Content uploaded by Carolina Ribeiro Silva
Author content
All content in this area was uploaded by Carolina Ribeiro Silva on Jun 29, 2023
Content may be subject to copyright.
Full Terms & Conditions of access and use can be found at
https://www.tandfonline.com/action/journalInformation?journalCode=tnzb20
New Zealand Journal of Botany
ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/tnzb20
New species of Jahnula (Jahnulales,
Dothideomycetes) and new records of aquatic
ascomycetes from estuarine beaches of the
Brazilian Amazon
Carolina Ribeiro Silva & Luis Fernando Pascholati Gusmão
To cite this article: Carolina Ribeiro Silva & Luis Fernando Pascholati Gusmão (2023): New
species of Jahnula (Jahnulales, Dothideomycetes) and new records of aquatic ascomycetes
from estuarine beaches of the Brazilian Amazon, New Zealand Journal of Botany, DOI:
10.1080/0028825X.2022.2157289
To link to this article: https://doi.org/10.1080/0028825X.2022.2157289
Published online: 15 Jan 2023.
Submit your article to this journal
View related articles
View Crossmark data
RESEARCH ARTICLE
New species of Jahnula (Jahnulales, Dothideomycetes) and
new records of aquatic ascomycetes from estuarine beaches
of the Brazilian Amazon
Carolina Ribeiro Silva
a
and Luis Fernando Pascholati Gusmão
b
a
Departamento de Micologia, Programa de Pós graduação em Biologia de Fungos, Universidade Federal de
Pernambuco, Recife, Brazil;
b
Departamento de Biologia, Programa de Pós Graduação em Botânica,
Universidade Estadual de Feira de Santana, Feira de Santana, Brazil
ABSTRACT
Estuaries are considered promising habitats for the presence of
marine and freshwater fungal species because of their favourable
environmental conditions. Janhula paraensis sp. nov. was found
on decaying wood collected in the intertidal zone in fluvial
beaches of Mosqueiro Island, Pará, Brazil. The new species differs
from others in the genus by its morphology including dark brown
ascospores, with thick and dark bands at septum when mature, a
hyaline cap at both ends and the absence of a mucilaginous
sheath. It was not possible to obtain molecular data for the new
species. New records of Boerlagiomyces websteri,Corollospora
pseudopulchella,Falciformispora lignatilis, and Savoryella lignicola
for South America and Annulusmagnus triseptatus for Brazil are
reported. Morphological descriptions and illustrations are
presented for all species.
ARTICLE HISTORY
Received 4 May 2022
Accepted 8 December 2022
First published online
16 January 2023
HANDLING EDITOR
Eric McKenzie
KEYWORDS
Fungal diversity; estuaries;
taxonomy; tropical
ascomycetes; wood
Introduction
Estuaries are coastal regions where rivers flow into the sea. These areas are influenced by
tides and salinity gradients and encompass several landforms, such as coastal bays,
lagoon inlets, and marshes (Pritchard 1952; Lessa et al. 2018). Estuaries are considered
a promising ecosystem type for fungal growth based on suitable temperatures and pres-
ence of diverse lignicolous substrates washed away by the tide (Raghukumar 2017).
The coastal zone of the Brazilian Amazon is characterised by high annual precipitation
and temperature regimes, discharge from dozens of estuaries, and high levels of nutrients
and organic matter (Pereira et al. 2009). The Paraense Coastal Zone is delimited into
three sectors according to physiographic characteristics, the Atlantic Coastal Salgado
Paraense, Insular Estuarine, and Continental Estuarine sectors (MMA 1996). The
latter sector is highly influenced by the Guajará Bay tides and the Pará river. Mosqueiro
Island is in the Continental Estuarine sector, in northeast Pará State. The predominant
vegetation types along river banks are dense ombrophilous forest and open alluvial
forest and the region has many estuarine sandy beaches subjected to meso-marine pro-
cesses (Lima et al. 2018).
© 2022 The Royal Society of New Zealand
CONTACT Carolina Ribeiro Silva crsilvabio@gmail.com
NEW ZEALAND JOURNAL OF BOTANY
https://doi.org/10.1080/0028825X.2022.2157289
Aquatic fungi live on submerged or partially submerged substrates and possess mor-
phological characteristics that help their dispersion in this environment (Shearer 1993;
Vijaykrishna et al. 2006).
Environmental factors interfere with the diversity of the aquatic fungal community
(Shearer 1972), especially temperature, which is an important determinant of the geographic
distribution of marine fungi (Jones 2000). Most species are restricted to tropical or temper-
ate regions, although some species are cosmopolitan (Hyde et al. 2005;Pangetal.2013).
Currently, 1857 species of fungi (including 136 species of microsporidia) are known to
inhabit the marine environment. They are distributed among 769 genera, 226 families, 88
orders, and 7 phyla. They are found on various decomposing substrates, associated with
leaves, trunks, plant pneumatophores and mangrove sediments; endophytes isolated
from algae; and phytoplankton parasites (Jones et al. 2019). In Brazil, approximately
90 species of sexual ascomycetes have been reported in the aquatic environment, includ-
ing 39 species in the marine environment and 52 in freshwater environments of the
north-eastern (15) and Amazonian (37) regions (Kohlmeyer 1969,1976,1984; Kohl-
meyer and Kohlmeyer 1971; Hughes 1974; Booth 1983; Barbosa et al. 2008,2013a,
2013b; Zelski et al. 2011; Cortez et al. 2016; Canto et al. 2020).
Jahnulales is predominantly an order of freshwater species and is composed of Ali-
quandostipitaceae and Manglicolaceae (Robert et al. 2005). However, several species
belonging to Manglicolaceae are found in marine habitats. Jahnula, (Kirschst.) Kirschst.
the main genus, was introduced by Kirschstein (1936), based on the previously described
Melanopsamma Kirschst. The aims of the present study were to expand our understand-
ing of the diversity and distribution of sexual ascomycetes in estuaries of the Amazon
region, and to illustrate and describe a new species of Jahnula from Brazil.
Materials and methods
Decomposed twigs were collected in the intertidal zone on Mosqueiro Island beaches.
Samples were washed in the river water to remove excess sand, placed in zip-lock bags
and transported to the Mycology Laboratory of the State University of Feira de
Santana, BA. The samples were processed according to Castañeda-Ruiz et al. (2016)
and regularly examined within 12 months using a Leica EC3 stereomicroscope for obser-
vation and the isolation of the reproductive structures of ascomycetes. Anatomical sec-
tioning was performed manually, and slides were prepared in distilled water, cotton blue,
and Melzer’s reagent. The slides were examined using an Olympus BX51 microscope
with Nomarski interference optics and photomicrographs of the new records were
obtained. The plant substrates with the collected specimens were deposited in the Her-
barium of the Universidade Estadual de Feira de Santana, Brazil.
Results
One new species Janhula paraensis sp. nov. is proposed, and five new records were collected.
Boerlagiomyces websteri Shearer & J.L. Crane, Corollospora pseudopulchella Nakagiri &
Tokura, Falciformispora lignatilis K.D. Hyde and Savoryella lignicola E.B.G. Jones & R.A.
Eaton represent new records for South America and Annulusmagnus triseptatus (S.W.
Wong, K.D. Hyde & E.B.G. Jones) J. Campb. & Shearer is a new record for Brazil.
2C. R. SILVA AND L. F. P. GUSMÃO
Taxonomy
Jahnula paraensis C.R. Silva & Gusmão, sp. nov. Figure 1.
