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Accepted by Eric McKenzie: 25 Mar. 2014; published: 20 Aug. 2014
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PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN
1179-3163
(online edition)
Copyright © 2014 Magnolia Press
Phytotaxa 176 (1): 092–101
www.mapress.com
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phytotaxa
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Article
http://dx.doi.org/10.11646/phytotaxa.176.1.11
Clavatispora thailandica gen. et sp. nov., a novel taxon of Venturiales
(Dothideomycetes) from Thailand
SARANYAPHAT BOONMEE
1
, JAYARAMA D. BHAT
2
, SAJEEWA S.N. MAHARACHCHIKUMBURA
1
&
KEVIN D. HYDE
1
1
Institute of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand; School of Science, Mae Fah
Luang University, Chiang Rai 57100, Thailand
2
Formerly at Department of Botany, Goa University, Goa 403 206, India
email: saranyaphat.khag@gmail.com; kdhyde3@gmail.com
Abstract
An unusual wood-inhabiting fungus was found in northern Thailand. The combination of characters of this taxon, viz.
setose, ostiolate ascomata, bitunicate asci, dark brown, wedge-shaped to clavate, muriform ascospores, and a
hyphomycetous asexual state producing ellipsoidal, septate, highly constricted, holoblastic conidia is not found in any
hitherto described genus of Dothideomycetes. An LSU phylogeny showed this taxon to be clearly placed in the family
Sympoventuriaceae, Venturiales. The name Clavatispora thailandica gen. and sp. nov., is introduced to accommodate
this taxon, which is described and illustrated and compared with other genera in this family.
Key words: bitunicate asci, hyphomycetous asexual state, molecular phylogeny, muriform ascospores,
Sympoventuriaceae
Introduction
Pleosporales is the largest order in the class Dothideomycetes that comprises 41 families and 181 genera (Hyde et
al. 2013). In this order, the family Pleosporaceae contains several genera, e.g. Clathrospora, Lewia,
Platysporoides, Pleospora, and Pyrenophora that have brown muriform ascospores (Sivanesan 1984, Barr 1987,
1990a, Zhang et al. 2012, Hyde et al. 2013). Species with muriform ascospores are also found in Aigialaceae,
Cucurbitariaceae, Halojulellaceae, Montagnulaceae, Platystomaceae, Pleomassariaceae, Shiraiaceae and
Teichosporaceae, thus this character has evolved on numerous occasions even outside Pleosporales (e.g.
Gloniopsis, Hysterobrevium, Hysterographium and other genera in Hysteriales). Barr (1990a) placed species with
muriform ascospores in 12 families within the Pleosporales, i.e. Arthopyreniaceae, Cucurbitariaceae,
Dacampiaceae, Hysteriaceae, Leptosphaeriaceae, Lophiostomataceae, Micropeltidaceae, Phaeosphaeriaceae,
Phaeotrichaceae, Pleomassariaceae, Pleosporaceae and Pyrenophoraceae. Species in these families have
differing ascomata characters and asexual states (Barr 1990a, 1990b, Zhang et al. 2012, Hyde et al. 2013).
In our ongoing study of Dothideomycetes of Thailand (Boonmee et al. 2011, 2012, Chomnunti et al. 2011,
2012a,b, Liu et al. 2011, 2012), we collected an intriguing species with ascospores resembling Alternaria-like
conidia and bitunicate asci indicative of Pleosporaceae. The superficial, solitary, ascomata with dark, septate setae,
forming on a basal subiculum, and bright-coloured ostiole, indicated that this fungus differed from other
dictyosporous genera in families of the Pleosporales (Simmons 1986, Barr 1987, 1990a, 1992). The fungus also
developed a Trichocladium-like asexual state in culture. The aim of this paper is to introduce this unusual
ascomycete, which is described as Clavatispora thailandica gen. nov., sp. nov. It is only described and illustrated
but also compared with morphologically similar genera. We sequenced our isolate to establish the phylogenetic
placement of our taxon. The trees generated using LSU region place our new genus in the Sympoventuriaceae,
Venturiales.
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Materials and methods
Collection, examination and isolation of fungi
Dead herbaceous stems were randomly collected from Doi Tung forest, Chiang Rai, Thailand. Microscopic
studies were carried out and the fungus illustrated following the procedures outlined in Boonmee et al. (2011).
