Mycologia: Phaeochorellaceae/Diaporthales: A New Fungal Family and A Re-Appraisal of Phaeochorella Species

Mycologia
Phaeochorellaceae/Diaporthales: a new fungal family and a re-appraisal of

Phaeochorella species
--Manuscript Draft--
Manuscript Number: mycologia-D-18-00198R1
Article Type: Original Research
Manuscript Classifications: Morphology/Development; Plant Pathogens; Systematics
Full Title: Phaeochorellaceae/Diaporthales: a new fungal family and a re-appraisal of

Phaeochorella species
Corresponding Author: JOSE CARMINE DIANESE, Ph.D.

Universidade de Brasilia Instituto de Ciencias Biologicas
BRASÍLIA, DISTRITO FEDERAL BRAZIL
Other Authors: Debora Cervieri Guterres, Dr. Sci.
Samuel Galvão-Elias, MSc
Maria do Desterro Mendes dos Santos, Dr. Sci.
Bruno Cézar Pereira de Souza, MSc
Camila Pereira Almeida, BSc
Danilo Batista Pinho, Dr. Sci.
Robert Neil Gerard Miller, PhD
Abstract: Species of Phaeochorella are biotrophic leaf parasites with tropical distribution,
traditionally accepted in the family Phyllachoraceae, Phyllachorales, when considered
based on morphological features. Molecular evidence shown here resolves the
relationship of the genus Phaeochorella within the Sordariomycetes, based on a
multilocus analysis of partial nuclear sequences of the nuclear rDNA (nuclear large
subunit 28S and nuclear internal transcribed spacer ITS), the DNA-directed RNA
polymerase II second largest subunit (RPB2), and the 1-α translation elongation factor
(TEF1 α) gene. Phylogenetic analyses indicate that Phaeochorella belongs in the
Diaporthales rather than the Phyllachorales. Phaeochorella parinarii, the type species
of the genus, present on native hosts from the Brazilian Cerrado, formed a unique
clade with a species of Phaeoappendicospora with high support. Thus, a new family,
Phaeochorellaceae, Diaporthales including both genera, is herein proposed. With the
exception of P. parinarii and Phaeochorella zonata, all other species in Phaeochorella
(P. artocarpi, P. ciliata, P. machaerii) were excluded from the genus. Phylogenetic
analyses indicated that Phaeochorella belongs in the Diaporthales rather than the
Phyllachorales. Phaeochorella parinarii, the type species of the genus, also present on
native hosts from the Brazilian Cerrado, formed a unique clade with a
Phaeoappendicospora species with high support. Thus, a new family,
Phaeochorellaceae, including both genera, is herein proposed. With the exception of
P. zonata, the other species previously considered as belonging to Phaeochorella (P.
artocarpi, P. ciliata, P. machaerii) were excluded from the genus after appropriate
taxonomic treatment.
Keywords: Cerrado; Diaporthaceae; Fungal taxonomy; multilocus phylogeny; Neotropical

microfungi; Phyllachora-like species; Phyllachoraceae; 1 new epitypification; 1 new
taxon
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1 Short title: Phaeochorellaceae, new tropical family of the Diaporthales