MycoBank: 845497.
Holotype −Brazil, Pará State: Belém, Mosqueiro island, 1
o
03’45’’S, 48
o
20’14’’W, 30
December 2020, on intertidal wood, coll. L.T. Carmo (HUEFS 255201).
Diagnosis −Differs from Jahnula bipileata by its smaller ascospores with rough or irre-
gularly striated walls.
Etymology −The epithet is in reference to the Brazilian state where the specimen was
collected.
Ascomata 360 −500 × 700 −800 μm, superficial to immersed in the substrate (wood),
solitary, globose to subglobose, membranous, dark brown to black, with hyphae attached
to the substratum. Neck 160 −520 × 57 −70 μm, dark brown to black. Peridium variable
in thickness, composed of hyaline to brown cells. Pseudoparaphyses 2.5 μm wide,
filiform, hyaline, unbranched, sparsely septate. Asci 270 −410 × 25 −30 μm, 8-spored,
bitunicate, fissitunicate, cylindrical, pedicellate, with or without an ocular chamber
and sometimes with a faint ring. Ascospores 47.5 −52.5 × 17.5 −20 μm, mostly uniseri-
ate, ellipsoid-fusiform, straight or curved, smooth, immature hyaline, when mature
reddish brown to brown, l-septate, constricted at the septum, upper cell slightly wider,
and more apiculate than the lower cell, with thick and dark band at septum when
mature, without sheath but with 3.7 −6.2 μm wide, hyaline caps at both ends.
Habitat −Intertidal dead wood.
Comments: The order Jahnulales was introduced to Dothideomycetes based on phylo-
genetic analyses (Pang et al. 2002). It includes Aliquandostipitaceae (Inderbitzin et al.
2001) composed of eight genera: Aliquandostipite Inderb., Ascagilis K.D. Hyde, Brachio-
sphaera Nawawi, Jahnula, Megalohypha A. Ferrer & Shearer, Neojahnula W. Dong,
H. Zhang & K.D. Hyde, Pseudojahnula W. Dong, H. Zhang & K.D. Hyde and Xylomyces
Goos, R.D. Brooks & Lamore and Manglicolaceae (Suetrong et al. 2011) with a single
genus, Manglicola Kohlm. & E. Kohlm.
Based on phylogenetic analyses and morphological examinations, Dong et al. (2020)
introduced two new genera, Neojahnula and Pseudojahnula to accommodate Jahnula
australiensis K.D. Hyde and J. potamophila K.D. Hyde & S.W. Wong, respectively.
They also resurrected Ascagilis to accommodate seven species, including four species
of Jahnula characterised by mucilaginous pads on the ascospores,a noticeable character-
istic of the genus not found in the proposed new species reinforcing the need for further
phylogenetic analyses to elucidate the systematic placement of J. systyla K.D. Hyde &
S.W. Wong as they believe it to be related to another genus. Thus, the genus Jahnula
sensu lato has 12 accepted species (Index Fungorum 2021).
J. paraensis exhibits characteristics common to the genus Jahnula, such as cylindrical asci
with 8 ellipsoid-fusiform, brown, l-septate ascospores with upper cell slightly wider, and
more apiculate than the lower cell, without sheath or appendages, but with hyaline caps
at both ends. The slightly wider, and more apiculate features of the upper cell than the
lower cell has also been observed in six other species,J.apiosporaA. Carter, Raja &
Shearer,J.aquatica(Kirschst.) Kirschst., J. bipileata Raja & Shearer,J.granulosaK.D.
Hyde & S.W. Wong,J.purpureaJ. Fourn., Raja & Shearer and J. rostrata Raja & Shearer
(Hyde and Wong 1999; Raja and Shearer 2006;Rajaetal.2008;Fournieretal.2015). The
NEW ZEALAND JOURNAL OF BOTANY 3
dimensions of the asci and ascospores of J. paraensis are similar to J. appendiculata Pinruan,
K.D. Hyde & E.B.G. Jones, however, the ascospores are characterised by a mucilaginous
sheath or prominent appendages (Pinruan et al. 2002). The only species with a hyaline
cap at both ends of the ascospore is J. bipileata, but the ascospores are smaller than those
of other species and the walls are rough or irregularly striated (Raja and Shearer 2006).
Figure 1. Jahnula paraensis (HUEFS 255201, holotype). A, Ascoma on the natural
substrate. B, Pseudoparaphyses. C–F, Asci. G–J, Ascospores. Arrow indicates caps. Scale bars A =
0.2 mm. B–D=20μm. E–J=10μm.
4C. R. SILVA AND L. F. P. GUSMÃO
New records for South America and Brazil
Sordariomycetes
Diaporthomycetidae, Annulatascales, Annulatascaceae
Annulusmagnus triseptatus (S.W. Wong, K.D. Hyde & E.B.G. Jones) J. Campb. &
Shearer, Mycologia 96 (4): 826 (2004). Figure 2.
Ascomata semi-immersed, solitary or in small groups, subglobose to globose, black. Peri-
dium 26–31 µm thick, composed of an outer layer of dark brown angular and irregular
cells, a middle layer of brown cells, and an inner layer of hyaline elongated cells. Para-
physes hyaline, simple, septate, 3.7–5 µm wide. Asci unitunicate, cylindrical, short ped-
icellate, non-amyloid apical ring, 2.5–3.7 µm long, bipartite, 106–131 × 7.5–10 µm.
Ascospores fusiform, smooth, hyaline, 3-septate, without a thin mucilaginous sheath,
15–21 × 5–7.5 µm.
Examined material: Brazil. Pará State: Belém, Mosqueiro island, 29 April 2019, on inter-
tidal wood, coll. L.T. Carmo (HUEFS 255192).
Known distribution: Australia (Wong et al. 1999), Brazil (this study), Brunei (Wong
et al. 1999; Fryar et al. 2004), Canada (Campbell and Shearer 2004), Costa Rica (Huhn-
dorf et al. 2004), France (Reblova et al. 2010), Hong Kong (Wong et al. 1999; Wong and
Hyde 2001), Malaysia (Ho et al. 2001), Norway (Nordén and Jordal 2015), USA (Camp-
bell and Shearer 2004), Thailand (Luo et al. 2015) and Venezuela (Campbell and Shearer
2004).
Comments: The monotypic genus Annulusmagnus (type species Annulatascus triseptatus)
was described from submerged freshwater wood in Brunei (Wong et al. 1999). The Brazi-
lian specimen shows, as in the holotype, the absence of the ascomata with a neck with a
hyaline apex, ascospores not constricted at the septa, and the absence of a mucilaginous
sheath. These characters, in addition to molecular analyses, supported the establishment
of a new genus to accommodate Annulatascus triseptatus (Campbell and Shearer 2004).
This species occurs in submerged freshwater wood in tropical (Shearer and Raja 2010)
and temperate (Nordén and Jordal 2015) regions and it has been also recorded on terres-
trial grasses and Cyperaceae (Wong and Hyde 2001). It occurs in the Americas, is equally
distributed in tropical and temperate regions, and has been recorded in Atlantic, Pacific,
and Indian Oceans samples. Li et al. (2003) reported eight new bioactive compounds,
annularins A–H, along with the known (-)-(S)p- hydroxyphenyllactic acid isolated from
extracts of A. triseptatus and showed that these compounds have antifungal and antibac-
terial activity. The species has four records in the collection at the Westerdijk Fungal Bio-
diversity Institute (CBS 127687, 127688, 128831, 131483), three records at the Mae Fah
Luang University Culture Collection (MFLUCC 170462, 170469, 181335), and two
records at the Illinois Natural History Survey (SMH 2359, 4832). It has 33 sequences depos-
ited in GenBank (ITS, LSU, SSU, Tef1, RPB2, and β-tubulin), which have been included in
phylogenetic studies (Raja et al. 2003;Huhndorfetal.2004; Miller and Huhndorf 2005;
Reblova et al. 2010; Luo et al. 2015,2019; Dayarathne et al. 2016;Vuetal.2019). This is
the first record of the species in Brazil.
