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PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN
1179-3163
(online edition)
Copyright © 2014 Magnolia Press
Phytotaxa 176 (1): 120–138
www.mapress.com
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phytotaxa
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Article
http://dx.doi.org/10.11646/phytotaxa.176.1.13
A re-assessment of Elsinoaceae (Myriangiales, Dothideomycetes)
RUVISHIKA S. JAYAWARDENA
1,3,4
, HIRAN A. ARIYAWANSA
2,3,4
, C. SINGTRIPOP
3,4
, YAN MEI LI
5
, JIYE
YA N
1
, XINGHONG LI
1*
, S. NILTHONG
4
& KEVIN D. HYDE
3,4
1
Institute of Plant and Environment Protection, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, People’s
Republic of China
2
The Engineering and Research Center for Southwest Bio-Pharmaceutical Resources of National Education Ministry of China,
Guizhou University, Guiyang 550025, Guizhou Province, People’s Republic of China
3
Institute of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
4
School of Science, Mae Fah Luang University, Chiang Rai. 57100, Thailand
5
International Fungal Research and Development Centre, Key Laboratory of Resource Insect Cultivation & Utilization State Forestry
Administration, The Research Institute of Resource Insects, Chinese Academy of Forestry, Kunming 650224, People’s Republic of
China
* Email: lixinghong1962@163.com
Abstract
The family Elsinoaceae is a relatively poorly known, but important family within Myriangiales, Dothideomycetes. The
genera of this family are mostly plant pathogens and causes disease, such as apple and grape scab. In this paper we revisit
the family by examining generic types and analysis of molecular sequence data available in GenBank. Elsinoaceae and
Myriangeaceae are morphologically and phylogenetically well-supported families in Myriangiales. In Elsinoaceae, 3 to
10 asci form in locules in light coloured pseudoascostromata, which form typical scab-like blemishes on leaf or fruit
surfaces, while Myriangeaceae forms raised, superficial, black ascostromata with single asci in each locule, genera may
or may not form scab-like lesions. Elsinoe is the type of the family Elsinoaceae and is characterized by forming scab-like
blemishes on leaves with few to numerous bitunicate, fissitunicate, globose asci forming inside each locule with a
pseudoascostroma containing fungal and host tissues. Following examination of generic type material, Molleriella is
retained in Elsinoaceae as it has characters similar to Elsinoe in forming scab-like lesions with pseudoascostromata
containing few to numerous bitunicate asci inside each locule. Beelia, Butleria, Hemimyriangium, Hyalotheles,
Micularia, Saccardinula, Stephanatheca and Xenodium are excluded from Elsinoaceae and their relative placement in
Dothideomycetes is discussed. Fresh collections of these genera are needed so that molecular sequence data can be
obtained and analysed to resolve their placement in families or orders of Dothidoemycetes.
Key words: Elsinoaceae, Elsinoe, Molleriella, morphology, phylogeny
Introduction
The family Elsinoaceae is a relatively poorly known, but important family of Dothideomycetes. Genera of this
family are mostly plant pathogens that cause scab and sunken spots on many economically important plants, such
as citrus, grapes, mango, peppers and legumes, reducing their fruit and vegetable values (Mchau et al. 1998, Condé
et al. 1997, Ellis & Erincik 2008, Hyun et al. 2001). Species also infect Protea species, reducing the value of the
flowers. Elsinoaceae is characterized by immersed to erumpent pseudoascostromata composed of pale gelatinous
thin-walled hyphae or pseudoparenchymatous cells with locules in a single layer or irregularly scattered. Asci are
bitunicate, fissitunicate, saccate to globose and with 3 to 10 arranged in locules. Ascospores are trans-septate or
sometimes muriform, and hyaline to brown (Kirk & Cannon 2008). The known asexual stages of Elsinoaceae are
acervular coelomycetes with polyphialidic conidiogenous cells (Sutton & Pollok 1973). Lumbsch & Huhndorf
(2010) listed Elsinoaceae as comprising ten genera viz Elsinoe, Beelia, Butleria, Hemimyriangium, Hyalotheles,
Micularia, Molleriella, Saccardinula, Stephanotheca and Xenodium. Li et al. (2011) excluded Beelia, Saccardinula
and Stephanotheca based on morphological characters.
Phytotaxa 176 (1) © 2014 Magnolia Press
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121
A RE-ASSESSMENT OF ELSINOACEAE
Molecular sequence data is presently only available for Elsinoe and its asexual state Sphaceloma. The first
molecular work on Elsinoaceae was that of Tan et al. (1996) who investigated the genetic differences among the
citrus scab pathogens Elsinoe fawcettii Bitanc. & Jenkins, E. australis Bitanc. & Jenkins from South America and
Sphaceloma fawcetii var. scabiosa (McAlpine & Tyron) Jenkins from Australia. Swart et al. (2001), Hyun et al.
(2001), Schoch et al. (2006, 2009), Boehm et al. (2009), Kerry et al. (2011) and Hyde et al. (2013) have also
carried out higher level molecular studies on Dothideomycetes, which included strains of genus Elsinoe.
Historic overview of Elsinoaceae
The family Elsinoaceae was introduced by Saccardo & Trotter (1913). Many classical treatments (Boedijn
1961) have placed Elsinoaceae in synonymy with Myriangiaceae. Höhnel (1909) however, was convinced that the
former constituted a separate family because of the difference in the habit and also based on developmental studies.
von Arx & Müller (1975) were not of the same opinion based on morphological characters, and placed Elsinoe
with another 15 genera in Myriangiaceae (Table 1). Barr (1979) and Eriksson (1981) were of the opinion that two
separate families should be maintained for Elsinoaceae and Myriangiaceae with the latter predominantly found on
branches and former restricted to foliar pathogens. Molleriella and Micularia were previously placed in
Saccardiaceae, also a family of Myriangiales. This family was characterized by superficial stroma and asci arising
at one level and Saccardiaceae are no longer considered as a family in order Myriangiales.
Elsinoaceae has also been referred as Plectodiscellaceae a family established by Woronichin (1914) based on
Plectodiscella piri. Jenkins (1932) treated Plectodiscella as a synonym of Elsinoe and also confirmed that the
conidial states of Elsinoe belong to Sphaceloma and not to other asexual genera (e.g. Cladosporium,
Gloeosporium) to which they had been assigned (Frederick et al. 1947). Elsinoaceae was placed in the order
Myriangiales by Frederick et al. (1947) and this was followed by Lumbsch & Huhndorf (2007, 2010). Recent
studies based on combine gene analysis of LSU, SSU, RPB1 and RPB2 concluded that Elsinoaceae and
Myriangeaceae forms two distinct sub-clades with high bootstrap support in the order Myriangiales (Schoch et al.
