Fungal Diversity

Classification of marine Ascomycota, anamorphic taxa and Basidiomycota

Jones, E.B.G.1*, Sakayaroj, J.1, Suetrong, S.1, 3, Somrithipol, S.1 and Pang, K.L.2

¹Bioresources Technology Unit, Phylogenetics Laboratory, National Center for Genetic Engineering and Biotechnology,
113 Paholyothin Road, Khlong 1, Khlong Luang, Pathum Thani 12120, Thailand
2
Institute of Marine Biology, National Taiwan Ocean University, No. 2 Pei-Ning Road, Keelung 20224, Taiwan
3
Department of Microbiology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla, 90112, Thailand.

Jones, E.B.G., Sakayaroj, J., Suetrong, S., Somrithipol, S. and Pang, K.L. (2009). Classification of marine Ascomycota,
anamorphic taxa and Basidiomycota. Fungal Diversity 35: 1-187.

A comprehensive classification of the filamentous marine fungi is outlined, with reference to recent molecular
phylogenetic analyses. The classification includes 530 species (in 321 genera) to order level: Ascomycota 424 species
(in 251 genera), anamorphic fungi 94 species (in 61 genera) and Basidiomycota 12 species (in 9 genera). The
Halosphaeriales is the largest order of marine fungi with 126 species in 53 genera, of which 35 are monotypic. Several
taxa are of uncertain position and cannot be assigned to any higher taxonomic ranks. The decadel index shows that most
marine fungi were described in the period 1980-1989 (135) and 1990-1999 (156), with 43 new species and 25 new
genera from the past eight years. Keys are provided to the major taxa, genera and species. One new species is described
in this paper.

Key words: fungal classification, marine fungi, molecular phylogeny, rDNA, new taxa

Article Information
Received 1 December 2008
Accepted 23 December 2008
Published online 24 March 2009
*Corresponding author: E.B.G. Jones; e-mail: remispora@gmail.com

Introduction reliable recent figure is that of Hyde et al.

(2000) who listed 444 “higher” obligate marine
Although a few synoptic keys are fungi. However, a number of taxa were not
available for the identification of obligate listed, while a further 43 new species in ten
marine fungi (Kohlmeyer and Volkmann- new genera have been described over the past
Kohlmeyer, 1991a; Hyde et al., 2000), there eight years. A further 15 new genera have been
has been no attempt to present a classification introduced to accommodate species rejected
for them since the book of Kohlmeyer and from existing genera. Other species not
Kohlmeyer (1979). This monograph addresses previously regarded as marine are included in
this issue. this monograph. The total now stands at 530
The number of marine fungi described and new taxa continue to be described from
has increased dramatically since the pioneer this habitat.
study of Barghoorn and Linder (1944) on ligni- Kohlmeyer and Kohlmeyer (1979) stated
colous species. Currently some 1,500 names that: “the decrease of new descriptions of
can be found in the literature, but many of marine fungi during the period 1970-1977
these are inadequately described, or may be indicates that the most common species have
facultative terrestrial species or synonyms of been named and that considerable additions of
existing taxa (Jones and Mitchell, 1996). new taxa in the future are unlikely.” However,
However, Schaumann (pers. comm.) estimates as new substrata and geographical locations are
there are some 6,000 marine species, but this being examined for fungi, the number of new
figure is not supported by data. The most species continues to rise (Fig. 1): 135 in 1980-

1
Fig. 1. Decadel increase in the number of marine fungi.

1989; 156 in 1990-1999 and 43 already marine Ascomycota: Decorospora gaudefroyi

recorded for the period 2000-2008. This is (as Pleospora gaudefroyi) (Yusoff et al.,
certainly noticeable for mangrove species (42 1994b), Julella avicenniae (Au et al., 1999a)
species in Kohlmeyer and Kohlmeyer, 1979; and the genus Massarina (Au et al., 2001;
over 170 in Hyde et al., 2000), and tropical Read et al., 1994, 1997a, b). Although
locations continue to yield a wide range of ultrastructural characters helped in the
new taxa (Jones et al., 2006; Koch et al., delineation of a number of genera (e.g.
2007). Kis-Papo (2005) has reviewed data on Corollospora) others were found to be more
the number of marine fungi and settles on 467 difficult to resolve (e.g. Halosarpheia).
species. Over the last decade, the use of
Identification and classification of the molecular techniques to examine the phylo-
marine higher fungi have followed traditional geny of organisms has advanced considerably
avenues of the evaluation and significance of (Spatafora and Blackwell, 1994). Molecular
morphological characters at the light micro- techniques have also been applied to examine
scope level. For the Ascomycota, ultra- the relationships of a number of marine taxa at
structural characters at the transmission and the ordinal level and at the genus/species level.
scanning electron microscope levels have also The earliest study was on the phylogeny of
been used (Jones, 1995). These were Halosphaeriopsis mediosetigera (as a repre-
specifically applied to ascospore appendage sentative of the Halosphaeriales), and the
ontogeny of the Halosphaeriales, where Microascales, with both orders sharing a
delineation of genera was questioned, e.g. common ancestor (Spatafora and Blackwell,
Corollospora (Kohlmeyer, 1972a; Jones et al., 1994). This relationship has also been
1983a), and Ceriosporopsis (Johnson et al., supported by other studies (Chen et al., 1999;
1987). Kong et al., 2000).
Exploratory studies using TEM and Spatafora et al. (1998) showed that the
SEM were also used for selected bitunicate genus Lulworthia did not group within the
2
 Fungal Diversity

Halosphaeriales clade, also noted by Campbell Origin of marine fungi

(1999) and Chen et al. (1999). Subsequently,
Kohlmeyer et al. (2000) erected the new order Many theories have been advanced to
Lulworthiales to accommodate the genera account for the origin of marine fungi, and in
Lindra and Lulworthia, while Inderbitzin et al. particular the ascomycetes, the Floridean
(2004) showed that the genera Spathulospora hypothesis being the favoured one (Denison
and Haloguignardia had affinities with the and Carroll, 1966; Kohlmeyer, 1975a;
Lulworthiales. Campbell et al. (2005) erected Demoulin, 1985). Spathulospora was
two monotypic genera within the order to considered to be closely related to the
accommodate Lulworthia-like species that did Laboulbeniales, a group in turn thought to be
not form a monophyletic group with related to the Rhodophyta (Kohlmeyer, 1973b).
Lulworthia fucicola, the type species Furthermore, Kohlmeyer (1975a) suggested
(neotypified by Campbell, 2005). Many taxa in that Spathulospora, with its marine occurrence
the Halosphaeriales have also been sequenced and as a parasite of the red alga Ballia, “was
to infer phylogenetic relationships between close to the hypothetical ancestor of
morphologically similar taxa. Studies by Kong Ascomycetes”. However, in a little cited paper,
et al. (2000) and Abdel-Wahab et al. (2001b) Walker et al. (1979) questioned the view that
confirmed the polyphyly of the genus Halo- Spathulospora belonged to an ancestral group,
sarpheia and this led to the erection of a or along with the Laboul-beniales and
number of new genera (Campbell et al., 2003; Uredinales, could be considered “living
Pang et al., 2003a, b). fossils”.
Many genera in the Ascomycota have That marine fungi are an ecological group
not been referred to a family or even an order has never been in dispute, but did they evolve
(Hawksworth et al., 1995; Kirk et al., 2001). in the sea or were they secondly adapted to life
However, the use of molecular techniques has in the marine milieu? While the latter view has
enabled the resolution of a number of genera. gained in acceptance, there has been little
Tam et al. (2003) have shown that the solely evidence to support it. Spatafora et al. (1998)
known marine bitunicate genera, Aigialus, however have advanced the discussion by
Helicascus, Julella and Paraliomyces, with demonstrating that the Halosphaeriales at least
unknown ordinal affinity, can now be has made the transition from terrestrial to
classified in the Pleosporales. Similar marine habitats with the consequent loss of
resolution is possible for the basidiomycetes dehiscing asci. They also concluded that this
Calathella, Digitatispora, Halocyphina, was not a solitary episode, as the Lulworthiales
Mycaureola and Nia (Binder et al., 2001; was also a transitional group, arising indepen-
Hibbett and Binder, 2001, 2002; Binder et al., dently of the Halosphaeriales (Kohlmeyer et al.,
2006). However, many taxa remain to be 2000).
examined before their assignment to an order Sakayaroj (2005) and Schoch et al. (2006)
can be made. In this monograph we have indicated that within the unitunicate
reviewed the published literature so as to ascomycetes seven and three lineages,
propose the best taxonomic assignment for respectively, have migrated into the sea.
“obligate” marine fungi. However, conflict Furthermore five lineages of marine bitunicate
may arise between traditional classifications ascomycetes are also indicated from analyses of
and the need to incorporate phylogenetic molecular data (Suetrong et al., unpublished
analysis, as in the case of the marine data) and may be intermediate forms with many
Basidiomycota. retaining active ascospore discharge. The latter
We have supplied a key within each group is particularly prevalent in mangrove
genus with more than one species, and added habitats where ascus discharge can take place
notes on others that may be useful for further during the intertidal period. Likewise, other
research and understanding of their ecology. groups can be shown to have evolved from

3
terrestrial to marine habitats: at least three There is no data to indicate when marine
lineages of the Basidiomycota (Binder et al., fungi evolved from terrestrial species,
2006), as well as ascomycetes with although Vijaykrishna et al. (2006) predicted
cleistothecial ascomata, but for which no that fungi became adapted to freshwater some
phylogenetic data is available: e.g. Biflua and 390 million years ago. Many lineages of
Marisolaris (Koch and Jones, 1989). marine fungi have been noted, while
The most primitive true marine fungi are freshwater fungi occur in only three classes
the Chytridiomycota, or as their sometimes (Vijaykrishna et al., 2006).
referred “the lower fungi”. They are
characterized by uniflagellate zoospores that Substrata supporting marine fungi
require water for dispersal. They occur in
aquatic habitats, both marine and freshwater, Early records of marine fungi were on
and also in terrestrial habitats such as forests, drift and decaying algae (Cotton, 1909;
agriculture, desert soils, and acidic bogs Sutherland, 1915, 1916a, b), but the study of
(James et al., 2006a, b) and highly adapted Barghoorn and Linder (1944) highlighted the
multiflagellate anaerobic rumen chytrids (Ho existence of a diverse fungal communities
and Barr, 1995; Ho, 2007). The Chytridio- occurring on driftwood. Subsequently dead
mycota have long been regarded as the attached or drift mangrove wood was shown to
ancestral group of other fungi (Barr, 1992; support a wide range of taxa, that differed
Margulis and Schwartz, 1998; Slack et al., appreciably from wood in coastal and oceanic
1999). Molecular studies confirm the early waters (Kohlmeyer, 1984; Hyde and Jones,
evolution of the fungi with the chytrids as the 1989a, b, 1992a; Jones, 2000). Other marine
ancestral group (James et al., 2000). As most substrata have also been investigated for the
molecular studies of chytrids have been of taxa occurrence of marine fungi: coral rocks
from freshwater or terrestrial origin, it is (Kohlmeyer and Volkmann-Kohlmeyer,
unclear whether ancestral forms were marine 1987b), mangrove leaves, hydrozoan tubes,
(James et al., 2006b). However, it seems likely intertidal marsh grasses (Kohlmeyer, 1972b;
that they evolved in freshwater habitats. Most Gessner and Kohlmeyer, 1976; Cuomo et al.,
marine chytrids are parasitic on algae, are few 1982, 1985), man-made materials (Jones and
in number and diversity and may well be Le Campion-Alsumard, 1970) and further
secondary invaders of marine habitats. Some observations on algicolous marine fungi
of these are clearly derived from terrestrial (Kohlmeyer and Volkmann-Kohlmeyer, 2003b;
groups, e.g., Rhizophydium littoreum (James, Zuccaro and Mitchell, 2006). Morrison-
pers. comm.). Undoubtedly marine chytrids Gardiner (2002) isolated a wide range of fungi
warrant further study to determine their from Australian coral reefs, many were typical
ancestral phylogeny, as little is known about terrestrial genera but these could not be
their phylogenetic position. identified using the available taxonomic keys,
Recently, Bass et al. (2007) have and might well represent new taxa. Many other
recovered novel lineages of chytrids from substrata await investigation: tropical marine
environmental DNA from marine ecosystems. grasses, mollusk shells and soft rocks (Golubic
The origin of Rozella and the related group et al., 2005; Raghukumar, 2008).
microsporidia, and Thalassochytrium
gracilariopsidis, parasitic on the alga Materials and Methods
Gracilariopsis sp. (Nyvall et al., 1999) raises
interesting questions as to the origin of marine Specimen collection and incubation
chytrids.
James et al. (2006b) state that the Various substrata supporting marine
earliest fungi were primarily aquatic and fungi, as mentioned earlier in “Substrata
lacked aerial spore dispersal, with at least four supporting marine fungi”, were collected
independent losses of the flagellum, and randomly at different coastal areas in Thailand
giving rise to fungi with aerial dispersal of and other countries e.g. Bahamas, China,
their spores.
4
 Fungal Diversity

Denmark, Guam (Micronesia, USA), England was then transferred with a Pasteur pipette
and Wales (UK). Samples were placed in onto the isolation agar medium. The spores on
plastic bags in order to avoid moisture loss. the agar were left to dry in a closed plate at
Samples with a thick sediment layer or other room temperature and incubated overnight. A
debris were washed thoroughly with running low nutrient medium (e.g. corn meal agar)
tap water. Surface fouling organisms were with antibiotics added (a mixture of Penicillin
scrapped off, followed by rinsing with tap G and Streptomycin in 1g/l) was used in this
water. Samples were then incubated in a procedure. Germinating spores were “picked
plastic box and kept moist by spraying with up” and transferred to a fresh agar plate. At
sterile seawater. least 5-10 isolates of each species were
prepared. Their general colony morphology
Microscopic examination and growth rate were compared to ensure that
the isolates obtained were the same species.
Initial examination was carried out using Axenic cultures were kept at BIOTEC Culture
a stereomicroscope with magnifications Collection (BCC) (Thailand), City University
between 10-40×. The surface of the wood was of Hong Kong (CY) and University of
sliced away in order to locate the buried Portsmouth (PP) (UK).
ascomata or pycnidia. Spore mass contents
were scooped out on a slide for examination. Molecular and phylogenetic analysis
The observation of sporulating structures was
examined in sterile seawater under a Fungal isolates were obtained from
compound microscope. The ascoma structure, BIOTEC Culture Collection (Thailand), City
ascomal wall, the presence or absence of University of Hong Kong (Hong Kong SAR)
catenophyses, paraphyses, pseudoparaphyses and University of Portsmouth (UK) and
and periphyses, ascus structure and cultured into GYP sea water broth (4 g/l
morphology of ascospores are the most glucose, 4 g/l yeast extract, 2 g/l peptone).
important clues for the identification of Mycelium (~100 mg) was harvested by
ascomycetes (Vrijmoed, 2000). For filtration, washed twice with sterile distilled
anamorphic fungi, the morphology of water, blotted dry by filter paper and
conidiomata and the mode of conidiation are immediately frozen in liquid nitrogen.
vital for identification, in addition to the Mycelial pellets were ground into fine powder
conidial characteristics. For basidiomycetes, using a mortar and pestle, and DNA was
the morphology of the basidiomata serves as extracted using the DNeasy Plant DNA
an essential feature for identification. Extraction Kit (QIAGEN) according to the
Therefore, keys are provided to the major manufacturer’s instructions.
phyla in this monograph. Nuclear ribosomal rRNA genes were
amplified using the following primers: small
Isolation of fungi subunit (SSU)- NS1, NS2, NS3, NS5, NS6,
NS8 (White et al., 1990) and large subunit
The routine isolation procedure was (LSU)- JS1, JS5, JS8, LROR, LR7, NL3, NL4,
obtained by single spore isolation (Jones and NL4R (Bunyard et al., 1994; Landvik, 1996).
Hyde, 1988; Choi et al., 1999; Vrijmoed, PCR reactions were performed in 50 μl using
2000). Fruiting bodies were picked up with FINNZYMES, DyNAzyme II DNA
fine forceps or needles and transferred to a Polymerase Kit (Macherey-Nagel, Product
small volume of sterile seawater on a glass code F-551S) in a Perkin Elmer thermal cycler.
slide. The fruiting bodies were then crushed to The amplification cycle consisted of an initial
release the spores. The spore suspension was denaturation step of 94°C for 2 min followed
then agitated to ensure a homogenous spore by 35 cycles of (i) denaturation (94°C for 1
distribution and checked under a compound min), (ii) annealing (55°C for 1.5 min) and (iii)
microscope at low magnification for appro- elongation (72°C for 2.5 min) and a final 10
priate density and identity. The suspension
5
min elongation step at 72°C. The PCR included the anamorphic name under its
products were analyzed by agarose gel teleomorph name. The anamorphic name is
electrophoresis and purified using a also included in Section C, with the
NucleoSpin Plant DNA Purification Kit teleomorph in a lower font, thus enabling the
(Macherey-Nagel, Catalogue No. 740 570. 50) reader to cross-reference the taxonomic names
according to the manufacturer’s instructions. in current use. Molecular sequences where
PCR products were sent to Macrogen Inc., available in the GenBank, are denoted by the
Korea, for direct sequencing. prefix ◙ to each species.
Returned sequences were checked for Each section is divided into higher-level
ambiguity and assembled. Sequences were classification and orders according to that
programme-aligned in Clustal W 1.6 outlined by Hibbett et al. (2007) and where
(Thompson et al., 1994) and manually known, the families (Hibbett, 2006; Spatafora
adjusted in Se-Al v1.0a1 (Rambaut, 1999) and et al., 2006; Zhang et al., 2006; Cannon and
BioEdit version 5.0.6. and 6.0.7 (Hall, 2001, Kirk, 2007). Genera and species of unknown
2004). The tree construction procedure was affinities are referred to incertae sedis.
performed in PAUP* 4.0b10 in Macintosh Anamorphic fungi are listed alphabetically
and Window versions (Swofford, 2002). under hyphomycetes and coelomycetes, but as
SSU and LSU rRNA gene sequences in common practice, no lower taxonomic rank
were analyzed individually using equally is given. However, teleomorphs are listed
weighted maximum parsimony method under species where they are known.
(heuristic searches with a stepwise starting We appreciate that the rapid progress in
tree, a random stepwise addition of 10 molecular phylogeny may change the
replicates and TBR branch-swapping placement of some of the taxa listed here, but
algorithm). Gaps were treated as missing data. hope it will help researchers focus on taxa
Combined SSU and LSU dataset was needing further evaluation and resolution.
analyzed using equally weighted parsimony Many of the taxa listed are known only from
and weighted parsimony approaches. their original description, and attention is
Weighted parsimony analysis was performed drawn to these in the hope that further efforts
using a step matrix to weight nucleotide can be made to collect them. Many may occur
transformations based on the transition : in specific niches that would require further
transversion (ti:tv) ratio estimated from the exploration.
dataset using maximum likelihood score in Classification of many marine fungi
PAUP* (Swofford, 2002). Finally, 1,000 remains a confused and an unresolved issue,
replicates of bootstrapping analysis and is particularly acute for the Ascomycota,
(Felsenstein, 1985) were performed on each the largest group. This is well demonstrated by
dataset (full heuristic searches, stepwise the fact that some 70 genera are referred to as
addition of sequence, 100 replicates of taxa incertae sedis in this monograph. Clearly
random addition of sequence and TBR much effort is required to improve on this state
branch-swapping algorithm). of affairs.
In this treatise we primarily deal with
Layout of the classification of the marine species that have been labelled as obligate
Ascomycota and Basidiomycota marine fungi and those marine derived taxa
isolated from submerged substrata or
Three fungal groups are treated: Section sediments. The latter is not comprehensive but
A: Basidiomycota, Section B: Ascomycota and a start must be made to recognize them as true
Section C: anamorphic species (hyphomycetes marine fungi. However, we have not included
and coelomycetes). Most of the latter have no facultative taxa as they are not found under
known teleomorphs, but this aspect is submerged conditions. As much as we dislike
advancing with the aid of molecular these arbitrary designations, it has been
techniques (Chatmala et al., 2002; Shenoy et necessary to limit the scope of this work. A
al., 2007). Where anamorph/teleomorph clear distinction of what is obligate/facultative
connections have been established, we have depends largely on personal opinion, and this
6
 Fungal Diversity

applies very much to species saprophytic on inland lakes, e.g. Salton Sea (Anastasiou,
decaying culms of maritime grasses, such as 1963a, b), we have excluded those reported
Spartina species, Juncus roemerianus, from hypersaline salterns (Gunder-Cimerman
Phragmites communis, mangrove fungi, et al., 2000) and the Dead Sea where some 70
especially those on the palm Nypa fruticans filamentous species have been isolated, none
which can occur in almost freshwater; and taxa typically marine (Buchalo et al., 1998; Nevo et
isolated from marine sediments (often brackish al., 2003; Wasser et al., 2003; Kis-Papo et al.,
water habitats) (Udea, 1980, 1995a, b; Udea 2003; Kis-Papo, 2005).
and Udagawa, 1983). Because the latter group Some mangrove fungi are regarded as
has been isolated onto agar media, they are obligately marine but also occur on parts of the
largely ignored by marine mycologists and trees not inundated by seawater, e.g. Julella
simply labelled facultative. The fact that some avicenniae, reported on drift/submerged
are repeatedly isolated from such habitats mangrove wood (Hyde, 1992c) but frequently
argues for a re-evaluation of their status, but collected on damaged twigs of Avicennia
this remains outside the scope of this treatise marina above the high tide water mark (Jones,
(Jones, 2000). Kohlmeyer and Volkmann- personal observation). Conversely, Mauritiana
Kohlmeyer (2003c) are critical of recent rhizophorae (Poonyth et al., 2000b) was
studies where fungi have been isolated from described from terrestrial Rhizophora
coral reefs (Kendrick et al., 1982; Höller et al., mucronata, but has also been collected on
2000; Verbist et al., 2000; Morrison-Gardiner, intertidal mangrove wood (Alias, pers. comm.).
2002; Nieves-Rivera, 2002; Raghukumar, We have therefore adopted a broad
2008) because they were isolated onto media, interpretation of what we consider to be
rather than observed sporulating on the obligately marine and this may differ
substratum. significantly from those of others. We have
Care must be taken as some marine fungi listed a few species that we consider to be
may be present in the substratum and do not borderline species that may also be able to
sporulate under the conditions under survive exposure to seawater. This has been
observation (Pang and Mitchell, 2005). It is extended to include taxa isolated from
possible that some of these fungi may exist in sediments, but cannot be consistently rejected
a similar way to terrestrial endophytes because of their mode of isolation. However,
(Zuccaro et al., 2003; Zuccaro and Mitchell, we excluded those considered as halotolerant
2005). Also there is evidence emerging that (Fletcher, 1975; Kohlmeyer et al., 2005).
these “so-called” terrestrial species may have Kohlmeyer and Kohlmeyer (1979), Kohlmeyer
evolved into marine forms, and further and Volkmann-Kohlmeyer (1991a), and Hyde
molecular studies are required to elucidate this and Sarma (2000) have provide synoptic keys
(Alker et al., 2001; Zuccaro et al., 2004). for the identification of marine fungi. All new
Kohlmeyer and Volkmann-Kohlmeyer in taxa published since these publications (some
their papers on fungi growing on Juncus 43 species) are illustrated by line drawings and
roemerianus, have attempted to characterize photographs.
fungi according to their position on the culms
and this reflects their degree of inundation by Marine lichens
seawater: obligate: 6-52 cm above the rhizome
e.g. Phaeosphaeria roemeriani; facultative: Lichens have been largely ignored in the
15-56 cm above the rhizome e.g. Floricola marine mycology literature. Johnson and
striata; and terrestrial or halotolerant: 45- Sparrow (1961) provide a general account of
120 cm above the rhizome, e.g. Septoriella their occurrence, but without a taxonomic
unigalerita (Kohlmeyer et al., 1997; treatment of the taxa. Species included
Kohlmeyer and Volkmann-Kohlmeyer, 2000). Arthopyrenia sublitoralis (on limpet shells),
However, as noted, there is a considerable Lichina pygmaea, L. confinis, Verrucaria
overlap in their position with respect to the ditmarsica, V. maura, V. microspora, and V.
rhizome. Although we include fungi from striatula (all on rock), the discussion focusing
7
on lichen zonation. Kohlmeyer and Kohlmeyer considerable emphasis on the later, while for
(1979) document submarine lichens and filamentous fungi morphology still pays a
lichen-like associations, listing 18 species in significant role (Statzell-Tallman et al., 2008).
the genera Arthopyrenia, Lichina, Stigmidium, Therefore yeasts are best left to specialist’s
and Verrucaria, on Littorina, barnacles and texts, e.g., Kurtzman et al., 5th edition of The
algae. Verrucaria maura can also be found on Yeasts, a Taxonomic Study (2009). The study
marine wood piles and ironwork (Fletcher, of yeasts has also been plagued by the
pers. comm.). However, there is no taxonomic arguments as to whether they are obligate or
treatment of these taxa. Jones (1976) did not facultative marine. As for filamentous fungi
include marine lichens in his treatise on this is an arbitrary division and emphasis
marine fungi, while Hyde et al. (2000) list should be placed on habitat and niches within
only two. The best account of marine lichens the marine ecosystem. Yeasts isolated from the
is that of Fletcher (1973a, b) who details some sea can grow equally well on freshwater media
80 names of littoral and supralittoral lichens. (Fell, pers. comm.).
However, he was unable to examine all of The number of yeasts documented has
these (Fletcher, pers. comm.). Erichsen (1930) steadily increased and with an estimated 1,500
described some twelve marine Verrucaria species. Taxa such as Leucosporidium spp.,
species from the Elbe Estuary in the 1930’s, Rhodosporidium spp., Candida austromarina,
this list is conservative. Therefore the number C. natalensis, Kwoniella mangroviensis and
of lichens listed in this monograph is still Sympodiomyces parvus are undoubtedly
conservative. autochonous species as they are recovered in
Nearly all of the species listed here have relatively high numbers from ocean samples
all been collected in the intertidal zone and (Lachance and Starmer, 1998). Their role in
thus subject to inundation by seawater. They nature is to break down a wide range of
are cosmopolitan, especially polar to temperate, organic matter, including lignin. Yeast
but rarely recorded from the tropics (Harada, communities in the open ocean are extensive
1995). Hawksworth (2000) com-ments on the and much greater that filamentous fungi. This
marine and freshwater lineages of lichens, in may be accounted for by the ratio of surface to
particular the genera Lichina, Pyrenocollema volume of the yeasts, enabling greater uptake
and Verrucaria. This aspect will be considered of nutrients (Fell, pers. comm.). In mangrove
later in this volume. swamps there are a large number of yeasts per
unit of water, actively involved in recycling
mangrove leachates and in turn acting as a
Marine yeasts food source for filter feeding invertebrates
(Statzell-Tallman et al., 2008). As with
Marine yeasts also have fared poorly in filamentous fungi, many yeasts remain to be
texts dealing with marine fungi: Johnson and described, especially in mangrove swamps
Sparrow (1961) and Kohlmeyer and (Fell et al., 2004). Statzell-Tallman et al.
Kohlmeyer (1979) list eight and twenty three (2008) reported 55 species of ascomycetes and
species, respectively of obligate marine yeasts. 58 species of basidiomycetes yeasts from three
Kohlmeyer and Kohlmeyer (1979) list a mangrove habitats, 50% of which are
further 140 facultative yeasts. Most of these undescribed.
have been isolated by plating out seawater on Many marine yeasts are also secondary
to various media or from marine animals and invaders of the sea, and a number of
sediments (van Uden and Castello-Branco, phylogenetic lineages have been reported
1963; Meyers et al., 1967; Fell, 1976). More (Jones and Choeyklin, 2008). For example,
recent treatment of marine fungi have been basidiomycete lineages: 1. Tremellomycetes,
confined to filamentous species (Kohlmeyer Cystofilobasidiales: Cystofilobasidium
and Volkmann-Kohlmeyer, 1991a; Hyde et al., bisporidii, C. capitatum (Fell et al., 2001),
2000) and we follow this in this volume. Rhodosporidium diobovatum, Rh.
Techniques for the study of yeasts are paludigenum, and Rh. sphaerocarpum (Fell et
polyphasic, phenotypic and molecular, with al., 2001); 2. Agaricostilbomycetes,
8
 Fungal Diversity

Agaricostilbales: Sterigmatomyces halophilus Calathella..................................................... 15

(Kurtzman and Fell, 2006); 3. Halocyphina ................................................. 15
Microbotryomycetes, Sporidiobolales: Nia................................................................. 15
Sakaguchia dacryoidea, Leucosporidiales:
Leucosporidium spp. (Fell et al., 2006; Physalacriaceae ............................................. 16
Kurtzman and Fell, 2006) and, ascomycete Physalacria ................................................... 16
lineage: Saccharomycetes, Saccharomy- Mycaureola ................................................... 16
cetales: Saccharomyces spp., Metschnikowia
spp. Agaricomycetes incertae sedis
Russulales

Key to the major phyla of marine fungi Digitatispora clade

Digitatispora ................................................. 17
1. Meiospores primarily exogenous, born on
basidia or similar structures,
Section A: Basidiomycota .................. 14 Peniophoraceae ............................................. 17
1. Meiospores produced endogenously, Haloaleurodiscus .......................................... 17
formed in asci,
Section B: Ascomycota....................... 21
1. Mitospores exogenous, or in pycnidia, Phylum: ASCOMYCOTA
Section C: Anamorphic fungi Subphylum: Pezizomycotina
(hyphomycetes and coelomycetes) ............. 146 Class: Dothideomycetes
Subclass: Dothideomycetidae
The following classification is based on Pleosporomycetidae
the publication “A higher-level phylogenetic
classification of the Fungi” by Hibbett et al. Subclass: Dothideomycetidae
(2007) for higher order ranks and Cannon and 1. Capnodiales 2. Dothideales
Kirk (2007) for familial placement.

Classification of the marine fungi 1. Capnodiales

Mycosphaerellaceae...................................... 23
Phylum: BASIDIOMYCOTA Mycosphaerella............................................. 23
Subphylum: Ustilaginomycotina Sphaerulina ................................................... 24
Class: Ustilaginomycetes Pharcidia....................................................... 24
Subclass: Ustilaginomycetidae
1.Urocystales 2. Ustilaginales 2. Dothideales
Dothideaceae................................................. 25
1. Urocystales Scirrhia ......................................................... 25
Urocystaceae ................................................. 14
Flamingomyces ............................................ 14 Dothideales incertae sedis
Botryosphaeriaceae ....................................... 25
2. Ustilaginales Amarenomyces .............................................. 25
Ustilaginaceae .............................................. 14 Belizeana....................................................... 26
Parvulago..................................................... 14 Capillatospora .............................................. 26
Passeriniella ................................................. 26
Subphylum: Agaricomycotina Thalassoascus ............................................... 27
Class: Agaricomycetes
Subclass: Agaricomycetidae Dothideomycetidae family incertae sedis

Agaricales Lautosporaceae ............................................. 27

Lachnellaceae............................................... 15 Lautospora .................................................... 27
9
Planistromellaceae ........................................ 28 Byssothecium................................................. 47
Loratospora................................................... 28
Pleosporales incertae sedis
Zopfiaceae..................................................... 28 Aigialus ......................................................... 48
Coronopapilla ............................................... 28 Biatriospora .................................................. 48
Caryospora ................................................... 28 Didymella...................................................... 48
Pontoporeia................................................... 29 Halotthia ....................................................... 49
Heleiosa ........................................................ 49
Subclass: Pleosporomycetidae Julella............................................................ 49
Kirschsteiniothelia ........................................ 51
Pleosporales Leptosphaerulina .......................................... 51
Lineolata ....................................................... 52
Didymosphaeriaceae ..................................... 29 Massariosphaeria ......................................... 52
Didymosphaeria ............................................ 29 Salsuginea ..................................................... 53
Tirisporella ................................................... 53
Testudinaceae................................................ 30 Wettsteinina................................................... 54
Verruculina ................................................... 30
Dothideomycetes incertae sedis
Leptosphaeriaceae......................................... 30
Leptosphaeria ............................................... 30 Hysteriales
Hysteriaceae .................................................. 55
Lophiostomataceae........................................ 33 Gloniella ....................................................... 55
Decaisnella.................................................... 33
Herpotrichia.................................................. 33 Patellariales
Lophiostoma.................................................. 35 Patellariaceae ................................................ 55
Massarina ..................................................... 35 Banhegyia ..................................................... 55
Paraliomyces................................................. 36 Patellaria ...................................................... 55
Platystomum.................................................. 37
Quintaria....................................................... 37 Jahnulales
Hypostromataceae......................................... 55
Melanommataceae ........................................ 37 Manglicola .................................................... 55
Acrocordiopsis .............................................. 37
Astrosphaeriella............................................ 41 Class: Eurotiomycetes
Bicrouania..................................................... 41 Subclass: Eurotiomycetidae
Caryosporella ............................................... 42 1. Onygeniales 2. Eurotiales
Trematosphaeria ........................................... 42
1. Onygeniales
Monoblastiaceae............................................ 43
Gymnoascaceae............................................. 56
Ascocratera ................................................... 43
Gymnascella.................................................. 56
Phaeosphaeriaceae ........................................ 43
Carinispora ................................................... 43 2. Eurotiales
Lautitia.......................................................... 43 Trichocomaceae ............................................ 59
Phaeosphaeria .............................................. 44 Eupenicillium ................................................ 59

Pleosporaceae................................................ 45
Decorospora.................................................. 45 Subclass: Chaetothyriomycetidae
Helicascus ..................................................... 46 1. Chaetothyriales
Falciformispora ............................................ 46 2. Pyrenulales
Pleospora ...................................................... 46 3. Verrucariales
Tremateia ...................................................... 47
1. Chaetothyriales
Teichosporaceae............................................ 47 Herpotrichellaceae ........................................ 59
10
 Fungal Diversity

Capronia ....................................................... 59 Class: Arthoniomycetes

2. Pyrenulales Arthoniales
Pyrenulaceae ................................................. 60
Pyrenographa................................................ 60 Roccellaceae ................................................. 69
Xenus............................................................. 60 Halographis .................................................. 69

Requinellaceae .............................................. 60 ARTHONIOMYCETIDAE family incertae

Mauritiana .................................................... 60 sedis

Xanthopyreniaceae........................................ 62 Melaspileaceae ............................................. 69

Collemopsidium ............................................ 62 Melaspilea.................................................... 69

3. Verrucariales Sordariomycetes
Verrucariaceae .............................................. 63 Subclass
Mycophycias.................................................. 63 Xylariomycetidae
Verrucaria..................................................... 64 Hypocreomycetidae
Sordariomycetidae

Class: Laboulbeniomycetes Subclass: Hypocreomycetidae

1. Hypocreales
Laboulbeniales 2. Coronophorales
Laboulbeniaceae............................................ 66 3. Halosphaeriales
Laboulbenia .................................................. 66
1. Hypocreales
Bionectriaceae............................................... 72
Class: Lecanoromycetes Emericellopsis............................................... 72
Subclass: Lecanoromycetidae Halonectria ................................................... 73
Heleococcum................................................. 73
Lecanorales Kallichroma .................................................. 73
Dactylosporaceae .......................................... 66
Dactylospora................................................. 66 Hypocreaceae ................................................ 74
Neocosmospora............................................. 74
Class: Leotiomycetes Payosphaeria ................................................ 74
Subclass: Lecanoromycetidae Pronectria ..................................................... 74

Helotiales Hypocreales incertae sedis

Helotiaceae.................................................... 68 Torpedospora................................................ 75
Amylocarpus ................................................. 68 Juncigena ...................................................... 75
Swampomyces ............................................... 75
Vibrisseaceae ................................................ 68 Etheirophora ................................................. 76
Vibrissea ....................................................... 68
Laetinaevia.................................................... 68 2. Coronophorales
Nitschkiaceae ................................................ 77
Class: Lichinomycetes Groenhiella ................................................... 77

Lichinales 3. Halosphaeriales
Lichinaceae ................................................... 69 Halosphaeriaceae .......................................... 77
Lichina .......................................................... 69 Alisea............................................................. 80

11
Aniptodera..................................................... 81 Tunicatispora ............................................. 108
Anisostagma .................................................. 82
Antennospora ................................................ 82 Subclass: Sordariomycetidae
Appendichordella .......................................... 82 1. Diaporthales
Arenariomyces .............................................. 83 2. Chaetosphaeriales
Bathyascus .................................................... 83 3. Sordariales
Bovicornua .................................................... 84 4. Ophiostomatales
Carbosphaerella ........................................... 84
Ceriosporopsis .............................................. 84 1. Diaporthales
Chadefaudia .................................................. 86 Valsaceae ................................................... 112
Corallicola .................................................... 86 Cryptovalsa ................................................ 112
Corollospora ................................................. 86 Diaporthe ................................................... 112
Cucullosporella............................................. 89 Gnomonia................................................... 112
Haligena........................................................ 89
Halosphaeria................................................. 90 Melanconidaceae........................................ 113
Halosarpheia sensu stricto ........................... 91 Hypophloeda .............................................. 113
Halosarpheia sensu lato ............................... 91
Halosphaeriopsis .......................................... 92 Diaporthales incertae sedis
Haiyanga....................................................... 92 Argentinomyces.......................................... 113
Havispora...................................................... 93
Iwilsoniella.................................................... 93 2. Sordariales
Lautisporopsis............................................... 94 Lasiosphaeriaceae ...................................... 113
Lignincola ..................................................... 94 Biconiosporella .......................................... 113
Limacospora ................................................. 94
Luttrellia ....................................................... 95 Chaetomiaceae ........................................... 113
Magnisphaera ............................................... 96 Chaetomium ............................................... 113
Marinospora ................................................. 97 Zopfiella ..................................................... 114
Moana ........................................................... 97
Morakotiella.................................................. 97 Sordariales incertae sedis
Nais ............................................................... 98 Abyssomyces .............................................. 114
Natantispora ................................................. 98 Savoryella .................................................. 114
Naufragella ................................................... 98
Nautosphaeria............................................ .. 99 3. Chaetosphaeriales
Neptunella .................................................. 100 Chaetosphaeriaceae.................................... 115
Nereiospora................................................ 100 Chaetosphaeria .......................................... 115
Nimbospora................................................ 100
Nohea ......................................................... 101 4. Ophiostomatales
Oceanitis .................................................... 101 Lanspora .................................................... 116
Ocostaspora ............................................... 102
Okeanomyces ............................................. 102 Subclass: Xylariomycetidae
Ondiniella .................................................. 103
Ophiodeira ................................................. 103 Xylariales
Panorbis ..................................................... 104 Cainiaceae .................................................. 117
Pseudolignincola........................................ 104 Arecophila.................................................. 117
Remispora .................................................. 104 Atrotorquata............................................... 117
Saagaromyces ............................................ 105
Sablecola.................................................... 106 Clypeosphaeriaceae.................................... 117
Thalassogena ............................................. 106 Apioclypea.................................................. 117
Thalespora ................................................. 106 Ommatomyces ............................................ 118
Tirispora .................................................... 108
Trailia ........................................................ 108 Diatrypaceae .............................................. 118
Trichomaris................................................ 108 Cryptosphaeria .......................................... 118
12
 Fungal Diversity

Eutypa ........................................................ 118 3. Magnaporthales

Eutypella .................................................... 118 Magnaporthaceae ....................................... 137
Pedumispora .............................................. 119 Buergenerula.............................................. 138
Gaeumannomyces ...................................... 140
Hyponectriaceae......................................... 119 Pseudohalonectria ..................................... 140
Frondicola.................................................. 119
Phragmitensis............................................. 119 4. Phyllachorales
Phyllachoraceae ......................................... 133
Xylariaceae................................................. 119 Phyllachora................................................ 133
Anthostomella ............................................ 119 Polystigma.................................................. 134
Astrocystis .................................................. 120
Fasciatispora ............................................. 120 Phyllachorales incertae sedis
Halorosellinia ............................................ 121 Mangrovispora........................................... 136
Nemania ..................................................... 121 Marinosphaera........................................... 136
Nipicola...................................................... 123 Phycomelaina............................................. 136
Xylariales incertae sedis Unitunicate Ascomycota family incertae
Adomia ....................................................... 123 sedis
Lanceispora................................................ 123
Linocarpon ................................................. 124 Hispidicarpomycetaceae ............................ 137
Neolinocarpon............................................ 124 Hispidicarpomyces..................................... 137
Oxydothis ................................................... 125
Phomatospora ............................................ 126 Spathulosporaceae...................................... 137
Retrostium .................................................. 137
Sordariomycetes incertae sedis
1. Lulworthiales Mastodiaceae.............................................. 140
2. Koralionastetales Turgidosculum ........................................... 140
3. Magnaporthales Mastodia .................................................... 140
4. Phyllachorales
Papulosaceae .............................................. 141
1. Lulworthiales Papulosa .................................................... 141
Lulworthiaceae........................................... 127
Kohlmeyeriella ........................................... 127 Unitunicate Ascomycota genera incertae
Lindra......................................................... 127 sedis
Lulwoana.................................................... 128
Lulwoidea................................................... 128 Aquamarina................................................ 141
Lulworthia.................................................. 128 Aropsiclus .................................................. 142
Rostrupiella................................................ 131 Biflua.......................................................... 142
Haloguignardia.......................................... 131 Crinigera.................................................... 142
Dryosphaera .............................................. 142
Spathulosporaceae...................................... 131 Eiona .......................................................... 142
Spathulospora ............................................ 131 Hapsidascus ............................................... 143
Marisolaris................................................. 143
2. Koralionastetales Orcadia ...................................................... 143
Koralionastetaceae ..................................... 132 Rhizophila .................................................. 144
Koralionastes ............................................. 132 Saccardoella .............................................. 144
Pontogeneia ............................................... 133
ANAMORPHIC ASCOMYCETES ....... 146

13
 Section A: cell wall and pigmented teliospores (Bauer et
PHYLUM: BASIDIOMYCOTA al., 2007).

Key to the Basidiomycota

USTILAGINALES
1. Basidiome reduced, parasite, hyphal septum lacking Ustilaginaceae
a dolipore and no parthenosomes ............................. .
.................................................. Ustilaginomycotina
Parvulago R. Bauer, M. Lutz., Piatek, Vánky
1. Visible basidiome, saprophytes, symbionts or & Oberw., Mycol. Res. 111: 1203, 2007 ..... (1)
parasites, basidiospores ballistosporic or statimo- ◙ P. marina (Durieu) R. Bauer, M. Lutz.,
sporic, clamp-connections present or absent.............. Piatek, Vánky & Oberw., Mycol. Res. 111:
......................................................Agaricomycotina
1203, 2007.
Ustilago marina Durieu, Annls Sci. Nat. Bot., sér.
Subphylum: USTILAGINOMYCOTINA 5, 5: 134, 1866.
Sporulation is at the base of the host
USTILAGINOMYCETES plant culms, in the intercellular space
USTILAGINOMYCETIDAE underneath the 2-layered epidermis, forming
Two orders with marine species bulbous swelling, sori not covered by a
presidium (after Bauer et al., 2007). This
1. Parasitic on dicotyledonous hosts .......... Urocystales species is known from Eleocharis parvula
1. Parasitic on members of the Poaceae ... Ustilaginales
(Cyperaceae) and was recently collected at the
Bassin d’Arcachon, Gironde, France (Bauer et
UROCYSTALES
al., 2007). It is the only member of the
Urocystaceae
Ustilaginaceae occurring on the Cyperaceae
and sporulating at the base of the culms of the
Flamingomyces R. Bauer, M. Lutz., Piatek,
host plant under the epidermis (Bauer et al.,
Vánky & Oberw., Mycol. Res. 111: 1202,
2007).
2007.............................................................. (1)
◙ F. ruppiae (Feldmann) R. Bauer, M. Lutz.,
Subphylum: AGARICOMYCOTINA
Piatek, Vánky & Oberw., Mycol. Res. 111:
1203, 2007.
Melanotaenium ruppiae Feldmann, Rev. Gén. AGARICOMYCETES
Bot. 66: 36, 1959. AGARICOMYCETIDAE
Sori dark on stems and leaves of the host,
initially covered by the host epidermis, Two orders with marine representatives, with
rupturing at maturity, spores dark, single with reduced basidiomes:
no germ pores. Teliospores produced singly,
smooth, spore wall comprising and electron- 1. A group with large pileate-stipitate basidiomes.........
................................................................. Agaricales
opaque exosporium and an electron- 1. A group of pileate to resupinate basidiomes .............
transparent endosporium. Teliospores germi- .......................................... Russulales incertae sedis
nate apically. Initial collection of this species
was on Ruppia maritima (Ruppiaceae), at the AGARICALES
Etang du Canet, Pyrénées Orientales, France
(Feldmann, 1959), with a subsequent collec- After Matheny et al. (2006) and equivalent to
tion at Bassin d’Arcachon, Gironde, France euagarics clade (Hibbett and Binder, 2001;
(Bauer et al., 2007). The new genus was Binder et al., 2006).
placed in the Urocystales (Bauer et al., 2007),
but earlier placed in the Ustilaginales 1. Basidioma resupinate, basidiospores filiform ............
(Begerow et al., 2006; Matheny et al., 2006). ................................................................Mycaureola
This was confirmed by the molecular study of 1. Basidioma not resupinate, ascospores not filiform ....
................................................................................. 2
Bauer et al. (2007). The genus is characterized
by the formation of haustoria with an electron- 2. Basidioma cyphelloid, funnel-shaped ...................... 3
opaque, vesicular matrix coating the fungal
14
 Fungal Diversity

2. Basidioma sub-globose, puff ball-like, no stalk, basidiospores subglobose, unicellular, smooth,

basidiospores with appendages ............................ Nia hyaline, nonamyloid, accumulating at the tip
2. Basidioma globose on a stalk, basidiospores non-
appendaged .............................................Physalacria of the basidiome and washed away by the
incoming tide. Detailed account of the
3. Basidioma white, surface hairs smooth, basidia 13- basidiome and the release of basidiospores is
22 × 5-9.5 µm........................................Halocyphina given by Nakagiri and Ito (1991). Classified in
3. Basidioma yellowish to ochre yellow, cup-like, wall the Polyporales, Cyphellaceae (Kirk et al.,
with surface hairs, brownish, dextrinoid, minutely
encrusted, basidia 60-80 × 5-8.5 µm......... Calathella 2001) while Ginns and Malloch (1977)
recorded it as a cyphelloid basidiomycetes.
Lachnellaceae Calathella mangrovei shares a common
ancestor with Nia and Cyphellopsis (Hibbett
After Matheny et al. (2006) and equivalent to and Binder, 2001). Molecular data confirm its
Nia clade (Hibbett and Binder, 2001; Binder et assignment to the euagaric clade and Nia clade
al., 2006) (Binder et al., 2001).