Hypocreomycetidae, Microascales, Halosphaeriaceae
Corollospora pseudopulchella Nakagiri & Tokura., Transactions of the Mycological
Society of Japan 28 (4): 428 (1988). Figure 3.
NEW ZEALAND JOURNAL OF BOTANY 5
Ascomata superficial, solitary, globose to subglobose, occasionally in grains of sand,
smooth and metallic, short papillate, ostiolate, carbonaceous, black, 18 × 12.5 µm. Peri-
dium brown, cells more or less spherical, 6.2–7.5 µm thick. Asci deliquesce early.
Ascospores fusiform, slightly curved, hyaline, cell attenuated both ends, 7-septate, con-
stricted at the septa, 75–97.5 × 7.5–10 µm (excluding apical appendages). Filiform
Figure 2. Annulusmagnus triseptatus A, Ascomata surface. B, Longitudinal section with two layers of
ascomal wall. C, Paraphyses. D, Ascus. E, Apical ring bipartite. F, Detail pedicellate ascus. Scale bars A
= 0.2 mm. B, E, F = 10 µm. C, D = 20 µm.
6C. R. SILVA AND L. F. P. GUSMÃO
appendages, hyaline, in the middle region (15–25 µm long) and at the ends (7.5–15 µm
long).
Known distribution: Brazil (this paper), Cuba (Samón and Enríquez 2010), India
(Nambiar and Raveendran 2010), Iraq (Al-Saadoon and Abdullah 2001), Japan (Nakagiri
and Tokura 1987), Mexico (Velez et al. 2015), Puerto Rico (Nieves-Rivera and Santos-
Figure 3. Corollospora pseudopulchella A, B, Ascomata surface. C, Detail ascospore both ends. D–F,
Ascospores. Scale bars A, B = 0.2 mm. C = 10 µm. D–F = 20 µm.
NEW ZEALAND JOURNAL OF BOTANY 7
Flores 2005), South Africa (Steinke and Jones 1993; Steinke and Lubke 2003) and Thai-
land (Jones et al. 2006).
Examined material: Brazil, Pará State: Belém, Mosqueiro island, 10 August 2017, on
intertidal wood, coll. L.T. Carmo (HUEFS 255194).
Comments:Corollospora is one of the largest fungal genera in marine habitats, with
approximately 23 species (Kohlmeyer and Kohlmeyer 1979; Abdel-Wahab et al. 2009).
The most similar species to C. pseudopulchella is C. pulchella Kohlm., I. Schmidt &
N.B. Nair, but its ascospores are longer and rounded at both ends. C. colossus Nakagiri
& Tokura is also similar with respect to the septation and length of ascospores, however,
its ascospores are wider (Kohlmeyer and Volkmann-Kohlmeyer 1991).
C. pseudopulchella is found exclusively in marine habitats (mangrove, estuary, intertidal
zones and beaches) and sandy habitats (Steinke and Jones 1993; Steinke and Lubke 2003;
Nambiar and Raveendran 2010; Velez et al. 2015). It has been associated with wood
(Nieves-Rivera and Santos-Flores 2005; Nambiar and Raveendran 2010; Velez et al.
2015) and sea foam (Steinke and Lubke 2003). It has been recorded mostly in the tropical
zone, in the Atlantic, Pacific, and Indian Oceans. Studies of the metabolic potential of this
species are lacking. Two isolates have been deposited at the NITE Biological Resource
Centre (NBRC 32112, 32113) and six sequences have been deposited in GenBank
(LSU, SSU, and RPB1) and were included in phylogenetic analysis (Schoch et al.
2012). This is the first record of the species in South America.
Hypocreomycetidae, Savoryellales, Savoryellaceae
Savoryella lignicola E.B.G. Jones & R.A. Eaton, Trans. Br. Mycol. Soc. 52: 161 (1969).
Figure 4.
Ascomata immersed, partly immersed or superficial, subglobose or ellipsoidal, with a
long neck, ostiolate, papillate, dark brown, 125–255 × 100–162 µm. Peridium thin,
brown, cells forming textura angularis, 12.5–21.2 µm thick. Asci unitunicate, clavate to
cylindrical, slightly truncate apex, 100–170 × 15–20 µm. Ascospores ellipsoidal, versico-
loured, central cells darker and terminal cells hyaline, 3-septate when mature, not con-
stricted at the septa, 22.5–30 × 10–12.5 µm.
Known distribution: Andaman Islands (Chinnaraj 1993b), Australia (Hyde 1993), Brazil
(this study), Brunei (Hyde 1990; Ho et al. 1997), China (Cai et al. 2002; Luo et al. 2004),
Cuba (Enríquez et al. 2011), England (Jones and Eaton 1969), Egypt (Abdel-Aziz 2010),
Hong Kong (Ho et al. 1997), India (Udaiyan 1989; Maria and Sridhar 2003; Jadav and
Borse 2017), Indonesia (Hyde 1989), Iraq (Al-Saadoon and Abdullah 2001), Japan (Ho
et al. 1997;Tsuietal.2003), Macau (Vrijmoed et al. 1994), Malaysia (Hyde 1994; Alias
and Jones 2000), Maldives (Chinnaraj 1993a), Mauritius (Poonyth et al. 1999), Mexico
(Gonzalez et al. 2001), Philippines (Jones et al. 1988; Alias et al. 1999), Portugal (Figueira
and Barata 2007), Seychelles (Hyde and Goh 1998), Singapore (Leong et al. 1991; Jones and
Hyde 1992), South Africa (Ho et al. 1997), Sri Lanka (Koch 1982), Taiwan (Pang et al.
2011), Thailand (Jones and Hyde 1992; Sulistyowati et al. 2010), USA (Shearer and Von
Bodman 1983), and U.K. (Hyde 1993).
Material examined: Brazil, Pará State: Belém, Mosqueiro island, 10 August 2017, on
intertidal wood, coll. L.T. Carmo (HUEFS 255195).
Comments: In all 13 species of the genus Savoryella (Jaklitsch and Réblova 2015; Zhang
et al. 2019) the ascospores are ellipsoidal, 3-septate, with two brown central cells and two
8C. R. SILVA AND L. F. P. GUSMÃO
hyaline ends cells (Jones et al. 2016). The ascospores of S. lignicola are similar to those of
S. aquatica K.D. Hyde, S. fusiformis W.H. Ho, K.D. Hyde & Hodgkiss, and S. longispora
E.B.G. Jones & K.D. Hyde, but differ in dimensions (Jones and Hyde 1992; Ho et al.
1997). S. yunnanensis Dong Wei, Dayar. & K.D. Hyde has larger asci, and ellipsoidal
or irregular and wider ascospores (Dayarathne et al. 2019) than those of S. lignicola.
