. Asci. N–Q. T. eucalypti (K-M 262955). N. Leaf spot
with sporocarps. O. Immature asci. P, Q. Spermatogenous cells with spermatia. Scale bars: D = 400 μm, A, H, K, N = 200 μm, G = 80 μm, all others = 10 μm.
278
pimple-like spermatogonia, followed by elongate, occasionally Diagnosis: Leaf spots brown, not extending through leaf surface,
Y-shaped, black hysterothecia opening by a longitudinal slit, are circular, containing both hysterothecia and pycnidia of sper-
formed on the lesion surface and are easily removed with a matogonial morph. Hysterothecia 300–400 × 100 μm; asci
needle. Distinctive 2-celled, guttulate, ascospores are formed in 18–28 × 8–9 μm, ascospores 8–10 × 3–4 μm.
bitunicate asci; ascospores with one cell slightly broader and
Typus: Australia, Victoria, on fading leaves of Eucalyptus sp.,
shorter than the other. The spermatogonia have a basal layer of
Mrs Martin No. 682 (lectotype of Leptostromella eucalypti K-M
phialidic spermatogenous cells that form a mass of needle-like
262955, designated by Swart 1988); Queensland, Imbil, Imbil
spores; all three morphs can occur on the same leaf spot.
State Forest, S26°28’28.3” E152°36’28.3”, on E. cloeziana, Jan.
Notes: The nomenclature and taxonomy of the fungus previ- 2005, A.J. Carnegie, AC0512 (epitype designated here CBS H-
ously known as “Aulographina eucalypti” is confused. The 24026, MBT388173, culture ex-epitype CBS 145894 = CPC
sexual morph was discussed by Müller & von Arx (1962), and 12986).
the spermatogonial morph by Petrak & Sydow (1924) and
Notes: The holotype of this species was annotated by H.J. Swart in
Swart (1988). This pathogen, which causes target spot disease
1975 as “probably identical to the fungus associated with Aulog-
on eucalypts, has had several names applied to it in the past.
raphina eucalypti on E. obliqua in Australia, and might be the
For many years it was treated as Aulographina eucalypti
spermatial state”. The latter view was subsequently show to be
(Cooke & Massee) Arx & E. Müll. However, the genus Aulog-
correct (Wall & Keane 1984, Carnegie & Keane 2003). Leaf spots
raphina is based on A. pinorum occurring on Pinus spp. in
(3–6 mm diam) are confined to one side of the leaf, circular, dark
Europe. It is not congeneric with the eucalypt fungus, as it lacks
brown, raised with margin and some dark brown hyphae seen
the typical spermatogonial and asexual morphs observed on
radiating outward across the leaf surface. The holotype also
Eucalyptus. Furthermore, the cultures that are available (CBS
contains pycnidia of Phaeothyriolum sp., and Blastacervulus
174.90, 302.71, 655.86) cluster with Catenulostroma (Ter-
eucalypti in adjacent leaf spots. Furthermore, some hysterothecia
atosphaeriaceae) and Venturiaceae (in the case of CBS
were observed (asci 20–27 × 8–9 μm, ascospores
655.86). Müller & von Arx (1962) considered why the name
8–9 × 3–4 μm), allowing us to link this name to fresh collections.
Aulographina was best suited (over Aulographum, Leptos-
tromella and Lembosiopsis), but they ignored the asexual Thyrinula eucalyptina Petr. & Syd., Annls Mycol. 22: 373. 1924.
names. Given the fact that Aulographina is no longer available Fig. 110.
for this fungus, the holomorph is best treated under the name Synonyms: Lembosiopsis eucalyptina Petr. & Syd., Annls Mycol.
introduced for the spermatogonial morph, Thyrinula euca- 22: 372. 1924.
lyptina, as this species is also the type of the genus Thyrinula Aulographum eucalypti Cooke & Massee, Grevillea 18(no. 85): 6.
(Petrak & Sydow 1924). For more details relating to its biology 1889.
and host range, see Wall & Keane (1984) and Carnegie & Aulographina eucalypti (Cooke & Massee) Arx & E. Müll.,
Keane (2003). Isolates established from single ascospores Sydowia 14: 332. 1960.
(on PDA and OA) produce both the spermatogonial and Lembosiopsis australiensis Hansf., Proc. Linn. Soc. N.S.W. 79:
ascomatal morphs in older cultures. 105. 1954.
Thyrinula dunnii Crous & Carnegie, sp. nov. MycoBank Diagnosis: Leaf spots brown, not extending through leaf surface,
MB832053. Fig. 110. circular, containing both hysterothecia and pycnidia of sper-
matogonial morph. Hysterothecia 300–400 × 100–180 μm; asci
Etymology: Name reflects the host species Eucalyptus dunnii
33–40 × 11–14 μm, ascospores (9–)12–13 × (3.5–)4–5 μm.
which it was collected from.
Descriptions and illustrations: Petrak & Sydow (1924), Müller &
Diagnosis: Leaf spots brown, not extending through leaf surface,
von Arx (1962), Swart (1988).
circular, containing both hysterothecia and pycnidia of sper-
matogonial morph, as well as hyphomycetous asexual morph. Typus: Australia, Victoria, Lilydale, on Eucalyptus sp., 24 Mar.
Hysterothecia 300–400 × 100–120 μm; asci 23–28 × 8–11 μm, 1886, Mrs Martin No. 444 (holotype of Aulographum eucalypti
ascospores 9–12 × 3.5–5 μm. K-M 262956); South Australia, Cape Jervis Peninsula, on
Typus: Australia, New South Wales, Nana Glen, Wedding Bells Eucalyptus cosmophylla, Jan. 1924, G. Samuel (holotype of
State Forest, Crabtree plantation, S30°8’45.6” E153°6’21.8”, on Lembosiopsis australiensis WARI 2070, also slide IMI 72980).
E. dunnii, Jan. 2005, A.J. Carnegie, AC0398 (holotype CBS H- South Africa, Mpumalanga Province, Vosman’s Beacon, Forest
24046, culture ex-type CBS 145893 = CPC 12977). Jessievale, on E. gigantea, 18 May 1923, leg. District Forest
Officer (lectotype of Lembosiopsis eucalyptina PREM 17266,
Notes: Asci and ascospores are somewhat intermediate to those selected by Doidge, Bothalia 5: 198. 1950; selected here as
of T. eucalypti and T. eucalyptina. Phylogenetically, this species lectotype of Thyrinula eucalyptina, sexual morph on same
is distinct from other Thyrinula species currently known from material MBT388174); Limpopo Province, Tzaneen, on Euca-
sequence data (Fig. 12). Some intraspecific variation exists for lyptus sp., 22 Jul. 2018, P.W. Crous, HPC 2467 (epitype of
this species, but we refrain from introducing cryptic species given Thyrinula eucalyptina designated here CBS H-24027,
the low internal support values in this clade. MBT388175, culture ex-epitype CBS 145897 = CPC 35990).
Thyrinula eucalypti (Cooke & Massee) H.J. Swart, Trans. Brit. Notes: Examination of the holotype of Aulographum eucalypti
Mycol. Soc. 90: 286. 1988. Fig. 110. found asci to be 27–30 × 9–13 μm, and ascospores
Basionym: Leptostromella eucalypti Cooke & Massee, Grevillea 10–12.5 × 4 μm. Ascospores of Thyrinula eucalyptina (as
19(no. 92): 91. 1891. Lembosiopsis eucalyptina) are cited as 9–12.5 × 4–5 μm
CROUS ET AL.
(Petrak & Sydow 1924), and those of Lembosiopsis australiensis Diagnosis: Conidiophores 1–2-septate, 10–15 × 5–7 μm.
as 10–13 × 3.5–4.5 μm (Hansford 1954). A freshly collected Conidiogenous cells 5–7 × 5–6 μm. Conidia subcylindrical to
specimen from South Africa (HPC 2467) proved similar in ellipsoidal, 0–6-septate, 10–35 × 5–7 μm.
morphology, asci being 33–40 × 11–14 μm, ascospores (9–)
12–13 × (3.5–)4–5 μm, and is herewith designated as epitype.
Typus: Uruguay, La Turrita, on E. dunnii, Jan. 2005, M.J.
Thyrinula parasitica (Crous) Crous, comb. nov. MycoBank
Wingfield (holotype CBS H-19766, cultures ex-type CBS
MB832054.
