Studies on Parmulariaceae I. A phylogeny based on available sequence data; introducing Parmulariales ord. nov., and Hemigraphaceae, Melaspileellaceae and Stictographaceae fam. nov.

Abstract

The family Parmulariaceae comprises three polyphyletic genera, but with very little data in GenBank and is presently placed in the order Asterinales. In this study, we re-analyze the available sequence data for taxa of the family and re-examine the type species of Hemigrapha, Inocyclus and Parmularia. The phylogenetic tree generated from maximum likelihood and Bayesian analyses of combined LSU-SSU sequence data demonstrate the relationships among Hemigrapha, Inocyclus and Parmularia species, and the relations of Buelliella, Karschia, Labrocarpon, Lembosia, Melaspileella, Melaspileopsis and Stictographa. We introduce Parmulariales ord. nov. to accommodate Parmulariaceae and the order Asterinales accommodates Asterinaceae, Asterotexaceae, Hemigraphaceae fam. nov., Melaspileellaceae fam. nov. and Stictographaceae fam. nov. Notes for each new order and families are provided. We confirm that Asterinaceae sensu lato is distant from Asterinaceae sensu stricto in the phylogenic analysis. The classification presented here is provisional, as more species are needed to re-collected and sequenced. We expect further support for our ordinal and familial lineages, as well as further novel lineages.

Full text

Phytotaxa 369 (2): 063–079
http://www.mapress.com/j/pt/
Copyright © 2018 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Sinang Hongsanan: 7 Aug. 2018; published: 13 Sept. 2018
https://doi.org/10.11646/phytotaxa.369.2.1
63
Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0
Studies on Parmulariaceae I. A phylogeny based on available sequence data;
introducing Parmulariales ord. nov., and Hemigraphaceae, Melaspileellaceae and
Stictographaceae fam. nov.
DONG-QIN DAI1, LI-ZHOU TANG1,2,*, CHAO LIU1, HAI-BO WANG1 & KEVIN D. HYDE3
1Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food Engineering, Qu-
jing Normal University, Qujing, Yunnan 655011, People’s Republic of China
2State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yun-
nan 650223, People’s Republic of China
3Centre of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
Corresponding author: tanglizhou@163.com
Abstract
The family Parmulariaceae comprises three polyphyletic genera, but with very little data in GenBank and is presently placed
in the order Asterinales. In this study, we re-analyze the available sequence data for taxa of the family and re-examine the
type species of Hemigrapha, Inocyclus and Parmularia. The phylogenetic tree generated from maximum likelihood and
Bayesian analyses of combined LSU-SSU sequence data demonstrate the relationships among Hemigrapha, Inocyclus and
Parmularia species, and the relations of Buelliella, Karschia, Labrocarpon, Lembosia, Melaspileella, Melaspileopsis and
Stictographa. We introduce Parmulariales ord. nov. to accommodate Parmulariaceae and the order Asterinales accommodates
Asterinaceae, Asterotexaceae, Hemigraphaceae fam. nov., Melaspileellaceae fam. nov. and Stictographaceae fam. nov.
Notes for each new order and families are provided. We confirm that Asterinaceae sensu lato is distant from Asterinaceae
sensu stricto in the phylogenic analysis. The classification presented here is provisional, as more species are needed to re-
collected and sequenced. We expect further support for our ordinal and familial lineages, as well as further novel lineages.
Keywords: 4 new taxa, taxonomy, phylogeny, biotrophs, lichenicolous fungi
Introduction
The family Parmulariaceae comprises more than 30 genera with extremely varied characters and is polyphyletic (Gua-
timosim et al. 2015). Genera such as Aldona, Aldonata, Dictyocyclus and Hemigrapha are not typical of the family
when compared with the type (Inácio & Cannon 2008, Tian et al. 2015). There are few studies relating to the phylog-
eny of Parmulariaceae (Guatimosim et al. 2015, Hyde et al. 2016, Liu et al. 2017), as genera of this family are uncul-
turable, therefore it is hard to obtain their DNA sequence data. However, sequence data is available for Hemigrapha,
Inocyclus and Parmularia with DNA extracted from fresh specimens. The family is presently included in Asterinales
(Hyde et al. 2013, Guatimosim et al. 2015), although this order affiliation needs verification.
Asterinales is an important order, commonly known as black mildews, forming dark, superficial colonies on liv-
ing leaves (Hosagoudar et al. 2013a). This order is characterized by orbicular, dark, flattened thyriothecia with central
dehiscent X- or Y- shaped openings. The thyriothecia of Asterinales usually contain few asci which are globose to oval,
or clavate. Ascospores are commonly ellipsoid, conglobate and initially hyaline, becoming brown to dark brown when
mature (Hosagoudar et al. 2013b, Hongsanan et al. 2014).
The order Asterinales previously included three families, viz. Asterinaceae, Aulographaceae and Parmulariaceae
(Hyde et al. 2013). However, although somewhat similar, characters of these families provide few clues as to the natu-
ral placement of the genera within these families. Based on both phylogeny and morphology, Hongsanan et al. (2014)
revised the order and accepted only a single family Asterinaceae. Subsequently, Ertz & Diederich (2015) sequenced
several lichenicolous genera (Buelliella, Hemigrapha, Karschia, Labrocarpon and Stictographa), as well as two sap-
robic genera (Melaspileella and Melaspileopsis) and placed these genera in Asterinales genera incertae sedis on the

