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219
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN
1179-3163
(online edition)
Copyright © 2014 Magnolia Press
Phytotaxa 176 (1): 219–237
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Article
http://dx.doi.org/10.11646/phytotaxa.176.1.22
The status of Myriangiaceae (Dothideomycetes)
ASHA J. DISSANAYAKE
1,2
, RUVISHIKA S. JAYAWARDENA
1,2
, SARANYAPHAT BOONMEE
2
, KASUN M.
THAMBUGALA
2,3
, QING TIAN
2
, AUSANA MAPOOK
2
, INDUNIL C. SENANAYAKE
2
, JIYE YAN
1
*, YAN
MEI LI
4
, XINGHONG LI
1
, EKACHAI CHUKEATIROTE
2
& KEVIN D. HYDE
2
1
Institute of Plant and Environment Protection, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, People’s
Republic of China
email: jiyeyan@gmail.com
2
Institute of Excellence in Fungal Research and School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
email: kdhyde3@gmail.com
3
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou Academy of Agricultural Sciences, Xiaohe District, Guiyang City,
Guizhou Province 550006, People’s Republic of China
4
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
Abstract
The family Myriangiaceae is relatively poorly known amongst the Dothideomycetes and includes genera which are
saprobic, epiphytic and parasitic on the bark, leaves and branches of various plants. The family has not undergone any
recent revision, however, molecular data has shown it to be a well-resolved family closely linked to Elsinoaceae in
Myriangiales. Both morphological and molecular characters indicate that Elsinoaceae differs from Myriangiaceae. In
Elsinoaceae, small numbers of asci form in locules in light coloured pseudostromata, which form typical scab-like
blemishes on leaf or fruit surfaces. The coelomycetous, “Sphaceloma”-like asexual state of Elsinoaceae, form more
frequently than the sexual state; conidiogenesis is phialidic and conidia are 1-celled and hyaline. In Myriangiaceae,
locules with single asci are scattered in a superficial, coriaceous to sub-carbonaceous, black ascostromata and do not
form scab-like blemishes. No asexual state is known. In this study, we revisit the family Myriangiaceae, and accept ten
genera, providing descriptions and discussion on the generic types of Anhellia, Ascostratum, Butleria, Dictyocyclus,
Diplotheca, Eurytheca, Hemimyriangium, Micularia, Myriangium and Zukaliopsis. The genera of Myriangiaceae are
compared and contrasted. Myriangium duriaei is the type species of the family, while Diplotheca is similar and may
possibly be congeneric. The placement of Anhellia in Myriangiaceae is supported by morphological and molecular data.
Because of similarities with Myriangium, Ascostratum (A. insigne), Butleria (B. inaghatahani), Dictyocyclus (D.
hydrangea), Eurytheca (E. trinitensis), Hemimyriangium (H. betulae), Micularia (M. merremiae) and Zukaliopsis (Z.
amazonica) are placed in Myriangiaceae. Molecular sequence data from fresh collections is required to confirm the
relationships and placement of the genera in this family.
Key words: Anhellia, Ascostratum, Butleria, Dictyocyclus, Diplotheca, Eurytheca, Hemimyriangium, Micularia,
morphology, Myriangium, saprobes, Zukaliopsis
Introduction
The family Myriangiaceae is a relatively poorly known, but interesting family of Dothideomycetes (Ascomycota)
comprising the genera Anhellia, Diplotheca, Eurytheca and Myriangium (Lumbsch & Huhndorf 2007, 2010). The
species are generally saprobic in nature, occurring on dead leaves, stems or bark of various host plants (Gäumann
1928). Distribution of Myriangiaceae is chiefly tropical or sub-tropical (Barr 1979). The consistency of the
ascostromata is often suggestive of the Hypocreales, although in Myriangium itself they are more cartilaginous to
gelatinous or generally brittle (Gäumann 1928). The family is characterized by only one ascus in each locule,
superficial, erumpent, scattered, solitary or gregarious, generally dark ascostromata, apedicellate or short
pedicellate asci and smooth to verruculose ascospores with 1–9-transverse septa, 0–5-longitudinal septa, rarely
having a mucilaginous sheath (only in Diplotheca). The asexual states of the family are unknown. We are
DISSANAYAKE ET AL.
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Phytotaxa 176 (1) © 2014 Magnolia Press
monographing the genera of all families of Dothideomycetes. By providing modern descriptions with
comprehensive micrographs we hope to stimulate recollection of genera and species so that sequence data from
fresh specimens can be generated and analyzed to provide a natural classification. In this paper we have added six
genera (Ascostratum, Butleria, Dictyocyclus, Hemimyriangium, Micularia and Zukaliopsis) to the Myriangiaceae,
as they have similar morphological characters.
Historical overview of Myriangiaceae
Myriangiaceae is a family of saprobes, but with some genera being parasitic and epiphytic on leaves. The family
was introduced by Nylander (1854) and included two species, Myriangium duriaei Mont. & Berk. (1845) and M.
curtisii Berk. & Mont. (1845). Millardet (1870) placed the family near the Tuberaceae. Many classical treatments
have placed Elsinoaceae in synonymy with Myriangiaceae, but Höhnel (1909) was convinced that the former
constituted a separate family. Petch (1924) provided a thorough review of Myriangium. Gäumann et al. (1928)
divided Myriangiaceae into two groups, based on the formation of the asci. In one group, the asci are scattered
irregularly over the whole ascostromata, while in the other group they are localized in definite regions of the
ascostromata, which are differentiated into sterile and fertile parts. No other authors have followed this grouping.
Clements & Shear (1931) placed the family Myriangiaceae in the order Dothideales, due to the presence of
ascostromata containing numerous locules, while von Arx (1963) treated Myriangiaceae in Myriangiales. von Arx
& Müller (1975) treated Myriangiaceae as a parasitic family on higher plants or on other fungi, especially
ascomycetes. According to Petch (1924), Myriangium duriaei, M. curtisii, M. montagnei Berk. (1845) and M.
thwaitesii Petch (1924) are entomogenous, but this needs confirming. Barr (1979) and Eriksson (1981) were of the
opinion that two separate families should be maintained as Elsinoaceae and Myriangiaceae with the latter
predominantly occurring on branches, while Elsinoaceae is restricted to foliar pathogens.