Calathella D.A. Reid, Persoonia 3: 122, 1964. Nia R.T. Moore & Meyers, Mycologia 51: 874,
...................................................................... (1) 1959.............................................................. (3)
◙ C. mangrovei E.B.G. Jones & Agerer, Bot. N. epidermoidea M.A. Rosselló & Descals,
Mar. 35: 259, 1992. Mycol. Res. 97: 68, 1993.
Basidiomes cyphelloid, pedunculate, N. globospora Barata & Basilio, Mycol. Res.
superficial on wood, yellowish with a reddish 101: 687, 1997.
tinge, older specimens ochre-yellow, leathery, ◙ N. vibrissa R.T. Moore & Meyers,
tomentose, solitary or gregarious, mycelium Mycologia 51: 874, 1959 (Type species).
with clamp connections, sterile hairs form a Basidiomes subglobose, superficial,
ring around the mouth of the basidiocarp that cylindrical, pedicellate, light coloured
are simple, brownish, non-septate, dextrinoid, becoming yellow, pink or orange, soft tissue,
round and minutely encrusted, basidia smooth, peridium ruptures at maturity, with
suburniform to cylindrical, hyaline, with 4 clamp connections, basidia subglobose to oval
sterigmata, basidiospores hyaline, unicellular, with 4-8 basidiospores, hyaline, lacking
smooth-walled, elliptical and accumulate at the sterigmata, basidiospores ovoid to ellipsoidal,
tip of the basidiocarps (Figs 3b, c). Initially unicellular, hyaline, with variable number of
referred to the Cyphellaceae, but molecular appendages, one terminal the others lateral.
sequences suggest it is better placed in the Initially N. vibrissa was classified as a
euagaric clade. A wide spread tropical species, deuteromycete (Moore and Meyers, 1959) but
often occurring on freshly cut and exposed with the demonstration of basidia and clamp
branches of the mangrove tree Bruguiera. connections and a dolipore septum, it clearly
belongs in the homobasidiomycetes (Doguet,
1967, 1968; Brooks, 1975). Subsequently it
Halocyphina Kohlm. & E. Kohlm., Nova has been referred to the Melanogastrales
Hedw. 9: 100, 1965 ...................................... (1) (Torrendiaceae: Dring, 1973, or Melanogastra-
◙ H. villosa Kohlm. & E. Kohlm., Nova Hedw. ceae: Doguet, 1967); Nidulariaceae (Rossello
9: 100, 1965 (Type species). et al., 1993) and the Niaceae (Jűlich, 1981).
Basidiomes cyphelloid, initially turbi- Binder et al. (2001) placed N. vibrissa in the
nate or clavate becoming funnel-shaped, euagarics clade, forming a sister group with
pedunculate, superficial, white or yellowish, Henningsomyces candidus. In a subsequent
soft, thin-walled, tomentose, solitary generally paper, Hibbett and Binder (2001) confirm the
gregarious, basidiome apex covered by sterile placement of N. vibrissa in the euagarics clade,
hairs, mycelium with clamp-connections, along with two other marine basidio-mycetes:
basidia clavate to cylindrical, 4-spored, C. mangrovei and H. villosa. They opined that
hyaline, sterigmata present and deliquescing, the shift from a terrestrial to an aquatic habitat
can be accounted for by three or four
15
independent transitions, the three shift Basidiomes stipitate-capitate, solitary to
scenarios giving rise to the terrestrial species gregarious, capitulum globose to subglobose to
Cyphellopsis anomala (i.e. derived from a pyxie-like, white becoming yellow, stipe
marine ancestor). central, cylindrical, pruinose, arising from a
Nia and Halocyphina are strongly short cushion (Fig. 2), basidia 4-spored,
supported in a clade (bootstrap value of 100%) clavate with sterigmata, basidiospores
with C. mangrovei and two terrestrial species: elongate-ellipsoid, smooth, hyaline, non-
Cyphellopsis anomala and Favolaschia amyloid, thin-walled, hymenium with gloeo-
intermedia. Hibbett and Binder (2002) cystidia Physalacria maipoensis produces a
speculate that Physalacria maipoensis may “capitate” fruiting body with a globose head
represent an early stage in the transition from and a short stalk (Fig. 2), and molecular data
terrestrial to the marine environment for these assign it to the euagarics within a clade
basidiomycetes. This is based on the terrestrial comprising Henningsomyces candidus and a
habitat of most Physalacria species, with Ph. sister clade to Schizophyllum commune but not
maipoensis often found in the intertidal zone within the family Marasmiaceae (Binder et al.,
of tropical mangroves. However, we have 2001). Inderbitzin and Desjardin (1999)
collected Physalacria species on palm rachis consider the genus as representing “reduced”
submerged or in amphibious habitats in a peat agarics allied to the genus Gloiocephala in the
swamp in Thailand (Pinnoi and Jones, pers. Tricholomataceae.
comm.).
There is evidence to suggest that N. Mycaureola Maire & Chemin, Comptes. rendu
vibrissa is a species complex and further hebd. Séanc. Acad. Sci., Paris, 175: 321, 1922.
studies are required to resolve this (Jones and ...................................................................... (1)
Jones, 1993; Binder and Hibbett, 2001). ◙ M. dilseae Maire & Chemin, Comptes.
Calathella, Halocyphina and Nia rendu hebd. Séanc. Acad. Sci., Paris, 75: 321,
consistently group together and all are adapted 1922 (Type species).
for life in aquatic habitats (Hibbett and Binder, Basidiomes globose, hemispherical to
2001). All have reduced basidiomes, possibly ellipsoidal, sessile, white, smooth, ostiolate,
as an adaptation to an aquatic environment with up to 12 formed in a circle around the
where large fruit bodies would not survive edges of decaying algal tissue, basidia are
(Jones, 1988). Hibbett (2007) considers the cylindrical to subclavate, no sterigmata,
minute forms of cyphelloid basidiomycetes to lacking cystidia, each basidium produces 4
be related to selection of spore production sigmoid basidiospores, unicellular, hyaline,
from minimal substrates. smooth-walled, lacking a sheath or appendages
(Porter and Farnham, 1986; Stanley, 1992).
1. Basidiomata lack sterile hairs/appendages (4.8-8 μm) Originally referred to the Ascomycota,
............................................................ N. globospora but ultrastructural studies confirm its
1. Basidiomata with sterile hairs/appendages .............. 2
assignment to the Basidiomycota (Porter and
2. Basidiospores 11-12 × 7-8 μm, appendages 25- Farnham, 1986). Mycaureola, a monotypic
35μm .........................................................N. vibrissa genus, is parasitic on the red alga Dilsea
2. Basidiospores 6-7 × 3-4 μm, appendages 21-28 μm.. carnosa and has a cyphelloid, gasteroid
........................................................ N. epidermoidea fruiting body that suggests assignment to the
Cyphellaceae euagaricoid clade. Molecular
Physalacriaceae
sequences indicate that the species nested
within Rhizomarasmius pyrrhocephalus and
After Matheny et al. (2006) and Binder
Gloiocephala phormiorum and is close to
et al. (2006).
Xerula and Oudemansiella species, in the
Physalacriaceae clade (Binder et al., 2006).
Physalacria Peck, Bull. Torrey bot. Club 9: 2,
However, its closest terrestrial relative
1882.............................................................. (1)
could not be identified with confidence
◙ Ph. maipoensis Inderb. & Desjardin,
(Binder et al., 2006). Gloiocephala aquatica,
Mycologia 91: 666, 1999.

16
 Fungal Diversity

Fig. 2. Physalacria maipoensis saprophytic on stems of Acanthus ilicifolius.

(Desjardin et al., 1995), a freshwater species, ◙ D. marina Doguet, Comptes. rendu hebd.
is in the same clade as M. dilseae, indicating Séanc. Acad. Sci., Paris, 254: 4338, 1962
that there is a second lineage of marine (Type species).
Agaricales (Binder et al., 2001, 2006). Basidiomes form irregular colonies on
Mycaureola dilseae is seasonal in its the wood surface, mycelium with clamp
occurrence and restricted to temperate areas connections, resupinate, hyaline to gray, soft,
occurring during late autumn in the UK when basidia cylindrical or subclavate, basidia
seawater temperatures are low (Stanley, 1992). elongate, no sterigmata, four-spored, hyaline,
basidiospores tetraradiate, one basal arm and
three radiating apical arms, hyaline, deciduous.
AGARICOMYCETES incertae sedis A preliminary molecular study of this genus
using mitochondrial rDNA sequence suggests
RUSSULALES this species should be placed in the russuloid
clade (Hibbett, pers. comm.). However, further
After Hibbett et al. (2007) and equivalent to molecular studies are required to confirm this
russuloid clade (Hibbett and Thorn, 2001; placement.
Binder and Hibbett, 2002).
1. Apical branch of basidiospores up to 25 μm long, 8
Two genera with marine species: μm diam .................................................. D. lignicola
1. Apical branch of basidiospores up to 41 μm long, 4
μm diam .................................................... D. marina
1. Resupinate thallus small, basidiospores tetraradiate ..
..............................................................Digitatispora
1. Resupinate thallus extensive, basidiospores not Peniophoraceae
tetraradiate..................................... Haloaleurodiscus
Haloaleurodiscus N. Maek., Suhara & K.
Digitatispora clade (Hibbett and Thorn, 2001) Kinjo, Mycol. Res. 109: 826, 2005 .............. (1)
Digitatispora Doguet, Comptes. rendu hebd. ◙ H. mangrovei N. Maek., Suhara & K. Kinjo,
Mycol. Res. 109: 827, 2005 (Type species).
Séanc. Acad. Sci., Paris, 254: 4338,1962.... (2) Basidiome resupinate, irregular fruit
D. lignicola E.B.G. Jones, Mycotaxon 27: 155, body on wood, pinkish cream, becoming pale
1986. dull-cream to greyish-white when dry,
gloeocystidia present, basidia narrowly
clavate 65-75 × 9-10.5 µm, with a basal clamp

17
connection and four sterigmata, basidiospores basidiomycetes have received little attention
ellipsoid to cylindrical, 13.5-15.5 × 8-9.5 µm, to date and further investigation for their
warted, thin to slightly thick-walled, amyloid adaptation to semi-aquatic habitats is
and lacking appendages. Haloaleurodiscus warranted.
mangrovei was described from decaying and Trees at Khanom mangrove, southern
decorticated branch of a living Sonneratia Thailand, were badly affected by butt rot
alba tree (Maekawa et al., 2005). Unlike some caused by basidiomycetes. The trees were
marine basidiomycetes, it has ellipsoid to multi branched as the result of fungal attack
cylindrical basidiospores 13.5-15.5 × 8-9.5 (Fig. 6). Two species of Phellinus were
µm, warted, thin to slightly thick-walled, identified by Dr. T. Hattori, Japan: Phellinus
amyloid, and lacking appendages (Fig. 3a). mangrovicus and Ph. swieteniae, and known to
Maekaea et al. (2005) suggest that the be potent wood decay species. Phellinus
thickening of the basidiospore walls after mangrovicus is of special interest as it has not
discharge from the sterigmata could be the been collected since it was described by
first acquired features to evolve and represent Imazeki, over 70 years ago. Therefore a
a unique evolutionary transition from modern description, with illustrations, is
terrestrial to marine environments. Although therefore required as well as sequence data to
H. mangrovei resembles Aleurodiscus sensu determine its phylogenetic relationship with
lato, phylogenetically it nestles in the root of other species in the genus.
the Peniophorales clade (18S, 28S rDNA Although there is no documentation of
sequences) (Maekawa et al., 2005). butt rot of Xylocarpus, there is a report of butt
and heart rot in another mangrove tree,
OTHER BASIDIOMYCOTA Rhizophora apiculata. Mwangi (2001)
reported that old Rh. apiculata trees were
Clipson et al. (2001) list five other attacked by two pathogenic terrestrial
basidiomycetes in their checklist of European polypores: Phellinus pachyphloeus and Ph.
marine fungi, but these are maritime occurring rimosus in Kenya.
in sand dunes and not obligate marine fungi Considerable progress has been made to
(Psalliota litoralis, Laccaria trullisata, resolve the taxonomic assignment of the
Inocybe decipiens (= Inocybe dunensis), marine basidiomycetes. The genera Calathella,
Tulostoma macrocephalum). Other maritime Halocyphina, and Nia all appear to be related
basidiomycetes have been reported, but (euagaric clade) but have very different
although these are undoubtedly tolerant to sea basidiomatal and spore morphologies.
spray, they are never intertidal or submerged. Similarly, Physalacria and Mycaureola nestle
Similarly, a Coprinus species has been distantly in the Physalacriaceae. Many marine
collected on cut branches of Avicennia marina basidiomycetes have a reduced basidiome,
and floating in the water in Mai Po Mangrove, regarded as an adaptation to marine conditions
Hong Kong (Jones, unpublished data). (Jones, 1988).
Another basidiomycete found on decaying
plant material (particularly Acanthus TOTAL BASIDIOMYCOTA ................. (12)
ilicifolius), and wood is an Aegerita species
(Sadaba et al., 1995), whose identity has not
been determined (Fig. 5).
At the time of writing we have collected
12 basidiomycetes on the intertidal bases of
the palm Nypa fruticans, e.g. Grammothele
fuligo (Fig. 4). Agarics have also been found
on mangrove soils, especially Coprinus,
Cortinarius and Mycena species; they are
short lived, fruit during the intertidal period
and shed their spores before the tide returns
(Jones, personal observation). These
18
 Fungal Diversity

a b c

Fig. 3. Basidiospores of a. Haloaleurodiscus mangrovei (after Maekawa et al., 2005), b., c. Calathella
mangrovei, Basidium, sterigmata and basidiospore (b), and basidiospores (c). Bars a, c = 5 μm; b = 10 μm.

Fig. 4. Grammothele fuligo saprophytic on the petiole base of the brackish water palm Nypa fruticans (Photo
by Rattaket Choeyklin).

19
 Fig. 5. Aegerita propagules on senescent stems of Acanthus ilicifolius, Mai Po Mangrove, Hong Kong SAR.

Fig. 6. Butt heart rot of Xylocarpus granatum tree in a Khanom mangrove in southern Thailand.

20
 Fungal Diversity

Section B: acospores hyaline to slightly pigmented, 1-septate

PHYLUM: ASCOMYCOTA occasionally 3-septate, sometimes enclosed in a
sheath, anamorphs hyphomycetes or Coelomycetes ..
PEZIZOMYCOTINA .......................... Capnodiales (Mycosphaerellaceae)

1. Ascomata with bitunicate asci.................................. 2 PLEOSPOROMYCETIDAE

1. Ascomata with unitunicate asci................................ 5
One order with marine taxa
2. Lichen forming ascomycetes.................................... 4
2. Non-lichenized ascomycetes.................................... 3 Ascomata perithecial, multiloculate, thyrothecium or
cleistothecial-like, globose, thick-walled, immersed or
3. Ascomata perithecial..............Dothideomycetes (21) erumpent, well-developed ostiole, interascal tissue
3. Ascomata predominantly cleistothecial, cellular or trabeculate pseudoparaphyses, asci
gymnothecial, globose, asci evanescent..................... cylindrical, fissitunicate, apical apparatus, ascospores
................................................. Eurotiomycetes (56) brown, septate, muriform, often with a gelatinous sheath.
Anamorphs hyphomycetes .........................Pleosporales
4. Ascomata apothecial, marine species with no known
phycobiot ..............................Lecanoromycetes (66) Key to marine taxa in the Dothideomycetes
4. Ascomata apothecial, various phycobionts ................
................................................. Eurotiomycetes (56)
4. Thallus varied, sometimes poorly developed or 1. Ascomata perithecial................................................ 2
absent ......................................Arthoniomycetes(69) 1. Ascomata cleistothecial ......Pontoporeia biturbinata
1. Ascomata apothecial .............................................. 56
5. Parasitic on insects ...........Laboulbeniomycetes (66)
5. Ascomycetes not parasitic on insects ....................... 6 2. Ascospores 1-septate................................................ 3
2. Ascospores multi-septate ....................................... 24
6. Lichen forming ascomycetes with cyanophycean 2. Ascospores muriform............................................. 47
photobionts............................... Lichinomycetes (69)
6. Non-lichenized ascomycetes.................................... 7 3. Ascospores hyaline .................................................. 4
3. Ascospores brown.................................................. 17
7. Ascomata perithecial.............. Sordariomycetes (72)
7. Ascomata apothecial .................. Leotiomycetes (68) 4. Ascospores with central septum............................... 5
4. Ascospores with eccentric septum ......................... 14
DOTHIDEOMYCETES 5. Ascospores lacking appendages............................... 6
5. Ascospores with appendages ................................... 8
Two subclasses with marine taxa
6. Growing on mangrove wood .............Acrocordiopsis
1. Ascomycetes lacking paraphyses, pseudo- 6. Growing on algae..................................................... 7
paraphyses or paraphysoids, asci bitunicate globose,
ellipsoidal or subcylindrical ...... Dothideomycetidae 7. Hosts Fucales or Laminariales ........... Thalassoascus
1. Ascomycetes with cellular or trabeculate 7. Host the red alga Chondrus crispus .. Lautitia danica
pseudoparaphyses, bitunicate, asci, cylindrical,
clavate, oblong saccate............. Pleosporomycetidae 8. Ascospores with bipolar cilia-like appendages,
ascospores 16.5-24 × 5.5-7 µm, on Juncus................
DOTHIDEOMYCETIDAE .................................................... Heleiosa barbatula
8. Ascospores with a fine halo of fine hairs, 18-27 × 8-
Two orders with marine taxa 14 µm..................................Capillataspora corticola
8. Ascospores with mucilaginous appendages............. 9
1. Ascomata variable in morphology and structure,
formed by lysigenous locules within stromatic tissue, 9. Ascospores with lenticular appendage on side, 17-28
interascal tissue lacking, asci variable morphology, × 8-12 µm .............................. Paraliomyces lentifer
develop in basal fascicle, thick-walled, generally 9. Ascospores lacking a lenticular appendage ........... 10
fissitunicate, rarely with apical apparatus, ascospores
hyaline to brown, septate constricted at the central 10. Ascospores with tuberculate wall and surrounded
septum, occasionally muriform...............Dothideales by a sheath, 17-25 × 10-15 µm ...............................
............................................ Belizeana tuberculata
1. Ascomata small, immersed in host tissue, single or 10. Ascospores not tuberculate ................................ .11
superficial, or imbedded in a pseudoparenchymatal
stroma, asci ovoid to saccate to subcylindrical, 11. Ascomata in a stroma................. Scirrhia annulata

21
11. Ascomata not stromatic....................................... 12 25. Ascospores pale yellowish to brown................... 28

12. Ostiole slit-like ...................................Lophiostoma 26. Ascomata superficial, crater-like, on mangrove

12. Ostiole not slit-like.............................................. 13 wood................................. Ascocratera manglicola
26. Ascomata immersed............................................ 27
13. Ascomata lenticular, ascospores 18.5-27 × 4-6 µm,
on Nypa ............................ Astrosphaeriella nypae 27. On Juncus, ascospores with a sheath, ascospores
13. Ascomata subglobose, on mangrove wood ............. 45-59 × 5.5-10 µm ...............Loratospora aestuarii
................................................................Massarina 27. On Nypa palm, ascospores with a sheath and apical
cellular appendages, 31-42 × 7.5-12.5 µm..............
14. Ascospores with a sheath ....................... Didymella ............................................ Herpotrichia nypicola
14. Ascospores without a sheath ......... Mycosphaerella 27. On mangrove wood, ascospores with or without a
14. Ascomata superficial with stalk, ascospores 80- sheath .....................................................Massarina
109 × 18-34 µm............Manglicola guatemalensis
14. Ascomata immersed in substratum...................... 15 28. Ascospores versicolored ..................................... 29
28. Ascospores uniformly coloured .......................... 31
15. Ascomata in mangrove wood .............................. 16
15. Ascomata on algae ..................................Pharcidia 29. Ascospores striated, 31-38 × 6-9 µm ......................
................................. Astrosphaeriella striataspora
16. Ascospores with a delicate sheath, multiloculate .... 29. Ascospores not striated ....................................... 30
............................................................... Helicascus
16. Ascospores with bipolar germ tubes, no sheath,
30. Ascospores longer than 45 µm........... Passeriniella
ascospores 59-72 × 24-30 µm .................................
30. Ascospores 28-34 × 10-14 µm, on Spartina and
............................................... Salsuginea ramicola
driftwood............................. Byssothecium obiones
17. Ascomata in a loose stroma, ascospores 37-60 ×
31. Ascomata with pseudoparaphyses....................... 32
16-26 µm, on Posidonia........Halotthia posidoniae
31. Ascomata lacking pseudoparaphyses, ascospores
17. Ascomata not stromatic....................................... 18
yellowish, 50-70 × 10-14 µm..................................
...............................................Wettsteinina marina
18. Ascomata mammiform, ascospores with bipolar
germ tubes, 22-30 × 10-12 µm ................................
32. Pseudoparaphyses trabeculate............................. 33
..................................... Caryosporella rhizophorae
32. Pseudoparaphyses cellular .................................. 34
18. Ascomata not mammiform and lacking bipolar
germ tubes ........................................................... 19
33. Ascospores fuscous, not eu-septate, 30-41 × 10-16
19. Ascomata superficial or semi-immersed ............. 20 µm .............................Trematosphaeria mangrovei
19. Ascomata immersed in substratum...................... 21 33. Ascospores black, eu-septate, 30-44 × 12-17 µm ..
........................................Caryospora australiensis
20. Ascomata reddish-brown, subglobose, superficial
ascospores 24-32 × 12-15 µm ................................ 34. Ascomata with thick peridial wall....Leptosphaeria
.............................................. Bicrouania maritima 34. Ascomata with thin peridium..........Phaeosphaeria
20. Ascomata brown, semi-immersed, ascospores 14-
21 × 5-8 µm.............. Kirschsteiniothelia maritima 35. Ascospores hyaline ............................................. 36
35. Ascospores brown............................................... 38
21. Ascospores verrucose.......................................... 22
21. Ascospores not verrucose.................................... 23 36. Ascomata superficial on mangrove wood,
ascospores with a sheath and apical appendage, 42-
22. Ascospores brown 18-23 × 9-11 µm ....................... 50 × 7.5-10 µm.............. Falciformispora lignatilis
................................... Didymosphaeria lignomaris 36. Ascomata immersed............................................ 37
22. Ascospores dark brown 16-23 × 7-11 µm ...............
..................................................Verruculina enalia
37. Pseudoparaphyses trabeculate, ascospores 5-
septate, 51-80 × 14-22 µm .......Quintaria lignatilis
23. Ascospores striate, 23-33 × 9-12 µm ......................
37. Paraphyses present, ascospores 3-6-septate ............
............................................ Lineolata rhizophorae
............................................................ Sphaerulina
23. Ascospores not striate, 36-60 × 16-24 µm ..............
....................................... Coronopapilla mangrovei
38. Ascospores with septa at the ends, 55-83 × 16-25
24. Ascospores 1-3-septate........................................ 25 µm ..........................................Biatriospora marina
24. Ascospores more than 3-septate .......................... 35 38. Septa uniformly spread along ascospores ........... 39

25. Ascospores hyaline.............................................. 26 39. Ascomata superficial........................................... 40

22
 Fungal Diversity

39. Ascomata immersed, or under a clypeus, µm ............................................ Pleospora pelagica

occasionally erumpent......................................... 42 Ascospores 24-38 × 10-13 µm .............P. spartinae
52. Growing on other substrata ................................. 53
40. Ascospores lacking appendages, 30-37.5 × 7.5-
11.5 µm .............................Leptosphaeria nypicola 53. On Juncus ........................................................... 54
40. Ascospores with appendages............................... 44 53. On mangrove wood............................................. 55

41. Ascomata large on Nypa palm, single polar 54. Asci with a gel-cap, ascospores with pronounced
appendage, ascospores 31-52.5 × 5-10.5 µm .......... sheath, 26-40 × 11-19.5 µm. ..Tremateia halophila
...........................................Tirisporella beccariana 54. Asci lacking a gel cap, ascospores with no
41. Ascospores with a sheath, 7-14-septate................... gelatinous sheath, 18-22.5 × 5-6.5 µm....................
....................................................Massariosphaeria .....................................................Julella herbatilis

42. Ascospores lacking a sheath or appendages........ 43 55. Ascomata with hyphal mycelium, ascospores with
42. Ascospores with a sheath or appendages............. 44 a faint sheath, 20-28.5 × 8-11 µm ...........................
................................. Leptosphaerulina mangrovei
43. Ascomata on Ammophila, ascospores 5-8-septate 55. Ascomata lacking ascomatal hyphae, ascospores
35-55 × 12-16 µm .....Amarenomyces ammophilae 28-36 × 12-16 µm ..................... Julella avicenniae
43. Ascomata on other substrata ...........Phaeosphaeria
56. Ascospores with polar setae, hyaline to pale brown,
44. Ascospores versicolor with hyaline end-cells, 15-27 × 6-10 µm .................... Banhegyia setispora
ascospores 34-48 × 7-10 µm ................................... 56. Ascospores lacking polar appendages or a
..............................Trematosphaeria lineolatispora prominent sheath................................................. 57
44. Ascospores uniformly brown. ............................. 45
57. Ascospores hyaline, 5-8-septate, 18-41 × 11-11.5
45. Ascospores 4-7-septate........................................ 46 µm ...................................... Gloniella clavatispora
45. Ascospores with more than 7-septate ...................... 57. Ascospores hyaline, 4-9-septate, 24-48 × 6-24 µm.
............................................................. Carinispora ..................................................... Patellaria atrata
*The two marine Paraphaeosphaeria species are not
46. Ascospores 3-7-septate....................Phaeosphaeria obligately marine.
46. Ascospores 4-5-septate.......... Paraphaeosphaeria*

47. Ascospores hyaline and thick-walled ... Lautospora DOTHIDEOMYCETES

47. Ascospores brown ............................................... 48 DOTHIDEOMYCETIDAE
48. Ascospores with a sheath with 2-3 subconical CAPNODIALES
gelatinous appendages, 33-48 × 12-21 µm.............. Mycosphaerellaceae
......................................... Decorospora gaudefroyi
48. Ascospores with a sheath but lacking subconical Mycosphaerella Johanson, Oefvers., Foerh. K.
appendages .......................................................... 49 Sven. Vetensk.-Akad. 41: 163, 1884 ........... (1)
M. pneumatophorae Kohlm., Ber. Dtsch. Bot.
49. Ascospores with lighter end-cells, trabeculate
hamathecium ............................................. Aigialus
Ges. 79: 32, 1966.
49. Ascospores uniformly brown .............................. 50 M. salicorniae (Auersw.) Lindau, Hilfsb.
Sammeln Ascomyc.: 103, 1903.
50. On driftwood associated with sand ..................... 51 Sphaeria salicorniae Auersw., Bot.
50. On other substrata ............................................... 52 Trauschveerein, 1863.
Lizonia salicorniae (Auersw.) Auersw., Bot.
51. Ascospores 5-septate, end-cells slightly paler......... Trauschveerein, 1869.
................................. Platystomum scabridisporum Sphaerella salicorniae (Auersw.) Auersw.,
51. Ascospores 8-9-septate, uniformly dark brown....... Gonnermann & Rabenhorst Mycol. Eur. No. 5: 16, 1869.
............................................... Decaisnella formosa Sphaerella peruviana Speg., An. Soc. Cient.
Argent. 12: 115, 1881.
52. Growing on algae, ascospores 28-29.5 × 13-13.5 M. staticicola (Pat.) Dias, Mem. Soc. Brot. 21:
µm ........................................Pleospora gracilariae 72, 1970.
Ascospores 25-35 × 12-17 µm ............. P. pelvetiae Sphaerella staticicola Pat., Cat. Raisonné Plantes
52. Growing on salt marsh plant Triglochin, cellilaires Tunésie, Paris p. 104, 1897.
ascospores 45-65 × 16-25 µm .................................
........................................Pleospora triglochinicola
M. suaedae-australis Hansf., Proc. Linn. Soc.
52. Growing on Spartina ascospores 35-52 × 10-15 N.S.W. 79: 122, 1954.

23
 Ascomata globose, subglobose, ellip- appendages (Fig. 7). These species are in need
soidal, immersed in the host, conical, of modern taxonomic treatment. Although S.
membranous, ostiolate, epapillate or a short orae-maris is accepted as an obligate marine
papilla, pale coloured, solitary or gregarious, fungus (Kohlmeyer and Volkmann-Kohlmeyer,
pseudoparaphyses absent, asci elongate- 1991a), the marine niche of S. albispiculata
cylindrical to short clavate, short pedunculate, has been queried (Kohlmeyer and Kohlmeyer,
thick-walled, bitunicate, no apical apparatus, 1979). The latter was described by Tubaki
ascospores ellipsoidal to elongate, 1-septate, (1957) from driftwood on Tane Island, Kyushu,
hyaline, with a mucilaginous sheath. Pycnidia Japan and does not appear to have been
or spermatogonia may be present. A well reported in the literature since. However, Jones
characterized genus, primarily of circa 500 (unpublished data) has collected it on
terrestrial species causing leaf spot disease of a driftwood in Friday Harbour, USA. It can be
wide range of hosts. Marine taxa are generally distinguished from S. orae-maris by its well-
on the salt marsh plants Armeria, Limonium, developed, bushy, white, thick and hairy neck.
Salicornia and Suaeda. Ascomata immersed, Kohlmeyer and Kohlmeyer (1979) query
generally lacking pseudoparaphyses, asci short if these species belong in Sphaerulina as
clavate and ascospores hyaline, one septate pseudoparaphyses and bitunicate asci were not
often with a gelatinous sheath, e.g. M. conclusively demonstrated in the type material.
salicorniae. Mycosphaerella pneumatophorae
occurs on the “bark” of pneumatophores of
Avicennia species, with recent records from
Asian mangroves (Jones, unpublished data).
Mycosphaerella species occurring on algae are
now referred to Mycophycias.

1. Mainly on mangrove pneumatophores (Avicennia),

ascospores 14-21 × 6-8.5 µm ....M. pneumatophorae
1. On other marine or marsh plants, ascospores
narrower than 6 µm.................................................. 2

2. Ascospores longer than 18 µm, saprobic on Suaeda

australis ...................................M. suaedae-australis
2. Ascospores shorter than 18 µm ................................ 3
Fig. 7. Sphaerulina albispiculata. Ascospore 5-septate,
3. Ascospores 8-18 × 2.6 µm, saprobic on Salicornia not constricted at the septa. Bar = 10 μm
and Suaeda ......................................... M. salicorniae
3. Ascsopores 12-15 × 4-6 µm, saprobic on Armeria
1. Ascomata with a prominent bushy, white hairy neck,
and Limonium....................................... M. staticicola
ascospores 25-30 × 5-6 µm, 5-6-septate ....................
.......................................................... S. albispiculata
Sphaerulina Sacc., Michelia 1: 399, 1878 .. (2) 1. Ascomata with a short papillate neck, ascospores 26-
S. orae-maris Linder, Farlowia 1: 413, 1944. 32 × 5-8 µm, 3-septate ......................... S. orae-maris
S. albispiculata Tubaki, Publs. Seto Mar. Biol.
Lab. 15: 366, 1957. Pharcidia Körber., Parerga Lichenologica,
Ascomata globose, subglobose, Breslau, p. 469-470, 1865. .......................... (3)
immersed, ostiolate, papillate, membranous, P. balani (G. Winter) Bausch, Pubbl. Stn. Zool.
hyaline to light brown, periphyses present or Napoli 15: 379, 1936.
absent, paraphyses present or absent, filiform, Epicymatia balani G.Winter ex Har., Jour. De
simple, septate, asci clavate to cylindrical, Bot. 1: 233, 1887.
short pedunculate, unitunicate?, apically P. laminariicola Kohlm., Bot. Mar. 16: 209,
thickened, without an apical apparatus, 1973.
ascospores 3-6-septate, slightly constricted at P. rhachiana Kohlm., Bot. Mar. 16: 210, 1973.
the septa, hyaline, smooth-walled, no sheath or

24
 Fungal Diversity

Ascomata globose to ellipsoidal, small, DOTHIDEALES incertae sedis

solitary, ostiolate, epapillate, periphysate, dark BOTRYOSPHAERIALES
brown to black, paraphyses septate, ramose, Botryosphaeriaceae
reticulate, in a gelatinous matrix, asci clavate
to cylindrical, thick-walled, no apical appa- Amarenomyces O.E. Erikss., Op. Bot. Soc.
ratus, persistent, ascospores ellipsoidal to Bot Lund 60: 124, 1981 ............................... (1)
obovoid, 1-septate, slightly constricted at the A. ammophilae (Lasch.) O.E. Erikss. Op. Bot.
septum, hyaline, with or without appendages Soc. Bot. Lund 60: 124, 1981 (Type species).
(gelatinous cap-like and at both ends). Sphaeria ammophilae Lasch, Flora, Jena 8: 282,
1850.
Pharcidia contains some 90 names, but many Leptosphaeria ammophilae (Lasch) Ves. De Not.,
are referred to Stigmidium, or lichen genera Comment. Soc. Critt. Ital. 1: 236, 1863.
such as Arthopyrenia, and Lichenodiplis. They Phaeosphaeria ammophilae (Lasch) Kohlm. & E.
occur on brown algae, Laminaria digitata Kohlm., Icones Fungorum Maris, Plate 55, 1965.
(Zuccaro and Mitchell, 2005) or shells of Leptosphaeria littoralis Sacc., Michelia 1: 38,
1877.
marine animals, e.g. molluscs (Kohlmeyer and Phaeosphaeria littoralis (Sacc.) L. Holm, Symb.
Kohlmeyer, 1979). Bot. Upsal. 14: 121, 1957.
Sphaeria subuletorum Berk. & Br., Ann. Mag.
1. Ascomata on barnacles and other marine shells......... Nast Hist. Soc. Ser. 2, 9: 382, 1852.
.................................................................... P. balani Leptosphaeria sabuletorum (Berk. & Br.) von
1. Ascomata on algae ................................................... 2 Höhn. Hedwigia 60: 141, 1918.
Metasphaeria sabuletorum (Berk. & Br.) Sacc.,
2. Ascospores 18-25 × 6-8 µm, with polar cap-like Syll. Fung. 2: 180, 1883.
appendages.......................................P. laminariicola Sphaeria subulectorum Berk. & Br. 1952.
2. Ascospores 12-20 × 4-5 µm, without appendages ..... Montagnula perforans (Roberge ex Desm.)
..............................................................P. rhachiana Aptroot, Mycosphaerella and its Anamorphs: Conspetus
of Mycosphaerella, 2: 150, 2006.
Paradidymella perforans (Roberge ex Desm.)
1. DOTHIDEALES Munk, Dansk bot. Ark. 17: 179, 1957.
Dothideaceae Sphaeria perforans Roberge ex Desm., Ann. Sci.
Nat., Bot, ser. 2, 19: 23.
Scirrhia Nitschke ex Fuckel, Jahrb. Tarospora perforans (Roberge ex Desm.) Höhn
Nassauischen Vereins Naturk., Wiesbaden 23- Anamorph: Amarenographium meta-
24: 220, 1870 ............................................... (1) bleticum (Trail) O.E. Erikss.
S. annulata Kohlm., Volkm.-Kohlm., & O.E. Camarographium metableticum (Trail) Grove,
British Stem- and Leaf Fungi (Coelomycetes) 2: 108,
Erikss., Can. J. Bot. 74: 1835, 1996. 1937.
Described from senescent culms of Diplodina ammophilae Trail, Scot. Nat. I, P 76
Juncus roemerianus, it occurs 28-121 cm Ascomata subglobose, ellipsoidal to
above the rhizome within the range regarded pyriform, immersed, clypeate, ostiolate,
as obligate to facultative is present throughout papillate, carbonaceous, black, solitary or
the year and grows well on saltwater agar gregarious, pseudoparaphyses septate, simple,
(Kohlmeyer et al., 1996). Diagnostic features in a gelatinous matrix, asci clavate or subcylin-
are the linear stromata, 1-3 mm long, generally drical, short pedunculate, thick-walled, bituni-
superficial, multiloculate with ascomata in cate, without an apical apparatus, ascospores,
longitudinal rows, asci clavate, ascospores 3- ellipsoidal to fusiform, 5-8-septate, slightly
septate, brown, with a thin sheath, and 46-60 × constricted at the septa, yellowish to pale-
9-11.5 µm. Taxonomic position needs verifica- brown, with a mucilaginous sheath constricted
tion at the molecular level to resolve its at the central septum (Kohlmeyer and
phylogenetic position. Kohlmeyer, 1979). Assignment of this species

25
at the generic level is subject to debate. The taxonomic position needs verification at the
genus was erected based on this species molecular level, but few collections of this
(Eriksson, 1981) but Leuchtmann (1984) and ascomycete have been made, and may escape
Kohlmeyer and Kohlmeyer (1965) placed it in detection because of its inconspicuous
Phaeosphaeria. perithecia.

Belizeana Kohlm. & Volkm.-Kohlm., Bot. Passeriniella Berl., Icon. Fung. (Abellini) 1:
Mar. 30: 195, 1987....................................... (1) 51, 1891........................................................ (2)
B. tuberculata Kohlm. & Volkm.-Kohlm., Bot. P. mangrovei Maria & K.R. Sridhar, Indian J.
Mar. 30: 196, 1987 (Type species). Forst. 25: 319, 2002.
Ascomata subglobose to ampulliform, P. savoryellopsis K.D. Hyde & Mouzouras,
immersed to erumpent, ostiolate, epapillate or Trans Br. Mycol. Soc. 91: 179, 1988.
with a short papilla, carbonaceous, black, Ascomata globose to subglobose,
solitary or gregarious, hyaline cells filling immersed, ostiolate, papillate, dark brown to
ostiolar canal, pseudoparaphyses simple, rarely black, coriaceous, solitary or gregarious,
branching or anastomosing, asci cylindrical, periphysate, pseudoparaphyses hyaline,
short pedunculate, thick-walled, fissitunicate, branched, septate, asci cylindrical, pedun-
without an apical apparatus, with a small culate, bitunicate, with an ocular chamber,
ocular chamber, ascospores ellipsoidal, 1- ascospores ellipsoidal, versicolour, smooth-
septate, constricted at the septum, hyaline, walled, 3-septate, constricted at the septa,
thick-walled, two-layered, tuberculate central cell larger and brown, end cells small
ornamentations between the two layers, the and hyaline, lacking a sheath or appendages
outer gelatinizing and forming a sheath (Fig. 8). Perplexing species that require to be
(Kohlmeyer and Volkmann-Kohlmeyer, studied at the molecular level, since P. obiones
1987c). Although tentatively referred to the has been transferred to Byssothecium (Barr,
Pleosporaceae, Pleosporales by Kohlmeyer 2002), however this has not been universally
and Volkmann-Kohlmeyer (1987c), we prefer accepted (Index Fungorum).
to leave it in the Dothideales incertae sedis, Currently there is confusion over the
until a more suitable family suggests itself. placement of P. obiones (= Leptosphaeria
The ascospores of B. tuberculata are unusual discors). Molecular studies show that L.
in having a verrucose wall sourrounded by a discors does not belong in Leptosphaeria or
sheath, which becomes sticky in water. A Phaeosphaeria (Khashnobish and Shearer,
similar species has been collected in Malaysia 1996a, b), while the type species of
but further collections are necessary to Passeriniella has a chequered history
determine if it is a new species. (Kohlmeyer and Volkmann-Kohlmeyer,
1991a). Barr (2002) regards P. obiones as best
Capillataspora K.D. Hyde, Can. J. Bot. 67: placed in Byssothecium and we accept that
2522, 1989................................................... (1) view in this monograph. Passeriniella
C. corticola K.D. Hyde, Can. J. Bot. 67: 2522, savoryellopsis and P. mangrovei conform
1989 (Type species). neither to the generic characteristics of
An inconspicuous species occurring on Byssothecium or Passeriniella (Maria and
the bark of the prop roots of Rhizophora Sridhar, 2002). In P. savoryellopsis the
apiculata, with globose to ovoid ascomata, hamathecium tissue, asci, 3-septate ascospores
hyaline to pale brown, coriaceous, periphysate, (Barr, 2002 regards them as uniseptate with a
solitary and hairy around the ostiole (Hyde, short terminal papilla), that are large, deeply
1989a). Asci are thick walled, saccate to constricted at the central septum are not
clavate without an apical apparatus. features of either Byssothecium or
Ascospores 18-27 × 8-13.5 µm, 1-septate, Passeriniella. Further collections and a
constricted at the septum, hyaline surrounded molecular study are warranted before the two
by small fine, hair-like appendages. Its species can be referred to a new genus.

26
 Fungal Diversity

1. Asci with 8 ascospores, 44-62 × 17-24 μm ................ DOTHIDEOMYCETIDAE family incertae

............................................................. P. mangrovei sedis
1. Asci with 4 ascospores, 64-88 × 24-28 μm ................
........................................................P. savoryellopsis Lautosporaceae

Lautospora K.D. Hyde & E.B.G. Jones, Bot.

Mar. 32: 479, 1989....................................... (2)
L. gigantea K.D. Hyde & E.B.G. Jones, Bot.
Mar. 32: 479, 1989 (Type species).
L. simillima Kohlm., Volkm.-Kohlm. & O.E.
Erikss., Bot. Mar. 38: 169, 1995.
Ascomata subglobose to fusiform,
immersed, lying horizontal to wood surface,
coriaceous, ostiolate, light brown to brown,
Fig. 8. Passeriniella mangrovei. Versicolorous
solitary, pseudoparaphyses cellular, septate,
ascospore. Bar = 10 μm. asci cylindrical, four-spored, thick-walled,
bitunicate, pedunculate, with an ocular
Thalassoascus Oll., C.R. Hebd. Séances Acad. chamber and ring, ascospores fusiform,
Sci. 182: 1348-1349, 1926 ........................... (3) muriform, 4-7 longi-septa, hyaline, very thick-
T. cystoseirae (Oll.) Kohlm., Mycologia 73: walled, no sheath or appendages (Hyde and
837, 1981. Jones, 1989d) (Figs. 9, 10). Lautospora
Melanopsamma tregoubovii var. cystoeirae Oll., simillima is a second species assigned to the
Ann. Inst. Océangor. (Paris) (N.S.) 7: 172, 1930. genus, occurring on the culms of Juncus
T. lessoniae Kohlm., Mycologia 73: 837, 1981. roemerianus (Kohlmeyer et al., 1995b). It is
T. tregoubovii Oll., Compt. Rend. Habd. an obligate marine species occurring 3-12 cm
Séancs Acad. Sci 182: 1348, 1926 (Type above the rhizome. These two species appear
species). to be very similar and can only be
Melanopsamma tregoubovii (Oll.) Oll., Bull. Inst.
Océangor. 522: 3, 1928.
distinguished by ascospore dimensions, which
Melanopsamma tregoubovii var. cutleriae Oll., overlap. These species are known from few
Ann. Inst. Océangor. (Paris) (N.S.) 7: 172, 1930. collections and further studies are required to
Ascomata subglobose to ellipsoidal, determine if they are distinct species and also
superficial, subiculate, ostiolate, epapillate, their phylogenetic positions within the
periphysate, subcarbonaceous to leathery, Dothideomycetes. No anamorph has been
black, gregarious, pseudoparaphyses filiform, reported.
asci cylindrical to clavate, pedunculate, thick-
walled, ascospores ellipsoidal, 1-septate, 1. On mangrove wood, ascospores 140-195 × 36-45
constricted at the septum, hyaline, thick- µm............................................................L. gigantea
1. On Juncus roemerianus, ascospores 127-210 × 28-
walled. A little known genus with 1-septate 57 µm......................................................L. simillima
ascospores, differentiated primarily by
ascospore measurements and their algal hosts
and warrants further study.

1. On Cystoseira spp., ascomata stalked ........................

.............................................................T. cystoseirae
1. On other none Fucales, ascomata sessile ................. 2

2. Ascospores less than 28 μm in diam., on Lessonia

(Laminariales) .........................................T. lessoniae
2. Ascospores wider than 28 μm, on Aglaozonia spp. Fig. 9. Lautospora gigantea. Ascospores hyaline,
and Zanardinia (Cutleriales) .............. T. tregoubovii muriform and thick-walled. Bars = 20 μm.

27
 Coronopapilla avellina Kohlm. & Volkm.-
a b c Kohlm., Mycol. Res. 94: 687, 1990.
Ascomata immersed under a stroma,
subglobose, ostiolate, papillate, clypeate,
coriaceous, light-coloured, single or grega-
rious, pale brown, periphysate, pseudopara-
physes trabeculate, anastomosing filaments in
a gelatinous matrix, asci cylindrical, long
pedunculate, thick-walled, fissitunicate, J-,
without an apical apparatus, with an ocular
Fig. 10. Lautospora simillima. a. Mature ascospores in chamber, ascospores ellipsoidal, 1-3-septate,
ascoma. b. Thick-walled ascus. c. Muriform and thick- constricted at the central septum, thick-walled
walled ascospores. Bars = 50 μm. and two layered, no sheath or appendages
Planistromellaceae (Kohlmeyer and Volkmann-Kohlmeyer,
1990a). The peridium is hyaline and soft,
Loratospora Kohlm. & Volkm.-Kohlm., Syst. ascospores are not ornamented and lack a
Ascomycetum 12: 10, 1993 ......................... (1) sheath, characters that separate it from
L. aestuarii Kohlm. & Volkm.-Kohlm., Syst. Caryospora. Originally described by Hyde
Ascomycetum 12: 10, 1993 (Type species). (1989c) as a Caryospora species, but it lacks
This species occurs on Juncus the characteristic features of that genus:
roemerianus culms in an intermediate position erumpent to superficial ascomata, peridium
(10-61 cm above the rhizome) between that carbonaceous, and ascospores surrounded by a
regarded as obligately and facultatively marine gelatinous sheath (Kohlmeyer and Volkmann-
(Kohlmeyer and Volkman-Kohlmeyer, 1993c). Kohlmeyer, 1990a). Coronopapilla was
Ascomata black immersed in the culms, initially referred to the Didymosphaeriaceae
carbonaceous, ostiolate, neck with periphyses, (Kohlmeyer and Volkmann-Kohlmeyer,
asci clavate, thick-walled, fissitunicate without 1990a).
an apical apparatus, with an ocular chamber, J-,
while ascospores are hyaline, 3-septate Caryospora De Not. Micromyc. Ital. Novi 9: 7,
surrounded by a thin mucilaginous sheath. 1855.............................................................. (1)
Loratospora can be compared with Lepto- C. australiensis M.A. Abdel-Wahab & E.B.G.
sphaerulina, Monascostroma and Wettsteinina Jones, Mycoscience 41: 379, 2000.
but they differ in the lack of periphyses, their The only marine species assigned to this
small ascomata, their thin walled ascomatal genus and collected on wood associated with
wall and asci that are ovoid to saccate. sand in Australia (Abdel-Wahab and Jones,
Barr (1996) erected the family 2000). Ascomata large 870-1000 µm, conical
Planistromellaceae for six genera in the to subglobose, immersed to erumpent,
Dothideales, including Loratospora, because carbonaceous, papillate, periphysate, solitary
their locules open schizogenously by a or gregarious, pseudoparaphyses trabeculate
periphysate ostiole. anastomosing above the asci, asci cylindrical
to subcylindrical with a wide opercular
Zopfiaceae chamber, ascospores dark brown to black, the
end-cells paler, thick-walled, 3-eu-septate and
Coronopapilla Kohlm. & Volkm.-Kohlm., lacking a sheath (Fig. 11a). It differs from
Mycol. Res. 94: 686, 1990 .......................... (1) Caryospora species in possessing cylindrical
C. mangrovei (K.D. Hyde) Kohlm. & Volkm.- asci and ascospores lacking a sheath. In
Kohlm., Bot. Mar. 34: 19, 1991 (Type species). Caryospora asci are usually saccate and not
Caryospora mangrovei K.D. Hyde, Trans. Mycol. cylindrical as in C. australiensis.
Soc. Jpn. 30: 336, 1989.