120122 = CPC 12111, CPC 12112, 12113).
Basionym: Alysidiella parasitica Crous, Fungal Diversity 23: 326.
2006. Notes: Because the epithet “eucalypti” is occupied, a new name
is introduced for this species. Although the sexual morph was not
Diagnosis: Conidiogenous cells 4–13 × 4–6 μm. Conidia dry, in
observed, one collection from Colombia (CPC 31921) was ob-
branched or simple acropetal chains, ellipsoidal to subcylindrical,
tained as single ascospores that were ejected onto the agar
0–13-septate, 8–30 × 5–7 μm.
surface showing that a sexual morph exists for this species.
Description and illustration: Summerell et al. (2006). Phylogenetically, this species is distinct from other Thyrinula
species currently known from sequence data (Fig. 12).
Mountain, on Eucalyptus sp., Jan. 2006, P.W. Crous (holotype Tracylla (Sacc.) Tassi, Bulletin Labor. Orto Bot. de R. Univ. Siena
CBS H-19742, cultures ex-type CBS 120088 = CPC 12835, CPC 6: 62. 1904.
12836, 12837).
Conidiomata pycnothyrial, superficial, rounded to oval or irreg-
Notes: One collection from Colombia (CPC 36157) was obtained ular in outline; upper shield connected to the immersed mycelium
as single ascospores, although the sexual morph could not be by a central supporting column of cells; central zone of the shield
located on the leaves. Phylogenetically, this species is distinct of textura angularis in several cell layers; peripheral zone of the
from other Thyrinula species currently known from sequence shield of textura prismatica with radially elongated cells; margin
data (Fig. 12). Some intraspecific variation exists for this species, of the shield entire or invaginated; brown to dark brown. Co-
but we refrain from introducing cryptic species pending the nidiophores reduced to conidiogenous cells, lining the concavity
collection of a broader global sampling of the fungus and its formed by the pycnothyrial shield and the central column of
relatives. supporting cells, invested in mucus. Conidiogenous cells
discrete, almost colourless to colourless, smooth, with visible
Thyrinula uruguayensis Crous, nom. nov. MycoBank
periclinal thickening. Conidia naviculate to lunate or broadly
MB832055.
ellipsoidal, unicellular, hyaline, smooth, bearing one or two ap-
Basionym: Heteroconium eucalypti Crous & M.J. Wingf., Fungal
pendages, arising as cellular extensions of the conidium body
Planet 10: 1. 2006.
and not separated from it by septa; appendages polar, un-
Synonym: Alysidiella eucalypti (Crous & M.J. Wingf.) Cheew. &
branched, attenuated, flexuous (from Nag Raj 1993).
Crous, Mycol. Progr. 11: 63. 2012.
Etymology: Name reflects the country where this species was
Type species: Tracylla spartinae (Peck) Tassi
collected, Uruguay.
Fig. 111. Tracylla aristata (CBS 141404). A. Sporulation on PDA. B. Conidiophores. C, D. Conidiogenous cells. E, F. Conidiogenous cells giving rise to conidia. G. Conidia.
Scale bars 10 μm (from Hernandez-Restrepo et al. 2016).
280
Note: The genus Tracylla is regarded to be of minor importance, media, and once germinating, resemble germinating ascospores,
containing species that are saprobic or weakly pathogenic. which probably explains the original confusion. Unfortunately,
living cultures of U. juvenis are no longer available and the fungus
Tracylla aristata (Cooke) Tassi, Bulletin Labor. Orto Bot. de R. must be recollected to resolve its relationship to other species of
Univ. Siena 6: 62. 1904. Fig. 111. Uwebraunia. Mycosphaerella juvenis has since been reduced to
Basionym: Leptothyrium aristatum Cooke, Grevillea 20(no. 93): synonymy with Teratosphaeria nubilosa (Crous et al. 2004b).
6. 1891. Li et al. (2011) showed that Uwebraunia and Dissoconium are
sister genera within Dissoconiaceae. Species of Dissoconium
Diagnosis: Commonly found on dead leaf tips of leaf litter;
commonly co-occur with species of Teratosphaeria on the same
regarded as of minor importance, probably saprobic or weakly
leaf spots. However, Jackson et al. (2004) demonstrated that
pathogenic. Conidiomata black pycnothyria. Conidia exuded in
D. dekkeri can infect E. globulus leaves, and that it was not a
slimy mass, naviculate to lunate or subcylindrical,
hyperparasite of T. cryptica or T. nubilosa. Nothing is known
13–17 × 2–3 μm, base obtuse, apex acute, giving rise to flex-
regarding the biology of other species of Dissoconium or their
uous apical appendage 6.5–14 μm long.
possible role as plant pathogens.
Description and illustration: Hernandez-Restrepo et al. (2016).
Uwebraunia australiensis (Crous & Summerell) Crous, Per-
Typus: Australia, Victoria, on Eucalyptus sp., Mrs. Martin No. soonia 28: 123. 2012.
752 (holotype K-M); Victoria, Toolangi State Forest, Basionym: Dissoconium australiensis Crous & Summerell,
S37°33’25.3” E145°31’55.9”, on E. regnans, 9 Nov. 2014, P.W. Fungal Diversity 26: 156. 2007.
Crous, J. Edwards & P.W.J. Taylor (epitype CBS H-22654,
Diagnosis: Conidia (20–)23–25(–27) × (3–)4(–5) μm, solitary,
culture ex-epitype CBS 141404 = CPC 25500).
pale olivaceous-brown, smooth, ellipsoid to obclavate, 1-septate,
Tracylla eucalypti Crous, Persoonia 40: 365. 2018. apex obtuse, base obconic-truncate, hilum unthickened,
1–1.5 μm wide.
Diagnosis: Pycnothyria, brown, round, occurring on leaf litter.
Conidia solitary, hyaline, aseptate, smooth, guttulate, falcate, Description and illustration: Crous et al. (2007c).
apex subobtusely rounded, base truncate, 1–1.5 μm diam, (12–)
17–19(–20) × (2.5–)3 μm. Typus: Australia, Queensland, Cairns, near Kuranda,
S16°560 23.300 , E145°320 34.600 , on E. platyphylla, 26 Aug. 2006,
Typus: Colombia, Cali, on E. urophylla, Jul. 2010, M.J. Wingfield 120729 = CPC 13282).
Uwebraunia commune (Crous & Mansilla) Crous, Persoonia
31806).
28: 123. 2012.
Uwebraunia Crous & M.J. Wingf., Mycologia 88: 446. 1996. Basionym: Dissoconium commune Crous & Mansilla, Stud.
Mycol. 50: 203. 2004.
Ascomata pseudothecial, immersed, globose, unilocular, papil-
Synonym: Mycosphaerella communis Crous & Mansilla, Stud.
late, ostiolate, canal periphysate; wall consisting of 3–4 layers of
Mycol. 50: 203. 2004.
brown textura angularis; inner layer of flattened, hyaline cells.
Pseudoparaphyses absent. Asci fasciculate, 8-spored, bituni- Diagnosis: Leaf spots amphigenous, sub-circular to circular,
cate. Ascospores ellipsoid-fusoid, 1-septate, hyaline, with or 4–12 mm diam, medium brown, surrounded by a thin, raised,
without mucoid sheath. Mycelium internal and external, con- concolourous border. Ascospores 2–3-seriate, overlapping, hy-
sisting of branched, septate, smooth, hyaline to pale brown hyaline, guttulate, thick-walled, straight to slightly curved, obovoid
phae, that anastomose, forming nets in culture. Conidiophores with subobtuse ends, medianly or unequally 1-septate, widest in
separate, arising from hyphae, subcylindrical, subulate or lage- middle of apical cell, or close to the apex of the apical cell,
niform to cylindrical, tapering to a bluntly rounded or truncate constricted at the septum, tapering towards both ends, but more
apex, straight to once geniculate, smooth, medium brown, 0–2- prominently towards the lower end, (12–)13–15(–17) × (3.5–)
septate; loci terminal and lateral, visible as slightly thickened, 4–4.5 μm in vivo. Conidia terminal, pale olivaceous, smooth,
darkened scars on a rachis; proliferation sympodial but also obclavate with obtuse apex and obconic-truncate base, 0–1-
appearing to be percurrent. Primary conidia solitary, pale septate, constricted at the septum, straight or curved,
olivaceous-brown, smooth, ellipsoid to obclavate, 1-septate; hila 20–30 × 4–5 μm (avg. 25 × 4.5 μm); hila inconspicuous.
somewhat darkened. Secondary conidia developing adjacent to
primary conidia, pale olivaceous to subhyaline, aseptate, smooth,
pyriform; conidium discharge active, usually with both conidial Typus: Spain, Pontevedra, Lourizan, Areeiro, on E. globulus, Dec.
types being discharged simultaneously. One or more secondary 2002, J.P. Mansilla (holotype of M. communis and D. commune
conidia anastomosing with primary conidium once discharged. CBS H-9900, culture ex-type CBS 114238 = CPC 10440).