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2 2
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5 3 Title: Phaeochorellaceae, Diaporthales: a new fungal family and a re-appraisal of
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7 4 Phaeochorella species.
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10 5
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12 6 Debora Cervieri Guterresa, Samuel Galvão-Eliasb, Maria do Desterro Mendes dos Santosc,
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14 7 Bruno Cézar Pereira de Souzaa, Camila Pereira de Almeidad, Danilo Batista Pinhoa,
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17 8 Robert Neil Gerard Millera,b, and José Carmine Dianesea,b
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19 9
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21 a
22 10 Departamento de Fitopatologia, Universidade de Brasília, 70910-900, Brasília, Distrito
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24 11 Federal, Brazil; bDepartamento de Biologia Celular, Universidade de Brasília, 70910-900,
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27 12 Brasília, Distrito Federal, Brazil; cDepartamento de Ecologia, Universidade de Brasília,
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29 13 70910-900, Brasília, Distrito Federal, Brazil, dDepartamento de Farmácia, Universidade
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14 de Brasília, 70910-900, Brasília, Distrito Federal, Brazil
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34 15 Corresponding author: jcarmine@gmail.com
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36 16
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39 17 ABSTRACT
40
41 18 Species of Phaeochorella are biotrophic leaf parasites with tropical distribution,
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43
44 19 traditionally accepted in the family Phyllachoraceae, Phyllachorales, when considered
45
46 20 based on morphological features. Molecular evidence shown here resolves the relationship
47
48
49
21 of the genus Phaeochorella within the Sordariomycetes, based on a multilocus analysis of
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51 22 partial nuclear sequences of the nuclear rDNA (nuclear large subunit 28S and nuclear
52
53 23 internal transcribed spacer ITS), the DNA-directed RNA polymerase II second largest
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55
56 24 subunit (RPB2), and the 1-α translation elongation factor (TEF1 α) gene. Phylogenetic
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58 25 analyses indicate that Phaeochorella belongs in the Diaporthales rather than the
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61 26 Phyllachorales. Phaeochorella parinarii, the type species of the genus, present on native
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64 Corresponding author: jcarmine@gmail.com
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27 hosts from the Brazilian Cerrado, formed a unique clade with a species of
1
2 28 Phaeoappendicospora with high support. Thus, a new family, Phaeochorellaceae,
3
4
5 29 Diaporthales including both genera, is herein proposed. With the exception of P. parinarii
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7 30 and Phaeochorella zonata, all other species in Phaeochorella (P. artocarpi, P. ciliata, P.
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31 machaerii) were excluded from the genus.
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12 32 KEY WORDS: Cerrado; Diaporthaceae; fungal taxonomy; multilocus phylogeny;
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14 33 Neotropical microfungi; Phyllachora-like species; Phyllachoraceae; 1 new epitypification;
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17 34 1 new taxon
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19 35
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21
22 36 INTRODUCTION
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24 37 Phaeochorella (Henn.) Theiss. & Syd. is a genus of biotrophic leaf parasites traditionally
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27 38 included in the family Phyllachoraceae, Phyllachorales, with five species all known from
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29 39 the tropics: P. parinarii (Henn.) Theiss. & Syd., P. zonata Petr., and Phaeochorella
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31
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40 artocarpi T.S. Ramakr. & K. Ramakr, P. ciliata Bat. & J.L. Bezerra, P. machaerii Bat. &
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34 41 Peres; (Theissen and Sydow 1915; Petrak 1947; Ramakrishnan and Ramakrishnan 1948;
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36 42 Batista and Peres 1960; Batista and Bezerra 1961).
37
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39 43 Phaeochorella parinarii was originally described by Hennings (1901) as Cocconia
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41 44 parinarii Henn., on leaves of the tree species Parinari curatellifolia (Oliv.) R. Grah. (=Pa.
42
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44 45 mobola Oliv.), a South African member of the pantropical family Chrysobalanaceae.
45
46 46 Based on C. parinarii, the genus Phaeochorella Theissen & Sydow was established
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48
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47 (Theissen and Sydow 1915) in the family Phyllachoraceae, subfamily Scirrhinieae, tribe
50
51 48 Catacaumineae. The genus Phaeochora was considered the closest genus to
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53 49 Phaeochorella as it also has brown 1-celled ascospores but differs by forming spores in
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56 50 aparaphysate asci. In addition to the type species, Theissen and Sydow (1915) included
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58 51 Phaeochorella. clypeata (G. Winter) Theiss. & Syd. (=Auerswaldia clypeata G. Winter).
59
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61 52 Their diagnosis related Phaeochorella to Catacauma, with a stroma considered to be
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3
53 phyllachoroid, but with brown 1-celled spores and paraphysate asci, indicating that, as in
1
2 54 Catacauma species, fungal development occurred between the cuticle/epidermis and the
3
4
5 55 palisade layer of the host leaf. Doidge (1942, 1950) reported Phaeochorella parinarii as
6
7 56 common in Parinari species in South Africa, and segregated it from Phyllachora based on
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9
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57 the brown ascospores. The author indicated that among the genera then included in the
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12 58 Phyllachoraceae with pigmented ascospores, Phaeochorella showed smaller ascospores
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14 59 with a paler coloration than those of Phaeochora, differing from Sphaeorodothis with
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17 60 globose ascospores not oblong to elliptical as in Phaeochorella.
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19 61 Petrak (1947) added the species Phaeochorella zonata Petrak, amended the genus
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21
22 62 description; described the asexual state of P. parinarii, namely Phomachorella parinarii
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24 63 Petrak, on Pa. capensis, and finally, focusing on the morphology of the ascospores,
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27 64 excluded P. clypeata from the genus.
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29 65 Later, three new species were described on hosts other than members of the
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31
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66 Chrysobalanaceae. Phaeochorella artocarpii was described on leaves of Artocarpus
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34 67 lacucha (Moraceae) by Ramakrishnan and Ramakrishnan (1948) in India; P. machaerii on
35
36 68 Machaerium scleroxylon (Fabaceae) by Batista and Peres (1960), and P. ciliata on
37
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39 69 Jaracatia dodecaphylla (Caricaceae) by Batista and Bezerra (1961); the latter two from
40
41 70 Brazil. Phaeochorella sphaerospora Chardón, on leaves of Machaerium angustifolium in
42
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44 71 Colombia (Chardón and Toro 1930; Dennis 1970), was later accepted as a synonym of
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46 72 Phyllachora conica (Chardón) Petr. by Cannon (1991), who considered the coloration of
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48
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73 the ascospores as lacking taxonomic value to differentiate the two species, although he did
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51 74 not examine Chardón and Toro (1930)’s holotype, then presumably lost.
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53 75 The type species, Phaeochorella parinarii, based on Cocconia parinari Henn., was
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56 76 described on the subshrub host (Parinari capensis), and later on a tree [Pa. curatellifolia
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58 77 (=Pa. mobola)] in South Africa (Hennings 1901). The species was also present in Angola,
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61 78 Namibia, Republic of Congo, Zambia, and Zimbabwe (Doidge 1950; Arx and Müller
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79 1954; Whiteside 1966, Parker 1978), with reports on other Parinari species from
1
2 80 Malaysia (Wiehe 1953; Peregrine and Siddiqi 1972). In Brazil, P. parinarii is known on
3
4
5 81 another subshrub (Pa. obtusifolia Hook. f.), as illustrated by Medeiros (1994) and Souza
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7 82 (2017), and presented in an unpublished index of Cerrado fungi incorporated in Mendes et
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83 al. (1998). The genus Parinari Aubl. is monophyletic, originated in Africa and more
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12 84 diversified in the Neotropica (Bardon et al. 2013), but following Gondwanan breakup it
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14 85 became a pantropical genus reaching Southern Africa, Neotropica from Brazil to Central
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17 86 America with 39 accepted species names as shown in Plant List (2019)
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19 87 (http://www.theplantlist.org/browse/A/Chrysobalanaceae/Parinari, retrieved on January
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22 88 14, 2019). Besides widespread presence of Parinari species, in Africa, South and Central
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24 89 America, in Australasia occurred 816 records mostly from Queensland, Australia
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27 90 represented by one species, Pa. nonda F.Muell. ex Benth. Parinari species are also
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29 91 present in Fiji, Indonesia, Malaysia, Papua New Guinea, Samoa, Solomon Islands,
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92 Thailand, and Tokelau Island (Atlas of Living Australia 2019)
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34 93 (https://bie.ala.org.au/species/http://id.biodiversity.org.au/node/apni/2918944, accessed on
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36 94 January 14, 2019). In addition, for Southeast Asia, Pa. costata (Korth.) Blume is present
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39 95 in Myanmar and The Philippines, according to Kew´s World Checklist of Selected Plant
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41 96 Families (https://wcsp.science.kew.org/namedetail.do?name_id=355493, retrieved on
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44 97 January 14, 2019).
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46 98 There has been no record of Phaeochorella for the entire Australasia, and only one
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99 unidentified Lasmeniella species has been associated with Pa. nonda in a single site in
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51 100 Australia (Hyde & Alcorn 1993). This fungus may be morphologically characterized as an
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53 101 asexual diaporthaceous ascomycete, showing 1-celled dark brown conidia, produced by
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56 102 conidiogenous cells inserted in black stromatic conidioma. Although appearing in a recent
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58 103 review of the dematiaceous coelomycetes (Wijayawardene et al. 2016), the genus
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104 Lasmeniella was not phylogenetically evaluated, as sequence data was unavailable, such
1
2 105 that its taxonomic placement remains uncertain.
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5 106 In Brazil, the genus Parinari is most diverse within the Atlantic and Amazonian rain
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7 107 forests, with mostly arboreal species (e.g. Pa. excelsa, Pa. alvimii, Pa. campestris, Pa.
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108 klugi, etc.), but in the Cerrado biome the dominant plant species is Pa. obtusifolia, the
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12 109 only known host of P. parinarii to date in Brazil (Rede speciesLink 2019).
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14 110 Currently, there is no molecular data available for Phaeochorella, with species defined by
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17 111 morphological characters. In this context, the present study aims to determine the
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19 112 phylogenetic position of P. parinarii, based on analysis of material collected from the
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22 113 Brazilian Cerrado on leaves of Pa. obtusifolia.
23
24 114 MATERIAL AND METHODS
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27 115 Morphological studies–. Macro-morphological examination of fungarium and fresh
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29 116 specimens was performed using a model 205C Leica stereomicroscope, followed by
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117 sectioning on a CM 1850 Leica freezing microtome. Resulting thin sections (20–30 µm)
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34 118 used for semi-permanent lactoglycerol mounts. Visualization, photography and
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36 119 measurements were performed on a DM 2500 Leica light microscope provided with
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39 120 Nomarski microscopy, and coupled to a DFC 490 Leica digital camera. Size estimations
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41 121 for all structural components were based on up to 30 measurements.
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44 122 DNA extraction, amplification and sequencing–. Genomic DNA extraction was performed
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46 123 from material taken from hydrated ascomata using a Wizard® Genomic DNA Purification
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49
124 Kit (Promega, Madison, WI, USA), following the manufacturer’s instructions. DNA
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51 125 quantification was performed by comparison to a Low DNA Mass Ladder (Invitrogen,
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53 126 CA, USA) following electrophoresis on 1% agarose gels. PCR amplifications were
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56 127 performed on a DNA Engine (PTC-200) Peltier Thermal Cycler (Bio-Rad Laboratories,
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58 128 CA, USA).
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129 Four partial nuclear gene sequences (two ribosomal and two protein-coding genes) were
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2 130 amplified and sequenced: the nuclear large subunit ribosomal DNA (nuc 28S rDNA), and
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5 131 the nuclear internal transcribed spacer (ITS1–5.8S–ITS2 rDNA), amplified with primers
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7 132 V9G (Hoog and van den Ende 1998), LR5 (Vilgalys and Hester 1990), and ITS3 (White
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9
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133 et al. 1990). The DNA-directed RNA polymerase II second largest subunit (RPB2) was
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12 134 amplified using primers 5F2 (Sung et al. 2007) and 7cR (Liu et al. 1999); and the
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14 135 translation elongation factor 1-alpha (TEF1) with primers EF1–938F and EF1–2218R
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17 136 (Rehner and Buckley 2005).
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19 137 PCR thermocycling was performed following DNA denaturation at 94 C for 1 min 30 s,
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22 138 with 35 cycles of DNA denaturation at 94 C for 30 s. Primer annealing was performed at
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24 139 53 C (18S, 28S and ITS), 56 C (TEF 1α) and 57 C (RPB2) for 30 s, extension at 72 C for
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27 140 45 s, with a final extension at 72 C for 5 min, followed by storage at 10 C. PCR products
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29 141 were checked on 1 % agarose gels, then treated with ExoSAP-IT® PCR Product Cleanup.
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31
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142 Sequencing was conducted at Macrogen Inc. USA (http://www.macrogen.com).
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34 143 Electropherograms were checked, with ambiguous positions clarified considering forward
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36 144 and reverse sequences. Newly generated sequences were assembled and annotated using
37
38
39 145 the software GENEIOUS 9.0.5 (Kearse et al. 2012), then deposited in GenBank
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41 146 (http://www.ncbi.nlm.nih.gov) (Table 1).
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44 147 Taxa sampling and alignment–. To approximate the phylogenetic position of P. parinarii,
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46 148 an exploratory analysis (Supplementary FIG. 1) was performed including a wide range of
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48
49
149 members belonging in the Sordariomycetes. Based on these preliminary results our taxa
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51 150 sampling could be later restricted to Diaporthales. The dataset recently used by
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53 151 Senanayake et al. (2017) was partially repeated here to construct a three gene matrix (28S,
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55
56 152 RPB2, and TEF 1α) with one hundred taxa, including our newly generated sequences of
57
58 153 P. parinarii and outgroups.
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61
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154 Nucleotide sequences of ribosomal markers were aligned using E–INS–i strategy (built in
1
2 155 MAFFT 7.305 (Katoh and Standley 2013) and manually checked in ALIVIEW (Larsson
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5 156 2014). RPB2 and TEF1α coding sequences were paired with the software Multiple
6
7 157 Alignment of Coding Sequences (MASCE) (Ranwez et al. 2011). Codon position
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158 assignment was confirmed by translating nucleotide sequences into predicted amino acids,
11
12 159 checking for stop-codon absence, and comparison with annotated reference sequences of
13
14 160 Neurospora crassa AF107789, Podospora anserina AF107790, and Chaetomium elatum
15
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17 161 AF107791 for RPB2 (Liu et al. 1999), and Trichoderma reesei Z23012 for TEF1(Nakari
18
19 162 et al 1993). GenBank accession numbers for all sequences are shown in Table 1. To best
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22 163 contemplate the full variation of coding sequences, both nucleic and amino acids were
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24 164 addressed in phylogenetic analysis through a mixed scheme of MrBayes as in Kim et al.
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27 165 (2003) as an example of complex partitioned models available in
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29 166 http://mrbayes.sourceforge.net/wiki/index.php/Kim.nex. As the implementation of mixed
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167 schemes are not available for Maximum Likelihood optimization in RAXML, only nucleic
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34 168 acids were used in our analysis. In both phylogenetic reconstructions, matrices of RPB2
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36 169 and TEF1 were partitioned in first, second and third codon positions.
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39 170 Phylogenetic analysis–. Maximum Likelihood (ML) analysis using RAXML 8.2.9
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41 171 (Stamatakis 2014) started with a randomized stepwise addition parsimony tree under a
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44 172 GTR+GAMMA model. Branch support values were calculated based in 1000 bootstrap
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46 173 replications (BS) under the same model. Bayesian Inference (BI) was performed using
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174 MRBAYES 3.2.6 (Ronquist and Huelsenbeck 2003), with appropriate evolutive models
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51 175 calculated to each partition in JMODELTEST 2 under Bayesian information criteria (Darriba
52
53 176 et al. 2012). The 4by4 and mixed mode were used for nucleic and amino acid partitions,
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56 177 respectively. Two independent chains were run, each one initiating from random trees and
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58 178 four simultaneous independent chains at 2×106 generations. Trees were sampled every
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61 179 1000th generation. Four rate categories were used to approximate the gamma distribution.
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180 Average standard deviation of split frequencies (ASDSF) and Effective Sample Size
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2 181 (ESS) were used as an acceptance criterion of Bayesian analyses. A total of 25% of all
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5 182 sampled trees were discarded as burn-in, while the remaining 75% were employed to
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7 183 estimate the Bayesian Posterior Probabilities (BPP) for branches. Both RAXML,
8
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184 JMODELTEST, and MRBAYES were conducted in CIPRES Science Gateway 3.1 (Miller et al.
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12 185 2010).
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14 186
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17 187 RESULTS
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19 188 Nucleotide matrix description─. The nucleotide matrix of Diaporthales and outgroups,
20
21
22 189 which included three partial genes (28S, RPB2, and TEF1), comprises 3183 aligned sites,
23
24 190 including gaps. Individual matrix length and the evolutionary models selected for each
25
26
27 191 locus are shown in Supplementary Table 1.
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29 192 Phylogenetic analyses─. Markov Chains of MrBayes Bayesian Inference converged at
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193 3,761,000 generations (~ 18 % of full chains, ASDSF 0.0099) and ESS greater than 200
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34 194 for all parameters. The results revealed that specimens of P. parinarii form a
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36 195 monophyletic lineage in BI and ML analyses (BPP = 1.00; BS = 100, respectively) within
37
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39 196 Diaporthales (BPP = 1.00; BS = 93), with Phaeoappendicospora thailandensis Senan.,
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41 197 Q.R. Li & K.D. Hyde as a sister lineage of P. parinarii, receiving high phylogenetic
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44 198 support (BPP = 1.00; BS = 97) (FIG. 1). Nucleotide matrices and phylogenetic trees are
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46 199 available in TreeBASE (study number 23429).
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49
200
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51 201 TAXONOMY
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53 202 Diaporthales was introduced by Nannfeldt (1932) to accommodate ascomycetes with
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56 203 ascomata immersed in stroma, with non-allantoid ascospores, sometimes producing short
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58 204 stalked unitunicate cylindrical to clavate asci with a well defined apical apparatus,
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61 205 sometimes floating freely in the ascomata. Asexual morph were described as
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206 coelomycetes generally stromatic, showing phialidic or annelidic conidiogenous cells.