DAI ET AL.
64 • Phytotaxa 369 (2) © 2018 Magnolia Press
basis of their phylogenic relationships close to taxa of Asterinales. Guatimosim et al. (2015) re-collected Parmularia
styracis Lév., the type species of Parmulariaceae and designated it as an epitype. The authors placed Parmulariaceae
in Asterinales, as the phylogenetic tree generated from Bayesian analysis of LSU sequence data, showed the Parmu-
laria strains clustered with several new collections of Asterinales sequenced by Guatimosim et al. (2015), viz. Asterina
melastomatis Lév., A. crysophylli, Prillieuxina baccharidincola (Rehm) Petr. and Lembosia abaxialis Firmino & R.W.
Barreto. These taxa, however were treated as Asterinaceae sensu lato by Liu et al. (2017), as they separated from a
separate set of Asterina species provided by different research groups. Asterotexis was previously placed in Asteri-
naceae (Inácio & Cannon 2008, Guerrero et al. 2011) and Guatimosim et al. (2015) had introduced a new family As-
terotexaceae and a new order Asterotexales to accommodate this genus. Subsequently, Liu et al. (2017) synonymized
Asterotexales under Asterinales, as strains of Asterotexaceae clustered with species of Asterinaceae sensu stricto.
Members of Asterinales are presently confused, with several genera incertae sedis groups. In this study, the
morphology of Hemigrapha, Inocyclus and Parmularia are re-examined based on herbarium specimens. The family
placements of the lichenicolous genera Buelliella, Hemigrapha, Karschia, Labrocarpon and Stictographa, as well as
the saprobic genera Melaspileella and Melaspileopsis are partially resolved based on phylogenetic analysis. The place-
ments of these genera are discussed and Parmulariales ord. nov., Hemigraphaceae, Melaspileellaceae and Stictogra-
phaceae families nov. are introduced.
Material and methods
Examination of specimens
Specimens (including types) of type species of Hemigrapha, Inocyclus and Parmularia were obtained from G, FH and
S (abbreviations according to Index Herbariorum 2018). The study methods followed are those reported in Dai et al.
(2014, 2017). Fruiting bodies were observed and photographed by Zeiss Stereo Discovery V8 with AxioCam ERc 5
s. Hand sections of the ascomata/ascostromata were mounted in distilled water on glass slides for microscopic studies
and photomicrography. The morphologic characters were observed by differential interference contrast (DIC) using a
Nikon ECLIPSE 80i compound microscope and photographed by Canon 600D digital camera fitted to the microscope
(Dai et al. 2017). Ascomata, asci and ascospores etc. were measured using a Tarosoft (R) Image Frame Work program.
Images used for figures were processed with Adobe Photoshop CS3 (Adobe Systems Inc., The United States). Index
Fungorum (2018) numbers are provided for newly taxa and Facesoffungi numbers (Jayasiri et al. 2015) are provided
for each taxa as well.
Phylogenetic analysis
DNA sequence data (LSU and SSU) of relevant taxa determined in Hyde et al. (2013), Guatimosim et al. (2015),
Ertz & Diederich (2015) and Liu et al. (2017) were download from GenBank. Selected taxa belonging to Asterinales,
Botryosphaeriales, Capnodiales, Cladoriellales, Hysteriales, Jahnulales, Lichenoconiales, Lichenotheliales,
Microthyriales, Myriangiales, Mytilinidiales, Natipusillales, Patellariales, Phaeotrichales, Tubeufiales, Valsariales
and Venturiales were included in the phylogenetic analysis to show the relationships among Buelliella, Hemigrapha,
Inocyclus, Karschia, Labrocarpon, Melaspileella, Melaspileopsis, Parmularia and Stictographa. Sequences were
aligned in MAFFT v. 7.215 (Katoh & Standley 2013) and manually edited in BioEdit (Hall 1999).
Maximum-likelihood (ML) analysis was performed in raxmlGUI v.1.0. (Stamatakis 2006, Silvestro & Michalak
2011) with 1000 bootstrap replicates. The PHYLIP format of combined alignment was exchanged and loaded from
website http://sing.ei.uvigo.es/ALTER/. The best-fit nucleotide substitution model (GTR+G) for sequence data was
selected by the online tool Findmodel (http://www.hiv.lanl.gov/content/sequence/findmodel/findmodel.html).
Bayesian analyses were running using MrBayes v. 3.0b4 (Ronquist & Huelsenbeck 2003). The best-fit model
of evolution was obtained by using MrModeltest v. 2.3 (Nylander 2004). Posterior probabilities (PP) (Rannala &
Yang 1996, Zhaxybayeva & Gogarten 2002) were determined by Markov Chain Monte Carlo sampling (MCMC)
in MrBayes v. 3.0b4. Six simultaneous Markov chains were run for 7,000,000 generations and trees were sampled
every 100th generation (Dai et al. 2017). The burn-in was set to 0.25, and the run was automatically stopped when the
average standard deviation of split frequencies reached below 0.01 (Maharachchikumbura et al. 2015).
Trees were viewed in TreeView (Page 1996), and additionally layouts were made with Adobe Illustrator CS v. 5.
Maximum-likelihood bootstrap values (MLBP) greater than 50 % and Bayesian posterior probabilities (BYPP) greater
than 0.80 are given. The sequences used in this study are listed in table 1. The combined alignment and phylogenetic
tree were submitted at TreeBASE (http://purl.org/phylo/treebase/phylows/study/TB2:S22032).

STUDIES ON PARMULARIACEAE Phytotaxa 369 (2) © 2018 Magnolia Press • 65
TABLE 1. Sequences data with GenBank accession numbers used in this study are provided.
Organism Strain GenBank accession numbers
LSU SSU
Aliquandostipite khaoyaiensis SS3321 EF175650 EF175628
Apiosporina collinsii CBS 118973 GU301798 GU296135
Asterina cestricola TH 591 GU586215 GU586209
Asterina crysophylli VIC 42823 KP143738
Asterina cynometrae MFLU 13-0373 KX845436
Asterina fuchsiae TH 590 GU586216 GU586210
Asterina melastomatis VIC 42822 KP143739
Asterina phenacis TH 589 GU586217 GU586211
Asterina siphocampyli ppMP 1324 HQ701140
Asterina weinmanniae TH 592 GU586218 GU586212
Asterina zanthoxyli TH 561 GU586219 GU586213
Asterina cynometrae MFLU 13-0373 KX845436
Asterotexis cucurbitacearum PMA M-0141224 HQ610510
Asterotexis cucurbitacearum VIC 42814 KP143734
Aulographum hederae CPC 21373 KM386981
Aulographum hederae MFLUCC 13-0001 KM386980
Bambusaria bambusae MFLUCC 12-0851 KP687812 KP687962
Batistinula gallesiae VIC 42514 KP143736
Botryosphaeria fusispora MFLUCC 10-0098 NG_042722 JX646823
Buelliella physciicola Ertz 19173 KP456148
Buelliella physciicola Ertz 18113 KP456147
Buelliella poetschii Ertz 18115 (BR) KP456149
Buelliella poetschii Ertz 18116 (BR) KP456150
Caliciopsis pinea AFTOL-ID 1869 DQ678097 DQ678043
Capnodium coffeae CBS 147.52 NG_027576 DQ247808
Cladoriella eucalypti CPC 10953 DQ195790 DQ195801
Cladoriella rubrigena CBS 124760 GQ303304
Dissoconium aciculare CBS 204.89 GU214419 GU214523
Dothiorella thailandica MFLUCC 11-0438 JX646813 JX646829
Glonium circumserpens CBS 123342 FJ161208 FJ161168
Glonium circumserpens CBS 123343 FJ161200 FJ161160
Hemigrapha atlantica Ertz 14014 (BR) KP456151
Hysterium angustatum MFLU 16-1179 KX611364 KX611365
Inocyclus angularis VIC 39747 KP143731
Inocyclus angularis VIC 39748 KP143732
Inocyclus angularis VIC 39749 KP143733
Jahnula aquatica R68.1 EF175655 EF175633
Jahnula bipileata F49.1 EF175657 EF175635
Karschia cezannei Ertz 19186 KP456154
Karschia talcophila Ertz 16749 KP456155
Labrocarpon canariense Ertz 16907 KP456158
Labrocarpon canariense Ertz 16308 KP456157
......continued on the next page