Type specimens are deposited at the herbarium of Mae Fah Luang University (MFLU), Chiang Rai, Thailand and
cultures are deposited at Mae Fah Luang University Culture Collection (MFLUCC), BIOTEC Culture Collection
(BCC) and IFRD culture collections, International Fungal Research & Development Centre, Kunming, China, the
latter under MTA no. 4/2010.
Single spore isolation was carried out following the methods outlined by Chomnunti et al. (2011). Germinating
ascospores were transferred to fresh malt extract agar (MEA, Difco Laboratories, Detroit, Michigan, USA) and
incubated at 28°C for one week in darkness. When colonies appeared, subcultures were made onto fresh MEA
media at room temperature. Morphology and cultural characters of the fungus were recorded based on MEA
cultures after 60 days (Boonmee et al. 2011, 2012).
Molecular procedures
Fresh mycelium was scraped from the surface of colonies grown on MEA for 60 days at 28°C. DNA
extraction, PCR amplification and sequencing of LSU region using primer pair LROR/LR5 was performed under
the conditions described in Boonmee et al. (2011). Sequencing was done in Shanghai Sangon Biological
Engineering Technology & Services Co., China and new sequences generated in this study were blasted to search
for related taxa in GenBank database (www.ncbi.nlm.nih.gov/blast/).
Phylogenetic analyses
Sequence data were aligned with Clustal W (Thompson et al. 1994) and manually adjusted using BioEdit (Hall
1999). Most of the taxa used in this study are derived from Schoch et al. (2009) and Zhang et al. (2011). The LSU
dataset for showing the placement of Clavatispora thailandica comprises 40 taxa including Acrospermum
compressum (Acrospermales) as an outgroup (Table 1). Poorly aligned regions on nucleotide multiple alignments
were removed using Gblock 0.91b (Castersana 2000), following nonconservative settings; the maximum number
of contiguous nonconserved positions was set to 10, the minimum length blocks set to 5, allowing for gap positions
in half the sequences. The construction of the maximum likelihood (ML) analysis using RAxML version 7.6.3
(Stamatakis 2006, Stamatakis et al. 2008) as part of the “RAxML-HPC BlackBox on TG tool” was performed at
the CIPRES Science Gateway V. 3.3 (http://www.phylo.org/portal2/, Miller et al. 2010). RAxML rapid
bootstrapping and subsequent ML search used distinct model/data partitions with joint branch length optimization,
executing 1,000 rapid bootstrap inferences and thereafter a thorough ML search. All free model parameters were
estimated by RAxML and ML estimate of 25 per site rate categories. Likelihood of final tree were evaluated and
optimized under GAMMA. GAMMA Model parameters were estimated to an accuracy of 0.1000000000 Log
Likelihood units. Every 100th tree was saved. Phylogenetic trees were illustrated in Treeview (Page 1996).
Bayesian posterior probability analysis was constructed with the MrBayes 3.1.2 run on XSEDE at the CIPRES
webportal (Ronquist & Huelsenbeck 2003), using the parameter setting of 2 parallel runs, 4 chains, run for
4,000,000 generations, sample frequency every 1,000 generations and all other parameters were left as default. The
50% majority rule consensus tree was created from the remaining trees in Treeview (Page 1996).
Results
Phylogenetic studies
The RAxML phylogenetic tree (Fig. 1) was constructed through analysis of the LSU gene region and comprised 40
taxa and 754 aligned characters out of 4% of ambiguous characters. The RAxML phylogram received the best
scoring tree with a final ln value of -4136.348470. The isolate of Clavatispora thailandiaca (MFLUCC 100107)
clustered with Scolecobasidiella avellanea (CBS 772.73) and grouped within the family Sympoventuriaceae with
high support values (98% BS and 1.00 PP). Additionally, taxa from the genera Ochroconis and Scolecobasidium
formed a sister-group to Clavatispora. To confirm the placement of new genus we built a further tree including
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Dothideomycetes and some species of Trichocladium (Sordariales) (tree not shown). Results indicate that
Clavatispora belongs to order Venturiales and is well separated from taxa in Pleosporales.
TABLE 1. Taxa included in the phylogenetic study.