Colonies not forming yellow pigment, nor sclerotia in culture.
Uwebraunia dekkeri (de Hoog & Hijwegen) Crous, Persoonia
Type species: Uwebraunia juvenis Crous & M.J. Wingf. 28: 123. 2012. Fig. 112.
Basionym: Dissoconium dekkeri de Hoog & Hijwegen, Mycol.
Symptoms: Associated with amphigenous, pale brown leaf spots;
Res. 95: 679. 1991.
mostly co-occurring with species of Teratosphaeria.
Synonyms: Uwebraunia lateralis Crous & M.J. Wingf., Mycologia
Notes: The proposed connection between Uwebraunia juvenis 88: 454. 1996.
and Mycosphaerella juvenis was incorrect (Crous & Wingfield Mycosphaerella lateralis Crous & M.J. Wingf., Mycologia 88: 454.
1996). Conidia of Uwebraunia actively discharge onto agar 1996.
CROUS ET AL.
Fig. 112. Uwebraunia dekkeri. A, B. Disease symptoms. C. Close-up of leaf spot with immersed pseudothecia. D, E. Asci. F. Ascus and spermatia. G. Germinating ascospore.
H. Colony on PDA. I. Colony shooting conidia on SNA. J. Typical conidial germinating pattern on MEA. K–N. Primary conidia, with smaller, obovoid secondary conidia. Scale
bars = 10 μm.
Mycosphaerella shimabarensis H.C. Evans & P.F. Cannon, Ascospores multiseriate, overlapping, hyaline, guttulate, thin-
Mycoscience 50: 187. 2009. walled, straight or slightly curved, fusoid-ellipsoidal with an
obtuse apex, widest in middle of apical cell, medianly 1-septate,
not constricted at septum, tapering toward both ends, (7–)
diam, grey-brown, surrounded by raised borders, medium brown
8–14(–16) × 2–2.5(–3) μm. Conidia terminal, solitary, pale
on the adaxial surfaces concolourous on the lower surfaces.
282
olivaceous, smooth, obclavate, apex obtuse, base obconic- prominent taper towards lower end, mucilaginous sheath visible
truncate, medianly 1-septate, (15–)17–21(–35) × (2–) around spore. Ascospore germination from both ends in two
3.5–4(–4.5) μm; hilum subtruncate, unthickened, not darkened patterns (remaining hyaline): Type I (Crous 1998), growing parallel
or refractive, flattened, 1.5–2 μm wide. to the long axis of the spore, with lateral branches parallel or
perpendicular to the long axis of spore, irregular in width, con-
Descriptions and illustrations: Crous & Wingfield (1996), Crous
stricted at the median septum of the spore, slightly distorting; Type
(1998).
B (Crous 1998), germ tube growing parallel to the long axis of the
Material examined: South Africa, Northern Province, Tzaneen, Magoebaskloof, spore, regular in width, not distorting or becoming constricted at
on E. grandis × E. saligna, Oct. 1994, G. Kemp (holotype of U. lateralis PREM septum. Spermatogonia, when present, amphigenous, dark
brown, subepidermal to erumpent, globose to subglobose. Sper-
Uwebraunia juvenis Crous & M.J. Wingf., Mycologia 88: 446. matia hyaline, smooth, rod-shaped, with obtuse ends.
1996.
Type species: Virosphaerella pseudomarksii (Cheewangkoon
Diagnosis: Conidia terminal, solitary, pale olivaceous, smooth, et al.) Videira & Crous (≡ Mycosphaerella irregularis Chee-
obclavate with obtuse apex and obconic-truncate base, 1- wangkoon et al.)
septate, becoming prominently constricted at septum when
Note: Although species of Virosphaerella are associated with leaf
mature, straight or curved, (25–)26–30(–40) × (4–)
spots on eucalypt leaves, nothing is known regarding their
4.5–5.5(–6) μm; hilum not thickened, not darkened or refractive,
1.5–2 μm wide. biology or role as pathogens.
Virosphaerella irregularis (Cheewangkoon et al.) Videira &
Descriptions and illustrations: Crous & Wingfield (1996), Crous
Crous, Stud. Mycol. 87: 378. 2017. Fig. 113.
(1998).
Basionym: Mycosphaerella irregularis Cheewangkoon et al. (as
Typus: South Africa, Kwazula-Natal Province, Pietermaritzburg, ‘irregulari’), Persoonia 21: 83. 2008.
on E. nitens, Jan. 1995, M.J. Wingfield (holotype PREM 51915). Synonym: Paramycosphaerella irregularis (Cheewangkoon
et al.) Guatimosim et al., Persoonia 37: 127. 2016.
Virosphaerella Videira & Crous, Stud. Mycol. 87: 377. 2017.
Ascomata amphigenous or epiphyllous, black, subepidermal to Diagnosis: Leaf spots amphigenous, subcircular to oval, pale
erumpent, ovoid, globose or subglobose, apical ostiole, wall brown with grey centres, 5–12 mm diam, surrounded by a thin,
consisting of 2–3 layers of medium brown textura angularis. Asci medium brown margin. Ascospores bi- to triseriate, overlapping,
aparaphysate, fasciculate, subsessile, subcylindrical to narrowly hyaline, guttulate, thin-walled, straight to slightly curved, fusoid-
obovoid, straight to slightly curved, 8-spored. Ascospores bi- to ellipsoidal with obtuse ends, widest just above the septum, or
triseriate, overlapping, hyaline, guttulate, thin-walled, straight to in the middle of the apical cell, medianly 1-septate or slightly
slightly curved, fusoid, fusoid-ellipsoidal with obtuse ends, medi- longer in the basal cell, slightly constricted at septum, tapering
anly 1-septate or slightly longer in the basal cell, slightly con- toward both ends, but with more prominent taper towards
stricted at septum, widest just above the septum, or in the middle lower end, at times with a mucous-like coating, (8–)
of the apical cell, tapering toward both ends, but with more 9–11(–13) × 2.5–3(–3.5) μm.
Fig. 113. Virosphaerella spp. A–C. V. irregularis. A. Leaf spot. B, C. Section through ascomata. L. Germinating ascospores. N–K. V. pseudomarksii. D–F. Asci. G–J.
Ascospores. K, L. Germinating ascospores. Scale bars: B, C = 100 μm, all others = 10 μm.
CROUS ET AL.
Description and illustration: Cheewangkoon et al. (2008). Description and illustration: Carnegie et al. (2011).
Typus: Thailand, Udonthani, on Eucalyptus sp., Jul. 2007, R. Typus: Australia, Queensland, Walkamin, on E. alba, 11 Nov.
Cheewangkoon (holotype CBS H-20135, culture ex-type CBS 2008, A.J. Carnegie, M. Ramsden & G.S. Pegg (holotype BRIP
123242 = CPC 15408); ibid., cultures CPC 15431, 15432. 52586, culture ex-type BRIP 52586 = CBS 130521).