The study species has been recorded in wood from marine and freshwater habitats
Figure 4. Savoryella lignicola A, Ascomata surface. B, C, Longitudinal section of ascomal wall. D, Ascus
immature. E–H, Ascospores. Scale bars A = 0.2 mm. B = 100 µm. C, E–H = 20 µm. D = 20 µm.
NEW ZEALAND JOURNAL OF BOTANY 9
(Ho et al. 1997; Poonyth et al. 1999), mangrove roots (Hyde 1990), seawater (Abdel-Aziz
2010) and brackish water-cooling towers (Jones and Eaton 1969; Eaton and Jones 1971a,
1971b) and has a wide distribution (Dayarathne et al. 2019). It is rarely recorded in the
Americas but has been reported in the Atlantic, Pacific, and Indian Oceans. S. lignicola
produces the extracellular cellulolytic enzymes endoglucanase, cellobiohydrolase, and β-
glucosidase (Pointing et al. 1999). Cellulase and xylanase for wood decomposition were
detected in an in vitro assay (Bucher et al. 2004). One isolate of the species is deposited in
the BIOTEC Culture Collection (NF 00204) and one in the National Taiwan Ocean Uni-
versity (NTOU 791). There are 11 sequences deposited in GenBank (ITS, LSU, SSU, Tef1,
and RPB2), and phylogenetic studies have been reported (Boonyuen et al. 2011; Reblova
et al. 2016; Dayarathne et al. 2019). This is the first record of the species in South
America.
Dothideomycetes
Tubeufiales, Tubeufiaceae
Boerlagiomyces websteri Shearer and J.L. Crane, Mycologia 87: 876 (1996). Figure 5.
Ascomata superficial, solitary, obpyriform, brown, 490–550 × 335–350 µm. Setae dark
brown. Peridium with two layers; outer layer 25–46 um thick, textura angularis,
brown and inner layer 5–15 um thick, fusiform cells, hyaline. Pseudoparaphyses 2.5–
3.7 µm wide, may be constricted at the septa. Asci fissitunicate, two spores, cylindrical,
rounded at the apex, short base, 157–260 × 25–52 µm. Ascospores oblong, rounded at
the ends, at first hyaline becoming dark brown, dictyosporous, nine transverse septa
and one to three vertical septa, surrounded by a gelatinous sheath mainly in the initial
stages, 72.5–117 × 22.5–50 µm.
Known distribution: Brazil (this study), Hungary (Toth 2009), USA (Shearer and Crane
1995; Raja et al. 2009) and Thailand (Kodsueb et al. 2006).
Material examined: Brazil, Pará State: Belém, Mosqueiro island, 29 April 2019, on inter-
tidal wood, coll. L.T. Carmo (HUEFS 255196).
Comments: The genus Boerlagiomyces includes eight species with muriform brown
ascospores and bristles on the walls of the ascomata (Kodsueb et al. 2006; Pande
2008). B. websteri differs from B. velutinus (Penz. & Sacc.) Butzin in shape, size,
number of septa, and colouration of the ascospores, while B. effusus (Syd., P. Syd. &
E.J. Butler) J.L. Crane, Shearer & M.E. Barr has ascomata with subiculum and ascus
with eight obovoid ascospores (Shearer and Crane 1995). The newly found specimen
had slightly larger asci than those of the type specimen (66–92(–107) × 26–35 µm).
B. websteri has been recorded in woody and freshwater plant detritus (Shearer and
Crane 1995; Raja et al. 2009), and decomposing cherry fruits (Toth 2009). There are
only two records of the species including this one, in tropical regions and it has
been recorded in Atlantic and Indian Oceans. B. websteri produces cellulase, endoxyla-
nase, β-xylosidase, polyphenol oxidase, peroxidase, tyrosinase, and laccase (Abdel-
Raheem and Shearer 2002). This species has a specimen deposited in the BIOTEC
Culture Collection (BCC 3834), only one sequence deposited in GenBank (LSU), and
a phylogenetic study (Kodsueb et al. 2006). This is the first record of the species in
South America.
Pleosporomycetidade, Pleosporales, Trematosphaeriaceae
Falciformispora lignatilis K.D. Hyde, Mycological Research 96(1):27 (1992). Figure 6.
10 C. R. SILVA AND L. F. P. GUSMÃO
Ascomata erumpent and eventually superficial, solitary or gregarious, subglobose, ostiolate,
not very evident papilla, black, 80–225 × 142–300 µm. Peridium 21–31 µm thick, compris-
ing an inner layer of elongate hyaline cells and an outer layer of angular to rounded brown
cells. Pseudoparaphyses septate, up to 2.5 µm wide. Asci fissitunicate, clavate to cylindrical,
short pedunculate, 127.5–162 × 25–30 µm. Ascospores fusiform, straight or slightly curved,
Figure 5. Boerlagiomyces websteri A, Ascomata surface. B, Longitudinal section of ascomal
wall. C, Ascus with two immature ascospores. D, Ascus with two mature ascospores. E–H, Ascospores.
Scale bars A = 0.2 mm. B = 20 µm. C–H=50µm.
NEW ZEALAND JOURNAL OF BOTANY 11
hyaline, 6-septate, slightly constricted at septa, surrounded by a thin mucilaginous sheath,
with a single scythe-like appendage 62.5–75 × 15–17.5 µm.
Known distribution: Brazil (this study), India (Devadatha and Sarma 2018), Mexico
(Hyde 1992), Taiwan (Pang and Jheng 2012), Thailand (Suetrong et al. 2009) and USA
(Raja and Shearer 2008).
Figure 6. Falciformispora lignatilis A, Ascomata surface. B, C, Longitudinal section of ascomal
wall. D, Ascus. E–H, Ascospores. Arrow indicates single scythe-like appendage. Scale bars A = 0.2
mm. B = 50 µm. C = 10 µm. D–H=20µm.
12 C. R. SILVA AND L. F. P. GUSMÃO
Material examined: Brazil, Pará State: Belém, Mosqueiro island, 10 August 2017, on
intertidal wood, coll. L.T. Carmo (HUEFS255197).
Comments: Hyde (1992) described F. lignatilis from intertidal mangrove wood col-
lected in Mexico. Falciformispora comprises five species, including two human patho-
gens (Ahmed et al. 2014; Eldridge et al. 2014; Hyde et al. 2020). Based on
morphological and phylogenetic characters, Suetrong et al. (2011a) proposed a new
family Trematosphaeriaceae and included the genus Falciformispora, based on SSU,
LSU, TEF-1-alpha, and RPB2 markers. F. lignatilis is similar to F. aquatica D.F. Bao,
K.D. Hyde & H.Y. Su, however, its asci and ascospores are larger and its appendage
is at the apical part of the ascospore (Hyde et al. 2019). F. uttaraditense Boonmee &
Huanraluek does not have a sickle-shaped appendage (Hyde et al. 2020). The dimen-
sions of the newly found specimen are larger than those described by Hyde (1992),
but consistent with observations by Raja and Shearer (2008) of freshwater material col-
lected in Florida (USA); this variation can probably be explained by sample site charac-
teristics. The species has been recorded on wood in freshwater (Raja and Shearer 2008)
and on mangrove (Hyde 1992; Devadatha and Sarma 2018). Other features like the
presence of discrete papilla and gel matrix integrating asci and pseudoparaphyses,
reported by Raja and Shearer (2008), Suetrong et al. (2011a), and Devadatha and
Sarma (2018), were not described in Hyde (1992). Most records are for tropical
regions and are related to the Atlantic and Indian Oceans. There is no study of the
metabolic potential of the genus. Two isolates have been deposited in the BIOTEC
Culture Collection (BCC 21117, 21118) and ten sequences have been deposited in
GenBank (ITS, LSU, SSU, and TEF-1-alpha), with some phylogenetic studies (Schoch
et al. 2009; Suetrong et al. 2009,2011a; Ahmed et al. 2014). This is the first record
of the species in South America.
