Asperisporium and Pantospora (Mycosphaerellaceae): epitypifications and phylogenetic placement.
ABSTRACT The species-rich family Mycosphaerellaceae contains considerable morphological diversity and includes numerous anamorphic genera, many of which are economically important plant pathogens. Recent revisions and phylogenetic research have resulted in taxonomic instability. Ameliorating this problem requires phylogenetic placement of type species of key genera. We present an examination of the type species of the anamorphic Asperisporium and Pantospora. Cultures isolated from recent port interceptions were studied and described, and morphological studies were made of historical and new herbarium specimens. DNA sequence data from the ITS region and nLSU were generated from these type species, analysed phylogenetically, placed into an evolutionary context within Mycosphaerellaceae, and compared to existing phylogenies. Epitype specimens associated with living cultures and DNA sequence data are designated herein. Asperisporium caricae, the type of Asperisporium and cause of a leaf and fruit spot disease of papaya, is closely related to several species of Passalora including P. brachycarpa. The status of Asperisporium as a potential generic synonym of Passalora remains unclear. The monotypic genus Pantospora, typified by the synnematous Pantospora guazumae, is not included in Pseudocercospora sensu stricto or sensu lato. Rather, it represents a distinct lineage in the Mycosphaerellaceae in an unresolved position near Mycosphaerella microsora.
- SourceAvailable from: Pedro W Crous[Show abstract] [Hide abstract]
ABSTRACT: Cercosporoid fungi (former Cercospora s. lat.) represent one of the largest groups of hyphomycetes belonging to the Mycosphaerellaceae (Ascomycota). They include asexual morphs, asexual holomorphs or species with mycosphaerella-like sexual morphs. Most of them are leaf-spotting plant pathogens with special phytopathological relevance. The only monograph of Cercospora s. lat., published by Chupp (1954), is badly in need of revision. However, the treatment of this huge group of fungi can only be accomplished stepwise on the basis of treatments of cercosporoid fungi on particular host plant families. The present first part of this series comprises an introduction, a survey on currently recognised cercosporoid genera, a key to the genera concerned, a discussion of taxonomically relevant characters, and descriptions and illustrations of cercosporoid species on other fungi (mycophylic taxa), Pteridophyta and Gymnospermae, arranged in alphabetical order under the particular cercosporoid genera, which are supplemented by keys to the species concerned. The following taxonomic novelties are introduced: Passalora austroplenckiae comb. nov., P. backmanii comb. nov., P. condensata comb. nov., P. gymnocladi comb. nov., P. thalictri comb. nov., Pseudocercospora davalliicola sp. nov., P. chamaecyparidis comb. nov., P. cratevicola nom. nov., P. gleicheniae comb. nov., P. lygodiicola sp. nov., P. lygodiigena nom. nov., P. nephrolepidigena sp. nov., P. paraexosporioides sp. nov., P. pini-densiflorae var. montantiana comb. et stat. nov., P. pteridigena sp. nov., P. ptisanae sp. nov., P. sciadopityos sp. nov., P. subramanianii nom. nov., P. thujina comb. nov., and Zasmidium australiense comb. nov.IMA fungus. 12/2013; 4(2):265-345.
- [Show abstract] [Hide abstract]
ABSTRACT: Pseudocercospora is a large cosmopolitan genus of plant pathogenic fungi that are commonly associated with leaf and fruit spots as well as blights on a wide range of plant hosts. They occur in arid as well as wet environments and in a wide range of climates including cool temperate, sub-tropical and tropical regions. Pseudocercospora is now treated as a genus in its own right, although formerly recognised as either an anamorphic state of Mycosphaerella or having mycosphaerella-like teleomorphs. The aim of this study was to sequence the partial 28S nuclear ribosomal RNA gene of a selected set of isolates to resolve phylogenetic generic limits within the Pseudocercospora complex. From these data, 14 clades are recognised, six of which cluster in Mycosphaerellaceae. Pseudocercospora s. str. represents a distinct clade, sister to Passalora eucalypti, and a clade representing the genera Scolecostigmina, Trochophora and Pallidocercospora gen. nov., taxa formerly accommodated in the Mycosphaerella heimii complex and characterised by smooth, pale brown conidia, as well as the formation of red crystals in agar media. Other clades in Mycosphaerellaceae include Sonderhenia, Microcyclosporella, and Paracercospora. Pseudocercosporella resides in a large clade along with Phloeospora, Miuraea, Cercospora and Septoria. Additional clades represent Dissoconiaceae, Teratosphaeriaceae, Cladosporiaceae, and the genera Xenostigmina, Strelitziana, Cyphellophora and Thedgonia. The genus Phaeomycocentrospora is introduced to accommodate Mycocentrospora cantuariensis, primarily distinguished from Pseudocercospora based on its hyaline hyphae, broad conidiogenous loci and hila. Host specificity was considered for 146 species of Pseudocercospora occurring on 115 host genera from 33 countries. Partial nucleotide sequence data for three gene loci, ITS, EF-1α, and ACT suggest that the majority of these species are host specific. Species identified on the basis of host, symptomatology and general morphology, within the same geographic region, frequently differed phylogenetically, indicating that the application of European and American names to Asian taxa, and vice versa, was often not warranted. New genera - Pallidocercospora Crous, Phaeomycocentrospora Crous, H.D. Shin & U. Braun; New species - Cercospora eucommiae Crous, U. Braun & H.D. Shin, Microcyclospora quercina Crous & Verkley, Pseudocercospora ampelopsis Crous, U. Braun & H.D. Shin, Pseudocercospora cercidicola Crous, U. Braun & C. Nakash., Pseudocercospora crispans G.C. Hunter & Crous, Pseudocercospora crocea Crous, U. Braun, G.C. Hunter & H.D. Shin, Pseudocercospora haiweiensis Crous & X. Zhou, Pseudocercospora humulicola Crous, U. Braun & H.D. Shin, Pseudocercospora marginalis G.C. Hunter, Crous, U. Braun & H.D. Shin, Pseudocercospora ocimi-basilici Crous, M.E. Palm & U. Braun, Pseudocercospora plectranthi G.C. Hunter, Crous, U. Braun & H.D. Shin, Pseudocercospora proteae Crous, Pseudocercospora pseudostigmina-platani Crous, U. Braun & H.D. Shin, Pseudocercospora pyracanthigena Crous, U. Braun & H.D. Shin, Pseudocercospora ravenalicola G.C. Hunter & Crous, Pseudocercospora rhamnellae G.C. Hunter, H.D. Shin, U. Braun & Crous, Pseudocercospora rhododendri-indici Crous, U. Braun & H.D. Shin, Pseudocercospora tibouchinigena Crous & U. Braun, Pseudocercospora xanthocercidis Crous, U. Braun & A. Wood, Pseudocercosporella koreana Crous, U. Braun & H.D. Shin; New combinations - Pallidocercospora acaciigena (Crous & M.J. Wingf.) Crous & M.J. Wingf., Pallidocercospora crystallina (Crous & M.J. Wingf.) Crous & M.J. Wingf., Pallidocercospora heimii (Crous) Crous, Pallidocercospora heimioides (Crous & M.J. Wingf.) Crous & M.J. Wingf., Pallidocercospora holualoana (Crous, Joanne E. Taylor & M.E. Palm) Crous, Pallidocercospora konae (Crous, Joanne E. Taylor & M.E. Palm) Crous, Pallidoocercospora irregulariramosa (Crous & M.J. Wingf.) Crous & M.J. Wingf., Phaeomycocentrospora cantuariensis (E.S. Salmon & Wormald) Crous, H.D. Shin & U. Braun, Pseudocercospora hakeae (U. Braun & Crous) U. Braun & Crous, Pseudocercospora leucadendri (Cooke) U. Braun & Crous, Pseudocercospora snelliana (Reichert) U. Braun, H.D. Shin, C. Nakash. & Crous, Pseudocercosporella chaenomelis (Y. Suto) C. Nakash., Crous, U. Braun & H.D. Shin; Typifications: Epitypifications - Pseudocercospora angolensis (T. Carvalho & O. Mendes) Crous & U. Braun, Pseudocercospora araliae (Henn.) Deighton, Pseudocercospora cercidis-chinensis H.D. Shin & U. Braun, Pseudocercospora corylopsidis (Togashi & Katsuki) C. Nakash. & Tak. Kobay., Pseudocercospora dovyalidis (Chupp & Doidge) Deighton, Pseudocercospora fukuokaensis (Chupp) X.J. Liu & Y.L. Guo, Pseudocercospora humuli (Hori) Y.L. Guo & X.J. Liu, Pseudocercospora kiggelariae (Syd.) Crous & U. Braun, Pseudocercospora lyoniae (Katsuki & Tak. Kobay.) Deighton, Pseudocercospora lythri H.D. Shin & U. Braun, Pseudocercospora sambucigena U. Braun, Crous & K. Schub., Pseudocercospora stephanandrae (Tak. Kobay. & H. Horie) C. Nakash. & Tak. Kobay., Pseudocercospora viburnigena U. Braun & Crous, Pseudocercosporella chaenomelis (Y. Suto) C. Nakash., Crous, U. Braun & H.D. Shin, Xenostigmina zilleri (A. Funk) Crous; Lectotypification - Pseudocercospora ocimicola (Petr. & Cif.) Deighton; Neotypifications - Pseudocercospora kiggelariae (Syd.) Crous & U. Braun, Pseudocercospora lonicericola (W. Yamam.) Deighton, Pseudocercospora zelkovae (Hori) X.J. Liu & Y.L. Guo.Studies in Mycology 06/2013; 75(1):37-114. · 9.30 Impact Factor
Article: Alternaria redefined.[Show abstract] [Hide abstract]
ABSTRACT: Alternaria is a ubiquitous fungal genus that includes saprobic, endophytic and pathogenic species associated with a wide variety of substrates. In recent years, DNA-based studies revealed multiple non-monophyletic genera within the Alternaria complex, and Alternaria species clades that do not always correlate to species-groups based on morphological characteristics. The Alternaria complex currently comprises nine genera and eight Alternaria sections. The aim of this study was to delineate phylogenetic lineages within Alternaria and allied genera based on nucleotide sequence data of parts of the 18S nrDNA, 28S nrDNA, ITS, GAPDH, RPB2 and TEF1-alpha gene regions. Our data reveal a Pleospora/Stemphylium clade sister to Embellisia annulata, and a well-supported Alternaria clade. The Alternaria clade contains 24 internal clades and six monotypic lineages, the assemblage of which we recognise as Alternaria. This puts the genera Allewia, Brachycladium, Chalastospora, Chmelia, Crivellia, Embellisia, Lewia, Nimbya, Sinomyces, Teretispora, Ulocladium, Undifilum and Ybotromyces in synonymy with Alternaria. In this study, we treat the 24 internal clades in the Alternaria complex as sections, which is a continuation of a recent proposal for the taxonomic treatment of lineages in Alternaria. Embellisia annulata is synonymised with Dendryphiella salina, and together with Dendryphiella arenariae, are placed in the new genus Paradendryphiella. The sexual genera Clathrospora and Comoclathris, which were previously associated with Alternaria, cluster within the Pleosporaceae, outside Alternaria s. str., whereas Alternariaster, a genus formerly seen as part of Alternaria, clusters within the Leptosphaeriaceae. Paradendryphiella is newly described, the generic circumscription of Alternaria is emended, and 32 new combinations and 10 new names are proposed. A further 10 names are resurrected, while descriptions are provided for 16 new Alternaria sections. New combinations - Alternaria abundans (E.G. Simmons) Woudenb. & Crous, Alternaria alternariae (Cooke) Woudenb. & Crous, Alternaria atra (Preuss) Woudenb. & Crous, Alternaria bornmuelleri (Magnus) Woudenb. & Crous, Alternaria botrytis (Preuss) Woudenb. & Crous, Alternaria caespitosa (de Hoog & C. Rubio) Woudenb. & Crous, Alternaria cantlous (Yong Wang bis & X.G. Zhang) Woudenb. & Crous, Alternaria caricis (E.G. Simmons) Woudenb. & Crous, Alternaria cinerea (Baucom & Creamer) Woudenb. & Crous, Alternaria didymospora (Munt.-Cvetk.) Woudenb. & Crous, Alternaria fulva (Baucom & Creamer) Woudenb. & Crous, Alternaria hyacinthi (de Hoog & P.J. Mull. bis) Woudenb. & Crous, Alternaria indefessa (E.G. Simmons) Woudenberg & Crous, Alternaria leptinellae (E.G. Simmons & C.F. Hill) Woudenb. & Crous, Alternaria lolii (E.G. Simmons & C.F. Hill) Woudenb. & Crous, Alternaria multiformis (E.G. Simmons) Woudenb. & Crous, Alternaria obclavata (Crous & U. Braun) Woudenb. & Crous, Alternaria obovoidea (E.G. Simmons) Woudenb. & Crous, Alternaria oudemansii (E.G. Simmons) Woudenb. & Crous, Alternaria oxytropis (Q. Wang, Nagao & Kakish.) Woudenb. & Crous, Alternaria penicillata (Corda) Woudenb. & Crous, Alternaria planifunda (E.G. Simmons) Woudenb. & Crous, Alternaria proteae (E.G. Simmons) Woudenb. & Crous, Alternaria scirpinfestans (E.G. Simmons & D.A. Johnson) Woudenb. & Crous, Alternaria scirpivora (E.G. Simmons & D.A. Johnson) Woudenb. & Crous, Alternaria septospora (Preuss) Woudenb. & Crous, Alternaria slovaca (Svob.-Pol., L. Chmel & Bojan.) Woudenb. & Crous, Alternaria subcucurbitae (Yong Wang bis & X.G. Zhang) Woudenb. & Crous, Alternaria tellustris (E.G. Simmons) Woudenb. & Crous, Alternaria tumida (E.G. Simmons) Woudenb. & Crous, Paradendryphiella salina (G.K. Sutherl.) Woudenb. & Crous, Paradendryphiella arenariae (Nicot) Woudenb. & Crous. New names - Alternaria aspera Woudenb. & Crous, Alternaria botryospora Woudenb. & Crous, Alternaria brassicae-pekinensis Woudenb. & Crous, Alternaria breviramosa Woudenb. & Crous, Alternaria chlamydosporigena Woudenb. & Crous, Alternaria concatenata Woudenb. & Crous, Alternaria embellisia Woudenb. & Crous, Alternaria heterospora Woudenb. & Crous, Alternaria papavericola Woudenb. & Crous, Alternaria terricola Woudenb. & Crous. Resurrected names - Alternaria cetera E.G. Simmons, Alternaria chartarum Preuss, Alternaria consortialis (Thüm.) J.W. Groves & S. Hughes, Alternaria cucurbitae Letendre & Roum., Alternaria dennisii M.B. Ellis, Alternaria eureka E.G. Simmons, Alternaria gomphrenae Togashi, Alternaria malorum (Ruehle) U. Braun, Crous & Dugan, Alternaria phragmospora Emden, Alternaria scirpicola (Fuckel) Sivan. New sections, all in Alternaria - sect. Chalastospora Woudenb. & Crous, sect. Cheiranthus Woudenb. & Crous, sect. Crivellia Woudenb. & Crous, sect. Dianthicola Woudenb. & Crous, sect. Embellisia Woudenb. & Crous, sect. Embellisioides Woudenb. & Crous, sect. Eureka Woudenb. & Crous, sect. Infectoriae Woudenb. & Crous, sect. Japonicae Woudenb. & Crous, sect. Nimbya Woudenb. & Crous, sect. Phragmosporae Woudenb. & Crous, sect. Pseudoulocladium Woudenb. & Crous, sect. Teretispora Woudenb. & Crous, sect. Ulocladioides Woudenb. & Crous, sect. Ulocladium Woudenb. & Crous, sect. Undifilum Woudenb. & Crous. New genus - Paradendryphiella Woudenb. & Crous.Studies in Mycology 06/2013; 75(1):171-212. · 9.30 Impact Factor
© 2011 Nationaal Herbarium Nederland & Centraalbureau voor Schimmelcultures
You are free to share - to copy, distribute and transmit the work, under the following conditions:
Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work).