Virosphaerella pseudomarksii (Cheewangkoon et al.) Videira Notes: Based on a megablast search of NCBI’s GenBank
& Crous, Stud. Mycol. 87: 378. 2017. Fig. 113. nucleotide database, the closest hits using the ITS sequence
Basionym: Mycosphaerella pseudomarksii Cheewangkoon et al., had highest similarity to Hyalozasmidium aerohyalinosporum
Persoonia 21: 83. 2008. [GenBank NR_156220.1; Identities = 497/514 (97 %), 7 gaps
Synonym: Paramycosphaerella pseudomarksii (Cheewangkoon (1 %)], Hyalozasmidium sideroxyli [GenBank NR_156372.1;
et al.) Guatimosim et al., Persoonia 37: 127. 2016. Identities = 423/446 (95 %), 5 gaps (1 %)], and Madagascar-
omyces intermedius [GenBank MH863356.1; Identities = 483/
Diagnosis: Ascospores bi- to triseriate overlapping, hyaline,
516 (94 %), 10 gaps (1 %)]. Closest hits using the LSU
guttulate, thin-walled, straight to slightly curved, fusoid with
sequence are Paramycosphaerella marksii [GenBank
obtuse ends, widest in the middle of the asymmetrical apical cell,
DQ204758.1; Identities = 815/822 (99 %), 1 gap (0 %)], Para-
medianly 1-septate or with slightly longer basal cell; tapering
mycosphaerella intermedia [GenBank NG_059428.1; Identi-
toward both ends, but with more prominent taper towards lower
ties = 813/822 (99 %), 1 gap (0 %)], and Hyalozasmidium
end, (12–)14–17(–18.5) × (2.5–)3(–3.5) μm.
aerohyalinosporum [GenBank NG_059440.1; Identities = 812/
Description and illustration: Cheewangkoon et al. (2008). 822 (99 %), 1 gap (0 %)]. Closest hits using the rpb2 sequence
had highest similarity to Hyalozasmidium sideroxyli [GenBank
Typus: Thailand, Chiang Mai, Mae Tang, on Eucalyptus sp., Jun.
MF951505.1; Identities = 668/805 (83 %), 10 gaps (1 %)],
2007, R. Cheewangkoon (holotype CBS H-20134, culture ex-
Paramycosphaerella marksii [GenBank MF951573.1; Identi-
type CBS 123241 = CPC 15410); ibid., cultures CPC 15435,
ties = 624/774 (81 %), 4 gaps (0 %)], and Paramycosphaerella
15436.
intermedia [GenBank MF951568.1; Identities = 623/774 (80 %),
Walkaminomyces Crous & Carnegie, gen. nov. MycoBank 4 gaps (0 %)].
MB832059.
Xenomycosphaerella Quaedvl. & Crous, Persoonia 33: 24. 2014.
Etymology: Named after the town where it was collected in
Ascomata pseudothecial, dark brown, subepidermal to erum-
Australia, Walkamin.
pent, globose, with an apical ostiole; wall of 2–3 layers of me-
Leaf spots amphigenous, irregular, small (5 mm diam) to large dium brown textura angularis. Asci aparaphysate, fasciculate,
spreading blights with irregular margins. Ascomata pseudothe- bitunicate, subsessile, obovoid to broadly ellipsoidal, straight to
cial, amphigenous, immersed, becoming erumpent, black, slightly curved, 8-spored. Ascospores bi- to multiseriate, over-
globose. Asci aparaphysate, fasciculate, bitunicate, subsessile, lapping, hyaline, thin- or thick-walled, straight to slightly curved,
obclavate to ellipsoidal, straight to incurved, 8-spored. Asco- fusoid-ellipsoidal with obtuse ends, widest in middle of the apical
spores multiseriate, hyaline, guttulate, straight, fusoid-ellipsoidal cell, medianly or unequally 1-septate, tapering towards both
with obtuse ends, medianly 1-septate. ends, but more prominently towards the lower end.
Type species: Walkaminomyces medusae (Carnegie & G.S. Type species: Xenomycosphaerella elongata (Crous & M.J.
Pegg) Crous & Carnegie Wingf.) Quaedvlieg & Crous
Notes: Based on DNA phylogenetic data, Mycosphaerella Symptoms: Leaf spots amphigenous, spreading inward from the
medusae represents a new genus in the Mycosphaerellaceae, leaf margins, irregular to subcircular, medium brown, surrounded
for which the name Walkaminomyces is introduced. Walkami- by thick, dark brown border.
nomyces is presently monotypic, known from a single species
collected on leaves of E. alba in Queensland, where it Notes: Xenomycosphaerella is morphologically similar to
was associated with prominent leaf spotting. This taxon is Mycosphaerella s. lat., and best distinguished based on DNA
characterised by having a distinct germination pattern, germi- data. Xenomycosphaerella elongata is considered to be a
nating with 4–6 snake-like germ tubes per ascospore (Carnegie pathogen of minor importance.
et al. 2011).
Xenomycosphaerella elongata (Crous & M.J. Wingf.)
Walkaminomyces medusae (Carnegie & G.S. Pegg) Crous & Quaedvlieg & Crous, Persoonia 33: 24. 2014. Fig. 114.
Carnegie, comb. nov. MycoBank MB832057. Basionym: Mycosphaerella elongata Crous & M.J. Wingf.,
Basionym: Mycosphaerella medusae Carnegie & G.S. Pegg, Fungal Diversity 26: 163. 2007.
Australas. Pl. Path. 40: 376. 2011.
Diagnosis: Leaf spots amphigenous, irregular, small (5 mm diam) 3–13 mm diam, medium brown, with a thin, raised, dark brown to
to large spreading blights with irregular margins, greyish brown red-brown border. Ascospores germinate from both ends, with
to reddish brown with a prominent purple to purplish green germ tubes parallel to the long axis of the spore; spore not
diffuse border. Ascospores multiseriate, overlapping, hyaline, darkening, nor distorting, becoming up to 5 μm wide.
guttulate, straight, fusoid-ellipsoidal with obtuse ends, obviously
widest in middle of apical cell, medianly 1-septate, not con- Description and illustration: Crous et al. (2007c).
stricted at septum or slightly so, tapered to both ends but more Typus: Venezuela, El Pi~nal Lotes farm near Acarigua, on
prominently to apical end, 10–12 × 3–4 μm. E. camaldulensis × E. urophylla, Oct. 2006, M.J. Wingfield
284
Fig. 114. Xenomycosphaerella elongata (CBS H-19824). A. Leaf spot. B–D. Asci and ascospores. E–G. Germinating ascospores. Scale bars = 10 μm (from Crous et al.
2007c).
(holotype CBS H-19824, cultures ex-type CBS 120735 = CPC angularis. Asci fasciculate, bitunicate, subsessile, hyaline,
13378, CPC 13379, 13380). smooth, 8-spored, obovoid to ellipsoid, aparaphysate, straight to
slightly curved. Pseudoparaphyses absent. Ascospores tri- to
Xenosonderhenia Crous, Persoonia 28: 175. 2012.
multiseriate, hyaline, smooth, fusoid-ellipsoid, widest in apical
Conidiomata pycnidial, black, globose, substomatal, erumpent, cell, one third from apex, tapering towards both ends, not
with central ostiole, lined with periphyses; wall of 2–3 layers of constricted at median septum.
brown textura angularis. Conidiophores reduced to con-
idiogenous cells. Conidiogenous cells lining the inner cavity, Type species: Xenosonderhenia syzygii Crous
subcylindrical to doliiform; finely verruculose, pale brown, Note: Xenosonderhenia represents a distinct genus in Mycos-
proliferating apically with several percurrent proliferations. phaerellaceae that is associated with leaf spots on Myrtaceae
Conidia subcylindrical, brown, finely verruculose, apex obtuse, (Eucalyptus, Syzygium).
base truncate with visible scar, euseptate, but septa with visible
central pore. Conidia of synasexual morph intermingled in same Xenosonderhenia eucalypti Crous & M.J. Wingf., Persoonia
conidioma, but conidiogenous cells proliferating percurrently or 33: 241. 2014. Fig. 115.
sympodially; conidia hyaline to subhyaline, narrowly obclavate, Diagnosis: Leaf spots amphigenous, dark brown, 10–20 mm
apex subobtuse, base truncate, straight to curved, transversely diam, with dark brown border. Co-occurring on leaf spots with
multi-septate. Synasexual morph also hyphomycetous, devel- Zasmidium eucalyptigenum. Ascospores tri- to multiseriate, hy-
oping in aerial mycelium; conidiophores subcylindrical, straight aline, smooth, fusoid-ellipsoid, widest in apical cell, one third
to curved, septate, hyaline to subhyaline, proliferating sympo- from apex, tapering towards both ends, not constricted at median
dially at apex. Conidiophores solitary or fasciculate or forming septum, (17–)18–20(–22) × (3–)4 μm.
on a reduced stroma. Ascomata black, erumpent, globose,
solitary, with central ostiole; wall of 2–3 layers of brown textura Description and illustration: Crous et al. (2014c).