DAI ET AL.
66 • Phytotaxa 369 (2) © 2018 Magnolia Press
TABLE 1 (Continued)
Organism Strain GenBank accession numbers
LSU SSU
Lembosia albersii MFLU 13-0377 KM386982
Lembosia xyliae MFLU14-0004 KT283685
Lempholemma polyanthes 11294-L1(2/2) AF356691
Leptoxyphium fumago CBS 123.26 GU214430 GU214535
Lichenoconium aeruginosum JL359-09 HQ174269
Lichenoconium lecanorae JL382-10 HQ174263
Lichenothelia convexa L1608 KC015070 KC015085
Lichenothelia convexa L1609 KC015071 KC015086
Melaspileella proximella G.M. 2015-04-29 KY654708
Melaspileella proximella G.M. 2014-05-25 KY654747
Melaspileopsis cf. diplasiospora Ertz 16247 KP456164
Melaspileopsis cf. diplasiospora Ertz 16624 KP456165
Mendogia macrostroma MFLU 13-0642 KU863104 KU872109
Microthyrium microscopicum CBS 115976 GU301846 GU296175
Microthyrium propagulensis IFRD 9037 KU948989
Myriangium duriaei CBS260.36 NG_027579 NG_013129
Myriangium hispanicum CBS 247.33 KX887067 GU371744
Myrmaecium rubrum CBS 109505 GU456324 GU456303
Natipusilla decorospora L_A236_1A HM196369 HM196376
Natipusilla naponensis L_AF217_1B HM196372 HM196379
Neodeightonia microspora MFLUCC 11-0483 KU863099 KU872105
Parmularia styracis VIC 42447 KP143728
Parmularia styracis VIC 42450 KP143729
Parmularia styracis VIC 42587 KP143730
Patellaria cf. atrata BCC 28876 GU371828 GU371836
Patellaria cf. atrata BCC 28877 GU371829 GU371837
Peltula auriculata AFTOL-ID 892 DQ832330
Phaeotrichum benjaminii CBS 541.72 AY004340 AY538349
Prillieuxina baccharidincola VIC 42817 KP143735
Psiloglonium simulans CBS 206.34 FJ161178 FJ161139
Rhytidhysteron rufulum MFLUCC 14-0577 KU377565 KU377570
Stictographa lentiginosa Ertz 17570 KP456170
Stictographa lentiginosa Ertz 17447 KP456169
Teratosphaeria fibrillosa CBS 121707 KF902075 GU296199
Trichodelitschia bisporula CBS 262.69 GU348996 GU296202
Tubeufia filiformis MFLUCC 16-1135 KY092411
Tubeufia javanica MFLUCC 12–0545 KJ880036 KJ880035
Valsaria insitiva CBS 123098 GU460204 GU456310
Venturia inaequalis ATCC 60070 EF114712 EF114737
Venturia populina CBS 256.38 GU323212 GU296206