2006, 2009, Boehm et al. 2009, Hyde et al. 2013).
We have been studying the genera of Dothideomycetes in order to provide a natural classification of this large
class (Boonmee et al. 2011, Liu et al. 2012, Wu et al. 2011, Zhang et al. 2012, Hyde et al. 2013). Some studies
have been based exclusively on morphological characterization and some have integrated molecular analysis
(Ariyawansa et al. 2013, Chomnunti et al. 2011, 2012, Liu et al. 2012, Wu et al. 2010, Zhang et al. 2012). The
present study revisits the family Elsinoaceae by examining the types of all genera included by Lumbsch &
Huhndorf (2010). We loaned generic types and re-describe and illustrate the species and suggest placements and
requirements for future work. We also provide a new phylogenetic tree for Myriangiales based on available
sequence data.
TABLE 1. Placement of genera in Elsinoaceae by different authors
.
Müller & von Arx (1975) Barr (1987) Kirk et al. (2001) Lumbsch & Huhndorf (2007, 2010) This paper (2014)
Hyalotheles
Micularia
Xenodium
Molleriella
Saccardinula
Beelia
Elsinoe
Stephanotheca
Diplotheca
Myriangium
Butleria
Anhelia
Cookella
Pycnoderma
Uleomyces
Anhelia
Butleria
Diplotheca
Elsinoe
Elsinoe
Beelia
Butleria
Hemimyriangium
Hyalotheles
Micularia
Molleriella
Saccardinula
Stephanotheca
Xenodium
Beelia
Butleria
Elsinoe
Hemimyriangium
Hyalotheles
Micularia
Molleriella
Saccardinula
Stephanotheca
Xenodium
Elsinoe
Molleriella
JAYAWARDENA ET AL.
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Phytotaxa 176 (1) © 2014 Magnolia Press
Materials and methods
Type specimens were loaned from BISH, L, LPS, PAD and S (abbreviations follow Index Herbarium, 2013).
Ascomata were rehydrated in water and/or 5% KOH prior to examination and sectioning following the methods
described by Chomnunti et al. (2011). Hand sections were mounted in water for microscopic studies and
photomicrography. Figures of Sphaceloma ampelinum de Bary were redrawn from Sutton & Pollok (1973) and
Molleriella mirabilis winter was redrawn from Höhnel (1909) using transparent drawing papers and drawing pens. The
details of the fungi were captured in a Nikon ECLIPSE 80i compound microscope with a Canon 450D digital camera
accessory. Measurements were made with the Tarosoft (R) Image Frame Work program and images used for figures
were processed with Adobe Photoshop CS3 Extended version 10.0 software (Adobe Systems, The United States).
Sequence alignment and phylogenetic analysis
The internal transcribe spacer (ITS), large and small subunits of the nuclear ribosomal RNA genes (LSU, SSU)
and two protein coding genes, second largest subunit of RNA polymerase II (RPB2) and translation elongation
factor-1 alpha (TEF1) were included in the analysis. All sequences were obtained from GenBank and are listed in
Table 2. Sequences were aligned using Bioedit version 7.0.9.0 (Hall 1999) and ClustalX v. 1.83 (Thompson et al.
1997). The alignments were checked visually and improved manually where necessary. Phylogenetic analyses were
carried by using PAUP v. 4.0b10 (Swofford 2002) for Maximum-parsimony (MP).
Maximum-parsimony analysis was performed to gain the most parsimonious tree. Trees were inferred using the
heuristic search option with 1000 random sequence additions. Maxtrees were setup to 5000 and branches of zero
length were collapsed and all multiple parsimonious trees were saved. Descriptive tree statistics for parsimony (Tree
Length [TL], Consistency Index [CI], Retention Index [RI], Relative Consistency Index [RC] and Homoplasy Index
[HI] )were calculated for trees generated under different optimality criteria. Kishino-Hasegawa tests (KHT) (Kishino
and Hasegawa 1989) were performed in order to determine whether trees were significantly different. Maximum
parsimony bootstrap values (MPBP) equal or greater than 50% are given above each node (Fig 1).
TABLE 2. GenBank accession numbers.
Species Voucher/Culture SSU LSU RPB2 TEF1 ITS
Aliquandostipite khaoyaiensis CBS 118232 AF201453 GU301796 FJ238360 GU349048 -
Amniculicola immersa CBS 123083 GU456295 FJ795498 GU456358 GU456273 -
Amniculicola parva CBS 123092 GU296134 FJ795497 - GU349065 -
Apiosporina collinsii CBS 118973 GU296135 GU301798 - GU349057 -
Bimuria novae-zelandiae CBS 107.79 AY016338 AY016356 DQ470917 DQ471087
Botryosphaeria dothidea CBS 115476 DQ677998 DQ678051 DQ677944 - DQ767637
Capnodium coffeae CBS 147.52 DQ247808 DQ247800 DQ247788 DQ471089 AJ244239
Capnodium salicinum CBS 131.34 DQ677997 DQ678050 - DQ677889 AJ244240
Cochliobolus heterostrophus CBS 134.39 AY544727 AY544645 DQ247790 DQ497603 -
Didymella exigua CBS 183.55 EU754056 EU754155 GU371764 - GU237794
Dothidea sambuci DAOM 231303 AY544722 AY544681 DQ522854 DQ497606 AY883094
Dothiora cannabinae CBS 737.71 DQ479933 DQ470984 DQ470936 DQ471107 AJ244243
Elsinoe brasiliensis CPC 18528 JN940567 JN940394 - - JN943501
Elsinoe centrolobi AFTOL-ID 1854 DQ678041 DQ678094 - DQ677934 -
Elsinoe fawcettii CPC 18570 JN940565 JN940385 - - JN943503
Elsinoe fawcettii CPC 18535 JN940559 JN940382 - - JN943496
Elsinoe mimosae CPC 18518 JN940564 JN940387 - - JN943505
Elsinoe phaseoli CBS 165.31 DQ678042 DQ678095 - DQ677935 -
Elsinoe veneta CBS 150.27 DQ767651 DQ767658 - DQ767641 -
Elsinoe verbenae CPC 18561 JN940562 JN940391 - - JN943499
Gloniopsis praelonga CBS 112415 FJ161134 FJ161173 FJ161113 FJ161090 -
Glonium circumserpens CBS 123343 FJ161160 FJ161200 FJ161126 - FJ161108
Guignardia bidwellii CBS 237.48 DQ678034 DQ678085 DQ677983 -
-
...... continued on the next page
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A RE-ASSESSMENT OF ELSINOACEAE
Results
Phylogenetic analysis
Phylogeny based on combined ITS, SSU, LSU, RPB2 and TEF1 gene datasets
The combined ITS, SSU, LSU, RPB2 and TEF1 data set utilized 60 taxa with Schismatomma decolorans as the
out group taxon. The maximum parsimony dataset consists of 5,846 total characters of which 2,974 characters were
constant, 1,300 variable characters were parsimony-uninformative and 1,572 characters were parsimony-
informative. Kishino-Hasegawa (KH) test showed length= 8935 steps, CI=0.487, RI=0.573, RC= 0.279 and
HI=0.513. Eleven MP trees were generated and the first of the most parsimonious tree was selected, (Fig. 1).