CROUS ET AL.
Fig. 115. Xenosonderhenia eucalypti (CPC 24247). A. Disease symptoms. B. Leaf spot. C–E. Asci with ascospores. F. Ascospores. G. Germinating ascospores. Scale
bar = 10 μm.
Typus: Mozambique, Forestas de Niassa, on E. urophylla, 2 chain, hyaline to subhyaline, smooth, oblong, cylindrical to long-
Feb. 2014, M.J. Wingfield (holotype CBS H-21991, culture ex- obclavate, base medium-long obconically truncate, apex rounded,
type CBS 138858 = CPC 24247). 15–50 × 5–6 μm, 0–4-septate, eu- or distosepta, sometimes
slightly constricted at the septa, hila slightly thickened and darkened.
Xenosonderhenioides Videira & Crous, Stud. Mycol. 87: 368.
2017. Description and illustration: Videira et al. (2017).
Mycelium composed of hyaline to pale brown hyphae, smooth, Typus: Indonesia, on Eucalyptus sp., 26 Mar. 2008, M.J.
septate, branching. Conidiophores micro- to macronematous, Wingfield (holotype CBS H-19824, culture ex-type CBS
subhyaline to pale brown, smooth to rough, simple, sometimes 142239 = CPC 15066).
branched, straight to sinuous. Conidiogenous cells integrated,
Zasmidium Fr., Summa Veg. Scand. 2: 407. 1849. Fig. 117.
terminal or intercalary, hyaline to pale brown, proliferating sym-
Synonyms: Periconiella Sacc., Atti Ist. Veneto Sci. Lett. Arti 3:
podially, polyblastic, with rim-like conidiogenous loci, slightly
727. 1885. (Type: Periconiella velutina (G. Winter) Sacc. 1885).
thickened and darkened. Conidia solitary, rarely catenate in a
Biharia Thirum. & Mishra, Sydowia 7: 79. 1953 (Type: Biharia
single chain, hyaline to subhyaline, smooth, oblong, cylindrical to
vangueriae Thirum. & Mishra 1953).
obclavate, straight, base medium-long obconically truncate, apex
Stenellopsis B. Huguenin, Bull. Trimestriel Soc. Mycol. France
rounded, aseptate or eu- or distoseptate hila thickened and
81: 695. 1966 (Type: Stenellopsis fagraeae B. Huguenin 1966).
darkened and protruding at the base or at both ends when
Verrucisporota D.E. Shaw & Alcorn, Austral. Syst. Bot. 6: 273.
catenate.
1993 (Type: Verrucisporota proteacearum (D.E. Shaw & Alcorn)
Type species: Xenosonderhenioides indonesiana C. Nakash., D.E. Shaw & Alcorn 1993).
Videira & Crous Verrucispora D.E. Shaw & Alcorn, Proc. Linn. Soc. New South
Wales 92: 171. 1967, nom. illeg. (Art. 53.1).
Note: Although Xenosonderhenioides appears to be of minor
importance as a foliar pathogen, very little is known regarding its Ascomata pseudothecial, amphigenous, single, black, erum-
biology. pent, globose, with apical ostiole, and prominent periphyses
lining the ostiolar channel; wall of 2–3 layers of medium brown
Xenosonderhenioides indonesiana C. Nakash. et al., Stud.
textura angularis. Asci aparaphysate, fasciculate, bitunicate,
Mycol. 87: 368. 2017. Fig. 116.
subsessile, obovoid to ellipsoid, straight or slightly incurved, 8-
Diagnosis: Conidiophores micro- to macronematous, subhyaline to spored. Ascospores tri- to multiseriate, overlapping, hyaline,
pale brown, smooth to finely verruculose. Conidiogenous cells in- guttulate, thin-walled, straight to slightly curved, fusoid-
tegrated, terminal and intercalary, hyaline to pale brown, smooth, ellipsoidal with obtuse ends, medianly 1-septate, tapering to-
proliferating sympodially. Conidia solitary, rarely catenate in a single wards both ends. Saprobic or mostly biotrophic, usually
286
Fig. 116. Xenosonderhenioides indonesiana (CPC 15066). A–E. Observations in vitro. A. Culture on OA. B, C. Conidiophore and conidiogenous cell. D. Conidiogenous cell
and conidia. E. Conidia. Scale bars = 10 μm (from Videira et al. 2017).
Fig. 117. Zasmidium spp. A–F. Z. eucalyptorum (CBS H-19689). A. Leaf spot. B, C. Asci. D. Ostiolar periphyses. E. Ascospores. F. Germinating ascospores. G–K.
Z. pseudoparkii. G–I. Conidiophores giving rise to conidia. J. Conidia. K. Germinating ascospores. Scale bars = 10 μm.
CROUS ET AL.
foliicolous, symptomless or causing various lesions, ranging Description and illustration: Crous et al. (2014c).
from yellowish discolourations to distinct leaf spots. In plant
Typus: Mozambique, Forestas de Niassa, on E. urophylla, 2
pathogenic species, mycelium mostly immersed as well as
Feb. 2014, M.J. Wingfield (holotype CBS H-21992, culture ex-
superficial, rarely only immersed; hyphae branched, septate,
type CBS 138860 = CPC 24251).
hyaline or almost so to pigmented, pale olivaceous to brown,
wall thin to somewhat thickened, immersed hyphae smooth or Zasmidium eucalyptorum (Crous & M.J. Wingf.) Quaedvlieg &
almost so to faintly rough, external hyphae distinctly verruculose Crous, Persoonia 33: 24. 2014. Fig. 117.
to verrucose (in culture immersed hyphae usually smooth or Basionym: Mycosphaerella eucalyptorum Crous & M.J. Wingf.,
almost so, aerial hyphae verruculose). Stromata lacking to well- Stud. Mycol. 55: 112. 2006.
developed, pigmented. Conidiophores solitary, arising from su-
Diagnosis: Leaf spots amphigenous, irregular to sub-circular,
perficial hyphae, lateral, occasionally terminal, in vivo (in plant
2–20 mm diam, medium brown, with raised, brown borders,
pathogenic taxa) sometimes also fasciculate, arising from in-
and thin, red-purple margins. Ascospores tri- to multiseriate,
ternal hyphae or stromata, semimacronematous to macro-
overlapping, hyaline, guttulate, thin-walled, straight to slightly
nematous, in culture occasionally micronematous, cylindrical,
curved, fusoid–ellipsoidal with obtuse ends, medianly 1-septate,
filiform, subuliform, straight to strongly geniculatesinuous,
widest in middle of apical cell, not constricted at the septum,
mostly unbranched, aseptate, i.e. reduced to conidiogenous
tapering towards both ends, but more prominently towards the
cells, to pluriseptate, subhyaline to pigmented, pale olivaceous
lower end, (12–)14–15(–17) × (3.5–)4(–4.5) μm in vivo; some
to medium dark brown, wall thin to somewhat thickened,
ascospores with slightly asymmetrical apical cells.
smooth to verruculose; conidiogenous cells integrated, terminal,
occasionally intercalary, rarely pleurogenous, or conidiophores Description and illustration: Crous et al. (2006d).
reduced to conidiogenous cells, mostly polyblastic, sympodial,
Typus: Indonesia, on E. urophylla, Mar. 2004, M.J. Wingfield
with conspicuous, somewhat thickened and darkened-refractive,
planate loci. Conidia solitary or catenate, in simple or branched
11174).
acropetal chains, shape and size variable, ranging from amero-
to scolecosporous, aseptate to transversely plurieuseptate, Zasmidium pseudoparkii (Crous & M.J. Wingf.) Crous & U.
subhyaline to pigmented, pale olivaceous to brown, wall thin to Braun, Schlechtendalia 20: 102. 2010. Fig. 117.
somewhat thickened, smooth or almost so to usually distinctly Basionym: Stenella pseudoparkii Crous & M.J. Wingf., Stud.
verruculose (in plant pathogenic species without superficial Mycol. 55: 128. 2006.
mycelium always verruculose), hila somewhat thickened and
darkened-refractive, planate, conidial secession schizolytic. (see
3–7 mm diam, pale brown, with a raised border. Conidia soli-
Braun et al. 2013).
tary to catenulate in simple chains, medium brown, verruculose,
Type species: Racodium cellare Pers., Neues Mag. Bot. 1: 123. cylindrical or narrowly obclavate, with subobtuse apex, and
1794 [= Zasmidium cellare (Pers.) Fr.] long obconically subtruncate base, straight to curved, 1–5-
septate, 20–50 × 2.5–3 μm; hila thickened, darkened and
Note: The generic concept of Zasmidium was widened by Videira
refractive.
et al. (2017), and the genus now includes many species capable
of causing foliar diseases of Eucalyptus. Description and illustration: Crous et al. (2006d).