The order Myriangiales was introduced by Stärback (1899) to accommodate species having crustose
ascostromata and muriform ascospores, and several apparently unrelated forms, with the distinguishing characters
of angiocarpous fruiting bodies with asci arranged irregularly in colourless locules and released by the breaking
away of the covering layer. The families Myriangiaceae and Saccardiaceae (Höhnel 1909) were treated by von Arx
(1963) as comprising the Myriangiales. Luttrell (1973) distinguished Atichiaceae (Raciborski 1900) and
Saccardiaceae in Myriangiales indicating that the asci are borne in discoid ascostromata in a hymenium-like layer
and are separated from each other by paraphyses-like filaments often forming an epithecium. von Arx & Müller
(1975) gave a new description for the order with asci borne singly, often at different levels in small loculi immersed
in a stromatic body, which opens by rupture or dehiscence. Barr (1987) distinguished the family Myriangiaceae and
included taxa that formed a pseudoparenchymatous or plectenchymatous ascostromata with interascal tissue
persisting between mono-ascal cavities. Kirk et al. (2008) included three families, Cookellaceae, Elsinoaceae and
Myriangiaceae in Myriangiales, while Lumbsch & Huhndorf (2010) accepted only Elsinoaceae and Myriangiaceae
in the order based on phylogenetic results.
Various treatments have been proposed for the genera in family Myriangiaceae (Table 1). Nylander (1854)
placed a single genus Myriangium, with two species under this family, but von Arx & Müller (1975) placed 16
genera in the family. After differentiating Elsinoaceae as a separate family from Myriangiaceae based on molecular
evidence, Schoch et al. (2006) maintained only Anhellia, Diplotheca, Eurytheca and Myriangium in Myriangiaceae.
This classification was followed in Lumbsch & Huhndorf (2007, 2010). The familial position of Anhellia was,
however, considered uncertain by Lumbsch & Huhndorf (2010), but in the phylogenetic analyses by Pinho et al.
(2012) its inclusion in this family was supported. The placement of Eurytheca is also confused. Theissen & Sydow
(1917) placed Eurytheca in the family Saccardiaceae (parts of the Saccardiaceae were classified by Müller & von
Arx (1965) within the family Schizothyriaceae, while Lumbsch & Huhndorf (2010) referred the genus to the
Myriangiaceae). Jayawardena et al. (2014) separated Myriangiaceae from Elsinoaceae in their phylogenetic analysis
with high bootstrap support value (82%). They analyzed 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) genes from 16 taxa of Myrangiales
and the resulting tree supported Elsinoaceae and Myriangiaceae as distinct families. In this study we re-examine
Anhellia, Diplotheca, Eurytheca and Myriangium and provide full descriptions, photographic plates and notes
suggesting where these genera should be placed. Ascostratum, Butleria, Dictyocyclus, Hemimyriangium, Micularia
and Zukaliopsis were also studied and are considered to also belong in Myriangiaceae. Further molecular sequence
data from fresh collections is required to confirm the relationships and placement of the genera in Myriangiaceae.
Phytotaxa 176 (1) © 2014 Magnolia Press
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THE STATUS OF MYRIANGIACEAE
TABLE 1. Genera included in Myriangiaceae by various authors.
Materials and methods
Type specimens of Anhellia tristis, Ascostratum insigne, Butleria inaghatahani, Dictyocyclus hydrangea,
Diplotheca tunae, Eurytheca trinitensis, Hemimyriangium betulae, Micularia merremiae, Myriangium duriaei and
Zukaliopsis amazonica were obtained from L, S, PC, BPI and IMI. Fruiting bodies were rehydrated in 5% KOH
prior to examination and sectioning. Specimens were examined under a stereo microscope and fine forceps were
used to remove one or two ascostromata, which were mounted in water or cotton blue. Observations and
photographs were made with a Nikon Eclipse 80i, and a Discovery v-8 microscope. Hand sections were cut with a
sharp razor blade. The sections were transferred to a drop of water or a drop of cotton blue for examination and
photography. Cross section of Myriangium duriaei was redrawn from Gäumann (1928).
Results
Taxo n o m y
Myriangiaceae Nylander, Mémoires de la Société Impériale des Sciences Naturelles de Cherbourg 2: 9 (1854)
MycoBank: MB81866
Synonym: Phymatosphaeriaceae Speg., Anal. Soc. cient. argent. 26(1): 57 (1888)
Saprobic on bark, leaves and branches, some genera epiphytic or parasitic on leaves. Sexual state: Ascostromata
superficial, scattered, solitary or aggregated, coriaceous to sub-carbonaceous, semi-immersed or immersed to
erumpent, generally dark, globose to oval, sometimes surrounded by the remains of the ruptured epidermis, multi-
loculate, locules generally in the upper layer or scattered throughout the ascostromata. Loculues with single ascus,
ostiolate. Ostiole minute or asci pushed out through pseudoparenchymatous cells. Paraphyses absent. Cells of
ascostromata comprising pseudo-parenchymatous cells of pale yellow to brown pigmented textura angularis,
textura globusa or textura intricata. Asci 8-spored, bitunicate, fissitunicate, globose to subglobose, apedicellate or
with a minute pedicel, apically rounded with indistinct ocular chamber. Ascospores hyaline to sub-hyaline or
brown, oblong or fusiform with slightly acute ends, with 1–9-transverse septa, with 0–5-longitudinal septa,
muriform, smooth-walled to verruculose, rarely having a sheath. Asexual state: Unknown.