Non-commercial: You may not use this work for commercial purposes.
No derivative works: You may not alter, transform, or build upon this work.
For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be
waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights.
Persoonia 27, 2011: 1–8
Mycosphaerella and related fungi are classified in Capnodiales
(Dothideomycetes, Ascomycota) and include thousands of spe-
cies (Crous et al. 2007). These fungi have diverse ecological
roles, especially as saprophytes and parasites, and numer-
ous species are of agricultural significance (Crous 2009). As
plant pathogens, these fungi are found on plant taxa across
the embryophytes (Farr & Rossman 2011) with most species
exhibiting host specificity (Crous 2009).
In terms of morphology, generic concepts for Mycosphaerella
and its related anamorphs have been challenging as the
number of variable and overlapping characters has led to dif-
ferent classifications that emphasize different characters (Baker
et al. 2000, Crous & Braun 2003, Crous et al. 2007, 2009b).
With progress towards phylogenetic hypotheses based on
DNA sequence data, it has become apparent that older mor-
phological classifications are riddled with non-monophyletic
groups and unexpected bedfellows (Crous et al. 2007, 2009b,
The backbone of our understanding of the phylogeny of this
group is skewed in large part, but not entirely, towards sampling
from fungi associated with hosts in two plant families, Myrtaceae
and Proteaceae (Crous et al. 2007, 2009a, b, Crous 2009). Fur-
thermore, the number of genetic loci that have been examined
is relatively small. While the type species of Mycosphaerella,
M. punctiformis, has been placed in a phylogenetic context (Verkley
et al. 2004), type species of large, related genera have yet to
be placed (Crous & Braun 2003, Crous et al. 2007, 2009b).
During the course of conducting work related to plant protec-
tion and quarantine, we obtained new collections via port
interceptions that were identified as Asperisporium caricae
and Pantospora guazumae. These collections represent the
generic types of Asperisporium (Ellis 1971) and Pantospora
(Deighton 1976), respectively. Asperisporium is a small genus
that includes roughly 12 species (Kirk et al. 2008). It shares
many morphological features with Passalora such as pigmented
conidia and thickened and darkened conidiogenous loci while
it is differentiated by verrucose conidia (Crous & Braun 2003,
Schubert & Braun 2005), but this distinction has been consid-
ered doubtful (Crous & Braun 2003, Schubert & Braun 2005).
Asperisporium caricae is responsible for an important leaf and
fruit spot disease of Carica papaya (papaw or papaya) (Stevens
1939) that is commonly referred to as black spot, blight or ‘rust’
of pawpaw (Ellis & Holliday 1972). The synnematous Panto
spora is a monotypic genus that causes a leaf spot disease of
Guazuma ulmifolia (Deighton 1976). Deighton (1976) and Crous
& Braun (2003) both noted its similarity to Pseudocercospora,
especially in regards to the dictyospores in the type species
of Pseudocercospora, P. vitis, and the latter authors formerly
classified it in that genus. Since neither Asperisporium nor
Pantospora have been placed phylogenetically (Crous & Braun
2003, Schubert & Braun 2005), we generated DNA sequence
data from the ITS and nLSU, conducted analyses, present
phylogenetic placements of these genera, and discuss them in
the context of existing phylogenetic studies (Crous et al. 2007,
2009b). To further stabilize the application of Asperisporium
and Pantospora, we herein epitypify the type species of these
genera with collections associated with living cultures and DNA
sequence data. Morphological descriptions of the designated
epitypes and associated ex-epitype cultures are presented, and
Asperisporium and Pantospora (Mycosphaerellaceae):
epitypifications and phylogenetic placement
A.M. Minnis1, A.H. Kennedy2, D.B. Grenier3, S.A. Rehner1, J.F. Bischoff3
1 Systematic Mycology & Microbiology Laboratory, USDA-ARS, B010A,
10300 Baltimore Ave., Beltsville, MD 20705, USA.
2 Molecular Diagnostics Laboratory, USDA-APHIS, B580, Powder Mill Rd.,
Beltsville, MD 20705, USA.
3 Systematic Mycology & Microbiology Laboratory, USDA–APHIS, B010A,
10300 Baltimore Ave., Beltsville, MD 20705, USA;
corresponding author e-mail: Joe.Bischoff@aphis.usda.gov.
Abstract The species-rich family Mycosphaerellaceae contains considerable morphological diversity and includes
numerous anamorphic genera, many of which are economically important plant pathogens. Recent revisions and
phylogenetic research have resulted in taxonomic instability. Ameliorating this problem requires phylogenetic place-
ment of type species of key genera. We present an examination of the type species of the anamorphic Asperisporium
and Pantospora. Cultures isolated from recent port interceptions were studied and described, and morphological
studies were made of historical and new herbarium specimens. DNA sequence data from the ITS region and nLSU
were generated from these type species, analysed phylogenetically, placed into an evolutionary context within
Mycosphaerellaceae, and compared to existing phylogenies. Epitype specimens associated with living cultures
and DNA sequence data are designated herein. Asperisporium caricae, the type of Asperisporium and cause of a
leaf and fruit spot disease of papaya, is closely related to several species of Passalora including P. brachycarpa.
The status of Asperisporium as a potential generic synonym of Passalora remains unclear. The monotypic genus
Pantospora, typified by the synnematous Pantospora guazumae, is not included in Pseudocercospora sensu stricto
or sensu lato. Rather, it represents a distinct lineage in the Mycosphaerellaceae in an unresolved position near
Article info Received: 9 June 2011; Accepted: 1 August 2011; Published: 9 September 2011.
Persoonia – Volume 27, 2011
historical collections and descriptions were studied to confirm
conspecificity of the new collections.
MATERIALS AND METHODS
Morphology and herbarium material
Dried herbarium material was rehydrated and viewed in 3 %
KOH (Largent et al. 1977), and microscopic observations of
cultures were made of material mounted in 3 % KOH or buffered
Shear’s mounting fluid (Graham 1959). Herbarium acronyms
follow Thiers (2011). See Farr & Rossman (2011) for additional
information about collections housed at BPI.
Isolates in pure culture were grown in plastic Petri plates on
2 % Difco potato-dextrose agar (PDA) and BBL Sabouraud dex-
trose agar (SDA), which were both prepared according to the
manufacturers’ instructions. Cultures were incubated at 24 °C
with a 12 h light/dark regimen. A subset of these cultures was
transferred to a 12 h black light/dark regimen also at 24 °C after
approximately 2–2.5 wk to promote sporulation. Sporulation
was also promoted by exposing 1 mo old cultures that were
otherwise incubated in the dark at ambient room temperature
to 1 h of UV at an intensity setting of 60 with a Fisher Biotech
Transilluminator FBTIV-816 at approximately 2–3 d intervals.
Terminology for colour includes general terms from author
notes as well as standard terminology with the sample refer-
ence code in parentheses from Kornerup & Wanscher (1967).