Zasmidium eucalypticola U. Braun et al., Stud. Mycol. 87: 358. Typus: Colombia, on Eucalyptus sp., 1995, M.J. Wingfield
2017. (holotype CBS H-19702, culture ex-type CBS 110999 = CPC
1087).
Diagnosis: Conidiophores micro- to macronematous, pale
olivaceous brown to olivaceous brown, septate, straight to Zasmidium pseudovespa (Carnegie) U. Braun et al., Stud.
slightly curved, 38–63 × 3–3.5 μm. Conidiogenous cells inte- Mycol. 87: 362. 2017.
grated, apical, polyblastic, proliferating sympodially, with a short Basionym: Mycosphaerella pseudovespa Carnegie, Mycologia
rachis (ramichloridium-like). Conidia solitary, pale olivaceous 99: 468. 2007.
brown, verruculose, ovoid to cylindrical, base obconically trun-
Diagnosis: Leaf spots amphigenous, subcircular, 3–7 mm diam,
cate and apex rounded, 7.5–20 × 2.5–4 μm, 0–1-septate, hila
single, yellow-brown with thin red-brown to purple margin,
thickened and darkened.
associated with wasp gall, often with raised centre and empty
Description and illustration: Videira et al. (2017). gall, wasp pupae rarely evident but exit hole often observed on
abaxial surface of lesion. Ascospores 2–3-seriate, fusoid with
Typus: Brazil, Minas Gerais, Viçosa, Paraíso, on Eucalyptus sp.,
obtuse ends, slightly tapered to basal end, slightly constricted or
1 Mar. 2008, coll. A.C. Alfenas, isol. P.W. Crous (holotype CBS
not so, widest in middle of apical cell, straight, (10.5–)
H-22959, culture ex-type CBS 142186 = CPC 15149).
11.5–13.5(–14.5) × 3–4 μm.
Zasmidium eucalyptigenum Crous & M.J. Wingf., Persoonia
33: 241. 2014.
Typus: Australia, New South Wales, Urbenville, Reid Plantation,
Diagnosis: Leaf spots amphigenous, dark brown, 10–20 mm
native regeneration within plantation boundary, on E. biturbinata,
diam, with dark brown border. Co-occurring on leaf spots with
14 Apr. 2005, A.J. Carnegie (holotype DAR 77432, culture ex-
Xenosonderhenia eucalypti. Conidia brown, verruculose, straight
type CBS 121159 = DAR 77432).
to curved, solitary or in branched chains, subcylindrical, apex
obtuse, base tapering to a truncate hilum, 1–1.5 μm diam, 1–9- Zasmidium xenoparkii (Crous & M.J. Wingf.) Crous & U. Braun,
septate, 30–120 × (2.5–)3 μm. Schlechtendalia 20: 103. 2010.
288
Basionym: Stenella xenoparkii Crous & M.J. Wingf., Stud. Mycol. resulted in the emergence of “new” diseases of eucalypts not
55: 129. 2006. known on these trees in their native environment. The most
relevant example of this situation is found in the myrtle rust
fungus Austropuccinia psidii that is native on Myrtaceae in South
2–10 mm diam, pale brown, with a raised border and thin, red-
America (Coutinho et al. 1998, Glen et al. 2007) and has become
purple margin. Conidia catenulate in branched chains, medium
one of the most important pathogens of Myrtaceae in Australia,
brown, verruculose, cylindrical or narrowly obclavate, with sub-
where these trees are native (Berthon et al. 2018, Carnegie &
obtuse apex, and subtruncate base, straight to curved, 0–2-
Pegg 2018). Many species of Eucalyptus and Corymbia are
septate, 12–50 × 3–5 μm; hila thickened, darkened and
included in the host range of A. psidii although they are not
refractive.
amongst the most susceptible hosts in Australia (Giblin &
Description and illustration: Crous et al. (2006d). Carnegie 2014, Pegg et al. 2014, Carnegie 2015, Berthon
et al. 2018, Winzer et al. 2019), being more relevant to planta-
Typus: Indonesia, on E. grandis, Mar. 1996, M.J. Wingfield
tion forestry in other parts of the world (Ferreira 1983).
(holotype PREM 54968, isotype CBS H-19703, cultures ex-
Many of the fungi known to occur on exotically grown eu-
type CBS 111185 = CPC 1300, CPC 1299, 1301).
calypts, including pathogens, saprophytes and those of un-
known biology, have likely originated where these trees are
native in Australia. This study is focussed specifically on the
DISCUSSION fungi that are found on living eucalypt foliage. Despite the fact
there are great numbers of genera that occur in this niche, only
Eucalypts display an incredibly wide diversity in their native a selection of species of the 110 genera have been included. Of
range including more than 800 species growing in almost every those chosen for treatment, only the species that have been
conceivable different environmental habitat (Potts & Pederick associated with leaf spots (in most cases pathogenicity re-
2000, Ladiges et al. 2003, Thornhill et al. 2019). It is conse- mains to be proven) have been treated. It is most likely that
quently not surprising these remarkable trees would also provide some of the fungi thought to be saprobes could emerge as
the substrates for an even larger diversity of fungi. These occur pathogens in the future. For example, it is well-recognised that
on tissues of every part of the trees (seeds, capsules, leaves, some fungi occurring as endophytes and thought to be sap-
twigs, branches, stems and leaves) that have been investigated robes or minor pathogens have emerged as major constraints
(Sankaran et al. 1995, Park et al. 2000). Clearly the very large to tree health where these trees are planted in new environ-
number of fungi described from these trees, as illustrated in this ments different to those associated with their natural habitat.
study, have only been minimally sampled. Many thousands of One of the most interesting and emerging examples of this
fungal taxa are yet to be discovered associated with eucalypts situation is found in the Cryphonectriaceae and Botryos-
and these will include not only those that can be grown in culture phaeriaceae introduced into new environments, most likely as
but also unculturable fungi as illustrated in the metabarcoding asymptomatic infections on planting material (Gryzenhout et al.
study of Kemler et al. (2013). 2010, Crous et al. 2016a).
Very little is known regarding the biology of most of the fungi This study was chiefly driven by curiosity of mycologists and
occurring on eucalypts. Those species causing diseases have forest pathologists that have for many years been inspecting
attracted most attention in past studies (Keane et al. 2000). This eucalypts, seeking to determine which organisms were causing
is because they have been established in many parts of the specific leaf spots, blotch or blight diseases. An additional
world in intensively managed plantations as a major component complication on eucalypts, however, is that other than the iconic
of the global timber (especially paper and pulp) industry Calonectria blight, or Teratosphaeria leaf disease, leaf spots
(Doughty 2000, Turnbull 2000). In this regard, fungal pathogens represent a niche to a large number of fungi as secondary
have become an increasingly important and growing constraint to pathogens and saprophytes that commonly co-occur with the
eucalypt forestry world-wide (Keane et al. 2000). Other than well- primary causal agents of eucalypt leaf diseases. Their origin is
known pathogens including the rust Austropuccinia psidii and mostly unknown but we assume that many represent fungi that
species in the Mycosphaerellaceae and Teratosphaeriaceae have resulted in asymptomatic endophytic infections and that
infecting leaves and shoots, important fungal pathogens of eu- they are able to grow and sporulate on stressed and dying tis-
calypts include those causing vascular wilts (e.g. Ceratocysti- sues that resulted from infections by primary pathogens. This
daceae), stem canker diseases (Cryphonectriaceae, often also complicates efforts to identify the causal agents of
Botryosphaeriaceae) and root diseases (e.g. Armillaria, Gano- eucalypt leaf and shoot diseases.