1
2 207 Chadefaud (1960) accepted three families within the order (Diaporthaceae,
3
4
5 208 Melanconidaceae and Gnominiaceae), followed by Wehmeyer (1975). Barr (1978),
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7 209 Monod (1983) and Eriksson (2001) later modified the morphological circumscription of
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9
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210 the Diaporthales. With the incorporation of molecular information, the Diaporthales
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12 211 enlarged its range. Based on analysis of LSU nrDNA sequences Castlebury et al. (2002)
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14 212 accepted Diaporthaceae, Gnomoniaceae, Melanconidaceae and Cytosporaceae in
15
16
17 213 Diaporthales, and Gryzenhout et al. (2006) using sequences of ITS, β-tubulin1 and 2 and
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19 214 LSU, introduced in Diaporthales the Cryphonectria–Endothia complex as family
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22 215 Cryphonectriaceae. Rossman et al. (2007) accepted nine families within the order based
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24 216 on the available molecular and morphological data. Multilocus phylogeny was applied to
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27 217 specific taxa within Diaporthales, as Sogonov et al. (2008) who focused the family
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29 218 Gnominiaceae; Cheewangkoon et al. (2010) detailed the phylogeny of the genus
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31
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219 Cryptosporiopsis; Alvarez et al. (2016) revised the family Schizoparmaceae;
33
34 220 and Voglmayr and Jaklitsch (2014) resurrected the family Stilbosporaceae.
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36 221 In conclusion, using modern taxonomic approach based on morphology and multilocus
37
38
39 222 phylogeny the order was expanded (Walker et al. (2010, 2012), Mejía1 et al. (2011, Crous
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41 223 et al. 2012, Maharachchikumbura et al. 2015), and finally 14 families were confirmed in
42
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44 224 the Diaporthales (Senanayake et al. 2017, Guterres et al. 2018).
45
46 225 Thus, a diversity of genera, outside the classical circumscription of the order, were
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226 incorporated into Diaporthales, e.g. Apiosphaeria Hönh., previously accepted in
50
51 227 Phyllachoraceae, Phyllachorales (Guterres et al. 2018), Tirisporella E.B.G. Jones et al.,
52
53 228 morphologically fitted in the Dothideomycetes due to its bitunicate asci (Suetrong et al.
54
55
56 229 2015), and Phaeoappendicospora Senan. et al. (Senanayake et al. 2017), showing bi-
57
58 230 appendiculate ascospores and asci produced in black papillate ascoma. In this context, the
59
60
61 231 genus Phaeochorella (Henn.) Theiss. & Syd. a classical member of the Phyllachoraceae,
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10
232 Phyllachoralles is here considered for a re-evaluation through morphological and

