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Submitted 12 February 2014, Accepted 27 March 2014, Published online 28 April 2014
CorrespondingAuthor:Davi A. C. de Almeida – e-mail – daviaugusto@gmail.com 379
Brazilian Semi-Arid Ascomycetes I: New and interesting records of
hysteriaceous ascomycetes
Almeida DAC1*, Gusmão LFP1 and Miller AN2
1Universidade Estadual de Feira de Santana, Av. Transnordestina, S/N – Novo Horizonte, 44036-900. Feira de
Santana, BA, Brazil. daviaugusto@gmail.com
2Illinois Natural History Survey, University of Illinois, 1816 S. Oak St., Champaign, IL 61820
Almeida DAC, Gusmão LFP and Miller AN 2014 – Brazilian Semi-Arid Ascomycetes I: new and
interesting records of hysteriaceous ascomycetes. Mycosphere 5(2), 379–391, Doi
10.5943/mycosphere/5/2/11
Abstract
During an investigation of ascomycetes carried out in one area of Caatinga and three
enclaves of the Atlantic Forest in the semi-arid region of Brazil, we found ten interesting species of
hysteriaceous ascomycetes. Psiloglonium clavisporum and Rhytidhysteron opuntiae are new
records for South America and Anteaglonium abbreviatum, Hysterium angustatum and
Hysterobrevium smilacis are new records for Brazil. All species are described, illustrated and
discussed.
Key words – biodiversity – taxonomy – tropical microfungi
Introduction
The ascomycetes traditionally referred to as hysteriaceous fungi (Clements 1909) are
characterized by navicular, carbonaceous ascomata, known as hysterothecia, that dehisce by a
longitudinal slit and contain pseudoparaphyses and bitunicate asci (Boehm et al. 2009b). They are
cosmopolitan, usually saprobes or weak parasites occurring on dead leaves, bark, wood, stems or
twigs in terrestrial habitats. Due to their similar ascomatal morphology, they were historically
classified under the same family, Hysteriaceae (Bisby 1923). Later the species possessing laterally
compressed ascomata with an evaginated slit (mytilinidioid forms) were placed in a separate
family, Mytilinidiaceae (Barr 1990a). However, phylogenetic studies based on DNA and amino
acid sequence data revealed that the hysterothecium evolved several times within
Pleosporomycetidae (Boehm et al. 2009a; Boehm et al. 2009b). Although these studies supported
the classification of Hysteriaceae and Mytilinidaceae in their own orders, Hysteriales and
Mytiliniales, respectively, they also revealed the existence of a distinct lineage possessing
dichotomously branched hysterothecia that are laterally anastomosed along their length and occur
on a conspicuous subiculum to which the family Gloniaceae incertae sedis was erected (Boehm et
al. 2009b). Other genera with hysterothecia such as Farlowiella and Hysterographium were
classified as Pleosporomycetidae incertae sedis (Boehm et al. 2009a; Mugambi & Huhndorf 2009),
and the new family Anteagloniaceae was proposed for Anteaglonium in Pleosporales (Hyde et al.
2013).
The diversity of hysteriaceous ascomycetes in Brazil is poorly known. The first record of a
hysteriaceous ascomycete for Brazil was Rhytidhysteron rufulum (Spreng.) Speg. (as R. brasiliense
Mycosphere 5 (2): 379–391 (2014) ISSN 2077 7019
www.mycosphere.org Article
Mycosphere
Copyright © 2014 Online Edition
Doi 10.5943/mycosphere/5/2/11
380
and R. viride) reported by Spegazzini (1881b). Zogg (1962) added one more record, providing a
description and illustration of Ostreichnion curtisii (Duby) M.E. Barr (as Glonium curtisii). Later,
Batista et al. (1967) reported Hysterographium bonariense Speg., considered doubtful by Zogg
(1962), and Rhytidhysteron rufulum (as R. fuscum). Recently, Vitória (2012) reported the
occurrence of Gloniopsis praelonga (Schwein.) Underw. & Earle on a decaying rachis of Elaeis
guineensis Jacq. and on a decaying petiole of Mauritia flexuosa L.f. in northeastern Brazil.
The present work is the first of a series of studies regarding the diversity of
Dothideomycetes and Sordariomycetes in the Caatinga biome in the semi-arid region of Brazil. The
goal of these studies is to contribute to the knowledge about the diversity of ascomycetes in this
region of Brazil through the description and illustration of new and interesting taxa.