28
 Fungal Diversity

assigning it to Zopfiaceae, with Barr (1979a)

a b and Hawksworth (1979) referring it to the
Massarinaceae and Testudinaceae, respectively.
We prefer at this stage to retain P.
biturbinata in the Zopfiaceae because of ascus
and ascospore morphology, however we do
not consider it well placed in Zopfia. Malloch
and Cain (1972) offer few details as to why the
species should be transfered to Zopfia. Recent
collections of the species have enabled a better
Fig. 11. a. Caryospora australiensis. Three-septate understanding of its phylogenetic relationship
ascospore. b. Platystomum scabridisporum. Muriform
ascospores. Bars a = 10 μm; b = 20 μm.
(Suetrong et al., unpublished data).

Pontoporeia Kohlm., Nova Hedw. 6: 5-6, DOTHIDIOMYCETES

1963.............................................................. (1) PLEOSPOROMYCETIDAE
◙ P. biturbinata (Durieu & Mont.) Kohlm., PLEOSPORALES
Nova Hedw. 6: 5, 1963 (Type species).
Sphaeria biturbinata Durieu & Mont., Explo. Sci. The number of families in the
Algérie, Bot. 1: 497. 1849. Pleosporales is unresolved with Schoch et al.
Amphisphaeria biturbinata (Durieu & Mont.) (2006) referring to only six (Leptosphae-
Sacc., Syll. Fung. 1: 729, 1882.
Zopfia biturbinata (Durieu & Mont.) Malloch &
riaceae, Lophiostomataceae, Phaeosphae-
Cain, Can. J. Bot. 50: 67, 1972. riaceae, Pleosporaceae, Sporormiaceae,
Ascomata large 805-1375 µm high, 805- Testudinaceae), while Myconet (2007) and
1120 µm diam., globose, dark brown to black, Kirk et al. (2001) list 17 and 19, respectively.
thick-walled, pseudoparaphysate, asci bituni- Other families have been proposed and are
cate, clavate with no apical apparatus, long cited in Kirk et al. (2001) as Pleosporales
tapering pedicel and ascospores thick-walled, incertae sedis. We refer marine taxa to nine
dark to blackish brown, 1-septate with a families within the order. Greater resolution of
prominent hyaline germ pore, 66-90 × 32-44 the familial position of these taxa requires
µm (Fig. 12). Described from the rhizomes of greater taxon sampling and a multigene
Posidonia oceanica (Kohlmeyer, 1963), and approach.
appears to be host specific as is Halotthia
posidoniae. Cuomo et al. (1985) found both P. 1. Didymosphaeriaceae
biturbinata and H. posidoniae common on
Posidonia oceanica rhizomes with a frequency Didymosphaeria Fuckel, Jahrb. Nassau.Ver.
of occurrence of 52% and 78%, respectively, Naturkd. 23-24: 140, 1870 ........................... (1)
but not on Cymodocea nodosa, another D. lignomaris Strongman & J.D. Mill., Proc.
seagrass growing in the same locality. Our N. S. Inst. Sci. 35: 102, 1985.
examination of P. oceanica from Cyprus A doubtful Didymosphaeria species
(collections in December, 2007 and February, infrequently collected and requires further
2008) noted that H. posidoniae was quite investigation (Strongman et al., 1985; Peña
common, but that P. biturbinata was rare. and Arambarri, 1998a). Aptroot (1995)
Currently some 16 Zopfia names are monographed the genus Didymosphaeria with
known, but many of these have been 550 described names, accepted 7 species and
transferred to other genera and families: transferred 100 species to other genera. He
Ulospora, Neotestudina, Lepidosphaeria, regarded D. lignomaris miss-placed in
Testudina (Testudinaceae), Zopfiofoveola Didymosphaeria and suggested placement in
(Zopfiaceae). Taxonomic assignment of Zopfia Lojkania cf enalia (Barr, 1990a). However,
is problematic with Von Arx and Műller (1975) this has not been universally accepted.

29
2. Testudinaceae L. nypicola K.D. Hyde & Alias, Mycol. Res.
103: 1414, 1999.
Verruculina Kohlm. & Volkm.-Kohlm., L. pelagica E.B.G. Jones, Trans. Br. Mycol.
Mycol. Res. 94: 689, 1990 .......................... (1) Soc. 45: 105, 1962.
◙ V. enalia (Kohlm.) Kohlm. & Volkm.- L. peruviana Speg., An. Soc. Cient. Argent.
Kohlm., Mycol. Res. 94: 689, 1990 (Type 12: 179, 1881.
species). Ascomata conical, subglobose,
Didymosphaeria enalia Kohlm., Ber. Deutsch. obpyriform, ostiolate, papillate, coriaceous or
Bot. Ges. 79: 28, 1966.
Lojkania enalia (Kohlm.) M.E. Barr, N. Amer.
carbonaceous, light brown to brown to black,
Fl., Ser. 2. 13: 56, 1990. solitary or gregarious, periphysate, pseudo-
Ascomata subglobose, ampulliform or paraphyses septate, branched with gelatinous
ellipsoidal, immersed, ostiolate, papillate, walls, asci cylindrical or subclavate-clavate-
clypeate, carbonaceous, black solitary or fusiform, short pedunculate, thick-walled,
gregarious, pseudoparaphyses trabeculate, lacking or with an apical apparatus, ascospores
anastomosing, in a gelatinous matrix, asci ellipsoidal, fusiform or cylindrical, 3 or more-
cylindrical, short pedunculate, J-, without an septa, slightly constricted at the septa, hyaline
apical apparatus, with an ocular chamber, to pale yellow-brown, with or without a sheath.
thick-walled, fissitunicate, ascospores ellip- Morphological characters that best describe
soildal, 1-septate, slightly constricted at the Leptosphaeria are: “superficial, conoid
septum, dark-brown, verrucose, lacking a ascomata that are strongly thickened toward
sheath or appendage (Fig. 13). the base, thick-walled scleroplec-tenchyma in
A species originally described as a the peridium, ascospores that are uniformly
Didymosphaeria species, but transferred to pale brown or hyaline and hosts that are
Lojkania (Barr, 1990a) and subsequently to primarily dicotyledons” (Khashnobish and
Verruculina (Kohlmeyer, and Volkmann- Shearer, 1996b). The genus is polyphyletic and
Kohlmeyer, 1990a) with its immersed clypeate in need of a thorough study of wider range of
ascomata with a dark peridium, a gelatinous taxa (Cannon and Kirk, 2007).
matrix around the pseudoparaphyses, asci that Currently five species are marine
are stipitate with an ocular chamber and occurring on mangrove substrata, generally
verruculose dark brown ascospores wood and maritime plants. Shoemaker and
(Kohlmeyer and Volkmann-Kohlmeyer, Babcock (1989) and Kohlmeyer and
1990a). Initially referred to the Volkmann-Kohlmeyer (1991a) retain
Didymosphaeriaceae, Melanommatales by Leptosphaeria orae-maris in Leptosphaeria,
Kohlmeyer and Volkmann-Kohlmeyer (1990a), while Khashnobish and Shearer (1996a, b),
sequence data place it in the Testudinaceae as have transferred it to Phaeosphaeria.
the most basal clade of the Pleosporales along Morphologically Leptosphaeria and
with Lepidosphaeria nicotiae and Ulospora Phaeosphaeria are difficult to delineate with
bilgramii (Schoch et al., 2006). confidence as is evidenced by the frequent
transfer of species from one genus to the other.
3. Leptosphaeriaceae Khashnobish and Shearer (1996a) opined that
the only useful characters in the delineation of
Leptosphaeria Ces. & De Not., Comment. Soc. these two genera were: “shape of ascoma,
Crittogam. Ital. 1: 234, 1863 ........................ (5) ascomal position relative to the substrate and
L. australiensis (Cribb & J.W. Cribb) G.C. the type of cells making up the peridium”.
Hughes, Syesis 2: 132, 1969. However, phylogenetically the two genera
Metasphaeria australiensis Cribb & J.W. Cribb, cannot be separated, with Phaeosphaeria
Univ. Queensl., Pap. Dept. Bot. 3: 79, 1955. forming a natural group, while Leptosphaeria
L. avicenniae Kohlm. & E. Kohlm., Nova did not form a monophyletic group
Hedw. 9: 98, 1965. (Khashnobish and Shearer, 1996b).

30
 Fungal Diversity

a b e

c d g

Fig. 12. Pontoporeia biturbinata. a. Front view of mature ascomata (cleistothecial) of Pontoporeia biturbinata on
rhizomes of the sea grass Posidonia oceanica. b. Pseudoparaphyses and immature ascus. c-d. Thick-walled, ascus
with long tapering pedicel. e-i. Ascospores bicelled and thick-walled. Bars a = 100 µm; b-i = 10 µm.

31
 a b

c d

f g

h i

Fig. 13. Verruculina enalia. a. Ascomata submerged in mangrove wood. b. Thin-walled pseudoparaphyses. c.
Cylindrical asci and pseudoparaphyses. d. Ascus cylindrical with an apical pore. e-i. Ascospores bicelled and
constricted at the septum. Bars a = 100 µm; b-i = 10 µm.

32
 Fungal Diversity

The families Leptosphaeriaceae and asci cylindrical, pedunculate, fissitunicate,

Phaeosphaeriaceae are closely related as is with a refractive ring in the endoascus,
evident from recent sequence data ascospores fusiform to elongate, muriform,
(Khashnobish and Shearer, 1996a; Cámara et distoseptate, 8-13 transverse septa, 1-6
al., 2002; Kodsueb et al., 2006; Schoch et al., longitudinal septa, slightly constricted at the
2006; Suetrong et al., unpublished data). septa, golden-brown, smooth-walled, no
Hibbett et al. (2007) refrained from sheath or appendages (Abdel-Wahab and
classification below ordinal level therefore Jones, 2003) (Fig. 14). Known only from the
offer no solution as to the validity of retaining type locality Mornington Peninsula, Australia,
the two families in their present form. on driftwood associated with sand, where it
Molecular data from Cámara et al. (2002) was common (Abdel-Wahab and Jones, 2003).
support the separation of Leptosphaeria and It conforms to the generic charac-teristics of
Phaeosphaeria, and consider that the large ascomata, wide peridium, refractive
significant morphological characters phylo- apical ring surrounding an ocular chamber in
genetically are: peridial characters, anamorphs the ascus and distoseptation in immature
and plant hosts. Kodsueb et al. (2006) question ascospores that are brown and smooth-walled.
whether the Phaeosphaeriaceae is a synonym Molecular data shows that D. formosa is well
for the Leptosphaeriaceae and advocate a re- placed in the Lophio-stomataceae with high
evaluation based on wider sampling and bootstrap support, forming a sister group
multigene sequence analyses. Cannon and comprising Massarina and Lophiostoma
Kirk (2007) accept both families. species (Fig. 17) (Suetrong et al., 2009 in
press).
1. Ascospores with a wide sheath ............................... 2
1. Ascospores with an indistinct sheath or lacking a Herpotrichia Fuckel, Fungi rhenani exsic. No.
sheath ....................................................................... 3 2171, 1868.................................................... (1)
H. nypicola K.D. Hyde & Alias, Mycol. Res.
2. Ascospores 18-25 × 6-8 µm, on mangrove wood,
tropical ................................................. L. avicenniae
103: 1412, 1999.
2. Ascospores 28-44 × 8-12 µm, on wood, temperate ... Ascomata globose, superficial, ostiolate,
................................................................. L. pelagica short papillate, gregarious, black, pseudo-
paraphyses filiform, numerous, septate,
3. Ascospores with 3 septa .......................................... 4 anastomosing above the asci, in a gelatinous
3. Ascospores with 3-5 septa, 30-37.5 × 7.5-11.5 µm,
indistinct sheath, on Nypa ........................L. nypicola
matrix, asci cylindrical-clavate, pedunculate,
thick-walled, bitunicate, with an ocular
4. Ascospores 12-16 × 4-5.5 µm, olive brown, on chamber, ascospores fusiform, 1-septate
Salicornia.............................................. L. peruviana becoming 3-septate, constricted at the central
4. Ascospores 19-27 × 6-9 µm, hyaline, on wood ......... septum, basal part longer and narrower than
..........................................................L. australiensis
the apical part, hyaline becoming pale brown,
with cellular apical appendages (Hyde et al.,
4. Lophiostomataceae 1999b).
The only species known from a marine
Decaisnella Fabre, Annls. Sci. Nat. Bot. Sér. 6, habitat, it has black, globose, superficial
9: 112, 1879 ................................................. (1) papillate ascomata often clustered together,
◙ D. formosa Abdel-Wahab & E.B.G. Jones, pseudoparaphyses filiform, numerous, septate
Can. J. Bot. 81: 598, 2003. and anastomosing above the asci, asci
Ascomata subglobose, immersed, bitunicate, cylindric-clavate with an ocular
clypeate, ostiolate, coriaceous, black, solitary chamber, ascospores 1-3-septate, hyaline to
or gregarious, periphysate, pseudoparaphyses pale brown becoming dark brown at maturity,
trabeculate, unbranched at the base, anasto- with apical cellular appendages (Hyde et al.,
mosing above the asci, in a gelatinous matrix, 1999b).

33
 a b

c d e f

g h i

Fig. 14. Decaisnella formosa. a-b. Asci and thin-walled pseudoparaphyses. c. Apical region of ascus with a pore.
d-i. Muriform brown ascospores. Bars a-i = 10 µm.

34
 Fungal Diversity

Described from the intertidal petiole of These species have been transferred
Nypa fruticans. The genus Herpotrichia from Massarina based on morphological and
groups in Lophiostomataceae group 2 of a molecular evidence (Liew et al., 2002).
multigene analyses by Schoch et al. (2006) Lophiostoma asiana and L. mangrovei have
and further resolution at the family level been transferred to Astrosphaeriella (Hyde et
requires greater species sampling. al., 2000).

Lophiostoma Ces. & De Not., Comm. Soc. 1. Ascospores 24-33 × 6-10 μm, on the fern
Acrostichum ...........................................L. acrostichi
Crittog. Ital. 1: 219, 1863............................. (3) 1. Ascomycetes on mangrove wood ............................ 2
The taxonomic position of marine
Astrosphaeriella, Lophiostoma and Massarina 2. Ascospores 28-39 × 7-10 μm..........L. armatisporum
species has been re-evaluated at the 2. Ascospores 22-28 (-33) × 4.5-6.5 μm ........................
ultrastructural and molecular level (Read et al., ........................................................... L. rhizophorae
1997a, b; Aptroot, 1998; Hyde and Aptroot,
1998; Hyde et al., 2002; Liew et al., 2002; Massarina Sacc., Syll. Fung. (Abellini) 2: 153,
Schoch et al., 2006). Earlier delineation of the 1883............................................................ (10)
genera based on slit-like versus rounded ?M. beaurivagea Poonyth, K.D. Hyde,
ostioles was found not to be consistent within Aptroot & Peerally, Fungal Diver. 3: 139,
a genus at the molecular level (Hyde et al., 1999.
2002). Marine species have been transferred M. cystophorae (Cribb & J.W. Herb.) Kohlm.
between these genera and the current & E. Kohlm., Marine Mycology. The Higher
assignment is based on the sequence data of Fungi: 427, 1979.
Liew et al. (2002). Otthiella cystophorae Cribb & J.W. Herb., Univ.
Queensl., Pap. Dept. Bot. 3: 10, 1954.
Melanopsamma cystophorae (Cribb & J.W.
L. acrostichi (K.D. Hyde) Aptroot & K.D. Herb.) Meyers, Mycologia 49: 485, 1957.
Hyde, Fungi in Marine Environments. Fungal M. lacertensis Kohlm. & Volkm.-Kohlm.,
Diversity Press: 106, 2002. Aust. J. Mar. Freshw. Res. 42: 92, 1991.
Massarina acrostichi K.D. Hyde, Mycol. Res. 93:
437, 1989.
?M. mauritiana Poonyth, K.D. Hyde, Aptroot
L. armatisporum (K.D. Hyde, Vrijmoed, & Peerally, Fungal Diver. 3: 141, 1999.
Chinnaraj & E.B.G. Jones) Liew, Aptroot & ◙ M. phragmiticola O.K. Poon & K.D. Hyde,
K.D. Hyde, Fungi in Marine Environments. Bot. Mar. 41: 145, 1998.
Fungal Diversity Press: 106, 2002. ◙ M. ramunculicola K.D. Hyde, Mycologia
Massarina armatispora K.D. Hyde, Vrijmoed, 83: 839, 1992.
Chinnaraj & E.B.G. Jones, Bot. Mar. 35: 325, 1992. M. ricifera Kohlm., Volkm.-Kohlm. & O.E.
L. rhizophorae (Poonyth, K.D. Hyde, Aptroot Erikss., Mycologia 87: 537, 1995.
& Peerally) Aptroot & K.D. Hyde, Fungi in ?M. rhizophorae Poonyth, K.D. Hyde,
Marine Environments. Fungal Diversity Press: Aptroot & Peerally, Fungal Diver. 3: 144,
108, 2002. 1999.
Massarina rhizophorae Poonyth, K.D. Hyde, M. thalassiae Kohlm. & Volkm.-Kohlm., Can.
Aptroot & Peerally, Fungal Divers. 3: 144, 1999. J. Bot. 65: 575, 1987.
Ascomata subglobose to elongate, ◙ M. velataspora K.D. Hyde & Borse,
immersed to erumpent, often strongly flattened, Mycotaxon 27: 161, 1986.
carbonaceous, papillate with a round or slit- Ascomata subglobose to obpyriform,
like ostiole, black, solitary to gregarious, immersed in the substratum under a
pseudoparaphyses cellular, in a gelatinous pseudostroma, or erumpent, ostiolate,
matrix, asci cylindrical, thick-walled, epapillate, clypeate, coriaceous, dark brown to
bitunicate, short pedunculate, with an ocular black, solitary or gregarious, periphysate,
chamber and faint ring, ascospores broad paseudoparaphyses trabeculate, anastomosing
fusiform, 1-septate, hyaline, with or without a filaments, in a gelatinous matrix, asci
sheath that may be drawn out terminally. obclavate, clavate, cylindrical, short

35
pedunculate, thick-walled, J-, ocular chamber Further ecological studies are required to
without or with an apical apparatus, determine if species marked ? are truly marine.
ascospores ellipsoidal, 1-3-septate, slightly
constricted at the septa, with a mucilaginous 1. Ascospores 1-septate................................................ 2
1. Ascospores 3-septate................................................ 5
sheath sometime drawn out to form apical
appendages e.g. M. ramunculicola (Kohlmeyer 2. Ascospores with polar appendages or cupulate
and Kohlmeyer, 1979; Read et al., 1992b). mucilaginous pad ..................................................... 3
Species assigned to Massarina have 2. Ascospores with a sheath......................................... 4
undergone extensive revision as the result of
3. Ascospores 28-37.5 × 4.7-6.5 µm, with cupulate,
morphological and molecular evaluation (Liew mucilaginous appendages at both ends, apical
et al., 2002). Of 160 Massarina names in the smaller than the basal appendage, on Phragmites ....
literature, Aptroot (1998) only retained 43 taxa, .......................................................M. phragmiticola
while others have been transferred to 3. Ascospores 50-65 (-73) × 15-23 (-25) µm, polar
Lophiostoma as the result of molecular drawn out polar appendages, equal in length, on
algae.................................................. M. cystophorae
evidence (Hyde and Aptroot, 1998; Hyde et al.,
2002; Liew et al., 2002). Aptroot (1989) 4. Ascospores 28-44 (-47) × 10-15 µm, sheath entire ...
considers that M. ricifera may be better placed ............................................................ M. lacertensis
in Wettsteinina, and that M. lacertensis does 4. Ascospores 35-42.5 × 12.5-18 µm, polar cap extends
not belong in Massarina because ascomata are into the enveloping sheath ............M. ramunculicola
4. Ascospores 22-28 (-33) × 4.5-6.5 µm, sheath
immersed in an extensive thick, black stroma. confined to ascospore tip, extending 2-6 µm, ending
We retain these species here until further bluntly................................................M. rhizophorae
molecular studies are undertaken.
Ascospores in marine Massarina species 5. Ascospores 45-56 × 14-19 µm........... M. velataspora
generally have a mucilaginous sheath, often 5. Ascospores length shorter than 45 µm..................... 6
elaborated into appendages (Read et al., 1994, 6. Ascospores 28-44 (-47) × 10-15 µm, on wood ........ 7
1997a, b; Au and Vrijmoed, 2002; Hyde et al., 6. Ascospores 19-25 × 5.5-7 µm, on Juncus..................
2002). In M. acrostichi, M. lacertensis, M. ..................................................................M. ricifera
ramunculicola, M. thalassiae and M.
velatospora there are well developed sheaths, 7. Ascospores 28-44 (-47) × 10-15 µm..........................
................................................................ M. thalassiae
which is multilayered in M. ricifera. Polar 7. Ascospores narrower than 10 µm ............................ 8
caps to the ascospores are found in M.
phragmiticola, while in M. cystophorae polar 8. Ascospores 26-34 × 8-9.5 µm, on Bruguiera
appendages are present. In M. ramunculicola, gymnorhiza wood............................... M. mauritiana
the exosporial mucilaginous sheath contains a 8. Ascospores 18-21 × 6-8 µm, on Hibiscus tiliaceus ...
..........................................................M. beaurivagea
fibrillar component. Prior to the release of
ascospores, the fibrillar component penetrates Paraliomyces Kohlm., Nova Hedw. 1: 81,
the delimiting membrane at the spore poles to 1959.............................................................. (1)
form a polar cap (Read et al., 1997b), a unique ◙ P. lentifer Kohlm., Nova Hedw. 1: 81, 1959
feature in the genus. Ascospores of Massarina (Type species).
thalassiae possess a well-developed exosporial Stromata black immersed in the
mucilaginous sheath and at the spore poles, a substratum, ascomata subglobose to pyriform,
polar chamber is formed within the episporium, immersed, ostiolate, papillate or epapillate,
which projects into the sheath. However, this carbonaceous, black, solitary, periphysate,
does not penetrate through the sheath or the pseudoparaphyses filiform, numerous, asci
delimiting membrane (Read et al., 1994). cylindrical, short pedunculate, thick-walled,
Massarina ricifera also has two types of bitunicate, without an apical apparatus,
sheaths, one cap-like around one end of the ascospores ellipsoidal to subfusiform, 1-
ascospore, and a more extensive diffuse sheath septate, constricted at the septum, hyaline
around the entire spore. It differs from other becoming brown, with a mucilaginous sheath
marine Massarina species and requires further and a lenticular appendage at the central
investigation at the TEM and molecular level. septum.

36
 Fungal Diversity

A monotypic genus, occurring on with Lophiostoma, as evidenced by the weak

submerged wood and geographically well bootstrap support. Barr (1990a) referred
distributed in the tropics (Tam et al., 2003). Platystomum to the Platystomataceae,
The genus can be assigned to the Pleo- Melanommatales, however, Platystomum
sporales, Lophiostomataceae with confidence compressum has been treated as a
as it forms a clade with Lophiostoma Lophiostoma species by Holm and Holm
crenatum and L. caulium (Tam et al., 2003). (1988) and this was supported by Eriksson and
Liew et al. (2002) have evaluated the Hawksworth (1991). However, ascospore
monophyly of the genus Massarina and morphology is significantly different from
transferred a number of taxa to Lophiostoma. Lophiostoma and we retain it as a separate
Included in their analysis was L. caulium, also genus.
used by Tam et al. (2003), and this falls into a
well characterized Lophiostoma group with Quintaria Kohlm. & Volkm.-Kohlm., Bot.
narrow, fusiform ascospores. Although P. Mar. 34: 34, 1991......................................... (1)
lentiferus has many features in common with ◙ Q. lignatilis (Kohlm.) Kohlm. & Volkm.-
other marine Lophiostoma and Massarina Kohlm., Bot. Mar. 34: 35, 1991 (Type species).
species they are not congeneric. Ascomata obpyriform, immersed,
Ultrastructurally, the ascospores of P. ostiolate, papillate, carbonaceous, black,
lentifer differ significantly from those of other solitary or gregarious, pseudoparaphyses
marine Lophiostoma and Massarina species septate, branching and anastomosing, asci
(Read et al., 1992). The mucilaginous sheath cylindrical, pedunculate, bitunicate, with an
in Paraliomyces is thick (340-380 nm) with apical plate, ascospores fusiform, 5-septate,
numerous electron-opaque granules. The constricted at the septa, hyaline, no sheath or
lateral lentiform appendage is located at the appendage (Fig. 16). The genus is
central septum and comprises longitudinally differentiated from Trematosphaeria by
oriented fibrils in an amorphous matrix and having completely immersed ascomata with
attached to the episporium by electron-dense rounded bases, black incrustations lining the
strands that pass through the sheath, features sides of the ostiolar canal, a non-amyloid plate
not observed in Massarina and Lophiostoma in the ascus and hyaline ascospores.
species.
5. Melanommataceae
Platystomum Trevis., Bull. Soc. R. Bot. Belg.
16: 16, 1877 ................................................. (1) Acrocordiopsis Borse & K.D. Hyde,
◙ Platystomum scabridisporum Abdel- Mycotaxon 34: 536, 1989 ............................ (2)
Wahab & E.B.G. Jones, Mycoscience 41: 384, ◙ A. patilii Borse & K.D. Hyde, Mycotaxon
2000. 34: 536, 1989 (Type species).
A newly described marine species from A. sphaerica Alias & E.B.G. Jones, Fungal
Australian driftwood associated with sand. Diver. 2: 39, 1999.
Ascomata subglobose, immersed, erumpent, Both Acrocordiopsis species were
papillate, ostiolate, periphysate, black and reported on mangrove wood with large, black,
coriaceous, peridium two-layered, trabeculate carbonaceous, conical ascomata seated on a
pseudoparaphyses, asci cylindrical with an black stromata, epapillate, peridium thick
ocular chamber, ascospores fusiform, composed of 2-3 layers, pseudoparaphyses
muriform, 5-8-transverse septate and 1-3- abundant, asci cylindrical, bitunicate with an
longitudinal septate, constricted at the septa, apical thickening and an ocular chamber, with
brown with rough or verrucose spore wall hyaline to yellowish, 1-septate ascospores,
surface (Figs. 11a, 15). Sequence data place it lacking a sheath or appendages (Fig. 18)
in the Lophiostomataceae as a sister group to (Borse and Hyde, 1989; Alias et al., 1999).
Lophiostoma species (Fig. 17) (Suetrong et al., Preliminary data confirms its assignment to the
2009 in press). However it is not congeneric Pleosporales.

37
 a b

c d e f

g h

i j

k l

Fig. 15. Platystomum scabridisporum. a-b. Cylindrical immature asci and pseudoparaphyses. c. Cylindrical asci. d.
Ascus tip with an apical pore. e-l. Muriform ascospores. Bars a-l = 10 µm.

38
 Fungal Diversity

a b

c d

Fig. 16. Quintaria lignatilis. a. Black ascomata immersed in mangrove wood. b. Thin-walled pseudoparaphyses. c.
Bitunicate ascus with short pedicel. d. Ascus cylindrical with uniseriate ascospores. e-g. Hyaline 5-septate ascospores.
Bars a = 100 µm; b-g = 10 µm.

39
Fig. 17. Phylogram generated from step matrix parsimony analysis from combined SSU, LSU rDNA, RPB2 and EF-1-
alpha sequences. Parsimony bootstrap value greater than 50% and Bayesian Posterior Probabilities greater than 0.95 are
given above and below each clade, respectively.

40
 Fungal Diversity

1. Ascomatal wall 2-layered, ascospores elongate, less Astrosphaeriella species are most similar
than 16 µm in width ..................................... A. patilii to Trematosphaeria and Caryospora species in
1. Ascomatal wall 3-layered, ascospores spherical-
ellipsoidal, wider than 15 µm................ A. sphaerica ascomatal features. Astrosphaeriella species
generally occur on monocotyledons while
Trematosphaeria occur on wide range of
a b plants, ascospore length-width ratios are
higher in Astrosphaeriella species than
Trematosphaeria. Astrosphaeriella differs
from Lophiostoma species in the narrower
trabeculate pseudoparaphyeses and with
carbonaceous ascomata (Hyde and Fröhlich,
1999). Astrosphaeriella asiana and A.
mangrovei were transferred from Lophiostoma
on the basis of ascomatal structural
morphology, narrower pseudoparaphyses and
molecular sequencing data (Hyde et al., 2002).

1. Ascospores with a sheath......................................... 2

1. Ascospores lacking a sheath, hyaline, 31-38 × 6-9
Fig. 18. Acrocordiopsis patilii. a. Cylindrical asci and µm, brown with pallid end cells, on Nypa .................
thin-walled pseudoparaphyses. b. Asci with hyaline, 1- ........................................................... A. striataspora
septate ascospores. Bars a-b = 10 µm.
2. On Nypa, ascospores 18.5-27 × 4-6 µm, hyaline to
pale yellow...................................................A. nypae
Astrosphaeriella Syd. & P. Syd., Annls. 2. On mangrove wood.................................................. 3
Mycol. 11: 260, 1913 ................................... (4)
◙ A. asiana (K.D. Hyde) Aptroot & K.D. 3. Ascospores (36-) 37.5-55 × 7-11 µm, fusiform, dark
brown ................................................... A. mangrovei
Hyde, Nova Hedw. 70: 145, 2000. 3. Ascospores 28-40 × 5-8 µm, broadly fusiform,
Lophiostoma asiana K.D. Hyde, Mycotaxon 55: brown .........................................................A. asiana
285, 1995.
◙ A. mangrovei (Kohlm. & Vital) Aptroot & Bicrouania Kohlm. & Volkm.-Kohlm. Mycol.
K.D. Hyde, Fungi in Marine Environments. Res. 94: 685, 1990 ....................................... (1)
Fungal Diversity Press: 106, 2002. B. maritima (H. Crouan & P. Crouan) Kohlm.
Lophiostoma mangrovei Kohlm. & Vital,
Mycologia 78: 487, 1986.
& Volkm.-Kohlm., Mycol. Res. 94: 685, 1990
A. nypae K.D. Hyde, Bot. J. Linn. Soc. 110: (Type species).
Didymosphaeria maritima (H. Crouan & P.
96, 1992. Crouan) Sacc., Syll. Fung. 1: 703, 1882.
A. striataspora (K.D. Hyde) K.D. Hyde, Bot. J. Sphaeria maritima P. Crouan & H. Crouan,
Linn. Soc. 110: 97, 1992. Florule du Finistére, Paris: 27, 1867 (non Sphaeria
Trematosphaeria striatispora K.D. Hyde, Bot. J. maritima Cooke & Plowr., 1877, nom. Illegit.)
Linn. Soc. 98: 142, 1988. Ascomata superficial, lacking a clypeus,
Ascomata immersed and subepidermal, subglobose to ellipsoidal, ostiolate, peri-
becoming superficial, conical or hemispheri- physate, short papillate, subcarbonaceous,
cal, brown to black, solitary or gregarious, blackish brown, pseudoparaphyses present,
ostiole central, papillate, pseudoparaphyses asci cylindrical, short pedunculate, without an
trabeculate in a gelatinous matrix, asci ocular chamber or apical apparatus, ascospores
cylindrical, cylindrical-clavate, pedunculate, ellipsoidal, 1-septate, constricted at the septum,
thick-walled, bitunicate, with an ocular reddish-brown, thick-walled and smooth
chamber and faint ring, ascospores elongate- (Kohlmeyer and Volkmann-Kohlmeyer,
fusiform, often tapering at their apices, 1 to 1990a). It was referred to the Melanom-
multiseptate, slightly constricted at the septa, mataceae, Melanommatales by Kohlmeyer and
hyaline, brown to reddish-brown, often with Volkmann-Kohlmeyer (1990a) but requires
mucilaginous sheaths or appendages. further study at the molecular level to confirm
41
its ordinal status. Originally described as a characterized species from senescent, decaying
Sphaeria species growing on the salt marsh mangrove wood (McKeown et al., 2001).
plant Halimione portulacoides, which was Although Kohlmeyer (1968a) refered to 25
later transferred to Didymosphae-ria, but it Trematosphaeria species, Bois (1985)
clearly does not belong in that genus because accepted only a few.
of its superficial ascomata. The genus has been assigned to various
families: Pleosporaceae (Kohlmeyer, 1968),
Caryosporella Kohlm., Proc. Indian Acad. Sci. and Melanommataceae (Barr, 1990a). We
(Plant Sci.) 94: 355, 1985 ............................ (1) consider the latter the most appropriate at this
C. rhizophorae Kohlm., Proc. Indian Acad. time. The taxonomic position of Tremato-
Sci. (Plant Sci.) 94: 356, 1985 (Type species). sphaeria is not fully resolved as the genus is
A monotypic genus described from dead polyphyletic (Schoch et al., 2006), with T.
wood of intertidal roots and branches of heterospora grouping with two other
mangrove trees (Rhizophora mangle), and Lophiostoma species (Lophiostomataceae
probably related to Caryospora (Kohlmeyer, clade 1), while T. pertusa is in the
1985). Characterized by large ascomata (750- Lophiostomataceae clade 2, along with
900 µm high, 830-1050 µm wide), superficial Herpotrichia and Pleomassaria species
on a thin black stroma, ostiolate, periphysate, (Schoch et al., 2006). However, verified
carbonaceous, thick peridium (90-160 µm), strains of T. pertusa form a robust cluster with
trabeculate pseudoparaphyses, asci bitunicate, Bimuria novae-zelandiae, Phaeodothis winteri,
thick-walled, J-, with an apical apparatus, Montagnula opulenta and Massarina eburnea,
ascospores 1-septate, dark brown, verrucose, a sister group with the Pleosporaceae,
and thickened at their apices. Kohlmeyer Phaeosphaeriaceae and Delitschiaceae with
(1985) referred it to the Massariaceae, moderate bootstrap support (Zhang et al.,
Melanommatales, but if it is considered related 2008).
to Caryospora, then an alternative taxonomic
group is required. Resolution of its taxonomic 1. Ascospores with wall striation and sheath ............... 2
position warrants study at the molecular level. 1. Ascospores lacking wall striation and sheath, 30-
(35.5)-41 × 12-13 (-16.5) µm, 3-septate, brown, light
brown at extreme poles ........................ T. mangrovei
Trematosphaeria Fuckel, Jahrb. Nassau. Ver.
Naturkd. 23-24: 161, 1869-1870.................. (3) 2. Ascospores 20-25 × 4-6 µm, fusiform or ellipsoid, 3-
T. lineolatispora K.D. Hyde, Mycol. Res. 96: septate, light brown, striated, with a sheath ...............
28, 1992. ............................................................ T. malaysiana
T. malaysiana Alias, McKeown, S.T. Moss & 2. Ascospores 34-48 × 7-10 µm, 5-septate, cinnamon
brown, striate sheath ....................... T. lineolatispora
E.B.G. Jones, Mycol. Res. 105: 616, 2001.
T. mangrovei Kohlm., Mycopath. Mycol.
Appl. 34: 1, 1968.
Ascomata subglobose, obpyriform,
immersed, ostiolate, papillate, coriaceous to
carbonaceous, black, solitary or gregarious,
necks with hyaline thin hyphae, pseudopara-
physes unbranched, trabeculate, in gelatinous
matrix, asci cylindrical, pedunculate, thick-
walled, fissitunicate, with an eccentric apical
plate, J-, ascospores fusiform multi-septate,
slightly constricted to constricted at the septa,
hyaline, smooth-walled, lacking a sheath or
Fig. 19. Trematosphaeria malaysiana. Three-septate
appendages (Fig. 19) (Kohlmeyer, 1984; ascospore with striations along the entire spore. Bar = 5
McKeown et al., 2001). Three well μm.

42
 Fungal Diversity

6. Monoblastiaceae ascospores 7-8-septate, central cells larger, and

with a sheath (Fig. 20). The two species have
Ascocratera Kohlm., Can. J. Bot. 64: 3036, yellow to pale-brown ascospores with mea-
1986.............................................................. (1) surements that overlap, but differ in the degree
A. manglicola Kohlm., Can. J. Bot. 64: 3036, of septation and the morphology of the sheath
1986 (Type species). (Hyde, 1992a, 1994b). Carinispora resembles
Kohlmeyer (1986a) regards this as a Phaeosphaeria but differs in that it is a
common bitunicate ascomycete in the upper saprobe on the palm Nypa, in wall structure,
intertidal zone. Ascomata on a thin stroma on the ocular chamber in the ascus and in the
the wood surface, crater-like, large (1100-1400 morphology of the ascospore gelatinous sheath.
µm diam.), black, carbonaceous, periphysate, Infrequently collected despite intensive
trabeculate pseudoparaphyses in a gelatinous sampling of the host plant (Pilantanapak et al.,
matrix, asci bitunicate, thick-walled with an 2005; Jones et al., 2005a).
apical apparatus, ascospores hyaline, 3-septate
and constricted at the central septum and 1. Ascospores 42-66 × 7-10.5 µm, 8-9 celled, with a
keel-shaped sheath .......................................C. nypae
surrounded by a gelatinous sheath. 1. Ascospores 43-54 × 8-9 µm, 7-8 celled, with a
Ascospores of A. manglicola can be narrow sheath.......................................C. velatispora
confused with those of Massarina velatospora.
In our molecular study, A. manglicola forms a Lautitia S. Schatz, Can. J. Bot. 62: 31, 1984....
sister group to Aigialus species in a clade that ...................................................................... (1)
also includes Quintaria lignatilis and L. danica (Berl.) S. Schatz, Can. J. Bot. 62:31,
Decaisnella formosa. However it shares few 1984 (Type species).
morphological characters (black, ostiolate Leptosphaeria danica Berl., Icones Fung. 1: 87,
ascomata, trabeculate pseudoparaphyses, 1892.
Leptosphaeria chondri Rosenv., Bot Tidsskr. 27:
pedunculate, 8-spores, uniseriate, cylindrical 35, 1906.
asci, septate, ellipsoidal to muriform Leptosphaeria marina Rostr., Bot. Tidsske. 17:
ascospores) with these taxa (Suetrong et al., 234, 1889.
2009, in press). Didymosphaeria danica (Berl.) I.M. Wilson &
Knoyle, Trans. Br. Mycol. Soc. 44: 55, 1961.
Sphaerella chondri H.L. Jones, Oberlin Coll Kab.
7. Phaeosphaeriaceae 9: 3, 1898.
Guignardia chondri (H.L. Jones) Estee, Publ.
The phylogenetic position of the families Ataz. Zool. Napoli 15: 378, 1936.
Phaeosphaeriaceae and Leptosphaeriaceae has Ascomata ampulliform to subglobose,
been discussed above, and will not be repeated immersed in cystocarps and tetracarps of
here. Chondrus crispus, ostiolate, papillate,
coriaceous, pale-coloured, clypeus black,
Carinispora K.D. Hyde, Bot. J. Linn. Soc. 110: gregarious, pseudoparaphyses thin, septate,
97, 1992........................................................ (2) simple or branched, asci subclavate to
C. nypae K.D. Hyde, Bot. J. Linn. Soc. 110: subcylindrical, short pedunculate, thick-walled,
99, 1992 (Type species). bitunicate, ascospores elongate fusiform, 1-
C. velatispora K.D. Hyde, Sydowia 46: 259, septate, slightly constricted at the septum,
1994. hyaline, no sheath or appendages. Lautitia
A genus known only from the palm danica grows on the cystocarps of the red alga
Nypa fruticans, ascomata are large circa 0.8 Chondrus crispus throughout the year, but is
mm, crust-like, with a central ostiole, occur more prevalent during November in British
beneath the epidermis with an overlaying coastal waters (Stanley, 1992). Ascomata are
clypeus. They lack periphyses and the globose and produced in the cortex of the alga
peridium is light brown composed of thin- beneath the epidermis. Ascospores are 33-40 ×
walled cells, pseudoparaphyses filiform 5-8 µm. The relationship of the species within
numerous and in a gel. Asci are clavate to the family and order warrants further
cylindrical, with an ocular chamber and investigation.
43
Phaeosphaeria I. Miyake, Bot. Mag. 23: 93, Ascomata subglobose, pyriform to
1909............................................................ (11) ellipsoidal, immersed, ostiolate, papillate,
Ph. capensis T.D. Steinke & K.D. Hyde, coriaceous, dark brown, solitary or gregarious,
Mycoscience 38: 101, 1997. pseudoparaphyses septate, filamentous, in a
Ph. gessneri Shoemaker & C.E. Babc., Can. J. gelatinous matrix, branched, anastomosing,
Bot. 67: 1567, 1989. asci clavate to cylindrical, short pedunculate,
◙ Ph. halima (T.W. Johnson) Shoemaker & bitunicate, fissitunicate, ocular chamber
C.E. Babc., Can. J. Bot. 67: 1514, 1989. present, J-, lacking with or without an apical
Leptosphaeria halima T.W. Johnson, Mycologia apparatus, ascospores fusiform to ellipsoidal,
48: 502, 1956. multi- septate, slightly constricted at the septa,
Ph. macrosporidium (E.B.G. Jones) yellowish to pale brown, with or without a
Shoemaker & C.E. Babc., Can. J. Bot. 67: gelatinous sheath, lacking appendages.
1532, 1989. Phaeosphaeria species are generally
Leptosphaeria macrosporidium E.B.G. Jones,
Trans. Br. Mycol. Soc. 45: 103, 1962.
parasites of grasses, sedges, rushes and other
Ph. neomaritima (R.V. Gessner & Kohlm.) monocotyledons, although many of the marine
Shoemaker & C.E. Babc., Can. J. Bot. 67: species are saprophytes, with anamorphs
1572, 1989. (where known) in Stagonospora. A number of
Leptosphaeria neomaritima R.V. Gessner & new species have recently been added to this
Kohlm., Can. J. Bot. 54: 2032, 1976. (nom. nov. superfl.) genus, and others transferred from
Sphaeria maritima Cooke & Plowr., Grevillea 5: Leptosphaeria: Ph. albopunctata, Ph. halima,
120, 1877. (nom. illegit.) Ph. macrosporidium, Ph. neomaritima, Ph.
Leptosphaeria maritima Sacc., Syll. Fung.
(Abellini) 2: 72, 1883.
orae-maris and Ph. spartinae (Shoemaker and
Ph. olivacea Kohlm., Volkm.-Kohlm. & O.E. Babcock, 1989; Khashnobish and Shearer,
Erikss., Bot. Mar. 40: 299, 1997. 1996a, b). Most of these transfers are made on
Ph. orae-maris (Linder) Khashn. & Shearer, the basis of the colour of the ascospores. The
Mycol. Res. 100: 1351, 1996. characters delineating Leptosphaeria and
Leptosphaeria oraemaris Linder, Farlowia 1: Phaeosphaeria are discussed under the former
413, 1944. genus. Phaeosphaeria has ascomata which
Ph. roemeriani Kohlm., Volkm.-Kohlm. & range from small to large and are generally
O.E. Erikss., Can. J. Bot. 76: 470, 1998. superficial on the substratum, with generally
◙ Ph. spartinae (Ellis & Everh.) Shoemaker & monocotyledonous hosts. The position of Ph.
C.E. Babc., Can. J. Bot. 67: 1573, 1989. typharum is questionable with Leuchtmann
Leptosphaeria spartinae Ellis & Everh., J. Mycol. (1984) rejecting its inclusion in Phaeo-
1: 43, 1885. sphaeria. Shoemaker and Babcock (1989)
Leptosphaeria sticta Ellis & Everh., J. Mycol. 1:
43, 1885.
retain it in the genus for the present.
Phaeosphaeria roemeriani is a rare species
◙ Ph. spartinicola Leuchtm. Mycotaxon 41: 2, growing on Juncus roemerianus, between 6-52
1991. cm above the rhizome, and is almost
◙ Ph. typharum (Desm.) L. Holm., Symb. Bot. permanently inundated by seawater
Ups. 14: 126, 1957. (Kohlmeyer and Kohlmeyer, 1998e) (Fig. 21).
Sphaeria scirpicola var. typharum Desm.,
Platentes Crypt. France ed. 2: 1778, 1849. 1. Ascospores 3-septate................................................ 2
Sphaeria typharum (Desm.) Raben., Herb. Myc. 1. Ascospores more than 3-septate............................... 8
ed 2: 731, 1858.
Pleospora typharum (Desm.) Fuckle, Symb. 2. On Juncus ................................................................ 3
Mycol. 137, 1870. 2. On other substrata .................................................... 4
Leptosphaeria typharum (Desm.) Karst., Bidr.
Känn. Finl. Nat. Folk 23: 100, 1873.
Sphaeria perpusilla var. typharum Auers., 3. Ascospores 16-21 × 4-5 µm, fusiform ......................
Rabenhorst Fungi Europaeae: 831, 1865. ............................................................... Ph. olivacea
3. Ascospores 23-35 × 9-13 µm, fusiform .....................
Anamorph: Scolecosporiella typhae
......................................................... Ph. Roemeriani
(Oudem.) Petrak, Ann. Mycol. 19: 31, 1921.

44
 Fungal Diversity

a b

Fig. 20. Carinispora nypa. a. 7-septate, yellow to pale-brown ascospore with a keel-shaped sheath (arrows). b. Clavate
ascus with an ocular chamber. Bars a = 10 µm; b = 20 µm.