Species Culture no. GenBank Accession number
LSU
Acrospermum compressum M 151 EU940084
Apiosporina collinsii CBS 118973 GU301798
Apiosporina morbosa dimosp EF114694
Caproventuria hanliniana ATCC 96019 AF050290
Caproventuria hystrioides CBS 117727 EU035459
Clavatispora thailandiaca MFLUCC 100107 KF770458
Coleroa robertiani CBS 458.64 JQ036231
Fusicladium africanum CPC 12828 EU035423
Fusicladium amoenum CBS 254.95 EU035425
Fusicladium intermedium CBS 110746 EU035432
Fusicladium proteae CPC18282 JN712551
Gibbera conferta CBS 191.53 GU301814
Metacoleroa dickiei medipc EF114695
Metacoleroa dickiei Kruys503 (UPS) DQ384100
Ochroconis calidifluminalis IFM54738 AB385698
Ochroconis constricta NBRC9375 AB564619
Ochroconis gallopava IFM54737 AB272164
Ochroconis humicola NBRC32054 AB564618
Ochroconis tshawytschae CBS100438 GU328005
Protoventuria barriae CBS 300.93 JQ036232
Protoventuria major CBS 114594 JQ036233
Scolecobasidiella avellanea CBS 772.73 EF204505
Scolecobasidium cateniphorum P050 EU107309
Scolecobasidium terreum P043 EU107306
Scolecobasidium tricladiatum P051 EU107286
Scolecobasidium variabile P041 EU107310
Scolecobasidium verruculosum P044 EU107308
Spilocaea pomi CBS176.42 GU348998
Sympoventuria capensis CPC 12839 DQ885905
Sympoventuria capensis CBS 120136 DQ885906
Tyrannosorus pinicola CBS 124.88 DQ470974
Venturia alpina CBS 373.53 EU035446
Venturia asperata ATCC34052 EF114711
Venturia carpophila ICMP5402 AY849967
Venturia chlorospora Kruys 502 UPS DQ384101
Venturia inaequalis CBS 815.69 GU301878
Venturia inaequalis ATCC 60070 EF114712
Venturia populina CBS 256.38 GU323212
Venturia pyrina ATCC38995 EF114714
Veronaeopsis simplex CBS 588.66 EU041877
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CLAVATISPORA THAILANDICA GEN. ET SP. NOV. FROM THAILAND
FIGURE 1. RAxML phylogenetic tree based on the LSU sequence data. Bootstrap support values for maximum likelihood (≥ 50%
BS) and Bayesian posterior probabilities (≥ 0.95 PP) are shown in the tree (values below these thresholds not shown). The tree was
rooted with Acrospermum compressum (Acrospermales) as outgroup. The new taxon is highlighted in blue and types are in bold.
Taxo n o m y
Clavatispora S. Boonmee & K.D. Hyde, gen. nov. MycoBank: MB 805923
Etymology:—“clavatispora,” name recognizes the clavate ascospores.
Type species:—Clavatispora thailandica S. Boonmee & K.D. Hyde
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Saprobic on dead stems of herbaceous plants. Sexual state: Ascomata superficial, solitary, scattered on substrate,
developing on subiculum of brown hyphae, subglobose to globose, dark brown, covered with dark brown, thick-
walled, septate setae, and strands of radiating fused hyphae at base, with a barely raised, bright, central ostiole.
Peridium comprising 6–8 layers of dark brown cells of textura angularis, almost black at outside. Hamathecium of
ca 2 µm wide, anastomosing, septate, branched pseudoparaphyses, hyaline, embedded in gelatinous matrix. Asci
bitunicate, subclavate to broadly obovoid, thick-walled, with a short pedicel, apically rounded, lacking an ocular
chamber. Ascospores overlapping 2–3-seriate, clavate, asymmetrical, yellowish to reddish brown, becoming dark
brown at maturity, muriform, constricted at septa, tapering towards a subacute base, with basal cell pale brown or
brownish, smooth-walled. Asexual state: directly developing on MEA. Mycelium superficial, pale to moderately
dark brown, composed of septate, branched, thin- to thick-walled, smooth-walled, 1–3 µm wide hyphae.
Conidiophores erect, developing on hyphae, brown or light brown, septate, smooth, sometimes branched.
Conidiogenous cells holoblastic, subglobose. Conidia ellipsoidal to subglobose, pale brown to brown, guttulate,
rounded at apex, sub-acute at base, trans-septate, constricted and darkened at septa, smooth-walled.
Type species:—Clavatispora thailandica S. Boonmee & K.D. Hyde, sp. nov. MycoBank: MB 805924 (Figs 2–3)
Holotype:—THAILAND. Chiang Rai: Mae Fah Luang, Doi Tung, elev. ca. 1000 m., on dead stems of unidentified herbaceous
plants, 10 June 2009, Saranyaphat Boonmee DT04 (MFLU 100038!)—extype living culture = MFLUCC 100107 = BCC
38863 = IFRD 2170.