STUDIES ON PARMULARIACEAE Phytotaxa 369 (2) © 2018 Magnolia Press • 67
Results and discussion
Phylogeny
A data set of combined LSU-SSU sequence data of 82 strains, including the outgroup taxon Caliciopsis pinea (AFTOL-
ID 1869), was used to determine the relationship among Hemigrapha, Inocyclus and Parmularia. The lichenicolous
Buelliella, Karschia, Labrocarpon and Stictographa, as well as saprobic Melaspileella and Melaspileopsis genera
are placed in Asterinales. The phylogenetic tree generated from maximum likelihood (ML) and Bayesian analyses of
LSU-SSU sequence data demonstrate that Parmulariaceae forms a sister clade with Asterinaceae sensu lato with high
bootstrap support (MLBS/BYPP 100/1.00) and forms a separate lineage from Asterinales (Fig. 1). The phylogenetic
analysis (Fig. 1) based on the representative sequences of the major orders in Dothideomycetes support Parmulariaceae
as an individual entity. A new order Parmulariales is therefore established to accommodate this family.
Guatimosim et al. (2015) mentioned that Parmulariaceae is polyphyletic, perhaps because the authors considered
the case that Inocyclus was phylogenetically far from Parmulariaceae. However, on the basis of re-examination of
isotype of the type species Inocyclus psychotriae (Syd. & P. Syd.) Theiss. & Syd., Inocyclus is not morphologically
and representatively similar with Parmulariaceae.
Asterinales forms a strongly supported clade (MLBS/BYPP 98/1.00) in the data matrix. Two major lineages can
be defined within Asterinales. One that contains taxa of Stictographaceae, with the genera Karschia, Labrocarpon
and Melaspileopsis as well as Buelliella physciicola Poelt & Hafellner. Currently, Buelliella is not included in
Stictographaceae, until sequences from the type species are available. The relationships among these genera are poorly
supported, probably because of lack of protein genes, and this result is similar to that of Ertz & Diederich (2015).
Other groups of Asterinales are Asterinaceae sensu stricto, Asterotexaceae, Hemigraphaceae and Melaspileellaceae,
together with several genera incertae sedis. The clade of Asterinaceae sensu stricto is stable and strongly supported
(MLBS/BYPP 99/1.00, Fig. 1). Strains of type species of Melaspileellaceae form a branch at the base of Asterinales.
Hemigrapha was previously placed in Parmulariaceae and then removed to Asterinales incertae sedis by Ertz &
Diederich (2015). It forms a single branch within Asterinales and therefore Hemigraphaceae is introduced here.
Inocyclus, a genus also previously assigned to Parmulariaceae forms a sister clade with two Lembosia species (L.
xyliae X. Y. Zeng et al. and L. albersii Henn.). The type species of Inocyclus has not been sequenced, therefore
Inocyclus is retained in Asterinales incertae sedis. Lembosia was previously included in Asterinaceae (Hongsanan et
al. 2014) based on the sequences obtained from a fresh specimen L. albersii, however few strains were included in
the phylogenic tree in Hongsanan et al. (2014). According to our study, two Lembosia species are phylogenetically
separate from Asterinaceae (Fig. 1). Lembosia needs further collections with molecular data of type species to establish
its relationship with Asterinaceae.
Taxonomy
Asterinales M.E. Barr ex D. Hawksw. & O.E. Erikss., Syst. Ascom. 5(1): 177 (1986)
Synonyms: Asterotexiales Firmino, O.L. Pereira & Crous [as ‘Asterotexiales’], Persoonia 35: 238 (2015)
Hemigraphaceae D.Q. Dai & K.D. Hyde, fam. nov.
Index Fungorum number: IF554062; Facesoffungi number: FoF 03910
Biotrophic on thallus of lichens. Sexual morph: Ascostromata solitary to gregarious, in groups, superficial, black to
dark brown, coriaceous, stellate, irregularly opening from the centre to margin, conical in section. Peridium composed
of two parts; outer part, a black layer, composed of thick-walled cells of textura angularis; inner part thin, composed
of light brown cells of textura angularis. Hamathecium composed of a few brown, unbranched, filamentous, septate,
pseudoparaphyses around asci. Asci 8-spored, bitunicate, clavate to cylindric-clavate, subglobose, with an ocular
chamber and a short pedicel. Ascospores 3-seriate to irregularly arranged, brown, ellipsoid, 1-septate, with larger upper
cell and narrower lower cell, smooth-walled. Asexual morph: Undetermined.
Type genus: Hemigrapha (Müll. Arg.) R. Sant. ex D. Hawksw., Kew Bull. 30(1): 9 (1975)

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68 • Phytotaxa 369 (2) © 2018 Magnolia Press
FIGURE 1. Maximum likelihood phylogenetic tree generated by RAxML (GTR+G model) based on analysis of combined LSU and SSU
sequence data. ML values (>50 %) resulting from 1000 bootstrap replicates and Bayesian posterior probabilities greater than 0.80 are given
at the nodes. The strain numbers are noted after the species names. The tree is rooted to Dendrographa decolorans (DUKE 0047570).

STUDIES ON PARMULARIACEAE Phytotaxa 369 (2) © 2018 Magnolia Press • 69
Notes:—The genus Hemigrapha, typified by H. asteriscus (Müll. Arg.) R. Sant. ex D. Hawksw (Basionym:
Melanographa asteriscus Müll. Arg.), was introduced by Hawksworth (1975). This genus is a communalistic taxon
having a biotrophic life-style (Diederich & Wedin 2000, Ertz & Diederich 2015). Inácio & Cannon (2008) revised
the genera of Parmulariaceae but did not include Hemigrapha in the family. However, Lumbsch & Huhndorf (2010),
Hyde et al. (2013) and Wijayawardene et al. (2014) included this genus in Parmulariaceae. The lichenicolous fungi,
Hemigrapha is similar to Viegasella (Tian et al. 2015) in having star-shaped ascomata, but differs in its smooth-
walled ascospores (Viegasella has verrucose ascospores). Hemigrapha is characterized by dark, superficial, star-
shaped ascomata forming on a thallus of lichens, clavate to subglobose asci and brown, ellipsoid, 2-celled ascospores
(Diederich & Wedin 2000). Based on the morphology of the re-examined type in this paper, Hemigrapha is not
similar with the family type of Parmulariaceae. Ertz & Diederich (2015) included H. atlantica in their phylogenetic
analysis and indicated that this genus belongs to order Asterinales based on LSU sequence data, without assigning the
family placement. In this study, Hemigrapha atlantica Diederich & Wedin forms a single clade clustering within the
order Asterinales (Fig. 1). Hemigrapha atlantica phylogenetically forms as a sister species with the generic type H.
asteriscus, seen in Ertz & Diederich (2015). Therein a new family Hemigraphaceae is introduced to accommodate this
single genus in Asterinales.
Type species: Hemigrapha asteriscus, Figure 2.
FIGURE 2. Hemigrapha asteriscus (G 00292584, syntype). a. Herbarium material. b–d. Appearance of ascostromata on host surface. c,
d. Ascostromata in the water. e–j. Section of ascostroma. h. Asci with hamathecial tissues. i–m. Asci. n, o. Ascospores. Scale bars: a = 20
mm, b–d = 500 µm, e–g = 50 µm, h–o = 5 µm.