Phylogenetic trees obtained from maximum parsimony analyses yielded trees with similar overall topology at
subclass and family relationship in agreement with previous work based on maximum parsimony (Schoch et al.
2006, 2009, Boehm et al. 2009).
TABLE 2 (continued)
Species Voucher/Culture SSU LSU RPB2 TEF1 ITS
Hysterobrevium mori CBS 123336 FJ161164 FJ161204 - -
-
Jahnula aquatica R68-1 EF175633 EF175655 - -
-
Jahnula bipileata F49-1 EF175635 EF175657 - -
-
Kalmusia scabrispora MAFF 239517 AB524452 AB524593 AB539093 AB539106 -
Leptosphaerulina australis CBS 317.83 GU296160 GU301830 GU371790 GU349070 GU237829
Lophium mytilinum CBS 269.34 DQ678030 DQ678081 DQ677979 - DQ677926
Macrovalsaria megalospora CBS 178150 FJ215707 FJ215701 - - -
Montagnula opulenta CBS 168.34 AF164370 DQ678086 - DQ677984 -
Mycosphaerella punctiformis CBS 113265 DQ471017 DQ470968 DQ470920 DQ471092 EU167569
Myriangium duriaei CBS 260.36 AY016347 DQ678059 DQ677954 DQ677900 -
Myriangium hispanicum CBS 247.33 GU296180 GU301854 GU371744 GU349055 -
Mytilinidion mytilinellum CBS 303.34 FJ161144 FJ161184 FJ161119 - FJ161100
Phaeocryptopus gaeumannii CBS 267.37 EF114722 EF114698 - - EU700365
Phaeotrichum benjaminii CBS 541.72 AY016348 AY004340 DQ677946 DQ677892
Phoma exigua CBS 431.74 EU754084 EU754183 GU371780 GU349080 FJ427001
Pleospora herbarum CBS 191.86 DQ247812 DQ247804 DQ247794 DQ471090 EF452449
Pyrenophora phaeocomes DAOM 222769 DQ499595 DQ499596 DQ497614 DQ497607 JN943649
Rhytidhysterium rufulum CBS 306.38 GU296191 FJ469672 - GU349031 -
Schismatomma decolorans DUKE 0047570 AY548809 AY548815 - DQ883715 DQ883725
Sphaceloma arachidis CPC 18533 JN940548 JN940372 - - JN943485
Sphaceloma arachidis CPC 18529 JN940547 JN940374 - - JN943484
Sphaceloma asclepiadis CPC 18544 JN940558 JN940383 - - JN943495
Sphaceloma asclepiadis CPC 18532 JN940556 JN940380 - - JN943493
Sphaceloma asclepiadis CPC 18583 JN940557 JN940381 - - JN943494
Sphaceloma bidentis CPC 18586 JN940555 JN940379 - - JN943492
Sphaceloma erythrinae CPC 18530 JN940566 JN940392 - - JN943502
Sphaceloma erythrinae CPC 18540 JN940549 JN940388 - - JN943486
Sphaceloma krugii CPC 18531 JN940551 JN940375 - - JN943489
Sphaceloma krugii CPC 18537 JN940553 JN940376 - - -
Sphaceloma krugii CPC 18554 JN940552 JN940377 - - -
Sphaceloma sesseae CPC 18549 JN940561 JN940393 - - JN943498
Sphaceloma terminaliae CPC 18538 JN940560 JN940371 - - JN943497
Stylodothis puccinioides CBS 193.58 - AY004342 - DQ677886 -
Trichodelitschia bisporula CBS 262.69 GU349000 GU348996 GU371802 GU349020 -
Trichodelitschia munkii Kruys201 DQ384070 DQ384096 - - -
Venturia inaequalis CBS 594.70 GU296205 GU301879 - GU349022 -
Venturia populina CBS 256.38 GU296206 GU323212 - - EU035467
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Phytotaxa 176 (1) © 2014 Magnolia Press
Phylogenic analysis
The combined ITS, SSU, LSU, RPB2 and TEF1 gene dataset of 15 families in the class Dothideomycetes is
shown in Fig. 1. The analysis confirms that the families Elsinoaceae and Myriangeaceae appear to be well-resolved
with high bootstrap support (82%). However, in our analysis we used eight sexual strains of Elsinoe species, seven
asexual strains identified as Sphaceloma species and two Myriangium species. Once more data becomes available
for other genera in these families and trees are better populated we cannot predict if these families will continue to
receive strong support. Problematic species were Elsinoe veneta (Burkh.) Jenkins and E. brasiliensis Bitanc. &
Jenkins which clustered separately from the main Elsinoaceae clade in the phylogenetic tree (Fig. 1).
FIGURE 1. Phylogram generated from maximum parsimony analysis based on combined multi-gene sequences (ITS, SSU, LSU,
RPB2 and TEF1), showing the phylogenetic relationships of the family Elsinoaceae. The values noted are parsimony bootstraps
(>50%). The tree is rooted with Schismatomma decolorans.
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A RE-ASSESSMENT OF ELSINOACEAE
Taxo n o m y
Myriangiales
The order Myriangiales was introduced by Starbäck (1899) for the species characterized by having crustose
ascostromata and muriform ascospores, based on the type species, Myrangium duriaei Mont. & Berk. (Miller 1938,
Hyde et al. 2013). Multigene phylogenetic studies revealed that the order Myriangiales always clusters in
Dothideomycetes (Boehm et al. 2009, Schoch et al. 2009, Zhang et al. 2012, Hyde et al. 2013). This order is
characterized by pulvinate, irregular ascostromata in which the asci are irregularly arranged in one or more layers
in locules. Locules may contain single or multiple asci within each locule. Asci have a minute pedicel and
indistinct ocular chambers. Ascospores are irregularly arranged and are liberated only by the breakup of the
stromatal layers above them. Asexual states are coelomycetous. Kirk et al. (2008) included three families under
order Myriangiales, Cookellaceae, Elsinoaceae and Myriangeaceae. Lumbsch & Huhndorf (2010) accepted only
Elsinoaceae and Myriangeaceae in Myriangiales based on phylogenetic results. Elsinoaceae is morphologically
different from Myriangeaceae as 3 to 10 asci form in locules in light coloured pseudoascostromata, which form
typical scab-like blemishes on leaf or fruit surfaces. No molecular data is available for Cookellaceae and its
placement in Myriangiales cannot be confirmed.