Discussion
All species of Jahnula commonly occur in submerged wood in freshwater environments
(Suetrong et al. 2011a; Huang et al. 2018). The genus has a subtropical and tropical dis-
tribution, except for J. apiospora and J. dianchia S.K. Huang & K.D. Hyde,which are
restricted to temperate regions (Huang et al. 2018; Dong et al. 2020). Only
J. appendiculata has been recorded in South America in Peru (Shearer et al. 2015).
Prior to that, only Ascagilis seychellensis (K.D. Hyde & S.W. Wong) W. Dong, Doilom
& K.D. Hyde (as Jahnula seychellensis) had been recorded in South America in Brazil
(Barbosa et al. 2013a).
Representatives of Dothideomycetes occur on a variety of marine substrates, mainly
woody, in intertidal and mangrove regions, and may have an active mechanism to dis-
perse ascospores (Suetrong et al. 2009), as in Boerlagiomyces websteri and Falciformispora
lignatilis.
In Sordariomycetes, the family Halosphaeriaceae (containing Corollospora pseudopul-
chella) is the largest among marine ascomycetes (Maharachchikumbura et al. 2016).
Among species found in this study, only C. pseudopulchella is considered an obligate
marine fungus, together with S. lignicola, which has also been described in marine habi-
tats. All other species are considered facultative marine or freshwater fungi and are
recorded in estuaries for the first time.
NEW ZEALAND JOURNAL OF BOTANY 13
Acknowledgments
The authors thank Luana T. do Carmo for collecting the material on Mosqueiro Island, Dr. Huzefa
Raja from University of North Carolina at Greensboro, for comments and help and Taimy Cantillo
(PPGM/UEFS) for revision and comments. The Programa de Pós-graduação em Biologia de
Fungos (PPGBF/UFPE) and the Conselho Nacional de Desenvolvimento Científico e Tecnológico
Development (CNPq) are thanked for financial support and grant (proc.140871/2018-7 and proc.
312984/2018-9, respectively).
Disclosure statement
No potential conflict of interest was reported by the author(s).
Funding
This work was supported by CNPQ [grant no proc. 312984/2018-9,proc.140871/2018-7].
ORCID
Carolina Ribeiro Silva http://orcid.org/0000-0001-8922-9159
Luis Fernando Pascholati Gusmão http://orcid.org/0000-0003-3288-3971
References
Abdel-Aziz FA. 2010. Marine fungi from two sandy Mediterranean beaches on the Egyptian north
coast. Botanica Marina. 53(3):283–289.
Abdel-Raheem A, Shearer CA. 2002. Extracellular enzyme production by freshwater ascomycetes.
Fungal Diversity. 11:1–19.
Abdel-Wahab MA, Nagahama T, Abdel-Aziz FA. 2009. Two new Corollospora species and one
new anamorph based on morphological and molecular data. Mycoscience. 50(3):147–155.
Ahmed SA, Van de Sande WWJ, Stevens DA, Fahal A, Van Diepeningen AD, Menken SBJ, de
Hoog GS. 2014. Revision of agents of black-grain eumycetoma in the order Pleosporales.
Persoonia. 33:141–154.
Alias SA, Jones EBG. 2000. Colonization of mangrove wood by marine fungi at Kuala Selangor
mangrove stand, Malaysia. Fungal Diversity. 5:9–21.
Alias SA, Jones EBG, Torres J. 1999. Intertidal fungi from the Philippines, with a description of
Acrocordiopsis sphaerica sp. nov. (Ascomycota). Fungal Diversity. 2:35–41.
Al-Saadoon AH, Abdullah SK. 2001. Some interesting ascomycetes from Iraq. Iraqi Journal of
Biology. 1:125–134.
Barbosa FR, Gusmão LFP, Raja HA, Shearer CA. 2008.Annulatascus apiculatus sp. nov., a new
freshwater ascomycete from the semi-arid Caatinga biome of Brazil. Mycotaxon. 106:403–407.
Barbosa FR, Gusmão LFP, Raja HA, Shearer CA. 2013a. New species and new records of fresh-
water ascomycetes from Brazil and Costa Rica. Mycologia. 105(2):335–343.
Barbosa FR, Raja HA, Shearer CA, Gusmão LFP. 2013b. Some freshwater fungi from the Brazilian
semi-arid region, including two new species of hyphomycetes. Cryptogamie, Mycologie. 34
(3):243–258.
Boonyuen N, Chuaseeharonnachai C, Suetrong S, Sri-Indrasutdhi V, Sivichai S, Jones EBG, Pang
KL. 2011. Savoryellales (Hypocreomycetideae, Sordariomycetes): a novel lineage of aquatic
ascomycetes inferred from multiple-gene phylogenies of the genera Ascotaiwania,
Ascothailandia and Savoryella. Mycologia. 103:1351–1371.
Booth T. 1983. Lignicolous marine fungi from São Paulo, Brazil. Canadian Journal of Botany. 61
(2):488–506.
14 C. R. SILVA AND L. F. P. GUSMÃO
Bucher VVC, Pointing SB, Hyde KD, Reddy CA. 2004. Production of wood decay enzymes, loss of
mass, and lignin solubilization in wood by diverse tropical freshwater fungi. Microbial Ecology.
48(3):331–337.
Cai L, Tsui CKM, Zhang KQ, Hyde KD. 2002. Aquatic fungi from lake fuxian, yunnan, China.
Fungal Diversity. 9:57–70.
Campbell J, Shearer CA. 2004.Annulusmagnus and Ascitendus, two new genera in the
Annulatascaceae. Mycologia. 96(4):822–833.
Canto ESM, Cortez ACA, Monteiro JS, Barbosa FR, Zelski S, Souza JVBD. 2020. Composition and
diversity of fungal decomposers of submerged wood in two lakes in the Brazilian Amazon State
of Para. International Journal of Microbiology. 2020:1–9.
Castañeda-Ruiz RF, Heredia G, Gusmão LFP, Li DW. 2016. Fungal diversity of central and South
America. In: Li D.W, editor. Biology of microfungi. Switzerland: Springer International
Publishing; p. 197–217.
Chinnaraj S. 1993a. Manglicolous fungi from atolls of Maldives, Indian ocean. Indian Journal of
Marine Sciences. 22(2):141–142.
Chinnaraj S. 1993b. Higher marine fungi from mangroves of Andaman and Nicobar Islands.
Sydowia. 45(1):109–115.
Cortez ACA, Sanches MA, Zelski SE, Souza JVBD. 2016. A comparison of the freshwater fungal
community during the non-rainy and rainy seasons in a small black water lake in Amazonas,
Brazil. Journal of Food, Agriculture and Environment. 14(2):156–161.
Dayarathne MC, Maharachchikumbura SSN, Jones EBG, Dong W, Devadatha B, Yang J,
Ekanayaka AH, De Silva W, Sarma VV, Al-Sadi AM, et al. 2019. Phylogenetic revision of
Savoryellaceae and evidence for its ranking as a subclass. Frontiers in Microbiology. 10
(840):1–26.