Nylander (1854) von Arx & Müller (1975) Barr (1987) Lumbsch & Huhndorf (2007, 2010) This paper (2014)
Myriangium Anhellia
Beelia
Butleria
Cookella
Diplotheca
Elsinoe
Hyalotheles
Micularia
Molleriella
Myriangium
Pycnoderma
Pycnodermina
Saccardinula
Stephanotheca
Uleomyces
Xenodium
Angatia
Myriangium
?Anhellia
Diplotheca
?Eurytheca
Myriangium
Anhellia
Ascostratum
Butleria
Dictyocyclus
Diplotheca
Eurytheca
Hemimyriangium
Micularia
Myriangium
Zukaliopsis
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Key to genera of Myriangiaceae
1. Locules located only at the upper layer of the ascostromata ................................................................................................ 2
Locules scattered irregularly throughout the ascostromata.................................................................................................... 5
2. Ascospores with only transverse septa......................................................................................................... Hemimyriangium
Ascospores muriform............................................................................................................................................................. 3
3. Saprobic on bark ...................................................................................................................................................Myriangium
Parasitic on leaves .................................................................................................................................................................. 4
4. Ascostromata on upper surface of the leaf............................................................................................................. Zukaliopsis
Ascostromata on lower surface of the leaf..................................................................................................................Anhellia
5. Ascospores hyaline................................................................................................................................................................. 6
Ascospores brown ....................................................................................................................................................... Butleria
6. Ascospores with only transverse septa................................................................................................................................... 7
Ascospores muriform............................................................................................................................................................. 9
7. Ascospores with only one transverse septum........................................................................................................... Micularia
Ascospores with 3–9-transverse septa ................................................................................................................................... 8
8. Ascostromata connected to the host with a prominent stalk ................................................................................ Ascostratum
Ascostromata embedded in the host, erumpent through ruptured epidermis........................................................... Eurytheca
9. Ascospores covered with a mucilaginous sheath....................................................................................................Diplotheca
Ascospores without a mucilaginous sheath..........................................................................................................Dictyocyclus
Type genus:—Myriangium Montagne & Berkeley, London Journal of Botany 4: 72 (1845)
Possible synonyms
Phymatodiscus Speg., Boln Acad. nac. Cienc. Córdoba 23: 484 (1919)
Phymatosphaeria Pass., Nuovo G. bot. ital. 7: 188 (1875)
Pyrenotheca Pat., Bull. Soc. bot. Fr. 33: 155 (1886)
Saprobic on bark of Acer, Citrus, Fraxinus, and other plants. Sexual state: Ascostromata superficial, scattered,
solitary or aggregated, coriaceous to sub-carbonaceous, erumpent, dark brown to black, globose to oval, multi-
loculate, locules scattered throughout the ascostromata but most are concentrated at the upper region, containing
only one ascus in each locule, ostiolate. Ostioles minute. Paraphyses absent, Cells of ascostromata comprising
thick-walled pigmented cells of textura angularis. Asci 8-spored, bitunicate, fissitunicate, globose to subglobose,
apedicellate, apically rounded with indistinct ocular chamber. Ascospores hyaline to subhyaline, elliptical to
oblong, fasciculate, muriform, usually with 4–7-transverse septa and one or more longitudinal septa, smooth-
walled, lacking a sheath. Asexual state: Unknown.
Notes:—The genus Myriangium was erected by Montage & Berkeley (1845), with M. duriaei as the type
species. Montage & Berkeley (1845) added M. curtisii and placed both species in the lichen family, Collemaceae.
Nylander (1854) erected the family Myriangeaceae (Miller 1940). The generic concept of Myriangium included
multi-loculate apothecia with asci forming in single locules (Miller 1938). Wolf & Wolf (1947) considered that M.
duriaei, M. curtisii, M. montagnei and M. thwaitesii, which are included in Myriangium, are parasitic on scale
insects. Miller (1938) studied the morphology and cytology of M. duriaei and M. curtsii and found that both occur
as parasites on scale insects attacking Nyssa sylvatica, Carya illinoensis and other broad-leaved trees in south-
eastern United States. In this study we have examined the type material of Myriangium and it does not appear to be
associated with any scale insects, although this should be confirmed in fresh collections. There are 50 species
epithets for Myriangium in Index Fungorum (2013) and the genus requires a restudy. There have been few
molecular studies on this genus. Lumbsch & Lindemuth (2001) used LSU and SSU gene data to analyze the major
lineages of Dothideomycetes in which the Myriangium species were included in the order Myriangiales. Schoch et
al. (2006) used M. duriaei and four Elsinoe spp. in their phylogenetic analyses and found that these Elsinoe species
belong to a different family, Elsinoaceae. This concept was confirmed by Schoch et al. (2009). Jayawardena et al.
(2014) used M. duriarei and M. hispanicum J.B. Martínez (1931) in their phylogenetic analysis to differentiate
Myriangiaceae from Elsinoaceae with a high bootstrap value (82%). Fresh collections and molecular studies are
needed to substantiate these results.
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THE STATUS OF MYRIANGIACEAE
Type species:—Myriangium duriaei Montagne & Berkeley, London Journal of Botany. 4: 73 (1845) MycoBank:
218324 (Fig. 1)
Saprobic on bark. Sexual state: Ascostromata 1.5–1.8 × 1.1–1.4 mm ( = 1.6 × 1.3 mm, n = 5), superficial,
scattered, solitary or aggregated, coriaceous to sub-carbonaceous, erumpent, dark brown to black, globose to oval,
multi-loculate, locules scattered throughout the ascostromata but concentrated mainly at the upper region, locules
with one ascus, ostiolate. Ostioles minute. Cells of the ascostromata comprising several layers of thick-walled pale
yellow pigmented cells of textura angularis. Paraphyses not observed. Asci 28–35 × 23–28 μm ( = 30.4 × 25.2
μm, n = 13), 8-spored, bitunicate, fissitunicate, globose to subglobose, apedicellate, apically rounded with
indistinct ocular chamber. Ascospores 20–25 × 7–10 μm ( = 22 × 9 μm, n = 12), hyaline, oblong or fusiform with
slightly acute ends, muriform, with 4–7-transverse septa, 1–5-longitudinal septa, smooth-walled, lacking a sheath.
Asexual state: Unknown.
Material examined:—ALGERIA. MC Durieu de Maisonneuve, on Lentiscos vivos (= Pistacia lentiscus,
Anacardiaceae) (PC, NL 8414!, holotype).
FIGURE 1. Myriangium duriaei (holotype). a. Ascostromata on host surface. b–c. Locules in cross-section of the ascostromata (b
redrawn from Gäumann 1928). d. Apedicellate ascus with 8 ascospores. e–f. Asci stained in cotton blue. g. Muriform, hyaline
ascospore. h. Ascospore stained in cotton blue. Scale bars: b = 500 μm, c–e = 30 μm, f = 20 μm, g–h = 10 μm.
x
x
x
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Anhellia Raciborski, Parasitische Algen und Pilze Javas 2: 10 (1900)
Parasitic on leaves. Sexual state: Ascostromata superficial, scattered, solitary or gregarious, carbonaceous,
immersed, dark brown to black, hemispherical or globose to oval, connected to the host by erumpent hypostroma,
multi-loculate, locules scattered in the upper layer of the ascostromata, one ascus in each locule, ostiolate. Ostioles
minute, asci pushed out through pseudoparenchymatous cells. Paraphyses absent. Cells of the ascostromata
comprising pale yellow to brown textura angularis. Asci 8-spored, bitunicate, fissitunicate, globose to saccate,
pedicellate, apically rounded with indistinct ocular chamber. Ascospores hyaline or yellowish brown, elliptical to
oblong, muriform, with 4–5-transverse septa, 1–2-longitudinal septa, constricted at the septa, smooth-walled,
lacking a sheath. Asexual state: Unknown.