derma). The greatest known diversity of fungi on eucalypts are Calonectria spp. stand out as some of the most important and
those described from leaves and shoots and those occurring on widely sampled fungi infecting the leaves and shoots of euca-
dead plant parts, mainly leaf litter. The majority of these are lypts. This is due to the fact that they rapidly emerged as
thought to be saprophytes, although in reality their ecological important leaf-infecting pathogens (Cylindrocladium leaf blight) of
significance is mostly unknown. Some are likely in future to be eucalypts grown in intensively managed plantations in the tropics
discovered as important sources of medicinal and industrial and sub-tropics (Park et al. 2000). No new species of Calonectria
products, while others will have roles that have yet to be have been introduced in the present study. This is mainly
imagined. because this genus has been, and continues to be well studied
The transfer of eucalypt germplasm to new environments for (Crous 2002, Lombard et al. 2010a,b,c, Alfenas et al. 2015,
amenity and forestry planting dates back more than 100 years Pham et al. 2019). Calonectria spp. are soil-borne fungi where
(Burgess & Wingfield 2017). Most of this germplasm trade has some species have adapted the ability to infect eucalypts (Crous
been via seed (Jimu et al. 2015) but there is also evidence that 2002, Alfenas et al. 2015). In this regard, most species infecting
plants have been moved globally (Andjic et al. 2011). This has eucalypts are likely native to the areas where they have emerged
CROUS ET AL.
as pathogens. But with expanding agriculture and dating back Brazil, Chile and South Africa (Park et al. 2000), we have not
many centuries, soil has also been moved globally. Thus, been able to grow it in culture, presumably due to its biotrophic
intriguing discoveries relating to the origin of some Calonectria habit. The DNA data generated in this study emerged from direct
spp. are likely to emerge in the future. This is particularly relevant sequencing from the inner contents of ascomata. Thyriopsis,
as genomes of these fungi are sequenced, allowing the easy which occurs on needles of Pinus spp., has thyrothecia that open
development of molecular markers to track the global distribution via linear fissures that are sometimes Y-shaped, which clearly
and likely origins of these fungi. distinguish it from Neothyriopsis. Neothyriopsis also has well-
Forest pathologists well-versed with older literature might developed haustoria that are absent in Thyriopsis.
ponder the question as to the fate of iconic eucalypt leaf-infecting Neotrichosphaeria, which is based on N. eucalypticola, was
genera such as Mycosphaerella (Crous 1998). Subsequent to initially described from symptomatic Eucalyptus leaves in
the one fungus one name initiative (Wingfield et al. 2012, Australia (Sivanesan & Shivas 2002b). Neotrichosphaeria is
Rossman et al. 2015), Mycosphaerella was reduced to synon- distinguished from Trichosphaeria (Reblova & Gams 2016) in
ymy under the older, well studied genus name Ramularia lacking a periphysate ostiole, having numerous, long setae,
(Videira et al. 2016). Molecular phylogenetic studies further paraphyses that dissolve during maturation, asci with a visible
revealed that these mycosphaerella-like species actually repre- discharge mechanism, and hyaline, aseptate ascospores. Neo-
sented countless genera, accommodated in the Mycosphaer- trichosphaeria is associated with amphigenous, subcircular,
ellaceae (Videira et al. 2017) and Teratosphaeriaceae brown leaf spots with irregular margins (Sivanesan & Shivas
(Quaedvlieg et al. 2013). Many of these genera give rise to 2002b), but is rarely encountered, and thus regarded to be of
similar disease symptoms and have mycosphaerella-like sexual less importance.
morphs. They are thus best distinguished based on the The family Nowamycetaceae and genus Nowamyces are
morphology of their asexual morphs, in combination with mo- introduced here with two new species, N. piperitae and
lecular data, which has become essential to enable accurate N. globulus. Although N. piperitae is associated with irregular leaf
species identification (Videira et al. 2017). spots on E. piperita, N. globulus co-occurs on older leaf spots on
In this study, we have not attempted to discuss all the new E. globulus along with Teratosphaeria nubilosa, and it is
species or epitypes designated. Yet, it is prudent to comment on assumed to be of minor relevance. Walkaminomyces is intro-
some of the newly introduced genera. Lembosiniella, which duced as new genus to accommodate W. medusae, a foliar
causes prominent round, brown leaf spots on Eucalyptus, is quite pathogen of E. alba. Furthermore, two species of Teratosphaeria
distinct from Lembosina, which is a saprobe occurring on leaves are newly treated in this study i.e. T. delegatensis on
and stems of Rhododendron (Crous et al. in prep.). Lembosi- E. delegatensis, and T. pseudonubilosa on E. globulus. Both
niella forms predominantly hypophyllous, black, superficial hys- T. delegatensis (Park & Keane 1984), and T. pseudonubilosa
terothecia, which are somewhat reminiscent of Aulographina (Perez et al. 2014) are known species that were previously
eucalypti (but see below). Although only two species of Lem- described, but are either newly combined, or validated in
bosiniella are treated here, many different species have been Teratosphaeria.
encountered on eucalypts during the course of this study, but This study includes a significant body of work on the fungi
most have proven to be biotrophic in nature, with ascospores not occurring on eucalypt foliage. Yet these fungi remain poorly
germinating or dying upon germination in pure culture, sug- studied. The boundaries of many genera remain unclear and
gesting that further collections and research are still required to require detailed examination. For example, Pachysacca spp. are
elucidate the species in this genus. commonly associated with amphigenous leaf spots, with less
A debate regarding the generic boundaries of Microthyrium/ well-developed stromata developing on the opposite side of the
Phaeothyriolum deserves comment, especially because no leaf and with the fungus forming dendritic or large circular stro-
molecular data were available for this complex prior to the pre- mata. Unfortunately, the spots associated with this fungus are
sent study. Phaeothyriolum is associated with chlorotic leaf relatively uncommon, being confined to a few trees in a plan-
spots, that at times form a red margin with clusters of black tation or native bush, and all attempts to cultivate it have to date
thyrothecia with irregular ostioles. In contrast, Microthyrium is proven unsuccessful. Another generic complex that is poorly
associated with leaf litter, and has regular, round ostioles. understood due to the lack of cultures and thus DNA-sequence
Although there are clearly several genera involved in this com- data is “Stigmina”. While Stigmina (based on S. platani) has been
plex, the distinction of “occurring on leaf litter” from those species shown to be a synonym of Pseudocercospora (Braun & Crous
occurring on living leaves is rather subtle, as the pathogens will 2006, Crous et al. 2013), the various “Stigmina” species occur-
eventually also occur on litter, and Phaeothyriolum is one such ring on eucalypts lack cultures or DNA data, and thus their
example. taxonomy cannot be resolved until their taxa are recollected.
Sonderhenia (pycnidial conidiomata, distoseptate conidia with A major fungal complex associated with leaf-spots on euca-
a central pore, brown, percurrently proliferating conidiogenous lypts is that accommodating the leaf pathogen Aulographina
cells, and a teratosphaeria-like sexual morph), was initially eucalypti (target spot or corky spot). Aulographina (based on
introduced with two species, S. eucalyptorum and A. pinorum occurring on Pinus spp. in Europe) is not congeneric
S. eucalypticola, with S. radiata added as a third species in this with the eucalypt-infecting fungus. This suggests that the name
study. Neosonderhenia is morphologically similar, chiefly being of its spermatogonial morph, Thyrinula (Petrak & Sydow 1924),
distinguished based on phylogeny, and the fact that all species should rather be used for this pathogen. As shown here, Thyr-
have conidiomatal paraphyses. Although species in these inula also has a hyphomycetous asexual morph, previously
genera have similar disease symptoms, they can still readily be described as Alysidiella. The genus Thyrinula represents several
distinguished based on conidial morphology. species, namely T. dunnii (from Australia), T. eucalypti (formerly
Neothyriopsis is associated with corky leaf spots on Euca- referred to as Aulographina eucalypti s. str., common in
lyptus leaves. Although the pathogen is known from Australia, Australia), T. eucalyptina (originally described from South Africa,
290
but also in Australia and Zimbabwe), T. parasitica (described Eucalypt leaf and shoot-infecting fungi are likely to become
from South Africa, but also found in Australia, Colombia and the increasingly important in the future. This has been an ongoing
UK), and T. uruguayensis (from Colombia, Madagascar, South and emerging trend as eucalypt plantation forestry has expanded
Africa and Uruguay). Further studies are now underway to globally (Andjic et al. 2011). While quarantine to reduce this
elucidate all the species associated with corky spot disease. This threat is being strongly promoted, this continues to fail in many
will make it possible to understand which species in the complex parts of the world (Eschen et al. 2015, Klapwijk et al. 2016,
have been introduced along with eucalypts to countries in Africa, McTaggart et al. 2016, Burgess & Wingfield 2017). This is
the Americas, Europe, and Asia. partially due to the enormous difficulty of applying effective
One of the most important yet relatively poorly known genera quarantine for the huge volumes of planting material that is
of leaf and shoot blight pathogens is Quambalaria. These appear traded globally. Potential pathogens of eucalypts are moved in
to be specifically Australian fungi that when first discovered were plants of other genera related to these trees. This is for example
thought to be ascomycetes. They were later shown to be Ba- the likely pathway of movement and introduction of the myrtle
sidiomycetes related to the smut fungi (de Beer et al. 2006). Two rust fungus into countries such as South Africa (Burgess &
species of Quambalaria, Q. pitereka and Q. eucalypti infect the Wingfield 2017). Latent pathogens with wide host ranges such
shoots and leaves of eucalypts (Simpson 2000, Pegg et al. as those in the Botryosphaeriaceae are found in the skins of fruit
2008). They are separated by their host genera with such as mangoes (Mangifera indica) and avocados (Persea
Q. pitereka specifically infecting species of Corymbia and americana) both of which are moved globally in large volumes
C. eucalypti only infecting species of Eucalyptus. Both of these and have the potential to directly infect eucalypts or to reach
fungi have been accidentally introduced into new environments these trees via various “host bridges”.