Sequence data:—LSU = KF770458, SSU = KF770457 and TEF1a = KF770459.
Etymology:—The specific epithet “thailandica” refers to the country from where the fungus was collected.
Saprobic on dead stems of herbaceous plants. Sexual state: Ascomata (115–)130–135 µm high ×
(120–)135–150(–160) µm diam ( = 130 × 140 µm, n = 5), superficial, solitary, scattered on substrate, developing
on subiculum of brown hyphae, subglobose to globose, dark brown, covered with dark brown, thick-walled, septate
setae, and strands of radiating fused hyphae at base, with a barely raised, bright central ostiole under light
microscope. Peridium 16–21 µm wide, comprising 6–8 layers of dark brown cells of textura angularis, almost
black at outside. Hamathecium of ca 2 µm wide, anastomosing, septate, branched pseudoparaphyses, hyaline,
embedded in gelatinous matrix. Asci 90–110 × 24–30 µm ( = 100 × 28 µm, n = 20), 8-spored, bitunicate,
fissitunicate, subclavate to broadly obovoid, thick-walled, with a short pedicel, apically rounded, lacking an ocular
chamber. Ascospores 32–44 × 10–13 µm ( = 37 × 11 µm, n = 20), overlapping 2–3-seriate, clavate, slightly
curved, asymmetrical, yellowish to reddish brown, becoming dark brown at maturity, 4–7(–8) transversely septate,
with 1–3 vertical septa in some cells, constricted at septa, tapering towards a subacute base, with basal cell pale
brown or brownish, smooth-walled. Asexual state: single spore isolate growing on MEA. Mycelium slightly raised,
pale to moderately dark brown, composed of septate, branched, thin- to thick-walled, smooth-walled, 1–3 µm wide
hyphae. Conidiophores erect, developing on hyphae, brown or light brown, septate, smooth, sometimes branched,
(5–)10–15 µm long. Conidiogenous cells holoblastic, subglobose. Conidia (14.5–)15–18(–21) × 5–7(–8) µm ( =
17 × 6 µm, n = 20), ellipsoidal to subglobose, pale brown to brown, guttulate, rounded at apex, sub-acute at base, 3-
septate, 1–2-septate when immature, constricted and darkened at septa, smooth-walled.
Cultural characteristics:—Ascospores germinating on WA within 12 h and germ tubes produced from spore
cells. Colonies on MEA slow growing, reaching 7 mm diameter in 1 week at 28°C, low convex, slightly effuse
hairy, edge entire, dark brown. Aerial mycelium, radiating outwards, superficial, septate.
Notes:—The dark brown ascomata, bitunicate asci and muriform brown ascospores of Clavatispora
thailandica are similar to characteristics of many genera of Pleosporales and some taxa in Capnodiales and
Hysteriales (Barr 1990, Boehm et al. 2009, Zhang et al. 2012, Hyde et al. 2013). Clavatispora can be distinguished
from other genera by its solitary ascomata, covered by superficial, dark setae, a bright ostiole and hyphomycetous
asexual state. The hyphomycetous conidia of the asexual state, derived from single ascospores, developed directly
on MEA within 60 days. The asexual conidia which formed chains, resemble species of Trichocladium, i.e. T.
constrictum (Goh & Hyde 1999), but C. thailandica differs in having branched conidiophores, light pigmented
conidia and cells with conspicuous guttules (Fig. 3E–L). Trichocladium constrictum has not been sequenced,
however, T. asperum (generic type) and T. opacum are not phylogenetically related to C. thailandica (data not
shown).
x
x
x
x
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CLAVATISPORA THAILANDICA GEN. ET SP. NOV. FROM THAILAND
FIGURE 2. Sexual state of Clavatispora thailandica (MFLU100038, holotype). A. Substrate and ascomata appearance. B. Squash
mount of ascoma. C. Section of ascoma. Note the fused hyphae forming hairs at the base. D. Setae. E. Peridium. F. Pseudoparaphyses.
G–H. Immature bitunicate asci with young developing ascospores. I–K. Mature asci. Fissitunicate dehiscence arrowed in H and I. L.
Characteristic dehiscence of asci after release of ascospores. M–P. Young and mature ascospores. Scale bars: A–C, G–L = 50 µm, D–E
= 20 µm, F = 5 µm, M–P = 10 µm.
Sequence data placed Clavatispora in the order Venturiales but it differs in morphology from most other
genera. The only genus in Venturiales with setose ascomata similar to Clavatispora is Trichodothis (Zhang et al.