Notes:—Anhellia was introduced by Raciborski (1900) in the “Myriangiearum” and comprised a single
species, A. tristis. This genus remained monotypic until von Arx (1963) added A. calami (Racib.) Arx, A.
escharoides (Syd.) Arx, A. purpurascens (Rehm.) Arx, A. lantanae (Henn.) Arx and A. nigra (Viégas) Arx. Luttrell
(1973) placed this genus in Myriangiaceae based on its discoid to hemiglobose ascostromata, with stalk erumpent
through epidermis from an intraepidermal hypostroma. Anhellia tristis, A. escharoides and A. tetracerae (Hansf.)
Arx, with obviously larger dimensions than A. tabebuiae Inácio & Dianese have been found on Vacc in iu m
teysmannianum and Tetracera alnifolia (Boedijn 1961, von Arx 1963). Anhellia calami has been reported on
Calamus sp. (von Arx 1963). Anhellia nectandrae differs from A. escharoides by having paler ascospores and
narrower asci, while A. tetracerae and A. tristis are different in having hyaline ascospores (Raciborski 1900, von
Arx 1963). Anhellia nectandrae was the first Anhellia species reported parasitizing a member of Lauraceae and
differs morphologically and phylogenetically from other previously described species (Pinho et al. 2012). Anhellia
tabebuiae is a first record of Anhellia on Tabebuia species (Pinho et al. 2012). The genus currently comprises nine
species, which are mainly tropical in distribution (von Arx & Müller 1975, Kirk et al. 2008) and is characterized by
its dark ascostromata, bearing many-celled ascospores inside bitunicate, fissitunicate asci mostly in the upper layer,
connected with the host by an erumpent, discoid hypostroma (von Arx & Müller 1975, Pereira 2003). Pinho et al.
(2012) investigated the phylogenetic position of Anhellia (Myriangiales) by analysis of nucleotide sequences of the
ribosomal large subunit (LSU) and ITS regions. Twenty-eight aligned sequences were obtained from GenBank and
this was the first attempt to resolve the placement of the genus Anhellia within Myriangiales. The genus Anhellia is
strongly supported in Myriangiaceae in this phylogenetic analysis.
Type species:—Anhellia tristis Raciborski, Parasitische Algen und Pilze Javas 2: 10 (1900) MycoBank: 219955 (Fig. 2)
Parasitic on leaves. Sexual state: Ascostromata 0.8–1.1 × 0.2–0.3 mm ( = 1 × 0.25 mm, n=5), superficial on the
lower surface, scattered, solitary, carbonaceous, immersed, flattened, brown to black, globose to oval, connected to
the host by an erumpent hypostroma, multi-loculate, locules scattered in the upper layer of the ascostromata, one
ascus in each locule, ostiolate. Ostioles minute, asci pushed out through pseudo-parenchymatous cells. Cells of the
ascostromata comprising several layers of thin-walled pale yellow to brown textura angularis cells. Paraphyses
absent. Asci 34–89 × 18–33 µm ( = 57 × 28 µm, n = 15), 8-spored, bitunicate, fissitunicate, globose to saccate,
pedicel short and broad, indistinct ocular chamber. Ascospores 20–32 × 6–12 µm ( = 27 × 10 µm, n = 33),
hyaline, elliptical to oblong, muriform, 4–5-transverse septa, 1–2-longitudinal septa, constricted mainly at the first
forming septum, with slightly acute or tapered ends, smooth-walled, lacking a sheath. Asexual state: Unknown.
Material examined:—INDONESIA. Java, Gedeh, on leaves of Vaccinium teysmannia (Ericaceae), M.
Raciborski (S, F47931!, isotype).
Ascostratum Sydow & P. Sydow, Annales Mycologici 10(1): 41 (1912)
Saprobic on bark in terrestrial habitats. Sexual state: Ascostromata superficial, scattered, solitary or aggregated,
carbonaceous, semi-immersed to erumpent, black, globose to sub-globose, ruptured epidermis, multi-loculate,
locules scattered in the middle region of the ascostromata, ostiolate. Ostioles minute, ascospores release via
breakdown of the cells of ascostromata. Hamathecium lacking paraphyses. Cells of the ascostromata comprising
flattened textura intricate. Asci 8-spored, bitunicate, fissitunicate, oval to globose, short pedicellate or apedicellate,
apically rounded with indistinct ocular chamber. Ascospores hyaline, ellipsoid to fusiform, obtuse at both ends,
7–9-transverse septa, longitudinal septa absent, smooth-walled, lacking a sheath. Asexual state: Unknown.
x
x
x
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THE STATUS OF MYRIANGIACEAE
Notes:—Ascostratum was introduced by Sydow & Sydow (1912) and was typified by Ascostratum insigne
Syd. & P. Syd. The genus was placed in Dothideomycetes, genera incertae sedis by Lumbsch & Huhndorf (2010).
Ascostratum is characterized by ascostromata with numerous locules scattered in the middle region of the
ascostromata, semi-immersed or erumpent, asci oval to globose, and ascospores hyaline with 7–9 transverse septa.
Ascostratum shows similarities with Anhellia by releasing ascospores via breakdown of the cells of ascostromata.
Ascostratum includes only two species (Index Fungorum 2013) and phylogenetic analysis is lacking for either
species and is needed to establish the natural classification of this genus. Here we provide detailed description and
illustrations as a contribution for future work. We refer Ascostratum to the family Myriangiaceae based on its
morphological similarity.
FIGURE 2. Anhellia tristis (isotype). a–b. Herbarium material. c–d. Ascostromata on host substrate. e–f. Cross section of the
ascostromata showing locules mostly at the periphery. g. Immature ascus stained in cotton blue. h. Mature ascus. i–j. Mature asci
stained in cotton blue. k. Dehiscence of mature ascus. l. Smooth-walled ascospore. m–n. Ascospores stained in cotton blue. Scale bars:
c = 200 μm, d–f = 100 μm, g–k = 10 μm, l–n = 5 μm.
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Type species:—Ascostratum insigne Sydow & P. Sydow, Annales Mycologici 10(1): 41 (1912) MycoBank: 214383
(Fig. 3)
Saprobic on bark of Euphorbia. Sexual state: Ascostromata 0.65–1.2 × 0.55–0.85 mm ( = 0.95 × 0.70 mm, n =
10), superficial, scattered, solitary or aggregated, carbonaceous, semi-immersed to erumpent, black, oval to
globose, ruptured epidermis, multi-loculate, locules spread around the middle region of the ascostromata, ostiolate.
Ostiole minute, ascospores release via breakdown of the cells of ascostromata. Hamathecium lacking paraphyses.