Materials & Methods
Study area and morphological examination
Samples of dead twigs and decaying wood were collected from March 2011 to May 2013 in
the Caatinga biome of the Brazilian semi-arid region in the following four areas: Paraiba State,
Mata do Pau-Ferro State Ecological Reserve (6º58’14”S, 35º44’55”W), Ceará State, Ubajara
NationalPark(3º50’24’’S,40º54’17’W), Bahia State, SerradaJibóia(12º50’57”S, 39º28’31”W),
and Piauí State, Serra das Confusões National Park (9º13’22”S, 43º29’39”W). The former three
areas are enclaves of Atlantic Forest located on mountain tops, while the fourth is characterized by
caatinga vegetation. Ascomata were measured and photographed using a QColor3 Olympus digital
camera mounted on a MZ7.5 Leica dissecting microscope. Ascomata were then sectioned with a
razor blade and the centrum was placed in a drop of water on a glass slide and covered with a cover
slip. Measurements and digital images of the pseudoparaphyses, asci and ascospores were made
using a BX51 Olympus microscope equipped with brightfield and Nomarski interference optics and
a QColor3 Olympus digital camera. Images were processed using Adobe Photoshop CS6 and
ImageJ. Specimens are deposited in the Herbarium of the State University of Feira de Santana
(HUEFS).
Results
Taxonomy
Anteaglonium abbreviatum (Schwein.) Mugambi & Huhndorf, Syst. Biodiv. 7(4): 460. 2009.
Figs. 1a–d.
≡Hysterium abbreviatum Schwein., Trans. Am. Phil. Soc., New Series 4(2): no. 2094. 1832.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twigs or decorticated wood of unidentified plant. Sexual state:
Hysterothecia erumpent to superficial, carbonaceous, black, subglobose to oblong, straight, smooth
or slightly striate laterally, with a longitudinal slit, sulcus shallow, gregarious, linear or rarely lying
at irregular angles, occurring on a black thin crust, tending to darken the substratum, without KOH
extractable pigment, 0.3–1 × 0.1–0.3 mm, 0.1–0.2 mm high. Pseudoparaphyses aseptate, hyaline,
branched above the asci, 1–2 µm wide. Asci bitunicate, cylindrical, short pedicillate, 8-spored,
obliquely to irregularly uniseriate, 37–75 × 4–5 µm. Ascospores ellipsoid to obovoid, straight,
hyaline, smooth, 1-septate, constricted at the septum, guttulate, 4.5–7(–8.5) × 2.5–4 µm. Asexual
state: Unknown.
Geographical distribution – Argentina, Chinese Taipei, Costa Rica, Ghana, India, New
Zealand, Papua New Guinea, Republic of Korea, Sierra Leone, South Africa, Spain, Turkey, United
States (GBIF 2013), Chile, Ecuador, Italy, Puerto Rico (Farr et al. 2013), Kenya (Mugambi &
Huhndorf 2009).
Material examined – Brazil, Piauí, Caracol, Serra das Confusões National Park, on
decorticated twig of unidentified plant, 28 Mar 2011, D.A.C. Almeida (HUEFS 42833); on
381
decorticated wood of unidentified plant, 28 Mar 2011, D.A.C. Almeida (HUEFS 42842); Ceará,
Ubajara, Ubajara National Park, on decorticated twig of unidentified plant, 4 May 2012, D.A.C.
Almeida (HUEFS 155111, HUEFS 141565); on decorticated wood of unidentified plant, 20 May
2013, D.A.C. Almeida & A.N. Miller (HUEFS 194240).
Notes – The characteristics of the Brazilian specimens are in agreement with those
presented by Lohman (1937), which based his description on the type specimen of A. abbreviatum
(as Glonium abbreviatum), Zogg (1962) and Lorenzo & Messuti (1998). Mugambi & Huhndorf
(2009) reported the production of a KOH extractable pigment from the ascomata of A.
abbreviatum, however this characteristic was not reported by the above mentioned authors and not
observed in the Brazilian specimens. Although Anteaglonium abbreviatum is widely distributed
throughout the world having been found in North, Central and South America, Africa, Asia, Europe
and Oceania (Farr et al. 2013; GBIF 2013), this study represents the first record for Brazil.
Gloniella adianti (Kunze) Petr. [as 'adianthi'], Annls. Mycol. 29(1/2): 120. 1931. Figs. 1e–h
≡Hysterium adianti Kunze [as 'adianthi'], Flora Jena 13: 371. 1830.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on decorticated wood of unidentified plant. Sexual state: Hysterothecia
superficial, carbonaceous, black, navicular, straight, smooth laterally, with a longitudinal slit that is
at first closed, opening slightly at maturity, gregarious, linear, tending to darken the substratum,
without KOH extractable pigment, 0.4–1.5 mm × 0.1–0.3 mm, 0.1–0.2 mm high. Pseudoparaphyses
aseptate, hyaline, branched, anastomosed distally, forming a pigmented epithecium above the asci,
1–1.5 µm wide. Asci bitunicate, cylindrical to clavate, short pedicillate, 8-spored, obliquely
uniseriate, 70–104 × 10.5–17 µm. Ascospores ellipsoid to fusiform, rarely clavate, slightly curved
or straight, hyaline, smooth, 3-septate, constricted at the septum, supra-median cell slightly broader
than other cells, guttulate, surrounded by a gelatinous sheath, 15–21 × 4.5–8 µm. Asexual state:
Unknown.