1
2 233 multilocus phylogenetic analyses. When P. parinarii was phylogenetically coupled with
3
4
5 234 the recently described monotypic Phaeoappendicospora (type species Ph. thailandensis
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7 235 Senan. et al. (Senanayake et al. 2017), their phylogenetic proximity suggested the need for
8
9
10
236 the establishment of a new family herein described, with phylogeny shown in FIG. 1.
11
12 237 Even though Phaeoappendicospora is morphologically and ecologically distant from
13
14 238 Phaeochorella, comprising one saprobic species on dead stems of Quercus sp.
15
16
17 239 (Fagaceae), with only the sexual morph known. The type species, Ph. thailandensis, is
18
19 240 characterized by: “stromata forming a thin weft of pale brown hyphae around the upper
20
21
22 241 part of the perithecia; ascomata immersed, aggregated, sub-globose to globose,
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24 242 coriaceous, black, ostiolate, papillate; papilla black, cylindrical, lateral, periphysate;
25
26
27 243 periphyses hyaline, filamentous; peridium comprising few layers of black, thick-walled,
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29 244 cells of textura angularis; hamathecium comprising few, septate, hyaline paraphyses
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31
32
245 attached to the base, longer than asci; asci 8-spored, unitunicate, ellipsoid, with a short
33
34 246 pedicel, inconspicuous, flat, J-refractive ring at the lower end of the thickened apical wall,
35
36 247 apex narrow and blunted; ascospores ellipsoidal to ovoid, with broadly rounded ends, pale
37
38
39 248 brown, 1-septate, not constricted at the septa, thick and smooth-walled, guttulate, with
40
41 249 short, hyaline, rounded appendages at both ends; asexual morph undetermined”
42
43
44 250 (Senanayake et al. 2017).
45
46 251 Phaeochorella is presently comprised by only two species: P. parinarii and P. zonata,
47
48
49
252 that together with monotypic Phaeoappendicospora, as indicated by our molecular
50
51 253 phylogeny, constitute a new family within Diaporthales, described below.
52
53 254 Phaeochorellaceae Guterres, Galvão-Elias & Dianese fam. nov.
54
55
56 255 MYCOBANK MB825364
57
58 256 Type genus: Phaeochorella (Henn.) Theiss. & Syd. Annales mycologici 13: 405. 1915.
59
60
62
63
65
11
258 Saprotrophic on plant debris or biotrophic on living leaves. Pseudostromata dark brown to
1
2 259 black, subepidermal on living leaves or present on plant debris. Ascomata perithecial,
3
4
5 260 globose, subglobose, to pyriform, ostiolate; ostioles periphysate; periphyses filiform, thin,
6
7 261 numerous. Asci unitunicate, thin-walled, cylindrical, subclavate or elliptic, paraphysate;
8
9
10
262 paraphyses filiform. Ascospores elliptic to cylindrical, dark brown, 0 to 1-septate.
11
12 263 Phaeochorella (Henn.) Theiss. & Syd. Annales mycologici 13: 405. 1915.
13
15
16
17 265 Leaf parasites with shiny black, irregularly shaped immersed pseudostromata, dispersed
18
19 266 on adaxial face of leaves, especially along veins, occupying the subepidermal layer,
20
21
22 267 bearing immersed perithecial ascomata, on top of a basal layer of olive-brown
23
24 268 prosenchymatous tissue; wall textura prismatica; covered by a black, carbonaceous layer
25
26
27 269 as a subepidermal clypeus. Ascomata perithecial, pyriform, globose or subglobose, with a
28
29 270 short cylindrical to sub-papillate periphysate neck. Asci numerous, cylindrical,
30
31
32
271 cylindrical-cuneiform to subclavate, paraphysate. Paraphyses hyaline, septate, thin-
33
34 272 walled. Ascospores 1-celled, oval or elliptical, brown to dark brown with a sub-hyaline to
35
36 273 hyaline equatorial band.
37
38
39 274
40
41 275 Type: Phaeochorella parinarii (Henn.) Theiss. & Syd. Annales mycologici. 13: 405.
42
43
44 276 1915. FIGS. 2−4.
45
47
48
49
278 ≡Cocconia parinarii Henn., Botanische Jahrbücher für Systematik Pflanzengeschichte
50
51 279 und Pflanzengeographie 30:257. 1901.
52
54
55
56 281 =Phomachorella parinarii Petr., Sydowia 1:126. 1947 (name given to the asexual morph).
57
59
60
61
62
63
65
12
283 Pseudostromata 2−4 mm wide, epiphyllous, circular or irregular, solitary or gregarious,

1
2 284 black, shiny, subepidermal, clypeate, multiloculate. Ascomata 158–350 μm diam,
3
4
5 285 immersed in pseudostromata, globose to ovoid, ostiolate with a short periphysate neck.
6
7 286 Asci 75–96 × 10–13 μm, cylindrical to subclavate, rounded at apex, thin-walled, without
8
9
10
287 conspicuous apical structures, paraphysate. Paraphyses 1.5–4 μm thick, hyaline, septate,
11
12 288 simple. Ascospores 9–13 × 4.5–7.5 μm, 1-celled, uniseriate, seldom biseriate, elliptical,
13
14 289 with rounded tips, brown with a hyaline median band. Asexual morph often lacking,
15
16
17 290 pycnidial immersed in same pseudostroma as sexual stage. Conidiophores in basal layer
18
19 291 of conidioma, simple, surrounded by a mucous matrix. Conidiogenous cells phialidic.
20
21
22 292 Conidia 8 × 3.5 μm, 1-celled, oblong or elongate-ellipsoid, fusiform, hyaline, thin-walled.
23
24 293 Hosts: Parinari capensis, Pa. curatellifolia (=Pa. mobola), and Pa. obtusifolia.
25
26
27 294 Distribution: Angola, Brazil, Malaysia, Namibia, South Africa, Republic of Congo,
28
29 295 Zambia, Zimbabwe.
30
31
32
296 Specimens examined─. Cocconia parinarii Henn. ANGOLA, K'nebe, on leaves of Pa.
33
34 297 capensis Harv., Oct 23 1899, Hugo Baum, Herbarium CUP! F-03689(2-99) (Isotype).
35
36 298 Phaeochorella parinari (Henn.) Theiss. & Syd. NAMIBIA, Zwischein Klubangi und
37
38
39 299 Mondongo, on leaves of Pa. curatellifolia (=Pa. mobola), May 05 1900, Hugo Baum,
40
41 300 Herbarium B! 2065, Reliquiae Petrakianae, Intitute für Botanik-Graz. SOUTH AFRICA,
42
43
44 301 Pretoria, on leaves of Pa. capensis, Apr 29 1911, E.M. Doidge, Herbarium B! 1472,
45
46 302 Reliquiae Petrakianae, Intitute für Botanik-Graz. BRAZIL. FEDERAL DISTRICT.
47
48
49
303 Brasília: Asa Norte, University of Brasília, Campus Darcy Ribeiro, -15.772778 -
50
51 304 47.865556, on living leaves of Parinari obtusifolia, May 7 2015, DC Guterres 114, UB
52
53 305 Mycol. Coll. 23270 (MBT 383151: epitype here designated). GenBank Accessions: ITS
54
55
56 306 = MK050523; 28S = MK050523; RPB2 = MK050190; TEF1 = MK050188.
57
58 307 Ibid: Brasília, Embrapa CPAC, Reserva do Mirante, -15.6168194 -47.7190111, on living
59
60
61 308 leaves of P., June 25 2015, BCP Souza 28, UB Mycol. Coll.23397. GenBank: ITS =
62
63
65
13
309 MK050524; 28S = MK050524; RPB2 = MK050191; TEF1 = MK050189. Don Bosco
1
2 310 Ecological Park, -15.6168194 -47.7190111, on living leaves of Pa. obtusifolia, Oct 10
3
4
5 311 2016, DC Guterres 115, UB Mycol. Coll.23806.
6
7 312 Additional specimens examined for morphological studies are in Supplementary Table 2.
8
9
10
313 The genus Phaeochorella Theiss. & Syd. (type species P. parinarii (Henn.) Theiss. &
11
12 314 Syd., based on Cocconia parinarii Henn.) (Theissen and Sydow 1915), was found in
13
14 315 Africa on the herbaceous Pa. capensis, and on an arboreal host, Pa. curatellifolia (= Pa.
15
16
17 316 mobola) (Chrysobalanaceae) (Hennings 1901). However, the fungus is known from Brazil
18
19 317 (Medeiros 1994; Mendes et al. 1998; Souza 2017) on a subshrub host, Pa. obtusifolia,
20
21
22 318 with distribution limited to the Brazilian Cerrado, as retrieved from GBIF
23
24 319 (https://www.gbif.org/species/2985023) on May 18 2018.
25
26
27 320 Phaeochorella zonata Petrak, Sydowia 1(1–3): 122. 1947.
28
30
31
32
322 This species was then described by Petrak (1947) as showing “spots solitary to irregularly
33
34 323 or loosely dispersed, minute, orbicular to elliptical, often irregular, pale ochraceous to
35
36 324 reddish-brown, 0.75–2 mm diam, with dark well-defined margin; stromata epiphyllous in
37
38
39 325 center of solitary spots, varying from minute punctiform, 200–400 µm diam reaching up
40
41 326 to 1.4 mm diam, orbicular to elliptical, often irregular, black, verruculose; texture poorly
42
43
44 327 defined, for the most part, except around the perithecia; clypeus epidermal, sub-
45
46 328 carbonaceous, dark brown, opaque, pseudoparenchymatous; perithecia globose to
47
48
49
329 ellipsoidal, depressed, 140–180 µm wide, ostiolate; ostiole papilliform; asci 100–140 ×
50
51 330 15–20 µm, cylindrical to cylindrical-clavate, sub-sessile to short stipitate, 8-spored;
52
53 331 ascospores 17–22 × 10–15 µm, broadly ovoidal to ellipsoidal, dark brown, with an
54
55
56 332 equatorial sub-hyaline band; paraphyses few, immersed in mucous layer, evanescent,
57
58 333 mostly inconspicuous.”
59
60
61
62
63
65
14
334 Type: PHILIPPINES. LUZON: Prov. Rizal: Bosoboso, on leaves of undetermined