Elsinoaceae Höhn. ex Sacc. & Trotter [as 'Elsinoёaceae'], Sylloge Fungorum (Abellini) 22: 584 (1913)
Synony ms:
Myxomyrangiaceae (Theiss.) Theiss.
Plectodiscellaceae Woron., Mykol. Zentbl. 4: 232 (1914)
Saccardinulaceae G. Arnaud, Annls Sci. Nat., Bot., sér. 10 7: 647 (1925)
Parasitic or saprotrophic on plant leaves and fruits causing scab and sunken scab-like blemishes. Sexual state:
Pseudoascostromata usually spread around host veins, solitary, aggregated, or gregarious, wart-like or scab-like
blemishes, pulvinate, superficial, globose to subglobose, white, pale yellow to brown, multi-loculate, locules
scattered in upper part of pseudoascostromata. Cells of pseudoascostromata comprising host cells and inter-
dispersed light coloured fungal hyphae, opening by unordered break down of the surface layer. Locules with 3–10
asci inside each locule, ostiolate. Ostioles minute. Pseudoparaphyses absent. Asci 8-spored, bitunicate,
fissitunicate, saccate to globose, with a minute pedicel, and indistinct ocular chamber. Ascospores irregularly
arranged, oblong or fusiform with slightly acute ends, with 2–3 transverse septa, hyaline, smooth-walled, lacking a
sheath. Asexual state: coelomycetous “Sphaceloma”. Lesions circular, dark brown raised margin, cream-brown.
Acervuli subepidermal, pseudoparenchymatous. Conidiophores hyaline to pale-brown, polyphialidic.
Conidiogenous cells formed directly from the upper cells of the pseudoparenchyma, monophialidic to polyphidalic,
integrated or discrete, determinate, hyaline to pale brown, lacking a thickened region around the phialide channel.
Conidia hyaline, unicellular, ellipsoidal, aseptate, biguttulate.
Lumbsch & Huhndorf (2010) included ten genera in Elsinoaceae. Li et al. (2011), Hyde et al. (2013) and
Dissanayake et al. (2014) however, revised the familial positions of Beelia, Butleria, Hyalotheles,
Hemimyriangium, Saccardinula, Stephanotheca and Xenodium and supported the separation of Elsinoaceae from
the family Myriangiaceae based on phylogenetic analysis. The families are also morphologically distinct. In
Elsinoaceae several asci form in locules that develop in pseudoascostromata, that comprises host cells and
interdispersed light coloured fungal hyphae. In Myriangiaceae, asci develop singly in locules that are generally
scatted in ascostromata. The pseudoascostromata of Elsinoaceae are generally superficial, pulvinate and comprise
of light coloured fungal hyphae and darkened host cells. Saccardinula shows similar characters to the family
Brefeldiellaceae where it is placed (Hyde et al. 2013). Stephanotheca shows characteristic of the family
Asterinaceae where it is retained, whereas Xenodium has unitunicate asci and thus excluded from
Dothideomycetes. Hyalotheles has been placed in Dothideomycetes genera incertae sedis. Butleria and
Hemimyrangium are placed in the family Myriangeaceae by Dissanayake et al. (2014). Fresh collections are
needed for epitypification and obtaining sequence data.
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Typ e : — Elsinoe Racib., Parasitische. Algen und Pilze Java’s (Jakarta) 1: 14 (1900)
Possible synonyms
Sphaceloma de Bary, Ann. Oenol. 4: 165 (1874)
Bitancourtia Thirum & Jenkins, Mycologia 45(5): 781 (1953)
Isotexis Syd., in Sydow & Petrak, Annls mycol. 29 (3/4): 261 (1931)
Plectodiscella Woron., Mykol Zentbl 4: 232 (1914)
Uleomycina Petr., Sydowia 8(1–6): 74 (1954)
Kurosawaia Hara, List of Japanese Fungi: 172. Ed. 4 (1954)
Manginia Viala & Pacotter, C.r. hebd, Séanc. Acad. Sci., Paris 139: 88 (1904)
Melanobasidium Maubl., Bull. Soc. Mycol. Fr. 22: 69 (1906)
Melonobasis Clem. & Shear, Gen. fung., Edn 2 (Minneapolis): 224, 403 (1931)
Melanodochium Syd., Annls mycol. 36 (4): 310 (1938)
Melanophora Arx, Verh. K. ned, Akad, wet., tweede sect. 51(3): 43 (1957)
Parasitic on plant leaves and fruits causing scab and sunken scab-like blemishes. Sexual state:
Pseudoascostromata usually spread around host veins, solitary, aggregated, or gregarious, wart-like or scab-like
blemishes, pulvinate, superficial, globose to subglobose, white, pale yellow to brown, multi-loculate, locules
scattered in upper part of pseudoascostromata. Cells of pseudoascostromata comprising host cells and inter-
dispersed light coloured fungal hyphae opening by unordered break down of the surface layer. Locules with
numerous 3–8 asci inside each locule, ostiolate. Ostiole minute. Pseudoparaphyses absent. Asci 8-spored,
bitunicate, fissitunicate, saccate to globose, apedicellate, with indistinct ocular chamber. Ascospores irregularly
arranged, oblong or fusiform with slightly acute ends, with 2–3 transverse septa, hyaline, smooth-walled, lacking a
sheath. Asexual state: Ceolomycetous “Sphaceloma” Acervuli subepidermal, pseudoparenchymatous.
Conidiophores hyaline to pale-brown, polyphialidic. Conidiogenous cells formed directly from the upper cells of
the pseudoparenchyma, monophialidic to polyphialidic, lacking a thickened region around the phialidic channel,
terminal, integrated, determinate, hyaline to pale brown. Conidia hyaline, unicellular, ellipsoidal, aseptate,
biguttulate.
Type species:—Elsinoe canavaliae Racib. [as 'canavalliae'], Parasitische. Algen und Pilze Java’s (Jakarta) 1: 14
(1900)
≡ Uleomyces canavaliae (Racib.) G. Arnaud, Annales des Sciences Naturelles Botanique 10 5: 685 (1925) MycoBank: 217658
(Figs 2, 3)
Parasitic on leaves, forming scab on lower leaf surface. Sexual state: Pseudoascostromata 1–5 × 5–8 mm in diam.