Geographical distribution – Brazil (GBIF 2013), England, Ireland (Farr et al. 2013),
Portugal (Zogg 1962).
Material examined – Brazil, Ceará, Ubajara, Ubajara National Park, on decorticated wood
of unidentified plant, 15 Nov 2012, D.A.C. Almeida (HUEFS 192136).
Notes – Gloniella is currently classified in Hysteriaceae (Lumbsch & Huhndorf 2010).
Zogg (1962) accepted six species in his account of the genus with an additional seven species later
added (Joshi & Patwardhan 1971; Sivanesan et al. 1988; Pande & Rao 1991; Steinke & Hyde 1997;
Checa et al. 2007). Since the size of the ascospores of many species of Gloniella are identical or
nearly so, Steinke & Hyde (1997) and Boehm et al. (2009a) suggested that many species may be
conspecific. Two species considered doubtful by Zogg (1962) were recognized by Barr (2009), G.
abietina Syd and G. lapponica (P. Karst.) Sacc. Recently, Messuti & Lorenzo (2009) reviewed six
type species of Gloniella from Chile and Argentina described by Spegazzini, all of them considered
doubtful species by Zogg (1962). As a result, two species were maintained as doubtful, G.
antarctica Speg. and G. gilliesi Speg., and three species were recognized, G. australis Speg., G.
chilensis Speg. and G. multiseptata Speg. Gloniella australis var. minor was considered a synonym
of G. australis. Although Messuti & Lorenzo (2009) did not examine the type of G. araucana
Speg., they also maintained it as a doubtful species.
The Brazilian specimen has longer asci and wider ascospores than presented by Zogg
(1962) for G. adianti (asci (25)30–40(45) µm long; ascospores 3–5 µm wide). Other differences are
the occurrence of the ascomata on a substratum tending to darken and the presence of a sheath in
the ascospores of the Brazilian specimen. However, since the morphology and length of the
ascospores are identical to those reported by Zogg (1962), we consider the observed differences to
be variations within the species and not sufficient to warrant a new taxon. Thus, we prefer to assign
our material to G. adianti and expand the concept of this species. Obviously, a review of the genus
and its species based on molecular data is necessary to clarify its phylogenetic placement within
Dothideomycetes and establish species boundaries within the genus.
382
Hysterium angustatum Alb. & Schwein., Consp. Fungi (Leipzig): 55. 1805. Figs. 1i–l
For synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twigs of unidentified plant. Sexual state: Hysterothecia superficial
with base immersed, carbonaceous, black, subglobose to elongate, straight or flexuous, smooth
laterally, with a longitudinal slit, sulcus shallow, gregarious, lying at irregular angles, without KOH
extractable pigment 0.2–1.4 × 0.1–0.3 mm, 0.1–0.3 mm high. Pseudoparaphyses septate, hyaline,
branched above the asci, 1–1.5 µm wide. Asci bitunicate, cylindrical, short pedicillate, 8-spored,
irregularly biseriate, 57.5–88 × 9–11 µm. Ascospores ellipsoid, slightly curved or straight, pale
brown, concolorous, smooth, 3-septate, constricted at the septum, 14–16.5 × 4–6.5 µm. Asexual
state: Unknown.
Geographical distribution – Algeria, Argentina, Australia, Belgium, Bulgaria, Chile,
Chinese Taipei, Denmark, France, Germany, Greece, India, Ireland, Luxembourg, Malay
Peninsula, Morocco, Pakistan, Poland, Russia, Spain, Turkey, United Kingdom (Farr et al. 2013;
GBIF 2013), Austria (Dämon et al. 2000), Italy, Japan, Norway, Portugal, Sweden, Ukraine, United
States (as H. acuminatum, GBIF 2013), Libya (as H. angustatum var. cerotoniae, Farr et al. 2013),
Mexico (Medel & Guzman 1999) New Zealand, Kenya (Mugambi & Huhndorf 2009), South
Africa (Lee & Crous 2003).
Material examined – Brazil, Paraíba, Areia, Mata do Pau-Ferro State Ecological Reserve, on
twigs of unidentified plant, 7 Nov 2011, D.A.C. Almeida (HUEFS 131189); 4 Jul 2012, D.A.C.
Almeida (HUEFS 192060).