Voucher cultures were deposited at the Centraalbureau voor
DNA extraction, PCR amplification, and sequencing
DNA was extracted from fresh mycelium using the Qiagen
DNeasy Plant Mini Kit (Gaithersburg, Maryland). DNA sequence
data were generated from the nuclear encoded ribosomal ITS
region (ITS1, 5.8S, ITS2), the nuclear encoded ribosomal large
subunit (28S nLSU), and a portion between amino acid motifs
5–7 of the nuclear DNA-dependant RNA Polymerase II gene’s
large subunit (RPB2) for Asperisporium caricae and Pantospora
guazumae. PCR cocktails for all reactions contained 0.2 µM of
each forward and reverse primer, GoTaq Flexi Buffer (Promega;
Madison, Wisconsin), 0.2 mM dNTPs, 2.0 mM Mg2+, and 5U/µL
Pomega GoTaq. Primers used for PCR and sequencing were
fRPB2-5F and fRPB2-7cR (Liu et al. 1999) for RPB2, ITS5
and ITS4 (White et al. 1990) for the ITS, and LROR (Monclavo
et al. 2000) and LR7 (Vilgalys & Hester 1990) for the nLSU.
Thermal cycling conditions for RPB2 and nLSU were those
of Malkus et al. (2006) and Reeb et al. (2004), respectively.
Thermal cycling conditions for the ITS were: 95 °C for 60 s; 35
cycles at 95 °C for 15 s, 55 °C for 20 s, and 72 °C for 1 min;
and a final extension at 72 °C for 3 min. Cycle sequencing was
conducted using BigDye v. 3.1 (Applied Biosystems; Foster City,
California) sequencing chemistry. Resulting fluorescent-labelled
fragments were sequenced on an ABI 3730 capillary sequencer.
Electropherograms were edited in the program Geneious Pro
v. 5 (Drummond et al. 2010). Sequences were submitted to
Data matrix and phylogenetic analysis
For the purpose of determining preliminary phylogenetic posi-
tions of A. caricae and P. guazumae among members of Myco
sphaerellaceae and related genera, their nLSU sequences
were manually incorporated into the alignment of Crous et al.
(2009b) and analyzed phylogenetically using Maximum Parsi-
mony (MP) as the optimality criterion in the program TNT v. 1.1
(Goloboff et al. 2008).
Based on these results (not presented), a selection of taxa
was made to provide the appropriate phylogenetic context for
a fine-scale placement of these taxa within Mycosphaerella
ceae. Additional taxa were included based on similarity and
determined using BLAST results of Genbank. In all cases,
the ITS and the nLSU sequences retrieved from GenBank to
represent a particular taxon and combined for analysis were
generated from the same culture.
Multiple sequence alignment of the ITS and the nLSU was con-
ducted within the program Geneious Pro v. 5 (Drummond et al.
2010) using MUSCLE v. 3.6 (Edgar 2004), and adjusted manu-
ally. Insertions and deletions (indels) within the concatenated
ITS and nLSU matrix were coded using the ‘simple indel coding’
method of Simmons & Ochoterena (2000) as implemented in
Gapcode.py (Ree 2007). The resulting alignment was depos-
ited into TreeBASE as accession number SN11747. Bayesian
Inference (BI) phylogenetic analysis was conducted on the
concatenated ITS/nLSU matrix, including indel characters, in
MrBayes v. 3.1.2 (Huelsenbeck & Ronquist 2001, Ronquist &
Huelsenbeck 2003). Indel data were analyzed as ‘restriction’
data type and DNA sequence data were analyzed as ‘DNA’ data
type. The GTR+I+G model of DNA sequence evolution was
determined as the best fit using the Akaike Information Criterion
(AIC; Posada & Buckley 2004) in MrModeltest v. 2.2 (Nylander
2004) and implemented in the BI analysis. The coding param-
eter for the restriction data type was set to variable. All other
parameters were left as default. The posterior probability (pp)
distribution of trees was estimated based on the results of two
independent runs of 1 million generations each of the Markov
Chain Monte Carlo (MCMC) simulation, which sampled trees
every 100 generations until the standard deviation of split fre-
quencies reached less than 0.01. The burn-in was determined
using the program Tracer v. 1.5 (Rambaut & Drummond 2007).
The remaining trees were combined and used to build a 50 %
majority rule consensus within the program FigTree v. 1.3.1
Data matrix and phylogenetic analysis
The combined ITS, nLSU, plus indel matrix contained 1 304
characters, 53 of which were indels. Of these 1 251 nucleotide
characters, 202 were variable (16.1 %) and 126 were informa-
tive (10.1 %). Thirty-one indel characters were informative
(58.5 %). The ITS alignment was 504 nucleotide positions in
length, 130 of which were variable (25.8 %) and 81 informative
(16.1 %). The nLSU alignment was 747 nucleotide positions,
72 of which were variable (9.6 %) and 45 informative (6.0 %).
The ITS alignment contributed 46 indels, 28 of which were
informative (60.9 %). The nLSU alignment contributed seven
indels, three of which were informative (42.9 %).
Preliminary MP analysis of the nLSU suggested that A. caricae
and P. guazumae occupy phylogenetic positions within the clade
with 95 % bootstrap support (Crous et al. 2009b) containing
Dothiostroma (clade 7), which is sister to the clade comprised of
Pseudocercospora-like fungi (clade 6), Phaeophleospora (clade
5), and Lecanosticta (clade 4). Our combined analysis of ITS,
nLSU, and indels resulted in strong support (pp = 1.0) for this
clade and an increased level of resolution among its members
relative to nLSU alone (Fig. 1). Within this clade, P. guazumae
occupies an unresolved position with Mycosphaerella microsora
sister to a well-supported clade (p = 0.94) containing Passalora
brachycarpa, M. ellipsoidea, M. aurantia, M. buckinghamiae, M.
africana, and A. caricae. The latter four species are members
of a well-supported (pp = 1.0) polytomy sister to Passalora
A.M. Minnis et al.: Asperisporium and Pantospora
Asperisporium Maubl., Lavoura 16: 207/212. 1913 ‘1912’, and
Bull. Trimestriel Soc. Mycol. France 29: 357. 1913. Note:
Maublanc published his article in two languages in Lavoura
and separately in Bull. Trimestriel Soc. Mycol. France. We
were unable to confirm which printed issuance was first.
Typus genericus. Asperisporium caricae (Speg.) Maubl.
Asperisporium caricae (Speg.) Maubl., Lavoura 16: 207/212.
1913 ‘1912’, and Bull. Trimestriel Soc. Mycol. France 29:
357. 1913. — Fig. 2
1886. Note: This species was published as ‘Cercospora? caricae’ to indicate
doubt as to the generic classification, but it is still valid according to ICBN
Art. 34.1 (McNeill et al. 2006).
≡ Fusicladium caricae (Speg.) Sacc., Rend. Congr. Bot. Palermo: 58.
[≡ Pucciniopsis caricae (Speg.) Höhn., Centralbl. Bakteriol., 2. Abth.: 60:
5. 1923, nom. illeg., non Earle 1902. Isonym (Speg.) Seaver, Scientific Survey
Basionym. Cercospora caricae Speg., Anales Soc. Ci. Argent. 22: 215.
of Porto Rico and the Virgin Islands, Vol. VIII, Part 1: 104. 1926.] Note: This
combination was published as ‘Pucciniopsis? caricae’ to indicate doubt as
to the generic classification, but it is still valid according to ICBN Art. 34.1.
= Scolicotrichum caricae Ellis & Everh., J. Mycol. 7: 134. 1892 as ‘Sco
= Epiclinium cumminsii Massee, Bull. Misc. Inform. Kew 1898: 133.
= Pucciniopsis caricae Earle, Bull. New York Bot. Gard. 2: 340. 1902.