( = 3.2 × 6.5 mm, n=10), spreading around the host veins, solitary, aggregated, or gregarious, wart-like or scab-
like blemishes, pulvinate, superficial, globose to subglobose, white, pale yellow or occassionally brown, in section
with numerous locules distributed inside the upper part of pseudoascostromata, with numerous asci within each
locule. Cells of pseudoascostromata comprising host cells and inter-dispersed light coloured fungal hyphae.
Locules with 3–8 asci inside each locule, ostiolate. Ostiole minute. Pseudoparaphyses absent. Asci 16–22 ×16–21
μm ( = 19.7 × 18.9 μm, n=20), 8-spored, bitunicate, fissitunicate, saccate to globose, apedicellate, with indistinct
ocular chamber. Ascospores 10–14 × 3–5 μm ( = 12.3 × 4 μm, n=40) irregularly arranged, oblong or fusiform with
slightly acute ends, with 2–3 transverse septa, hyaline, smooth-walled, lacking a sheath. Asexual state:
“Sphaceloma”, Acervuli sub-epidermal, pseudoparenchymatous. Conidiophores hyaline to pale-brown,
polyphialidic. Conidiogenous cells formed directly from the upper cells of the pseudoparenchyma, monophialidic
to polyphidalic, integrated or discrete, determinate, hyaline to pale brown, lacking a thickened region around the
phialide channel. Conidia 8–16 × 3–5 μm ( = 14.1 × 4 μm, n = 20), hyaline, unicellular, ellipsoidal, aseptate,
biguttulate.
Material examined:—Philippines. Laguna Province: Mount Maquiling, near Los Baños, on Canavalia
ensiformis (Fabaceae), Baker, August 1913 (S, F66900!, isotype).
Elsinoe was established by Raciborski (1900) including three species (E. canavaliae, E. antidesmae Racib., E.
meninspermacearum Racib.). von Arx & Müller (1975) placed Elsinoe in Myriangiaceae based on its immersed or
erumpent, pulvinate or irregular ascomata and being parasitic on higher plants causing scab. Later, the genus was
placed in the family Elsinoaceae (Barr 1979, Kirk et al. 2001, Lumbsch & Huhndorf 2007, 2010). There are 139
x
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species epithets for Elsinoe (Index Fungorum 2013) and they are generally parasites on leaves, stems, and fruits.
The asexual state of Elsinoe is “Sphaceloma” (Wijayawardena et al. 2012). Bitancourt & Jenkins (1946) described
Sphaceloma manihoticola Bitanc. & Jenkins on Manihot esculenta Crantz (Zeigler & Lozano, 1983). The
relationship between Sphaceloma and Elsinoe has been resolved by analyzing rDNA sequence data
(Cheewangkoon et al. 2010). There are 168 species epithets for Sphaceloma in Index Fungorum (2013). Most of
the Sphaceloma species are pathogens that affect flowers, fruits, leaves and stems, causing characteristic scab
lesions on the organs that they attack as well as necrotic spots on leaves.
FIGURE 2. Elsinoe canavaliae (isotype). a. Herbarium material. b. Pseudoascostromata on host substrate. c–d. Section of
pseudoascostroma. e. Asci stained with cotton blue reagent in a section of pseudoascostromata. f–g. Ascus eight irregularly arranged
ascospores. h. Fissitunicate dehicnese of the ascus. i. Smooth, hyaline ascospores. j. Ascospore stained with cotton blue . Scale bars:
c–d = 100 μm, e = 50 μm, f–h = 20 μm, i–j = 10 μm.
Elsinoe is an important plant pathogenic genus causing scab and anthracnose. This genus is widely known in
association with various Citrus species. But it also causes diseases in Malus, Rubus, Vitis species and several other
plants and effects plant families such as Moraceae, Piperaceae, Sapindaceae, Anacardiaceae, Myrtaceae and
Vitaceae. Elsinoe fawcettii Bitanc. & Jenkins and E. australis Bitanc. & Jenkins cause scab disease of Citrus sp.
(Hanlin 1989, Timmer et al. 1996, Hyun et al. 2001, Nelson 2008) and there are many other important pathogens of
Elsinoe and its asexual states (Table 3). Elsinoe takoropuku G.S. Ridl & Ramsfield is a recently introduced species
(Ridley & Ramsfield 2006), but differs from E. canavaliae markedly as it forms ascostromata on twigs of
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Pittosporum tenuifolium Gaertn instead of scabs on leaves. It contains locules each containing single asci. The asci
are however, thought to be more similar to Elsinoe type even though it has many characters similar with
Myriangiaceae.
FIGURE 3a. Sphaceloma sp. (THAILAND, Chiang Rai, near Phu Chi Fa, on Citrus sp., 28 January 2013, MFLU 12-2214). a–b.
Conidiomata on host substrate. c–d. Section of conidiomata. e–g. Phialidic conidiogenous cells. h–i. Conidia. Scale bars: e–i = 50 μm,
c–d = 20 μm, k = 10 μm.
FIGURE 3b. Sphaceloma ampelinum material redrawn from Sutton & Pollok (1973) a. Conidia. b. Phialidic conidiogenous cells. c.
V.S. of confluent acervuli. Scale bars: a–b = 10 μm, c = 100 μm.
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TABLE 3. Some crops affected by Elsinoe species
.
Species named as Sphaceloma shoul d be synonymized under Elsinoe if their relationships are confirmed by molecular data
Schoch et al. (2006, 2009) and Boehm et al. (2009) showed that there is an obvious subclade among the
species of Myriangiaceae (named Elsinoaceae). However, only four Elsinoe strains and one Myriangium strain
were used in Schoch et al. (2006) and no Elsinoe strains were used in Schoch et al. (2009). Thus, molecular data
does convincingly resolve the two families. In the study on the taxonomy of the species associated with scab
disease of Proteaceae, Swart et al. (2001) analyzed ITS sequence of six Elsinoe species, E. banksiae Pascoe &
Crous, E. leucospermi L. Swart & Crous, E. proteae Crous & L. Swart, Elsinoe sp. (from Citrus), Elsinoe sp. (from
Banksia), Sphaceloma protearum L. Swart & Crous; the molecular analysis supported five species and four were
described in their paper. Hyun et al. (2001) have done molecular analysis of several Elsinoe isolates causing scab
disease of Citrus sp. in Jeju Island in Korea. Kerry et al. (2011) used molecular identification for Sphaceloma
perseae and its absence. Phylogenetic analysis in for this paper shows that E. veneta and E. brasiliensis clustered
separately from the Elsinoaceae clade. The basionym of Elsinoe veneta is Plectodiscella veneta Burkh. Jenkins
(1932) however, considered Plectodiscella veneta to be a species of Elsinoe and transferred it to E. veneta. In
Elsinoe veneta and E. brasiliensis the pseudoascostromata are multi-loculate, pulvinate, scab-like structures, asci
are globose, bitunicate and fissitunicate and ascospores hyaline with three septa which resembles Elsinoaceae. In
Elsinoe veneta and E. brasiliensis there is one ascus per locule (Saccardo 1925–1928). In the type genus Elsinoe,
multiple asci can be found in each locule (Hyde et al. 2013). Further studies are needed to clarify their taxonomic
placement.