Dayarathne MC, Maharachchikumbura SSN, Phookamsak R, Fryar SC, To-anun C, Jones EBG,
Al-Sadi AM, Zelski SE, Hyde KD. 2016. Morpho-molecular characterization and epitypification
of Annulatascus velatisporus. Mycosphere. 9:1389–1398.
Devadatha B, Sarma VV. 2018.Pontoporeia mangrovei sp. nov, a new marine fungus from an
Indian mangrove along with a new geographical and host record of Falciformispora lignatilis.
Current Research in Environmental & Applied Mycology. 8(2):238–246.
Dong W, Wang B, Hyde KD, McKenzie EHC, Bhat DJ, Raja HA, Tanaka K, Abdel-Wahab MA,
Abdel-Aziz FA, Doilom M, et al. 2020. Freshwater dothideomycetes. Fungal Diversity.
105:319–575.
Eaton RA, Jones EBG. 1971a. The biodeterioration of timber in water–cooling towers. I. Fungal
ecology and the decay of wood at Connah’s Quay and Ince. Material und Organismen. 6:51–
80.
Eaton RA, Jones EBG. 1971b. The biodeterioration of timber in water-cooling towers. II. Fungi
growing on wood in different positions in a water-cooling system. Material und Organismen.
6:81–92.
Eldridge ML, Chambers CJ, Sharon VR, Thompson GR. 2014. Fungal infections of the skin and
nail: new treatment options. Expert Review of Anti-Infective Therapy. 12(11):1389–1405.
Enríquez DI, Gonzáez MC, Nuñez R, Delgado Y. 2011. Ascomicetes marinos aislados de playas en
la zona occidental de Cuba. Revista del Jardín Botánico Nacional. 32/33:293–298.
Figueira D, Barata M. 2007. Marine fungi from two sandy beaches in Portugal. Mycologia. 99
(1):20–23.
Fournier J, Raja HA, Shearer CA. 2015. Freshwater Ascomycetes: Jahnula purpurea (Jahnulales,
Dothideomycetes), a new species on submerged wood from Martinique Island, Lesser
Antilles. MycoKeys. 9:29–36.
Fryar SC, Booth W, Davies J, Hodgkiss IJ, Hyde KD. 2004. Distribution of fungi on wood in the
tutong river, brunei. Fungal Diversity. 17:17–38.
Gonzalez MC, Hanlin RT, Ulloa M. 2001. A checklist of higher marine fungi of Mexico.
Mycotaxon. 80:241–253.
Ho WH, Hyde KD, Hodgkiss IJ. 1997. Ascomycetes from tropical freshwater habitats: the genus
Savoryella, with two new species. Mycological Research. 101:803–809.
NEW ZEALAND JOURNAL OF BOTANY 15
Ho WH, Hyde KD, Hodgkiss IJ. 2001. Fungal communities on submerged wood from streams in
Brunei, Hong Kong, and Malaysia. Mycological Research. 105(12):1492–1501.
Huang SK, Maharachchikumbura SSN, Jeewon R, Bhat DJ, Chomnunti P, Hyde KD, Lumyong S.
2018. Morphological and molecular taxonomy of Jahnula dianchia sp. nov. (Jahnulales) from
submerged wood in Dianchi Lake, Yunnan China. Mycological Progress. 17:547–555.
Hughes GC. 1974. Geographical distribution of the higher marine fungi. Veroeff. Inst.
Meeresforsch. Bremerhaven. 5:419–441.
Huhndorf SM, Miller AN, Fernández FA. 2004. Molecular systematics of the Sordariales: the order
and the family Lasiosphaeriaceae redefined. Mycologia. 96(2):368–387.
Hyde KD. 1989. Intertidal mangrove fungi from north Sumatra. Canadian Journal of Botany.
67:3078–3082.
Hyde KD. 1990. A study of the vertical zonation of intertidal fungi on Rhiziphora apiculata at
Kampong Kapok mangrove, Brunei. Aquatic Botany. 36(3):255–262.
Hyde KD. 1992. Intertidal mangrove fungi from the west coast of Mexico, including one new
genus and two new species. Mycological Research. 96:25–30.
Hyde KD. 1993. Tropical Australian freshwater fungi. V. Bombardia sp., Jahnula australiensis sp.
nov., Savoryella Lignicola and S. Aquatica sp. nov. Australian Systematic Botany. 5:161–167.
Hyde KD. 1994. The genus Savoryella from freshwater habitats, including S. grandispora sp. nov.
Mycoscience. 35:59–61.
Hyde KD, Cai L, Jeewon R. 2005. Tropical fungi. In: Dighton J, White JF, Oudemans P, editor. The
Fungal Community: its organization and role in the ecosystem. New York: CRC Press;
p. 93–109.
Hyde KD, Dong Y, Phookamsak R, Jeewon R, Bhat DJ, Jones EBG, Liu NG, Abeywickrama PD,
Mapook A, Wei D, et al. 2020. Fungal diversity notes 1151–1276: taxonomic and phylogenetic
contributions on genera and species of fungal taxa. Fungal Diversity. 100:5–277.
Hyde KD, Goh TK. 1998. Fungi on submerged wood in the riviere St marie-louis, the Seychelles.
South African Journal of Botany. 64(6):330–336.
Hyde KD, Tennakoon DS, Jeewon R, Bhat DJ, Maharachchikumbura SSN, Rossi W, Leonardi M,
Lee HB, Mun HY, Houbraken J, et al. 2019. Fungal diversity notes 1036–1150: taxonomic
and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity.
96:1–242.
Hyde KD, Wong SW. 1999. Tropical Australian freshwater Fungi. XV. The ascomycete genus
Jahnula, with five new species and one new combination. Nova Hedwigia. 68(3–4):489–509.
Inderbitzin P, Landvik S, Abdel-Wahab MA, Berbee ML. 2001. Aliquandostipitaceae, a new family
for two new tropical ascomycetes with unusually wide hyphae and dimorphic ascomata.
American Journal of Botany. 88:52–61.
Index Fungorum. 2021. Index Fungorum. Access: 14/12/2021. http://www.indexfungorum.org/
names/Names.asp.
Jadav RT, Borse KN. 2017. Some fresh water ascomycetes from Nagpur district of Maharashtra,
India. Bioscience Discovery. 8(3):422–425.
Jaklitsch WM, Réblova M. 2015.Savoryellaceae. Index Fungorum. 209:1.
Jones EBG. 2000. Marine fungi: some factors influencing biodiversity. Fungal Diversity. 4:53–73.
Jones EBG, Eaton RA. 1969. Savoryella lignicola gen. et sp.nov. from water-cooling towers.
Transactions of the British Mycological Society. 52:161–174.
Jones EBG, Hyde KD. 1992. Taxonomic studies on savoryella jones et eaton (ascomycotina).
Botanica Marina. 35:83–91.
Jones EBG, Pang KL, Abdel-Wahab MA, Scholz B, Hyde KD, Boekhout T, Ebel R, Rateb ME,
Henderson L, Sakayaroj J, et al. 2019. An online resource for marine fungi. Fungal Diversity.
96:347–433.
Jones EBG, Pilantanapak A, Chatmala I, Sakayaroj J, Phongpaichit S, Choeyklin R. 2006. Thai
marine fungal diversity. Songklanakarin Journal of Science and Technology. 28(4):687–708.
Jones EBG, To-anun C, Suetrong S, Boonyuen N. 2016. Mycosphere Essays 12. Progress in the
classification of the watercooling tower ascomycete Savoryella and a tribute to John Savory: a
review. Mycosphere. 7(5):570–581.