Types and typifications
Lectotypus of Cercospora caricae designated by Chupp, A
Monograph of the Fungus Genus Cercospora: 106. 1953:
Paraguay, Guarapi, on leaves of Carica papaya, Feb. 1881,
coll. B. Balansa, No. 2739 (LPS).
Epitypus of Cercospora caricae hic designatus: Brazil, Inter-
cepted at USA, Washington, Seattle, entering from Brazil, on
fruit of Carica papaya, 16 Apr. 2010, coll. C. Weight, isolated by
J.F. Bischoff from BPI 880773, epitype is a dried culture on SDA
(BPI 881135); ex-epitype CBS 130298; GenBank accession
nos: ITS (JN190955), LSU (JN190953), RPB2 (JN190951).
Fig. 1 A 50 % majority rule Bayesian Inference phylogram resulting from analysis of the ITS, the nLSU, and associated indels. The scale bar is proportional
to the amount of character evolution on the tree. The ITS and nLSU Genbank accession numbers are provided on terminals, in that order, after each taxon
name. Only a single accession is labelled on terminals where the Genbank record contains concatenated ITS and nLSU sequences. Taxon names in bold
Mycosphaerella microsora EU167599, EU167599
Lecanosticta acicola GU214663, KQ852598
Passalora sp. GU214668, GQ852623
Asperisporium caricae JN190955, JN190953
Mycosphaerella madeirae AY725553, DQ204756
Dothistroma pini AY808302, GQ852596
Mycosphaerella aurantia AY725531, DQ246256
Mycosphaerella africana AF173314, GQ852601
Pantospora guazumae JN190956, JN190954
Mycosphaerella ellipsoidea AY725545, GQ852602
Mycosphaerella keniensis AF173300, GQ852610
Mycosphaerella endophytica DQ302953, GQ852603
Mycosphaerella laricina EU167595, EU167595
Mycosphaerella buckinghamiae EU707855
Passalora daleae EU040236, EU040236
Ramularia miae DQ885902
Phaeophleospora eugeniae FJ493188, FJ493206
Phaeophleospora eugeniicola FJ493190, FJ493208
Passalora ageratinae GU214639, GU214453
Ramularia endophylla EU167569
Passalora brachycarpa GU214664
Persoonia – Volume 27, 2011
Fig. 2 Cultures and microscopic features of Asperisporium caricae. a–d. Ex-epitype (CBS 130298) at approximately 1 mo at 24 °C with a 12 h light/dark
regimen: a. PDA; b. reverse on PDA; c. SDA; d. reverse on SDA. — e. Lectotype packet, No. 2739 (LPS). — f–m. Ex-epitype (CBS 130298) on SDA; f–h.
conidiophores and conidia; i–l. conidia; m. spermatia. — Scale bars = 10 µm for all.
A.M. Minnis et al.: Asperisporium and Pantospora
and no. 3855, in the protologue without indicating either as type.
These are syntypes according to ICBN Art. 9.4. Chupp (1953)
lectotypified the species via ICBN Art. 7.11 when he indicated
the word type for no. 2739 at LPS.
Notes — Spegazzini (1886) cited two collections, no. 2739
Description of the epitype (preserved culture) and
collection from which it was isolated
Fructicolous with spots scattered, 3–4 mm diam. Sporodochia
formed on stromata, immersed becoming erumpent, punctiform,
blackish to black. In culture on SDA, sporodochia of loosely to
densely arranged conidiophores produced on darkly pigmented
stromata. Conidiophores macronematous, mononematous,
simple or less commonly branched, more or less straight to
slightly sinuous, smooth, brownish, septate, 58–168 × 5–10
µm. Conidiogenous cells integrated, terminal, polyblastic,
sympodial, cylindrical to clavate, at times slightly geniculate,
conidiogenous loci thickened, darkened. Conidia 14–22 × 8–13
µm, solitary, broadly ellipsoid, ellipsoid, obovate, pyriform, or
oblong and somewhat clavate, smooth becoming verrucose,
hyaline becoming brownish, each typically 0–1-septate with one
more or less median septum, rarely with 2 septa, hila thickened,
darkened, occasionally on an apiculus-like base. Spermogonia
with spermatogenous cells cylindrical, lageniform, or ampulli-
form, smooth, hyaline. Spermatia 6–10 × 1–2.5 µm, solitary,
cylindrical, clavate, ellipsoid, sigmoid, or stocking-shaped,
apices obtuse, bases truncate, smooth, hyaline, aseptate, at
times with a frill-like structure below basal septum.
Culture characteristics — Colonies on PDA 9–10 mm after
1 mo at 24 °C with a 12 h light/dark regimen; mycelium forming
a raised mound composed of tiers of smaller mounds, surface
slightly velutinous, near dark green (30F3–4), at times por-
tions covered with whitish, short erect hyphae or whitish aerial
hyphae; with scattered black spherical structures, spermogo-
nia, 80–160 µm diam; margin lobed, whitish to concolorous;
reverse concolorous. Surface mycelium with hyphae branching,
walls smooth, hyaline to brownish, septate, 3–6.5 µm diam.
Sporulation within approximately 1 mo, but precise time not
noted. Conidial production on PDA sparse in comparison to
that on SDA.
Colonies on SDA 8–9 mm after 1 mo at 24 °C with a 12 h light/
dark regimen; mycelium forming a raised mound composed of
tiers of smaller mounds, surface velutinous, near dark green
(30F3) to greenish grey (30F2), at times portions covered with
whitish, short erect hyphae, dense lanose to cottony whitish
hyphae, or whitish aerial hyphae; with scattered black spheri-
cal structures, spermogonia, 80–160 µm diam; margin lobed,
more or less concolorous; reverse near brownish orange (6C7).
Surface mycelium with hyphae branching, walls smooth, hyaline
to brownish, septate, 3–6.5 µm diam. Sporulation confirmed
at 3 wk.
Colonies on both media showing slightly increased growth rates
on a 12 h black light/dark regimen. Sporulation more or less
as above. Colonies exposed to UV light via the transilluminator
showed increased growth rate as submerged, uncoloured
hyphae with a corresponding lack of mycelium above the agar
rarely Carica chilensis (≡ Vasconcellea chilensis) (Caricaceae)
(Ellis & Holliday 1972, Crous & Braun 2003). This fungus is
widely distributed in subtropical and tropical regions with reports
from Africa, Asia, North America, Oceania, and South America
(Crous & Braun 2003, Farr & Rossman 2011).
Habitat & Distribution — Leaves and fruits of Carica papaya,
Carica papaya, 24 June 1921, coll. W.H. Whetzel (BPI 424787). – Brazil, Inter-
cepted at USA, Washington, Seattle, entering from Brazil, on fruit of Carica
Specimens examined. Bermuda, Paget, Agricultural Station, on leaves of
papaya, 16 Apr. 2010, coll. C. Weight (BPI 880773); culture isolated by J.F.
Bischoff from BPI 880773, dried culture on SDA (BPI881135, designated
epitype). – ColomBia, Intercepted at New York, J.F.K.I.A., #24116, on fruit of
Carica papaya, 4 Jan. 1969, coll. C. Smock (BPI 424779). – CuBa, Santiago
de Las Vegas, Experiment Station, on leaves of Carica papaya, 8 May
1915, coll. R.A. Jehle (BPI 424780). – dominiCan repuBliC, Valle del Cibao,
Santiago Prov., Santiago, Hato del Yaque, Gardens, on leaves of Carica
papaya, 19 Jan. 1931, coll. R. Ciferri, Mycoflora Domingensis Exsiccata
311 (BPI 424783). – Honduras, Jamastran, on leaves of Carica papaya, 10
Mar. 1964, coll. A.S. Muller (BPI 424789). – mexiCo, Intercepted at Texas,
Laredo, #008382, on fruit of Carica papaya, 20 Feb. 1975, coll. S. Kendall
(BPI 424778). – puerto riCo, Yauco, on leaves of Carica papaya, 30 Mar.