Several studies have been conducted on Elsinochrome which are non-host selective, light-activated,
polyketide-derived toxins produced by Elsinoe species (Chung & Liao 2008). Further molecular studies are needed
on Elsinoe species and also on its asexual state to resolve the correct placement of this taxon. Phylogenetic analysis
shows that Sphaceloma species cluster with Elsinoe species (Fig. 1) and this is also supported by morphological
data (Cheewangkoon et al. 2010). Based on the available molecular and morphological data it may be necessary to
synonymize Sphaceloma under Elsinoe. However, the type species of Elsinoe nor Sphaceloma have been
sequenced and therefore they should not be synonymized at this time. Therefore we list Sphaceloma as a possible
synonym only.
Molleriella G. Winter, Boletim da Sociedade Broteriana, Coimbra, sér 1, 4: 199 (1886)
Possible synonyms:
Agrona Hӧhn., Sber, Akad, Wien, Math-naturw, Kl., Abt. 1, 118: 362 [88 repr.] (1909)
Capnodiopsis Henn., Hedwigia 41: 298 (1902)
Elachophyma Petr., in Sydow & Petrak, Annls. Mycol. 29 (3/4): 258 (1931)
Elenkinella Woron., Bot. Mater. Insr. Sporov. Rast, Glavn, Bot. Sada RSFSR 1: 33 (1922)
Nostocotheca Starbäck, Bih, K. svenska VetenskAkad. Handl., Afd. 3 25 (no. 1): 20 (1899)
Saprotrophic on plant leaves causing scab-like lesions on lower leaf surface. Sexual state: Pseudoascostromata
solitary, aggregated or gregarious, pulvinate, superficial, globose to oval, pale yellow, multi-loculate, locules
Species Crop Reference
Elsinoe sp. Proteaceae sp. Swart 2001
Sphaceloma*symphoricarpi Barrus & Horsfall Snowberry Kudela & Krejza 2006
Elsinoe mangiferae Bitanc. & Jenkins Mango Conde et al. 1997
Sphaceloma rosarum (Pass.) Jenkins Rose Jenkins 1947
Elsinoe phaseoli Jenkins Lima bean Jenkins 1947
Sphaceloma manihoticola Bitanc. & Jenkins Cassava Zeigler & Lozano 1983, Reeder et al. 2009
Sphaceloma perseae Jenkins Avocado Kerry et al. 2011
Elsinoe solidaginis Jenkins & Ukkelberg Goldenrod Jenkins & Ukkelberg 1935
Elsinoe ampelina Shear Grape Jenkins 1942, Ellis & Enrick 2008
Sphaceloma sacchari T.C. Lo Sugarcane Lo 1964
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scattered in upper part of pseudoascostromata. Cells of pseudoascostromata comprising host cells and inter-
dispersed light coloured fungal hyphae opening by unordered break down of the surface layer. Locules with 4–10
asci inside each locule, ostiolate. Ostiole minute, Paraphyses absent. Asci 8-spored, bitunicate, fissitunicate,
globose to subglobose, thick-walled, with a minute pedicel and an ocular chamber. Ascospores irregularly
arranged, oblong to sub-clavate, 6–8-septate, hyaline. Asexual state: Unknown.
Typ e : —Molleriella mirabilis G. Winter, Boletim da Sociedade. broteriana, Coimbra, sér 1, 4: 199 (1886)
MycoBank No: 528389 (Fig 4)
FIGURE 4. Molleriella mirabilis (type) material redrawn from Höhnel (1909). a. Herbarium material. b. Pseudoascostromata on host
substrate. c. Section through pseudoascostroma. d. Asci. e–f. Immature asci. g. Mature ascus. h. Immature ascus stained in cotton blue
reagent. i–k. Irregularly arranged hyaline 6–8-septate ascospores. i. Ascospore stained in cotton blue. Scale bars: c–d = 100 μm, e–l =
10 μm.
Saprotrophic on plant leaves. Sexual state: Pseudoascostromata 5–8 × 2–3 mm ( = 7.5 × 3.1 mm, n = 5) spread on
lower surface, solitary, aggregated, or gregarious, pulvinate, superficial, pale yellow, multi-loculate, locules
scattered in the upper part of the pseudoascostromata. Cells of pseudoascostromata comprising host cells and inter-
dispersed light coloured fungal hyphae. Locules with 4–10 asci inside each locule, ostiolate. Ostiole minute.
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Paraphyses not observed. Asci 17–37 × 9–21 μm ( = 31.4 × 16.1 μm, n = 10) 8-spored, bitunicate, fissitunicate,
globose to sub globose, solitary or aggregated, with minute pedicel and an ocular chamber. Ascospores 18–24 ×
6–8 μm ( = 22.7 × 7.3 μm, n = 10) irregularly arranged, smooth, oblong to sub-clavate, often curved, 6–8-septate,
muriform, hyaline, no sheath. Asexual state: Unknown.
Material examined:—AFRICA. S. Thomé Insel, pr. Bate-pá, on Convolvulaceae, A. Moller, June 1885 (S,
F51162!, type)
The genus Molleriella was introduced by G. Winter (1886) in the class Discomycetes and later placed in the
family Phymatophaeriaceae by Engler et al. (1887, 1897) and Hieronymus and Hennings (1901). Boedijn (1961)
placed Molleriella in the family Saccardiaceae, while Arnaud (1918) and Bessy (1950) had placed Molleriella in
Myriangiaceae. On the other hand, Kirk et al. (2001) placed Molleriella in the family Elsinoaceae and Lumbsch &
Huhndorf (2007, 2010) followed this. At present there are four species epithets listed in Index Fungorum (2013).
This genus differs from the type Elsinoe by being saprotrophic and differs in having globose to subglobose sessile
asci. This genus is similar to family type Elsinoe in having multi-loculate, pulvinate scab-like blemish
pseudoascostromata, without paraphyses, and minute ostioles, plus globose asci with indistinct ocular chambers.
This genus is retained in the family Elsinoaceae but, fresh collections and molecular analysis are needed to
establish natural placement of this genus.