Notes – Hysterium angustatum differs from H. pulicare Pers., the most similar species,
mainly by concolorous and smaller ascospores. Recent phylogenetic analyses based on four nuclear
genes demonstrated that they are closely related (Boehm et al. 2009b). Lohman (1933) and Zogg
(1962) described larger asci (105–120 × 10–16 µm and 100–120 × 11–14 µm, respectively) than
found in the Brazilian specimens, while Ellis & Everhart (1892) reported wider asci with similar
lengths (75–80 × 12–15 µm). Ellis & Everhart (1892) and Lohman (1933) described H. angustatum
with larger ascospores (15–22 × 6–7 µm and 22–26 × 6.5–8(9) µm, respectively) than observed in
this study. The measurements of the asci and ascospores of the Brazilian material, however, are
very similar to those stated by Lee & Crous (2003). The hysterothecia described here have shallow
slits, differing from the deep slits reported by the above authors. Although H. angustatum has a
wide distribution, this is the first record for Brazil.
Hysterobrevium mori (Schwein.) E. Boehm & C.L. Schoch, in Boehm, Mugambi, Miller,
Huhndorf, Marincowitz, Spatafora & Schoch, Stud. Mycol. 64: 62. 2009. Figs. 1m–p
≡Hysterium mori Schwein., Syn. Fung. Amer. bor.: no. 2087. 1832.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twig of unidentified plant. Sexual state: Hysterothecia superficial,
carbonaceous, black, ellipsoid to elongate, with obtuse ends, straight or lightly flexuous, smooth or
lightly striated laterally, with a longitudinal slit, sulcus shallow or deep, gregarious, linear,
occurring on a black thin crust, without KOH extractable pigment, 0.3–1.7 × 0.1–0.2 mm, 0.1–0.2
mm high. Pseudoparaphyses septate, hyaline, branched apically above the asci, 1.5–2.5 µm. Asci
bitunicate, cylindrical, short pedicillate, 8-spored, irregularly biseriate, 65.5–95 × 12–16 µm.
Ascospores ellipsoid or clavate, straight, pigmented, smooth, with 4–5 transverse septa and one
longitudinal septum usually associated with mid-cells and sometimes present obliquely in end cells,
15–20.5 × 6–7 µm. Asexual state: Aposphaeria-like (Boehm et al. 2009a).
Geographical distribution – Germany (GBIF 2013), Argentina, Australia, Brazil, Canada, Costa
Rica, Denmark, Ecuador, France, Ghana, India, Luxembourg, Norway, Pakistan, Portugal, Spain,
Sweden, United Kingdom, United States (as Hysterographium mori, GBIF 2013), Austria (as
Hysterographium pumilionis, GBIF 2013), Italy (as Hysterium grammodes, GBIF 2013), Malta (as
Hysterium vulgare, GBIF 2013), Morocco (as Hysterographium grammodes, GBIF 2013),
Venezuela (as Hysterographium punctiforme, GBIF 2013), Greece (Farr et al. 2013), Kenya
(Boehm et al. 2009a), Tunisia (as Hysterographium ziziphi, Patouillard 1897).
383
Fig. 1 – a–d Anteaglonium abbreviatum. a Hysterothecia. b Ascus with ascospores. c–d Ascospores. e–h
Gloniella adianti. e Hysterothecia. f Ascus with ascospores. g–h Ascospores. i–l Hysterium angustatum. i
Hysterothecia. j Asci with ascospores. k–l Ascospores. m–p Hysterobrevium mori. m Hysterothecia. n Ascus
with ascospores. o–p Ascospores. Bars = 0.5 mm (a, e, i, m), 10 µm (b, f, j, n), 5 µm (c, d, g, h, k, l, o, p).
Material examined – Brazil, Piauí, Caracol, Serra das Confusões National Park, on twig of
unidentified plant, 28 Mar 2011, D.A.C. Almeida (HUEFS 42835).
Notes – Hysterobrevium was recently erected by Boehm et al. (2009a) to accommodate
three species transferred from other genera based on molecular analyses. Hysterobrevium mori
(type species) was transferred from Hysterographium, while H. smilacis (Schwein.) E. Boehm &
C.L. Schoch and H. constrictum (N. Amano) E. Boehm & C.L. Schoch were transferred from
Gloniopsis. The measurements of the Brazilian specimen agree with Boehm et al. (2009a) and Ellis
& Everhart (1892), but some morphological variations were observed such as hysterothecia with
obtuse rather than acuminate ends and ascospores that do not constrict at the median septum.