4. On Spartina.............................................................. 5 8. Pleosporaceae
4. On other substrata .................................................... 6
Decorospora Inderb., Kohlm. & Volkm.-
5. Ascospores 12-18 × 5-8 µm, yellow-brown, also on Kohlm., Mycologia 94: 657, 2002 ............... (1)
driftwood.................................................. Ph. halima ◙ D. gaudefroyi (Pat.) Inderb., Kohlm. &
5. Ascospores 23-35 × 9-13 µm, yellow-brown to
brown, only on Spartina................... Ph. spartinicola
Volkm.-Kohlm., Mycologia 94: 657, 2002
(Type species).
Pleospora gaudefroyi Pat., Tabulae Analticae
6. Ascospores less than 8 µm wide, golden-brown,
Fungorum, Paris 2: 40, 1886.
echinate 17-24 × 6-8 µm ................... Ph. orae-maris
Pleospora salsolae Fuckel var. schoberiae Sacc.,
6. Ascospore wider than 8 µm ..................................... 7 Michelia 2: 69, 1880.
Pleospora schoberiae (Sacc.) Berl., Icon. Fung.
7. Ascospores (21-) 24-39 (-35) × 8-12 µm, reddish- 2: 23, 1895.
brown, on Typha ..................................Ph. typharum Pleospora lignicola J. Webser & M.T. Lucas,
7. Ascospores 45-68 (-72) × 10-14 µm, hyaline to pale Trans Br. Mycol. Soc. 44: 431, 1961.
yellow, on wood, Juncus and Spartina ..................... Pleospora salicorniae Jaap, Verh. Bot. Ver.
................................................. Ph. macrosporidium Prov., Brandenburg 49: 16, 1907.
Pleospora herbarum (Fr.) Rabenh. var.
8. On wood, ascospores 29-36 × 7-8 µm, 6-7-septate, salicorniae (Jaap) Jaap, Ann. Mycol. 14: 17, 1916.
clavate ................................................... Ph. capensis Ascomata subglobose to ellipsoidal,
8. On salt marsh grasses............................................... 9 immersed, ostiolate, epapillate or with a short
papilla, carbonaceous, black, solitary to
9. Ascospores with no sheath, 35-40 (-52) × 9-11 (-14), gregarious, pseudoparaphyses septate, ramose,
5-septate, yellow-brown, on Spartina ........................ asci clavate, short pedunculate, thick-walled,
............................................................. Ph. spartinae
bitunicate, without an apical apparatus,
9. Ascospores with a sheath ....................................... 10
ascospores ellipsoidal, muriform, brown, with
10. Ascospores 68-85 × (10-) 15-20 µm, 6-7-septate, a mucilaginous sheath slightly constricted at
yellowish-brown, on Spartina ............. Ph. gessneri the center and drawn out at each apex into 2-3
10. Ascospores (30-) 32-45 × (6-) 8-14 µm, 3-5- subconical extensions (Inderbitzin et al., 2002).
septate, yellowish-brown, on Juncus....................... This genus was introduced to accommodate
.....................................................Ph. Neomaritima Pleospora gaudefroyi as it forms a sister taxon

45
 assigned to the Pleosporales, Pleosporaceae
with confidence, as it forms a well supported
clade with Kirschsteiniothelia elaterascus
(Shearer, 1993a; Tam et al., 2003). Common
characters for taxa in this clade include:
persistent, anastomosing hyphal-like
pseudoparaphyses, a coiled endoascus, which
uncoils when the spores are released. The
genus is monophyletic, with H. nypae found
on the palm Nypa fruticans that differs from
the type species in having smaller ascospores,
a verrucose wall and a persistent sheath
(Hyde, 1991b).

1. Ascospores 25-35 × 12-15 µm, ascospore wall

verrucose with a persistent sheath, on the palm Nypa
Fig. 21. Phaeosphaeria roemeriani. Three-septate fruticans ..................................................... H. nypae
ascospore. Bar = 5 μm. 1. Ascospores 30-55 × 17-25 µm, wall smooth,
ascospores lacking a sheath, on mangrove wood.......
........................................................... H. kanaloanus
to Alternaria alternata, Cochliobolus sativus,
Pleospora herbarum, Pyrenophora tritici- Falciformispora K.D. Hyde, Mycol. Res. 96:
repentis and Setosphaeria rostrata within the 26, 1992........................................................ (1)
Pleosporaceae (Inderbitzin et al., 2002). The ◙ F. lignatilis K.D. Hyde, Mycol. Res. 96: 27,
genus differs from Pleospora at the molecular 1992 (Type species).
and morphological level, especially the well A little known species described from
developed gelatinous sheath drawn into 2-4 mangrove wood from Mexico (Hyde, 1992b),
subconical extensions (Yusoff et al., 1994b). characterised by ascomata that are black, soft-
walled, superficial with a rounded ostiole,
Helicascus Kohlm., Can. J. Bot. 47: 1471, bitunicate asci with an ocular chamber, wide
1969.............................................................. (2) and cellular pseudoparaphyses and fusiform,
◙ H. kanaloanus Kohlm., Can. J. Bot. 47: hyaline, 6-8-septate ascospores, slightly
1471, 1969 (Type species). constricted at the septa, surrounded by a thin
◙ H. nypae K.D. Hyde, Bot. Mar. 34: 314, mucilaginous sheath and single scythe-like
1991. appendage at its base.
Ascomata ampulliform, lenticular,
horizontally arranged under a black Pleospora Rabenh. ex. Ces. & De Not.,
pseudoclypeus, immersed, ostiolate, carbo- Comment. Soc. Crittogam. Ital. 1: 217, 1863 ...
naceous, centrum in 3-5 locules, papillate, ...................................................................... (6)
periphysate, pseudoparaphyses numerous, ◙ P. gracilariae E.G. Simmons & S. Schatz,
persistent, asci subcylindrical to oblong- Mem. N. Y. Bot. Gard. 49: 305, 1989.
clavate, pedunculate, thick-walled, bitunicate, ◙ P. pelagica T.W. Johnson, Mycologia 48:
with an apical apparatus, ascospores 504, 1956.
obovoidal, 1-septate, constricted at the P. pelvetiae G.K. Sutherl., New Phytol. 14: 41,
septum, dark-brown, spore wall thick, 2- 1915.
layered, no appendages, lacking or with a ◙ P. spartinae (J. Webster & M.T. Lucas)
sheath. A well characterized genus, the Apinis & Chesters, Trans. Br. Mycol. Soc. 47:
ascomata with many locules, a coiled 432, 1964.
endoascus, thick-walled brown ascospores Pleospora vagans var. spartinae J. Webster &
with a prominent germ pore growing on M.T. Lucas, Trans. Br. Mycol. Soc. 44: 427, 1961.
mangrove wood. Initially classified in the ◙ P. triglochinicola J. Webster, Trans. Br.
Dothideomycetes incertae sedis, it can now be Mycol. Soc. 53: 481, 1969.

46
 Fungal Diversity

Anamorph: Stemphylium triglochinicola Anamorph: Phoma sp.

B. Sutton & Piroz., Trans. Br. Mycol. Soc. 46: Ascomata immersed, no ostiole,
519, 1963. subcarbonaceous, brown, pseudoparaphyses
Marine Pleospora species occur on wide septate, thickened at their tips, asci clavate,
range of hosts, Tamarix aphylla twigs, short pedunculate, thick-walled, ascospores
seaweeds and salt marsh plants. Ascomata ellipsoidal, yellowish-brown, muriform, with 3
solitary or gregarious, globose, leathery to transverse septa and 2-4 longitudinal septa,
subcarbonaceous, immersed, ostiolate, end cells slightly paler, with a thick gelatinous
papillate, pseudoparaphyses numerous, asci sheath (“perispore”). Forms pycnidia in culture
cylindrical to clavate, pedunculate, thick- (Kohlmeyer et al., 1995b).
walled, ascospores ellipsoid, clavariform or This ascomycete occurs on senescent
fusiform, muriform, with transverse and culms of Juncus roemerianus and is regularly
longitudinal septa, generally constricted at the immersed at high tide in salt marshes, although
septa, yellowish or pale brown with gelatinous regarded as a facultative species by Kohlmeyer
sheaths. et al. (1995b). Distinguished from similar
Although Pleospora species frequently genera (Lewia, Diademosa), by the apical cap
have anamorphs, the only marine species with to the ascus, J- ocular chamber, ascospores
an anamorph is P. triglochinicola with a wide mucilaginous sheath (up to 25 μm),
(Stemphylium triglochinicola). The genus ascomata not papillate, and with a Phoma-like
Pleospora is polyphyletic (Kodsueb et al., anamorph.
2006) and as evidenced by the transfer of the
marine P. gaudefroyi to Decorospora. Further 9. Teichosporaceae
studies are required to determine if marine
species are correctly assigned to this genus as Byssothecium Fuckel, Bot. Ztg. 19: 251, 1861
species such as P. bjoerlingii and P. iqbalii ...................................................................... (1)
show no affinity with other Pleospora species ◙ B. obiones (P. Crouan & H. Crouan) M.E.
(Kodsueb et al., 2006). Barr, Mycotaxon 82: 378, 2002.
Passeriniella obiones (P. Crouan & H. Crouan)
1. On algae ................................................................... 2 K.D. Hyde & Mouzouras, Trans. Br. Mycol. Soc. 91:
1. On other substrate .................................................... 3 183, 1988.
Pleospora obiones P. Crouan & H. Crouan,
2. Ascospores with pronounced gelatinous sheaths 28- Florule Finistère (Paris): 22, 1867.
29.5 × 3-13.5 μm, on Gracilaria.........Pl. gracilariae Leptosphaeria discors Sacc. & Ellis, Michelia 2:
2. Ascospores with 6-7 trans-septa, several longisepta, 567, 1882.
25-35 × 12-17 μm, on the brown alga, Pelvetia......... Leptosphaeria obiones (P. Croun & H. Crouan)
............................................................... Pl. pelvetiae Sacc., Syll.fung. (Abellini) 2 : 24, 1883.
Leptosphaeria obiones f. evolutior Grove, J. Bot.
3. Ascospores with 5 trans-septa, 1 longiseptum, 24-28 (Lond.) 71 : 281, 1933.
× 10-13 μm, on the marsh angiosperm Spartina ........ Didymosphaeria spartinae Grove, J. Bot (Lond.)
.............................................................. Pl. spartinae 71: 259, 1933.
3. Ascospores with 7 trans-septa, 1-3 longisepta, 45-66 Metasphaeria discors (Sacc. & Ellis) Sacc., Syll.
× 16-25 μm, on the salt marsh plant Triglochin ......... Fung. 2: 173, 1883.
..................................................... Pl. triglochinicola Passeriniella discors (Sacc. & Ellis) Apinis &
3. Ascospores with 7-9 trans-septa, 1 longiseptum, 35- Chesters, Trans. Br. Mycol. Soc. 47: 432, 1964.
52 × 10-15 μm, on Spartina ....................Pl. pelagica Passeriniella incarcerata Berl., Icon. Fung 1: 51,
1892.
Ascomata subglobose or ellipsoidal,
Tremateia Kohlm., Volkm.-Kohlm. & O.E. immersed to erumpent, ostiolate,
Erikss., Bot. Mar. 38: 165, 1995 .................. (1) subcarbonaceous, dark brown to black,
T. halophila Kohlm., Volkm.-Kohlm. & O.E. gregarious. Pseudoparaphyses septate, ramose,
Erikss., Bot. Mar. 38: 166, 1995 (Type species). asci clavate to subcylindrical, short

47
pedunculate, thick-walled, without an apical each cell. In Aigialus the ascospores are brown,
apparatus, ascospores versicoloured, end cell muriform and lack germ slits. They also differ
hyaline, central cells brown, 3-septate, and significantly in ascoma morphology, ascus
constricted at the septa. This common shape, pseudoparaphyses morphology and in
ascomycete on decaying Spartina culms has the substrata on which they grow. Aigialus
been assigned variously to the genera: species also form a sister group to Ascocratera
Pleospora, Leptosphaeria, and Passeriniella manglicola (Suetrong, pers. comm.). The
(Hyde and Mouzouras, 1988; Barr, 2002). position of A. striatispora in the genus needs
Khashnobish and Shearer (1996a, b) showed to be re-evaluated. Aigialus rhizophorae is a
that based on molecular analysis Passeriniella nomen rejectum.
obiones did not belong in either Leptosphaeria
or Phaeosphaeria. Barr (2002) assigned the
1. Ascospores with 6 transverse-septate (rarely 8)....... 2
species to Byssothecium based on its 1. Ascospores with more than 8 transverse-septate ..... 3
versicolorous ascospores, two dark brown
central cells and hyaline terminal cells. 2. Ascospores 35-55 × 10-16 µm, wall smooth .............
............................................................. A. mangrovei
PLEOSPORALES incertae sedis 2. Ascospores 26-38 × 16-19.5 µm, wall with striations
............................................................ A. striatispora
Aigialus Kohlm. & S. Schatz, Trans. Br.
3. Ascospores 44-74 × 19-27 µm, (9) 10-11 (-12)
Mycol. Soc. 85: 699 ..................................... (4) tranverse-septa ............................................A. parvus
◙ A. grandis Kohlm. & S. Schatz, Trans. Br. 3. Ascospores 67-101 × 18-29 µm, with more than 13
Mycol. Soc. 85: 699, 1985 (Type species). transverse-septa......................................... A. grandis
◙ A. mangrovei Borse, Trans. Br. Mycol. Soc.
88: 424, 1987.
◙ A. parvus S. Schatz & Kohlm., Trans. Br. Biatriospora K.D. Hyde & Borse, Mycotaxon
Mycol. Soc. 85: 704, 1985. 26: 263, 1986 ............................................... (1)
◙ A. striatispora K.D. Hyde, Mycol. Res. 96: ◙ B. marina K.D. Hyde & Borse, Mycotaxon
1044, 1992. 26: 264, 1986 (Type species).
Ascomata globose, completely immersed Assigned to the Melanommatales by
in a black stroma, ostiolate, apapillate, Hyde and Borse (1986) its taxonomic position
carbonaceous, to coriaceous, black, gregarious, remains unresolved, characterized by its large
pseudoparaphyses trabeculate, unbranched and (over 850 µm long) submerged, elongate
anastomosing above the asci, asci cylindrical, ascomata in mangrove wood, bitunicate,
pedunculate, thick-walled, apical apparatus, branched pseudoparaphyses, cylindrical asci
ascospores ellipsoidal to broadly fusiform, with an apical apparatus and a long pedicel
muriform, slightly constricted at the septa, and brown to dark-brown, 2-4 septate at each
dark brown, with hyaline to light brown apical end of the ascospores that are fusiform with a
cells, subapical cells covered by a gelatinous globose end cell. Hyde and Borse (1986) refer
sheath. to these as end chambers or appendages, but
This genus was initially assigned to the no mucilage is released from them as in
Melanommatales (Kohlmeyer and Schatz, Lulworthia species. Molecular data confirm its
1985), but referred to the Massariaceae, position in the Pleosporales, but it can not be
Pyrenulales by Hawksworth et al. (1995). The assigned to any family in the order (Fig. 17)
genus is best accommodated in the (Suetrong et al., unpublished data).
Pleosporales, but further studies are required
Didymella Sacc., Michelia 2: 57, 1880................ (4)
with a greater number of taxa, to assign them
D. avicenniae S.D. Patil & Borse, Trans.
to a family with confidence (Tam et al., 2003).
Mycol. Soc. Jpn. 26: 271, 1985.
Sequence data confirms their placement within
◙ D. fucicola (G.K. Sutherl.) Kohlm.,
the Pleosporales and preliminary data suggests
Phytopath. Z. 63: 342, 1968.
an affinity with the Sporormiaceae (Suetrong, Didymosphaeria fucicola G.K. Sutherl., New
pers. comm.), however, the latter have brown, Phytol. 14: 189, 1915.
phragmosporous ascospores with germ slits to

48
 Fungal Diversity

Didymosphaeria pelvetiana G.K. Sutherl., New appendages (Fig. 22) (Kohlmeyer and
Phytol. 14: 186, 1915. Kohlmeyer, 1964-1969). Known only from
D. gloiopeltidis (Miyabe & Tokida) Kohlm. & material on the sea grass Posidonia oceanica
E. Kohlm., Marine Mycology. The Higher and sometimes confused with Pontoporeia
Fungi 382, 1979. biturbinata, but differs in that it has triangular
Guignardia gloiopeltidis Miyabe & Tokida, Bot.
Mag. 61: 118, 1948.
ascomata with the asci produced in a layer at
D. magnei Feldmann, Rev. Gén. Bot. 65: 414, the base of the ascomata, pseudoparaphyses
1958. filiform, branching, asci cylindrical, bitunicate,
Ascomata solitary, globose, immersed or and ascospores 1-septate dark brown and only
erumpent, ostiolate, papillate, dark in colour, slightly constricted at the septum. Along with
pseudoparaphyses filiform, asci cylindrical, Pontoporeia biturbinata the species was
short pedunculate, thickened apex, ascospores initially assigned to the Pleosporaceae and
ovoid to ellipsoidal, 1-septate, constricted at thought to be related to Caryospora,
the septum, hyaline with no sheaths or Herpotrichia and Otthia (Kohlmeyer, 1963).
appendages. Most species occur on the larger However, its relationship to other taxa in the
marine algae, with D. avicenniae the exception, family, is in need of investigation.
growing on mangrove wood. Revision of the
Heleiosa Kohlm., Volkm.-Kohlm. & O.E.
marine species is required.
Erikss., Can. J. Bot., 74: 1830, 1996........... (1)
H. barbatula Kohlm., Volkm.-Kohlm. & O.E.
1. On mangrove wood (Avicennia species), ascospores
Erikss., Can. J. Bot., 74: 1830-1832, 1996
25-32 × 10-15 µm, with a sheath ........D. avicenniae (Type species).
1. On algae ................................................................... 2 Ascomata globose, immersed, ostiolate,
epapillate, clypeate (2-3 under a small
2. Ascospores narrower than 4 µm, 8-16 × 2-4 µm, clypeus), coriaceous, hyaline at the base, light
parasite of Rhodymenia............................. D. magnei
2. Ascospores wider than 4 µm.................................... 3
brown at the sides, pseudoparaphyses
branched and anastomosing, septate, in a
3. Ascospores 16-23 × 6-8 µm, saprobic on brown gelatinous matrix, asci cylindrical, short
seaweeds, with a sheath ........................... D. fucicola pedunculate, thick-walled, refractive apical
3. Ascospores 14-22 × 4-6 µm, parasitic on the red alga apparatus over the ocular chamber, ascospores
Gloiopeltis, no sheath........................ D. gloiopeltidis
ellipsoidal to fusiform, 1-septate, slightly
constricted at the septum, pale brown, with 10
or more cilia-like polar appendages. The
Halotthia Kohlm., Nova Hedw. 6: 9, 1963 . (1)
species can be considered marginally marine
H. posidoniae (Durieu & Mont.) Kohlm.,
as it occurs 42-67 cm above the rhizomes of
Nova Hedw. 6: 9, 1963 (Type species).
Juncus roemerianus. Kohlmeyer et al. (1996)
Sphaeria posidoniae Durieu & Mont., Expl. Sci.
Algérie, Bot. 1: 502. 1849.
were unable to assign it to any family in the
Amphisphaeria posidoniae (Durieu & Mont.) Dothideomycetes although they referred to
Ces. & De Not., Comment. Soc. Crittog. Ital 1: 224, similarity to Appendispora frondicola (Hyde,
1863. 1994a). The latter species was placed in the
Stromata thick, black, subepidermal in Dothideales incertae sedis by Kirk et al.
host cortex. Ascomata broadly conical to (2001).
semiglobose, enclosed in a stroma, immersed
becoming erumpent, ostiolate, epapillate, Julella Fabre, Annls. Sci. Nat. Bot, sér. 6, 9:
carbonaceous, pseudoparaphyses septate, 113, 1879...................................................... (2)
ramose, persistent, asci cylindrical, attenuate at ◙ J. avicenniae (Borse) K.D. Hyde, Mycol.
the base, short pedunculate, thick-walled, Res. 96: 939, 1992.
bitunicate, persistent, ascospores ellipsoidal, Pleospora avicenniae Borse, Curr. Sci. India 56:
subcylindrical to obtuse-fusiform, 1-septate, 1109, 1987.
constricted at the septum, dark brown, wall J. herbatilis Kohlm., Volkm.-Kohlm. & O.E.
thickened at both ends, no sheath or Erikss., Bot. Mar. 40: 296, 1997.
49
 a b

c d e

f g h

i j k l

m n o p

Fig. 22. Halotthia posidoniae. a. Ascomata conical partly immersed in rhizomes of the sea grass Posidonia oceanica.
b. Surface of ascoma is rough. c-d. Ostiolar region of ascomata. f. Cylindrical ascus. g-h. Asci and pseudoparaphyses
arising from basal ascogenous tissue. i-k. Apical region of asci with an apical pore. l. Thin walled anastomosing
pseudoparaphyses. m-p. Thick-walled, 1-septate ascospores. Bars a = 500 µm; b, e = 200 µm; c-d, h = 100 µm; f-g, l
= 20 µm; i-k, m-p = 10 µm.

50
 Fungal Diversity

Ascomata subglobose, obpyriform to Microthelia maritima (Linder) Kohlm., Nova

ellipsoidal, immersed becoming erumpent, Hedw. 2: 322, 1960.
Microthelia linderi Kohlm., Trans. Br. Mycol.
ostiolate, papillate to epaplliate, coriaceous, Soc. 57: 483, 1971.
clypeate, brown to dark brown, solitary or Ascomata on driftwood, bark, or
gregarious, pseudoparaphyses septate, coniferous wood, ascomata small (57-128 µm
branching and anastomosing, asci clavate to high, 104-268 µm in diameter), semiglobose,
cylindrical, short pedunculate, thick-walled, superficial, ostiolate, short papillate,
bitunicate, fissitunicate, ocular chamber apical carbonaceous, black and gregarious.
ring J-, ascospores elipsoidal, muriform, 7-8- Pseudoparaphyses septate, anastomosing, asci
transeptate, 1-longiseptate, constricted at the clavate to elongate-ellipsoidal, pedunculate,
septa, golden-brown, with a gelatinous sheath. thick-walled lacking an apical apparatus, and
Julella avicenniae was originally ascospores brown, 1-septate and constricted at
described as a Pleospora species, but the septum.
transferred to Julella on the basis of it The genus Kirschsteiniothelia has been
possessing ascomata that develop on woody referred to the Pleosporaceae (Eriksson and
substrata, immersed beneath a clypeus, the Hawksworth, 1981; Kirk et al., 2001),
peridium with a single layer of elongated cells, Pleomassariaceae (Barr, 1993), and
and narrow pseudoparaphyses (Hyde, 1992c). questionably the Massarinaceae (Kodsueb et
However, it is frequently found on the aerial al., 2006). The genus appears to be
twigs of Avicennia marina in Morib mangrove, polyphyletic (Shearer, 1993a) and Schoch et al.
Malaysia damaged by a moth larva (Jones, (2006) are of the opinion that K. aethiops does
unpublished data). not belong in the Pleosporaceae. Kodsueb et al.
Although clustering within the (2006) show that K. elaterascus (a freshwater
Pleosporales, its relationship with other taxa in species) clusters with Massarina
the order cannot be inferred from data ramunculicola in a sister clade to the
currently available (Tam et al., 2003). In a Melanommataceae. However, K. elaterascus
new analysis, J. avicenniae forms a sister differs from K. maritima, and other
clade to one comprising Helicascus, Kirschsteiniothelia species, in ascus structure,
Massarina velataspora and Kirschsteiniothelia its unusual endoascus with a long coiled base
elaterascus (Suetrong et al., unpublished data). that uncoils during ascus dehiscence,
A second maritime species occurs on the ascospore measurements, the presence of an
senescent leaves of Juncus roemerianus, some ascospore sheath and its freshwater occurrence
23-118 cm above the rhizome. Kohlmeyer et (Shearer, 1993a).
al. (1997) regarded J. herbatilis as
facultatively marine. It is included in the key
below for comparison with J. avicenniae. Leptosphaerulina McAlpine, Fung. Diseas.
Stone-Fruit Trees: 103, 1902 ....................... (1)
L. mangrovei Inderb. & E.B.G. Jones,
1. On mangrove wood (Avicennia species), ascospores
28-36 × 12-16 µm, hyaline but generally brown........ Mycoscience 41: 233, 2000.
............................................................. J. avicenniae Ascomata erumpent on wood, dark
1. Facultatively marine on culms of Juncus brown, globose, papillate, ostiolate, mem-
roemerianus, ascospores 18-22.5 × 5-6.5 µm, branous, with dark brown septate, hyphae
hyaline.....................................................J. herbatilis
anchoring ascoma to substratum, pseudopara-
physes sparse, asci elongate to saccate, clavate,
Kirschsteiniothelia D. Hawksw., Bot. J. Linn. few in number, persistent in nature but
Soc. 91: 183, 1985 ....................................... (1) deliquescing in culture, ascospores light brown,
◙ K. maritima (Linder) D. Hawksw., Bot. J. muriform with 4-5 trans-septa and 1-2
Linn. Soc. 91: 193, 1985. longitudinal septa, thin-walled, ellipsoidal to
Amphisphaeria maritima Linder, Farlowia 1: 411, oval, apical cell may be hyaline, with
1944.

51
pronounced sheath constricted at the central structure around the ocular chamber and
septum (Fig. 23). ornamented ascospores Kohlmeyer and
The first marine species of Volkmann-Kohlmeyer, 1990a). Molecular data
Leptosphaerulina reported from submerged confirm its position in the Pleosporales, but
attached decaying branches of the mangrove cannot be assigned to any family within the
tree Kandelia candel (Inderbitzin et al., 2000), order (Suetrong et al., unpublished data). It
also found on the herbaceous mangrove plant forms a clade comprising Massarina species,
Acanthus ilicifolius (Jones, pers. observ.) Few and Leptosphaeria bicolor. Further taxon
Leptosphaerulina species occur in warmer sampling is required to resolve its position.
climates, while this fungus is the only species
found on wood and in the marine environment.
Its occurrence in the Pearl River Estuary may Massariosphaeria (E. Müll.) Crivelli, Diss.
be explained by the variable salinity of the Eidgenöss. Techn. Hochschule Zürich 7318:
water, which ranges from 5-24 ‰ 141, 1983...................................................... (2)
The genus includes some 50 names and ◙ M. typhicola (P. Karst.) Leuchtm., Sydowia
is generally referred to the Pleosporaceae 37: 168, 1984.
Leptosphaeria typhicola Karst. Mycol. Fenn. 2:
(Eriksson and Hawksworth, 1991; Kirk et al., 100, 1873.
2001), although Kodsueb et al. (2006) find Leptosphaeria baldingerae Fautrey & F. Lamb.,
that Leptosphaerulina and Macroventuria are Revue Mycol., Toulouse, 19: 53, 1897.
phylogenetically related with no affinities with Leptosphaeria cladii Cruchet Bull. Soc. Vaud.
the Pleosporaceae. These two genera lack Sci. Nat. 55: 161, 1923.
Phaeosphaeria baldingerae (Fautrey & F.
pseudoparaphyses, a taxonomic character that Lambotte) Hedjar., Sydowia 22: 87, 1969.
differentiates the Pleosporales from the Phaeosphaeria typhicola (P. Karst.) Hedjar.,
Dothideales. We therefore refer the genus to Sydowia 22: 86, 1969.
Dothideomycetes incertae sedis. Chaetomastia typhicola (P. Karst.) M.E. Barr,
Mycotaxon 34: 514, 1989.
◙ M. erucacea Kohlm., Volkm.-Kohlm., &
Lineolata Kohlm. & Volkm.-Kohlm., Mycol. O.E. Erikss., Can. J. Bot., 74: 1835, 1996.
Res. 94: 687, 1990 ....................................... (1) Ascomata subglobose, immersed,
◙ L. rhizophorae (Kohlm. & E. Kohlm.) ostiolate, epapillate to papillate, coriaceous,
Kohlm. & Volkm.-Kohlm., Mycol. Res. 94: light to dark brown, pseudoparaphyses septate,
688, 1990 (Type species). densely packed in a gelatinous matrix, asci
Didymosphaeria rhizophorae Kohlm. & E. cylindrical to clavate, short pedunculate, thick-
Kohlm., Icones Fungorum Maris, 4/5: 62a, 1967. walled, with an ocular chamber, ascospores
Lojkania rhizophorae (Kohlm. & E. Kohlm. M.E. fusiform, hyaline to light brown, 7-14-septate
Barr, N. Amer. Flora, Ser, 2, 13: 58, 1990. with a gelatinous sheath. Both species have
Ascomata obpyriform, immersed to pycnidia with globose, ovoid to ellipsoidal
superficial, ostiolate, papillate, subcarbona- hyaline conidia.
ceous to subcoriaceous, periphysate, and dark Massariosphaeria typhicola has been
brown to black, pseudoparaphyses trabeculate, assigned to different genera with Leuchtmann
in a gelatinous matrix, asci cylindrical, short (1984) considering it best placed in
pedunculate and ascospores 1-septate, Massariosphaeria, while Barr’s (1989) more
ellipsoidal, brown with surface sculpturing. recent review places it in Chaetomastia.
Originally described as Didymosphaeria Leuchtmann (1984) referred Massariosphaeria
rhizophorae (Didymosphaeriaceae, Melanom- to the Lophiostomatceae while Barr (1989)
matales), but transferred to this new genus places it in the Dacampiaceae. Later she
because it differs in the following features: no assigned Chaetomastia to the new family
clypeus, almost superficial ascomata, a Teichosporaceae (Pleosporales) (Barr, 2002).
coloured peridium, a hamathecium with a A molecular study is required to resolve the
gelatinous matrix, asci with an apical ring-like phylogeny of these genera.

52
 Fungal Diversity

a b

c d

Fig. 23. Leptosphaerulina mangrovei. a. Ascoma superficial on substratum. b. Immature asci. c. Ascus with ectoascus
ruptured (lower arrow) and extension of the endoascus (upper arrow). d. Line drawing of ascospore. Bars a-d = 10 μm.

chamber to the ascus and ascospores with

1. Ascospores 7-11-septate, 34-62 × 7-13 µm ............... prominent apical pores and lacking a
.............................................................. M. typhicola mucilaginous sheath. Fresh material was not
1. Ascospores 10-14-septate, 50-71.5 × 9-12 µm ..........
...............................................................M. erucacea available for a molecular study, so its
relationship to Helicascus can not be
determined at this stage.
Salsuginea K.D. Hyde, Bot. Mar. 34: 315,
1991.............................................................. (1) Tirisporella E.B.G. Jones, K.D. Hyde & Alias,
◙ S. ramicola K.D. Hyde, Bot. Mar. 34: 316, Can. J. Bot. 74: 1489, 1996.......................... (1)
1991 (Type species). T. beccariana (Ces.) E.B.G. Jones, K.D. Hyde
Ascomata fusiform, conical or & Alias, Can. J. Bot. 74: 1490, 1996 (Type
subglobose, immersed under a dark clypeus, species).
ostiolate, short neck, pseudoparaphyses thin- Sphaeria beccariana Ces., Atti Acad. Sci. Fis.
walled, numerous, anastomosing, asci Mat. Napoli 8: 20, 1880.
Melanomma cesatianum (Ces.) Sacc., Syll.
cylindrical, pedunculate, fissitunicate, with an Fungorum 2: 113, 1883.
apical apparatus, large ocular chamber and Gibberidea nipae Henn., Hedw. 47: 257, 1908.
prominent ring, ascospores 1-septate, Tryblidiella beccariana (Ces.) Sacc. Syll. Fung.
constricted at the septum, brown to black, 2: 758, 1883.
smooth-walled, with hyaline apical germ pores Anamorph: Phialophora cf. olivacea.
and lacking a sheath or appendages (Hyde, Probably one of the earliest ascomycetes
1991b). Salsuginea differs from another described from the marine environment on the
mangrove bitunicate ascomycete Helicascus in marine palm Nypa fruticans collected by
lacking a stroma, as the ascomata are formed Cesati (1880) in the Philippines and named
under a clypeus with a distinctive ocular Sphaeria beccariana. Hennings (1908)
53
collected an identical taxon Gibberidea nipae. Ascomata scattered, immersed,
This is a common species on the lower ellipsoidal, ostiolate, papillate, periphyses
bases of Nypa fronds and the large, lacking, glaborous, black, pseudoparaphyses,
carbonaceous, immersed to superficial thin, septate, with a slime layer, asci numerous,
ascomata easily detected by touch. cylindrical, short pedunculate, ascospores
Pseudoparaphyses are unbranched and narrowly fusiform, straight or slightly curved
constricted at the septa, asci bitunicate, 1-3-septate, constricted at the central septum,
fissitunicate, thick-walled with an apical hyaline, smooth with a sheath. Shoemaker and
apparatus comprising a canal and pore, while Babcock (1989) transferred this species from
ascospores are brown, 4-7 septate, verrucose Leptosphaeria to Wettsteinina, although they
with a polar appendage formed by the state “the type has hyaline ascospores that are
inversion of an apical sheath-like material (Fig. variable in shape, long and slender and 1-
24) (Jones et al., 1996). Referred to different septate when young, later broader and tardily
orders including the Patellariales, it is best left appearing 3-septate forming a ring-like septa”.
as Pleosporales incertae sedis until molecular Kodsueb et al. (2006) examined the
studies are undertaken to determine its phylogenetic relationship of three Wettsteinina
phylogenetic position. species and showed they were monophyletic,
but distinct from the Pleosporaceae. Schoch et
al. (2006) showed that W. lacustris also did
a b
not group within the Pleosporaceae clustering
with Massaria platani. Placement in the
Pleomassariaceae is suggested by Kodsueb et
al. (2006) but further studies are required with
wider taxon sampling.

c d DOTHIDEOMYCETES incertae sedis

1. Lirelliform ascomata, erumpent or superficial,

opening by an elongate slit, interascal tissue narrow,
cellular pseudoparaphyses, asci cylindrical,
fissitunicate, ascospores hyaline or brown, septate,
sometime with a mucilaginous sheath, anamorphs
varied ....................................................... Hysteriales

Fig. 24. Tirisporella beccariana. Light microscope 1. Stromata absent, ascomata erumpent, eventually
micrographs. a. Ascus with short pedicel. b, d. apothecial, interascal tissue narrow anastomosing
Ascospores with pale or hyaline polar cell to which the pseudoparaphyses, asci cylindrical, fissitunicate, I-,
appendage remains attached. c. Scanning electron with ocular chambers, ascospores hyaline or brown,
micrograph, ascospore with polar appendage (Ap) septate or muriform, usually no sheath, anamorphs.
formed by fragmentation of a sheath. Arrow highlight Anamorphs coelomycetes where known....................
verrucose spore wall. Bars a-b = 20 µm; c = 5 µm, d = ...............................................................Patellariales
10 µm.
1. Ascomata perithecial, superficial rarely immersed,
Wettsteinina Höhn., Sber. Akad. Wiss. Wien, on a stalk, mycelium wide, asci clavate or cylindrical,
Math.-naturw. Ki., Abt. 1 116: 126, 1907.... (1) pseudoparaphyses present, ascospores brown,
W. marina (Ellis & Everh.) Shoemaker & C.E. generally 1-septate, with a sheath, anamorphs rare....
................................................................. Jahnulales
Babc., Can. J. Bot. 67: 1596, 1989.
Leptosphaeria marina Ellis & Everh., J. Mycol.
1: 43, 1885.
Heptameria marina (Ellis & Everh.) Cooke,
Grevillea 18: 32, 1889.
Metasphaeria marina (Ellis & Everh.) Berl., Icon.
Fung. 1: 140, 1894.
Leptosphaeria treatiana Sacc., Syll. Fung. 10:
923, 1892.
54
 Fungal Diversity

HYSTERIALES Patellaria Fr., Syst. Mycol. (Lundae) 2: 158,

Hysteriaceae 1822.............................................................. (1)
P. atrata (Hedw.) Fr., Syst. Mycol. 2: 160,
Gloniella Sacc., Syll. Fung. (Abellini) 2: 765, 1822.
1883.............................................................. (1) Recently a Patellaria species was
G. clavatispora Steinke & K.D. Hyde, collected on mangrove wood in Hong Kong
Mycoscience 38: 7, 1997. (primarily on Kandelia candel) and Thailand.
A species collected at a number of sites Numerous Patellaria species have been
in South Africa (Steinke and Jones, 1993; described, primarily from temperate locations,
Steinke and Hyde, 1997a) and has dark-brown, but none from marine habitats. A brief
rounded or discoid to cupulate, flat description follows as it is frequently
hysterothecioid ascomata, that are superficial encountered on mangrove wood.
and gregarious. Pseudoparaphyses swollen at Ascomata arising singly, initially closed,
the tips, rarely branching, septate, hyaline and later opening by a pore to form a flat or
longer than the asci. Asci clavate, short pedicel, convex black disc, apothecioid, superficial,
bitunicate, fissitunicate and with an apical sessile, circular, subgelatinous when moist,
opercular chamber. Ascospores hyaline, 0.3-0.9 (width) × 0.6 mm (high), (x = 0.7 ×
smooth 5-8-septate and clavate. This 0.57 mm, n = 38). Margin entire, raised and
resembles a Patellaria species collected in incurved. Outer surface of stalk smooth.
Hong Kong on Kandelia candel, but differs in Hypothecium thin-walled with isodiametric
that the paraphyses are not branched. Saprobic colourless cells 16-40 µm (x = 29.3 µm, n = 9).
on intertidal Avicennia marina. Hamathecium paraphysoidal, hyaline,
branched above, 96-172 × 1-2 µm (x = 132.4 ×
1.4 µm, n = 50). Asci cylindric-clavate, with a
PATELLARIALES stipe, bitunicate, thick-walled, with ocular
Patellariaceae chamber in apical dome, fissitunicate, 8-
spored, J- in Meltzer’s reagent, 92-150 × 14-40
Banhegyia L. Zeller & Toth, Sydowia 14: 326, µm (x = 116 × 23 µm, n = 45). Ascospores
1960.............................................................. (1) irregularly biseriate, clavate, slightly curved,
B. setispora L. Zeller & Toth, Sydowia 14: not constricted at the septa, 4-9-septate,
327, 1960 (Type species). hyaline, no mucilaginous sheath, 24-48 × 6-24
Celidium proximellum var. uralensis Naoumov, µm (x = 37.6 × 10.2 µm, n = 70) (Fig. 27).
Bull. Soc. Mycol. France 30: 384, 1914.
No anamorph in culture.
Ascomata semiglobose, discoid,
Habitat: on badly decayed wood of Kandelia
apothecial-like, erumpent, leathery, dark
candel, Three Fathom Cove, Hong Kong SAR,
brown, pseudoparaphyses septate, simple,
China.
hyaline to light brown, extending beyond the
Distribution: Hong Kong SAR, China and
asci, apically club-shaped, asci clavate to
Thailand.
ellipsoidal, short pedunculate, no apical
apparatus, ascospores ellipsoidal, 1-septate, JAHNULALES
constricted at the septa, hyaline to pale brown Hypsostromataceae
with bristle-like appendages at either end of
the spore. Collected on coniferous wood with a Manglicola Kohlm. & E. Kohlm., Mycologia
restricted distribution (Kohlmeyer and 63: 840, 1971 ............................................... (1)
Kohlmeyer, 1979). No molecular data is ◙ M. guatemalensis Kohlm. & E. Kohlm.,
available to support its inclusion in the Mycologia 63: 841, 1971 (Type species).
Patellariaceae and fresh collections are Fresh collections of this unique
necessary to enable a re-evaluation of its ascomycete on Nypa fruticans in Thailand has
phylogenetic relationship. enabled a re-evaluation of its phylogenetic

55
relationship in the Dothideomycetes (Suetrong EUROTIOMYCETES
et al., 2009, in press). Ascomata 1100-1750
µm high, 290-640 µm around the center, 1. Saprobic, parasitic or mycorrhizal, ascomata
obtusely clavate to fusiform, stipitate, generally cleistothecial, globose, brightly coloured,
hamathecial elements lacking, asci evanescent,
peridium differentiated into several layers, scattered throughout ascoma, ascospores unicellular,
superficial on the substratum, epapillate, lenticular, sphaerical or elliptical...............................
coriaceous, solitary and olive brown. ..................................................... Eurotiomycetidae
Pseudoparaphyses narrow, numerous, septate
anastomosing, or simple, asci cylindrical, 1. Lichenized, parastic and saprobic, ascomata
thick-walled, few in number and developing at perithecial, superficial or immersed in a thallus, asci
clavate to cylindrical, hamathecium of
the base of the ascomata, ascospores fusiform, pseudoparaphyses, ascospores variable, hyaline to
apiculate, unequally 1-septate, constricted at brown, simple or muriform........................................
the septum, apical cell larger, orange-brown to ............................................Chaetothyriomycetidae
chestnut brown, basal cell turbinate, light
brown, with a gelatinous appendage (Fig. 25).
Phylogenetically Manglicola guatema- EUROTIOMYCETIDAE
lensis is the first marine member of the
Jahnulales, although the anamorphic fungus Two orders with marine taxa.
Xylomyces rhizophorae has also been reported
from marine locations (Kohlmeyer and 1. Peridium composed of thick-walled cells, ascomatal
appendages present .................................Onygenales
Volkmann-Kohlmeyer, 1998d). Recently a 1. Peridium thin, membranous, no ascomatal
Xylomyces species has been shown to be an appendages................................................Eurotiales
anamorphic species in the Jahnulales
(Campbell et al., 2007). Initially M.
guatemalensis was thought to be closely ONYGENALES
related to the Pleosporaceae or Venturiaceae Gymnoascaceae
(Kohlmeyer and Kohlmeyer, 1971) while
Huhndorf (1992) and Huhndorf et al., (1994) Gymnascella Peck, Ann. Rep. Reg. St. N.Y.
classified it in the Hypostromataceae incertae 35: 143, 1884. .............................................. (1)
sedis. G. littoralis (G.F. Orr) Currah, Mycotaxon 24:
Morphological and molecular evidence 87, 1985
places it in the Jahnulales with strong Plunkettomyces littoralis G.F. Orr, Mycotaxon
bootstrap support with Aliquandostipite 6: 34, 1977.
species as a sister group (Fig. 26) (Suetrong et Gymnascus littoralis (G.F. Orr) Arx, Persoonia,
13: 179, 1986.
al., 2009, in press). This is yet another marine Arachniotus littoralis (Orr) Arx, Persoonia 9:
lineage and is of particular interest as all other 397, 1977.
Jahnulales members are freshwater or peat
swamp species (Pang et al., 2002; Pinruan et Ascomata yellow to orange brown,
al., 2002). It has been hypothesised that globose, peridial hyphae, hyaline to yellow,
marine fungi are derived from terrestrial or simple or branched, smooth, slightly thick-
freshwater habitats that have migrated into the walled, asci globose, ascospores yellow-brown
sea (Shearer, 1993c; Jones, 2000). The to orange-brown, smooth, thick-walled, oblate
mangrove habitat of M. guatemalensis may with equatorial rim, 2.9-3.8 × 4.2-5.7 µm,
well form a link between lignicolous anamorph with arthro and aleurioconidia 2.0-
freshwater taxa and estuarine to marine 2.8 × 3.5-7 µm. Gymnascella has priority over
environments. Vijaykrishna et al. (2006) have Arachniotus for those with oblate ascospores.
examined the ancestry of freshwater taxa from A well-characterized member of the
terrestrial species and conclude this migration Onygenales (Currah, 1985).
occurred 390 million years ago.

56
 Fungal Diversity

a b

c d e f

g h i j k l

m n o p

Fig. 25. Manglicola guatemalensis. a. Mature ascomata of Manglicola guatemalensis on the surface of Nypa fruticans,
partially immersed in mud. b, d. Ascoma superficial seated on the substratum. c. Longitudinal section of ascoma with
stalk, asci and pseudoparaphyses. e. Narrow pseudoparaphyses. f. Cylindrical ascus. g. Ascus tip with ocular chamber.
h. Ascospore in ascus with apical and basal appendages (arrow). i-p. Bicelled ascospores. Bars a = 500 µm; b = 250
µm; c-d = 100 µm; e, h = 10 µm; f-g = 50 µm; i-p = 20 µm

57
Fig. 26. Phylogram generated from weighted parsimony analysis (step matrix) from combined SSU and LSU rDNA
sequences. Parsimony bootstrap value greater than 50% and Bayesian Posterior Probabilities greater than 0.95 are given
above and below each clade, respectively.

58
 Fungal Diversity

a b c

d e f g h

Fig. 27. Patellaria cf atrata. a. Superficial apothecium. b. Section through ascoma with hymenium. c, e.
Pseudoparaphyses with branched club-shaped tips. d. Immature thick-walled asci and pseudoparaphyses. f-g. Clavate
asci. h-i. Ascospores hyaline, clavate, curved 4-9 septate. Bars a-b = 200 µm; c-g = 25 µm; h-i = 10 µm.

EUROTIALES and transversely septate to muriform..........................