1916, coll. W.H. Whetzel and E.W. Olive (BPI 424777). – Venezuela, Federal
District, Valle de Puerto La Cruz, El Limón, on leaves of Carica papaya, 15
Jan. 1925, coll. H. Sydow (BPI 1112163).
is a cut portion of fruit. Unfortunately, it could not be well pre-
served before it was overgrown with anamorphic fungi including
Penicillium. Thus, we have designated a dried culture isolated
from this collection as the epitype. Conidiophores in culture
were longer than those observed on host tissues (Ellis 1971,
Ellis & Holliday 1972).
Ellis & Holliday (1972) listed three taxonomic synonyms. Exa-
mination of the protologues of these names supports the syno-
nymy. Maublanc (1913a, b) described a teleomorph that he
suggested was associated with Asperisporium caricae as
Sphaerella caricae. This morph was later classified as Myco
sphaerella caricae (Maubl.) Hansf., which makes it an il-
legitimate later homonym of Mycosphaerella caricae Syd. &
P. Syd. These teleomorphic names have been considered to be
conspecific and associated with the anamorphic Phoma caricae
(Sivanesan 1984). Others expressed doubt that A. caricae
has a known teleomorph since the connection has never been
proven (Crous & Braun 2003). Recently, M. caricae Syd. & P. Syd.
and M. caricae (Maubl.) Hansf. were listed separately with each
having different anamorphs, but the type of the latter was not
studied (Aptroot 2006). Mycosphaerella caricae Syd. & P. Syd.
was transferred to Stagonosporopsis with no mention of the
later homonym in the synonym list (Aveskamp et al. 2010).
No additional information about the life cycle of A. caricae in
regards to its ascal state and the synonymy of these historical
names can be added by this study. However, a structure of
unproven identity that is presumed to be a spermogonium was
found in culture. Its presumptive spermatogenous cells and
spermatia are reminiscent of those described by Crous (1998)
for Mycosphaerella crystallina. The presumptive spermatia did
not germinate on PDA. The presence of spermogonia suggests
that A. caricae has a sexual stage in its life cycle.
Notes — The collection from which the epitype was isolated
Pantospora Cif., Ann. Mycol. 36: 242. 1938.
Typus genericus. Pantospora guazumae Cif.
Micologia, Publicação No. 372: 13. 1962.
= Dictyocephala A.G. Medeiros, Universidade do Recife, Instituto de
referenced via the invalid Dictyocephala ulmifoliae (Obreg.-Bot.) A.G. Me-
deiros (ICBN Art. 10.3).
Typus genericus. Cercospora ulmifoliae Obreg.-Bot., which was indirectly
Pantospora guazumae Cif., Ann. Mycol. 36: 242. 1938. —
do Recife, Instituto de Micologia, Publicação No. 372: 13. 1962 as ‘ulmifolii’,
nom. inval. via ICBN Art. 33.4.
≡ Pseudocercospora ulmifoliae (Obreg.-Bot.) U. Braun & Crous, CBS
Biodiversity Series 1: 415. 2003. Note: Pseudocercospora guazumae (Syd.)
Deighton prevented a legitimate combination based on Pantospora guazu
mae (Crous & Braun 2003).
= Cercospora ulmifoliae Obreg.-Bot., Caldasia 1: 51. 1941.
≡ Dictyocephala ulmifoliae (Obreg.-Bot.) A.G. Medeiros, Universidade
Persoonia – Volume 27, 2011
Types and typifications
Lectotypus of Pantospora guazumae designated by Deighton,
Mycol. Pap. 140: 159. 1976: Dominican Republic, Valle del
Cibao, prov. Santiago, Hato del Yaque, on leaves of Guazuma
ulmifolia, 20 Apr. 1930, coll. R. Ciferri & A.M. Borgna Ciferri,
Batey no. 1, Mycoflora Domingensis Exsiccata 210 (IMI 59269,
Epitypus of Pantospora guazumae hic designatus: Mexico,
Intercepted at USA, Arizona, Nogales, entering from Mexico,
on leaf of Guazuma ulmifolia, 12 Feb. 2009, coll. J. Moore (BPI
880778); ex-epitype CBS 130299; GenBank accession nos: ITS
(JN190956), LSU (JN190954), RPB2 (JN190952).
number 210 in exsiccatae sets. These are syntypes accord-
ing to ICBN Art. 9.4. Deighton (1976) lectotypified the species
when he indicated that the collection at IMI was the type. This
specimen is now housed at K.
Notes — Ciferri (1938) distributed the original material as
Fig. 3 Cultures and microscopic features of Pantospora guazumae. a–d. Ex-epitype (CBS 130299) at approximately 1 mo at 24 °C with a 12 h light/dark
regimen: a. PDA; b. reverse on PDA; c. SDA; d. reverse on SDA. — e. Leaf spot on abaxial surface of Guazuma ulmifoliae (BPI 880778, designated epitype);
f. synnema on Guazumae ulmifoliae (BPI 880778, designated epitype); g–i. conidia from ex-epitype (CBS 130299) on PDA. — Scale bars = 1 mm for e, 10
µm for f–i.
A.M. Minnis et al.: Asperisporium and Pantospora
Description of the epitype
Leaf spots scattered, 1.5–2 mm diam, visible on both adaxial
and abaxial surfaces, typically circular, dark brown to blackish
with a distinct, lighter, occasionally purplish, margin on adaxial
surface, discoloured brown with a distinct, lighter, occasionally
purplish, margin on abaxial surface. Caespituli hypophyllous,
scattered within margin of leaf spots, conidiophores densely
aggregated forming synnemata. Synnemata basistromatic with
immersed stromata, erect, more or less even with hyphal tips
spreading apart at apex, up to 290 µm long, up to 40 µm wide
along non-apical portion. Conidiophores unbranched, more or
less straight to somewhat sinuous with apices more or less
obtuse, often interwoven, smooth, pale brown to brown, septate,
3–6 µm, widest towards synnematal apex. Conidiogenous cells
integrated, terminal, blastic, slightly verrucose and with annel-
lations, conidiogenous loci visible, not significantly thickened
or darkened. Conidia up to 43 × 13 µm, solitary, versiform, el-
lipsoid with short beaks to obclavate, slightly verrucose, brown,
with multiple transverse, longitudinal, and occasionally oblique
septa, hila visible but not thickened or darkened.
1 mo at 24 °C with a 12 h light/dark regimen; mycelium forming
a raised mound composed of tiers of smaller mounds, surface
lanose, near brownish orange (7C7) to light brown (7D7); mar-
gin lobed, more or less concolorous; reverse almost black or
black; medium becoming discoloured with a greyish red (7B6)
to orange (6B7) soluble pigment. Surface mycelium with hy-
phae branching, walls smooth, at times covered with orange
crystals, hyaline to brownish orange in Shear’s mounting fluid,
purple in 3 % KOH, septate, 5–6.5 µm diam. Sporulation not
Colonies on SDA 11–13 mm after 1 mo at 24 °C with a 12 h light/
dark regimen; mycelium forming a raised mound composed
of tiers of smaller mounds, surface lanose, multicoloured with
near brownish orange (7C7) to light brown (7D4) to orangish
tan to greyish hues; margin lobed, more or less concolorous;
reverse almost black or black; medium becoming discoloured
with a greyish red (7B6) to orange (6B7) soluble pigment. Sur-
face mycelium with hyphae branching, walls smooth, at times
covered with orange, pigmentary crystals, hyaline, orangish to
brownish orange in Shear’s mounting fluid, purple in 3 % KOH,
septate, 5–6.5 µm diam. Sporulation not observed.