2011). However, Trichodothis differs in having stromatic ascomata, oval asci and 2-celled, hyaline to pale brown,
ornamented ascospores. Molecular data are needed to confirm the placement of Trichodothis in Sympoventuriaceae
as it is rather atypical of the family. The presence of stromatic ascomata covered by spine-like setae, cylindrical
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asci, 2-celled and hyaline, smooth ascospores are also found in Gibbera conferta and Venturia compacta (Peck
1873, Petrak 1947). Furthermore, LSU data analysis paired G. conferta with an isolate of Fusicladium proteae in
the Venturiaceae clade (Fig. 1). Venturia compacta lacks DNA sequence data. Acantharia, Coleroa, Metacoleroa,
Protoventuria and Pyrenobotrys also have setose ascomata, but ascospores of these genera are 1-septate, hyaline or
coloured, with smooth or verrucose walls (Barr 1989, Carris & Poole 1993). Phylogenetic analysis based on LSU
sequence data shows that Clavatispora clusters with the hyphomycete genus Scolecobasidiella in
Sympoventuriaceae as well as Ochroconis, Scolecobasidium, Fusicladium and Veronaeopsis. The asexual state of
Clavatispora has 1–3-phragmoseptate, smooth conidia that are deeply constricted at septa, whereas in other genera
the conidia and conidiogenous apparatus are mostly verrucose to denticulate.
FIGURE 3. Asexual state of Clavatispora thailandica (MFLUCC100107, extype living culture). A. Germinating ascospore. B–C.
Colonies on MEA, upper surface and lower surface. D. Vegetative hyphae in culture. E–H. Conidiophores with conidia. I–L. Conidia.
Scale bars: A = 10 µm, B–C = 1 cm, D = 5 µm, E–L = 10 µm.
Discussion
Clavatispora thailandica, discovered on dead stems of an unidentified herbaceous plant in northern Thailand, is
characterized by its setiferous black ascomata, bitunicate asci with fissitunicate dehiscence, with a shrunken
ectotunica, endotunica and coloured plasmalemma layers, and clavate, dark brown, muriform ascospores. The
unique ascospore features suggest this species belongs in the Pleosporales. However, molecular studies place C.
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thailandica in the order Venturiales (Dothideomycetes), although its morphological characters are rather different
from all taxa in Venturiales (Zhang et al. 2012). Members of the order Pleosporales are distantly related (data not
shown). The phylogenetic analysis based on LSU sequence data show the placement of Clavatispora in family
Sympoventuriaceae, where it clusters with Scolecobasidiella avellanea (Fig. 1). Most taxa in Sympoventuriaceae
are hyphomyceteous, and, with the exception of Sympoventuria, the sexual state of most species is unknown (Crous
et al. 2007, Seifert et al. 2011, Samerpitak et al. 2014). Clavatispora also produced a hyphomyceteous state in
culture characterized by erect, coloured, septate, sometimes branched conidiophores and ellipsoidal to subglobose,
light coloured, guttulate, 3-septate, deeply constricted conidia (Fig. 3). The asexual state of Clavatispora, which is
Trichocladium-like, differs from all genera in the same clade i.e. Fusicladium, Ochroconis, Scolecobasidiella,
Scolecobasidium and Veronaeopsis (Abbott 1927, Ellis 1971, 1976, Fothergill 1996, Schubert et al. 2003, Seifert et
al. 2011, Martin-Sanchez et al. 2012, Zhang et al. 2012, Samerpitak et al. 2014). In the light of these unique
morpho-molecular features, the taxonomic affinity of Clavatispora thailandica as a new genus and species is
confirmed.
Acknowledgements
This work was supported by TRF/BIOTEC program Biodiversity Research and Training Grant BRT R_251181,
BRT R_253012 and partially by the Mushroom Research Foundation, Chiang Rai Province. The International
Fungal Research and Development Centre, The Research Institute of Resource Insects, the Chinese Academy of
Forestry (Bailongsi, Kunming 650224, China) and Mae Fah Luang University (grant for study Dothideomycetes
No. 56101020032) are also thanked for research facilities. Amy Rossman is thanked for her comments on the draft
manuscript and thanks are also extended to SMML-USDA for the use of laboratory facilities and for providing
Saranyaphat Boonmee with a scholarship to support her stay in USDA. We are grateful to Dr. Eric H.C. McKenzie
and Nalin N Wijayawardene for their helpful suggestions.
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