Trichocomaceae .........................................................Chaetothyriales

1. Majority are lichenized associated with green algae,

Eupenicillium F. Ludw., Lehrb. Nied. Krypt.: nonlichenized species referred to the
p. 256, 257, 263, 1892.................................. (1) Requienellaceae, thin thallus, ascomata immersed
◙ E. limosum S. Ueda, Mycoscience 36: 451, or superficial, ostiolate, papillate, pseudoparaphyses
1995. trabeculate, asci clavate, ascospores hyaline to
brown, transversely septate to muriform ....................
Anamorph: Penicillium limosum Ueda ................................................................Pyrenulales
Ascomata cleistothecial, globose to
subglobose, superficial, scattered, pale yellow, 1. Mostly lichenized, saxicolous, ascomata superficial
asci subglobose to ellipsoidal, evanescent at to immersed in the thallus, hamathecium often
maturity, ascospores subglobose 3-3.5 × 2.5-3 absent or evanescent tissue of gelatinized
pseudoparaphyses, ostiole covered with periphyses,
µm, hyaline spore wall roughened (Udea, asci fissitunicate or evanescent, ascospores hyaline
1995b). Isolated several times from marine to brown, septate to muriform ............ Verrucariales
sediments in Nahasaki, Japan (Udea, 1995b).
CHAETOTHYRIALES
Herpotrichellaceae
CHAETOTHYRIOMYCETIDAE
Three orders with marine taxa (Geiser et al., 2001).
Capronia Sacc., Syll. Fung. (Abellini) 2: 288,
1883 ..............................................................(1)
1. Primarily nonlichenized taxa, dark mycelium on C. ciliomaris (Kohlm.) E. Müll., Petrini, P.J.
substrata or inconspicuous immersed mycelium, Fisher, Samuels & Rossman, Trans Br. Mycol.
ascomata erumpent to superficial, sometimes setose, Soc. 88: 73, 1987.
short apical periphysoids, asci clavate, thickening of Herpotrichiella ciliomaris Kohlm., Nova Hedw.
the apical region, ascospores hyaline to pale grey 2: 313, 1960

59
 Ascomata globose to ovoid, superficial 1991), it is common on various mangrove tree
rarely immersed, ostiolate, epapillate, species, especially Rhizophora apiculata and
membranous, variable in colour from hyaline Sonneratia species (Alias et al., 1996; Jones
to light to dark blue to black, solitary or and Abdel-Wahab, 2005; Jones and Pugsili,
gregarious, apical paraphysoids merging with 2006). Aptroot (1991) states that the fungus
periphyses, asci cylindrical to subclavate, stains the wood purple, we have not observed
short pedunculate, bitunicate, thick-walled this. However, we have noted that the wood
apically, ascospores ellipsoidal to subovoid, around the ascomata is bleached white and
1-septate, constricted at the septum, hyaline, may be due to enzyme action. It is easily
with a crown of cilia-like appendages at each recognizable on mangrove wood by the
pole. Initially described as a Herpotrichiella it prominent white zones surrounding the raised
was transferred by Muller et al. (1987) to ascomata. Its position high in the intertidal
Capronia based on morphological zone exposes it to sunlight and long periods
characteristics. An ultrastructural study (Au et of drying out at low tides. Found in locations
al., 1999b) showed that the periphysoids arise not subject to routine submergence. Here
from the upper third of the ascomal wall, placed in the Pyrenulaceae, it has also been
extending through the ostiolar canal and referred to the Requienellaceae (Kirk et al.,
merging with the apical setae. The ascus has 2001).
no ocular chamber, but the endoascus is
thickened at the apex. Ascospores are
verrucose, while the cilia-like appendages Xenus Kohlm. & Volkm.-Kohlm.,
arise sub-terminally from the mesosporium Crytogamie Bot. 2: 367, 1992.......................(1)
through discontinuities in the episporium. A X. lithophylli Kohlm. & Volkm.-Kohlm.,
frequently collected species on bark in Friday Crytogamie Bot. 2: 368, 1992 (Type species).
Harbour, USA (Jones, 1985; Au et al., 1999b). Ascomata subglobose, superficial,
The taxonomic position of C. ciliomaris ostiolate, periphysate, epapillate, clypeate,
remains in question, as it is the only marine black and gregarious, pseudoparaphyses
species in the genus, with hyaline ascospores, trabeculate, branched and anastomosing, asci
1-septate with a crown of sub-terminal clavate, thick-walled, without an apical
appendages and no anamorph has been apparatus, ascospores 1-3-septate, slightly
reported for it. These are not characteristic constricted at the central septum, hyaline, no
features of Capronia. appendages or sheath (Kohlmeyer and
Volkmann-Kohlmeyer, 1992).
PYRENULALES A monotypic genus parasitic on the red
Pyrenulaceae alga Lithophyllum sp., attached to coral rock
collected in the Caribbean (Belize).
Pyrenographa Aptroot, Bibliotheca Kohlmeyer and Volkmann-Kohlmeyer (1992)
Lichenologica 44: 103, 1991........................ (1) referred it to the Dothideales incertae sedis,
◙ P. xylographoides Aptroot, Bibliotheca but noted similarities with the bitunicate
Lichenologica 44: 103, 1991. ascomycetes Arthopyrenia halodytes,
Ascomata developing under a dark Pharcidia laminariicola and P. rhachiana. Its
stroma, subglobose to fusiform, ostiolate, taxonomic position is far from clear but a
short papilla, periphysate, pseudoparaphyses concensus places it in the Pyrenulales.
branched, septate, asci clavate to cylindrical,
short pedunculate, thick-walled, bitunicate, Requienellaceae
with an apical ring, J-, ascospores ellipsoidal
to fusiform, 3-septate, not constricted at the Mauritiana Poonyth, K.D. Hyde, Aptroot &
septa, yellow brown, thick-walled, smooth Peerally, Fungal Diver. 4: 102, 2000 ............(1)
and lacking a sheath or appendages (Fig. 28). ◙ M. rhizophorae Poonyth, K.D. Hyde,
Originally described from mangrove wood Aptroot & Peerally, Fungal Diver. 4: 102,
collected in Queensland, Australia (Aptroot, 2000 (Type species).

60
 Fungal Diversity

a b

c d e

Fig. 28. Pyrenographa xylographoides. a. Ascoma within a black stroma. b. Longitudinal section through an ascoma on
a raised cushion of wood. c. Ascus clavate with immature ascospores. d. Pseudoparaphyses branched. e-h. Ascospores
brown 3-septate. Bars a-b = 200 µm; c-h = 10 µm.

61
 Ascomata globose to ovoid, immersed, C. sublitorale (Leight.) Grube & B.D. Ryan
ostiolate, short neck, pale brown, gregarious, Lich. Fl. Greater Sonoran Desert 1: 163, 2002.
pseudoparaphyses filamentous, septate, Verrucaria sublitoralis Leight, Lich. Fl. Gr.
branching, asci cylindrical to clavate, with an Britain: 435. 1871.
Arthopyrenia sublitoralis (Leight.) Arnold, Ber.
ocular chamber, bitunicate, thick-walled, short Bayer. Bot. Ges. 1 (suppl.): 121, 1891.
pedunculate, ascospores fusiform, dark brown Pyrenocollema sublitorale (Leight.) R.C. Harris,
the end cells paler, 9-13-distoseptate, septa ex Fletcher Lichenologist 24: 368, 1992.
thick, slightly constricted at the central septum, C. foveolatum (A.L. Sm.) F. Mohr, Mycol.
smooth walled and lacking a sheath or Res. 108: 529, 2004.
appendages (Fig. 29). This species has been Arthopyrenia foveolata A.L. Sm., J. Bot. 49: 43,
recovered from marine habitats (Jones, 1911.
unpublished data on branches of Hibiscus C. ostrearum (Vain.) F. Mohr, Mycol. Res.
tiliaceus immersed in the intertidal, Thailand 108: 530, 2004.
Lecanactis ostrearum Vain., Cat. Welwitsch. Afr.
and Guam) although generally growing on the Pl 2: 430, 1901.
more terrestrial parts of mangrove trees, Arthoniactis ostrearum (Vain.) Clem. & Shear,
especially Rhizophora mucronata. Poonyth et Gen. Fungi :319, 1931.
al. (2000b) referred the genus to the C. pelvetiae (G.K. Sutherl.) Kohlm., D.
Pyrenulales sensu stricto, as it has immersed Hawksw. & Volkm.-Kohlm., Mycol. Prog.
ascomata, interascal tissue composed of 3: 54, 2004.
branched pseudoparaphyses, thick-walled, Arthopyrenia pelvetiae (G.K. Sutherl.) D.
fissitunicate asci and brown, septate Hawksw., Lichenologist 12: 106, 1980.
Dothidella pelvetiae G.K. Sutherl., Trans. Br.
ascospores. Mycol. Soc 5: 154,1915.
Leiophloea pelvetiae (G.K. Sutherl.) Kohlm. &
Kohlm., Marine Mycology, the Higher Fungi, 376,
Xanthopyreniaceae 1979.
Placostroma pelvetiae (G.K. Sutherl.) Meyers,
Mycologia 49: 480, 1957.
Collemopsidium Nyl., Flora 64: 6, 1881. .... (6) Pyrenocollema pelvetiae (G.K. Sutherl.) D.
C. halodytes (Nyl.) Grube & B.D. Ryan Lich. Hawksw. J. Linn. Soc., Bot. 96: 10, 1988.
Fl. Greater Sonoran Desert Region 1: 163, Kohlmeyer et al. (2004) transferred
2002 (Type species). Pyrenocollema pelvetiae to Collemopsidium.
Verrucaria halodytes Nyl., Mém. Soc. Sci. Nat. Species assigned to Collemopsidium have
Cherbourg 5: 142, 1857.
Arthopyrenia halodytes (Nyl.) Arnold, Ber.
variously been placed in Verrucaria,
Bayer. Bot. Ges. 1 (suppl.): 121, 1891. Arthopyrenia and Pyrenocollema (Mohr et al.,
Pyrenocollema halodytes (Nyl.) R.C. Harris, 2004). Pyrenocollema orustensis may not be a
Bryologist 90: 164, 1987. synonym of Collemopsidium halodytes as this
Verrucaria consequens Nyl., Flora 47: 357, 1864. taxon may include several morphological
Arthopyrenia consequens (Nyl.) Arnold, Flora 53:
485, 1870.
entities (Fig. 30). The genus is assigned to the
Arthopyrenia kelpii Körb., Parerga Lich. 387, Xanthopyreniaceae (Eriksson et al., 2003), a
1865. family of uncertain position, although Grube
Verrucaria fluctigena Nyl., Flora 58: 14, 1875. and Ryan (2002) included it in the Dothideales
Arthopyrenia orustensis Erichsen, Nyt Mag. incertae sedis. The position of C. pelvetiae is
Naturv. 68: 159, 1930.
Pyrenocollema orustensis (Erichsen) A. Fletcher,
in doubt as it is considered to be an epiphyte
24: 368, 1992. rather than a lichenized ascomycete, and it has
C. elegans (R. Sant.) Grube & B.D. Ryan, much larger ascomata than other
Lich. Fl. Greater Sonoran Desert Region 1: Collemopsidium species (Mohr et al., 2004).
163, 2002. Lichenized photobionts are cyanobacteria
Pyrenocollema elegans R. Sant., Lichenologist (Hyella) (Harada, 2000) or non-lichenized
24: 7, 1992. Phaeophyceae (Pelvetia).

62
 Fungal Diversity

a b c

Fig. 29. Mauritiana rhizophorae. a. Cylindrical ascus with biseriate ascospores. b. Scanning electron micrographs of
ascospore. c-d. Fusiform ascospores with 12-13 distoseptate. Bars a, c-d = 10 µm; b = 5 µm. (Photos by Aisyah Alias).

1. Perithecia large (310-590 μm) on the seaweed VERRUCARIALES

Pelvetia canaliculata............................... C. pelvetiae Verrucariaceae
1. Perithecia smaller (less than 0.6 mm in diam.) not on
seaweeds .................................................................. 2
Mycophycias Kohlm. & Volkm.-Kohlm., Syst.
2. Thallus superficial on the substratum....................... 3 Ascomycetum 16: 2, 1998 ........................... (2)
2. Thallus immersed in the substratum ........................ 4 M. apophlaeae (Kohlm.) Kohlm. &Volkm.-
Kohlm., Syst. Ascomycetum 16: 3, 1998.
3. Thallus with black, carbonaceous ridges... C. elegans Mycosphaerella apophlaeae Kohlm., Bot. Mar.
3. Thallus lacking black ridges, often immersed in 24: 13, 1981
calcareous rock and shells...................... C. halodytes M. ascophylli (Cotton) Kohlm. & Volkm.-
Kohlm., Syst. Ascomycetum 16: 3, 1998 (Type
4. Involucrellum intermixed with the substratum, wide
spreading .................................................................. 5 species).
Mycosphaerella ascophylli Cotton, Trans. Br.
4. Involucrellum not containing substratal material........
Mycol. Soc. 3: 95, 1908.
..............................................................C. ostrearum
Anamorph: Septoria ascophylli Melnik
5. Perithecia superficial or semi-immersed, 0.15-0.55 & Ju. Petrov, Nov. Sist. Niz. Rast. 1966: 211,
mm diam., involucrellum well developed and 1966.
spreading laterally ................................ C. sublitorale Sphaerella ascophylli (Cotton) Sacc. & Trotter,
5. Perithecia immersed in pits in the substratum, 0.1- Sacc.: Syll. Fung. 22: 147, 1913.
0.24 mm diam., involucrellum lid-like, not Mycosphaerella pelvetiae G.K. Sutherl., New
spreading ............................................. C. foveolatum Phytol. 14: 34-35, 1915.
Key after Mohr et al. (2004). Sphaerella pelvetiae (G.K. Sutherl.) Sacc., Syll.
Fung. 24: 849, 1928.

63
 V. corallensis P.M. McCarthy, Aust. Lichenol.
63: 17, 2008.
V. ditmarsica Erichs., Schr. Naturw. Ver.
Schles.-Holst. 22: 90, 1937.
V. durietzii I.M. Lamb, Lilloa 14: 205, 1948.
V. fusconigrescens Nyl. (1872) In Bull. Soc.
Linn. Normand ser III, vol VI, 1872, p. 266 &
314.
V. halizoa Leight., Lichen Flora of Great
Britain & Ireland, 461, 1871.
V. halochlora H. Harada, Nova Hedw. 60: 74,
1995.
◙ V. maura Wahlenb., in Acharius Methodus:
Fig. 30. Habitat photomicrograph of Collemopsidium 19, 1803.
halodytes. (Photo by Anthony Fletcher). V. meridionalis P.M. McCarthy, Muelleria 8:
103, 1994.
Ascomata ovoid, ellipsoid to obpyriform, V. microsporoides Nyl., Bull. Soc. Bot. France
immersed in the host, ostiolate, epapillate, 8: 759, 1861.
periphysate, coriaceous, brown solitary or ◙ V. mucosa Wahlenb., In Acharius Methodus:
gregarious, periphysoid in the upper part of the 23, 1803.
ascoma, asci fusiform to clavate, pedunculate, V. psychrophila I.M. Lamb., Discovery Repts.
ascospores ellipsoidal, 1-septate, hyaline, no 25: 18, 1948.
appendages or sheaths. These two species have V. serpuloides I.M. Lamb., Discovery Repts.
been transferred from Mycosphaerella because 25: 20, 1948.
they have periphysate ostioles with the V. sessilis P.M. McCarthy, N.Z.J. Bot. 29: 285,
periphysoids arising from the upper peridium. 1991.
They form mycophycobioses with marine ◙ V. striatula Wahlenb., in Acharius,
macroalgae (Kohlmeyer and Volkmann- Methodus:Methodus: 21, 1803.
Kohlmeyer, 1998b). V. subdiscreta P.M. McCarthy, Muelleria 7:
327, 1991
V. tavaresiae R.L. Moe, Bull. California
1. Ascospores 15-20 × 4-5 μm, mycobiont of Lichen Soc. 4: 8, 1997.
Apophlaea lyallii ................................M. apophlaeae V. tessellatula Nyl., in Crombie, J.Bot.,
1. Ascospores 15-22 × 4-6 μm, mycobiont of
Ascophyllum nodosum and Pelvetia canaliculata .....
London 13: 335, 1875.
............................................................. M. ascophylli Crustose pyrenocarpous lichens
generally with a green unicellular photobiont
Verrucaria Schrad., Spicil. Fl. Germ. 108, (genus Dilabifilum = syn: Pseudopleuro-
1794............................................................ (24) coccus), thallus discrete, scattered, immersed
V. adguttata Zahlbr. (1941) Denkschr. Akad. or superficial, black ascomata, periphysate,
Wiss. Wein, math. –naturw. K1., CIV p. 250. asci and paraphyses deliquesce early, clavate
V. allantoidea H. Harada, Nova Hedw. 60: 75, to cylindroclavate, fissitunicate asci containing
1995. 8 hyaline, ovoid to subglobose to ellipsoidal
V. amphibia Clemente(1807) Apud Acharius, ascospores (McCarthy, 2001) (Figs. 31-33).
Syn. Lich. 1814, p. 94. syn. V. symbalana, Nyl. Many Verrucaria lichens are to be found in the
1873. littoral and supralittoral zone and are thus
V. aucklandica Zahlbr., Denkschr. Akad. thought to be salt-tolerant (Fletcher, 1975,
Wiss., Wein, Mathematische- 1980) but Verrucaria serpuloides has been
naturwissenschaftliche Klasse 104: 250, 1941. found on dredged-up stones from 30 m in
V. bubalina M. Mayrhofer & P.M. McCarthy, Antarctica (Lamb, 1973).
Muelleria 7: 344, 1991. Perithecial ascomata with an apical
V. ceuthocarpa Wahlenb., in Acharius, ostiole, with short pseudoparaphyses bordering
Methodus: 22, 1803. the upper part of the perithecial cavity and
64
 Fungal Diversity

hanging into this without touching the 3. Thallus white to pale-grey, often poorly developed,
hymenium (Gueidan et al., 2007). Bitunicate perithecia semi-immersed, to superficial, solitary to
2 - 3 together, 0.4 - 0.8 mm diam., involucrellum
asci dehiscence often by gelification of the thick and well developed .....................V. halochlora
outer wall.
Molecular studies of the family 4. Thallus not submerged............................................. 5
concluded they were a sister group to the non- 4. Thallus submerged, jet-black, involucrellum well-
lichenised order Chaetothyriales (Lindemuth developed, ascospores broadly ellipsoid 15- 17.5 ×
8-9.5 μm ............................................. V. serpuloides
and Lumbsch, 2001; Lumbsch et al., 2005).
Current studies indicate that the generic 5. Thallus with prominent, glossy, branched and
delineations of the Verrucariaceae were not swollen ridges (jugae).............................................. 6
monophyletic. In fact, Verrucaria is highly 5. Thallus without prominent black ridges (jugae) ...... 7
polyphyletic and is spread out in eleven clades.
Four lineages were identified by Gueidan et al. 6. Thallus brown ....................V. striatula ssp. australis
6. Thallus grass-green .................................. V. striatula
(2007), one constituting a marine group (V.
6. Thallus medium gray green to green black or grey-
mucosa, V. striatula). A second aquatic group black..................................................... V. corallensis
was identified with V. maura (marine), V. 6. Thallus pitch black, with black ridges ... V. amphibia
adriatica and V. scabra (freshwater),
suggesting that this genus migrated to the 7. Thallus continuous to sparingly rimose, lower to
marine environment on at least two separate mid-littoral species................................................... 8
7. Thallus strongly rimose to areolate, upper littoral to
occasions. supralittoral species ............................................... 10
Recently described species include V.
allantoidea, V. halochlora from Japan and V. 8. Ascospores 7-13 μm long ....................................... 9
corallensis, V. meridionalis, V. subiscreta 8. Ascospores 12-15 μm long .............. V. psychrophila
from Australia (McCarthy, 1991, 1994; 8. Ascospores 10-16 (-18) μm long ..V. microsporoides
Harada, 1995; McCarthy 2008) (Fig. 32).
9. Perithecia immersed, to 0.15 mm diam., exciple
Verrucaria tavaresiae is unique in having a colourless, hallus thick, hypothallus white, olive -
brown alga Petroderma maculiforme as a green to dark green-black in sun, on open rocks........
photobiont, while others have green algae ...…………………………………………V. mucosa
(Coccobotrys, Desmococcus, Dilabifilum, 9. Perithecia superficial, 0.2-0.34 mm diam., exciple
medium-grey to brown-black, thallus thin, filmy,
Myrmecia) or a xanthophyte (Heterococcus)
pale olive-green, usually in shaded crevices ..............
photobiont (Moe, 1997). Aquatic Verrucaria ...................................................................V. halizoa
species are generally cold-water species with 9. Perithecia superficial, 0.24-0.45 mm diam., black.....
various numbers recorded from different .......................................................... V. meridionalis
localities: Fidalgo Island Washington (7-9
10. Margin of thallus placodioid-dissected ...................
species), New England, (7 species), Great
.............................................................. V. durietzii
Britain (8 species), the Antarctic Peninsula (6- 10. Margin not dissected placodioid ......................... 11
8 species) and 10 from Scandinavia (Ryan,
1988; Taylor, 1982; Purvis et al., 1992; Lamb, 11. Ascospores 8-15 × 4-7 μm .................................. 12
1948; Santesson, 1993). There are many 11. Ascospores 12-26 × 6-15 μm .............................. 15
terrestrial Verrucaria species known from bark.
12. Thallus black, effuse or in blotches..................... 13
12. Thallus grey-brown to olive-green or green black,
1. Photobiont a brown alga, Petroderma maculiforme .. not effuse or in blotches ...................................... 14
..............................................................V. tavaresiae
1. Photobionts from other algal groups ........................ 2 13. Perithecia sessile, to 0.25 mm diam ........................
............................................................ V. adguttata
2. Ascospores allantoid, 12-14 × 2-3 μm V. allantoidea 13. Perithecia immersed, 0.1-0.16 mm diam.................
2. Ascospores not allantoid .......................................... 3 ....................................................... V. ceuthocarpa
13. Perithecia sessile, 0.02 mm diam ...... V. ditmarsica
3. Thallus epilithic and conspicuous grey-brown to
green-brown, medium-green, green-black or black .. 14. Thallus grey-brown to mid-green-black, perithecia
................................................................................. 4 0.22-0.45 mm diam .........................V. aucklandica

65
14. Thallus dark olive-green to green-black, perithecia ostiolate, hyaline to light-brown, solitary,
0.12-0.22 mm diam ..........................V. subdiscreta appendages born on a side branch, asci 4-
spored, elongate to clavate, thin-walled,
15. Thallus pale-buff to grey-brown.......................... 16
15. Thallus olive-brown, greenish-grey, dark grey
unitunicate, early deliquescing, ascospores 26-
brown to dark greenish-black .............................. 17 35 × 4 µm, elongate-fusiform, pointed at the
apex, rounded at the base, 1-septate, lower cell
16. Thallus pale-buff with conspicuous black cracks, smaller, and surrounded by a mucilaginous
perithecia 0.1-0.2 mm diam., not radially ridged, sheath. The marine status of this species needs
exciple colourless, ascospores 10-15 × 6-9 μm....... questioning as it was found at the base of the
.......................................................... V. tessellatula
16. Thallus buff-brown, grey-brown or green-grey,
elytra of the beetle Aepus robini, living in the
without prominent black cracks, perithecia 0.2-0.3 Laminaria zone (Kohmeyer and Kohlmeyer,
mm diam., often radially ridged, exciple brown to 1979).
brown-black, ascospores 14.5-23.5 × 7-11 μm .......
..............................................................V. bubalina
LECANOROMYCETES
17. Ascospores 9-16 (17) μm long ............................ 18
17. Ascospores 16-26 μm long, thallus olive brownish, LECANOROMYCETIDAE
green grey to dark-brown, prothallus distinct, LECANORALES
exciple brown black ................ V. fusconigrescens Dactylosporaceae

18. Thallus effuse, dull medium-green to green, black, Dactylospora Körb., Syst. Lich. Germ. 271,
areolate only around the perithecia, exciple brown- 1855.............................................................. (3)
black, 25-32 μm thick, ascospores 9-12 (- 16) × 6-
7 (8.2) μm................................................ V. sessilis
D. canariensis Kohlm. & Volkm.-Kohlm.,
18. Thallus strongly rimose to areolate, dark green to Mycotaxon 67: 248, 1998.
greenish black, exciple 10-20 μm thick, hyaline to ◙ D. haliotrepha (Kohlm. & E. Kohlm.)
brown-black, ascospores 12-20 (-22) × 6-8 (- 9) Hafellner, Nova. Hedw. 62: 111, 1979.
μm ............................................................V. maura Kymadiscus haliotrephus (Kohlm. & E. Kohlm.)
Kohlm. & E. Kohlm., Mycologia 63: 837, 1971.
Key modified from Galloway, D.J. 2007: Flora Buellia haliotrepha Kohlm. & E. Kohlm., Nova
of New Zealand. Volume II. Indigenous Hedw. 9: 90, 1965.
Tracheophyta - Monocotyledons except D. mangrovei E.B.G. Jones, Alias, Abdel-
Graminae. First electronic edition, Landcare Wahab & S.Y. Hsieh, Mycoscience 40: 317,
Research, June 2004. Transcr. A.D. Wilton 1999.
and I.M.L. Andres. Apothecia initially sub-globose,
becoming subglobose or discoid, flat or
LABOULBENIOMYCETES convex, superficial, sessile, leathery, dark
LABOULBENIOMYCETIDAE reddish-brown, becoming black, solitary,
LABOULBENIALES: sometimes gregarious, asci clavate, short
Laboulbeniaceae pedunculate, apically thick-walled, without an
apical apparatus, ascospores ellipsoidal or
Laboulbenia Mont. & C.P. Robin, Histoire obovoid, 1-septate, constricted at the septum,
Naturelle des Végétaux Paraites qui croissent with longitudinal or verrcuose ornamentations
sur l’Homme et sur les Naimaraux Vivants, (Fig. 34). No appendages except in D.
Braillière et Fils, Paris: 622, 1853 ............... (1) canariensis. Au et al. (1996) have illustrated
L. marina F. Picard, C. R. Séances Soc. Biol. the complexity of the ascospore wall in D.
Fil. 65: 484, 1908. haliotrepha, the wall consisting of a series of
This species has been described and ridges derived from outgrowths of the
illustrated by Kohlmeyer and Volkmann- mesosporium, and surrounded by the
Kohlmeyer (2003b) on which the following exosporium. The areas between the ridges are
summary is drawn from: Thallus 150-230 µm, filled with mucilage and when the exosporium
receptacles 105-112 × 48-53 µm, ascomata 76- ruptures, the mucilage is lost. Another
118 × 38-44 µm, elongate- ellipsoidal, sessile, observation is that the pseudoparaphyses are

66
 Fungal Diversity

a b

c d

Figs. 31. Habitat micrographs of: a. Verrucaria mucosa. b. V. striatula. c. V. amphibia and Collemopsidium halodytes
in background. d. V. maura (Photos by Anthony Fletcher).

a b

Fig. 32. Verrucaria maura. Habitat on shore in South Fig. 33. Ascospores of: a. Verrucaria allantoidea. b. V.
Wales. halochlora. Bars a-b = 5 μm.

67
surrounded by a hyphal sheath which stains Blumeria. However, it shows no
with ruthenium red. Hafellner (1979) morphological affinities with any of these,
suggested that the ascus in D. haliotrepha was with the exception of Blumeria, both having
one layered, but the study of Au et al. (1996) cleistothecial ascomata and a short stalk to the
confirms that it is bitunicate. ascus. The latter is not characteristic of the
Dactylospora canariensis was originally Plectomycete family Eurotiaceae, where it has
referred by Kohlmeyer (1967) and Kohlmeyer previously been assigned. Amylocarpus
and Kohlmeyer (1968) to Banhegyia uralensis encephaloides, groups with Neobulgaria
and B. setispora, respectively, but on re- premnopia (Leotiaceae) with low support, in a
examination was found to be a different sister clade comprising Blumeria graminis and
species from the original collection various members of the Erysiphales
(Kohlmeyer and Volkmann-Kohlmeyer, 1998c; (Hambleton and Sigler, 2005). However, it is
Kutorga and Hawksworth, 1997). It is the only distantly placed from Leotia species (Leotiales)
Dactylospora species with appendaged and further studies are needed to resolve the
ascospores and produces antheridia in culture. final taxonomic position of Amylocarpus
(Landvik et al., 1996).
1. Ascospores with appendages.............. D. canariensis
1. Ascospores lacking appendages............................... 2
Vibrisseaceae
2. Ascospores narrow (less than 7 µm), 10.9-17.2 ×
3.5-6.4 µm, verrucose spore wall .........D. mangrovei
Vibrissea Fr., Syst. Mycol., Index alphab. 2: 4,
2. Ascospores wider than 7 µm, 18-28 × 8-12 (-14.5) 31, 1822........................................................ (1)
μm, spore wall with longitudinal striations ................ V. nypicola K.D. Hyde & Alias, Mycol. Res.
........................................................... D. haliotrepha 103: 1419, 1999.
Apothecia superficial, reddish-brown,
sessile, discoid, paraphyses filiform, septate,
LEOTIOMYCETES
apically branching, swollen at the apex, asci
LEOTIOMYCETIDAE
cylindrical, short pedunculate, in a gelatinous
HELOTIALES
matrix, ascospores fasiculate, filiform,
Helotiaceae
unicellular, hyaline with inconspicuous
Amylocarpus Curr., Proc. R. Soc. Lond., mucilage (Hyde et al., 1999b). Occurs on the
9:119-123, 1857-1859.................................. (1) petiole base of Nypa fruticans intertidally in
◙ A. encephaloides Curr., Proc. R. Soc. Lond. brackish, estuarine habitats. Hyde et al. (1999b)
9: 119, 1859 (Type species). drew attention to its similarity to Vibrissea,
Plectolitus acanthosporum Kohlm., Nova Hedw. sections Apostemium and Microstemium,
2: 329, 1960. which are difficult to distinguish at the
Ascomata solitary or gregarious, cleisto- morphological level (Iturriaga, 1997).
thecial, globose or subglobose, erumpent to
superficial, coriaceous, variously coloured:
Dermateaceae
cream-yellow, yellow or reddish yellow,
paraphyses absent, asci broadly clavate or
Laetinaevia Nannf., Nova Acta R. Soc. Scient.
ellipsoidal, apiculate, pedunculate, unitunicate,
Upsal., ser. 4, 8: 190, 1932 .......................... (1)
thin-walled, without an apical apparatus, and
L. marina (Boyd) Spooner, Kew Bull. 38: 568,
deliquescing early, ascospores hyaline,
1984.
subglobose to ovoidal, unicellular, with 10-25 Orbilia marina Boyd, Trans. Br. Mycol. Soc. 3:
awl-shaped appendages distributed over the 116, 1908 (1909).
ascospore surface. (Calloria marina Phillips, in Smith (1908),
A genus of uncertain taxonomic status unpublished manuscript).
despite a molecular study by Landvik et al. Apothecia concave, becoming convex
(1996) who report it clustering with the and discoid, erumpent, superficial, sessile,
Cyttariales, Leotiales, Rhytismatales, light orange, becoming darker, solitary or
Thelebolus and the erysiphalean genus gregarious, paraphyses filamentous, branched,

68
 Fungal Diversity

septate with swollen tips, asci cylindrical- containing many species of photobiont
clavate, tapering at the base, unitunicate, thin- including Chlorophyceae and principally
walled, with an apical ring, ascospores Calothrix (Fletcher, pers. comm.) (Fig. 35).
ellipsoidal, 1-septate, smooth-walled, hyaline, However, at least nine names are marine,
lacking a sheath or appendages. Hosts usually mostly Antarctic, S. America. Only the above
cast brown seaweeds in the drift zone and two species are known from the Northern
strictly not an obligate marine species. Which Hemisphere (Fletcher, pers. comm.).
raises the question of when are fungi truly
marine? In this instance when are the 1. Lobes flatted, 1cm long, often prostrate, richly
seaweeds colonised by the fungus? Some 25 branched in one plane, shiny dark brown to black,
apothecia terminal, globose, ascospores uniseriate....
Laetinaevia species are listed in Index ................................................................ L. pygmaea
Fungorum of which L. marina is the only 1. Thallus terete, erect, tufted, 5 mm high, lobes dull,
marine fungus. olive-brown to black .................................L. confinis

ARTHONIOMYCETES
LICHINOMYCETES ARTHONIALES
LICHINALES Roccellaceae
Lichinaceae
Halographis Kohlm. & Volkm.-Kohlm., Can.
Lichina C. Agardh, Syn. Alg. Scand. Xii, 9, J. Bot. 66: 1138, 1988 .................................. (1)
1817.............................................................. (2) H. runica Kohlm. & Volkm.-Kohlm., Can. J.
L. confinis (O.F. Müll.) C. Agardh, Spec. alg. 1: Bot. 66: 1138, 1988 (Type species).
105, 1821. Ascomata lirelliform, simple or branched,
Clathroporina confinis Műll. Ahg., Englers
Bot.Jarb. 6: 403, 1885.
immersed in calcareous substrata, opening
Fucus pygmaeus f. minor Turner with a longitudinal slit, light brown, no
Lichen confinis O.F. Műll., Icon. Plant. Daniae. 5: periphyses, single or gregarious, paraphysoid,
5, 1782. anastomosing, septate, in a gelatinous matrix,
Lichina pumila sensu Gray A natural asci clavate, short pedunculate, thick-walled,
arrangement of British plants 1: 1-824, 1821.
Lichina pygmaea var minor (Turner) Hook.
with an ocular chamber, ascospores ellipsoidal
Neolichina confinis (Műll. Ahg.) Gyein., In Ann. to fusiform, 1-septate, not constricted at the
Mus. Nat. Huhgar. 32: 166, 1939. septum, smooth, hyaline, and lacking a sheath
Pygmaea confinis (O.F. Műll.) Kuntze, or appendages (Kohlmeyer and Volkmann-
Revis.gen.pl. (Leipzig, 2, 1891. Kohlmeyer, 1988b). A lichenoid species
Stereocaulon confine (O.F. Műll.) Hoffm.
Dutschl. Flora, p130, 1796.
known from Belize, Caribbean and the Great
L. pygmaea (Lightf.) C. Agardh, Flora Scotia Barrier Reef, Australia found on the lower side
2: 964, 1777. of subtidal coral slabs, on worm tubes and on
Fucus pygmaea Lightf., Flora Scotica 2: molluscan shells. Originally placed in the
964,1777. Opegraphales by Kohlmeyer and Volkmann-
Thallus fruticose, erect, tufted, in clumps, Kohlmeyer (1988b) it is referred here to the
becoming terete near the apices, 10 mm tall, Arthoniales (Kirk et al., 2001). However,
and 0.1-0.2 mm thick, shiny dark brown to Lumbsch and Huhndorf (2007) question this
black or dark olive-green, gelatinous when wet, assignment.
apothecia terminal, globose or flask-shaped,
photobiont Calothrix (Cyanophyceae) (From ARTHONIOMYCETIDAE family incertae
http://floraseries.landcareresearch.co.nz). A sedis
genus of some 13 species of which two are Melaspileaceae
marine. An important distinction between
these two species is the presence of a cortex in Melaspilea Nyl., Act. Soc. Linn. Bordeaux, sér.
L. pygmaea which makes it cartilaginous, A 21, 416, 1857............................................ (1)
while L. confinis has a loose hyphal weft

69
Fig. 34. Dactylospora haliotrepha. a. Apothecia on mangrove wood. b-d. Asci, pseudoparaphyses and ascospores. d.
Tips of the pseudoparaphyses staining with melzer. e. Dactylospora mangrovei SEM micrograph of ascospore with
corrugated surface. Bars a = 500 µm; b-d = 10 µm; e = 5µm

M. mangrovei Vrijmoed, K.D. Hyde & E.B.G. or “skirt” at the septum to which it is attached
Jones, Mycol. Res. 100: 293, 1996. (Vrijmoed et al., 1996). It is found commonly
Ascomata lirelliform, coriaceous, on mangrove wood and can be confused with
erumpent, dark coloured with an opening that Massarina species with its 1-septate, hyaline
runs the length of the ascoma, solitary or ascospores, surrounded by a mucilaginous
gregarious, pseudoparaphyses branched, sheath. However, in M. mangrovei, the sheath
anastomosing and in a gelatinous matrix, asci ruptures apically to form a skirt-like
clavate, thick-walled, with an ocular chamber, appendage around the spore. Currently some
wall staining blue in Melzer’s reagent, 132 species are assigned to the genus, and
ascospores 1-septate, ellipsoidal, hyaline, includes lichenized, lichenicolous and saprobic
becoming light brown, constricted at the taxa (Coppins, 1989). The genus is in need of
septum, with a mucilaginous sheath (Fig. 36). revision and placement of this species remains
The sheath ruptures at the apex to form a band unresolved.

70
 Fungal Diversity

a b

c d

Fig. 35. Habitat of a-b. Lichina confinis. c-d. L. pygmaea (Photos by Anthony Fletcher).

a b

c d

Fig. 36. Melaspilea mangrovei. a. Lirelliform ascomata on mangrove wood. b, d. Ascospores 1-septate markedly
constricted at the septum with a skirt-like equatorial appendage. c. Thick-walled ascus and pseudoparaphyses. Bars a =
100 µm; b, d = 5 µm; c = 10 µm.

71
 SORDARIOMYCETES 2. Ascospores 1-septate ........................... Etheirophora
2. Ascospores 3-septate . ......................... Torpedospora
Three subclasses with marine taxa
After Zhang et al. (2006). 3. Ascospores unicellular ............................................. 4
3. Ascospores septate ................................................... 6
Perithecial or derived cleistothecial
ascomata, unitunicate asci, basal or peripheral 4. Ascospores needle-shaped...................... Halonectria
in ascoma with a wide range of anamorphs. 4. Ascospores oval or globose ..................................... 5

1. Stromata well developed, mostly consisting only 5. Ascospores 6-11 × 4-7 µm ..................Payosphaeria
fungal tissue, black with thick wall, ostioles papillate, 5. Ascospores 13.5-17 × 10-11.5 µm ... Neocosmospora
periphysate, ascomata perithecial, interascal tissue
well developed, asci cylindrical with J+ apical
6. Ascospores 1-septate ................................................ 7
apparatus, ascospores brown to black, with germ
6. Ascospores 1-3-septate ............................................ 9
pores............................................. Xylariomycetidae
1. Ascomata in a pseudostroma or coloured stroma or
absent, lacking germ pores....................................... 2 7. On seaweed (Laminaria), ascospores, pale brown
verruculose, 13-20 × 7-9 µm .....................Pronectria
2. Ascoma perithecial, rarely cleistothecial, sometimes 7. On wood, ascospores hyaline. .................................. 8
stromatic and coloured, ostiole weakly to well
developed, interascal tissue apical paraphyses or 8. Ascospores 18-21 × 10-13 µm, Trichothecium-like
catenophyses or absent, asci thin-walled often anamorph, ascomata yellow to pale brown
deliquescing, ascus apical apparatus poorly ..............................................................Heleococcum
developed, generally J-, ascospores septate, variable 8. Ascospores 17-26 × 8-13 µm, no known anamorph,
morphology, hyaline to brown, anamorphs may be ascomata orange .....................................Kallichroma
present ..................................... Hypocreomycetidae
2. Ascomata perithecial rarely cleistothecial, rarely 9. Ascomata immersed in senescent leaves of Juncus
stromatic or in a pseudostroma, necks well developed, roemerianus, asci with an apical ring, ascospores 3-
interascal tissue poorly developed or absent, asci septate, fusiform to elongate ellipsoidal 26.5-34.5 ×
cylindrical or clavate often thick-walled but not 6-7 µm....................................................... Juncigena
fissitunicate, ascospores 0-1-septate, varied 9. Ascomata in wood, asci apically thickened,
anamorphs ..................................Sordariomycetidae ascospores 1-3-septate ...................... Swampomyces
HYPOCREOMYCETIDAE
Three orders with marine taxa Bionectriaceae

1. Ascomata in a stroma, perithecial, papillate or short The assignment of marine Nectria-like

necks, generally coloured, asci clavate to cylindrical, taxa to this family is debatable and sequences
ascospores 1-septate, hyaline to pale brown, of other genera are required before their true
prominent anamorphs ........................... Hypocreales
1. Ascomata rarely in a stroma..................................... 2 placement can be made. With the exception of
Emericellopsis, they lack anamorphs, while the
2. Ascomata rarely stromatic, with well developed often submerged ascomata with long necks are
necks, asci generally clavate, ascospores hyaline not typical of the Hypocreales, e.g. Halonectria.
rarely coloured, 1-many-septate, rarely with
anamorphs ...................................... Halosphaeriales
2. Ascomata dark, thick-walled, opening by an irregular Emericellopsis J.F.H. Beyma, Antonie van
lysigenous pore, asci clavate, long peduncle, Leeuwenhoek Ned. Tijdschr. Hyg. 6: 264, 1940.
ascospores hyaline to brown, allantoid ...................................................................... (2)
........................................................Coronophorales ◙ E. maritima Beliakova, Mikol. Fitopatol. 4:
530, 1970.
HYPOCREALES ◙ E. stolkiae D.E. Davidson & M. Chr., Trans.
Br. Mycol. Soc. 57: 385, 1971.
Key to the marine Hypocreales and
Hypocreales incertae sedis 1. Ascospores 5-9 × 3-4.2 µm, 3 triangular wing-like
1. Ascospores with wing-like appendages ..................... appendages with an attenuated tips, projecting 5.6-10
........................................................... Emericellopsis µm.............................................................E. stolkiae
1. Ascospores with polar appendages .......................... 2 1. Ascospores 4-5 × 7-8 µm, 2-3 wings, up to 10 μm
1. Ascospores without appendages............................... 3 long ........................................................ E. maritima

72
 Fungal Diversity

Anamorph: Acremonium species. Rossman et al. (1999) for the present as a

Ascomata cleistothecial, globose, member of the Bionectriaceae, but they point
glaborous, superficial, asci scattered, globose out that the immersed ascomata with long
to subglobose, thin-walled, hyaline, moderately necks and the elongate aseptate ascospores are
persistent, ascospores ellipsoidal to oval not typical of the Hypocreales. Sequence data
unicellular, dark green to pale brown, are required to resolve its taxonomic position.
surrounded by subhyaline wings, triangular
with an attenuated tip, finely spinulose Heleococcum C.A. Jørg., Bot. Tidsskr. 37: 417,
(Davidson and Christensen, 1971). 1922.............................................................. (1)
Emericellopsis species have been reported ◙ H. japonense Tubaki, Trans. Mycol. Soc.
from marine habitats (Udea, 1980, 1995a) but Jpn. 8: 5, 1967.
are generally regarded as facultatively marine. Anamorph: Trichothecium-like
This aspect requires to be challenged, as they Ascomata gregarious, globose,
are isolated from marine sediments, and often superficial, no ostiole, cleistothecial,
ruled out as truly marine. membranous, white becoming orange or pale
This genus is assigned to the brown, lacking paraphyses, asci globose or
Bionectriaceae, forming a third marine lineage subglobose, sessile, unitunicate, thin-walled,
within the family (Rossman et al., 2001). Udea deliquescing, ascospores broad ellipsoidal or
(1995a) isolated E. microspora from marine ovoidal, 1-septate, not constricted at the septum,
sediments and found that optimum growth was hyaline, lacking appendages or a sheath.
in 80% seawater. Artemczuk (1980) lists Molecular studies confirm the assignment of
Emericellopsis maritima (Fig. 37) from the genus to the Bionectriaceae, with affinities
sediments in the Black Sea, with ascospore to Roumegueriella rufula, another
measurements of 4-5 × 7-8 µm. Acremonium cleistothecial member of the Bionectriaceae
species are also frequently encountered on (Rehner and Samuels, 1995).
incubated wood from marine habitats (Jones,
unpublished data). Kallichroma Kohlm. & Volkm.-Kohlm.,
Mycol. Res. 97: 759, 1993 ........................... (2)
a b ◙ K. glabrum (Kohlm.) Kohlm. & Volkm.-
Kohlm., Mycol. Res. 97: 759, 1993.
Hydronectria tethys var. glabra Kohlm., Mar.
Ecol. (P.S.Z.N.I.) 5: 351, 1984.
◙ K. tethys (Kohlm. & E. Kohlm.) Kohlm. &
Volkm-Kohlm., Mycol. Res. 97: 759, 1993
(Type species).
Hydronectria tethys Kohlm. & E. Kohlm., Nova
Fig. 37. a. Emericellopsis maritima Ascospore. b. Hedw. 9: 95, 1965.
Conidium. Bars a-b = 5 μm. Ascomata solitary or gregarious,
subglobose, immersed sometimes erumpent,
Halonectria E.B.G. Jones, Trans. Br. Mycol. ostiolate, periphysate, lacking a papilla,
Soc. 48: 287, 1965 ....................................... (1) orange-brownish to orange-yellowish, peridium
H. milfordensis E.B.G. Jones, Trans. Br. thick, lacking paraphyses, asci clavate,
Mycol. Soc. 48: 287, 1965 (Type species). unitunicate, thin-walled, deliquescing,
Ascomata solitary or gregarious, globose ascospores ellipsoidal, fusiform or ovoid, 1-
or subglobose, usually immersed, ostiolate, septate, constricted at the septum, hyaline
paplliate, coriaceous, orange-coloured to pale lacking appendages or a sheath. Originally
brown, lacking paraphyses, asci clavate, short described as Hydronectria but transferred by
pedunculate, unitunicate, thin-walled, Kohlmeyer and Volkmann-Kohlmeyer (1993a)
deliquescing early, ascospores elongate, to Kallichroma because the type species of
fusiform or cylindrical, unicellular, and hyaline. Hydronectria is a lichen, with the alga
No appendages or sheath. A genus accepted by Trentepohlia, occurs on rocks in freshwater

73
and is temperate in its distribution. hyaline, asci long cylindrical, short
Kallichroma species are saprobic, marine pedunculate, persistent, unitunicate, thin-
(primarily on mangrove wood) and sub-tropical walled, no apical apparatus, ascospores round
to tropical. They also differ in ascoma, to oval, unicellular, hyaline, thin-walled,
paraphyses, ascus and ascospore morphology smooth and lacking a sheath or appendages. A
from Hydronectria. Kohlmeyer and Volkmann- poorly known species described from
Kohlmeyer (1993a) state the asci are persistent, mangrove wood samples from Malaysia and
but we frequently observe asci deliquescing. Singapore, but common on submerged test
Molecular data supports placement in the blocks at these locations with 61 collections
Bionectriaceae, but is the most distant genus in (Leong et al., 1990). Tentatively assigned by
the family (Rossman et al., 2001; Schroers, Leong et al. (1990) to the Hypocreales (as
2001). SEM studies show longitudinal ridges Nectriales) its taxonomic position needs further
running the length of the ascospores of K. study at the molecular level.
tethys (Hyde, 1986).
a
1. Ascospore wall smooth ........................... K. glabrum
1. Ascospore wall with longitudinal ridges......K. tethys

Hypocreaceae

Neocosmospora E.F. Sm., U.S.D.A. Div. Veg.

Pathol. Bull. 17: 45, 1899 ............................ (1)
N. tenuicristata S. Udea & Udagawa,
Mycotaxon 16: 387, 1983. b
Anamorph: Acremonium tenuicristatum S.
Udea & Udagawa.
Ascomata ovoid to pyriform, superficial,
scattered or aggregated, pale coloured
becoming pink or orange-red, glaborous with
hyaline to pale-yellow, unbranched, septate,
smooth-walled short, hyphal-like hairs, short
Fig. 38. Neocosmospora tenuicristata. a. Ascospore. b.
neck, periphysate, asci cylindrical, short Conidium. Bars a-b = 5 μm.
pedunculate, hyaline, no apical apparatus,
ascospores broadly ellipsoid to ellipsoid, Pronectria Clem., Gen. Fungi. 78, 282, 1931 ..
unicellular, yellowish brown, thick-walled, no ...................................................................... (1)
germ pore, surface ornamented (Fig. 38). The P. laminariae (O.E. Erikss.) Lowen,
only species known from marine habitats, Mycotaxon 39: 461, 1990.
isolated from marine sludge at Oomura Bay, Nectriella laminariae O.E. Erikss., Svensk. Bot.
Japan (Udea and Udagawa, 1983). Tidskr. 58: 233, 1964.
Characterized by its striated ascospores and a Ascomata solitary or gregarious, globose,
polyphialidic anamorph. ostiolate, papillate, periphysate, immersed,
light brown, paraphyses filamentous
Payosphaeria W.F. Leong, Bot. Mar. 33: 511, deliquescing, asci cylindrical, thin-walled,
1990...............................................................................................(1) unitunicate, with an apical pore, ascospores
P. minuta W.F. Leong, Bot. Mar. 33: 511, ellipsoidal, ovoid, fusiform, 1-septate, lacking
1990 (Type species). appendages or a sheath and hyaline. Initially
Ascomata globose to pyriform, described as a Nectriella species but transferred
superficial, ostiolate, papillate, lacking to this genus by Lowen (1990) and is an
periphyses, membranous, hyaline, solitary or accepted genus in the Bionectriaceae (Rossman
gregarious, paraphyses few, branched, septate, et al., 1999).