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70 • Phytotaxa 369 (2) © 2018 Magnolia Press
Facesoffungi number: FoF 02312
Basionym: Melanographa asteriscus Müll. Arg., Flora, Regensburg 65(33): 519 (1882)
Syntype:—G 00292584
Biotrophic on lichens. Sexual morph: Ascostromata 1–1.5 mm diam., solitary to gregarious, in groups, superficial,
black to dark brown, coriaceous, stellate, irregularly opening from the centre to margin, conical in section, 50–115
µm diam., 30–55 µm high. Peridium 7–10 µm thick, composed of two layers; outer part, black, composed of thick-
walled cells of textura angularis; inner part thin, composed of light brown cells of textura angularis. Hamathecium
composed of few brown, unbranched, 1–1.5 µm wide, filamentous, septate, pseudoparaphyses around asci. Asci 19–30
× 8–15 µm ( = 24.4 × 10.9 µm, n = 20), 8-spored, bitunicate, clavate to cylindric-clavate, subglobose, with an ocular
chamber and a short pedicel. Ascospores 8.5–12.5 × 3–4 µm ( = 10.1 × 3.7 µm, n = 20), 3-seriate to irregularly
arranged, brown, ellipsoid, 1-septate, with large upper cell and narrower lower cell, smooth-walled. Asexual morph:
Undetermined.
Material examined:—AUSTRALIA, New South Wales, Mount Kosciuszko, on thallus of Peltigera dolichorrhiza
(Lichen). Müller J. 1882 (G 00292584, syntype).
Melaspileellaceae D.Q. Dai & K.D. Hyde, fam. nov.
Index Fungorum number: IF554063; Facesoffungi number: FoF 03911
Saprobic on trees and shrubs. Sexual morph: Ascostromata solitary, superficial, dark to black, coriaceous, rounded.
Peridium outer layers composed of black, thick-walled cells; inner layers composed of light brown to hyaline cells
of textura angularis. Hamathecium of dense, hyaline, unbranched, filamentous, septate pseudoparaphyses around
asci, brown at the apices. Asci 8-spored, bitunicate, wide clavate to subglobose, with a round apex and a short
pedicel. Ascospores 2-seriate to irregularly arranged, hyaline, ellipsoid, 1-septate, with slightly larger upper cell, with
slightly narrower lower cell, smooth-walled, bearing 2–3 appendages which disappear when dry. Asexual morph:
Undetermined.
Type genus: Melaspileella (P. Karst.) Vain.
Notes:—Melaspileella was introduced by Vainio (1921) and was placed in Melaspileaceae within the order
Arthoniales (Index Fungorum 2018). Ertz & Diederich (2015) lectotypified the genus with Melaspileella proximella
(Nyl.) and provided SSU sequence data. LSU sequence data was earlier submitted in GenBank for its synonym
Banhegyia setispora L. Zeller & Tóth. Based on the phylogenetic analysis, Ertz & Diederich (2015) transferred
Melaspileella to Asterinales without assigning it to a family. In our study, the generic type Melaspileella proximella
forms a single clade close to Hemigrapha in Asterinales with high bootstrap support (MLBS/BYPP 97/1.00) (Fig.
1). Melaspileella is morphologically similar to Karschia in having rounded and dark ascomata, but is distinct in the
phylogenic analysis (Fig. 1). Herein, Melaspileellaceae is introduced to accommodate a single genus Melaspileella.
Type species: Melaspileella proximella (Nyl.) Ertz & Diederich
Stictographaceae D.Q. Dai & K.D. Hyde, fam. nov.
Index Fungorum number: IF554064; Facesoffungi number: FoF 03912
Lichenicolous on the thallus of lichen. Sexual morph: Ascomata solitary, scattered to clustered, superficial, cymbiform
to lirelliform, or rounded, a slit-like disc, black to dark brown, with a centre ostiole opening by a slit or star-shaped
fissure. Peridium thick, outer layers composed of black to dark brown, thick-walled cells of textura angularis; inner
layers composed of light brown to hyaline cells of textura angularis. Hamathecium of dense, hyaline, filamentous,
septate pseudoparaphyses, unbranched or occasionally branched at upper part around the asci. Asci 8-spored, bitunicate,
wide clavate to subglobose, with an ocular chamber, apically rounded and short pedicellate. Ascospores 2-seriate to
irregularly arranged, hyaline, becoming light brown to dark brown, ellipsoid, 1-septate, with slightly larger upper cell,
with slightly narrower lower cell, smooth-walled. Asexual morph: Undetermined.
Type genus: Stictographa Mudd, Man. Brit. Lich.: 226 (1861)
Notes:—Stictographaceae is introduced here to accommodate several lichenicolous genera or with saprobic life-
cycles, and is characterized by cymbiform to lirelliform, or a slit-like disc, black to dark brown ascomata, wide clavate
to subglobose asci and brown, ellipsoid ascospores with a single septum. Stictographa was collected on the exclusive
host Phaeographis dendritica Muell in western Europe. It is morphologically similar to Labrocarpon Etayo & Pérez-
Ortega in having cymbiform ascomata opening by a long slit. However, they can be distinguished in the phylogenetic
analysis (Fig. 1). The monotypic genus Labrocarpon was introduced by Pérez-Ortega & Etayo (2010) and is typified