1
2 335 Fabaceae, 1924, MS Clemens 1800.
3
4
5 336 Distribution: known only from the type.
6
7 337 Notes─. Phaeochorella parinarii is associated with members of the Chrysobalanaceae,
8
9
10
338 while P. zonata is parasitic on Fabaceae. They are morphologically distinct as evidenced
11
12 339 in the respective descriptions. Thus, P. parinarii shows larger pseudostromata and
13
14 340 ascomata, whilst P. zonata is described with larger asci and ascospores, as well as having
15
16
17 341 evanescent paraphyses. Both, however, show ascospores with a hyaline equatorial band
18
19 342 (Petrak 1947). Without access to living material of P. zonata, we still decided to accept
20
21
22 343 the two species within the genus, given the authoritative description by Petrak (1947).
23
24 344 In conclusion, our data favors the establishment of a new family, Phaeochorellaceae,
25
26
27 345 presently with two genera (Phaeochorella and Phaeoappendicospora), morphologically
28
29 346 dissimilar.
30
31
32
347 KEY TO GENERA AND SPECIES OF PHAEOCHORELLACEAE
33
34 348 1. Single-celled ascospores 2
35
36 349 1’. Two-celled bi-appendiculate ascospores Phaeappendicospora thailandensis
37
38
39 350 2. Ascospore, brown to dark brown, with a medium hyaline band; on leaves of Parinari
40
41 351 species (Chrysobalanaceae); pseudostromata 2–4 mm wide; ascomata perithecial, 158–
42
43
44 352 350 μm diam; asci 75–96 × 10–13 μm; ascospores 10–13 × 4.5–7.5 μm; reported from
45
46 353 Brazil and Africa Phaeochorella parinarii
47
48
49
354 2’. Ascospore, brown, with medium hyaline band; on leaves of unidentified Fabaceae;
50
51 355 stromata 200–400 µm diam up to 1.4 mm diam; ascomata perithecial, 140–180 µm wide;
52
53 356 asci 100–140 × 15–20 µm; ascospores 17–22 × 10–15 µm, reported from the Philippines
54
55
56 357 Phaeochorella zonata
57
58 358
59
60
61 359 EXCLUDED TAXA
62
63
65
15
360 The type materials of Phaeochorella machaerii and P. ciliata, kindly provided by the
1
2 361 URM Herbarium, were herein reevaluated (Batista and Peres 1960; Batista and Bezerra
3
4
5 362 1961), and excluded from Phaeochorella based on morphological examination of
6
7 363 specimens (FIGS. 5−6). Phaeochorella machaerii is here accepted as a synonym of
8
9
10
364 Phyllachora puncta (Phyllachoraceae) (FIG. 5). The holotype material of Phaeochorella
11
12 365 ciliata was not well preserved, although the asci were shown to be apparently bitunicate,
13
14 366 and the ascospores were one-celled without an equatorial hyaline band (FIG. 6). The same
15
16
17 367 is true for P. artocarpi that for the same reason does not belong in the genus. In addition,
18
19 368 these species were excluded from Phaeochorella based on our concept of the genus, and
20
21
22 369 the revision by Petrak (1947), when describing P. zonata (type not examined), and
23
24 370 considering ascospores with median sub-hyaline to hyaline transverse band produced in
25
26
27 371 unitunicate cylindrical to subclavate asci, in ostiolate pseudostromatic ascomata, as
28
29 372 reliable taxonomic characteristics.
30
31
32
373 1. Phaeochorella artocarpi T.S. Ramakr. & K. Ramakr. Indian Acad. Sci., Pl. Sci._28: 37.
33
34 374 1948. On living leaves of Artocarpus lacucha Roxb. (Moraceae). INDIA, TAMILNADU,
35
36 375 Nilgiris, Buliar, Nov 1946, T.S. Ramakrishnnan. IASD1196. MYCOBANK MB #289272
37
38
39 376 Notes─. We had no access to the type material of P. artocarpi, however, the detailed
40
41 377 description and illustration provided by Ramakrishnan and Ramakrishnan (1948) showed
42
43
44 378 that the fungus does not fit in Phaeochorella sensu Thiessen and Sydow (1915) and
45
46 379 Petrak (1947), as it lacks the brown ascospores bearing a hyaline equatorial band, and
47
48
49
380 shows amphigenous highly developed pseudostromata, instead of exclusively epiphyllous
50
51 381 clypeate irregular stromata.
52
53 382
54
55
56 383 2. Phaeochorella machaerii Bat. & Peres. Publicações do Instituto de Micologia do
57
58 384 Recife 277: 15. 1960. FIG. 5
59
60
61
62
63
65
16
385 MYCOBANK MB336090