Cells of the ascostromata composed of dark brown to black thick-walled flattened textura intricate cells. Asci
70–96 × 52–70 μm ( = 81 × 60 μm, n = 10), 8-spored, bitunicate, fissitunicate, oval to globose, short pedicellate
or apedicellate, apically rounded with indistinct ocular chamber. Ascospores 53–68 × 19–24 μm ( = 60 × 21 μm,
n = 10), hyaline, ellipsoid to fusiform, obtuse at each end, 7–9-transverse septa, longitudinal septa absent,
constricted at each septum, smooth-walled, lacking a sheath. Asexual state: Unknown.
Material examined:—SOUTH AFRICA. KwaZulu-Natal, Natal, Amanzintote, on Euphorbia bark
(Euphorbiaceae), E.M. Doidge, 10 July 1911 (S, F7426!, holotype).
FIGURE 3. Ascostratum insigne (holotype). a–c. Herbarium material. d–f. Ascostromata on host surface. g–i. Cross section of
ascostromata. j–l. Immature asci with distinct, short pedicel in j. m. Asci with ascospores. n–r. Ascospores. Scale bars: d = 1000 µm,
e–f = 200 µm, g = 50 µm , h–i = 20 µm , j–r = 10 µm.
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Butleria Saccardo, Annales Mycologici 12: 302 (1914)
Parasitic on leaves forming leaf spots. Sexual state: Ascostromata superficial, scattered, solitary or aggregated,
erumpent, black, globose to oblong, multi-loculate, numerous locules distributed at diơerent levels inside
ascostromata, one ascus inside each locule, ostiolate. Ostiole minute. Paraphyses absent. Cells of ascostromata are
textura angularis. Asci 8-spored, bitunicate, fissitunicate, globose to sub-globose, apedicellate, apically rounded
with indistinct ocular chamber. Ascospores brown, oblong to ovoid, 1-transverse septum, longitudinal septa absent,
smooth-walled, constricted at the septum, lacking a sheath. Asexual state: Unknown.
Notes:—Butleria is a monotypic genus established by Saccardo (1914). von Arx & Müller (1975) referred this
genus to Myriangiaceae, while Barr (1979) placed it under Elsinoaceae. Kirk et al. (2001), Lumbsch & Huhndorf
(2007, 2010), Li et al. (2011) and Hyde et al. (2013) have retained this genus under family Elsinoaceae. Butleria
has similarities with Myriangiaceae in having a single ascus in each locule. Therefore the genus can be better
placed under the family Myriangiaceae. New collections of this genus and molecular analysis are needed to clarify
the familial placement.
Type species:—Butleria inaghatahani Saccardo, Annales Mycologici 12: 302 (1914) MycoBank: 156750
Notes:—This species is described and illustrated in Li et al. (2011)
Dictyocyclus Sivan., W.H. Hsieh & Chi Y. Chen, Botanical Journal of the Linnean Society 126(4): 324 (1998)
Foliar epiphyte on the lower surface of leaves. Sexual state: Ascostromata superficial, scattered, solitary,
immersed, black, discoid, multi-loculate, opening by irregular fissures; locules arranged around a raised, sterile,
central column of ascostromata, ostiolate. Ostiole minute. Paraphyses absent. Cells of the ascostromata comprising
a single stratum of more or less thick-walled textura angularis brown cells. Asci 8-spored, bitunicate, fissitunicate,
obovoid, thick-walled, apedicellate, apically rounded. Ascospores hyaline, obovoid, partially overlapping, with
broadly to narrowly rounded ends, muriform, 3–5-transverse septa, one longitudinal or oblique septum in each of
the end cells, smooth-walled, lacking a mucilaginous sheath. Asexual state: Unknown.
Notes:—Dictyocyclus was introduced by Sivanesan et al. (1998) as a monotypic genus of the family
Parmulariaceae. Dictyocyclus shares most of the characteristics similar to Myriangium, including a single ascus
inside each locule and muriform ascospores.
Type species:—Dictyocyclus hydrangeae Sivan., W.H. Hsieh & Chi Y. Chen, Botanical Journal of the Linnean
Society 126(4): 324 (1998) MycoBank No: 444846 (Fig. 4)
Foliar epiphyte on the lower surface of Hydrangea integrifolia leaves, appearing as small black spots. Sexual state:
Ascostromata 0.4–0.5 × 0.5–0.6 mm ( = 0.45 × 0.55 mm, n = 5), superficial, scattered, solitary, immersed, black,
discoid, multi-loculate, opening by irregular fissures; locules arranged around a raised, sterile, central column of
ascostromata, ostiolate, Ostiole minute. Paraphyses absent. Cells of the ascostromata comprising thick-walled
lightbrown textura angularis. Asci 28–50 × 22–30 μm ( = 37 × 26 μm, n = 20), 8-spored, bitunicate, fissitunicate,
obovoid, thick-walled, apedicellate, apically rounded with indistinct ocular chamber. Ascospores 13–20 × 5–9 μm
( = 17 × 6 μm, n = 30), hyaline, obovoid, partially overlapping, thin-walled, broadly to narrowly rounded ends,
muriform, 3–5-transverse septa, one longitudinal or oblique septum in each of the end cells, smooth-walled,
lacking a sheath. Asexual state: Unknown.
Material examined:—TAIWAN. Taitung Hsien, on leaves of Hydrangea integrifolia (Hydrangeaceae), C.Y.
Chen 1995 (IMI, 371179!, holotype).
Diplotheca Starbäck, Botaniska Notiser: 30 (1893)
Parasitic on bark. Sexual state: Ascostromata superficial, scattered, aggregated, carbonaceous, immersed to
erumpent, black, globose to sub-globose, multi-loculate, locules generally located at the lower part of the
ascostromata, ostiolate, Ostioles minute. Paraphyses absent. Cells of the ascostromata comprising several layers of
thick-walled pale brown cells of textura globosa. Asci 8-spored, bitunicate, fissitunicate, globose or ellipsoid,
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minute pedicel, apically rounded with indistinct ocular chamber. Ascospores hyaline or pale gray, fusiform, both
ends bluntly tapered, muriform, 3–6-transverse septa, 1–3-longitudinal septa, constricted at septum, smooth-
walled, surrounded by a mucilaginous sheath. Asexual state: Unknown.
Notes:—Diplotheca was introduced by Starbäck (1893). Diplotheca tunae was transferred to Myriangium by
Petrak (1929), however, due to its parasitic nature, Miller (1940) suggested retainning the genus name Diplotheca.
Petrak (1951) compared the size of ascostromata and accepted the separation of D. tunae from Myriangium.
Lumbsch & Huhndorf (2007, 2010) placed Diplotheca in the family Myriangiaceae. Diplotheca is parasitic on
various plant species in Cactaceae, especially on Epiphyllum phyllanthus an epiphytic plant, forming scab-like
symptoms on most stems (Pereira & Barreto 2006).