STUDIES ON PARMULARIACEAE Phytotaxa 369 (2) © 2018 Magnolia Press • 71
by L. canariense (D. Hawksw.). In the phylogenetic tree (Fig 1), the type species of Labrocarpon clusters within
Stictographaceae. Karschia Körb. was described by Körber (1865) and is typified by K. talcophila (Ach.) Körb. Barr
(1979) placed Karschia in Patellariaceae Corda. Then it was transferred to Lichenotheliaceae, Lichenotheliales by
Thambugala et al. (2014a) based on the morphology observed from herbarium material. Ertz & Diederich (2015) re-
examined the type specimen and sequenced the type species. According to the phylogenetic analysis of Ertz & Diederich
(2015), Karschia forms a single clade close to Labrocarpon, Melaspileopsis and Stictographa. Our phylogenetic tree
(Fig. 1) shows the same result and Karschia is phylogenetically distant from Lichenotheliales. Thus, Karschia is placed
in Stictographaceae. Melaspileopsis was introduced by Ertz & Diederich (2015) and is typified by M. diplasiospora
(Nyl.) Ertz & Diederich. Index Fungorum (2018) lists the current name of the type species is Melaspilea diplasiospora
(Nyl.) Müll. Arg. However, the phylogenic analysis shows it does not belong to Melaspileaceae Walt. Watson, but
clusters close to the clade of Stictographa (Fig. 1). Melaspileopsis is therefore placed in Stictographaceae.
Type species: Stictographa lentiginosa (Lyell ex Leight.) Mudd, Man. Brit. Lich.: 226 (1861)
Other genera included:
Karschia Körb., Parerga lichenol. (Breslau) 5: 459 (1865)
Type species: Karschia talcophila (Ach.) Körb., Parerga lichenol. (Breslau) 5: 460 (1865)
Labrocarpon Etayo & Pérez-Ortega, in Pérez-Ortega & Etayo, Lichenologist 42(3): 271 (2010)
Type species: Labrocarpon canariense (D. Hawksw.) Etayo & Pérez-Ortega [as ‘canariensis’], in Pérez-Ortega & Etayo, Lichenologist
42(3): 272 (2010)
Melaspileopsis (Müll. Arg.) Ertz & Diederich, Fungal Diversity 71: 161 (2015)
Type species: Melaspileopsis diplasiospora (Nyl.) Ertz & Diederich, Fungal Diversity 71: 161 (2015)
Asterinales genera incertae sedis
Inocyclus Theiss. & Syd., Annls mycol. 13(3/4): 211 (1915)
Facesoffungi number: FoF 02315
Parasitic on upper surface of living leaves. Sexual morph: Ascostromata solitary, scattered or gregarious, superficial,
circular or ellipsoidal, flat, black, opening by radiating fissures to irregular slits, internal part of stromata thick and
containing a single locule, with a thin light brown tissue under the loculate layer, reaching leaf surface, composed
of dark brown cells of textura angularis; upper wall of ascostromata comprising dark brown tissue of cells of
textura prismatica. Locules immersed under the upper wall of stromata. Hamathecium composed of hyaline, septate,
filamentous pseudoparaphyses, extending above the asci. Asci 8-spored, bitunicate, subglobose, obovoid, clavate-
cylindrical, with a short pedicellate, thick-walled, with rounded apex, smooth, with a large apical chamber. Ascospores
irregularly arranged, initially hyaline and becoming pale brown to dark brown, ellipsoidal, apiospores, 1-septate, with
larger upper cell and shorter lower cell, thin-walled, smooth-walled. Asexual morph: See notes.
Notes:—Inocyclus was introduced by Theissen & Sydow (1915) for taxa forming black ascostromata opening
by radiating fissures to irregular slits on living leaves. It includes eight accepted species, of which two species (I.
discoideusis (Racib.) Arx and I. angularis Guatim. & R.W. Barreto) were reported on ferns (Polypodiaceae J. Presl &
C. Presl) (Guatimosim et al. 2014a). Inocyclus angularis has conidiomata intermixed with the ascomata, occupying the
same stromata located in the central region of the colonies and hyaline, aseptate, fusiform to clavate conidia reported
by Guatimosim et al. (2014a). However, the asexual morph of generic type is undetermined. Guatimosim et al. (2015)
included I. angularis in the phylogenetic analysis based on LSU sequences data. It is indicated that I. angularis is
phylogenetically separate from Parmulariaceae but close to species of Asterotexis, thus Guatimosim et al. (2015)
placed Inocyclus in Asterotexales incertae sedis. However, Asterotexales was synonymized under Asterinales by Liu
et al. (2017). Based on the phylogenic analysis in this study (Fig. 1), Inocyclus clusters within Asterinales and is close
to several Lembosia species. However, the type species of Inocyclus has not been sequenced yet, so we place Inocyclus
in Asterinales genera incertae sedis.
Type species: Inocyclus psychotriae (Syd. & P. Syd.) Theiss. & Syd., Annls mycol. 13(3/4): 211 (1915),
Figure 3, 4
Facesoffungi number: FoF 02316
Lectotype:—K 180637

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72 • Phytotaxa 369 (2) © 2018 Magnolia Press
FIGURE 3. Inocyclus psychotriae (K 180637, lectotype). a, b. Herbarium material. c, d. Ascostromata on leaves. E. Squash mount of
dark brown to black ascostromata. f, g. Hand section of ascostroma (Note: the peridium without apical cells). h–l. Ascospores immersed
in KOH. m–q. Asci with ascospores immersed in KOH. Scale bars: c =500 µm, d = 200 µm, e = 100 µm, f = 50 µm, g = 20 µm, h–l = 5
µm, m–q = 10 µm.
Parasitic on upper surface of living leaves. Sexual morph: Ascostromata 0.3–0.7 mm diam., 100–150 µm high,
solitary, scattered or gregarious, superficial, circular or ellipsoidal, flat, black, opening by radiating fissures to irregular
slits, internal part of stromata thick and containing a single locule, with a thin light brown tissue under the loculate layer,
reaching leaf surface, composed of dark brown cells of textura angularis; upper wall of ascostromata comprising 15–20
µm thick, dark brown tissue of cells of textura prismatica. Locules 140–160 diam., 100–125 µm high, immersed under
the upper wall of stromata. Hamathecium composed of hyaline, septate, 2–3 µm wide, filamentous pseudoparaphyses,
extending above the asci. Asci 23–38 × 8–10.5 µm ( = 29.7 × 9.8 µm, n = 20), 8-spored, bitunicate, subglobose,
obovoid, clavate-cylindrical, with rounded apex, with a short pedicellate, thick and smooth-walled, with a large apical
chamber. Ascospores 8–11.5 × 3–4 ( = 9.3 × 3.8 µm, n = 20), irregularly arranged, initially hyaline and becoming
pale brown to dark brown, ellipsoidal, apiosporous, 1-septate, with larger upper cell and shorter lower cell, thin-walled,
smooth-walled. Asexual morph: Undetermined.

STUDIES ON PARMULARIACEAE Phytotaxa 369 (2) © 2018 Magnolia Press • 73
FIGURE 4. Inocyclus psychotriae (NY 01102760, isolectotype). a. Herbarium material. b, c. Ascostromata on leaves. D. Squash mount
of dark brown to black ascostroma. e, f. Hand section of ascostroma (Note: the peridium without apical cells). g–h. Asci with ascospores
immersed in water. i. Asci with ascospores immersed in KOH. j–l. ascospores. Scale bars: b = 1 mm, c = 100 µm, d–f = 20 µm, g–l = 5
µm.
Material examined:—PHILIPPINES, Luzon, Antipolo, Rizal, on living leaves of Psychotria luzoniensis (Cham.
& Schltdl.) Fern.-Vill., 11 October 1912, M. Ramos, (K 180637, lectotype).
Other specimen examined:—PHILIPPINES, Luzon, Antipolo, Rizal, on living leaves of Psychotria luzoniensis
(Cham. & Schltdl.) Fern.-Vill., M. Ramos and E.D. Merrill (NY 01102760, isolectotype).
Parmulariales D.Q. Dai & K.D. Hyde, ord. nov.
Index Fungorum number: IF554065; Facesoffungi number: FoF 03913
Epiphytic or parasitic on living leaves of plants and ferns or lichenicolous. Colonies superficial, dark brown to black,
rounded to irregular. Hyphae usually spreading out from ascomata border. Sexual morph: Ascostromata solitary,
scattered, gregarious, or in groups, superficial to immersed, shield-like, elliptical, dark brown to black, carbonaceous
to membranaceous, strongly flattened. Locules immersed in ascostromata or in longitudinal ridges radiating from
the centre. Peridium composed of dark to hyaline, thick to thin cells of textura prismatica to textura angularis.