16 C. R. SILVA AND L. F. P. GUSMÃO
Jones EBG, Uyenco FR, Follosco MP. 1988. Fungi on driftwood collected in the intertidal zone
from the Philippines. Asian Mar. Biol. 5:103–106.
Kirschstein W. 1936. Beiträge zur Kenntnis der Ascomyceten und ihrer Nebenformen besonders
aus der Mark Brandenburg und dem Bayerischen Walde. Annales Mycologici. 34:180–210.
Koch J. 1982. Some lignicolous marine fungi from Sri Lanka. Nordic Journal of Botany. 2(2):163–
169.
Kodsueb R, Jeewon R, Vijaykrishna D, McKenzie EHC, Lumyong P, Lumyong S, Hyde KD. 2006.
Systematic revision of Tubeufiaceae based on morphological and molecular data. Fungal
Diversity. 21:105–130.
Kohlmeyer J. 1969. Marine fungi of Hawaii including the new genus Helicascus. Canadian Journal
of Botany. 47:1469–1487.
Kohlmeyer J. 1976. Marine fungi from South America. Mitteilungen aus dem Instituto Colombo-
Alemán de Investigaciones Científicas Punta de Betín. 8:33–39.
Kohlmeyer J. 1984. Tropical marine fungi. Marine Ecology. 5(4):329–378.
Kohlmeyer J, Kohlmeyer E. 1971. Marine fungi from tropical America and Africa. Mycologia. 63
(4):831–861.
Kohlmeyer J, Kohlmeyer E. 1979. Marine mycology. The higher fungi. New York (NY): Academic
Press; p. 48–54.
Kohlmeyer J, Volkmann-Kohlmeyer B. 1991. Illustrated key to the filamentous higher marine
fungi. Botanica Marina. 34(1):1–61.
Leong WF, Tan TK, Jones EBG. 1991. Fungal colonization of submerged Bruguiera cylindrica and
Rhizophora apiculata wood. Botanica Marina. 34:69–76.
Lessa GC, Santos FM, Souza Filho PW, Corrêa-Gomes LC. 2018. Brazilian estuaries: a geomorpho-
logic and oceanographic perspective. In: Brazilian Estuaries. Berlin (DE): Springer; p. 1–37.
Li C, Nitka MV, Gloer JB, Campbell J, Shearer CA. 2003. Annularins A−H: new polyketide metab-
olites from the freshwater aquatic fungus Annulatascus triseptatus. Journal of Natural Products.
66(10):1302–1306.
Lima LC, Neto ABB, Santos CRC, Braga AN, Nunes SCT. 2018. Longterm temporal analysis of
vegetation cover and soil use on Ilha de Mosqueiro, Belém, Pará, Amazonian Brazil. Revista
agro@mbiente On-line-line. 12(1):80–88.
Luo J, Yin JF, Cai L, Zhang KQ, Hyde KD. 2004.Freshwater fungi in Lake Dianchi, a heavily pol-
luted lake in Yunnan, China. Fungal Diversity. 16:93–112.
Luo ZL, Hyde KD, Liu JK, Maharachchikumbura SSN, Jeewon R, Bao DF, Bhat DJ, Lin CG, Li WL,
Yang J, et al. 2019. Freshwater sordariomycetes. Fungal Diversity. 99:451–660.
Luo ZL, Maharachchikumnura SSN, Liu XY, Li SH, Chen LJ, Su HY, Zhou DQ, Hyde KD. 2015.
Annulatascus saprophyticus sp. nov. and Pseudoannulatascus gen. nov. to accommodate
Annulatascus biatriisporus (Annulatascales Sordariomycetes) from Thailand. Phytotaxa. 239
(2):174–182.
Maharachchikumbura SN, Hyde KD, Jones EBG, McKenzie EHC, Jayarama B, Dayarathne M,
Huang SK, Norphanphoun C, Senanayake IC, Perera RH, et al. 2016. Families of sordariomy-
cetes. Fungal Diversity. 79:1–317.
Maria GL, Sridhar KR. 2003. Diversity of filamentous fungi on woody litter of five mangrove plant
species from the southwest coast of India. Fungal Diversity. 14:109–126.
Miller AN, Huhndorf SM. 2005. Multi-gene phylogenies indicate ascomal wall morphology is a
better predictor of phylogenetic relationships than ascospore morphology in the Sordariales
(Ascomycota, Fungi). Molecular Phylogenetics and Evolution. 35(1):60–75.
MMA. 1996. Perfil dos estados litorâneos do Brasil: subsídios à implantação do Programa
Nacional de Gerenciamento Costeiro. Brasília DF: PNMA/GERCO, 301. (Série
Gerenciamento Costeiro v. 9).
Nakagiri A, Tokura R. 1987. Taxonomic studies of the genus Corollospora (Halosphaeriaceae
Ascomycotina) with descriptions of seven new species. Transactions of the Mycological
Society of Japan. 28(4):413–436.
Nambiar GR, Raveendran K. 2010. Frequency and abundance of arenicolous marine fungi along
south Indian beaches. Journal of Scientific Research. 2(1):138–143.
NEW ZEALAND JOURNAL OF BOTANY 17
Nieves-Rivera AM, Santos-Flores CJ. 2005. Hongos acuáticos de los estuarios en Puerto
Rico: Boca del Río Manatí. The Journal of Agriculture of the University of Puerto Rico. 89-
(1-2):97–105.
Nordén B, Jordal JB. 2015. A checklist of Norwegian sordariomycetes. Agarica. 36:55–73.
Pande A. 2008. Ascomycetes of peninsular India. Jodhpur: Scientific Publishers.
Pang KL, Abdel-Wahab MA, Sivichai S, El-Sharouney HM, Jones EBG. 2002. Jahnulales
(Dothideomycetes, Ascomycota): a new order of lignicolous freshwater ascomycetes.
Mycological Research. 106(9):1031–1042.
Pang KL, Jheng JS. 2012. A checklist of marine fungi of Taiwan with a description of Kitesporella
keelungensis gen. et sp. nov. Botanica Marina. 55(5):459–466.
Pang KL, Jheng JS, Jones EBG. 2011. Marine mangrove fungi of Taiwan. Keelung Taiwan: National
Taiwan Ocean University.
Pang KL, Vrijmoed LLP, Jones EBG. 2013. Genetic variation within the cosmopolitan aquatic
fungus Lignincola laevis (Microascales, Ascomycota). Organisms Diversity & Evolution. 13
(3):301–309.
Pereira LCC, Dias JA, Carmo JA, Polette M. 2009. A zona costeira amazônica brasileira. Revista de
Gestão Costeira Integrada-Journal of Integrated Coastal Zone Management. 9(2):3–7.
Pinruan U, Jones EBG, Hyde KD. 2002. Aquatic fungi from peat swamp palms: Jahnula appendi-
culata sp. nov. Sydowia. 54(2):242–247.
Pointing SB, Buswell JA, Jones EBG, Vrijmoed LLP. 1999. Extracellular cellulolytic enzyme profiles
of five lignicolous mangrove fungi. Mycological Research. 103(6):696–700.
Poonyth AD, Hyde KD, Peerally A. 1999. Intertidal fungi in Mauritian mangroves. Botanica
Marina. 42:243–252.
Pritchard DW. 1952. Estuarine hydrography. Advances in Geophysics. 1:243–280.
Raghukumar S. 2017. Fungi in coastal and oceanic marine ecosystems. New York, NY, USA:
Springer.