Genera excluded from Elsinoaceae
Asterinaceae
The family Asterinaceae are small obligately biotrophic ascomycetes that are associated with living leaves of a
broad range of angiosperms from tropical and subtropical regions (Hyde et al. 2013). The important features of
Asterinaceae are the superficial, black, web-like colonies that form on the upper and lower surface of leaves,
thyriothecia that are closely attached to the host plant cuticle and open at maturity with central star-shaped fissures.
The globose bitunicate asci develop vertically in the ascomatal cavity, from the base to the dehiscent opening; this
is the important character of the family Asterinaceae which is different from the family Microthyriaceae, where
asci are embedded in mucilage and grow horizontally to the host surface and inclined from the thyriothecia rim
towards the ostiole. Ascospores are mostly ellipsoidal, 2-celled and hyaline when young and becoming brown at
maturity (Hyde et al. 2013).
Stephanotheca Syd. & P. Syd., The Phillippine Journal of Science: C Botany 9: 178 (1914)
Foliar epiphytes on plants. Sexual state: Ascomata superficial, globose to subglobose, base flattened, black. Upper
wall of thick-walled cells, forming a textura porrecta. Hamathecium lacking pseudoparaphyses. Asci 16–31 ×
16–25 µm ( = 25 × 19 µm, n = 20), 8-spored, bitunicate, broadly subglobose to obovoid, arranged around outer
layer of ascomata within a palisade layer, short pedicellate, apically rounded, without a distinct ocular chamber and
ring structures. Ascospores 10–26 × 3–8 µm ( = 16 × 5 µm, n = 5), multi-seriate, irregularly arranged, broadly
clavate to ellipsoid, ends rounded, hyaline, 3-septate, without distinct gelatinous sheath. Asexual state: Unknown.
Typ e : —Stephanotheca micromera Syd. & P. Syd., The Phillippine Journal of Science: C Botany 9(2): 179 (1914)
MycoBank No: 215191 (Fig 5)
Foliar epiphytes on lower surface of leaves of Taxotrophis ilicifolia. Sexual state: Ascomata 186–266 µm high ×
175–251 µm diam. ( = 232 × 212 µm, n = 5), superficial on lower leaf host tissues, globose to subglobose, base
flattened, black. Upper wall comprising thick-walled cells, forming a textura porrecta, lower wall poorly
developed. Hamathecium lacking pseudoparaphyses. Asci 16–31 × 16–25 µm ( = 25 × 19 µm, n = 20), 8-spored,
bitunicate, broadly subglobose to obovoid, arranged around outer layer of ascomata within a palisade layer, short
pedicellate, apically rounded, without a distinct ocular chamber and ring structures. Ascospores 10–26 × 3–8 µm (
x
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x
x
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= 16 × 5 µm, n = 5), multi-seriate, irregularly arranged, broadly clavate to ellipsoid, ends rounded, hyaline, 3-
septate, without distinct gelatinous sheath. Asexual state: Unknown.
Material examined:—PHILIPPINES. Palawan, Lake Manguao, on Taxotrophis ilicifolia (Moraceae), E.D.
Merril, April 1913, (S, F6581!, holotype).
FIGURE 5. Stephanotheca micromera (holotype). a. Material label. b–c. Appearance of thyriothecia on host. d. Squash mount of
thyriothecium. e. Cells of upper wall of thyriothecium. f. Arrangement of asci around ascomata. g–i. Asci. j–k. Immature ascospores.
Scale bars: b = 500 µm, c = 200 µm, d = 100 µm, f = 50 µm, e–g = 20 µm, h–k = 10 µm.
Stephanotheca was introduced by Sydow & Sydow (1914) and has only three epithets listed in Index
Fungorum (2013). Stephanotheca is characterized by small, black, superficial, rounded bodies with an elevated
centre. This genus was originally described as Hemisphaeriaceae, but later placed in Microthyriaceae as it
possesses thyriothecia (Stevens & Ryan 1939, Clements & Shear 1931). Lumbsch & Huhndorf (2007, 2010) placed
Stephanatheca in the family Elsinoaceae. Hyde et al. (2013) suggested this genus could be placed in the family
Asterinaceae. Stephanotheca forms solitary, gregarious, superficial, black thyriothecia with ovate to oblong,
longitudinally radiating asci, and hyaline, one-septate ascospores. Another family which might accommodate it is
x
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Elsinoaceae, but the asci are arranged in a single layer in thyriothecia which have a poorly developed base.
Stephanothecaceae may need resurrecting to accommodate Stephanatheca and similar genera if sequence data
becomes available. We retain the species in Asterinaceae even though it is atypical.
Myriangiaceae Nyl., Mémoires de la Société Impériale des Sciences Naturelles de Cherbourg 2: 9 (1854)
Saprobic and parasitic on bark, leaves and branches. Producing raised superficial, black ascostromata in which asci
develop singly in locules that are generally scattered in ascostromata.
Hemimyriangium J. Reid & Piroz., Canadian Journal of Botany 44: 650 (1966)
Typ e : — Hemimyriangium betulae J. Reid & Piroz. , Canadian Journal of Botany 44: 651 (1966)
This taxon appears to be more similar to Myriangium, the type of family Myriangeaceae in having superficial
ascostromata, with a single ascus in each locule, and in the arrangement of locules in the outer layer of the
ascostromata and also due to its saprophytic nature. Hemimyriangium is referred to family Myriangiaceae in
Dissanayake et al. (2014).
Butleria Sacc., Annales Mycologici 12: 302 (1914)
Typ e : — Butleria inaghatahani Sacc., Annales Mycologici 12: 302 (1914)
Butleria is a monotypic genus established by Saccardo (1914). von Arx & Müller (1975) referred this genus to
Myriangiaceae based on ascostromata and two-celled ascospores. Barr (1979) placed this genus under Elsinoaceae.
Kirk et al. (2001), Lumbsch & Huhndorf (2007, 2010), Li et al. (2011) and Hyde et al. (2013) have retained this
genus under the family Elsinoaceae. Butleria has similarities with Elsinoaceae in being a parasite on leaves, but
differs in having ascostromata on both sides of the leaves, with single asci with a small ocular chamber in each
locule and distinct 2-celled, brown ascospores. Butleria shows similarity to the family Myriangiaceae in having
globose, single asci in each locule, but differs in having brown ascospores. The genus has been placed in
Myriangeaceae by Dissanayake et al. (2014).