DAI ET AL.
74 • Phytotaxa 369 (2) © 2018 Magnolia Press
Hamathecium of hyaline, septate pseudoparaphyses with brown apices, or pseudoparaphyses lacking. Asci 8-spored,
bitunicate, broad cylindrical to subglobose, with an ocular chamber. Ascospores 2-seriate to irregular arranged, hyaline
to dark brown, oblong to ellipsoid, 1-septate, smooth- to verrucose-walled. Asexual morph: Undetermined.
Notes:—Parmulariaceae has had a varied history in terms of its higher-level placement (Inácio & Cannon 2008),
with placements in the orders Dothiorales, Microthyriales, Hemisphaeriales, Dothideales and Hysteriales (Müller &
von Arx 1962, Ainsworth 1971, Luttrell 1973, von Arx & Müller 1975, Barr 1979, Hawksworth et al. 1983, 1995).
The family has also been placed in Dothideomycetes family incertae sedis, as DNA sequence data was not available
(Eriksson 1999, Kirk et al. 2001). Kirk et al. (2008) followed this placement. Inácio & Cannon (2008) reviewed
the family with a detailed study, however they did not assign the family Parmtoulariaceae to any order. Lumbsch
& Huhndorf (2010) maintained Parmulariaceae in Dothideomycetes family incertae sedis. Hyde et al. (2013) re-
examined the types of bitunicate families and placed Parmulariaceae in the order Asterinales based on morphological
similarities with Asterinaceae. Guatimosim et al. (2015) sequenced the type species of type genus of Parmulariaceae
and placed this family in order Asterinales, as Parmulariaceae was phylogenetically close to several Asterina species.
However, Liu et al. (2017) placed above mentioned Asterina taxa in Asterinaceae sensu lato and their lineages are
separate from Asterinales sensu stricto. Thus, the ordinal placement of Parmulariaceae is undetermined. The family
Parmulariaceae is morphologically and phylogenetically distinct from families of Asterinales and justifies a separate
order in Dothideomycetes (Fig. 1). Thus, Parmulariales is recommended here to accommodate a single family
Parmulariaceae.
Type family: Parmulariaceae E. Müll. & Arx ex M.E. Barr, Mycologia 71(5): 944 (1979)
Facesoffungi number: FoF 02296
Epiphytic or parasitic on living leaves of plants and ferns or thallus of lichens. Colonies superficial, dark brown to
black, rounded to irregular. Hyphae spreading out from ascomata, brown, septate, branching, sinuous, usually lacking
appressoria. Sexual morph: Ascostromata solitary, scattered, or gregarious, superficial to immersed, shield-like,
elliptical to boat-shaped, dark brown to black, carbonaceous to membranaceous, strongly flattened, or longitudinal slits
slightly protuberant. Locules immersed in ascostromata or in longitudinal ridges radiating from the centre. Stromata
wall thin to thick, composed of black, large cells of textura prismatica. Wall of locules thin to thick, composed of dark
brown to hyaline smaller cells of textura angularis. Hamathecium of dark brown to hyaline, septate, pseudoparaphyses
or pseudoparaphyses lacking. Asci 8-spored, bitunicate, long ovate, broad cylindrical, broad-clavate, or obclavate,
globose to subglobose, short pedicellate, with a distinct ocular chamber, thick-walled. Ascospores 2–multi-seriate,
hyaline to dark brown, oblong to ellipsoid, 1-septate, smooth- to verrucose-walled. Asexual morph: Undetermined.
Notes:—The family Parmulariaceae was invalidly introduced by Müller & von Arx (1962) to accommodate
species with colonies forming on the surface of living plant tissues, mostly leaves, and was later validated by Barr
(1979). Parmulariaceae contains a range of ascomata types, asci and ascospores forms (Inácio & Cannon 2008), as
numerous genera were placed in this family. Lumbsch et al. (2007) included 34 genera in Parmulariaceae and Inácio
& Cannon (2008) included 35 genera. Kirk et al. (2008) included 34 genera with 24 synonyms as well as 119 species.
Inácio et al. (2012) introduced a new genus Antoniomyces Inácio, according to its similarity of morphology with
Parmulariaceae. Hyde et al. (2013) accepted 31 genera and excluded four genera viz. Coccodothis Theiss. & Syd.,
Dothidasteroma Höhn., Englerodothis Theiss. & Syd. and Perischizon Syd. & P. Syd. from this family. Guatimosim et
al. (2014b) introduced a new genus Rhagadolobiopsis Guatim. & R.W. Barreto based on morphology. Thambugala et
al. (2014b) re-examined the herbarium of type species of Placostromella Petr. and assign it in Parmulariaceae.
The family Parmulariaceae can be compared with Asterinaceae, however, it differs in having ascostromata
produce multi-locules which forming longitudinal ridges radiating from the centre surrounded by thick wall, and
formed a developed base, pseudoparaphyses, ellipsoidal to obclavate asci, and usually lack appressoria (Inácio &
Cannon 2008, Inácio 2012, Hyde et al. 2013). Species of Asterinaceae have superficial hyphae mostly with appressoria,
small thyriothecia with a thin upper wall layer and poorly developed base, subglobose asci, lack pseudoparaphyses,
(Pirozynski & Shoemaker 1970, Inácio 2012, Hongsanan et al. 2014). Parmulariaceae can also be compared with
Englerulaceae whose species are usually parasitic on living leaves. Englerulaceae however it differs in having small,
superficial, globose and deliquescent ascomata on mycelium (Dai et al. 2014).
Type genus: Parmularia Lév., Annls Sci. Nat., Bot., sér. 3 5: 286 (1846)
Facesoffungi number: FoF 02297
Epiphytic on upper surface of leaves. Colonies foliicolous, superficial, dark brown to black, circular. Hyphae
growing at base of ascomata, brown, septate, branching, sinuous, lacking appressoria. Sexual morph: Ascostromata
large, solitary to groups, gregarious, superficial, shield-like, dark brown to black, carbonaceous, flattened, with ridges
radiating from the centre to the outer rim; ridges containing elongated locules, which open by a longitudinal slit and