1
2 386 Specimens examined.—BRAZIL. MINAS GERAIS, Horto Florestal de Paraopeba, on
3
4
5 387 living leaves of Machaerium scleroxylon (Fabaceae), 12 Mar 1960, EP Heringer
6
7 388 (Holotype URM! 18692). DISTRITO FEDERAL: Brasília, Gama, -15,991211, -
8
9
10
389 48,087405, on living leaves of M. scleroxylon, 28 May 2010, MDM Santos #129 (UB
11
12 390 Mycol. Col. 21753).
13
14 391 Notes─. Phaeochorella machaerii was originally described as having bitunicate asci, but
15
16
17 392 re-examination of the holotype showed that it is a typical Phyllachora species, here
18
19 393 identified as Phyllachora puncta (Cooke) Cooke (Phyllachoraceae), which currently has
20
21
22 394 31 synonyms, as cited from Index Fungorum (http://www.indexfungorum.org, accessed
23
24 395 10 November 2018), and is widely known on the fabaceous genera Dalbergia and
25
26
27 396 Machaerium (Cannon 1991).
28
29 397 3. Phaeochorella ciliata Bat. & J. L. Bezerra. Memórias da Sociedade Broteriana 14: 41.
30
31
32
398 1961. MYCOBANK MB336089 FIG. 6
33
34 399 Specimen examined: BRAZIL. PERNAMBUCO: Carpina, on living leaves of Jaracatia
35
36 400 dodecaphylla) (Caricaceae), 31 Aug 1959, O. Soares (Holotype URM! 17529).
37
38
39 401 Notes─. Batista and Bezerra (1961) described this species as having bitunicate asci and
40
41 402 placed it in Phaeochorella probably based on the olive-brown color of the ascospores and
42
43
44 403 presence of paraphyses, which are also characteristic of several Phyllachora and Telimena
45
46 404 species. In our examination all ascospores were hyaline to pale brown, fusoid and highly
47
48
49
405 guttulate, and therefore not fitting in any genus within the Sordariomycetes, thus
50
51 406 explaining our decision in favor of the exclusion of this species from Phaeochorella.
52
53 407
54
55
56 408
57
58 409
59
60
61 410 DISCUSSION
62
63
65
17
411 The Phaeochorella specimens examined on Pa. capensis, Pa. curatellifolia and Pa.
1
2 412 obtusifolia, all show the characteristics of the type species of the genus Phaeochorella,
3
4
5 413 clearly differing from the excluded species (P. artocarpi, P. ciliata, P. machaerii), and
6
7 414 also from P. zonata. Theissen and Sydow (1915) described the type species of
8
9
10
415 Phaeochorella, P. parinarii, based on Cocconia parinarii Henn. (Hennings 1901) (FIG.
11
12 416 2). Only P. parinarii and P. zonata meet the criteria of the genus Phaeochorella, as
13
14 417 circumscribed by Petrak (1947), who emphasized the presence of the equatorial hyaline
15
16
17 418 band mentioned by Theissen and Sydow (1915) as a diagnostic morphological
18
19 419 characteristic of the genus. There is no molecular data to verify the phylogenetic position
20
21
22 420 of these two Phaeochorella species in relation to each other. However, we present a new
23
24 421 phylogeny for the type species P. parinarii, which in the absence of type material
25
26
27 422 available for sequencing, will be herein epitypified. When placed in an ample phylogeny
28
29 423 containing representative members of Sordariomycetes (Supplementary FIG.1), it was
30
31
32
424 shown to belong in the order Diaporthales, and not in the Phyllachorales. Now, on a more
33
34 425 restricted phylogeny, both Phaeochorella materials clustered with two representatives of
35
36 426 the genus Phaeoappendicospora, and together were segregated from other families of the
37
38
39 427 Diaporthales included in the analyses (FIG. 1).
40
41 428 Besides Phaeochorella, another genus, Apiosphaeria (type species A. guaranitica),
42
43
44 429 previously considered a member of the Phyllachoraceae, was recently shown through
45
46 430 molecular phylogenetic analyses to belong in Diaporthaceae (Guterres et al. 2018). In
47
48
49
431 addition, Phyllachora-like species that did not infect poaceous hosts, were shown to
50
51 432 belong in the new family Telimenaceae, also on the basis of DNA sequence analyses
52
53 433 (Mardones et al. 2017, 2018). Thus, Phyllachorales and Diaporthales have undergone
54
55
56 434 several changes (Castlebury et al. 2002; Rossman et al. 2007; Kirk et al. 2008; Guterres et
57
58 435 al. 2018). Using 28S sequences, six major lineages were elucidated (Castlebury et al.
59
60
61 436 2002), and with the increase of multilocus phylogenies a number of new families have
62
63
65
18
437 been recognized (Cheewangkoon et al. 2010; Crous et al. 2012; Suetrong et al. 2015,
1
2 438 Norphanphoun et al. 2016; Voglmayr et al. 2017). Senanayake et al. (2017) increased the
3
4
5 439 total number of families accepted in the Diaporthales to 21. Yang et al. (2018) recently
6
7 440 described a new family, Diaporthosporellaceae, based exclusively on a monotypical genus
8
9
10
441 Diaporthosporella (type species D. cercidicola) in China.
11
12 442 As P. parinarii formed a well-supported clade (BS = 97; BPP = 1) with the monotypic
13
14 443 Phaeoappendicospora (type species Ph. thailandensis), considered a genus incertae sedis
15
16
17 444 in the Diaporthales (Senanayake et al. 2017), the phylogeny performed suggests their
18
19 445 placement in a new family. In addition, Senanayake et al. (2017) described Ph.
20
21
22 446 thailandensis as a saprobic species, and showed its morphological distance from the plant
23
24 447 parasitic P. parinarii.
25
26
27 448 Thus, besides being two morphologically and ecologically different genera, they clustered
28
29 449 together within Diaporthales, but are phylogenetically distinct with high support (BS =
30
31
32
450 100 and BPP = 1).
33
34 451 Finally, Phaeoappendicospora, with the inclusion of the genus Phaeochorella in the
35
36 452 phylogenetic analyses of the order Diaporthales, formed a strongly supported clade,
37
38
39 453 recognizable as a valid family, herein designated as Phaeochorellaceae.
40
41 454
42
43
44 455 ACKNOWLEDGEMENTS
45
46
47 456 The authors thank CNPq (Conselho Nacional de Desenvolvimento Científico e
48
49
50 457 Tecnológico) for a Ph.D. fellowship given to the first author; CAPES (Coordenação de
51
52 458 Aperfeiçoamento de Pessoal de Nível Superior/Ministry of Education) for a Ph.D.
53
54
55
459 fellowship to the second author; CNPq for research grants to the penultimate and last co-
56
57 460 authors. We express our thanks to Leonor Costa Maia, curator of the Herbarium Pe.
58
59 461 Camille Torrend (URM) at Universidade Federal de Pernambuco. Thanks are given to Dr.
60
61
62
63
65
19
462 Teresa Iturriaga-Herbarium CUP Curator, and Dr. Robert Lücking-Herbarium B Curator
1
2 463 for lending such precious material as the Reliquiae Petrakianae of Cocconia parinarii and
3
4
5 464 Phaeochorella parinarii, respectively. To Prof. Mariza Sanchez (in memoriam) our
6
7 465 thanks for the excellence of her Fungarium assistance. This work received financial
8
9
10
466 support from the Fundação de Apoio à Pesquisa do Distrito Federal (FAP-DF) and CNPq
11
12 467 (PPBIO-Cerrado-MCTIC-CNPq).
13
14 468
15
16
17 469
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53 517 239–249. https://doi.org/10.3852/mycologia.98.2.239
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58 519 JC. 2018. Taxonomy, phylogeny, and divergence time estimation for Apiosphaeria
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2 521 https://doi.org/10.1080/00275514.2018.1465774
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658 LEGENDS
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4 Figure 1. The 50% rule-consensus tree of Diaporthales generated based on Bayesian
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6 Inference from DNA sequences for 28S, RPB2 and TEF1-α regions. Support values
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indicated on the branches represent Posterior Probabilities (BPP) of Bayesian Inference,
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11 and Bootstrap (BS) of Maximum Likelihood analyses. Thickened branches indicate BPP
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13 and BS higher than 0.98 and 95, respectively.
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18 Figure 2. Cocconia parinarii (≡ Phaeochorella parinari) isotype [Herbarium CUP-F-
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21 03689(2-99)]. A. Picture of isotype material. B. Leaf showing pseudostromata on the
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23 adaxial face. C. Section through a multilocular pseudostroma showing a periphysate
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26 ostiole (arrow). D. Ascospores. Bars: A= 10 mm, B = 5 mm, C = 100 µm, D = 10 µm.
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Figure 3. Phaeochorella parinarii. A. Label of Herbarium B exsiccate. B–C. Parinari
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33 curatellifolia showing fungal pseudostromata only on the adaxial face of the leaf. D.
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35 Superficial view of pseudostromata. E. Section through a pseudostroma showing an
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38 accumulated ascospore mass (arrows). F. Ascospore. G. Label of Herbarium B exsiccate.
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40 H. Pseudostromata on the adaxial face of Parinari capensis leaf. I. Superficial view of
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43 pseudostromata. J. Section through a pseudostroma containing a mass of ascospores
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45 (arrow). K. Periphysate ostiole erupting through the host epidermis, containing a group of
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ascospores. L. Ascospores. Bars: B−C = 5 mm, D = 500 µm, E = 100 µm, F = 10 µm, H =
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50 10 mm, I = 1 mm, J = 100 µm, K = 50 µm, L = 10 µm.
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56 Figure 4. Phaeochorella parinarii epitype on leaf of Parinari obtusifolia (UB Mycol.
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58
Coll. 23270). A–B. Pseudostromata. C. Section through a pseudostroma showing its
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61 intradermal insertion. D. Mature immersed perithecial ascoma with a periphysate ostiole.
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E. Ascoma with arrow showing textura prismatica of the wall. F. Immature asci and
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2 paraphyses (arrows). G. Septate paraphyses (arrows). H. Mature ascospores with typical
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5 sub-hyaline equatorial band. Bars: A = 2 cm, B = 5mm, C = 50 µm, D = 100 µm, E = 20
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7 µm, F = 10 µm, G = 20 µm, H = 10 µm.
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12 Figure 5. Phyllachora puncta on Machaerium scleroxylon (holotype URM! 18692). A–B.
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14 Pseudostromata. C. Section through a multilocilar peudostroma. D. Internal view of a
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17 locule with free ascospores. E. Paraphysate degraded asci. F. Ascospores. Bars: A = 2 cm,
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19 B = 0.5 mm, C = 100 µm, D = 10 µm, E–F = 10 µm.
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27 Figure 6. “Phaeochorella” ciliata on leaf of Jaracatia dodecaphylla (holotype URM!
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29 17529). A. Pseudostromata containing perithecia with punctiform ostioles. B–D. Asci and
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ascospores. Bars: A = 500 µm, B–D = 10 µm.
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36 Supplementary Figure 1. Phylogenetic tree of Sordariomycetes. Tree generated in
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39 Maximum Likelihood optimization from DNA sequences of 28S and RPB2 partial
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41 regions. Thickened branches indicate Bootstrap Support higher than 95 %.
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TABLE 1 REVISED. Fungal species used for molecular and/or
morphological study
TABLE 1. Details of fungal species used for molecular and/or morphological study, including the
GenBank accession numbers of the sequences used in the phylogenetic analyses.
ORDER
Taxon Country Host nrLSU RPB2 TEF1
Voucher
CALOSPHAERIALES
Calosphaeria africana
STEU 6182 EU367455
Calosphaeria pulchella
CBS 115999 Prunus sp. AY761075 GU180661
Jattaea sp.
MFLUCC 160534 Thailand Rhizophora mucronata KY034451 KY039317
TOGNINIALES
Phaeoacremonium vibratile
CBS 117115 France Fagus sylvatica DQ649065 HQ878611 KF764645
DIAPORTHALES
Alnecium auctum
CBS 124263 Austria Alnus glutinosa KF570154 KF570170 KF570200
Ambarignomonia petiolorum
CBS 121227 USA: VA Liquidambar EU255070 EU219307 EU221898
styraciflua
Amphiporthe hranicensis
CBS 119289 Austria Tilia platyphyllos EU199122 EU199137
Apiognomonia errabunda
AR 2813 Switzerland Fagus sylvatica NG027592 DQ862014 DQ313565
Apiosphaeria guaranitica
UB (Mycol. Coll.) 23795 Brazil Handroanthus MG573333 MG573316 MG573325
ochraceus
ochraceus
ochraceus
Celoporthe dispersa
CMW 9978 South Africa Syzygium cordatum HQ730854 HQ730841
Celoporthe eucalypti
CMW 26913 China Eucalyptus EC48 clone HQ730865 HQ730852
Celoporthe polygoni
MFLU 170965 Germany Rumex acetosa MF190063
Chiangraiomyces bauhiniae
MFLUCC 17 1669 Thailand Bauhinia sp. MF190064 MF377603 MF377598
MFLUCC 17 1670 Thailand Bauhinia sp. MF190065 MF377604 MF377599
Chromendothia citrina
AR 3445 Russia Quercus mongolica EU255074 EU219342 EU222013
Chrysoporthe cubensis
CMW 14394 Colombia Eucalyptus sp. JN940856 GQ290137
Coniella africana
CBS 114133 South Africa Eucalyptus nitens AY339293 KX833421 KX833600
Coniella australiensis
IMI 261318 South Africa Leaf litter AF408336 KX833497 KX833692
Coniella fragariae
CBS 110394 Peru: Iquitos Forest soil KJ710441 KX833499 KX833695
Coniella straminea
CBS 14922 USA Fragaria sp. AY339296 KX833506 AY339366
Coniella tibouchinae
CPC 18511 Brazil: Minas Gerais Tibouchina granulosa JQ281776 KX833507 JQ281778
Coniella wangiensis
CPC 19397 Australia Eucalyptus sp. JX069857 KX833509 KX833705
Coryneum arausiaca
MFLUCC 15 1110 Italy Quercus sp. MF190067 MF377610 MF377575
Cryptodiaporthe aesculi
AR 3580 Austria Aesculus AF408342 EU199138
hippocastanum
Cryptosporella hypodermia
CBS 116866 Austria Ulmus minor AF408346 EU199140
Cryptosporella suffusa
CBS 121077 Austria Alnus incana EU199124 EU199142
Cytospora centrivillosa
MFLUCC 17 1660 Italy Sorbus domestica MF190069 MF377601
Cytospora chrysosperma
CFCC 89630 China: Yulin, Salix psammophila KF765690 KF765706
Shaanxi
Cytospora hippophaes
CFCC 89640 China: Gannan, Hippophae KF765698 KF765714 KP310865
Gansu rhamnoides
Cytospora junipericola
BBH 42444 Italy Juniperus communis MF190071 MF377579
Cytospora nivea
CFCC 89643 China: Yulin, Salix psammophila KF765701 KF765717
Shaanxi
Cytospora sacculus
CFCC 89625 China: Gansu Juglans regia KR045725 KP310861
Province
Diaporthe cassines
CBS 136440 South Africa: Cassine peragua KF777208 KF777244
Western Cape
Province
Diaporthe eres
MFLUCC 14 0862 Russia Catalpa bignonioides MF190082 MF377596
MFLUCC 17 1661 Russia Catalpa bignonioides MF190083 MF377597
MFLUCC 17 1667 Russia Fraxinus MF190080 MF377594
pennsylvanica
MFLUCC 17 1668 Russia Fraxinus MF190081 MF377595
pennsylvanica
Diaporthe rudis
IT 1526 Italy Acer campestre MF190088 MF377576
MFLU 17 0895 Italy Acer campestre MF190089 MF377577
Discula destructiva
CBS 109771 USA Cornus nuttallii AF408359 EU199144
Ditopella biseptata
MFLU 17 0884 Italy Alnus glutinosa MF190090 MF377617
Ditopella ditopa
CBS 109748 Austria Alnus glutinosa EU199126 EU199145
Ditopellopsis sp.
CBS 121471 USA: NC Clethra alnifolia EU255088 EU219254 EU221936
Gnomonia gnomon
CBS 19953 Italy Corylus avellana AF408361 EU219295 EU221885
Gnomoniopsis racemula
AR 3892 USA: MN Epilobium EU255122 EU219241 EU221889
angustifolium
Hyaliappendispora galii
MFLUCC 161208 Italy Galium sp. MF190095
Juglanconis juglandina
D96 Austria Juglans regia KY427145 KY427195 KY427214
Juglanconis oblonga
MAFF 410216 Japan Juglans ailanthifolia KY427153 KY427203 KY427222
Juglanconis pterocaryae
MAFF 410079 Japan Pterocarya rhoifolia KY427155 KY427205 KY427224
Lamproconium desmazieri
MFLUCC 15 0870 Russia Tilia sp. KX430135 MF377605 MF377591
Mazzantia napelli
AR 3498 Austria Aconitum lycoctonum AF408368 EU219345 EU222017
Melanconiella chrysomelanconium
MFLU 17 0879 Italy Carpinus betulus MF190113 MF377619
Melanconiella ellisii
BPI 878343 Carpinus caroliniana JQ926339 JQ926406
Melanconis alni
AR 3500 Austria Alnus viridis AF408371 EU219300 EU221896
Melanconis marginalis
AR 3442 Canada Alnus rubra AF408373 EU219301 EU221991
Microascospora rubi
MFLU 17 0886 Italy Rubus ulmifolia MF190098 MF377611 MF377581
Occultocarpon ailaoshanense
LCM 52201 China: Yunnan Alnus nepalensis JF779852 JF779857 JF779862
Ophiodiaporthe cyatheae
YMJ 1364 Taiwan Cyathea lepifera JX570891 JX570893 KC465406
Ophiognomonia melanostyla
LCM 38901 Germany: Hesse Tilia cordata JF779854 JF779858
Ophiognomonia vasiljevae
AR 4298 USA: TN Juglans nigra EU255162 EU219331 EU221999
Phaeoappendispora
thailandensis
MFLU 12–2131 Thailand: Chiang Rai Quercus sp. MF190102 MF377613 MF377613
● Phaeochorella parinarii
UB (Mycol. Coll.) 23270 Brazil: DF Parinari obtusifolia MK050523 MK050190 MK050188
UB (Mycol. Coll.) 23397 Brazil: DF Parinari obtusifolia MK050524 MK050191 MK050189
Phaeocytostroma ambiguum
CPC 17077 South Africa Zea mays FR748100 FR748073
Pleuroceras pleurostylum
CBS 90679 Switzerland Salix helvetica EU255197 EU219311 EU221962
Pleuroceras tenellum
CBS 121082 USA: North Carolina Acer rubrum EU255202 EU199155 EU221907
Pseudoplagiostoma eucalypti
CPC 14161 Viet Nam Eucalyptus GU973604 GU973540
camaldulensis
Pseudoplagiostoma oldii
CBS 115722 Australia Eucalyptus GU973610 GU973565
camaldulensis
Pseudoplagiostoma variabile
CBS 113067 Uruguay Eucalyptus globulus GU973611 GU973566
Sirococcus tsugae
AR 4010 USA:Oregon Cedrus deodara EU255207 EU219289 EU221928
Stegonsporium acerophilum
CBS 117025 USA Acer saccharum EU039993 KF570173 EU040027
Stilbospora macrosperma
CBS 121883 Austria: Carpinus betulus JX517299 KF570196
Leithenbachtal
FIGURE 1 REVISED Click here to
access/download;Figure;FIG_1_PhaeochorellaceaePhylogeny
Figure 2 ISOTYPE Click here to access/download;Figure;Figure 2 Cocconia
parinari.png
Figure 3 OTHER MATERIALS DOCUMENTED Click here to access/download;Figure;Figure 3 other
materials.png
Figure 4 EPITYPE Click here to
access/download;Figure;FIG_4_Phaeochorella_parinarii_epitype
Figure 5 PHYLLACHORA PUNCTA ON MACHAERIUM Click here to access/download;Figure;FIG_5_Phaeochorella_machaerii.png
Figure 6 PHAEOCHORELLA CILIATA Click here to access/download;Figure;FIG_6_Phaeochorella_ciliata.png
EXPLORATORY PHYLOGENY Sordariomycetes
SUPPLLENTARY TABLE 1 REVISED Evolutive models data
SUPPLEMENTARY TABLE 1. EVOLUTIVE MODELS DATA.