74
 Fungal Diversity

HYPOCREALES incertae sedis Anamorph: Cirrenalia adarca Kohlm,

Volkm-Kohlm & O.E. Erikss.
Torpedospora Meyers, Mycologia 49: 496, Ascomata subglobose to pyriform,
1957.............................................................. (2) immersed, ostiolate, papillate, coriaceous,
◙ T. ambispinosa Kohlm., Nova Hedw. 2: 336, fuscous, solitary, periphysate, pseudo-
1960. paraphyses thin, branched, septate, asci
◙ T. radiata Meyers, Mycologia 49: 496, 1957 fusiform to cylindrical, short pedunculate, thin-
(Type species). walled, unitunicate, apical apparatus with an
Ascomata solitary, subglobose to apical ring, J-, ascospores fusiform to elongate-
ellipsoidal, immersed or superficial, ostiolate, ellipsoidal, 3-septate, constricted at the septa,
papillate or epapillate, subcarbonaceous to hyaline, no sheath or appendages. A salt marsh
coriaceous, dark brown, paraphyses ramose, fungus, with a Cirrenalia adarca anamorph,
deliquescing or persistent, asci clavate to which grows on the submerged bases of leaves
ellipsoidal, unitunicate, thin-walled, early (between 12-25 cm above the rhizomes) of
deliquescing, ascospores cylindrical to Juncus roemerianus, and thus regarded as
elongate-ellipsoidal, 3-septate, constricted at obligately marine. Originally Eriksson (1999)
the septum, hyaline but pale orange in a mass, considered it to belong in the Magna-
with appendages at one or both poles (Fig. 39) porthaceae. Thongkantha et al. (2009) found no
(Sakayaroj et al., 2005b). support for this. Using DNA sequences from
Although these species have appendaged protein coding and ribosomal nuclear loci,
ascospores and deliquescing asci, and Schoch et al. (2006) noted three subclades (1.
tentatively assigned to the Halosphaeriales, Torpedospora spp., 2. Swampomyces spp. and
they were later excluded from the order Etheirophora spp. and 3. Swampomyces sp.
(Kohlmeyer, 1972a). The morphology of the and Juncigena adarca) were associated with
ascomata is very different from those of the the Coronophorales with good support. The
Halosphaeriaceae and molecular data show data suggests that the 3-septate Swampomyces
they are a sister group to the Bionectriaceae, species may be congeneric with Juncigena
Hypocreales from LSU rDNA, or the clades adarca but further clarification of the
comprising the Phyllachorales, Halos- molecular data of S. triseptatus is required.
phaeriales, Microascales from SSU rDNA and
Swampomyces Kohlm. & Volkm.-Kohlm., Bot.
combined data set (Sakayaroj et al., 2005b). At
Mar. 30: 198, 1987....................................... (4)
the molecular level the genus clusters with
◙ S. armeniacus Kohlm. & Volkm.-Kohlm.,
Swampomyces (Sakayaroj et al., 2005b) and
Bot. Mar. 30: 200, 1987 (Type species).
this has been verified by Schoch et al. (2006),
◙ S. triseptatus K.D. Hyde & Nakagiri,
and confirms that they form a sister group to
Sydowia 44: 122, 1992.
the Hypocreales in the Hypocreomycetidae.
◙ S. aegyptiacus Abdel-Wahab, El-Shar. &
Torpedospora and Swampomyces also group
E.B.G. Jones, Fungal Diver. 8: 35, 2001.
with the genera Juncigena and Etheirophora,
◙ S. clavatispora Abdel-Wahab, El-Shar. &
but share few morphological characters
E.B.G. Jones, Fungal Diver. 8: 37, 2001.
(Schoch et al., 2006).
Ascomata pyriform, subglobose, globose,
coriaceous, centrum apricot coloured, solitary,
1. Ascospores with appendages at one end, longer than
30 µm .........................................................T. radiata
immersed, ostiolate, necks with periphyses,
1. Ascospores with appendages at both ends, shorter dark brown to black, paraphyses numerous,
than 25 µm ....................................... T. ambispinosa simple, hyaline, in a gel, asci cylindrical-
oblong, unitunicate, thin-walled, short
Juncigena Kohlm., Volkm.-Kohlm. & O.E. pedunculate, apically thickened, J-, persistent,
Erikss., Bot. Mar. 40: 291, 1997 .................. (1) ascospores clavate, ellipsoidal, 1-3-septate,
◙ J. adarca Kohlm., Volkm.-Kohlm. & O.E. hyaline, slightly to constricted at the septa, no
Erikss., Bot. Mar. 40: 291, 1997 (Type species). appendages or sheaths. All species found
predominantly on mangrove wood (Fig. 40)
75
 a b c

Fig. 39. Ascospores of a-b. Torpedospora radiata. c. Torpedospora ambispinosa. Bars a-b = 10 µm, c = 5 µm.

a b
(Abdel-Wahab et al., 2001a). The genus was
tentatively assigned to the Polystigmataceae
(Kohlmeyer and Volkmann-Kohlmeyer, 1987c),
but was left unclassified pending sequence data.
At the ultrastructure level, the ascus apex
consists of a large amorphous apical thickening,
but no central pore was observed, although
serial sections were made (Read et al., 1995).
Molecular data has not helped in resolving the
higher level taxonomic position of this genus, Fig. 40. Ascospores of a. Swampomyces clavatispora. b.
which groups with Torpedospora species S. aegyptiacus. Bars a-b = 5 μm.
(Sakayaroj et al., 2005b). Swampomyces
armeniacus shows closer affinity with ◙ E. unijubata Kohlm. & Volkm.-Kohlm.,
Etheirophora species, but further strains of S. Mycol. Res. 92: 415, 1989.
triseptatus need to be sequenced to resolve the Ascomata subglobose, elongate,
status of the genus (Schoch et al., 2006). immersed in wood, ostiolate, papillate, clypeate,
coriaceous, light-coloured, peri-physate,
1. Ascospores 1-septate, 13-20 × 6.9 µm S. armeniacus paraphyses septate, rarely branched in a
1. Ascospores with more than 1 septum....................... 2 gelatinous matrix, asci cylindrical to oblong,
pedunculate, J-, thin-walled, persistent, no
2. Ascospores clavate, 25-28 × 5-6 µm.. S. clavatispora
2. Ascospores ellipsoidal.............................................. 3 apical apparatus, ascospores ellipsoidal, 1-
septate, hyaline, with variable number of
3. Ascospores 15-19 × 6-8 µm, deeply constricted at the appendages. Kohlmeyer and Volkmann-
septa ....................................................S. aegyptiacus Kohlmeyer (1989) erected the genus for a
3. Ascospores 18-25 × 8-11 µm, weakly constricted at group of lignicolous marine fungi of uncertain
the septum, spore wall with granular ornamentation..
............................................................. S. triseptatus
taxonomic position and incorrectly assigned to
the Halosphaeriales by Hawksworth et al.
Etheirophora Kohlm. & Volkm.-Kohlm., (1995) and Kirk et al. (2001). Although
Mycol. Res. 92: 414, 1989 ........................... (3) Etheirophora species share a number of
E. bijubata Kohlm. & Volkm.-Kohlm., Mycol. features in common with the Halosphaeriales
Res. 92: 414, 1989 (Type species). (lignicolous, immersed ascomata, ostioles
◙ E. blepharospora (Kohlm. & E. Kohlm.) periphysate, asci unitunicate, with 1-septate
Kohlm. & Volkm.-Kohlm., Mycol. Res. 92: hyaline and appendaged ascospores) they differ
415, 1989. in having cylindrical, pedunculate, non
Keissleriella blepharospora Kohlm. & E. Kohlm., amyloid persistent asci with polar filamentous
Nova Hedw. 9: 97,1965. appendaged ascospores. Appendages are long

76
 Fungal Diversity

(12-18 µm), bristle-like, rigid, slightly curved HALOSPHAERIALES

and of undetermined origin. Molecular data
places them in the TBM clade with affinties to Halosphaeriaceae
the Coronophorales in the Hypocreomycetidae
(Schoch et al., 2006). The Halosphaeriales is one of the most
intensively studied marine ascomycete order at
1. Ascospores with appendages at both ends, 16-21.5 × the morphological, ultrastructural and
6-8 µm ..................................................... E. bijubata molecular level with 53 genera (of which 35
1. Ascospores with appendages at one end .................. 2 are monotypic) and 126 species. Nearly 50% of
the genera have been sequenced and found to
2. Ascospores up to 21 µm, on bark of Rhizophora form a monophyletic group within the
mangle, up to 7 appendages ...........E. blepharospora
2. Ascospores up to 29 µm, on other hosts, with more
Ascomycota (Fig. 41). Hibbett et al. (2007)
than 7 appendages ................................. E. unijubata places the Halosphaeriales within the
Microascales. However, Zhang et al. (2006)
CORONOPHORALES and Tang et al. (2007a) retain the order, and
this is followed in this volume. Tang et al.
Nitschkiaceae (2007a) undertook a multigene analysis of the
Groenhiella Jørg. Koch, E.B.G. Jones & S.T systematics of the Sordariomycetes. Three
Moss, Bot. Mar. 26: 265, 1983 .................... (1) subclasses were defined: Hypocreomycetidae,
G. bivestia Jørg. Koch, E.B.G. Jones & S.T. Sordariomycetidae and Xylariomycetidae as
Moss, Bot. Mar. 26: 265, 1983 (Type species). employed in this monograph. They noted that
Ascomata hemispherical-oblong tuberi- the Microascales is paraphyletic with
form, cleistothecial, superficial, leathery to Ceratocystis phylogenetically associated with
carbonaceous, brown to black, with a weak Faurelina, while Microascus and Petriella
subiculum, paraphyses numerous, septate, formed a separate clade and were basal to other
constricted at the septa, simple or branched, members of the Halosphaeriales (Tang et al.,
asci broadly clavate, long pedunculate, 2007a). In the LSU dataset the Halosphaeriales
unitunicate thin-walled at maturity, no apical and Microascales form subclades, with the
apparatus, generally persistent, ascospores latter in a basal position. In the SSU dataset the
broadly fusiform, 1-septate, slightly Microascales splits the Halosphaeriales into
constricted at the septum, hyaline later brown, two separate clades, but with weak support.
with appendages. Appendages formed by The Microascales are basal to the
fragmentation of a sheath forming apical and Halosphaeriales in the RPB2 and combined
equatorial appendages (Koch et al., 1983). datasets, but with too few taxa to satisfactorily
Hibbett et al. (2007) refer the resolve their taxonomic position (Tang et al.,
Nitschkiaceae to the Coronophorales, the order 2007a).
to which Koch et al. (1983) originally assigned
Key to the genera
Groenhiella. However, a molecular study is
required to validate its assignment to the 1. Ascospores aseptate ................................................. 2
Nitschkiaceae and to the Coronophorales. The 1. Ascospores septate ................................................... 8
most recent study is that of Petersen (1997) on
the ultrastructure of the ascospores and 2. Ascospores appendaged ........................................... 3
2. Ascospores lacking appendages............................... 4
confirms the exosporic origin of the
appendages, comprising fibrillar electron-dense 3. Ascospores with polar and 4 groups of equatorial
material in an electron-transparent matrix and hair-like appendages ..........................Nautosphaeria
circa 360 nm thick. As the ascus deliquesces 3. Ascospores with a single, uncoiling polar appendage
the exosporic sheath separates from the ....................................................................... Moana
episporium and then fragments to form the 4. Ascospores longer than 50 µm, filiform, range 50-
polar and equatorial appendages. 300 × 4-15 µm ........................................ Bathyascus

77
Fig. 41. One of three MPTs inferred from LSU rDNA sequences of all halosphaerialean taxa, generated with
maximum parsimony analysis. Bootstrap values higher than 50% are given above branches. Scale bar indicates 10
character state changes.

78
 Fungal Diversity

4. Ascospores shorter than 50 µm, spherical or 19. Polar appendages emerging from a hood-like
ellipsoidal................................................................. 5 structure......................................... Cucullosporella
19. Polar appendages not formed through a hood. .... 20
5. Ascospores spherical to round ................................. 6
5. Ascospores ellipsoidal, range 9-38 × 4-24 µm........... 20. Ascospores longer than 35 µm and wider than 20
..............................................................Chadefaudia µm ...................................................................... 22
20. Ascospores shorter and narrower than 35 µm and
6. Asci with an apical pore, ascopores globose to 20 µm, respectively ............................................. 21
subglobose........................................... Thalassogena
6. Asci lacking an apical pore ...................................... 7 21. Ascospores wider than 14 µm ..........Saagaromyces
21. Ascospores 12-14 µm wide...................Aniptodera
7. Ascomata cream-coloured, with long necks,
ascospores globose to ellipsoidal ..........Anisostagma 22. Ascospores wider than 30 µm .................................
7. Ascomata brown, necks short, ascospores spherical .. ...................................... Halsarpheia sensu stricto
................................................................Iwilsoniella 22. Ascospores narrower than 30 µm........................ 23
8. Ascospores 1-septate................................................ 9 23. Ascospores 21-31 × 8-11 µm .................. Panorbis
8. Ascospores 1 to many-septate ............................... 44
23. Ascospores 20-34 × 7-11 µm ............Natantispora
9. Ascospores with no appendages ............................ 10 23. Ascospores 10- 32 × 4-13µm ..................................
9. Ascospores with polar and/or equatorial appendages ........................................ Halosarpheia sensu lato
or with sheaths ...................................................... 14
24. Ascospores with two types of appendages .......... 25
10. Asci deliquescing early .................................. Nais 24. Ascospores with only one type of appendage ..... 28
10. Asci persistent. .................................................... 11
25. Ascospores with a sheath and polar and lateral or
11. Asci persistent, no retraction of the plasmalemma.. subpolar appendages ........................................... 26
............................................................................. 12 25. Ascospores with no sheath .................................. 27
11. Asci with plasmalemma retracted ....................... 13
26. Ascospores with polar hair-like and lateral sheath-
12. Ascus tip thimble-shaped, or slightly thickened...... like appendage .................................... Nimbospora
............................................................... Lignincola 26. Ascospores with a fragmenting sheath, and polar
12. Ascus tip lacking thimble-shaped apical thickening unfurling appendages ....................... Tunicatispora
.......................................................................Alisea
27. Polar strap-like mucilaginous and subpolar hair-
13. Ascospores without unfurling bipolar appendages.. like appendages ................................... Naufragella
...............................................................Aniptodera 27. Sub-polar hair-like appendages arise from a pad,
13. Ascospores with a faint sheath .............. Neptunella and on the opposite side long, sticky appendages
that uncoil in water ...................................... Nohea
14. Ascospores with a single polar appendage.......... 15
14. Ascospores with bipolar hamate appendage, or 28. Ascospores with polar or subpolar appendages... 29
polar and equatorial appendages ........................ 18 28. Ascospores with polar and/or lateral appendages 35
28. Ascospores with an exosporic sheath.................. 41
15. Ascospores with a hamate polar appendage........ 16
15. Ascospores with an ephemeral drop of polar 29. Ascospores with polar appendages ..................... 30
mucilage, becoming 2-4-septate on germination..... 29. Ascospores with subpolar appendages ................ 32
.......................................................... Okeanomyces
30. Ascospore appendages formed by fragmentation of
16. Ascomata formed beneath a stroma, ascospores 6- a sheath................................................................ 31
21× 6-8 µm .......................................... Ophiodeira 30. Appendages coiled around the ascospores,
16. Ascomata not stromatic....................................... 17 uncoiling in water............................... Morakotiella

17. Ascospores oval, 24-32 × 8-12 µm ..........Tirispora 31. Ascospore wing-like or radiating appendages.........
17. Ascospores filiform, 60-80 × 4-6 µm ascomata ......................................... Remispora sensu stricto
thick-walled .............................................Oceanitis 31. Ascospores appendages moustache shaped, each
apex surrounded by a large, subglobose cap, with
18. Ascospores with polar unfurling appendages ....... 9 delicate radiating striae, (16-) 20-28 (-35) × 7-12
18. Ascospore appendages with a different µm ............................................ Remispora galerita
morphology ......................................................... 24 31. Ascospores appendages spoon-shaped, delicate
and fibrillar, 22-34 × 8-12 µm.....Remispora crispa

79
32. Ascospore appendages with a spade-like tip ........... .......................................................Pseudolignincola
........................................................ Arenariomyces 47. Ascospores hyaline or brown with polar and
32 Ascospores appendages lacking a spade-like tip. 33 equatorial appendages ......................................... 48
47. Ascospores hyaline, with only polar appendages....
33. Ascospores with 2 sub-polar appendages................ ............................................................................ 50
..........................................................Antennospora 47. Ascospores with sheath, lacking polar and
33. Ascospores with more than 2 sub-polar equatorial appendages ......................................... 56
appendages ......................................................... 34
48. Appendages hair-like tufts, one polar and four
34. Ascospores with 3-4 sub-polar spoon-shaped equatorial............................................................. 49
appendages ..............................................Haiyanga 48. Polar appendages spine-like, equatorial
34. Ascospores with 5-7 appendages .........Corallicola appendages formed by fragmentation of an
exosporic sheath.................................Corollospora
35. Equatorial appendage ring- or annulus-like ........ 36
35. Equatorial appendages distinct ........................... 38 49. Catenophyses present, ascospores hyaline,
appendages string-like, lacking an equatorial pad...
36. Chamber-like polar appendage from which ............................................................... Havispora
mucilage is released ...... Ceriosporopsis tubulifera 49. Catenophyses lacking, ascospores with hyaline end
36. Polar appendages do not release mucilage .......... 37 cells and brown central cells, equatorial pad
present ................................................. Nereiospora
37. Equatorial appendage annulus-like ........Ondiniella
37. Equatorial appendage ring-like ....... Lautisporopsis 50. Ascospores with a single polar appendage.......... 51
50. Ascospores with bipolar appendages ................. 52
38. Equatorial appendages lunate, with a cup-like
polar appendage .........................Halosphaeriopsis 51. Appendage an ephemeral drop of mucilage ............
38. Appendages spoon-like or obclavate.................. 39 ......................................................... Okeanomyces
51. Tetraradiate appendages formed after release from
39. Appendages spoon-like, do not fragment................ the ascoma.............................................Thalespora
...........................................................Halosphaeria
39. Appendages become fibrillar at maturity ........... 40 52. Ascospores appendages hamate, unfurling in water
............................................................................ 53
40. Polar appendage longer than equatorial 52. Ascospore appendages not hamate...................... 55
appendages .........................................Ocostaspora
40. Appendages equal in length .................... Sablecola 53. Ascospores narrower than 7 µm...............Oceanitis
53. Ascospores wider than 7 µm ............................... 54
41. Appendages with a cup-like exosporic fragments
at their tips..........................................Marinospora 54. Ascospores verrucose, wider than 40 µm................
41. Appendages lacking cup-like fragments at their ........................................................... Magnisphaera
tips...................................................................... .42 54. Ascospores not verrucose, narrower than 40 µm ....
........................................ Halosarpheia sensu lato
42. Ascomata grayish-white, exosporic sheath
55. Ascospores appendages sub-polar, spine-like .........
envelopes the spore and horn-like polar appendage
....................................................... Arenariomyces
..............................................................Bovicornua
55. Ascospores appendages broad strap-like .Haligena
42. Ascomata brown to dark coloured, appendages not
55. Ascospores appendages round ............Trichomaris
horn-like .............................................................. 43
56. Ascospores 12-20 µm wide, central cells dark,
43. Exosporic sheath highly fibrillar (mucilaginous), appendages a fragmenting sheath net-like...............
circa 5 µm wide, appendage slug-like..................... ..................................................... Carbosphaerella
............................................................Limacospora 56. Ascospores hyaline, 7-11 µm wide .........................
43. Exosporic sheath compact, not fibrillar, polar ................................................... Appendichordella
appendage uncoiling in water..........Ceriosporopsis
Alisea J. Dupont & E.B.G. Jones, Mycol. Res.
44. Ascospores with no appendages.......................... 45
44. Ascospores with appendages............................... 47 in press, 2009 ............................................... (1)
◙ A. longicola J. Dupont & E.B.G. Jones,
45. Ascospores filamentous, broad at one end, tapering Mycol. Res. in press, 2009.
at the other....................................................Trailia Ascomata globose to obpyriform,
45. Ascospores not tapering at one end..................... 46 ostiolate, papillate, coriaceous, black,
superficial, solitary or gregarious, with a long
46. Ascospores 5-septate (rarely 9), asci with 4
ascospores ................................................Luttrellia cylindrical to conical neck circa 250 µm,
46. Ascospores 3-septate, asci with 8 ascospores.......... periphysate, asci clavate, long pedunculate,
80
 Fungal Diversity

thin-walled, unitunicate, persistent, ascospores periphysate, catenophyses present but

fusiform, slightly curved, 1-septate, thick- deliquescing, asci clavate, short pedunculate,
walled, hyaline but distinct appendages not unitunicate, thin-walled, but with an apical
reported. Molecular and morphological data pore and a retracting plasmalemma subapically,
confirm the position of this genus within the persistent or deliquescing, ascospores
Halosphaeriales (Dupont et al., 2009). On ellipsoidal, 1-septate, hyaline, thick-walled,
trawled sunken wood fragments collected at with or without apical appendages which unfurl
1,000 m depth in the Pacific Ocean off when mounted in water (Hyde and Jones,
Vanuatu Islands, the ascomata are large (600- 1989c; Campbell et al., 2003).
650 µm), with a thick peridial wall, no Initially this genus was well
paraphyses or catenophyses, asci unitunicate, circumscribed (Shearer and Miller, 1977), but
clavate with a long pedicel, persistent, with the addition of further species the
ascospores 32-35 × 3.2-3.6 µm, hyaline, situation has become confused. Originally
fusiform, 1-septate and with no appendages ascospores were characterized as thick-walled,
(Fig. 42). Ultrastructural data suggest there is 1-septate, lacking bipolar appendages, with
an exosporium which may form a thin persistent asci, with a retracting plasmalemma,
mucilaginous layer around the ascospore. and an apical pore. Since the assignment of
species with bipolar, unfurling appendages, the
differences between it and Halosarpheia have
become confused (Kong et al., 2000) (Fig. 41).
Any taxonomic changes proposed for
Halosarpheia must take into account the genus
Aniptodera. Marine Aniptodera species occur
on a wide range of substrata: mangrove and
driftwood, Nypa fruticans, Juncus roemerianus.
Some freshwater Aniptodera species may also
Fig. 42. Alisea longicola. Ascospore. Bar = 10 μm.
occur in saline habitats, for example A.
Aniptodera Shearer & M.A. Mill., Mycologia limnetica was reported by Nakagiri (1993b) on
69: 893, 1977 ............................................... (9) Bruguiera gymnorhiza wood collected in the
◙ A. chesapeakensis Shearer & M.A. Mill., Shiira River mangrove, Japan. Ascospores in A.
Mycologia 69: 894, 1977 (Type species). limnetica are thin-walled and released through
A. haispora Vrijmoed, K.D. Hyde & E.B.G. a fissure in the apical plate which splits at the
Jones, Mycol. Res. 98: 701, 1994. pore. We regard Aniptodera indica as a
A. intermedia K.D. Hyde & Alias, Mycol. Res. synonym of Tirispora unicaudata (Ananda and
103: 1409, 1999. Sridhar, 2002). Assignment of A. juncicola and
◙ A. juncicola Volkm.-Kohlm. & Kohlm., Bot. A. mangrovei to the genus needs to be tested at
Mar. 37: 109, 1994. the molecular level as they may be better
A. limnetica Shearer, Mycologia 81: 140, 1989. placed in other genera.
◙ A. longispora K.D. Hyde, Bot. Mar. 33: 335, The stability of ascospore appendages as
1990. a character in the delineation of genera might
A. mangrovei K.D. Hyde, Can. J. Bot. 64: be questioned in view of the behaviour of those
2989, 1986. of Aniptodera salsuginosa when mounted in
A. nypae K.D. Hyde, Sydowia 46: 257, 1994. water of different salinity. In freshwater the
A. salsuginosa Nakagiri & Tad. Ito, Mycol. appendages immediately detach but uncoil
Res. 98: 931, 1994. when mounted in salinities of 3-10 ‰
Ascomata globose to subglobose, (Nakagiri and Ito, 1994). The issue of
immersed or superficial, ostiolate, papillate, appendaged/non-appendaged ascospores in A.
membranous, hyaline to light brown to dark chesapeakensis also needs resolution, before
brown, neck variable in length cylindrical, the taxonomy of this genus can be resolved.

81
1. Ascospores with bipolar appendages ...................... 2 Antennospora Meyers, Mycologia 49: 501,
1. Ascospores lacking appendages............................... 3 1957.............................................................. (1)
2. Ascospores 37-45 × 12-14 μm, on mangrove wood .. ◙ A. quadricornuta (Cribb & J.W. Cribb) T.W.
............................................................. A. mangrovei Johnson, J. Elisha Mitchell Sci. Soc.74: 46,
2. Ascospores 14-20 × 4-7 μm, on mangrove bark ........ 1958 (Type species).
........................................................... A. salsuginosa Halosphaeria quadricornuta Cribb & J.W. Cribb,
Univ. Queensl. Pap. Dept. Bot. 3: 99, 1956.
3. Ascospores thick-walled .......................................... 4 Antennospora caribbea Meyers, Mycologia 49:
3. Ascospores thin-walled ............................................ 6 503, 1957.
Ascomata subglobose or ellipsoidal,
4. Ascospores with very thick-walled, on wide range of
immersed, superficial when growing on the
substrata, 21-37 × 7-15 μm .......... A. chesapeakensis
4. Ascospore walls less thick ....................................... 5 calcareous tubes of mollusks, ostiolate,
papillate, coriaceous or subcarbonaceous, dark
5. Ascospores 20-25 × 14-18 μm, on mangrove wood .. brown, solitary or gregarious, catenophyses
............................................................... .A. haispora deliquescing, periphysate, asci clavate,
5. Ascospores 24-31 × 8-12 μm, on Juncus roemerianus pedunculate, untiunicate, thin-walled, no apical
............................................................... A. juncicola
apparatus, deliquescing early, ascospores
6. Ascospores longer than 35 μm, 32-54 × 9-13.5 μm, ellipsoidal, 1-septate, slightly constricted at the
on mangrove wood............................... A. longispora septum, hyaline with polar appendages.
6. Ascospores shorter than 35 μm ................................ 7 Appendages subterminal, at each end of the
7. Ascospores 10.5-13 × 7-8 μm ..............A. intermedia
spore, at right angles to each other, cylindrical,
7. Ascospores 16-22 × 5-7 μm .........................A. nypae and attenuate (Figs 41, 43h). Their ultra-
7. Ascospores 20-25 × 8-10 μm ................. A. limnetica structure has been examined by Yusoff et al.
(1994c). Ascospores of A. salina and A.
Anisostagma K.R.L. Petersen & Jørg. Koch, quadricornuta are morphologically different
Mycol. Res. 100: 211, 1996 ......................... (1) with two round, sub-polar appendages in the
A. rotundatum K.R.L. Petersen & Jørg. Koch, latter species, while the former has 4-5 sub-
Mycol. Res. 100: 211, 1996 (Type species). terminal wing-like appendages (Yusoff et al.,
Ascomata globose to broadly ellipsoidal, 1994c). Consequently, and supported by
immersed to erumpent, coriaceous, ostiolate, molecular data, A. salina was transferred to
papillate, cream-coloured to pale brown, Haiyanga (Pang et al., 2008b).
solitary or gregarious, neck long (140-420 µm),
periphysate, catenophyses present, asci clavate, Appendichordella R.G. Johnson, E.B.G. Jones
pedunculate, unitunicate, thin-walled, lacking & S.T. Moss, Can. J. Bot. 65: 941, 1987 (l)
an apical apparatus, early deliquescing, A. amicta (Kohlm.) R.G. Johnson, E.B.G.
ascospores globose to ellipsoidal, unicellular, Jones & S.T. Moss, Can. J. Bot. 65: 941, 1987
hyaline, thin-walled without appendages or a (Type species).
sheath (Petersen and Koch, 1996). Sphaerulina amicta Kohlm., Nova Hedw. 4: 414,
Anisostagma rotundatum resembles 1962.
Thalassogena sphaerica, both possessing Haligena amicta (Kohlm.) Kohlm., Marine
sphaerical-ellipsoidal hyaline ascospores and Mycology: The Higher Fungi: 288, 1979.
with no appendages or sheaths. Two Ascomata globose to subglobose,
distinguishing characters are: peridium immersed, ostiolate, papillate, coriaceous, light
structure and ascus morphology. In brown to reddish-brown, solitary, periphysate,
Thalassogena the peridial wall is catenophyses deliquescing, asci clavate,
undifferentiated, while in Anisostagma it is two pedunculate, unitunicate, thin-walled, without
layered (inner layer of flattened thin-walled an apical apparatus, deliquescing, ascospores
cells and an outer layer forming a textura cylindrical or ellipsoidal, 3-septate, constricted
angularis). An apical pore is present in at the septa, hyaline and with a pronounced
Thalassogena, but lacking in Anisostagma sheath. A monotypic genus (previously
(Petersen and Koch, 1996). Whether these assigned to Sphaerulina and Haligena)
characters are sufficient to separate these two characterized by a gelatinous sheath to the
genera remains to be resolved.
82
 Fungal Diversity

ascospore, which is composed of thread-like 2. Ascospores cylindrical, 16-25 × 3-6 μm . A. parvulus

appendages arising from the episporium 2. Ascospores ellipsoidal ............................................. 3
(Johnson et al., 1987). Well placed in the
3. Ascospores consistently with 3 appendages, mainly
Halosphaeriales.
on sand, 24-38 × 7-16 μm .................... A. trifurcatus
3. Ascospores with 3-4 appendages, mainly on wood,
Arenariomyces Höhnk, VerVöff. Inst. 28-39 × 10-14 μm ...............................A. majusculus
Meersforsch. Bremerhaven 3: 28, 1954 ....... (4)
A. majusculus Kohlm. & Volkm.-Kohlm.,
Mycol. Res. 92: 411, 1989. Bathyascus Kohlm., Rev. Mycol. 41: 190,
A. parvulus Jørg. Koch, Nordic. J. Bot. 6: 497, 1977.............................................................. (4)
1986. B. avicenniae Kohlm., Bot. Mar. 23: 530, 1980.
◙ A. trifurcatus Höhnk, Veröff. Inst. B. grandisporus K.D. Hyde, Bot. Mar. 30: 413,
Meersforsch. Bremerhaven 3: 30, 1954 (Type 1987.
species). B. tropicalis Kohlm., Bot. Mar. 23: 532, 1980.
Halosphaeria trifurcata (Höhnk) Cribb & J.W.
Cribb, Univ. Queensl. Pap., Dept. Bot. 3: 99, 1956
B. vermisporus Kohlm., Rev. Mycol. 41: 191,
Peritrichospora trifurcata (Höhnk) Kohlm., Nova 1977 (Type species).
Hedw. 3: 89, 1961 Ascomata subglobose or ellipsoidal,
Corollospora trifurcata (Höhnk) Kohlm. Ber. immersed, ostiolate, papillate, coriaceous, dark
Dtsch. Bot. Ges. 75: 126, 1962 brown, solitary, neck lacking periphyses,
A. triseptatus Kohlm., Mar. Ecol. (P.S.Z.N.I.) catenophyses deliquescing, asci fusiform to
5: 333, 1984. clavate, unitunicate, thin-walled, early
Ascomata globose or subglobose, deliquescing, ascospores filiform, 0-1-septate,
immersed generally superficial, attached to hyaline and lacking appendages. This genus
substrata by subicula, with or without ostioles, was initially assigned to the Halosphaeriales
papillate or epapillate, carbonaceous, black or because of the deliquescent nature of the asci,
dark brown, solitary, catenophyses but prelimiary molecular studies show that it
deliquescing, asci fusiform to subclavate, short has little affinity with that order. Furthermore,
pedunculate, unitunicate, thin-walled, early B. tropicalis may be incorrectly placed in the
deliquescing, ascospores variable fusiform, genus because of the thin-walled nature of the
ellipsoidal or oblong, 1-3-septate, slightly ascoma, not a feature of the genus. Bathyascus
constricted at the septa, hyaline and superficially resembles Pseudohalonectria in
appendaged. Appendages subterminal at each ascospore morphology, but differ in the degree
end of the spores, with a bulbose base and long of spore septation, and cylindrical asci with a
attenuated arms (Jones et al., 1983a). well developed apical apparatus (Shearer,
The type species has been transferred to 1993b). A molecular study is required to
numerous genera, but molecular data confirms confirm whether B. tropicalis would be better
its position in the Halosphaeriales as a well assigned to Thalespora (Jones et al., 2006).
supported genus (Fig. 41). Jones et al. (1983a)
in a TEM study showed that the appendages 1. Ascospores with no septum, ascomata many layered
differed from those of Corollospora, and re- peridium................................................................... 2
established the genus. In Arenariomyces the 1. Ascospores with 1 septum, 70-100 × 8-10 μm,
subpolar appendages are outgrowths from the ascomata thin-walled ............................. B. tropicalis
mesosporium, while in Corollospora they
comprise both the mesosporium and 2. Ascospores thick-walled, a deep sea species, 50-72 ×
episporium (Jones et al., 1983a). Furthermore, 4-6 μm............................................... B. vermisporus
2. Ascospores thin-walled, a mangrove species........... 3
the base of the appendage is swollen with
electron-dense fibres on one side. 3. Ascospores 90-145 × 2.5-4 μm ............B. avicenniae
3. Ascospores 205-300 × 9-15 μm .......B. grandisporus
1. Ascospores 1-septate................................................ 2
1. Ascospores 3-septate, 27-34 × 6-8 μm. A. triseptatus

83
Bovicornua Jørg. Koch & E.B.G. Jones, Can. J. analysis. It is well delineated in the
Bot. 71: 346, 1993........................................ (1) Halosphaeriales and has affinties with
◙ B. intricata Jørg. Koch & E.B.G. Jones, Can. Remispora pilleata and R. maritima (Fig. 41).
J. Bot. 71: 347, 1993 (Type species).
Ascomata globose to subglobose, 1. Ascospores with transverse septa only.......................
.................................................. C. leptosphaerioides
ostiolate with short conical necks, greyish
1. Ascospores with both transverse and longitudinal
white, erumpent, membranous, gregarious, septa ............................................... .C. pleosporoides
centrum pseudoparenchyma breaks down, no
catenophyses, asci broadly clavate, pedunculate, Ceriosporopsis Linder Farlowia 1: 408, 1944
unitunicate, thin-walled, early deliquescing, no ...................................................................... (5)
apical pore, ascospores unequally 1-septate, C. caduca E.B.G. Jones & Zainal, Mycotaxon
slightly curved, constricted at the septum, 32: 238, 1988.
hyaline and appendaged. At each pole there is a C. cambrensis I.M. Wilson, Trans. Br. Mycol.
single appendage enclosed within an outer Soc. 37: 276, 1954.
sheath which swells when mounted in sea C. capillacea Kohlm., Can. J. Bot. 59: 1314,
water. A genus that shares many features with 1981.
Ceriosporopsis but differs in the degree of ◙ C. halima Linder, Farlowia 1: 408, 1944
elaboration of the ascospore appendages as (Type species).
seen at the TEM level (Yusoff et al., 1993). ◙ C. tubulifera (Kohlm.) P.W. Kirk ex Kohlm.,
The exosporic wall layer enrobes both the Can. J. Bot. 50: 1953, 1972.
spore and polar appendages, which arise as Halosphaeria tubulifera Kohlm. Nova Hedw. 2:
outgrowths of the spore. The exosporium is 312, 1960.
bipartite, the inner region has regular, parallel Ascomata subglobose to cylindrical,
electron-dense lamellations that radiate from immersed, ostiolate, papillate, coriaceous or
the episporium while the outer region consists subcarbonaceous, light brown to black, solitary
of electron-dense fibrillar material. Molecular or gregarious, catenophyses deliquescing, asci
results confirm the placement of Bovicornua in clavate, pedunculate, unitunicate, thin-walled,
the Halosphaeriales. It clusters with deliquescing early, ascospores ellipsoidal, 1-
Ceriosporopsis halima with strong support septate, hyaline, with appendages. Appendage
within the same clade as Marinospora, morphology variable depending on the species
Ceriosporopsis tubulifera and Ondiniella (Fig. (Johnson et al., 1987). Ceriosporopsis
41). cambrensis may have been described on the
basis of more than one species, as the original
Carbosphaerella I. Schmidt, Feddes Repert. 80: description is confused in certain details (Jones
108, 1969...................................................... (2) et al., 1995), consequently can be designated a
◙ C. leptosphaerioides I. Schmidt, Nat. doubtful species.
Naturschutz Mecklenburg 7: 9, 1969. Ceriosporopsis tubulifera differs in many
C. pleosporoides I. Schmidt, Feddes Repert. 80: details from the type species C. halima, but
108, 1969 (Type species). primarily in the origin and nature of the
Ascomata globose or subglobose, ascospore appendage (Figs. 41, 43a, 44a)
superficial, subiculate, ostiolate, papillate or (Johnson et al., 1987). It possesses a polar end-
epapillate, carbonaceous, black, lacking chamber from which mucilage is released, but
paraphyses, asci obpyriform, subglobose, ovoid, this does not arise from the mesosporium.
short pedunculate, lacking an apical apparatus, However, its phylogenetic position is placed
unitunicate, deliquescing early, ascospores within the Marinospora clade with good
ellipsoidal or ovoid, 3-septate or muriform, support (Fig. 41). However, it is not
brown, apical cell paler, with a pronounced monophyletic with the type species,
gelatinous sheath (Johnson et al., 1984). A well Ceriosporopsis halima. In other
delineated genus with its inclusion in the order Ceriosporopsis species the polar appendages
supported by molecular and morphological arise from the mesosporium (Johnson et al.,
evidence. The phylogeny of C. 1987). Ceriosporopsis circumvestita and C.
leptosphaerioides is confirmed by molecular sundica have been transferred to new genera:
84
 Fungal Diversity

a b c

d e f

g h

Fig. 43. Ascospores of various halosphaeriaceous ascomycetes. a. Ceriosporopsis tubulifera. (mucilage arrowed) b.
Lautisporopsis circumvestita. c, d. Cucullosporella mangrovei, apical collar arrowed. e. Halosarpheia trullifera. f.
Saagaromyces glitra. g. Haiyanga salina. h. Antennospora quadricornuta. Bars a, b, e, g = 10 µm, d, f = 25 µm, h =
20 µm.
Lautisporopsis and Limacospora, respectively 2. Polar ascospore appendage an end chamber
(Yusoff et al., 1994a; Jones et al., 1995). containing mucilage .............................. C. tubulifera
2. Polar ascospore appendage arising from the
1. Ascospores with equatorial appendages................... 2 mesosporium, without an end chamber......C. caduca
1. Ascospores lacking equatorial appendages .............. 3 3. Ascospore appendages >750 nm........... C. capillacea

85
3. Ascospore appendages <750 nm.............................. 4 3. Ascomata smaller than 300 µm in diam., ascospores
9-19 × 4-8 µm .................................. C. corallinarum
4. Ascospores 22.5-26 × 8-10 μm ..................C. halima 3. Ascomata greater then 300 µm im diam., ascospores
4. Ascospores 29-31.5 × 10.5-14.5 μm ...C. cambrensis 16-24 × 4-8 µm .................................C. polyporolithi

Chadefaudia Feldm., -Maz. Rev. Gén. Bot. 64: 4. Parasite of Rhodymenia, ascospores 12-16 × 4-6 µm
...................................................................C. marina
150, 1957...................................................... (6) 4. On other algal hosts ................................................. 5
C. balliae Kohlm., Mycologia 65: 244, 1973.
C. corallinarum (P. Crouan & H. Crouan) E. 5. On the alga Schizymenia, ascospores 12-18 × 1-4 µm
Müll. & Arx, The Fungi 4A: 116, 1973. .........................................................C. schizymeniae
Sphaeria corallinarum P. Crouan & H. Crouan, 5. On a wide range of algal hosts, ascospores 14-20 × 4-
Florule Finistère, Paris: 24, 1867. 7 µm.................................................C. gymnogongri
Physalospora corallinarum (P. Crouan & H.
Crouan) Sacc., Syll. Fung. 1: 448, 1882. Corallicola Volkm.-Kohlm. & Kohlm.,
Mycophycophila corallinarum (P. Crouan & H. Mycotaxon 44: 418, 1992 ............................ (1)
Crouan) Kohlm., Nova Hedw. 6: 128, 1963. C. nana Volkm.-Kohlm. & Kohlm.,
C. gymnogongri (Feldmann) Kohlm., Bot. Mar. Mycotaxon 44: 418, 1992 (Type species).
16: 202, 1973. Ascomata subglobose, superficial,
Macrophoma gymnogongri Feldmann, Bot. Soc. ostiolate, short papillate or epapilliate,
Hist. Nat. Afr. Nord. 31: 167, 1940.
Mycophycophila gymnogongri (Feldmann) Cribb
subiculate, coriaceous, dark brown, solitary or
& J.W. Cribb, Pap. Univ. Queensl. Dept. Bot. 4: 43, gregarious, catenophyses deliquescing, asci
1960. deliquescing early, unitunicate, thin-walled,
C. marina Feldmann-Maz. Rev. Gén. Bot. 64: ascospores ellipsoidal, 1-septate, hyaline,
150, 1957 (Type species). slightly constricted at the septum, with 5-7
C. polyporolithi (Bonar) Kohlm., Bot. Mar. 16: polar appendages at each pole. This species is
205, 1973. very similar to Arenariomyces and differs in
Mycophycophila polyporolithi Bonar, Mycologia that the centrum pseudoparenchyma has no pit
57: 379, 1965. connections; ascospore appendages do not
C. schizymeniae Stegenga & Kemperman, Bot. terminate in a spade-like structure; and it can
Mar. 27: 443, 1984. be found on dead coral rocks (Volkmann-
Ascomata subglobose, superficial or Kohlmeyer and Kohlmeyer, 1992). The genus
immersed, ostiolate, papillate or epapillate, should be examined at the molecular level to
coriaceous to carbonaceous, dark brown, determine if it is distinct from Arenariomyces
solitary or gregarious, paraphyses absent, (Jones et al., 1983a).
catenophyses deliquescing, asci subglobose to
clavate, unitunicate, thin-walled, deliquescing Corollospora Werderm., Notizbl. Bot. Gart.
early, ascospores ellipsoidal, 0-septate, hyaline Mus. Berlin-Dahlem 8: 248, 1922 ............. (21)
with gelatinous cap-like appendages at both ◙ C. anglusa Abdel-Wahab & Nagah.,
ends of the spore. The taxonomic position of Mycoscience, in press, 2009.
the genus needs re-evaluation, but fresh Anamorph: Varicosporina anglusa
material for examination is rarely available for Abdel-Wahab & Nagah.
study. It is doubtful if it belongs in the ◙ C. angusta Nakagiri & Tokura, Trans. Mycol.
Halosphaeriales. All species have unicellular, Soc. Jpn. 28: 417, 1988.
hyaline ascospores with small polar pad or cap- Anamorph: ?Varicosporina ramulosa
like appendages. All are parasitic on various Meyers & Kohlm.
algal genera. C. armoricana Kohlm. & Volkm.,-Kohlm. Can.
J. Bot. 67: 1281, 1989.
1. Ascospores longer than 25 µm: 29-38 × 14-22 µm.... ◙ C. baravispora Steinke & E.B.G. Jones sp.
................................................................... C. balliae nov. Fungal Diver. 35: 88, 2009.
1. Ascospores shorter than 25 µm ................................ 2
C. besarispora Sundari, Mycol. Res. 100: 1259,
2. Ascomata superficial, base flat ................................ 3
1996.
2. Ascomata immersed in algal thallus, base rounded . 4 C. californica Kohlm. & Volkm.-Kohlm., Bot.
Mar. 40: 225, 1997.

86
 Fungal Diversity

a b

c d

Fig. 44. Scanning electron micrographs of halosphaeriaceous ascomycetes. a. Ceriosporopsis tubulifera. b. Remispora
crispa. c. Saagaromyces abonnis (Photo by Theresa Baker). d. Tirispora unicaudata. Bars a-c = 5 µm, d = 10 µm.

C. cinnamomea Jørg. Koch, Nordic. J. Bot. 6: ◙ C. lacera (Linder) Kohlm., Ber. Dtsch. Bot.
498, 1986. Ges. 75: 126, 1962.
◙ C. colossa Nakagiri, Trans. Mycol. Soc. Jpn. Peritrichospora lacera Linder, Farlowia 1: 415,
28: 418, 1988. 1944.
◙ C. filiformis Nakagiri, Trans. Mycol. Soc. C. luteola Nakagiri & Tokura, Trans. Mycol.
Jpn. 28: 422, 1988. Soc. Jpn. 23: 102, 1982.
◙ C. fusca Nakagiri & Tokura, Trans. Mycol. Anamorph: Halosigmoidea luteola
Soc. Jpn. 28: 424, 1988. Nakagiri & Tubaki.
C. gracilis Nakagiri & Tokura, Trans. Mycol. ◙ C. maritima Werderm., Notizbl., Königl. bot.
Soc. Jpn. 28: 426, 1988. Gart. u. Museum zu Berlin 8: 248, 1922 (Type
C. indica Prasannarai, Ananda & K.R. Sridhar, species).
Arenariomyces cinctus Höhnk, Veröff. Inst.
J. Environ. Biol. 21: 235, 2000. Meeresforsch. Bremerhaven 3: 28, 1954.
◙ C. intermedia I. Schmidt, Natur Naturschutz Peritrichospora integra Linder, Farlowia 1: 414,
7: 6, 1970. 1944.
Anamorph: ?Varicosporina prolifera C. novofusca Kohlm. & Volkm.-Kohlm., Bot.
Nakagiri Mar. 34: 34, 1991.