Raja HA, Campbell J, Shearer CA. 2003. Freshwater ascomycetes: Cyanoannulus petersenii, a new
genus and species from submerged wood. Mycotaxon. 88:1–17.
Raja HA, Carter A, Platt HW, Shearer CA. 2008. Freshwater ascomycetes: jahnula apiospora (jah-
nulales, dothideomycetes), a new species from Prince Edward Island, Canada. Mycoscience. 49
(5):326–328.
Raja HA, Schmit JP, Shearer CA. 2009. Latitudinal habitat and substrate distribution patterns of
freshwater ascomycetes in the Florida Peninsula. Biodiversity and Conservation. 18(2):419–455.
Raja HA, Shearer CA. 2006.Jahnula species from north and central America, including three new
species. Mycologia. 98(2):319–332.
Raja HA, Shearer CA. 2008. Freshwater ascomycetes: new and noteworthy species from aquatic
habitats in Florida. Mycologia. 100(3):467–489.
Reblova M, Fournier J, Hyde KD. 2010.Achroceratosphaeria, a new ascomycete genus in the
Sordariomycetes, and re-evaluation of Ceratosphaeria incolorata. Fungal Diversity. 43(1):75–84.
Reblova M, Seifert KA, Fournier J, Štěpánek V. 2016. Newly recognised lineages of perithecial
ascomycetes: the new orders Conioscyphales and Pleurotheciales. Persoonia - Molecular
Phylogeny and Evolution of Fungi. 37:57–81.
Robert V, Stegehuis G, Stalpers J. 2005. The MycoBank engine and related databases. https://www.
MycoBank.org/.
Samón E, Enríquez D. 2010. Nuevos registros de hongos marinos en el litoral sur de Guantánamo,
Cuba. Serie Oceanológica. 7:61–68.
Schoch CL, Crous PW, Groenewald JZ, Boehm EWA, Burgess TI, de Gruyter J, de Hoog GS, Dixon
LJ, Grube M, Gueidan C, et al. 2009. A class-wide phylogenetic assessment of Dothideomycetes.
Studies in Mycology. 64:1–15.
Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque CA, Chen W. 2012. The
internal transcribed spacer as a universal DNA barcode marker for fungi. Fungal Barcoding
Consortium. Proceedings of the National Academy of Sciences of the United States of
America. 109:6241–6246.
18 C. R. SILVA AND L. F. P. GUSMÃO
Shearer CA. 1972. Fungi of the Chesapeake Bay and its tributaries. III. The distribution of wood-
inhabiting ascomycetes and fungi imperfecti of the Patuxent Rive. American Journal of Botany.
59(9):961–969.
Shearer CA. 1993. The freshwater ascomycetes. Nova Hedwigia. 56:1–3.
Shearer CA, Crane JL. 1995.Boerlagiomyces websteri a new ascomycete from fresh water.
Mycologia. 87:876–879.
Shearer CA, Raja HA. 2010. Freshwater Ascomycetes. (http://fungi.life.illinois.edu).Acesso:10/01/
2020.
Shearer CA, Von Bodman SB. 1983. Patterns of occurrence of Ascomycetes associated with
decomposing twigs in a midwestern stream. Mycologia. 75(3):518–530.
Shearer CA, Zelski SE, Raja HA, Schmit JP, Miller AN, Janovec JP. 2015. Distributional patterns of
freshwater ascomycetes communities along an Andes to Amazon elevational gradient in Peru.
Biodiversity and Conservation. 24:1877–1897.
Steinke TD, Jones EBG. 1993. Marine and mangrove fungi from the Indian ocean coast of South
Africa. South African Journal of Botany. 59(4):385–390.
Steinke TD, Lubke RA. 2003. Arenicolous marine fungi from Southern Africa. South African
Journal of Botany. 69(4):540–545.
Suetrong S, Boonyuen N, Pang KL, Ueapattanakit J, Klaysuban A, Sriiindrasutdhi V, Sivichai S,
Jones EBG. 2011. A taxonomic revision and phylogenetic reconstruction of the Jahnulales
(Dothideomycetes), and the new family Manglicolaceae. Fungal Diversity. 51(1):163–188.
Suetrong S, Hyde KD, Ahang Y, Bahkali AH, Jones EBG. 2011a. Trematosphaeriaceae fam. nov.
(Dothideomycetes Ascomycota). Cryptogamie Mycologie. 32:343–358.
Suetrong S, Schoch CL, Spatafora JW, Kohlmeyer J, Volkmann-Kohlmeyer B, Sakayaroj J,
Phongpaichit S, Tanaka K, Hirayama K, Jones EBG. 2009. Molecular systematics of the
marine Dothideomycetes. Studies in Mycology. 64:155–173.
Sulistyowati L, Moslem MA, Kevin D. 2010. Diversity of freshwater ascomycetes in freshwater
bodies at amphoe Mae chan, chiang Rai. Cryptogamie Mycologie. 3(3):323–333.
Toth S. 2009.Boerlagiomyces websteri (Ascomycota Tubeufiaceae) from Hungary first record
outside the USA. Mycologia Balcanica. 6:85–86.
Tsui CKM, Hyde KD, Fukushima K. 2003. Fungi on submerged wood in the koito river, Japan.
Mycoscience. 44(1):55–59.
Udaiyan K. 1989. Some interesting ascomycetes from water cooling towers. Kavaka. 17:11–16.
Velez P, Gonzalez MC, Capello-Garcia S, Rosique-Gil E, Hanlin RT. 2015. Diversity of marine
ascomycetes from the disturbed sandy beaches of Tabasco Mexico. Journal of the Marine
Biological Association of the United Kingdom. 95(5):897–903.
Vijaykrishna D, Jeewon R, Hyde KD. 2006. Molecular taxonomy, origins and evolution of fresh-
water ascomycetes. Fungal Diversity. 23:351–390.
Vrijmoed LLP, Hyde KD, Jones EBG. 1994. Observations on mangrove fungi from Macau and
Hong Kong with the description of two new ascomycetes: Diaporthe salsuginosa and
Aniptodera haispora. Mycological Research. 98(6):699–704.
Vu D, Groenewald M, de Vries M, Gehrmann T, Stielow B, Eberhardt U, Al-Hatmi A, Groenewald
JZ, Cardinali G, Houbraken J, et al. 2019. Large-scale generation and analysis of filamentous
fungal DNA barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal
species and higher taxon delimitation. Studies in Mycology. 92:135–154.
Wong KM, Hyde KD. 2001. Diversity of fungi on six species of Gramineae and one species of
Cyperaceae in Hong Kong. Mycological Research. 105(12):1485–1491.
Wong SW, Hyde KD, Jones EBG, Moss ST. 1999. Ultrastructural studies on the aquatic ascomy-
cetes Annulatascus velatisporus and A. triseptatus sp. nov. Mycological Research. 103:561–571.
Zelski SE, Raja HA, Miller AN, Barbosa FR, Shearer CA. 2011.Longicollum biappendiculatum gen.
et sp. nov., a new freshwater ascomycete from the Neotropics. Mycosphere. 2:539–545.
Zhang SN, Abdel-Wahab MA, Jones EBG, Hyde KD, Liu JKJ. 2019. Additions to the genus
Savoryella (Savoryellaceae), with the asexual morphs Savoryella nypae comb. nov. and
S. Sarushimana sp. nov. Phytotaxa. 408(3):195–207.
NEW ZEALAND JOURNAL OF BOTANY 19