384
Hysterobrevium smilacis (Schwein.) E. Boehm & C.L. Schoch, in Boehm, Mugambi, Miller,
Huhndorf, Marincowitz, Spatafora & Schoch, Stud. Mycol. 64: 63. 2009. Figs. 2a–d
≡Hysterium smilacis Schwein., Schr. Naturf. Ges. Leipzig 1: 49. 1822.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twig of unidentified plant. Sexual state: Hysterothecia superficial,
carbonaceous, black, navicular, straight or flexuous, smooth laterally, with a longitudinal slit,
sulcus slightly deep, subgregarious to scattered, lying at irregular angles, without KOH extractable
pigment, 0.3–1 × 0.1–0.4 mm, 0.1–0.3 mm high. Pseudoparaphyses septate, hyaline, branched
apically forming an epithecium above the asci, 1–1.5 µm wide. Asci bitunicate, cylindrical to
clavate, short pedicillate, 8-spored, irregularly biseriate, 80–134 × 13–18 µm. Ascospores ellipsoid,
straight, hyaline, smooth, (4–)5–6 transversally septate, with a longitudinal septa usually associated
with mid-cells and sometimes present obliquely in end cells, strongly constricted at the median
septum, slightly constricted at other cells, surrounded by a gelatinous sheath, 16.5–24 × 6–9 µm.
Asexual state: Aposphaeria-like (Boehm et al. 2009a).
Geographical distribution – Australia, Austria, Belgium, Czech Republic, Estonia, France,
Germany, India, Ireland, Italy, Morocco, Norway, Pakistan, Poland, Portugal, Russia, Spain,
Sweden, United States (GBIF 2013), Ukraine (Farr et al. 2013).
Material examined – Brazil, Ceará, Ubajara, Ubajara National Park, on twig of unidentified
plant, 4 May 2012, D.A.C. Almeida (HUEFS 155107).
Notes – The characteristics of the Brazilian specimen agree with the descriptions consulted
(Ellis & Everhart 1892; Boehm et al. 2009a). Although H. smilacis has a wide distribution, this
study is the first to discover this species in Brazil.
Oedohysterium insidens (Schwein.) E. Boehm & C.L. Schoch, in Boehm, Mugambi, Miller,
Huhndorf, Marincowitz, Spatafora & Schoch, Stud. Mycol. 64: 59 (2009) Figs. 2e–h
≡Hysterium insidens Schwein., Trans. Am. Phil. Soc., Ser. 2 4(2): 244. 1832.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twig of unidentified plant. Sexual state: Hysterothecia erumpent or
superficial, carbonaceous, black, oblong, straight, smooth laterally, with a longitudinal slit, deep,
gregarious, linear, occurring on a black thin crust, without KOH extractable pigment, 0.3–0.8 ×
0.1–0.3 mm, 0.1–0.3 mm high. Pseudoparaphyses septate, hyaline, branched above the asci, 1–1.5
µm wide. Asci bitunicate, cylindrical, short pedicillate, 8-spored, irregularly biseriate, 76–88 × 16–
17 µm. Ascospores ellipsoid to fusiform, straight to slightly curved, brown, smooth, transversely 6–
8-septate, constricted at septa, with a prominent swollen supra median cell, 25–35(–39) × 8–11 µm.
Asexual state: Septonema spilomeum Berk. (Boehm et al. 2009a).
Geographical distribution – Australia, Canada, Germany, India, Italy, Portugal, Spain,
South Africa, Switzerland, United Kingdom, United States (GBIF 2013), Brazil (as Hysterium
apiculatum, GBIF 2013), Chile (as Hysterium batucense, Farr et al. 2013), Argentina (as Hysterium
andicola, Spegazzini 1912), Morocco (as Hysterium atlantis, Maire & Werner 1937).
Material examined – Brazil, Piauí, Caracol, Serra das Confusões National Park, on twig of
unidentified plant, 28 Mar 2011, D.A.C. Almeida (HUEFS 42837).
Notes – Oedohysterium was erected by Boehm et al. (2009a) for three species based on
molecular data. Oedohysterium insidens (type species) and O. sinense (Teng) E. Boehm & C.L.
Schoch were transferred from Hysterium, while O. pulchrum (Checa, Shoemaker & Umaña) E.
Boehm & C.L. Schoch was moved from Hysterographium. The genus is characterized by
phragmospores with a prominent swollen supra median cell. Oedohysterium insidens differs from
O. sinense by smaller ascospores and from O. pulchrum by absence of longitudinal septa in the
ascospores. The features of the Brazilian specimen agree with Boehm et al. (2009a) and van der
Linde (1992), except the asci are shorter (130–150 µm versus 76–88 µm) and there is a black thin
crust under the ascomata. This black thin crust was observed by Schweinitz (1832) but not
mentioned by Ellis & Everhart (1892). The measurements are also similar to that reported by Ellis
& Everhart (1892), who examined the original type specimen, except the ascospores of the
385
Brazilian specimen are wider (6–8 µm versus 8–11 µm).