FIGURE 4. Dictyocyclus hydrangea (holotype). a. Herbarium material. b. Ascostroma on host surface. c–d. Squash mount of
ascostromata. e. Cross section through ascostroma. f. Immature ascus. g–i. Mature asci with 8 ascospores. j–m. Ascospores. Scale
bars: c = 200 µm, d–e = 50 µm, f = 10 µm, g–i = 20 µm, j–m = 10 µm.
Type species:—Diplotheca tunae (Spreng.) Starbäck, Botaniska Notiser: 30 (1893) MycoBank No: 247593 (Fig. 5)
Parasitic on bark. Sexual state: Ascostromata 0.2–0.3 × 0.25–0.3 mm ( = 0.25 × 0.3 mm, n = 5), superficial,
scattered, aggregated, carbonaceous, immersed to erumpent, black, globose to sub-globose, multi-loculate, locules
located at the lower part of the ascostromata, ostiolate. Ostioles minute. Paraphyses absent. Cells of the
ascostromata brown to dark brown, comprising several layers of thick-walled textura globosa. Asci 46–52 × 40–46
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µm ( = 48 × 41 µm, n = 20), 8-spored, bitunicate, fissitunicate, globose or slightly ellipsoid, minute pedicel,
apically rounded with indistinct ocular chamber. Ascospores 25–30 × 12–14 µm ( = 27 × 12 µm, n = 20), hyaline
or pale grey, oblong-oval to elliptical, both ends bluntly tapered, muriform, with 3-transverse septa, 1–3-
longitudinal septa, slightly constricted at the septa, smooth-walled, covered with a mucilaginous sheath. Asexual
state: Unknown.
Material examined:—PUERTO RICO. Mayaguez, on Opuntia sp. (Cactaceae), F.L. Stevens 1913 (BPI,
684149!, isotype).
FIGURE 5. Diplotheca tunae (isotype). a. Herbarium material. b. Ascostromata occurring on substrate. c. Partial section through
ascostroma. d. Peridium wall. e–f. Asci. g. Ascospores. Note colour change in 3% KOH, becoming pinkish or purple in c, f and g.
Scale bars: b = 500 µm, c = 100 µm, d–g = 20 µm.
Eurytheca De Seynes, Bulletin de la Société botanique de France 25: 88 (1878)
Saprobic on bark. Sexual state: Ascostromata superficial, scattered, solitary or aggregated, carbonaceous, erumpent
through the ruptured epidermis, black, orbicular to irregular, multi-loculate, locules scattered in the outer region of
the ascostromata, one ascus in each locule, ostiolate. Ostiole minute. Paraphyses absent. Cells of the ascostromata
comprising several layers of thick-walled textura globosa pigmented cells. Asci 8-spored, bitunicate, fissitunicate,
globose to saccate, short pedicellate, with indistinct ocular chamber. Ascospores hyaline, fusiform, 3–6-transverse
septa, longitudinal septa absent, not constricted at the septa, base or both base and apical end bluntly tapered,
smooth-walled, lacking a sheath. Asexual state: Unknown.
Notes:—The genus Eurytheca was erected by De Seynes (1878), and placed under family Saccardiaceae by
Höhnel (1917). Wolf & Wolf (1947) remarked that this genus belongs to “Myriangiaceen”. Lumbsch & Huhndorf
(2007) placed this genus under family Myriangeaceae. In the original description of the type species E. trinitensis
(Sydow & Sydow 1914), the ascospores are described as 10–14-septate while Theissen & Sydow (1917) describe
them as 4–5-septate. Lumbsch & Huhndorf (2007, 2010) placed Eurytheca in family Myriangiaceae. The spores
examined in the current study showed only 3–6-transverse septa. There are only three species epithets for this
genus (Index Fungorum 2013). Fresh collections and phylogenetic analysis are needed to confirm the familial
placement of this genus in Myriangiaceae.
Type species:—Eurytheca trinitensis Sydow & P. Sydow, Annales Mycologici 13(1): 40 (1915) MycoBank No:
221235 (Fig. 6)
Saprobic on bark. Sexual state: Ascostromata 1–2 × 0.6–1 mm ( = 1.5 × 0.9 mm, n = 5), superficial, scattered,
solitary or aggregated, carbonaceous, erumpent through the ruptured epidermis, black, orbicular to irregular, multi-
loculate, locules scattered in the outer region of the ascostromata, one ascus in each locule, ostiolate. Ostioles
minute. Paraphyses absent. Cells of the ascostromata comprising several layers of thick-walled textura globosa
pigmented cells. Asci 10–16 × 8–15 μm ( = 13 × 11 μm, n = 10), 8-spored, bitunicate, fissitunicate, globose to
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saccate, short pedicellate, with indistinct ocular chamber. Ascospores 8–23 × 2–7 μm ( = 14 × 5 μm, n = 10),
hyaline, fusiform, 3–6-transverse septa, longitudinal septa absent, not constricted at the septa, base or both base and
apical end bluntly tapered, smooth-walled, lacking a sheath. Asexual state: Unknown.
Material examined:—TRINIDAD. Tobago, Balanchisseuse, on Truncum arboris viventis (Bromeliaceae),
W.E. Broadway, 15 May 1908 (S, F11411!, holotype).
FIGURE 6. Eurytheca trinitensis (holotype). a. Herbarium material. b. Ascostromata on host substrate. c. Section of ascostroma. d–e.
Asci arrangement within hamathecium. f. Immature ascus. g–h. Mature asci with short pedicel bearing eight irregularly arranged
ascospores. i–k. Smooth-walled, hyaline ascospores. l. Ascospore stained with cotton blue. Scale bars: c–f = 100 mm, g–l = 10 mm.
Hemimyriangium Reid & Pirozynski, Canadian Journal of Botany 44: 650 (1966)
Saprobic on twigs forming spots. Sexual state: Ascostromata superficial, scattered, solitary or gregarious,
carbonaceous, semi-immersed to erumpent, black, globose to oval, multi-loculate, locules arrange in two layers in
the outer layer of the ascostromata, one ascus in each locule, ostiolate. Ostioles minute. Paraphyses absent. Cells of
the ascostromata comprised thick-walled textura angularis. Asci 8-spored, bitunicate, fissitunicate, globose,
apedicellate, apically rounded with indistinct ocular chamber. Ascospores hyaline, oblong to oval, 1–4- transverse
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septa, longitudinal septa absent, constricted at the septa, smooth-walled, lacking a sheath. Asexual State:
Unknown.