with Q. pitereka now found in plantations of Corymbia in China Host shifts are occurring where native pathogens in countries
(Zhou & Wingfield 2011, Chen et al. 2017) and Q. eucalypti outside the natural range of eucalypts are adapting the capacity to
causing serious diseases of Eucalyptus spp. in South Africa infect these trees (Burgess & Wingfield 2017). These newly
(Wingfield et al. 1993, Roux et al. 2006) and Brazil (Alfenas et al. adapted eucalypt pathogens such as those in the Cryphonec-
2001). Intriguingly, Q. eucalypti has also undergone a host shift triaceae and Ceratocystidaceae (Wingfield 2003, Gryzenhout et al.
to infect a native species of Myrtaceae in Uruguay (Perez et al. 2010) now threaten eucalypts and other Myrtaceae in their native
2008). These fungi are of growing importance globally and are range. Myrtle rust (Austropuccinia psidii), native to Myrtaceae in
likely to threaten global eucalypt plantation forestry in the future. South America, is a sobering reminder of when such host shift
Interestingly within this group is another closely related species, pathogens establish in Australia (Carnegie et al. 2016, Pegg et al.
Q. coyrecup, which causes a severe and devastating canker 2017, Carnegie & Pegg 2018). Importantly, these “new” eucalypt
disease of Corymbia calophylla in Western Australia (Paap et al. pathogens do not remain static but also have the capacity to adapt
2008). and change as has recently been shown for Austropuccinia psidii,
An important technology applied in eucalypt plantation which is undergoing sexual reproduction generating novel geno-
forestry during the past approximately 40 years has been artifi- types (McTaggart et al. 2018).
cially induced hybridisation between species particularly of Very little is known regarding the global patterns of
Eucalyptus. These trees have been propagated vegetatively and spread for even the most important eucalypt leaf pathogens.
have become a major component of clonal plantations particu- The two pathogens for which pathways of spread have been
larly in South America, Africa and Asia. This has made it possible reasonably well studied are Austropuccinia psidii mentioned
to enhance adaptation to specific environments, and capitalise above and Teratosphaeria nubilosa that has been subjected
on hybrid vigour (heterosis), to enhance desired fibre charac- to a number of population genetic studies (Perez et al.
teristics and to avoid disease problems. Hybridising different 2012). This situation will likely improve in the future as
species of eucalypts has already been seen to influence leaf and eucalypt leaf and shoot pathogens become increasingly
shoot diseases with hybrids acting as “bridges” to susceptibility important, and as tools to study pathways of introduction and
between species. This has been seen with Teratosphaeria global movement become increasingly available and refined
destructans, a species causing a serious leaf and shoot disease (Burgess & Wingfield 2017).
of Eucalyptus in South East Asia (Wingfield et al. 1996, Old et al. Beyond the primary pathogens, secondary pathogens and
2003) and South Africa (Greyling et al. 2016). In areas of South saprophytes are adapting to infect eucalypts globally. This is a
East Asia where the pathogen occurs, Eucalyptus brassiana is poorly understood subject, but where there are growing exam-
highly susceptible to infection by T. destructans but E. pellita is ples of newly emerging problems. The disease problems arise in
not. But recently-developed hybrids between E. pellita and some cases where damage to leaf tissue due to insects provides
E. brassiana can be severely infected by T. destructans (Wing- sites of infection or stress (e.g. drought) that allows endophytic
field, unpubl. data). The previously observed resistance in fungi such as those in the Botryosphaeriaceae to develop. An
E. pellita will break down due to the “bridge” of susceptibility interesting recent example has emerged in countries of South
provided by the hybrid between the two Eucalyptus species. A East Asia where damage due to Myridae such as species of
similar case was seen for hybrids of E. globulus and E. nitens in Helopeltis has enabled the previously relatively innocuous
Tasmania, Australia, with F1 hybrids being more susceptible to Coniella eucalypti to emerge as an important cause of leaf dis-
Teratosphaeria leaf disease than either parent (Dungey et al. ease (Wingfield, unpubl. data).
1997). An assessment of the importance of T. cryptica and Past experience clearly indicates that fungi infecting euca-
T. nubilosa in causing disease on juvenile leaves from this study lypts (not only leaves and shoots) will become increasingly
revealed that both species were equally important on E. globulus, important in the future and assumptions of their biology, patho-
T. cryptica was the only species on E. nitens, while both were genicity and ecology are highly likely to change as a result of
also present on the F1 hybrids but with T. cryptica predominating climate change. This calls for a much more intensive study of
(Carnegie & Ades 2002). these fungi with surveys of the many species that have yet to be
CROUS ET AL.
sampled and studied. This is clearly a huge task that will require Barber PA, Burgess T, Hardy GStJ, et al. (2005). Botryosphaeria species from
substantial global resources. But it is one that is sufficiently Eucalyptus in Australia are pleoanamorphic, producing Dichomera syna-
namorphs in culture. Mycological Research 109: 1347–1363.
important to justify robust investment. Our hope is that the pre- Barber PA, Carnegie AJ, Burgess TI, et al. (2008). Leaf diseases caused by
sent study will provide a foundation for increased work on these Mycosphaerella species in Eucalyptus globulus plantations and nearby
fungi in the future. native forest in the Green Triangle Region of Australia. Australasian Plant
Pathology 37: 372–381.
Barber PA, Crous PW, Groenewald JZ, et al. (2011). Reassessing Vermisporium
(Amphisphaeriaceae), a genus of foliar pathogens of eucalypts. Persoonia
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Beenken L (2017). Austropuccinia: a new genus name for the myrtle rust
Puccinia psidii placed within the redefined family Sphaerophragmiaceae
Supplementary data to this article can be found online at https:// (Pucciniales). Phytotaxa 297: 53–61.
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available online at www.studiesinmycology.org STUDIES IN MYCOLOGY 94: 125–298 (2019).

Foliar pathogens of eucalypts*

*Correspondence: P.W. Crous, p.crous@wi.knaw.nl

Key words: Corymbia, Eucalyptus, Foliar pathogen, New taxa, Taxonomy.

Available online 8 August 2019; https://doi.org/10.1016/j.simyco.2019.08.001.

Peer review under responsibility of Westerdijk Fungal Biodiversity Institute.

30–35(–37) × 8–9(–10) μm. Germinating ascospores dark

Notes: Lembosina eucalypti is reminiscent of Thyrinula eucalypti

Elsinoe jasminicola CBS 212.63 (KX886997.1)

1 Austroafricana parva CBS 110503 (KT186510.1)

Fig. 61. (Continued).

16–18 × 6–7 μm (Hansford 1956, Swart 1986c). This species is

Fig. 95. (Continued).

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