Ostreichnion curtisii (Duby) M.E. Barr, Mycotaxon 3(1): 86. 1975. Figs. 2i–l
≡Hysterium curtisii Duby, Mém. Soc. Phys. Hist. nat. Genève 16(1): 42. 1861.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twigs or woods of unidentified plant. Sexual state: Hysterothecia
superficial, carbonaceous, black, elongate conchate, straight, smooth laterally, with a longitudinal
slit, sulcus deep, subgregarious to scattered, lying at irregular angles, without KOH extractable
pigment. 0.3–0.7 × 0.1–0.3 mm, 0.2–0.3 mm high. Pseudoparaphyses hyaline, septate, branched
above the asci, 1–2.5 µm wide. Asci bitunicate, cylindrical, short pedicillate, 4–8-spored,
irregularly biseriate, 128–133 × 34–36 µm. Ascospores cylindrical, straight to slightly curved,
subhyaline to pale brown, smooth, ends broadly rounded, 1-septate, septum submedian, strongly
constricted at the septum, 59–62.5 × 14.5–17.5 µm. Asexual state: Unknown.
Geographical distribution – Brazil (as Glonium curtisii, Zogg 1962), United States (GBIF
2013).
Material examined – Brazil, Paraíba, Areia, Mata do Pau-Ferro State Ecological Reserve, on
twigs of unidentified plant, 3 Jul 2012, D.A.C. Almeida (HUEFS 192079); 18 Mar 2013, D.A.C.
Almeida (HUEFS 192178); Bahia, Santa Teresina, Serra da Jibóia, on wood of unidentified plant, 9
May 2013, D.A.C. Almeida & A.N. Miller (HUEFS 192249); Ceará, Ubajara, Ubajara National
Park, on twigs of unidentified plant, 4 May 2012, D.A.C. Almeida (HUEFS 131816, HUEFS
131817); 6 May 2012, D.A.C. Almeida (HUEFS 155083); 14 Nov 2012, D.A.C. Almeida (HUEFS
192109).
Notes – Only three taxa are accepted in Ostreichnion: O. curtisii, O. nova-caesariense
(Ellis) M.E. Barr and the type species, O. sassafras (Schwein.) M.E. Barr (Barr 1975; Boehm et al.
2009b). Ostreichnion curtisii is an atypical species in the genus and can be easily differentiated
from the other two species by ascospores with a single septum. In a recent phylogenetic study
based on molecular data of Hysteriaceae and Mytilinidiaceae, including O. sassafras and O.
curtisii, Boehm et al. (2009b) found that Ostreichnion grouped inside Hysteriaceae and transferred
it from Mytilinidiaceae. Another finding was that these two species are unrelated and the genus
should be considered artificial.
The features of our material agree with Barr (1975) except the asci are shorter (128–133 µm
versus 150–180(–220) µm). Zogg (1962) reported on a specimen of O. curtisii from Brazil with
smaller ascospores ((31–)35–45(–53) × (8–)9–11(–13) µm), but considered it as a variation within
the species.
Psiloglonium clavisporum (Seaver) E. Boehm, C.L. Schoch & Spatafora, Mycol. Res. 113(4): 469.
2009. Figs. 3a–d
≡Glonium clavisporum Seaver, Mycologia 17(1): 4. 1925.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twig of unidentified plant. Sexual state: Hysterothecia superficial,
carbonaceous, black, ellipsoid to elongate, straight or flexuous, smooth or striate laterally, with a
longitudinal slit, sulcus deep, gregarious, linear, without KOH extractable pigment, 0.5–5 × 0.1–0.4
mm, 0.1–0.5 mm high. Pseudoparaphyses aseptate, hyaline, branched above the asci, 1–1.5 µm
wide. Asci bitunicate, clavate, short pedicillate, 8-spored, obliquely uniseriate, 108.5–140 × 10–12
µm. Ascospores clavate, straight or lightly curved, hyaline, smooth, 1-septate, constricted at the
septum, 18–22.5 × 6–8.5 µm. Asexual state: Sporidesmium stygium Berk. & M.A. Curtis (Lohman
1937).
Geographical distribution – India (GBIF 2013), Japan (Hosoya & Tanaka 2007), Nicaragua
(as Glonium clavisporum, Seaver 1925), Saint Croix (as Glonium clavisporum, Seaver 1925),
United States (Boehm et al. 2009b).
Material examined – Brazil, Paraíba, Areia, Mata do Pau-Ferro State Ecological Reserve, on
twig of unidentified plant, 3 Apr 2013, D.A.C. Almeida (HUEFS 192176).