Notes:—The hemispherical shape of the ascostromata of Hemimyriangium has been assumed due to the
sporogenous layer, which has formed a hemispherical resin droplet. As the resin diffuses the ascostromata seems
flat in shape.
This genus was placed in the family Myriangiaceae by Reid & Pirozynski (1966). Though Lumbsch &
Huhndorf (2007) placed Hemimyriangium in the family Elsinoaceae it shows similarity to type genus Myriangium
having one ascus in each locule and the arrangement of locules in the outer layer of the ascostromata.
Hemimyriangium also bears some superficial resemblance to ascostromata of genus Anhellia (Reid & Pirozynski
1966). Piley & Larsen (1968) also were able to show that Hemimyriangium has a close phylogenetic relationship
with Myriangium and Diplotheca. This taxon does not appear to be a candidate for Elsinoaceae as it has a single
ascus in locules scattered throughout the outer layer of ascostromata. We suggest Hemimyriangium is better
referred to the family Myriangiaceae than in Elsinoaceae. Fresh collections and molecular analyses are needed to
clarify the familial position of this genus.
Type species:—Hemimyriangium betulae Reid & Pirozynski, Canadian Journal of Botany 44: 651 (1966)
MycoBank No: 331874 (Fig. 7)
Saprobic on twigs forming black superficial spots on drops of transparent, yellowish resinous secretions produced
by peltate glandular hairs of the host. Sexual state: Ascostromata 2–4 × 3–4 mm ( = 3.2 × 3.6 mm, n = 12),
superficial, scattered, solitary or gregarious, carbonaceous, semi-immersed to erumpent, black, hemispherical or
globose to oval, multi-loculate, locules arrange in two layers in the outer layer of the ascostromata, one ascus in
each locule, ostiolate. Ostioles minute. Paraphyses absent. Cells of the ascostromata comprising thick-walled
textura angularis. Asci 17–47 × 15–3 μm ( = 33.0 × 24 μm, n = 12), 8-spored, bitunicate, fissitunicate, sub-
globose to globose, apedicellate, apically rounded with indistinct ocular chamber. Ascospores 15–29 × 4–8 μm (
= 19 × 7 μm, n = 34), hyaline, oblong to oval, 1–3-transverse septa, longitudinal septa absent, constricted at the
septa, smooth-walled, lacking a sheath. Asexual state: Unknown.
Material examined:—SWEDEN. Österlövsta parish, Skyttskär, on the resinous warts of young Betula
pendula (= B. verrucosa, Betulaceae), K. & L. Holm, 1981 (S, F30030!, isotype).
Micularia Boedijn, Persoonia 2(1): 67 (1961)
Parasitic on the upper surface of leaves forming darkened areas. Sexual state: Ascostromata superficial, scattered,
solitary or gregarious, coriaceous, semi-immersed, dark brown to black, globose to sub-globose, multi-loculate,
locules scattered throughout the ascostromata, containing a single ascus in each locule, ostiolate. Ostioles minute,
with dark brown hairs around apex. Paraphyses absent. Cells of ascostromata comprising dark brown cells of
textura angularis. Asci 8-spored, bitunicate, fissitunicate, sub-globose to pyriform, short pedicellate, apically
rounded with small ocular chamber. Ascospores hyaline, oblong to ellipsoid, 1-transverse septum, longitudinal
septa absent, constricted at the septum, smooth-walled to verruculose, lacking a sheath. Asexual state: Unknown.
Notes:—Micularia was introduced by Boedijn (1961) and remained monotypic until Peres & Bezerra (1981)
added Micularia tabebuiae to the genus. Micularia is characterized by having bundles of small spreading hairs at
the apex (Boedijn 1961). Boedijn (1961) placed this genus in the family Saccardiaceae, while Lumbsch &
Huhndorf (2007, 2010) placed Micularia in the family Elsinoaceae. This placement was followed by Hyde et al.
(2013). Even though it is a parasite on leaves, the inclusion of this genus in Elsinoaceae causes confusion, and as it
has only one ascus in each locule we suggest to place it in Myriangiaceae.
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FIGURE 7. Hemimyriangium betulae (isotype). a. Herbarium material. b–c. Ascostromata on host substrate. d. Cross section of
ascostroma. e–f. Peridium of locules. g. Immature and mature asci stained with cotton blue. h. Immature ascus. i. Immature ascus
stained with cotton blue. j–k. Mature asci. l–n. Mature asci stained with cotton blue. o–p. Smooth to verrculose walled ascospores. q–s.
Ascospore stained with cotton blue. Scale bars: b = 2 mm, c = 200 μm, d = 400 μm, e = 100 μm, f–g = 40 μm, h–s = 10 μm.
Type species:—Micularia merremiae Boedijn, Persoonia 2(1): 67 (1961) MycoBank: 334354 (Fig. 8)
Parasitic on the upper surface of leaves forming black spots. Sexual state: Ascostromata 0.5–0.65 × 0.63–0. 75 mm
( = 0.6 × 0.7 mm, n = 5), superficial, scattered, solitary or gregarious, coriaceous, immersed, dark brown to black,
globose to sub-globose, multi-loculate, locules generally scattered throughout the ascostromata, comprising a single
ascus in each locule, ostiolate. Ostioles minute; dark brown hairs frequently set around the blunt or acute apex of the
ascostromata, 20–40 × 3–6 µm ( = 28 × 4 µm, n = 10). Paraphyses absent. Cells of ascostromata comprising dark
brown cells of textura angularis. Asci 20–30 × 19–23 µm ( = 26 × 21 µm, n = 10), 8-spored, bitunicate,
fissitunicate, sub-globose to pyriform, short pedicellate, apically rounded with small ocular chamber. Ascospores
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11–15 × 5–7 µm ( = 13 × 6 µm, n = 20), hyaline, oblong to ellipsoid, 1-transverse septum, longitudinal septa
absent, constricted at the septum, smooth-walled or verruculose, lacking a sheath. Asexual state: Unknown.
Material examined:—INDONESIA. Java, Hortus Bogor, on leaves of Merremia sp. (Convolvulaceae), K.B.
Boedijn, 17 May 1954, (L, 0793023!, holotype).
FIGURE 8. Micularia merremiae (holotype). a. Herbarium specimen. b–d. Superficial ascostromata on substrate. e. Section through
ascostroma. f. Dark brown hairs around apex of ascostroma. g. Cells of ascostroma. h–j. Immature and mature asci. k. Mature ascus
stained with cotton blue. l–p. Ascospores. Scale bars: b, c = 1 mm, d = 50 µm, e–f = 10 µm, g = 5 µm, h–p = 10 µm.