Partition
Parameter
LSU rpb2 1ª rpb2 2ª rpb2 3ª tef1 1ª tef 2ª tef 3ª
Evolutive model GTR+I+G TIM2+G TPM3uf+I+G TPM2uf+I+G TPM2uf+G F81+G GTR+G
Likelihood -6585,247 -2950,077 -1924,635 -12637,475 -747,340 -680,548 -1510,791
Matrix length 1326 1134 1134 1134 723 723 723
Base frequencies - - - - - - -
Freq. A 0,258 0,270 0,366 0,152 0,304 0,329 0,013
Freq. C 0,239 0,263 0,207 0,330 0,181 0,253 0,565
Freq. G 0,293 0,343 0,172 0,287 0,381 0,172 0,275
Freq. T 0,210 0,125 0,255 0,231 0,135 0,246 0,147
Transition rates - - - - - equal -
R (AC) 1,116 1,896 2,960 2,006 2,368 - 77,050
R (AG) 4,049 1,691 3,250 9,717 1,143 - 113,226
R (AT) 2,378 1,896 1,000 2,006 2,368 - 0,022
R (CG) 0,868 1,000 2,960 1,000 1,000 - 0,022
R (CT) 10,939 7,128 3,250 9,717 1,143 - 14,860
R (GT) 1,000 1,000 1,000 1,000 1,000 - 1,000
Prop. Invariable
0,559 - 0,544 0,030 - - -
sites
Gamma 0,637 0,269 0,746 3,002 0,113 0,084 0,768
SUPPLEMENTARY TABLE 2
Supplementary Table 2. Additional UB (Mycol. Coll.) accesses of Phaeochorella parinarii, collected in official and
private natural reserves in the Brazlian Cerrado.
Access number Host species Date State Municipality Locality Latitude Longitude
UB (Mycol. Coll.) 366 Parinari obtusifolia 11/20/1991 Distrito Federal Brasilia Água Limpa Reserve/ Univ. of Brasilia -47,9567 -15,7173
UB (Mycol. Coll.) 392 Parinari sp. 11/20/1991 Distrito Federal Brasilia Água Limpa Reserve/ Univ. of Brasilia -47,9567 -15,7173
UB (Mycol. Coll.) 2084 Parinari obtusifolia 9/20/1992 Distrito Federal Brasilia QI 10, conjunto 6, Lago Norte -47,9567 -15,7173
UB (Mycol. Coll.) 2255 Parinari obtusifolia 6/18/1992 Distrito Federal Brazlândia Road Taguatinga to Brazlândia -48,1352 -15,625
UB (Mycol. Coll.) 2364 Parinari obtusifolia 7/10/1991 Distrito Federal Planaltina Águas Emendadas Ecological Station -47,5322 -15,6697
UB (Mycol. Coll.) 2563 Parinari obtusifolia 11/15/1981 Distrito Federal Brasilia Biological Exp. Sta. Univ. of Brasília -47,9567 -15,7173
UB (Mycol. Coll.) 2566 Parinari obtusifolia 11/10/1992 Distrito Federal Brasilia PAD/DF Cerrado Fragments -47,9567 -15,7173
UB (Mycol. Coll.) 2850 Parinari obtusifolia 12/22/1992 Bahia Correntina São Pedro Farm -45,4133 -13,4776
UB (Mycol. Coll.) 2851 Parinari obtusifolia 12/22/1992 Bahia Correntina São Pedro Farm -45,4133 -13,4776
UB (Mycol. Coll.) 5743 Parinari obtusifolia 2/10/1993 Distrito Federal Brasilia Brasília Botanic Garden -47,9567 -15,7173
UB (Mycol. Coll.) 6091 Parinari sp. 5/11/1994 Distrito Federal Brasilia Biological Exp. Sta. Univ. of Brasília -47,9567 -15,7173
UB (Mycol. Coll.) 8942 Parinari sp. 6/27/1995 Distrito Federal Planaltina Águas Emendadas Ecological Station -47,5322 -15,6697
UB (Mycol. Coll.) 10109 Parinari sp. 6/27/1995 Distrito Federal Brasilia Brasília National Park -47,9567 -15,7173
Chapada dos
UB (Mycol. Coll.) 11700 Parinari sp. 4/14/1996 Mato Grosso Chap. Guimarães National Park -55,5047 -15,045
Guimarães
Chapada dos
Guimarães
Chapada dos
Guimarães
Chapada dos
Guimarães
Mato Grosso do
UB (Mycol. Coll.) 12090 Parinari sp. 8/16/1996 Campo Grande EMBRAPA/DEPATE Ecol. Reserve -54,4913 -20,6396
Sul
Mato Grosso do
UB (Mycol. Coll.) 12109 Parinari sp. 8/16/1996 Campo Grande EMBRAPA/DEPATE Ecol. Reserve -54,4913 -20,6396
Sul
Mato Grosso do
UB (Mycol. Coll.) 12142 Parinari sp. 8/17/1996 Terenos Estância Vanessa -55,0252 -20,4467
Sul
Mato Grosso do
UB (Mycol. Coll.) 12160 Parinari sp. 8/17/1996 Terenos Estância Vanessa -55,0252 -20,4467
Sul
Mato Grosso do
UB (Mycol. Coll.) 12380 Parinari sp. 8/19/1996 Campo Grande EMBRAPA Lagoinha Reserve -54,4913 -20,6396
Sul
Mato Grosso do
UB (Mycol. Coll.) 12802 Parinari sp. 8/22/1996 Campo Grande CINDACTA via BR163 Highway -54,4913 -20,6396
Sul
UB (Mycol. Coll.) 13300 Parinari sp. 12/03/1996 Rondônia Vilhena R. Vermelho Road, km 20 from Vilhena -60,2525 -12,088
UB (Mycol. Coll.) 13885 Parinari sp. 7/04/1997 Goiás Mineiros Road Taguatinga to Brazlândia -52,7647 -17,6138
UB (Mycol. Coll.) 13927 Parinari sp. 4/07/1997 Goiás Mineiros Emas National Park at Água Ruim -52,7647 -17,6138
UB (Mycol. Coll.) 14090 Parinari sp. 4/09/1997 Goiás Mineiros Km 37 Road Mineiros to Emas Nat. Park -52,7647 -17,6138
UB (Mycol. Coll.) 15395 Parinari sp. 9/25/1997 Minas Gerais Chapada Gaúcha Grande Sertão Veredas Nacional Park -45,5765 -15,2639
UB (Mycol. Coll.) 15820 Parinari obtusifolia 9/23/1997 Distrito Federal Brasilia Águas Emendadas Ecological Station -47,9567 -15,7173
UB (Mycol. Coll.) 16590 Parinari sp. 6/21/1998 Rondônia Vilhena Km 48, BR 364 Vilhena to Parecís -60,2525 -12,088
UB (Mycol. Coll.) 20731 Parinari sp. 9/12/2007 Distrito Federal Vargem Bonita Água Limpa Reserve/ Univ. of Brasilia -47,9567 -15,7173
UB (Mycol. Coll.) 20831 Parinari sp. 7/11/2007 Distrito Federal Brasilia Brasília National Park: Cachoeira -47,9567 -15,7173
UB (Mycol. Coll.) 21148 Parinari sp. 10/01/2009 Distrito Federal Brasilia Arboretum Univ. of Brasília, Asa Norte -47,9567 -15,7173
UB (Mycol. Coll.) 21322 Parinari obtusifolia 5/02/2010 Distrito Federal Brasilia Brasília National Park -47,9567 -15,7173
UB (Mycol. Coll.) 22414 Parinari obtusifolia 8/24/2010 Distrito Federal Brasilia Brasília National Park -47,9567 -15,7173
UB (Mycol. Coll.) 23270 Parinari obtusifolia 5/07/2015 Distrito Federal Brasilia Cerrado CPD Bld., Univ. of Brasilia -47,9567 -15,7173