Micularia Boedijn, Persoonia 2(1): 67 (1961)
Typ e : — Micularia merremiae Boedijn, Persoonia 2(1): 67 (1961)
Boedijn (1961) placed this genus in family Saccardiaceae, while Lumbsch & Huhndorf (2007, 2010) placed
Micularia in the family Elsinoaceae. This placement has followed by Hyde et al. (2013). Even though it is a
parasite on leaves, the inclusion of this genus into Elsinoaceae causes confusion, as it has only 1-septate
ascospores, hairs at the apex of the ascostromata and contains a single ascus in each locule. Dissanayake et al.
(2014) therefore placed Micularia in Myriangeaceae.
Brefeldiellaceae E. Müll. & Arx, in Müller & von Arx, Beiträge zur Kryptogamenflora der Schweiz 11(no. 2): 148
(1962)
Foliar epiphytes or parasites on leaves of various hosts. Thallus comprising a wide area of radiating cells with
darker raised areas under which the ascomata develop and in which saccate to cylindro-clavate asci are formed
(Hyde et al. 2013).
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Saccardinula Speg., Anales de la Sociedad cientifica argentina 19: 257(1885)
Typ e : — Saccardinula guaranitica Sacc Anales de la Sociedad cientifica argentina 19: 257 (1885)
Luttrell (1973) grouped Saccardinula under Saccardinulaceae based on its ascostromata grouping in a radiate,
superficial, cellular membrane (Li et al. 2011). von Arx & Müller (1975) removed this genus to Myriangiaceae and
Lumbsch & Huhndorf (2007) placed Saccardinula in the family Elsinoaceae. Li et al. (2011) suggested that
Saccardinula can be referred to either Microthyriaceae or Brefeldiellaceae, or Saccardinulaceae can be retained as
a distinct family. Saccardinula has globose asci without paraphyses, and forms thyriothecia. Formation of asci and
ascomata of Saccardinula are more similar to Brefeldiella which occurs on leaves and has a thallus comprising an
area of radiating cells with ascomata (Reynolds & Gilbert 2005, Wu et al. 2011, Li et al. 2011). Hyde et al. (2013)
placed Saccardinula in family Brefeldiellaceae, even though it is a less convincing member of the family as it has
globose asci and muriform spores, while the thallus is less-developed.
Dothideomycetes genera incertae sedis
Dothideomycetes is the largest class in Ascomycota. Lumbsch & Huhndorf (2010) in Outline of Ascomycota listed
over 150 genera in Dothideomycetes genera incertae sedis (Thambugala et al. 2014).
Hyalotheles Speg., Revista del Museo de La Plata 15(2): 11 (1908)
Typ e : — Hyalotheles dimerosperma Speg., Revta Mus. La Plata 15(2): 11 (1908)
Li et al. (2011) and Hyde et al. (2013) suggested that this taxon should be placed in Dothideomycetes genera
incertae sedis because it is not a suitable candidate in Elsinoaceae apart from the oblong to ovoid sessile asci.
Chaetothyriaceae Hansf. ex M.E. Barr, Mycologia 71(5): 943 (1979)
Epiphytic or biotrophic on leaves forming mycelium appressed to the host cuticle without penetrating host tissue.
Ascomata are surrounded by a thin pellicle of superficial mycelium forming black sooty mould-like areas on leaves
that are easily detached from the cuticle (Chomnunti et al. 2012).
Beelia F. Stevens & R.W. Ryan, Bulletin of the Bernice Pauahi Bishop Museum, Honolulu, Hawaii 19: 71 (1925)
Typ e:—Beelia suttoniae F. Stevens & R.W. Ryan, Bulletin of the Bernice Pauahi Bishop Museum, Honolulu,
Hawaii 19: 71 (1925)
Beelia was introduced by Stevens & Ryan (Stevens 1925) and placed Beelia in the family Microthyriaceae which
was accepted by Petrak (1953). von Arx & Müller (1975) transferred Beelia to Myriangiaceae based on its
dimidiate ascomata and muriform ascospores. Hawksworth et al. (1995) referred this genus to family Elsinoaceae
where it has been listed by Kirk et al. (2001) and Lumbsch & Huhndorf (2007, 2010). Li et al. (2011) excluded this
genus from Elsinoaceae as Beelia is a superficial biotroph on leaf surfaces which is a characteristic of the family
Chaetothyriaceae. The species appears to be most similar to Phaeosaccardinula suggesting that this genus should
be placed in Chaetothyriaceae.
Sordariomycetes genera incertae sedis
The Sordariomycetes is one of the largest classes in the Ascomycota and the majority of its species are
characterized by perithecial ascomata and in-operculate unitunicate asci. Lumbsch & Huhndorf (2010) in Outline
of Ascomycota, 119 genera were placed under Sordariomycetes genera incertae sedis (Zhang et al. 2006).
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Xenodium Syd., Annales Mycologici 33(1/2): 95 (1935)
Typ e : — Xenodium petrakii Syd., Annales Mycologici 33(1/2): 95 (1935)
Parasitic on leaves. Sexual state: Ascomata scattered, immersed or erumpent, brown to dark brown. Ostiole, with
paraphyses. Peridium comprising hyaline cells of textura porrecta. Paraphyses cylindrical to filiform. Asci 8-
spored, unitunicate, non- fissitunicate, oblong-clavate, apically rounded, short pedicellate. Ascospores irregularly
arranged, cylindrical to filiform, hyaline, 3-septate, hyaline and smooth walled. Asexual state: Xenodiella
Asexual state: Xenodiella Syd., Annales Mycologici 33(1/2): 98 (1935)
Typ e : — Xenodiella petrakii Syd., Annales Mycologici 33(1/2): 98 (1935)
Parasitic on leaves. Basal stroma globose, cells rounded, yellowish-brown. Conidiophore cells, simple or forked-
branched, branches are always 1-celled. Sterile hyphae distinctively curved. Conidia globose or ovate-round, hyaline.
Xenodium was included in the family Elsinoaceae by Lumbsch & Huhndorf (2007, 2010). Hyde et al. (2013)
suggested that this genus should be excluded from Dothideomycetes as it has unitunicate, non-fissitunicate asci. In
this paper we suggest that this genus should be placed under Sordariomycetes genera incertae sedis.
Acknowledgements
We are grateful to the Mushroom Research Foundation, Chiang Rai, Thailand. The herbaria S, PAD, L, BISH, LPS
are thanked for loan of specimens. MFLU grant number 56101020032 and BRG 5580009 are thanked for
supporting studies on Dothideomycetes. KD Hyde thanks The Chinese Academy of Sciences, project number
2013T2S0030, for the award of Visiting Professorship for Senior International Scientists at Kunming Institute of
Botany. Ruvishika S. Jayawardena thanks P. Jayawardena, S. Jayawardena, K.C. Mallikarathna and Samantha C.
Karunarathna for the support given.
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