Zukaliopsis Hennings, Hedwigia 43: 367 (1904)
Parasitic on leaves. Sexual state: Ascostromata superficial, scattered, solitary, coriaceous to sub-carbonaceous,
semi-immersed to erumpent, black to brown, sub-globose, multi-loculate, locules arranged in the outer margin of
the ascostromata, one ascus in each locule, ostiolate. Ostioles minute. Paraphyses absent. Cells of the ascostromata
comprising dark brown pseudoparenchymatous textura angularis. Asci 8-spored, bitunicate, fissitunicate, ovoid,
short pedicellate, narrowly pointed at the base, rounded at the apex with indistinct ocular chamber. Ascospores
hyaline, oblong to cylindro-clavate, muriform, with 4–6-transverse septa, 0–1-longitudinal septum, slightly
constricted at the septa, smooth-walled to veruculose, lacking a sheath. Asexual state: Unknown.
Notes:—Zukaliopsis was introduced by Hennings (1904) with Z. amazonica as type species and placed in the
family Perisporiaceae. Höhnel (1909) proposed that Zukaliopsis is morphologically related to Molleriella. von Arx
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(1963) synonymized Zukaliopsis with Molleriella and suggested that this genus resembles a transition to
Saccardinula Speg. (1885) more than Molleriella. Sydow & Theissen (1917) placed this genus into the family
Myriangiaceae providing a detailed description. We observed that Zukaliopsis has one ascus in each locule and,
therefore, suggest placing it in the family Myriangiaceae. New collections and molecular analysis are needed to
clarify the familial position of this genus.
Type species:—Zukaliopsis amazonica Hennings, Hedwigia 43(6): 367 (1904) MycoBank No: 213216 (Fig. 9)
Parasitic on leaves of Sapindaceae. Sexual state: Ascostromata 0.3–0.4 × 2–3 mm ( = 0.38 × 2.3 mm, n = 10),
superficial, scattered, solitary or aggregated, coriaceous to sub-carbonaceous, semi-immersed to erumpent, black to
brownish grey, ovoid to sub-globose, multi-loculate, numerous locules distributed throughout the upper part of the
ascostromata, one ascus in each locule, ostiolate. Ostioles minute. Paraphyses not observed. Cells of the
ascostromata comprising dark brown pseudoparenchymatous textura angularis. Asci 7–8 × 6–7 μm ( = 7.5 × 7
μm, n = 20), 8-spored, bitunicate, fissitunicate, ovoid to clavate, short pedicellate, narrowly pointed at the base,
apically rounded with indistinct ocular chamber. Ascospores 5–6 × 1–2 μm ( = 6 × 2 μm, n = 40), hyaline, oblong
to cylindro-clavate, muriform, with 4–6-transverse septa, 0–1-longitudinal septum, slightly constricted at the septa,
smooth-walled to verruculose, lacking a sheath. Asexual state: Unknown.
Material examined:—PERU. Iquitos, Rio Amazon, on Paullinia sp. (Sapindaceae), 1902, (S, F5604!,
isotype).
FIGURE 9. Zukaliopsis amazonica (isotype). a–b. Herbarium specimen. c–d. Ascostromata on natural substrate. e–f. Cross section of
ascostromata. g. Ascus in water. h–k. Asci in cotton blue. l–n. Ascospores in water. o. Ascospore in cotton blue. p. Ascospores in
Melzer ’s. Scale bars: c = 1 mm, d = 200 µm, e–f = 50 µm, g–p = 10 µm.
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Discussion
Starbäck (1899) introduced the order Myriangiales for the species characterized by crustose ascostromata and
muriform ascospores, similar to the type species Myriangium duriaei (Miller 1938). Phylogenetic studies based on
multigene analysis have indicated that Myriangiales always clusters with Dothideomycetes (Schoch et al. 2009,
Zhang et al. 2011). Kirk et al. (2008) included three families, Cookellaceae, Elsinoaceae and Myriangiaceae in
Myriangiales, while Lumbsch & Huhndorf (2010) accepted only Elsinoaceae and Myriangiaceae in the order based
on phylogenetic results. The order is characterized by pulvinate, irregular ascostromata in which the asci are
irregularly arranged in one or more layers in locules. Locules have single or multiple asci within each locule. Asci
are globose, with minute pedicels and indistinct ocular chambers. Ascospores which are irregularly arranged are
liberated only by the breakup of the stromatal layers above them. Asexual states are unknown.
Elsinoaceae is a family of plant pathogens that cause anthracnose and scab. Many classical treatments have
placed Elsinoaceae in synonymy with Myriangiaceae. However, Höhnel (1909) was convinced that the former
constituted a separate family. Eriksson (1981) agreed and maintained two separate families with Myriangiaceae
predominantly saprobic on branches and Elsinoaceae restricted to foliar pathogens. In Elsinoaceae the
pseudoascostromata form whitish, crust-like patches on leaves with a small number of asci in each locule. Asci are
globose and mostly 4–5 asci are embedded in one locule. Species of Schizothyriaceae Höhn. have similar
characters in the shape of the asci to those in Myriangiaceae, but differ as asci form in a single hymenial layer and
not in locules scattered in an ascostromata or pseudostromata (Hyde et al. 2013).
In this paper the generic types of the existing Myriangiaceae members, Anhellia tristis, Diplotheca tunae,
Eurytheca trinitensis and Myriangium duriaei, were re-examined and their placement in the family confirmed. The
generic types of Ascostratum insigne, Butleria inaghatahani, Dictyocyclus hydrangea, Hemimyriangium betulae,
Micularia merremiae and Zukaliopsis amazonica were also examined and we recommend placement in the family
Myriangiaceae because locules with single asci form in a well-developed dark ascostromata.
The position of Anhellia was uncertain within Myriangiaceae (Lumbsch & Huhndorf 2007, Krik et al. 2008).
Pinho et al. (2012) analysed nucleotide sequences of ribosomal large subunit (LSU) and ITS region gene data from
Anhellia nectandrae Pinho & Pereira (2012) showed it to cluster in Myriangiaceae. This and other genera that lack
sequence data need recollection so that molecular studies can confirm and resolve their relationships in
Myriangiales.
Acknowledgements
We thank the curators of S, L, PC, IMI and BPI herbaria for loaning specimens. Asha J. Dissanayake thanks MFLU
grant No. 56101020032, BRG5580009 and BAAFS for funding support and Samantha C. Karunarathna,
Dhanushka Udayanga, Sajeewa N. Maharachchikumbura, Nalin Wijayawardena, Hiran Ariyawansha and Tharanga
Rajakaruna for their help. 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.
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