Colonies on both media showing increased growth rates on a
12 h black light/dark regimen. Sporulation not observed. Colo-
nies exposed to UV via the transilluminator showed increased
growth rate, a significant increase in submerged hyphae with dark
pigmentation at margins with a corresponding lack of mycelium
above, and production of a few scattered synnemata. Mature
synnemata were observed approximately 1.5 wk after the UV
regimen was initiated. Synnemata in culture were of greater width
and fascicles of conidiophores were somewhat more loosely
arranged than on host leaves. Conidia 32–60 × 8–16 µm, versi-
form, ellipsoid with short beaks, obclavate, cylindrical-obclavate,
or more or less cylindrical, slightly verrucose, brown, with multiple
transverse, longitudinal, and occasionally oblique septa. Both
dictyosporous and scolecosporous conidia present, additional
longitudinal and/or oblique septa may develop over time. Other-
wise, synnemata similar in culture and on host leaves.
Culture characteristics — Colonies on PDA 12–13 mm after
vaceae). According to Farr & Rossman (2011), this fungus is
known from North America (Cuba, Dominican Republic) and
South America (Brazil, Colombia). This is the first report of this
species from Mexico.
Habitat & Distribution — Leaves of Guazuma ulmifolia (Mal
folia, June 1938, coll. C. GarcesOrejuela (BPI 445535). – dominiCan repu
Specimens examined. ColomBia, Rioclaro, near Cali, on Guazuma ulmi
BliC, Valle del Cibao, prov. Santiago, Hato del Yaque, on leaves of Guazuma
ulmifolia, 20 Apr. 1930, coll. R. Ciferri & A.M. Borgna Ciferri, Batey no. 1,
Mycoflora Domingensis Exsiccata 210 (IMI 59269, K(M) 169346, lecto-
type); (BPI 445536, syntype); Feb. 1932, coll. R. Ciferri (BPI 445537); 01
Sept. 1931, coll. R. Ciferri (BPI 445538); 01 Sept. 1931, coll. R. Ciferri (BPI
445539). – mexiCo, Intercepted at USA, Arizona, Nogales, entering from
Mexico, on leaf of Guazuma ulmifolia, 12 Feb. 2009, coll. J. Moore (BPI
880778, designated epitype).
herbarium material. The epitype specimen is not fully mature
as it possesses small leaf spots and lacks a large number of
mature conidia. In the interest of preserving the specimen, the
sample size of measured structures is relatively small. Discrep-
ancies with the observations of previous authors should not be
considered significant. As with and loosely following Medeiros
(1962), UV was found to stimulate the production of synnemata
Both Deighton (1976) and Crous & Braun (2003) considered Cer
cospora ulmifoliae as a synonym of Pantospora guazumae
even though they did not study Obregón-Botero’s original
material of C. ulmifoliae. Based on the descriptions provided
by Obregón-Botero (1941) and Chupp (1953), there can be no
doubt that the two names are synonymous. There is no known
sexual stage for Pantospora guazumae.
Notes — This epitype description is based on sparse, dried
Asperisporium has been thought of as a likely synonym of
Passalora since the two genera were separated on the ba-
sis of conidial surface ornamentation (Crous & Braun 2003,
Schubert & Braun 2005). These authors tentatively maintained
them as distinct due to the absence of DNA sequence data.
The phylogenetic analyses place Asperisporium caricae, the
type of the genus, in a relatively close relationship with several
species of Passalora including P. brachycarpa. Sequence data
are available for only a small number of the approximately 580
species of Passalora (Kirk et al. 2008) and the type species
of the genus, Passalora bacilligera, has not been sequenced
and placed phylogenetically. As Passalora currently stands, it
is a polyphyletic genus (Crous et al. 2009b). A final conclusion
cannot be made on the status of Asperisporium. The collective
works of Patil & Thirumalachar (1966), Ellis (1976), Barreto &
Evans (1995), Braun (2000a, b) and Braun & Crous (2007)
cover nearly all of the remaining species that are currently
classified in Asperisporium.
Crous & Braun (2003) formally classified Pantospora guazumae
in Pseudocercospora, noted the lack of molecular data for it and
explained why the presence of dictyospores may not be a distin-
guishing character at the rank of genus. Based on the phyloge-
netic analyses, Pantospora is not closely related to two clades
presented by Crous et al. (2009b), namely, Pseudocercospora
including the type species, P. vitis (clade 16) and Pseudo
cercospora-like (clade 14). Pantospora is in an unresolved
position near Mycosphaerella microsora. Pantospora appears
to be a distinct lineage.
In summary, the type species of Asperisporium, A. caricae,
and Pantospora, P. guazumae, have been placed phyloge-
netically in the Mycosphaerellaceae. Studies of their culture
characteristics were made and both species were epitypified
with herbarium material associated with living cultures and
DNA sequence data. The phylogenetic placement of these
genera demonstrates that previous generic concepts and the
perceived values of particular morphological features were not
necessarily congruent with phylogeny. Since this is becoming a
repeated result (Crous & Braun 2003, Crous et al. 2007, 2009b,
Crous 2009) and that sampling in terms of taxa and numbers of
genetic loci remains small for such a large family, we advocate
Persoonia – Volume 27, 2011
a conservative and patient approach towards the creation of
new taxonomic schemes and nomenclatural novelties in this
group of fungi.
Acknowledgements We thank Jorge A. Chayle and other staff at LPS for
providing information about and images of the original material of Asperispo
rium caricae and permission to reproduce the image of the lectotype packet.
Curators at K are acknowledged for their loan of the lectotype of Pantospora.
Pedro W. Crous and Uwe Braun are thanked for their comments about pre-
liminary phylogenetic trees and generic concepts. Amy Y. Rossman provided
helpful comments to improve the manuscript prior to its submission.
Aptroot A. 2006. Mycosphaerella and its anamorphs: 2. Conspectus of
Mycosphaerella. CBS Biodiversity Series 5: 1–231.
Aveskamp M, Gruyter H de, Woudenberg J, Verkley G, Crous PW. 2010. High-
lights of the Didymellaceae: A polyphasic approach to characterize Phoma
and related pleosporalean genera. Studies in Mycology 65: 1–60.
Baker WA, Partridge EC, Morgan-Jones G. 2000. Notes on hyphomycetes.
LXXVIII. Asperisporium sequoiae, the causal organism of conifer needle
blight, reclassified in Cercosporidium, with comments on the status of the
genus. Mycotaxon 76: 247–256.
Barreto RW, Evans HC. 1995. The mycobiota of the weed Mikania micrantha
in southern Brazil with particular reference to fungal pathogens for biological
control. Mycological Research 99: 343–352.
Braun U. 2000a. Miscellaneous notes on some micromycetes. Schlechten-
dalia 5: 31–56.
Braun U. 2000b. Annotated list of Cercospora spp. described by C. Spegaz-
zini. Schlechtendalia 5: 57–79.
Braun U, Crous PW. 2007. The diversity of cercosporoid hyphomycetes –
new species, combinations, names and nomenclatural clarifications. Fungal
Diversity 26: 55–72.
Chupp C. 1953. A monograph of the fungus genus Cercospora. Published
by the author, Ithaca, New York.
Ciferri R. 1938. Mycoflora domingensis exsiccata. Annales Mycologici 36:
Crous PW. 1998. Mycosphaerella spp. and their anamorphs associated with
leaf spot diseases of Eucalyptus. Mycological Memoirs 21: 1–170.
Crous PW. 2009. Taxonomy and phylogeny of the genus Mycosphaerella
and its anamorphs. Fungal Diversity 38: 1–24.
Crous PW, Braun U. 2003. Mycosphaerella and its anamorphs: 1. Names pub-
lished in Cercospora and Passalora. CBS Biodiversity Series 1: 1–571.