87
◙ C. portsaidica Abdel-Wahab & Nagah., material collected in South Africa (Steinke and
Mycoscience, in press, 2008. Jones, 1993). This is sufficiently different from
C. pseudopulchella Nakagiri & Tokura, Trans. other Corollospora species to warrant the
Br. Mycol. Jpn. 28: 428, 1988. erection of a new species, which is proposed
◙ C. pulchella Kohlm., I. Schmidt & N.B. Nair, here.
Ber. Dtsch. Bot. Ges. 80: 98, 1967.
Anamorph: Clavatospora bulbosa Corollospora baravispora Steinke & E.B.G.
(Anastasiou) Nakagiri & Tubaki, Bot. Mar. 28: Jones, sp. nov. .....................................Fig. 45d.
489, 1985. Mycobank:MB513019
◙ C. quinqueseptata Nakagiri, Trans. Br. Ascomata globosa vel subglobosa, superficiala,
Mycol. Jpn. 28: 430, 1988. nigra, carbonacia, solitaria vel gregaria, ostiolata,
papillata, subiculata. Subiculum texturam angularem
Ascomata globose or subglobose, formans. Centrum ascomatis immaturi cellulis
superficial, erumpent, subiculate, ostiolate or pseudoparenchymaticis, hyalinis, leptodermis, foveolatis,
lacking ostioles, papillate or epapillate, deliquescentibus. Paraphyses nullae. Asci octospori,
carbonaceous, pale brown to black, ellipsoidei, unitunicati, mox deliquescentes. Ascosporae
catenophyses deliquescing, asci fusiform or 36-40 × 18-22 μm, fusiformes, 1-septatae, ad septa
leviter constrictae, bruennei. Appendices bigeneres: (i)
subclavate, apiculate, short pedunculate, appendices polares, spiniformes, 10-15 µm longae, 1-2
unitunicate, thin-walled, lacking an apical µm diam. (ii) appendices circa . septum centralem 10-12
apparatus, deliquescing, ascospores various µm longae, peritrichiatae, per fragmenta exosporarum
fusiform, subellipsoidal, constricted at the effectae et appendices apicales 4-6 µm longae.
septum, 1-14-septate, muriform with Substratum: South Africa, Beachwood,
longitudinal and trans-septa, hyaline to pale Durban, Coll. E.B.G. Jones, driftwood buried
brown to dark brown, with appendages. Two in sand, 15 January 1992,
types of appendages: polar spine-like Holotype: culture PP6266, Sequence AF
appendages arising as outgrowths of the 491263, Fig. 45d.
mesosporium and episporium and secondary Etymology: from the Polynesian “baravi” =
fine hair-like appendages formed by beach, and spora, in reference to its habitat.
fragmentation of the exosporium (Jones et al., Ascomata globose, superficial, ostiolate,
1983a) (Figs. 45a-d). Delineation of species papillate, subiculate, carbonaceous, black,
has been largely based on ascospore solitary, attached to sand grains. Neck short, at
measurements, possession of polar spines and base of ascomata. Subiculum dark brown to
nature of the appendages (Prasannarai et al., black. Asci thin-walled, unitunicate
2000). However, Nakagiri and Tubaki (1986) deliquescing early. Ascospores 36-40 × 18-22
also demonstrated significant differences in μm, (excluding polar appendages) 1-septate,
ascomatal wall structure, number of cell wall constricted at the septum, pale brown and
layers and especially the columnar-like cells in appendaged. Polar appendages 15-30 μm long,
the outer layer (Sundari et al., 1996; Hsieh et with equatorial appendages formed by
al., 2007). fragmentation of an exosporic sheath.
Species are common sand dwelling fungi,
1. Ascospores light to dark brown ............................... 2
forming a cohesive morphological group, and 1. Ascospores hyaline .................................................. 7
well supported by molecular data (Campbell et
al., 2002). No distinct taxonomic groups can be 2. Ascospores 1-or more-septate, no trans-septa.......... 3
discerned within the genus, which is probably 2. Ascospores with trans-septa..................................... 6
paraphyletic, and continuously evolving. 3. Ascospores with polar spines................................... 4
However, recent analysis of molecular data 3. Ascospores lacking a polar spine 54-72 × 6-7.5 µm,
suggests the genus may comprise two well- 7-11-septate..........................................C. californica
defined groups (Jones et al., 2009; unpublished 4. Ascospores wider than 10 µm, 1-septate, 36-40 × 18-
data). Corollospora species have anamorphs 22 μm, pale brown ............................. C. baravispora
from three different genera: Clavatospora, 4. Ascospores narrower than 10 µm ............................ 5
Halosigmoidea and Varicosporina.
Campbell et al. (2002) provided a 5. Ascospores 1-septate, 18-25 × 6-9 µm, dark..............
......................................................... C. cinnamomea
sequence of Corollospora sp. isolated from
88
 Fungal Diversity

5. Ascospores 1-septate, 27-32 × 8-9 µm, brown........... Cucullosporella K.D. Hyde & E.B.G. Jones,
............................................................ C. portsaidica Mycotaxon 37: 200, 1990 ............................ (1)
◙ C. mangrovei (K.D. Hyde & E.B.G. Jones)
6. Ascospores 63-220 × 20-38 μm, 12-21 trans-septa,
polar appendages 25-75 μm ..........................C. fusca K.D. Hyde & E.B.G. Jones, Mycotaxon 37:
6. Ascospores 74-99 × 24-34 μm, 13 trans-septa, polar 200, 1990 (Type species).
appendages 52-86 μm ...........................C. novofusca Cucullospora mangrovei K.D. Hyde & E.B.G.
Jones, Bot. Mar. 29: 491, 1986.
7. Ascospores 1-septate................................................ 8 Ascomata subglobose, ovoid or
7. Ascospores with more than 1-septate..................... 10 ellipsoidal, immersed, ostiolate, papillate,
greyish brown to black, solitary or gregarious,
8. Ascospores wider than 8 µm, 22-33 × 8-10 µm, polar
appendages............................................. C. maritima
periphysate, catenophyses present, asci clavate,
8. Ascospores narrower than 8 µm............................... 9 pedunculate, unitunicate, thin-walled,
thickened at the apex with a lens-shaped
9. Ascospores 18-30 × 3-4 (-5), polar appendages 4-7 refractive region, persistent, ascospores fusoid
µm with Varicosporina anamorph ........... C. anglusa to ellipsoidal, 1-septate, not constricted at the
9. Ascospores 26-45 × 3-7 µm, polar appendages 6.5-12
µm ............................................................. C. gracilis
septum, hyaline with polar appendages (Hyde
and Jones, 1986a). This genus, along with
10. Ascospores with 3-septate................................... 11 Halosarpheia, has bipolar unfurling
10. Ascospores with more than 3-septate .................. 13 appendages, but differs from it in that the
11. Ascospores wider than 7 μm ............................... 12 appendages comprise two components, bundles
11. Ascospores narrower than 7 μm wide, 35-57 × 3- of fibrillar material in an amorphous matrix
7.5 μm, polar appendages 3-8 μm ......... C. angusta (Figs. 43c, d). Molecular sequences and
morphological observations confirm that this
12. Ascospores 29-44 × 6.5-8.5 μm, polar appendages
9-13 μm ........................................... C. armoricana genus is distinct from other ascomycetes with
12. Ascospores 25-34 × 7-12 μm, polar appendages 10- bipolar unfurling appendages (Alias et al.,
14 μm ................................................C. intermedia 2001) (Fig. 41).
13. Ascospores with polar appendage ....................... 18 Haligena Kohlm., Nova Hedw. 3: 87, 1961 (1)
13. Ascospores without polar appendages ................ 14
◙ H. elaterophora Kohlm., Nova Hedw. 3: 87,
14. Ascospores 5-septate, 50-85 × 5-8 μm .... C. luteola 1961 (Type species).
14. Ascospores with more than 5-septate .................. 15 Ascomata globose or ovoid, immersed or
15. Ascospores predominately 7-septate ................... 16
superficial, ostiolate, papillate or epapillate,
15. Ascospores with more than 7-septate .................. 17 coriaceous or subcarbonaceous, black, solitary
or gregarious, periphysate, catenophyses
16. Ascospores 60-108 × 13-26 μm ............. C. colossa deliquescing, asci clavate, apiculate,
16. Ascospores 52-112 × 7-16 μm ............ C. pulchella pedunculate, unitunicate, thin-walled, lacking
17. Ascospores 7-11-septate, 65-98 × 8-12 μm.............
an apical apparatus, early deliquescing,
..................................................C. pseudopulchella ascospores oblong ellipsoidal, 3-5-septate,
17. Ascospores 13-septate, 73-120 × 5-8 μm ................ constricted at the septa, hyaline, with polar
............................................................. C. filiformis appendages (Figs. 49a-e). Appendages initially
wrapped around the ascospores but separates
18. Ascospores 3-5-septate........................................ 19
to form long strap-like appendages (Johnson et
18. Ascospores over 5-septate................................... 20
al., 1987).
19 Ascospores 39-63 × 10-19 μm, polar appendages This is a genus that has undergone
10-14 μm .................................................. C. lacera several revisions with species referred to other
19. Ascospores 38-59 × 8-10 μm, polar appendages 5- genera: Appendichordella (Johnson et al.,
12 μm ........................................ C. quinqueseptata 1987), Magnisphaera and Ascosalsum
(Oceanitis), largely based on the morphology
20. Ascospores 7-8-septate, 100-163 × 25-38 μm, polar
appendages 10-38 μm...................... C. besarispora and ultrastructure of the ascospore appendages
20. Ascospores 3-12-septate, 45-102.5 × 7.5-17.5 μm, (Campbell et al., 2003; Dupont et al., 2009).
polar appendages 12-25 μm ..................... C. indica Molecular data indicate H. elaterophora is
89
 a b c

Fig. 45. Ascospores of: a. Corollospora intermedia, (SEM micrograph). b. C. anglusa. c. C. portsaidica. (Photos b, c
by Mohamed Abdel-Wahab). d. C. baravispora. Bars a-c = 10 µm, d = 20 µm.

well placed in the Halosphaeriales but it is not Although all species have bipolar
monophyletic. Haligena salina has therefore unfurling appendages they differ greatly in
been transferred to a new genus Morakotiella other characters. For example, degree of
(Sakayaroj et al., 2005a) (Fig. 41). ascospore septation; wall ornamentation (Jones
and Moss, 1978, reported a highly verrucose
Halosarpheia Kohlm. & E. Kohlm., Trans. Br. wall for H. spartinae); shape of the ascus;
Mycol. Soc. 68: 208, 1977 ........................... (3) length of the ascus pedicel; deliquescent or
Ascomata globose, subglobose, persistent asci; asci with or lacking an apical
obpyriform, ellipsoidal, immersed to pore; presence of catenophyses, and thickness
superficial, ostiolate, papillate, coriaceous, pale of the ascoma wall (Jones, 1995). Molecular
brown to black, catenophyses deliquescing, data suggest that the genus was in need of
necks variable in length, periphysate, asci division but care must be exercised as the
clavate, pedunculate, unitunicate, thin-walled, genus has many overlapping features with
retraction of the plasmalemma apically in some Aniptodera, but few species of that genus have
species, no apical apparatus, persistent, been sequenced to date. Paraphyly of the genus
ascospores broad-ellipsoidal, 1-septate, not must also not be ignored (Mitchell, pers.
constricted at the septum, with apical comm.). As the result of molecular studies
appendages, initially closely adpressed to the (Abdel-Wahab et al., 1999, 2001b; Campbell et
spore wall (hamate) later separating and al., 2003; Pang et al., 2003a,b; Pang and Jones,
uncoiling to form long polar thread-like, sticky 2004) a number of Halosarpheia species have
appendages (Kohlmeyer and Kohlmeyer, 1977) already been transferred to new genera. No
(Figs. 43e, 46). Kong et al. (2000) were the anamorphs have been reported for the marine
first to draw attention to the polyphyletic Halosarpheia species, however Anderson and
nature of this genus, and this has been Shearer (2002) report a Trichocladium
subsequently supported by other studies anamorph for the freshwater species
(Anderson et al., 2001). Halosarpheia heteroguttulata.
90
 Fungal Diversity

Halosarpheia sensu stricto 4. Ascospores 18-23 (-26) × 9-12 μm, ascus with short
◙ H. fibrosa Kohlm. & E. Kohlm., Trans. Br. pedicel, appendages cap-like, then filamentous .........
.................................................................. H. marina
Mycol. Soc. 68: 208, 1977 (Type species).
4. Ascospores 12-21 × 4-7.5 μm, ascus with a long
◙ H. trullifera (Kohlm.) E.B.G. Jones, S.T. pedicel, appendages well developed ..... H. kandeliae
Moss & Cuomo, Trans. Br. Mycol. Soc. 80:
200, 1983. 5. Ascospores 25-32 × 9-13 μm, on Juncus ...................
Remispora trullifera Kohlm., Nova Hedw. 6: 321, ............................................................H. culmiperda
1963. 5. Ascospores 25-35 × 7.5-10.5 μm, on Phragmites......
Halosphaeria trullifera (Kohlm.) Kohlm., Can. J. ....................................................... H. phragmiticola
Bot. 50: 1956, 1972.
◙ H. unicellularis Abdel-Wahab & E.B.G. We accept the species keyed out above as
Jones, Mycoscience 42: 255, 2001. Halosarpheia sensu stricto species, however,
the remaining must await further studies at the
1. Ascospores unicellular ..................... H. unicellularis molecular level. Halosarpheia kandeliae
1. Ascospores 1-septate................................................ 2
differs from many of the Halosarpheia species
2. Ascospores 32-44 × 18-24 μm .................. H. fibrosa in having asci with long drawn out stalks (tail-
2. Ascospores 23-32 (-36) × 14-18 μm. ..... H. trullifera like). Species with similar ascospore
measurements have asci with short pedicels.
Halosarpheia sensu lato Another unusual feature of H. kandeliae is the
H. bentotensis Jørg. Koch, Nordic. J. Bot. 2: polar appendages initially appears amorphous
165, 1982. and only later forms the characteristic thread-
H. culmiperda Kohlm., Volkm.-Kohlm. & O.E. like bipolar appendages (Abdel-Wahab et al.,
Erikss., Mycologia 87: 532, 1995. 1999). Although ascospore measurements of
◙ H. kandeliae Abdel-Wahab & E.B.G. Jones Halosarpheia culmiperda are similar to those
Mycol. Res. 103: 1500, 1999. of H. viscosa, it differs in ascus morphology.
◙ H. marina (Cribb & J.W. Cribb) Kohlm., Asci of H. culmiperda are similar with those of
Mar. Ecol. (P.S.Z.N.I.) 5: 345, 1984. the H. fibrosa group in that they are persistent,
Gnomonia marina Cribb & J.W. Cribb, Pap. Univ. clavate to oval, with a pore, and a stalk one
Queensl. Dept. Bot. 3: 100, 1956. third of the ascus length.
H. minuta W.F. Leong, Can. J. Bot. 69: 883,
1991. Species transferred to other genera:
H. phragmiticola Poon & K.D. Hyde, Bot. Mar.
41: 143, 1998. H. abonnis, H. ratnagiriensis to Saagaromyces
The position of H. bentotensis, H. K.L. Pang & E.B.G. Jones
culmiperda, H. kandeliae, H. marina, H. H. spartinae to Magnisphaera J. Campb., J.L.
minuta and H. phragmiticola remains Anderson & Shearer
unresolved. In a number of cases cultures of H. cincinnatula, H. unicaudata and H.
these species are not available for a molecular viscidula to Oceantitis J. Kohlm
study. Until such data is available it is pointless (Ascosalsum J. Campb., J.L. Anderson &
to speculate as to their future assignment at the Shearer)
generic level. H. retorquens and H. lotica to Natantispora J.
Campb., J.L. Anderson & Shearer
1. Ascospores 3-septate, 28-36 × 6-8 μm....................... H. viscosa to Panorbis J. Campb., J.L.
............................................................H. bentotensis
1. Ascospores 1-septate................................................ 2 Anderson & Shearer
H. aquatica and H. heteroguttulata to
2. Ascospores less than 15 μm long (10-14 × 4-6 μm)... Ascosacculus J. Campb., J.L. Anderson
.................................................................. H. minuta & Shearer, both freshwater species.
2. Ascospores longer than 15 μm long......................... 3

3. Species on wood ...................................................... 4

Halosphaeria Linder, Farlowia 1: 412, 1944. ..
3. Species on other substrata ........................................ 5 ...................................................................... (1)

91
◙ H. appendiculata Linder, Farlowia 1: 412, constricted at the septum, hyaline with a small
1944 (Type species). apical appendage and 3-4 crescent-shaped
Ascomata globose or subglobose, equatorial appendages. Appendages formed by
ellipsoidal, immersed erumpent, ostiolate, the fragmentation of an exosporic sheath
papillate, coriaceous or membranous, pale to (Moss and Jones, 1977; Jones et al., 1984). A
dark brown to black, solitary or gregarious, monotypic genus well placed in the
catenophyses deliquescing, asci clavate or Halosphaeriales and supported by both
subfusiform, pedunculate, unitunicate, thin- morphological and molecular evidence
walled, deliquescing early, ascospores ellip- (Spatafora and Blackwell, 1994; Sakayaroj et
soidal, 1-septate, hyaline, with 3-4 spoon- al., 2005a) (Fig. 41).
shaped equatorial appendages and one at each
end of the spore. Ultrastructure of the Haiyanga K.L. Pang & E.B.G. Jones, The
appendages have been elucidated by Jones et al. Raffles Bull. Zool. 19: 8, 2008 .................... (1)
(1984). ◙ H. salina (Meyers) K.L. Pang & E.B.G.
Kohmeyer (1972a) referred 12 species to Jones, The Raffles Bull. Zool. 19: 8, 2008.
this genus but subsequent studies have assigned Arenariomyces salina Meyers, Mycologia 49: 505,
them to other genera: 1957.
Remispora salina (Meyers) Kohlm., Mycologia
60: 262, 1968.
H. cucullata to Okeanomyces K.L. Pang &
Halosphaeria salina (Meyers) Kohlm., Can. J.
E.B.G. Jones Bot. 50: 1957, 1972.
H. quadricornuta to Antennospora Meyers Antennospora salina (Meyers) Yusoff, E.B.G.
H. salina to Haiyanga K.L. Pang & E.B.G. Jones & S.T. Moss, Mycol. Res. 98: 1003, 1994.
Jones Ascomata globose to subcylindrical,
H. mediosetigera to Halosphaeriopsis T.W. immersed, ostiolate, papillate, membranous or
Johnson coriaceous, dark brown to black, periphysate,
H. maritima, H. pilleata, H. quadri-remis, H. catenophyses, asci clavate to ellipsoidal, short
stellata to Remispora Linder pedunculate, thin-walled, unitunicate, no apical
H. trullifera to Halosarpheia Kohlm. & E. apparatus, deliquescing early, ascospores
Kohlm. ellipsoidal, 1-septate, slightly constricted at the
H. circumvestita to Lautisporopsis E.B.G. septum, hyaline with appendages at both ends
Jones, Yusoff & S.T. Moss of the spore. Appendages are obclavate, curved,
H. torquata to Ondiniella E.B.G. Jones, R.G. attenuate and observed to release a drop of
Johnson & S.T. Moss mucilage from their tips (Fig. 43g) (Zainal and
Currently only one species is accepted in Jones, 1984). In H. salina the appendages are
the genus (Pang et al., 2004b; Sakayaroj et al., outgrowths of the spore and attached to the
2005a). mesosporium by an isthmus of electron-dense
material. At the TEM level electron-dense
Halosphaeriopsis T.W. Johnson, J. Elisha striations run the entire length of the appendage,
Mitchell Sci. Soc. 74: 44, 1958.................... (1) these are confined to the outer layer of the
◙ H. mediosetigera (Cribb & J.W. Cribb) T.W. spore while the core appears amorphous
Johnson, J. Elisha Mitchell Sci. Soc. 74: 44, (Yusoff et al., 1994c).
1958 (Type species). Haiyanga salina is distantly related to
Halosphaeria mediosetigera Cribb & J.W. Cribb,
Antennospora quadricornuta and Halosphaeria
Univ. Queensl. Pap., Dept. Bot. 3: 100, 1956.
Anamorph: Trichocladium achrasporum appendiculata, but well placed in the
(Meyers & R.T. Moore) Dixon Halosphaeriales (Figs 41, 47). Haiyanga salina,
Ascomata subglobose or ellipsoidal, instead of clustering with Antennospora
immersed or erumpent, ostiolate, papillate, quadricornuta (a genus it was most recently
subcarbonaceous, dark brown to black, solitary assigned to), forms a moderately-supported
or gregarious, asci clavate, pedunculate, group with two isolates of Arenariomyces
unitunicate, thin-walled, lacking an apical trifurcatus (Fig. 47). However, the two genera
apparatus, deliquescing early, ascospores differ in their habitat, nature of the ascomata
ellipsoidal or subfusiform, 1-septate, slightly and most strickling in the origin and

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 Fungal Diversity

Fig. 46. Halosarpheia unicellularis. Ascospores with one polar appendage (arrowed). Bars = 10 μm.

ultrastructural detail of the appendages. (Pang et al., 2008a). The tufts of polar and
Morphological data also support the erection of equatorial appendages in Havispora resembles
a new genus for this species, especially the those of the genera Nautosphaeria and
ultrastructure and ontogeny of the ascospore Nereiospora. However the ascospores of N.
appendages (Pang et al., 2008b). Ascospore cristaminuta are unicellular while Nereiospora
appendages of Haiyanga salina are also similar species have ascospore with brown central cells.
to those of Remispora species, especially R. Nereiospora cristata has Monodictys pelagica
stellata and R. quadriremis. as its anamorph (Mouzouras and Jones, 1985).
The molecular result indicates clearly that this
Havispora K.L. Pang & Vrijmoed, Mycologia genus is well placed in the Halosphaeriales. It
100: 293, 2008 ............................................. (1) is phylogenetically nestled in the Nereiospora
◙ H. longyearbyenensis K.L. Pang & and Monodictys clade, although with low
Vrijmoed, Mycologia 100: 293, 2008. bootstrap support (Fig. 41).
Ascomata subglobose to ellipsoidal,
immersed, coriaceous, solitary or gregarious, Iwilsoniella E.B.G. Jones, Syst. Ascomycetum
lacking periphyses, catenophyses present 10: 8, 1991 ................................................... (1)
deliquescing, long necks, asci clavate, I. rotunda E.B.G. Jones, Syst. Ascomycetum
pedunculate, unitunicate, thin-walled, 10: 8, 1991 (Type species).
persistent, ascospores ellipsoidal, hyaline, thin- Ascomata subglobose, immersed to
walled, 3-septate, constricted at the septa, with superficial, ostiolate, papillate, membranous,
tufts of appendages (Pang et al., 2008a). dark brown, solitary or gregarious, no
Appendages polar and four at the catenophyses, asci clavate, short pedunculate,
central septum, string-like composed of lacking an apical pore, unitunicate, thin-walled,
intertwining strands (Fig. 53b). A new genus deliquescing early, ascospores round,
described from material collected at unicellular, hyaline without a visible sheath.
Longyearbyen, Norway on driftwood and However, at the scanning electron
characterized by the ellipsoidal thin-walled microscope level a sheath is apparent peeling
ascospores, 24-(30)-36 × 8-(11)-14 μm with away from the spore wall. It sporulates in
tufts of polar and equatorial appendages culture on cornmeal seawater agar. It was
initially string-like and composed of described from a test block exposed in a water
intertwining strands that separate in seawater cooling tower with brackish water circulating
93
hyaline, round, one-celled ascospores lacking L. tropica does not belong in the genus either,
appendages and deliquescing asci. This genus but lack of a consensus on the morphology of
requires further study to determine if it is its ascus prevents its transfer to a new genus
correctly assigned to the Halosphaeriales (Kohlmeyer, 1984; Kohlmeyer and Volkmann-
(Jones, 1991). Kohlmeyer, 1988a) (Fig. 41). Originally the
ascus was described as possessing an apical
Lautisporopsis E.B.G. Jones, Yusoff & S.T. pore (Kohlmeyer, 1984), but subsequently
Moss, Can. J. Bot. 72: 1558, 1994 ............... (1) Kohlmeyer and Volkmann-Kohlmeyer (1988a)
◙ L. circumvestita (Kohlm.) E.B.G. Jones, revised the description to exclude taxa with an
Yusoff & S.T. Moss, Can. J. Bot. 72: 1558, apical ascus pore from the genus. Molecular
1994 (Type species). data show no affinity between L. tropica (ascus
Halosphaeria circumvestita Kohlm., Nova Hedw. with an apical pore) and the type species,
2: 307, 1960.
Ceriosporopsis circumvestita (Kohlm.) Kohlm.,
consequently it should be transferred to a new
Can. J. Bot. 50: 1953, 1972. genus (Pang et al., 2003a). We have been
Ascomata subglobose, ovoid, ostiolate, unable to obtain fresh material of L. tropica
papillate, immersed to superficial, light to dark (with no apical pore), therefore reassignment
brown, coriaceous, solitary or gregarious, must await further molecular studies.
catenophyses, asci clavate, pedunculate, Lignincola nypae clearly does not belong in
unitunicate, thin-walled, no apical apparatus, this genus but is left here for the present until
deliquescing early, ascospores ellipsoidal, 1- fresh material is collected, isolated and
septate, slightly constricted at the septum, sequenced.
hyaline with a mucilaginous sheath extended to
form equatorial and polar appendages (Fig. 1. Ascospores ellipsoidal ............................................. 2
1. Ascospores cylindrical ................................. L. nypae
43b). A monotypic genus with the transfer of
Ceriosporopsis circumvestita to this genus 2. Ascospores 22-36 × 12-16 μm, thick-walled. ............
based on a scanning and transmission electron ................................................................... L. tropica
microscope studies of ascospore appendage 2. Ascospores 13-24 × 5-8 μm, thin-walled..... L. laevis
ontogeny (Yusoff et al., 1994a).
Limacospora Jørg. Koch & E.B.G. Jones, Can.
Lignincola Höhnk, Veröeff. Inst. Meeresforsch. J. Bot. 73: 1011, 1995 .................................. (1)
Bremerhaven 3: 216, 1955 ........................... (3) L. sundica (Jørg. Koch & E.B.G. Jones) Jørg.
◙ L. laevis Höhnk, Veröeff. Inst. Meeresforsch. Koch & E.B.G. Jones, Can. J. Bot. 73: 1013,
Bremerhaven 3: 216, 1955 (Type species). 1995 (Type species).
L. nypae K.D. Hyde & Alias, Mycol. Res. 103: Ceriosporopsis sundica Jørg Koch & E.B.G.
1417, 1999. Jones, Nordic J. Bot. 6: 339, 1986.
◙ L. tropica Kohlm., Mar. Ecol. (P.S.Z.N.I.) 5: Ascomata deeply immersed in wood, pale
355, 1984. brown, ostiolate, long necks, no periphyses,
Ascomata subglobose to ellipsoidal, asci clavate, tapering, pedunculate, unitunicate,
immersed to superficial, ostiolate, papillate, thin-walled, deliquescing early, interthecial
coriaceous, light brown to black, catenophyses filaments present, ascospores 1-0-septate,
deliquescing, asci clavate or subfusiform, hyaline, with a mucilaginous sheath.
pedunculate, unitunicate, thin-walled, Originally described as Ceriosporopsis
persistent, apiculate, asci released from the sundica, but segregated to a new genus based
ascoma through the neck, ascospores on ultrastructural characters, primarily the
ellipsoidal, 1-septate, hyaline, lacking ontogeny of the ascospore appendages (Jones
appendages (Pang et al., 2003a). The genus et al., 1995).
Lignincola has only one unifying character, the The polar appendages are extensions of
hyaline, 1-septate ascospores, lacking the sheath and not discrete mesosporial
appendages. Lignincola longirostris with an outgrowths characteristic of the genus
ascus with an apical pore and retraction of the Ceriosporopsis. Its relationship with
plasmalemma has been transferred to the new Ceriosporopsis, Marinospora and Bovicornua,
genus Neptunella (Pang et al., 2003a). Clearly requires evaluation at the molecular level.
94
 Fungal Diversity

Fig. 47. Phylogram of selected Halosphaeria species and related genera in the Halosphaeriales.

Luttrellia Shearer, Mycologia 70: 692, 1978 ... ostiolate, neck long, cylindrical hyaline and
...................................................................... (1) periphysate. Asci clavate to cylindrical, thin-
L. estuarina Shearer, Mycologia 70: 693, 1978 walled and deliquescing. Catenophyses present.
(Type species). Ascospores hyaline, multiseptate and lacking
Ascomata superficial, globose to sub- appendages. There is no known ana-morph.
globose, membranous, becoming pale brown, The only genus in the Halosphaeriales
95
 a b

Fig. 48. Scanning electron micrographs of selected ascomycetes with polar unfurling appendages. a, b. Oceanitis
cincinnatula. c. Magnisphaera spartinae. e. Natantispora retorquens. d, f. Oceanitis unicaudata. Bars a, c = 10 µm, b,
d-f = 2 µm.

with tetrasporic asci. Saprophytic on wood. An Pang & E.B.G. Jones, Nova Hedw. 77 (1-2):15,2003.
infrequently collected species and its position Ascomata globose to subglobose,
in the Halosphaeriales requires verification. immersed, ostiolate, papillate, dark brown to
black, thick-walled, periphysate, solitary or
Magnisphaera J. Campb., J.L. Anderson & gregarious, asci clavate, pedunculate,
Shearer, Mycologia 95: 546, 2003 ............... (1) unitunicate, thin-walled, deliquescing early,
◙ M. spartinae (E.B.G. Jones) J. Campb., J.L. ascospores ellipsoidal to fusiform, hyaline, 4-9-
Anderson & Shearer, Mycologia 95: 547, 2003 septate (generally 5), constricted at the septa,
(Type species). with polar appendages, initially closely
Haligena spartinae E.B.G. Jones, Trans Br. adpressed to the spore wall, then separating and
Mycol. Soc. 45: 245, 1962. uncoiling to form long thread-like appendages
Halosarpheia spartinae (E.B.G. Jones) Shearer &
(Fig. 48c). A new genus erected as the result of
J.L. Crane, Bot. Mar. 23: 608, 1980.
Matsusphaeria spartinae (E.B.G. Jones) K.L a re-evaluation of the genus Halosarpheia,

96
 Fungal Diversity

distinguished by its broadly acerose, septate 1. Ascospores 20-36 × 7-19 μm, with polar and
ascospores, constricted at the septa, with a equatorial appendages equal in size ..... M. calyptrata
verrucose spore wall, polar appendages arising
from a pore and supported by 18S rDNA Moana Kohlm. & Volkm.-Kohlm., Mycol. Res.
sequence data (Campbell et al., 2003) (Fig. 41). 92: 418, 1989 ............................................... (1)
M. turbinulata Kohlm. & Volkm.-Kohlm.,
Marinospora A.R. Caval., Nova Hedw. 11: Mycol. Res. 92: 418, 1989 (Type species).
548, 1966...................................................... (2) Ascomata subglobose, immersed,
◙ M. calyptrata (Kohlm.) A.R. Caval., Nova ostiolate, papillate, coriaceous, cream-coloured,
Hedw. 11: 548, 1966 (Type species). solitary, periphysate, catenophyses present,
Ceriosporopsis calyptrata Kohlm., Nova Hedw. 2: asci clavate, pedunculate, rounded at the apex,
301, 1960. lacking a pore, persistent, ascospores
Ceriosporella calyptrata (Kohlm.) Caval., Nova
Hedw. 10: 394, 1966.
subglobose, unicellular, hyaline with a single
◙ M. longissima (Kohlm.) A.R. Caval., Nova top-shaped appendage that unfurls to form
Hedw. 11: 548, 1966. tapering ribbons. The unique feature of this
Ceriosporopsis longissima Kohlm., Nova Hedw. species is the subglobose ascospores with a
4: 398, 1962. single polar top-shaped appendage within the
Ceriosporella longissima (Kohlm.) Caval., Nova ascus, which unfurls in water to produce a long
Hedw. 10: 394, 1966. ribbon 250-550 µm long, 1.5-3 µm wide,
Ascomata ellipsoidal, subglobose, which stains with methylene and cotton blue
immersed, ostiolate, papillate, subcarbonaceous (Kohlmeyer and Volkmann-Kohlmeyer, 1989).
or subcoriaceous, light brown to black, Its relationship with other members of the
periphysate, solitary or gregarious, Halosphaeriaceae remains to be determined. It
catenophyses deliquescing, asci clavate, is similar to Thalassogena sphaerica and
unitunicate, thin-walled, no apical apparatus, Anisostagma rotundatum but differs in that
deliquescing early, ascospores broadly these do not have polar appendages.
ellipsoidal, 1-septate, constricted at the septum,
hyaline, with equatorial and polar appendages: Morakotiella Sakay. Mycologia 97: 806, 2005
a single appendage at each pole, and 4 ...................................................................... (1)
radiating appendages at the central septum. ◙ M. salina (C.A. Farrant & E.B.G. Jones)
Primary appendage an outgrowth of the Sakay., Mycologia 97: 806, 2005 (Type
mesosporium and episporium, with a species).
fragmenting exosporic sheath forming tiny Haligena salina C.A. Farrant & E.B.G. Jones, J.
caps to the tips of the primary appendages Linn. Soc. Bot. 93: 406, 1986.
(Johnson et al., 1984). Ascomata immersed or partly immersed,
These species were initially referred to globose, ostiolate, papillate, coriaceous, black,
Ceriosporopsis, but subsequently transferred to solitary or gregarious, catenophyses present,
Ceriosporella and finally Marinospora. early deliquescing, asci fusiform to clavate,
Sequence data confirm that the genus is distinct pedunculate, thin-walled, unitunicate,
from Ceriosporopsis, although closely related deliquescing early, ascospores ellipsoidal, 1-
(Sakayaroj et al., 2004) (Fig. 41). septate, slightly constricted at the septum, (14-)
Morphologically its polar and equatorial 16-18 (-20) × 8-10 µm, hyaline with polar
appendages are distinctive with the cup-like appendages. Appendages polar, spoon-shaped
exosporic remnants at their tips (Johnson et al., at their base, attenuate, channelled, attached
1984). Marinospora longissima has been apically and arising as an outgrowth of the
regarded as a synonym of M. calyptrata spore wall (Figs. 49 f-j) (Farrant and Jones,
(Kohlmeyer and Kohlmeyer, 1979), but 1986). Referred initially to Haligena because
molecular data indicate they are distinct species of its polar appendages coiled around the
(Fig. 41). ascospore, but molecular sequences indicate it
is distantly related to Haligena elaterophora
1. Ascospores 20-34 × 8-18 μm, with longer polar
(Figs. 41, 47). Morphologically they differ in
appendages to equatorial appendages .M. longissima
97
the degree of ascospore septation, the ontogeny hyaline with a hamate appendage initially
of the polar appendages and ascoma structure closely adpressed to the spore wall, separating
(Sakayaroj et al., 2005a). and unraveling to form long drawn out thread-
like polar appendages (Fig. 48e). Campbell et
Nais Kohlm., Nova Hedw. 4: 409, 1962...... (1) al. (2003) segregated these two species from
◙ N. inornata Kohlm., Nova Hedw. 4: 409, Halosarpheia based on sequence data, although
1962 (Type species). distinguishing morphological features at the
Ascomata subglobose, immersed to generic level are not well established. From
superficial, ostiolate, papillate, coriaceous, dark combined 18S and 28S sequences Natantispora
brown to black, gregarious, long necks (100- species are distantly placed from Halosarpheia
620 µm), periphysate, catenophyses present, sensu stricto (Figs. 41, 47) (Abdel-Wahab et al.,
asci clavate, short pedunculate, unitunicate, 2001b).
thin-walled, no apical apparatus, deliquescing
early, ascospores broadly ellipsoidal, 1-septate, 1. Ascospores 26-38 × 10-14 μm, appendages not
slightly constricted at the septum, hyaline, extending to the middle septum ................... N. lotica
lacking appendages or a sheath. A genus 1. Ascospores 20-33 × 7-11 μm, appendages extend
characterized by hyaline bicelled ascospores beyond the middle septum ...................N. retorquens
with a characteristic arrangement of the
internal wall ornamentation along the septum Naufragella Kohlm. & Volkm.-Kohlm., Syst.
where small oil globules aggregate and lacking Ascomycetum 16: 10, 1998 ......................... (2)
appendages. It is inferred from phylogenetic N. delmarensis Kohlm. & Volkm.-Kohlm.,
analysis of the partial large subunit ribosomal Syst. Ascomycetum 16: 10, 1998 (Type
DNA sequences to be closely related to species).
Aniptodera, forming a moderately supported ◙ N. spinibarbata (Jørg. Koch) Kohlm. &
monophyletic group with a bootstrap value of Volkm.-Kohlm., Syst. Ascomycetum 16: 11,
89% (Pang et al., 2003a) (Figs. 41, 47, 55). 1998.
Two further species have been described: N. Remispora spinibarbata Jørg. Koch, Nordic J.
Bot. 8: 517, 1989.
glitra now transferred to Saagaromyces and a
Ascomata subglobose, immersed or
freshwater species N. aquatica. The latter
superficial, ostiolate, papillate, coriaceous,
differs from N. inornata in that ascospores
cream-coloured, solitary or gregarious, neck
develop appendages on release from the
periphysate, catenophyses, asci clavate,
ascomata (Hyde, 1992d).
pedunculate, unitunicate, thin-walled, early
deliquescing, no apical apparatus, ascospores
Natantispora J. Campb., J.L. Anderson &
ellipsoidal, 1-septate, not constricted at the
Shearer, Mycologia 95: 543, 2003 ............... (2)
septum, hyaline, with two types of appendages.
◙ N. lotica (Shearer) J. Campb., J.L. Anderson
Polar gelatinous appendages covering the
& Shearer, Mycologia 95: 543, 2003.
Halosarpheia lotica Shearer, Mycotaxon 20: 505,
apical region and along one side of the spore,
1984. becoming strap-like or forming a wide band;
◙ N. retorquens (Shearer & J.L. Crane) J. second type subpolar, a fragmenting gelatinous
Campb., J.L. Anderson & Shearer, Mycologia sheath forming a crown of hair-like filaments
95: 543, 2003 (Type species). born on flat cushions. Naufragella species are
Halosarpheia retorquens Shearer & J.L. Crane, frequently difficult to distinguish from
Bot. Mar. 23: 608, 1980. Remispora. Mounted in seawater, two types of
Ascomata globose, immersed to appendages can be discerned: 1). Gelatinous
superficial, ostiolate, papillate, membranous, strap-like polar appendages with faint striations,
black, long neck, periphysate, catenophyses initially covering the apices and one side of the
present deliquescing, asci clavate, short ascospores but swelling in water and separating
pedunculate, unitunicate, thin-walled, with an from the side of the spore and extending to 90
apical pore and retraction of the plasmalemma µm long; and 2). Sub-polar hair-like
at the tip of the ascus, deliquescing early, appendages arising from a small flat cushion
ascospores fusiform to ellipsoidal, 1-septate, and spreading out in water (Koch, 1989).

98
 Fungal Diversity

a b

f c d e

g h

i j

Fig. 49. Morphological features of Haligena species a-e. Haligena elaterophora (JS147) a. Ascus containing
ascospores; b-e. Ascospores multi-septate with bipolar, long, strap-like appendages. f-j. Morakotiella salina
(BCC12781) f. Ascospore forms a long thread-like appendage after release into water; g. Black, globose ascomata: h, j.
Ascospores one-septate with tightly coiled appendages around the ascospores; i. Cylindrical-clavate ascus. Bars a-e =
20 µm; g = 100 µm; f, h-j = 10 µm. Reproduced by permission of Mycologia.
The origin of these appendages has not 1. Ascospores 18-26 × 8-12 μm, with smooth apical
been elucidated. The appendaged ascospores of appendages........................................N. spinibarbata
Nohea umiumi superficially resemble those of
Naufragella, but differ in that in the former Nautosphaeria E.B.G. Jones, Trans Br. Mycol.
ascospores have two thick gelatinous Soc. 47: 97, 1964 ......................................... (1)
appendages attached to one side of the spore ◙ N. cristaminuta E.B.G. Jones, Trans Br.
and two tufts of subapical fibres on the other Mycol. Soc. 47: 97, 1964 (Type species).
side (Kohlmeyer and Volkmann-Kohlmeyer, Ascomata globose to subglobose,
1998a). Molecular data show that Naufragella immersed, ostiolate, papillate, coriaceous,
is distantly related to Remispora (Fig. 41). hyaline to pale brown, solitary or gregarious,
Morphologically the two Naufragella catenophyses deliquescing, asci clavate to
species are very similar and differ only slightly ellipsoidal, short pedunculate, unitunicate, thin-
in ascospore measurements, which overlap, and walled, lacking an apical apparatus,
in the nature of the apical appendages. deliquescing early, ascospores ellipsoidal,
Naufragella delmarensis is probably a unicellular, hyaline, with tufts of bristle-like
synonym of N. spinibarbata, but sequence data appendages apically at the spore equator.
are required for these two rare species before Sakayaroj et al. (2004) confirmed the
this is formally proposed (Koch, pers. comm.). assignment of this genus within the
Halosphaeriales based on partial 28S rDNA
1. Ascospores 18.5-21 × 8-9.5 μm, with striated apical sequences which grouped with Haligena
appendages.........................................N. delmarensis elaterophora and Remispora galerita (Fig. 41).
99
However, there is no homology between ◙ N. comata (Kohlm.) E.B.G. Jones, R.G.
appendage ontogeny in these genera. The hair- Johnson & S.T. Moss, J. Linn. Soc. Bot. 87:
like appendages in Nautosphaeria are similar 206, 1983. (Type species).
to those of Nereiospora species, however the Peritrichospora comata Kohlm., Nova Hedw. 2:
latter are 3-5-septate, central cells brown with 323, 1960.
Corollospora comata (Kohlm.) Kohlm., Ber.
hyaline to pale ends cells. Phylogenetically Dtsch. Bot. Ges. 75: 126, 1962.
they are not related (Sakayaroj et al., 2004). ◙ N. cristata (Kohlm.) E.B.G. Jones, R.G.
Johnson & S.T. Moss, J. Linn. Soc. Bot. 87:
Neptunella K.L. Pang & E.B.G. Jones, Mycol. 206, 1983.
Prog. 2: 35, 2003 .......................................... (1) Peritrichospora cristata Kohlm., Nova Hedw. 2:
◙ N. longirostris (Cribb & J.W. Cribb) K.L. 324, 1960.
Pang & E.B.G. Jones, Mycol. Progr. 2 : 35, Corollospora cristata (Kohlm.) Kohlm., Ber.
2003 (Type species). Dtsch. Bot. Ges. 75: 126, 1962.
Gnomonia longirostris Cribb & J.W. Cribb, Pap. Anamorph: Monodictys pelagica (T.W.
Dept. Bot. Univ. Qd. 3: 101, 1956. Johnson) E.B.G. Jones
Lignincola longirostris (Cribb & J.W. Cribb) Ascomata subglobose or ellipsoidal,
Kohlm., Mar. Ecol. (P.S.Z.N.I.) 5: 353, 1984. immersed or superficial, with or without
Ascomata subglobose, immersed to ostioles, papillate or epapillate, carbonaceous,
erumpent, ostiolate, with long necks (80-1125 black, solitary or gregarious, centrum cells
µm), membranous, hyaline, pale to brown, deliquescing, asci broadly fusiform or clavate,
solitary or gregarious, no periphyses or pedunculate, unitunicate, thin-walled, lacking
paraphyses, asci cylindrical-clavate, an apical apparatus, deliquescing early,
ellipsoidal-clavate, short pedunculate, ascospores 3-5-septate. Constricted at the septa,
unitunicate, thick-walled apically, with central cells brown, end cells hyaline, with
retraction of the plasmalemma at the apex, tufts of hair-like appendages apically and at the
apical pore, persistent to deliquescing, central septum. Jones et al. (1983a) erected the
ascospores elongate ellipsoidal, 1-septate, genus to accommodate species with polar and
slightly constricted at the septum, no sheath or equatorial appendages that are ontogenetically
appendages (Figs 41, 55). distinct from those of Corollospora.
Assignment of this species with its In the study by Campbell et al. (2002)
hyaline, bicelled ascospores lacking Nereiospora nestles within the Corollospora
appendages has proved difficult, initially clade. However, in a more extensive treatment
referred to Gnomonia, then Lignincola, but of the Halosphaeriales, Nereiospora is distantly
differing from the type species L. laevis in placed from Corollospora (Fig. 41). In
ascus morphology. Sequences of the large Nereiospora appendages arise from the
subunit rRNA gene, indicates that Lignincola mesosporium, while in Corollospora they are
laevis and L. longirostris have a close spine-like and formed by outgrowths of the
relationship, but they are not monophyletic. epi- and mesosporium, while the secondary
Neptunella has therefore been erected for L. appendages are formed by fragmentation of an
longirostris as it differs from L. laveis as exosporial layer (Jones et al., 1983a).
follows: 1). asci with retraction of the
plasmalemma, an apical thickening and pore; 1. Ascospores 32-54 × 12-17 μm, with 5 septa..............
2). an extra ascospore wall layer (the .................................................................. N. comata
exosporium) is present; and 3). asci are 1. Ascospores 24-41 × 8-16 μm, with 3 septa................
.................................................................. N. cristata
persistent and do not get released through the
ostiole, as in L. laevis. Thus there is Nimbospora Jørg. Koch, Nordic J. Bot. 2: 166,
morphological and molecular evidence to 1982.............................................................. (3)
separate these two taxa (Pang et al., 2003a). N. bipolaris K.D. Hyde & E.B.G. Jones, Can. J.
Bot. 63: 611, 1985.
Nereiospora E.B.G. Jones, R.G. Johnson & ◙ N. effusa Jørg. Koch, Nordic J. Bot. 2: 166,
S.T. Moss, J. Linn. Soc. Bot. 87: 204, 1983...(2) 1982 (Type species).