386
Fig. 2 – a–d Hysterobrevium smilacis. a Hysterothecia. b Ascus with ascospores. c–d Ascospores. e–h
Oedohysterium insidens. e Hysterothecia. f Ascus with ascosopores. g–h Ascospores. i–l Ostreichnion
curtisii. i Hysterothecia. j Ascus with ascospores. k–l Ascospores. Bars = 0.5 mm (a, e), 0.2 mm (i), 20 µm
(j), 10 µm (b, f, k, l), 5 µm (c, d, g, h).
Notes – Previously considered synonymous with Glonium by Zogg (1962), Psiloglonium
was reinstated under Hysteriaceae by Boehm et al. (2009b) based on molecular data to
accommodate species with unbranched, non-subiculate hysterothecia. Glonium was placed in its
own family, Gloniaceae, closely related to Mytilinidiaceae, and retained species with
dichotomously branched, laterally anastomosed, flabelliform or pseudo-stellate, subiculate
hysterothecia.
387
Due to the similarities between P. clavisporum and P. lineare (Fr.) Petr., Zogg (1962)
considered them synonyms (under Glonium) despite the differences presented by Lohman (1937)
such as ascospore length ((15–)16–18(–22) µm versus (10–)12–15 µm, respectively) and associated
anamorph (Sporidesmium stygium versus pycnidial Plenomous, respectively). We agree with
Lohman (1937) and Boehm et al. (2009b) that these differences are sufficient to maintain them as
separate entities. Psiloglonium simulans (W.R. Gerard) E. Boehm, C.L. Schoch & Spatafora was
also considered by Zogg (1962) as a synonym (under Glonium) of P. clavisporum. Besides the
morphological differences pointed out by Lohman (1937), Boehm et al. (2009b) provided
additional evidence from molecular data for the separation of P. simulans and P. clavisporum.
The features of the Brazilian specimen agree with Seaver (1925) and Lohman (1937),
except the asci are longer than those presented by Lohman (1937) (90–110 µm). This study
represents the first record of P. clavisporum for South America.
Rhytidhysteron opuntiae (J.G. Br.) M.E. Barr, Mem. N. Y. Bot. Gdn. 62: 72. 1990. Figs. 3e–h
≡Hysterographium opuntiae J.G. Br., Mycologia 45(6): 967. 1953.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat: Saprobe on twig of unidentified plant. Sexual state: Hysterothecia erumpent,
carbonaceous, black, oblong, straight or flexuous, smooth laterally, with a longitudinal slit, sulcus
deep, gregarious, lying at irregular angles, without KOH extractable pigment, 0.5–1.7 × 0.2–0.5
mm, 0.3–0.4 mm high. Pseudoparaphyses septate, hyaline, branched above the asci, 1.5–2.5 µm
wide. Asci bitunicate, cylindrical to clavate, short pedicillate, 8-spored, uniseriate, 150–207 × 10–
14 µm. Ascospores ellipsoid, ends obtuse, brown, smooth, transversally 3-septate, usually with one
longitudinal septum in mid-cells, slightly constricted at median septa, thick-walled, 20–25 × 7.5–10
µm. Asexual state: Unknown.
Geographical distribution – United States (as Hysterographium opuntiae, Brown 1953),
Kenya (Boehm et al. 2009a).
Material examined – Brazil, Piauí, Caracol, Serra da Confusões National Park, on twig of
unidentified plant, 28 Mar 2011, D.A.C. Almeida (HUEFS 42841).
Notes – Previously classified in the Patellariaceae (Samuels & Müller 1979), Rhytidhy-
steron is now a member of the Hysteriaceae (Boehm et al. 2009a). The phylogenetic analyses
carried out by Boehm et al. (2009a) indicate that the genus is artificial since the single isolate of R.
opuntiae from Kenya grouped outside the clade formed by R. rufulum (type species) and R.
hysterinum (Dufour) Samuels & E. Müll. They preferred, however, to not remove R. opuntiae from
Rhytidysteron until additional phylogenetic analyses with more isolates support this result.
The features of our material agree with Brown (1953) and Barr (1990b), except for the
longer asci (85–160 µm and 130–160 µm, respectively, vs. 150–207). This study is the first record
of R. opuntiae for South America.
Rhytidhysteron rufulum (Spreng.) Speg., Anal. Soc. C ient. Argent. 90(1-6): 177. 1921.
Figs. 3i–m
≡Hysterium rufulum Spreng., K. Svenska Vetensk-Akad. Handl. 46: 50. 1820.
For more synonyms see Index Fungorum (www.indexfungorum.org).