Crous PW, Braun U, Groenewald JZ. 2007. Mycosphaerella is polyphyletic.
Studies in Mycology 58: 1–32.
Crous PW, Summerell BA, Carnegie AJ, Wingfield MJ, Groenewald JZ.
2009a. Novel species of Mycosphaerellaceae and Teratosphaeriaceae.
Persoonia 23: 119–146.
Crous PW, Summerell BA, Carnegie AJ, Wingfield MJ, Hunter GC, Burgess
TI, Andjic V, Barber PA, Groenewald JZ. 2009b. Unravelling Mycosphae-
rella: do you believe in genera? Persoonia 23: 99–118.
Deighton FC. 1976. Studies on Cercospora and allied genera. VI. Pseudo-
cercospora Speg., Pantospora Cif. and Cercoseptoria Petr. Mycological
Papers 140: 1–168.
Drummond AJ, Ashton B, Buxton S, Cheung M, Cooper A, Heled J, Kearse M,
Moir R, Stones-Havas S, Sturrock S, Thierer T, Wilson A. 2010. Geneious
v. 5.1, Available from www.geneious.com.
Edgar RC. 2004. MUSCLE: multiple sequence alignment with high accuracy
and high throughput. Nucleic Acids Research 32: 1792–1797.
Ellis MB. 1971. Dematiaceous hyphomycetes. Commonwealth Mycological
Institute, Kew, Surrey, England.
Ellis MB. 1976. More dematiaceous hyphomycetes. Commonwealth Myco-
logical Institute, Kew, Surrey, England.
Ellis MB, Holliday P. 1972. Asperisporium caricae. CMI Descriptions of
Pathogenic Fungi and Bacteria 347: 1–2.
Farr DF, Rossman AY. 2011. Fungal databases, systematic mycology and
microbiology laboratory, ARS, USDA. Retrieved March 30, 2011 from
Goloboff PA, Farris JS, Nixon KC. 2008. TNT, a free program for phylogenetic
analysis. Cladistics 24: 774–786.
Graham SO. 1959. The effects of various reagents, mounting media, and
dyes on the teliospore walls of Tilletia controversa Kühn. Mycologia 51:
Huelsenbeck JP, Ronquist F. 2001. MrBayes: Bayesian inference of phylo-
geny. Bioinformatics 17: 754–755.
Kirk PM, Cannon PF, Minter DW, Stalpers JA with assistance. 2008. Ains-
worth & Bisby’s dictionary of the fungi, tenth edition. Cab International,
Kornerup A, Wanscher JH. 1967. Methuen handbook of color, 2nd ed.
Methuen & Co Ltd, London, UK.
Largent D, Johnson D, Watling R. 1977. How to identify mushrooms to genus
III. Microscopic features. Mad River Press, Eureka, CA.
Liu YJ, Whelen S, Hall BD. 1999. Phylogenetic relationships among asco-
mycetes: evidence from an RNA polymerase II subunit. Molecular Biology
and Evolution 16: 1799–1808.
Malkus A, Chang P-FL, Sabina MZ, Chung K, Shao J, Cunfer BM, Arseniuk
E, Ueng PP. 2006. RNA polymerase II gene (RPB2) encoding the second
largest protein subunit in Phaeosphaeria nodorum and P. avenaria. Myco-
logical Research 110: 1152–1164.
Maublanc A. 1913a ‘1912’. Sobre uma molestia do mamoeiro (Caryca
Papayal, L.)/Sur une maladie des feuilles du papayer “Carica papaya”.
Lavoura 16: 204–212.
Maublanc A. 1913b. Sur une maladie des feuilles du papayer (Carica
papaya). Bulletin Trimestriel de la Société Mycologique de France 29:
McNeill J, Barrie FR, Burdet HM, Demoulin V, Hawksworth DL, Marhold K,
Nicolson DH, Prado J, Silva PC, Skog JE, Wiersema JH, Turland NJ (eds).
2006. International code of botanical nomenclature (Vienna code): Adopted
by the Seventeenth International Botanical Congress, Vienna, Austria, July
2005. Gantner, Ruggell, Liechtenstein.
Medeiros AG. 1962. Dictyocephala novo género de fungos Dematiaceae. Uni-
versidade do Recife, Instituto de Micologia, Publicação No. 372: 1–24.
Monclavo JM, Lutzoni FM, Rehner SA, Johnson J, Vilgalys R. 2000. Phyloge-
netic relationships of agaric fungi based on nuclear large subunit ribosomal
DNA sequences. Systematic Biology 49: 278–305.
Nylander JAA. 2004. MrModeltest, v. 2. Program distributed by the author.
Evolutionary Biology Centre, Uppsala University.
Obregón-Botero R. 1941. Cuatro nuevos deuteromicetos Colombianos.
Caldasia 1: 49–51.
Patil BV, Thirumalachar MJ. 1966. Studies on some fungi of Masharashtra-
India-I. Sydowia 20: 33–38.
Posada D, Buckley TR. 2004. Model selection and model averaging in
phylogenetics: advantages of akaike information criterion and Bayesian
approaches over likelihood ratio tests. Systematic Biology 53: 793–808.
Rambaut A. 2009. FigTree v. 1.3.1. Computer program and documentation
distributed by the author at http://tree.bio.ed.ac.uk/software/.
Rambaut A, Drummond AJ. 2007. Tracer v. 1.5. Computer program and docu-
mentation distributed by the authors at http://beast.bio.ed.ac.uk/Tracer.
Ree RH. 2007. Gapcode.py, v. 2.0. Distributed by the author at http://www.
Reeb V, Lutzoni F, Roux C. 2004. Multilocus phylogenetic circumscription of
the lichen-forming fungi family Acarosporaceae and its position within the
Ascomycota. Molecular Phylogeny and Evolution 32: 1036–1060.
Ronquist F, Huelsenbeck JP. 2003. MrBayes 3: Bayesian phylogenetic infer-
ence under mixed models. Bioinformatics 19: 1572–1574.
Schubert K, Braun U. 2005. Taxonomic revision of the genus Cladosporium s.l.
4. Species reallocated to Asperisporium, Dischloridium, Fusicladium, Passa-
lora, Pseudoasperisporium and Stenella. Fungal Diversity 20: 187–208.
Simmons MP, Ochoterena R. 2000. Gaps as characters in sequence-based
phylogenetic analyses. Systematic Biology 49: 369–381.
Sivanesan A. 1984. The bitunicate ascomycetes and their anamorphs.
Gantner Verlag, Vaduz, Liechtenstein.
Spegazzini C. 1886. Fungi Guaranitici. Pugillus I. Anales de Sociedad
Científica Argentina 22: 186–224.
Stevens HE. 1939. Papaya diseases. Proceedings of the Florida State Horti-
cultural Society 52: 57–63.
Thiers B. 2011 [continuously updated]. Index Herbariorum: A global directory
of public herbaria and associated staff. New York Botanical Garden’s Virtual
Verkley GJM, Crous PW, Groenewald JZ, Braun U, Aptroot A. 2004. Myco-
sphaerella punctiformis revisited: morphology, phylogeny, and epitypification
of the type species of the genus Mycosphaerella (Dothideales, Ascomy-
cota). Mycological Research 108: 1271–1282.
Vilgalys R, Hester M. 1990. Rapid genetic identification and mapping of en-
zymatically amplified ribosomal DNA from several Cryptococcus species.
Journal of Bacteriology 172: 4238–4246.
White TJ, Bruns T, Lee S, Taylor JW. 1990. Amplification and direct sequenc-
ing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand
DH, Sninsky JJ, White TJ (eds), PCR protocols: A guide to methods and
applications. Academic Press, Inc., New York, New York: 315–322.