STUDIES ON PARMULARIACEAE Phytotaxa 369 (2) © 2018 Magnolia Press • 75
containing numerous asci; cells of ascostromata wall thick-walled and elongated and composed of black, amorphous
tissues in the stromatic crust. Locules in vertical section subglobose, immersed within ascostroma. Upper wall of
ascostromata thick, comprised of several black layers, composed of dark brown cells of textura prismatica. Cells
between locules composed of brown to hyaline cells of texture angularis. Hamathecium of hyaline, septate, filamentous
pseudoparaphyses, swollen, brown and verrucose at their apices. Asci 8-spored, thick-walled, bitunicate, cylindrical,
short-pedicellate, with a distinct ocular chamber. Ascospores 2–3-seriate, hyaline and becoming pale brown when
mature, ellipsoidal, 1-septate, with large and round upper cells, with narrow and long lower cells, constricted at the
septum, smooth-walled. Asexual morph: Undetermined.
Notes:—Parmularia is typified by P. styracis Lév., introduced by Léveillé (1846). Parmularia is similar to Mintera
in having elongate locules radiating from a central point. However, Mintera has mycelium with appressoria, and each
black ascostroma with a single locule which aggregate into star-shaped structures. In Parmularia, a single ascostroma
contains many locules. Parmularia comprises numerous species with 53 names in Index Fungorum (2018) and Kirk et
al. (2008) estimated there are 30 species. However, Inácio & Cannon (2008) compared only six Parmularia species, all
from Brazil and only one species from New Zealand. Parmularia is characterized by large, black, shield-like, flattened
ascostromata with ridges radiating from the centre to the outer rim, and the ridges comprising elongated, radiating
locules, filamentous pseudoparaphyses with brown and verrucose apices and ellipsoid, 1-septate ascospores (Inácio &
Cannon 2008, Hyde et al. 2013).
Type species: Parmularia styracis Lév., Annls Sci. Nat., Bot., sér. 3 5: 286 (1846). Figure 5
FIGURE 5. Parmularia styracis (S F21306). A. Herbarium material. B. Appearance of ascostromata on upper of leaf surface. c, e, f. Black
shield-like ascostroma. d. Dark brown ascostroma wall. G. Sections of ascostroma showing multi-locules. H. Wall cells of ascostroma. i,
j. Asci with ascospores. K. Pseudoparaphyses with brown swollen apices. l–n. Immature hyaline ascospores. Scale bars: b =10 mm, c =1
mm, d = 5 µm, e, f = 500 µm, g=100 µm, h = 50 µm, i–k = 10 µm, l–n = 10 µm.

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76 • Phytotaxa 369 (2) © 2018 Magnolia Press
Facesoffungi number: FoF 02298
Epiphytic on upper surface of leaves. Colonies foliicolous, superficial, dark brown to black, round. Hyphae
growing out from ascomata, brown, septate, branching, sinuous, lacking appressoria. Sexual morph: Ascostromata 1–
3 mm diam., solitary to gregarious, or in groups, superficial, shield-like, dark brown to black, carbonaceous, flattened,
with ridges radiating from the centre to outer rim; ridges containing elongated locules, which open by a longitudinal
slit and contain numerous asci, cells of ascostromata thick-walled and elongate and composed of black, amorphous
tissues in the stromatic crust. Locules in vertical section: 150–250 µm diam., 80–120 µm high, subglobose, immersed
in ascostromata. Upper wall of ascostromata 20–40 µm thick, comprising several black layers, composed of dark
brown cells of textura prismatica. Cells between locules composed of brown to hyaline cells of texture angularis.
Hamathecium of hyaline, septate, filamentous pseudoparaphyses, 50–80 × 2–3.5 µm, swollen, brown and verrucose
at their apices. Asci 45–70 × 12–16 µm ( = 58.2 × 13.9 µm, n = 20), 8-spored, thick-walled, bitunicate, broadly
cylindrical, short-pedicellate, with a distinct ocular chamber. Ascospores 13.5–16.5 × 4–6 µm ( = 15.6 × 5.1 µm, n =
20), 2–3-seriate, hyaline becoming pale brown when mature, ellipsoidal, 1-septate, with larger and rounder upper cells,
with narrow and longer lower cells, constricted at the septum, smooth-walled. Asexual morph: Undetermined.
Material examined:—BRAZIL, Brasilia, São Paolo, near Urbem Faxina, on leaves of Styracis sp., August 1901,
v. Wettstein & Schiffner (S F21306).
Notes:—Parmularia styracis described from leaves of Styrax L., collected in Brazil (Léveillé 1846). Guatimosim
et al. (2015) recollected and epitypified this species and provided a phylogenetic analysis based on LSU sequence data.
Inácio & Cannon (2008) recorded the ascostromata of P. styracis as 0.3–1.6 mm in diam.; however, Guatimosim et al.
(2015) indicated the epitype having 1–3 mm diam. ascostromata. We also found the ascostromata to be 1–3 mm diam.
on the material we examined. The holotype of this species was collected in Brazil on the leaves of Styracis sp. and was
deposited in PC (Index Herbariorum 2018), however, a later specimen collected from same country on same host was
examined in this study.
Acknowledgments
This work was supported by the Key Laboratory of Yunnan Province Universities of the Diversity and Ecological
Adaptive Evolution for Animals and plants on Yun-Gui Plateau, the National Natural Science Foundation of China
(No. NSFC 31760013, NSFC 31260087, NSFC 31460561, NSFC 31460179) and the Scientific Research Foundation
of Yunnan Provincial Department of Education (2017ZZX186). Dong-Qin Dai and Li-Zhou Tang would like to thank
Yunnan Province Universities of the Science and Technology Innovation Team for the exploitation and utilization of
endophytes for finance support. Dong-Qin Dai would like to thank the Thousand Talents Plan, Youth Project of Yunnan
Provinces for support and Dr. Jiang-Kui Liu for the suggestions about phylogeny. Yunnan Local Colleges Applied
Basic Research Projects (No. 2017FH001-034) is acknowledged by Dr. Chao Liu for support.
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