Mycologia: Phaeochorellaceae/Diaporthales: A New Fungal Family and A Re-Appraisal of Phaeochorella Species

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Mycologia: Phaeochorellaceae/Diaporthales: A New Fungal Family and A Re-Appraisal of Phaeochorella Species

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Mycologia

Phaeochorellaceae/Diaporthales: a new fungal family and a re-appraisal of

Manuscript Number: mycologia-D-18-00198R1

Article Type: Original Research

Manuscript Classifications: Morphology/Development; Plant Pathogens; Systematics

Full Title: Phaeochorellaceae/Diaporthales: a new fungal family and a re-appraisal of

Corresponding Author: JOSE CARMINE DIANESE, Ph.D.

Other Authors: Debora Cervieri Guterres, Dr. Sci.

Samuel Galvão-Elias, MSc

Maria do Desterro Mendes dos Santos, Dr. Sci.

Bruno Cézar Pereira de Souza, MSc

Camila Pereira Almeida, BSc

Danilo Batista Pinho, Dr. Sci.

Robert Neil Gerard Miller, PhD

Keywords: Cerrado; Diaporthaceae; Fungal taxonomy; multilocus phylogeny; Neotropical

1 Short title: Phaeochorellaceae, new tropical family of the Diaporthales

206 coelomycetes generally stromatic, showing phialidic or annelidic conidiogenous cells.

232 Phyllachoralles is here considered for a re-evaluation through morphological and

283 Pseudostromata 2−4 mm wide, epiphyllous, circular or irregular, solitary or gregarious,

334 Type: PHILIPPINES. LUZON: Prov. Rizal: Bosoboso, on leaves of undetermined

385 MYCOBANK MB336090

470 LITERATURE CITED

520 guaranitica, a Neotropical parasite on bignoniaceous hosts. Mycologia 110: 526–545.

571 Monod M. 1983. Monographie taxonomique des Gnomoniaceae (Ascomycetes) de l'ordre

622 combinational bootstrap approach. Molecular Phylogenetics and Evolution 44:1204–1223.

GenBank accession numbers of the sequences used in the phylogenetic analyses.

SUPPLEMENTARY TABLE 1. EVOLUTIVE MODELS DATA.

private natural reserves in the Brazlian Cerrado.

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