Habitat – Saprobe on twigs of unidentified plant. Sexual state: Hysterothecia erumpent,
carbonaceous, black, navicular, straight or flexuous, striate laterally, striations perpendicular to the
long axis, with a longitudinal slit, sulcus closed at first, then later opening to become irregularly
apothecioid at maturity with incurved margins, gregarious, linear, without KOH extractable
pigment, 0.8–2.5 × 0.25–1.2 mm, 0.4–0.8 mm high. Pseudoparaphyses septate, hyaline, branched
distally, 2–3 µm wide, covered by a black or red pseudoepithecium. Asci bitunicate, cylindrical to
clavate, short pedicillate, 8-spored, irregularly uniseriate, 220–248 × 15–21 µm. Ascospores ellip-
soid, ends rounded to subacute, straight or curved, central cells brown, concolorous or with median
cells darker than distal cells, smooth, 3-septate, slightly constricted at the septa, thick walled, 21–32
× 7.5–11 µm. Asexual state: Diplodia-like and Aposphaeria-like (Samuels & Müller 1979).
388
Fig. 3 – a–d. Psiloglonium clavisporum. a Hysterothecia. b Ascus with ascospores. c–d Ascospores. e–h
Rhytidhysteron opuntiae. e Hysterothecia. f Asci with ascospores. g–h Ascospores. i–m Rhytidhysteron
rufulum. I–j Hysterothecia. k Asci with ascospores and pseudoparaphyses. l–m Ascospores.
Bars = 1 mm (a, e, i), 0.5 mm (k), 20 µm (f, j), 10 µm (b), 5 µm (c, d, g, h, l, m).
Geographical distribution – Argentina, Cook Islands, Costa Rica, Dominica, France, India,
Japan, New Zealand, Philippines, Puerto Rico, Tonga, United States (GBIF 2013), China, Cuba,
Ghana, Jamaica, Malaysia, Papua New Guinea, Sierra Leone, Tanzania (as Tryblidiella rufula,
GBIF 2013), Brazil (Spegazzini 1881a), Micronesia (as Rhytidhysteron viride, GBIF 2013), Kenya
(Boehm et al. 2009a).
389
Material examined – Brazil, Piauí, Caracol, Serra das Confusões National Park, on twigs of
unidentified plant, 28 Mar 2011, D.A.C. Almeida (HUEFS 42836, HUEFS 42845, HUEFS 131197,
HUEFS 131198); 29 Mar 2011, D.A.C. Almeida (HUEFS 192084, HUEFS 192085); 30 Mar 2011,
D.A.C. Almeida (HUEFS 192082); Paraíba, Areia, Mata do Pau-Ferro State Ecological Reserve, on
twigs of unidentified plant, 8 Nov 2011, D.A.C. Almeida (HUEFS 131048, HUEFS 131049,
HUEFS 192070); 3 Jul 2012, D.A.C. Almeida (HUEFS 192057); 17 Apr 2013, D.A.C. Almeida
(HUEFS 192194); 18 Apr 2013, D.A.C. Almeida (HUEFS 192197, HUEFS 192199, HUEFS
192175); Ceará, Ubajara, Ubajara National Park, on twigs of unidentified plant, 4 May 2012,
D.A.C. Almeida (HUEFS 131253); 16 Nov 2012, D.A.C. Almeida (HUEFS 192128); 19 May
2013, D.A.C. Almeida & A.N. Miller (HUEFS 194207); Bahia, Santa Teresinha, Serra da Jibóia,
on twig of unidentified plant, 10 May 2013, D.A.C. Almeida & A.N. Miller (HUEFS 192253).
Notes – Murillo et al. (2009) performed an assessment of the intraspecific variability of R.
rufulum among several isolates from Costa Rica based on molecular (ITS rDNA), morphological
and chemical data. They found four distinct lineages that correlated with morphological differences
(hysterothecia margin, pseudoepithecium color, asci and ascospores size) and chemical
components. The authors, however, did not propose any nomenclatural changes, but highlighted the
necessity of including isolates from other regions and a revaluation of the type specimens
previously listed as synonyms of R. rufulum.
Our material agrees with the descriptions by Samuels & Müller (1979), Kutorga &
Hawksworth (1997) and Murillo et al. (2009). The Brazilian specimen, however, has longer asci
(220–248 vs. 77–100(–112) µm) and wider ascospores ((7.5–11 vs. 3.5–4.5(–6.5) µm) than
reported by Samuels & Müller (1979).
Acknowledgements
The authors thank the Brazilian Federal Agency for Support and Evaluation of Graduate
Education (CAPES – proc. 071/2012), the National Council for Scientific and Technological
Development (CNPq – proc. 14/2011), and the Program of Research on Biodiversity in the
Brazilian Semi-arid (PPBIO Semi-arid/Ministry of Technology and Science – proc. 558317/2009-
0) for financial support. The first author also thanks CAPES for the international scholarship to
perform part of his doctoral research at the University of Illinois Urbana-Champaign (CAPES –
proc. 071/2012).
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