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 Fungal Diversity

N. octonae Kohlm., Can. J. Bot. 63: 1122, has two types of ascospore appendages, but
1985. phylogenetically are distantly placed within the
Ascomata globose to subglobose, Halosphaeriales (Fig. 41). Naufragella and
ostiolate, papillate, membranous, hyaline, Remispora (sensu stricto) species have a
immersed, solitary or gregarious, necks common ancestor but are not monophyletic.
periphysate, no paraphyses, asci clavate, short Nohea forms a sister group to Aniptodera
pedunculate, unitunicate, thin-walled, chesapeakensis and Nimbospora effusa
deliquescing early, ascospores ellipsoidal, 1- (Campbell et al., 2002), while in a larger data
septate, not constricted at the septum, hyaline, set it nestles in a sister group to Neptunella
and appendaged. Ascospores in Nimbospora longirostris and is basal to a clade comprising a
have two types of appendages: a well range of taxa (Fig. 41).
developed exosporic mucilaginous sheath, and Kohlmeyer and Volkmann-Kohlmeyer
secondary fibrillar appendages at the spore (1991c) compare N. umiumi with the genera
equator that are formed as outgrowths of the Halosarpheia (does not possess tufts of hair-
mesosporium. These fibrillar appendages like appendages), Nautospora (probably mean
initially are contained within the sheath, but on Nautosphaeria), Nimbospora and Nereiospora
spore release into water, the outer region (all having tufts of hair-like appendages, but
ruptures and the appendages break through these lack the long unfurling appendages
(Read et al., 1993a). Sequence data places the forming sticky filaments), and any resemblance
genus in the Halosphaeriales in a sister group is therefore superficial. Molecular data show
to Panorbis viscosus and Halosarpheia marina no affinity between these genera (Fig. 41).
(Fig. 41) (Sakayaroj, 2005).
Oceanitis Kohlm., Rev. Mycol. 41: 193, 1977
1. Ascospores lacking hair-like appendages, sheath with ...................................................................... (4)
many folds, 22-20 (-31) × 12-16 (-18) μm................. ◙ O. scuticella Kohlm., Rev. Mycol. 41: 194,
................................................................. N. octonae
1. Ascospores with lateral tufts of hair-like appendages
1977 (Type species).
................................................................................. 2 ◙ O. cincinnatula (Shearer & J.L. Crane) J.
Dupont & E.B.G. Jones, Mycol. Res. in press,
2. Ascospores with one lateral group of appendages, 8- 2009.
10 μm ......................................................... .N. effusa Halosarpheia cincinnatula Shearer & J.L. Crane,
2. Ascospores with 2 or more lateral groups of Bot. Mar. 23: 613, 1980.
appendages, 18-27 × 8-12 μm.................N. bipolaris Falcatispora cincinnatula (Shearer & J.L. Crane)
K.L. Pang & E.B.G. Jones, Nova Hedw. 77: 14, 2003.
Nohea Kohlm. & Volkm.-Kohlm., Syst. Ascosalsum cincinnatulum (Shearer & J.L. Crane)
Ascomycetum 10: 121, 1991 ....................... (1) J. Campb., J.L. Anderson & Shearer, Mycologia 95: 546,
2003.
◙ N. umiumi Kohlm. & Volkm.-Kohlm., Syst.
◙ O. viscidula (Kohlm. & E. Kohlm.) J.
Ascomycetum 10: 122, 1991 (Type species).
Dupont & E. B.G. Jones, Mycol. Res. in press,
Ascomata subglobose, immersed or
2009.
superficial, ostiolate, papillate, coriaceous, Haligena viscidula Kohlm. & E. Kohlm., Nova
cream-coloured, solitary, neck 50-300 µm long, Hedw. 9: 92, 1965.
periphysate, catenophyses deliquescing, asci Halosarpheia viscidula (Kohlm. & E. Kohlm.)
clavate, pedunculate, unitunicate, thin-walled Shearer & J.L. Crane, Bot. Mar. 23: 608, 1980.
lacking an apical apparatus, deliquescing early, Falcatispora viscidula (Kohlm. & E.Kohlm.) K.L.
Pang & E.B.G. Jones, Nova Hedw. 77: 14, 2003.
ascospores ellipsoidal, 1-septate, hyaline, with Ascosalsum viscidulum (Kohlm. & E. Kohlm.) J.
two types of appendages. The first type are on Campb., J.L. Anderson & Shearer, Mycologia 95: 546,
one side of the spore, gelatinous and attached 2003.
to flat subapical pads, then unfurl to form long ◙ O. unicaudata (E.B.G. Jones & Le Camp.-
sticky filaments, the second type are subapical Als.) J. Dupont & E.B.G. Jones, Mycol. Res. in
comprising two bundles of fibres attached to press, 2009.
the spore wall. These appendages are Haligena unicaudata E.B.G. Jones & Camp.-Als.,
superficially similar to Naufragella, which also Nova Hedw. 19: 574, 1971.

101
 Halosarpheia unicaudata (E.B.G. Jones & genus Ascosalsum was erected to accommodate
Camp.-Als.) R.G. Johnson, E.B.G. Jones & S.T. Moss ex three species based on LSU rDNA phylogeny
Kohlm. & Volkm.-Kohlm., Bot. Mar. 34: 22, 1991.
Falcatispora unicaudata (E.B.G. Jones & Camp.- supported by morphological evidence
Als.) K.L. Pang & E.B.G. Jones, Nova Hedw. 77: 14, (Campbell et al., 2003). It differs from
2003. Halosarpheia in lacking catenophyses and
Ascosalsum unicaudatum (E.B.G. Jones & having cylindrical to falcate ascospores.
Camp.-Als.) J. Campb., J.L. Anderson & Shearer,
Mycologia 95: 550, 2003. 1. Ascospores with a single appendage........................ 2
Ascomata subglobose to ellipsoidal, on a 1. Ascospores with bipolar appendages, 38-89 × 3-7 μm,
thin hypostroma, ostiolate, epapillate, 5-16 septa predominantly 11-septate .........................
coriaceous, brown to dull orange, gregarious, ................................................................O. viscidula
paraphyses absent, asci clavate, unitunicate,
2. Ascospores 1-septate..............................O. scuticella
thin-walled, without an apical apparatus, 2. Ascospores 3 or more-septate .................................. 3
deliquescing, ascospores filiform to elongate
fusiform, 1-11-septate, hyaline, with bipolar 3. Ascospores 36-60 × 2.5-5 μm, 3-5-septate ................
appendages or at one end of the spore (Figs. ............................................................O. unicaudata
48a, d, f). Appendages initially closely 3. Ascospores 34-60 × 4-5 μm, 5-11-septate .................
adpressed to the spore wall, separating, and ..........................................................O. cincinnatula
uncoiling to form long thin narrow sticky
threads. A genus initially described with Ocostaspora E.B.G. Jones, R.G. Johnson &
needle-shaped, falcate, ascospores, 1-septate, S.T. Moss, Bot. Mar. 26: 353, 1983............. (1)
hyaline with an appendage at one end and not ◙ O. apilongissima E.B.G. Jones, R.G.
collected since its description on trawled Johnson & S.T. Moss, Bot. Mar. 26: 354, 1983
submerged wood at 3,975 m, in the Gulf of (Type species).
Angola, Atlantic Ocean (Kohlmeyer, 1977). Ascomata globose to subglobose, hyaline
A recent collection from submerged upper part dark brown or black, ostiolate,
wood at 1,000 m in the Pacific Ocean off papillate, membranous, immersed to partly
Vanuatu Islands has enabled further immersed, short necks lacking periphyses,
characterization of the genus (Dupont et al., solitary or gregarious, no paraphyses, asci
2009). Ascomata aggregated into a stroma, clavate to subcylindrical, short pedunculate,
with a thick peridium, 200 μm wide, composed unitunicate, thin-walled, no apical apparatus or
of polygonal cells with large lumina forming a pore, deliquescing early, ascospores 1-septate,
textura angularis, merging towards the center slightly constricted at the septum, hyaline, with
into flattened cells. Asci deliquesce early. The a single long appendage at each pole, and 6-8
cavity of the ascomata are filled with fusiform equatorial appendages. A monotypic genus,
ascospores, more or less grouped, 1-septate, that is widely distributed in temperate climates
hyaline, with a single polar appendage initially and occurs on submerged wood. It most closely
closely adpressed to the ascospore wall, but resembles Halosphaeria appendiculata with its
separating at maturity. Ascospore wall is two polar and equatorial appendaged ascospores
layered, a narrow episporium and a (Jones et al., 1983b). It differs in that the polar
mesosporium with an outer electron-dense appendages are much longer and easily become
zone, and an inner electron-transparent zone, fibrillar, features not found in H. appendiculata.
both layers forming the ascospore septum. The Molecular sequences of 18S and 28S confirm
polar appendage is composed of tightly coiled its placement in the Halosphaeriales and that is
filaments arising as outgrowths from the distantly related to H. appendiculata
mesosporium with a fibrous multilayered (Sakayaroj et al., unpublished data, see Fig. 41).
structure (Dupont et al., 2009).
Okeanomyces K.L. Pang & E.B.G. Jones, Bot.
Dupont et al. (2009), based on the
J. Linn. Soc. 146: 228, 2004 ........................ (1)
morphological similarity of O. scuticella to
◙ O. cucullatus (Kohlm.). K.L. Pang & E.B.G.
Ascosalsum species, and supported by
Jones, Bot. J. Linn. Soc. 146: 228, 2004 (Type
molecular evidence, transferred all Ascosalsum
species).
species to Oceanitis (Figs 41, 47, 55). The

102
 Fungal Diversity

Remispora cucullata Kohlm., Mycologia 56: 770, lacking an exosporium (Jones et al., 1984). A
1964. cold water species. Isolates were examined for
Halosphaeria cucullata (Kohlm.) Kohlm., Can. J.
Bot. 50: 1956, 1972. thier phylogenetic relationship. The LSU
Anamorph: Periconia prolifica rDNA phylogeny revealed that this genus is not
Anastasiou related to Halosphaeria (Kohlmeyer, 1972a),
Ascomata subglobose, ostiolate, papillate, but closely related to Ceriosporopsis tubulifera
coriaceous, immersed, brownish-black or and Marinospora species (Fig. 41).
brownish-red, solitary or gregarious,
catenophyses deliquescing, asci clavate, short Ophiodeira Kohlm. & Volkm.-Kohlm., Can. J.
pedunculate, unitunicate, thin-walled, without Bot. 66: 2062, 1988...................................... (1)
an apical apparatus, deliquescing early, ◙ O. monosemeia Kohlm. & Volkm.-Kohlm.,
ascospores cylindrical, 1-septate, slightly Can. J. Bot. 66: 2062, 1988 (Type species).
constricted at the septum, hyaline, with a single Ascomata ellipsoidal, immersed under a
cap-like or globose gelatinous appendage at thin black stroma, ostiolate, papillate, light
one spore apex, and deciduous. Frequently, brown, solitary or gregarious, necks long (60-
appendages not observed. A genus proposed by 300 µm long) and periphysate, asci clavate,
Pang et al. (2004b) to accommodate pedunculate, unitunicate, thin-walled,
Halosphaeria cucullata based on a recent deliquescing early, lacking an apical apparatus,
collection of the fungus in Malaysia. No ascospores ellipsoidal, 1-septate, not
ascospore appendage was observed in any of constricted at the septum, hyaline, with a single
the material examined but the characteristic polar appendage, initially hamate, separating
reddish ascomata and the production of the from the spore wall and eventually uncoiling to
anamorph Periconia prolifica from single form long, sticky filaments. Ophiodeira is a
ascospore isolates confirmed its identity. LSU monotypic genus that is reminiscent of
rDNA phylogeny revealed that this fungus is Halosarpheia species with its single polar
not related to Halosphaeria appendiculata, but appendage uncoiling to form sticky filaments.
groups with Thalespora appendiculata, but is It differs from the genera with unfurling polar
not monophyletic with it. They form a sister appendages in that the ascomata are formed
group to Ascosacculus (Fig. 55). beneath a thin stroma, has deliquescing asci,
absence of catenophyses and with only a single
Ondiniella E.B.G. Jones, R.G. Johnson & S.T. polar appendage (Kohlmeyer and Volkmann-
Moss, Bot. Mar. 27: 136, 1984 .................... (1) Kohlmeyer, 1988a). At the molecular level it
◙ O. torquata (Kohlm.) E.B.G. Jones, R.G. forms a sister clade with Halosarpheia-like
Johnson & S.T. Moss, Bot. Mar. 27: 136, 1984 species with a single polar appendage: O.
(Type species). scuticella, O. cincinnatula, O. unicaudata and
Halosphaeria torquata Kohlm., Nova Hedw. 2: O. viscidula with bipolar appendaged
311, 1060. ascospores (Dupont et al., 2009) (Figs 47, 55).
Ascomata elongate-cylindrical or The only feature separating it from these taxa is
subglobose, immersed or superficial, ostiolate, that the ascomata are formed beneath a thin
papillate, membranous, ranges from hyaline to stroma (applies also to O. scuticella). Tang et
pale brown, necks short and cylindrical, no al. (2007) have highlighted the congruence
periphyses, catenophyses deliquescing, asci between morphological and molecular
clavate to subfusiform, apiculate, pedunculate, classification schemes and this is becoming a
unitunicate, thin-walled, no apical apparatus, common observation in phylogenetic studies
deliquescing, ascospores broadly ellipsoidal, 1- (Binder et al., 2006).
septate, slightly constricted at the septum, Appendages such as those of O.
hyaline with appendages. Two types of scuticella and Ophiodeira monosemeia that do
appendages: spine-like polar appendages and not appear to readily uncoil in water, have been
an annulus-like equatorial appendage, arising observed for other taxa and may be attributed
as outgrowths of the episporium, with to their degree of maturation or the media they
amorphous material within the appendages and are mounted in. In Aniptodera salsuginosa, the
103
salinity of the water markedly affects the hyaline, smooth-walled, lacking a sheath or
unfurling of the polar appendages (Nakagiri appendages. Described from wood collected in
and Ito, 1994), while in a freshwater Thailand, it has clavate asci with a truncated
Halosarpheia species, amorphous material was thickened apex, a pore, the ascus plasmalemma
released, and connected to the spore by a fine is retracted and ascospores are 1-4-septate,
thread, prior to its organization into filaments hyaline, cylindrical and lacking appendages
(Jones, 2006). (Fig. 50) (Jones et al., 2006). This ascomycete
is morphologically similar to Lignincola nypae,
Panorbis J. Campb., J.L. Anderson & Shearer, but differs in the dimensions of ascomata, asci,
Mycologia 95: 544, 2003 ............................. (1) 1-4-septate ascospores and with a Humicola
◙ P. viscosus (I. Schmidt) J. Campb., J.L. anamorph. Phylogenetically LSU sequences
Anderson & Shearer, Mycologia 95: 544, 2003 place it in an unsupported clade with
(Type species). Antennospora, Cucullosporella, Halosarpheia
Halosphaeria viscosa I. Schmidt, Nat. and Lignincola tropica (Figs 41, 47, 55).
Naturschutz Mecklenburg 12: 70, 1974.
Halosarpheia viscosa (I. Schmidt) Shearer & J.L.
Crane, Bot. Mar. 23: 608, 1980.
Remispora Linder, Farlowia 1: 409, 1944... (8)
Ascomata globose, ostiolate, papillate, ◙ R. crispa Kohlm., Can. J. Bot. 59: 1317,
membranous, immersed or superficial, black, 1981.
neck with periphyses, catenophyses ◙ R. galerita Tubaki, Publ. Seto Mar. Biol. Lab.
deliquescing, asci clavate, unituicate, thin- 15: 362, 1967.
walled, deliquescing, with an apical pore, ◙ R. maritima Linder, Farlowia 1: 410, 1944
ascospores cylindrical to fusiform, 1-septate, (Type species).
Remispora lobata Höhnk, Veröeff. Inst.
hyaline, with bipolar appendages. Appendages Meeresforsch. Bremerhaven 3: 206, 1955.
closely adpressed to the spore wall, separating, Halosphaeria maritima (Linder) Kohlm., Can. J.
and uncoiling to form long sticky thread-like Bot. 50: 1956, 1972.
filaments. Panorbis viscosus resembles R. minuta E.B.G. Jones, K.L. Pang &
Natantispora retorquens morphologically Vrijmoed, Can. J. Bot. 82: 486, 2004.
although they are not related phylogenetically ◙ R. pilleata Kohlm., Nova Hedw. 6: 319,
(Campbell et al., 2003) (Fig. 47). However it is 1963.
morphologically not well delineated from Halosphaeria pilleata (Kohlm.) Kohlm., Can. J.
Natantispora, the key provided merely Bot. 50: 1957, 1972.
separating them on ascospore shape and ◙ R. quadri-remis (Höhnk) Kohlm., Nova
measurements: ellipsoid (less than 25 µm) in Hedw. 2: 332, 1960.
Palomyces quadri-remis Höhnk, Veröeff. Inst.
the former and dufoid-ellipsoid (over 25 µm) in Meeresforsch. Bremerhaven 3: 213, 1955.
the latter. More genera with polar unfurling Arenariomyces quadri-remis (Höhnk) Meyers,
appendages need to be sequenced to determine Mycologia 49: 505, 1957.
the validity of this genus. Halosphaeria quadri-remis (Höhnk) Kohlm., Can.
J. Bot. 50: 1957, 1972.
Pseudolignincola Chatmala & E.B.G. Jones, ◙ R. spitsbergenensis K.L. Pang and Vrijmoed,
Nova Hedw. 83: 225, 2006. ......................... (1) Mycologia, in press, 2009.
◙ P. siamensis Chatmala & E.B.G. Jones, ◙ R. stellata Kohlm., Nova Hedw. 2: 334, 1960.
Nova Hedw. 83: 226, 2006. Halosphaeria stellata (Kohlm.) Kohlm., Can. J.
Bot. 50: 1957, 1972.
Anamorph: Humicola siamensis
Ascomata globose, subglobose, ovoid,
Chatmala & E.B.G. Jones
immersed or superficial, ostiolate, papillate,
Ascomata globose, dark brown, deeply
coriaceous, hyaline to dark brown, solitary or
immersed in the wood, coriaceous, with a long
gregarious, catenophysate, necks periphysate,
(710-1100 µm) periphysate neck, solitary,
asci clavate broadly fusoid, pedunculate,
catenophyses present, asci clavate to slightly
apiculate, unitunicate, thin-walled, without an
cylindrical, long pedicellate, unitunicate, thin-
apical apparatus, deliquescing, ascospores 1-
walled, truncate at the apex with a refractive
septate, ellipsoid to rhomboidal, hyaline, with
thickening and retraction of the plasmalemma
polar appendages. Appendages morpho-
at the apex, ascospores cylindrical, 1-4-septate,
104
 Fungal Diversity

logically variable depending on the species appendage is more discrete (Manimohan et al.,
(Figs. 51, 52a-c, e, 53a). 1993b).
Molecular sequences indicate that
Remispora is polyphyletic with R. 1. Ascospores ellipsoidal ............................................. 2
spitsbergenensis, R. pilleata, R quadri-remis 1. Ascospores rhomboid, 24-36 × 12-20 µm..................
.................................................................. R. pilleata
and R. stellata forming a well supported clade,
while the type species, R. maritima (Fig. 41) 2. Ascospores with four or more appendages at each
forms sister group with R. pilleata and pole .......................................................................... 6
Carbosphaerella leptosphaerioides. Remispora 2. Ascospores with a single polar appendage............... 3
crispa and R. galerita are distantly placed (Fig.
41), the former grouping with Ocostaspora 3. Ascospores 18-30 (-32) × 8-13 µm, with wing-like
apilongissima and Morakotiella salina, while appendages............................................. R. maritima
3. Ascospore appendages variously shaped ................. 4
the latter forms a basal clade to the order with
Nautosphaeria cristaminuta and Haligena 4. Cap-like ascospore appendages with radiating strands,
elaterophora. Remispora minuta is a new 20-28 × 7-12 µm ....................................... R. galerita
species described from unidentified wood 4. Ascospore appendages moustache-like.................... 5
collected in Singapore and for which no
molecular data is available (Pang et al., 2004a). 5. Ascospores constricted at the septum, 17-26 × 6-10
The erection of new genera for R. crispa and R. μm..............................................................R. minuta
5. Ascospores not constricted at the septum, 22-34 × 8-
galerita are warranted, but further isolates are 12 µm...........................................................R. crispa
required to prove their monophyly.
Remispora was described by Barghoorn 6. Ascospores with 4 appendages at each apex............ 7
and Linder (1944) but all species were 6. Ascospores with 6 appendages at each apex, 24-30 ×
transferred to Halosphaeria by Kohlmeyer 8-12 µm......................................................R. stellata
(1972a). The genus is characterised by hyaline
to brown ascomata, with the peridium 7. Ascospores 20-28 × 7-13 µm, appendages long,
composed of one cell type, asci clavate that ribbon-like 19-24 × 3-6 µm long R. spitsbergenensis
7. Ascospores 18-20 (-34) × 8-12 µm, appendages
deliquesce early, and ascospores with polar
radiating, obclavate 12-21.5 × 2.5-4 µm ....................
appendages of variable morphology (Johnson .......................................................... R. quadri-remis
et al., 1984; Manimohan et al., 1993a). Initial
ultrastructural studies of R. maritima and R.
pilleata showed that the appendages were Saagaromyces K.L. Pang & E.B.G. Jones,
exosporic in origin and consisting of a fibrous Mycol. Progr. 2: 35, 2003 ............................ (3)
component in an amorphous matrix. ◙ S. abonnis (Kohlm.) K.L. Pang & E.B.G.
Fragmentation of the sheath giving rise to the Jones, Mycol. Progr. 2: 35, 2003.
wing-like polar appendages (Jones and Moss, Halosarpheia abonnis Kohlm., Mar. Ecol.
1978; Johnson et al., 1984). However, radiating (P.S.Z.N.I.) 5: 339, 1984.
appendages are found in R. quadri-remis and R. Littispora abonnis (Kohlm.) J. Campb., J.L.
stellata but the substructure was the same as in Anderson & Shearer, Mycologia 95: 549, 2003.
R. maritima and R. pilleata (Manimohan et al., ◙ S. glitra (J.L. Crane & Shearer) K.L. Pang &
1993a). In R. cispa, the appendages are closely E.B.G. Jones, Mycol. Progr. 2: 35, 2003.
Nais glitra J.L. Crane & Shearer, Trans. Br. Mcol.
adpressed to the spore wall and not wing-like Soc. 86: 509, 1986.
or radiating, but ultrastructurally are similar to ◙ S. ratnagiriensis (S.D. Patil & Borse) K.L.
R. stellata and R. quadri-remis. In these two Pang & E.B.G. Jones, Mycol. Progr. 2: 35,
species, appendages arise in a polar position 2003.
with radiating bundles of electron-dense stands Halosarpheia ratnagiriensis S.D. Patil & Borse,
exuded through pores in the episporium and Indian Botanical Reporter 1: 102, 1982.
embedded in an amorphous matrix. Similar Littispora ratnagiriensis (S.D. Patil & Borse) J.
observations were made for R. galerita, but Campb., J.L. Anderson & Shearer, Mycologia 95: 549,
2003.
differs from other species in that the polar

105
 Ascomata ellipsoidal, immersed, ostiolate, et al., 2004a). A new taxon described from
papillate, coriaceous, hyaline to brown, necks material collected on wood from a sandy beach
long (up to 1 mm), periphysate, catenophyses in southern China. This genus is characterized
present, asci clavate, long pedunculate, by its unique bipolar and quadri-equatorial
unitunicate, thin-walled, with or without apical appendages, which disintegrate when mounted
pore, persistent, retraction of the plasmalemma in seawater. The appendages are obclavate,
at the apex, ascospores ellipsoidal, 1-septate, tapered to a fine point, appear striated when
slightly constricted at the septum, hyaline, with released from the ascus but later become
or without polar appendages. Appendages large fibrillar (Pang et al., 2004a). The ascospores
if present, initially closely adpressed to the resemble those of Halosphaeria appendiculata
spore wall, separating, uncoiling to form long and Ocostaspora apilongissima with polar and
sticky filamentous threads (Fig. 44c). The four equatorial appendages. In O.
genus was erected to accommodate two species apilongissima appendages are striate, fragment
of Halosarpheia and Nais glitra (Fig. 43f), as into threads, but the polar appendages are much
they did not group within the Halosarpheia longer than the equatorial ones. In H.
sensu stricto clade, the type species of the appendiculata the appendages do not
genus (Pang et al., 2003b) (Fig. 55). breakdown but maintain their spoon-like
Morphologically they differ in the following morphology (Jones et al., 1984). The LSU
respects from Halosarpheia species: 1). Asci rDNA data confirms its placement in the
persistent, cylindrical-clavate, with a long stalk Halosphaeriales and is distantly related to
up to one third of the length of the ascus; 2). Halosphaeria appendiculata and Ocostaspora
Apical pore present in S. ratnagiriensis and S. apilongissima (Fig. 41).
glitra; and 3). Ascospores large, oval and
heavily guttulated. Therefore there is molecular Thalassogena Kohlm. & Volkm.-Kohlm., Syst.
and morphological evidence for their Ascomycetum 6: 223, 1987 ......................... (1)
segregation from the genus Halosarpheia. T. sphaerica Kohlm. & Volkm.-Kohlm., Syst.
(Pang and Jones, 2004). Ascomycetum 6: 225, 1987 (Type species).
A monotypic genus with cream-coloured
1. Ascospores with no appendages: 42-59 × 21-31 μm . ascomata, globose, immersed to superficial,
...................................................................... S. glitra
1. Ascospore with bipolar appendages......................... 2
ostiolate, papillate, coriaceous, with long necks
(230-570 μm), periphysate, with catenophyses,
2. Ascospores greater than 48 μm: 48-64 (-72) × 22-28 asci clavate, unitunicate, papillate, pedunculate,
μm ................................................... S. ratnagiriensis J-, flattened at the apex with an apical pore and
2. Ascospores shorter than 48 μm: 33-47 × 14-22 μm... hyaline subglobose ascospores without sheaths
..................................................................S. abonnis or appendages (Kohlmeyer and Volkmann-
Kohlmeyer, 1987a). This species has not been
Sablecola E.B.G. Jones, K.L. Pang & extensively collected and further studies are
Vrijmoed, Can. J. Bot. 82: 486, 2004 .......... (1) required to determine if it is correctly referred
◙ S. chinensis E.B.G. Jones, K.L. Pang & to the Halosphaeriales. Ascospores super-
Vrijmoed, Can. J. Bot. 82: 486, 2004 (Type ficially resemble Iwilsoniella rotunda, but the
species). species differs in ascomatal characteristics, the
Ascomata ellipsoidal to subglobose, latter with dark coloured ascomata, with short
immersed to partly immersed, yellow to brown, necks, asci lacking an apical pore and
ostiolate, papillate, necks short and periphysate, deliquescing early.
lacking catenophyses, asci clavate, pedunculate,
unitunicate, thin-walled, no apical apparatus, Thalespora Chatmala & E.B.G. Jones, Nova
persistent to deliquescing, ascospores Hedw. 83: 228, 2006.................................... (1)
ellipsoidal, hyaline, thin-walled, 1-septate. ◙ Th. appendiculata Chatmala & E.B.G. Jones,
With one appendage at each pole and four at Nova Hedw. 83: 229, 2006.
the equatorial septum (Fig. 53c). Appendages Ascomata ellipsoidal, immersed in wood,
flattened, attenuate and strap-like, with parallel ostiolate, papillate, coriaceous, unitunicate,
striations, later splitting into fine threads (Pang

106
 Fungal Diversity

a b c

Fig. 50. Pseudolignincola siamensis. a. Ascus with an apical pore (arrowed). b. Three-septate germinating ascospore. c,
d. Three-septate ascospores. Bars a-d = 20 μm.

thin-walled, light brown, solitary, paraphyses

absent, asci ellipsoidal, short pedunculate,
clavate, unitunicate, thin-walled, deliquescing
early, ascospores elongate-fusiform, rounded at
the pole, tapering toward the base, straight or
slightly curved, 1-septate, not constricted at the
septum, hyaline, with 2-4 terminal radial
appendages formed after release from the ascus
(Fig. 54). This species is phylogenetically close
to Okeanomyces cucullatus (Fig. 55), but they
differ in ascospore appendage morphology: a
single ephemeral mucilaginous drop/pad in the
latter but tetraradiate in Th. appendiculata and
formed after release from the ascomata (Fig. 54)
(Jones et al., 2006). Marine ascomycetes with
ascospores over 100 µm include Bathyascus,
Lindra and Lulworthia, the latter two genera
Fig 51. Remispora spitsbergenensis. Bar = 10 µm. are members of the Lulworthiales

107
while Bathyascus may not be correctly tapering, thick-walled, 1-4-septate, not
assigned to the Halosphaeriales (Jones, constricted at the septa, hyaline, bent double
unpublished data). and coiled within the ascus, no appendages or
sheath. An infrequently collected species
Tirispora E.B.G. Jones & Vrijmoed, Can. J. known from brown seaweeds, and further
Bot. 72: 1373, 1994...................................... (2) studies are required to establish its position in
◙ T. unicaudata E.B.G. Jones & Vrijmoed, the Halosphaeriales, as Sutherland (1915)
Can. J. Bot. 72: 1373, 1994 (Type species). referred it to the Hyponectriaceae. However,
Aniptodera indica Ananda & Sridhar, J. Envir. Rossman et al. (1999) excluded it from the
Biol. 22: 283, 2001. Hypocreales because of the long-necked
T. mandoviana V.V. Sarma & K.D. Hyde, ascomata, lack of apical paraphyses and
Aust. Mycol. 19: 52, 2000. blackening of the host thallus. We do not
Ascomata globose to subglobose, consider its placement in the Halosphaeriales
superficial, ostiolate, papillate, periphysate, as satisfactory, but is left here until collections
pale to dark brown, catenophyses present, asci are available for a molecular study.
clavate, short pedunculate, with a ring and an
apical plate, unitunicate, thick-walled apically Trichomaris Hibbits, G.C. Hughes & Sparks,
with retraction of the plasmalemma at the tip, Can. J. Bot. 59: 2123, 1981.......................... (1)
persistent, ascospores 1-septate, constricted at T. invadens Hibbits, G.C. Hughes & Sparks,
the septum, ellipsoid, hyaline with a single Can. J. Bot. 59: 2123, 1981 (Type species).
appendage at one pole (Fig. 44d). Initially Ascomata superficial on an extensive
adpressed to the spore wall but unfurls to form subiculum on crab carapace, thick-walled,
a long filamentous thread. Aniptodera indica, coriaceous, papilla lacking or short,
described by Ananda and Sridhar (2001), is aparaphysate, periphysate, asci elongate clavate,
reduced to synonymy with T. unicaudata. thin-walled, unitunicate, ascospores oblong to
Tirispora mandoviana was originally described ellipsoidal, 15-22 × 4.6-5 μm, 1-3- septate, bi-
from freshwater habitats (Sarma and Hyde, polar uncoiling appendages. Appendages
2000) but has recently been collected in Indian initially coiled around the ascospore, later
mangroves where salinity ranges from 1.1-34.1 uncoil and consist of two parts: a short
‰ (Maria and Sridhar, pers. comm.). proximal region is stiffer and a distal region
Phylogenetically T. unicaudata forms a which is flexible, sticky and stretching to a
sister group to Halosarpheia marina and great length (Hibbits et al., 1981). It differs
Panorbis viscosus, but with weak support (Fig. from those of the Halosarpheia complex in that
55). appendages are not hamate, adhering to the
ascospore wall, and later unfurling. A genus
1. Asci with retracted plasmalemma, an apical ring and
pore, catenophyses present, ascospores 24-32 × 8-12
requiring further study, at the molecular level,
μm ....................................................... T. unicaudata to determine whether it is correctly assigned to
1. No retraction of the plasmalemma but with an the Halosphaeriales.
indistinct apical pore, no catenophyses, ascospores
15-22 × 8-12 μm ............................... T. mandoviana Tunicatispora K.D. Hyde, Aust. Syst. Bot. 3:
712, 1990...................................................... (1)
Trailia G.K. Sutherl., Trans. Br. Mycol. Soc. 5: T. australiensis K.D. Hyde, Aust. Syst. Bot. 3:
149, 1915...................................................... (1) 712, 1990 (Type species).
T. ascophylli G.K. Sutherl., Trans. Br. Mycol. Buxetroldia bisaccata K.R.L. Petersen & Jørg.
Soc. 5: 149, 1915 (Type species). Koch, Mycol. Res. 101: 1526, 1997.
Ascomata subglobose, immersed in Ascomata globose to subglobose,
blackened area of algal host, ostiolate, immersed or semi-immersed, hyaline below to
papillate with long necks (140-450 µm long), light brown above, ostiolate, papillate,
hyaline, thin-walled, solitary or gregarious, no membranous, solitary or gregarious, necks
paraphyses, asci cylindrical to clavate, periphysate, catenophyses present deliquescing,
unitunicate, thin-walled, no apical apparatus, asci ovoid to clavate, unitunicate, thin-walled,
deliquescing early, ascospores filamentous, pedunculate, no apical apparatus,

108
 Fungal Diversity

a b

c d e

Fig. 52. Ascospores of Remispora and Naufragella species: a. R. pilleata. b. R. maritima. c, e R. stellata. d.
Naufragella spinibarbata. Bars a-e = 10 µm.

a b c

Fig. 53. Ascospores of: a. Remispora spitsbergenensis. b. Havispora longyearbyenensis. c. Sablecola chinensis.
Bars a-c = 10 μm.

109
 a b c d e

Fig. 54. Thalespora appendiculata. a-e. Stages in the development of ascospore polar appendages. Bars a-e = 20 μm.

deliquescing early, ascospores ellipsoidal, 1- 102 × 22-26 µm in T. australiensis (Hyde,

septate, slightly constricted at the septum, 1990). Thus the differences between the two
hyaline, with appendages. Two types of genera/species, revolve around the degree of
appendages: a thin skin-like sheath with a pore separation of the sheath from the ascospore
at the apex from which a viscous cap emerges. wall. Petersen and Koch (1997) show the
This cap-like appendage slowly unfurls to form sheath bulging away from the ascospore wall.
thread like appendages. This has also been demonstrated at the TEM
We regard Buxetroldia bisaccata as a level by McKeown et al. (1996) who examined
synonym of this species as the features collections of the fungus (made by Petersen)
separating them are not sufficient for a new from Denmark. Similarities in ascospore
genus. This species differs from other genera appendage morphology, especially the
with unfurling polar appendages in that the thickness of the basal part and its slowness to
ascospore wall comprises three layers: unfurl, are found in B. bisaccata, Halosarpheia
mesosporium, episporium and exosporium. The trullifera and T. australiensis. All possess a
latter has been shown by McKeown et al. sheath varying in its degree of separation from
(1996) to be loosely attached to the episporium, the spore wall and in its degree of persistence.
except at the central septum. The appendages The appendages in these species are cap-like,
comprise electron-dense fibrillar material, rather than hamate, initially amorphous and
within which there are tripartite strands similar only later unraveling to form extremely fine
to the episporium. The basal part of the thread-like. There is evidence that
appendage is stoute and arises through a Halosarpheia fibrosa may also have an
discontinuity in the exosporium. This differs exosporic sheath (Pang et al., 2006). The
from the porefields reported for Halosarpheia taxonomic assignment of these taxa needs to be
species (Alias et al., 2001). confirmed by a molecular study. Abdel-Wahab
Ascospore measurements of T. et al. (2001b) have shown that H. fibrosa, H.
australiensis overlap with those of Buxetroldia trullifera and H. unicellularis form a well
bisaccata, although the asci are greater in supported clade within the Halosphaeriales.
length (125-162 × 32-45 µm) as opposed to 70-

110
 Fungal Diversity

Fig. 55. Taxonomic position of the new genera Thalespora and Pseudolignincola.

111
 SORDARIOMYCETIDAE hyaline deliquescent, asci clavate, truncate,
long pedunculate, apical refractive non-
1. Diaporthales amyloid ring, ascospores allantoid, unicellular,
2. Chaetosphaeriales pale yellow to pale brown, 8-128 per ascus, no
3. Ophiostomatales appendages or sheaths. A largely terrestrial
genus, two are known from mangroves: C.
1. Generally stromatic, ascomata perithecial, long- halosarciicola on Halosarcia halocnemoides in
necked, interascal tissue absent, asci thick-walled, J-
apical ring, ascospores varied no germ slit,
Australia and C. mangrovei on Kandelia
anamorphic coelomycete...................... Diaporthales candel in Hong Kong. A third species C.
1. Ascomata not stromatic............................................ 2 suaedicola has been described from the salt
marsh plant Suaeda fruticosa (Inderbitzin et al.,
2. Paraphyses filiform and septate, ascomata superficial 1999) with ascospores measuring 5-7 × ca. 1
or immersed, perithecial, ostiolate or nonostiolate,
asci subglobose, clavate or cylindrical, thin-walled,
µm.
ascospores one-celled, hyaline or pigmented, with
1. Asci 8 spored, ascospores hyaline, smooth walled, 4-
germ slit, anamorphs phialidic .... Chaetosphaeriales
2. Interascal tissue absent or thin-walled and 7 × 1.5-2 μm ...................Cryptovalsa halosarciicola
1. Asci 64 spored, ascospores pale yellow-pale brown,
inconspicuous........................................................... 3
verrucose, 5.6-11 (-18) × 1.6-3.6 μm.........................
3. Ascomata perithecial, thick-walled, asci cylindrical ............................................ Cryptovalsa mangrovei
or clavate, ascospores with at least one dark cell with
germ pore, appendaged, generally lacking anamorphs Diaporthe Nitschke, Pyrenomycetes Germanici:
................................................................Sordariales 240, 1870...................................................... (1)
3. Ascomata perithecial rarely cleistothecial, thin- D. salsuginosa Vrijmoed, K.D. Hyde & E.B.G.
walled, long necked, ostiolar setae, asci small, Jones, Mycol. Res. 98: 699, 1994.
evanescent, formed in chains, ascospores small,
unicellular, appendaged or with eccentric wall
Ascomata in a group within a wide
thickening, anamorphs hyphomycetes ....................... spreading blackened zone on the wood,
.......................................................Ophiostomatales individual ascomata subglobose, immersed,
brown to black, coriaceous, with short necks
1. DIAPORTHALES periphysate, penetrating the host surface,
paraphyses hypha-like, filiform, tapering
1. Ascospores with appendages..................... Diaporthe
1. Ascospores lacking appendages or sheath ............... 2
distally, septate and hyaline, asci cylindrical,
unitunicate, thin-walled, short pedunculate,
2. Ascospores septate ............................... Hypophloeda flattened apex, J- subapical ring, ascospores 1-
2. Ascospores unicellular ............................................. 3 septate, not constricted at the septum, hyaline,
smooth walled, with a mucilaginous pad at
3. Ascospores allantoid, pale yellow to brown, small
(range 4-18 × 1.5-3.5 µm).......................Cryptovalsa
each pole which swells in water and is sticky.
3. Ascospores naviculoid, hyaline, (25) 30-35 (-40) × The only Diaporthe species known from
(6-) 8-10 (-14) µm............................Argentinomyces marine habitats, occurring on intertidal
mangrove wood, but no anamorph has been
Valsaceae linked to this species, although marine
Phomopsis spp. are known (Vrijmoed et al.,
Cryptovalsa Ces. & De Not. ex Fuckel, Jahrb. 1994). Ascospores are distinctive with a small
nassauischen Vereins Naturk., 23-24: 212, mucilaginous pad at each end, which swells in
1870.............................................................. (2) water and attaches the spores to substrata.
C. halosarciicola K.D. Hyde, Mycol. Res. 97:
799, 1993. Gnomonia Ces. & De Not., Soc. crittog Ital. 1:
C. mangrovei Abdel-Wahab & Inderb., Mycol. 231, 1863...................................................... (1)
Res. 103: 1628, 1999. G. salina E.B.G. Jones, Trans. Br. Mycol. Soc.
Ascomata in groups in a stroma, 45: 107, 1962.
immersed becoming erumpent, ectostroma This is a doubtful Gnomonia species and
effuse, ascomata subglobose to broadly known only from the type collection. It is
ellipsoidal, ostiolate, papillate, necks probably best referred to Halosarpheia (e.g. H.
periphysate, coriaceous, paraphyses septate trullifera). Clearly it does not belong in

112
 Fungal Diversity

Gnomonia because of ascus and ascospore

morphology and spore dimensions. Nomen a b
rejectum.

Melanconidaceae
Hypophloeda K.D. Hyde & E.B.G. Jones,
Trans. Mycol. Soc. Jpn. 30: 61, 1989 .......... (1)
H. rhizospora K.D. Hyde & E.B.G. Jones,
Trans. Mycol. Soc. Jpn. 30: 62, 1989 (Type c
species).
Ascomata solitary or gregarious; slightly
stromatic; periphysate; asci clavate, thin-walled,
with a refractive non-amyloid apical thickening,
persistent, with a tapering stalk; paraphyses Fig. 56. Argentinomyces navisculisporus. a. Clavate
ascus. b-c. Ascospores (After Peña and Arambari,
numerous, septate and broad; ascospores are
1997b). Bars a-c = 10 µm.
hyaline, cylindrical, 0-3-septate with no
appendages (Hyde and Jones, 1989b). 2. Ascospores 1-septate, apical cell dark brown to black
A monotypic genus that is infrequently .................................................................... Zopfiella
reported, occurring on mangrove prop roots 2. Ascospores 3-septate................................................ 3
with the ascomata just below the surface of the
bark. A molecular study is required to establish 3. Ascospores hyaline, with bipolar, semiglobose cap-
like polar appendage .............................Abyssomyces
its true taxonomic position.
3. Ascospores with brown cells............ Biconiosporella
3. Ascospores versicolorus............................Savoryella
DIAPORTHALES incertae sedis
Lasiosphaeriaceae
Argentinomyces N.I. Peña & Aramb., Biconiosporella Schaumann, Veröff. Inst.
Mycotaxon 65: 333, 1997 ............................ (1) Meereforsch. Bremerhaven 14: 24, 1972..... (1)
A. naviculisporus N.I. Peña & Aramb., B. corniculata Schaumann, Veröff. Inst.
Mycotaxon 65: 333, 1997 (Type species). Meereforsch. Bremerhaven 14: 24, 1972 (Type
Ascomata subglobose, immersed, species).
ostiolate, papillate, periphysate, membranous, Ascomata ovoid, pyriform, flask-shaped,
cream-coloured, solitary sometimes gregarious, superficial or immersed, ostiolate, papillate,
paraphyses absent, asci clavate short coriaceous, brown to black, solitary or
pedunculate, unitunicate, thin-walled, gregarious, periphysate, catenophyses present,
thickened at the apex, without a pore, persistent, asci cylindrical, short pedunculate, thin-walled,
ascospores ellipsoidal, 0-3-septate, slightly unitunicate, persistent, with an apical plate,
constricted at the septa, hyaline, without a ascospores biconical, unequally 3-septate,
sheath or appendages (Peña and Arambari, thick-walled, slightly constricted at the septa,
1997a) (Fig. 56). Although it has similarities dark brown, no sheath or appendages
with the Halosphaeriales, it clearly does not (Schaumann, 1972). Infrequently reported and
belong in that order, because of the thick- no recent taxonomic study on this species,
walled ascus apex, and persistent asci. It also often found on heavily decayed wood
shows similarities to the mangrove species associated with sand (Jones et al., 2005).
Rhizophila marina and Marinosphaera
mangrovei. Chaetomiaceae
Chaetomium Kunze, Mykologische Hefte
2. SORDARIALES (Leipzig) 1: 15, 1817.................................... (1)
1. Ascospores unicellular ..........................Chaetomium Ch. heteropilum N.J. Artemczuk Mykol.
1. Ascospores 2-3-septate ............................................ 2 Fitopatol. 14: 93, 1980.

113
 Ascomata globose, superficial, brown to 1. Ascospores with subapical germ pore, basal cell
fuscous, ostiolate, epapillate, with terminal broadly cylindrical, 16-22 × 10-13 µm ....... Z. latipes
1. Ascospores with apical pore, basal cell elongate
sterile hairs variously ornamented, straight cylindrical, 14-20 × 10-14 µm ...................Z. marina
curved or coiled, no paraphyses, asci clavate,
thin-walled, unitunicate, hyaline, long
SORDARIALES incertae sedis
pedunculate, no apical apparatus, deliquescing
early, ascospores ovoid, pip-like, and olive
brown. Isolated from sediments in the Black Abyssomyces Kohlm., Ber. Dtsch. Bot. Ges. 83:
Sea, ascomata with spine-like appendages that 505, 1970...................................................... (1)
are slightly bulbous at the base, longer hair-like A. hydrozoicus Kohlm., Ber. Dtsch. Bot. Ges.
appendages verruculose, ascospores 12.5-16 × 83: 505, 1970 (Type species).
8-9.8 μm. Ascomata subglobose or pyriform,
Chaetomium species are frequently noted superficial, ostiolate, papillate, coriaceous,
from marine habitats but whether they are true light brown, setose, solitary, short necks
marine species is not clear. For example, periphysate, asci subcylindrical or fusiform,
Chaetomium ramipilosum was isolated from short pedunculate, unitunicate, thin-walled, no
Ammophila arenaria plant material in a sand apical apparatus, persistent, ascospores
dune system and sporulated well on seawater subcylindrical, straight or slightly curved, 3-
media (Schaumann, 1973b), while Jones (1962) septate, not constricted at the septa, hyaline,
reported Chaetomium erectum and Ch. appendages gelatinous cap-like, at each end of
globosum on submerged test blocks exposed in the spore. Abyssomyces was collected at depths
the sea at Langstone harbour. of 631 and 641 m on hydrozoan, hydrorhiza
and hydrocaulon, attached to stony corals.
Zopfiella G. Winter, Rabenh. Krypt.-Fl. Ed2 (Kohlmeyer and Volkmann-Kohlmeyer,
1(2): 56, 1884............................................... (2) 2003b). Few collections are available for study
◙ Z. latipes (N. Lundq.) Malloch & Cain, Can. and the genus is in need of further study to
J. Bot. 49: 876, 1971. determine its phylogenetic relationship.
Tripterospora latipes Lundq., Bot. Not. 122: 592,
1969.
Anamorph: Humicola-like. Savoryella E.B.G. Jones & R.A. Eaton, Trans.
Z. marina Furuya & Udagawa, J. Jap. Bot. 50: Br. Mycol. Soc. 52: 161, 1969 ..................... (5)
249, 1975. ◙ S. appendiculata K.D. Hyde & E.B.G. Jones,
Anamorph: Humicola-like. Bot. Mar. 35: 89, 1992.
Ascomata globose to subglobose, ◙ S. lignicola E.B.G. Jones & R.A. Eaton,
immersed but usually superficial, lacking an Trans. Br. Mycol. Soc. 52: 161, 1969 (Type
ostiole, coriaceous, dark brown to black, species).
covered with septate branched hairs, solitary, ◙ S. longispora E.B.G. Jones & K.D. Hyde,
paraphyses present deliquescing early, asci Bot. Mar. 35: 84, 1992.
clavate, short pedunculate, unitunicate, with an S. melanospora Abdel-Wahab & E.B.G. Jones,
apical ring, ascospores ellipsoidal, 1-septate, Mycoscience 41: 387, 2000.
large apical olivaceous to dark brown cell and a ◙ S. paucispora (Cribb & J.W. Cribb) Jørg.
smaller hyaline basal cell, lacking appendages Koch, Nordic. J. Bot. 2: 169, 1982.
or a sheath. Both Z. latipes and Z. marina have Leptosphaeria paucispora Cribb & J.W. Cribb,
been reported with Humicola-like anamorphs Pap. Dept. Bot., Univ. Qd. 4: 41, 1960.
and conidia produced on peg-like structures on Ascomata subglobose or ellipsoidal,
undifferentiated hyphae, respectively (Guarro immersed to superficial, ostiolate, papillate,
et al., 1999) and widely distributed. Zopfiella membranous, pale brown to dark brown, necks
latipes has been recorded on Nypa fruticans in with periphyses, paraphyses deliquescing early,
Thailand at salinities of 5-21 ‰ (Pilantanapak solitary or gregarious, asci elongate to
et al., 2005) and on other seagrasses (Sakayaroj, cylindrical or clavate, short pedunculate,
unpublished data).

114
 Fungal Diversity

unitunicate, thin-walled, persistent with an mycetes until wider analysis of taxa and genes.
apical pore, ascospores 3-septate, constricted at While there is no known anamorph for
the septa, versicoloured, apical cells hyaline, Savoryella, Ascotaiwania has two: Helicoon
central cells brown, mucilaginous sheath farinosum (A. hughesii), and Monotosporella
around central cells, apical appendages in S. (A. sawadae, A. mitriformis) (Ranghoo and
appendiculata (Read et al., 1993b) (Fig. 57). Hyde, 1998; Sivichai et al., 1998; Boonyuen,
This genus has some similarities with members pers. comm.).
of the Halosphaeriales, in particular the nature
of the paraphyses and lack or poorly developed 1. Asci with 2 ascospores, 36-60 × 12-16.5 µm, on
apical pore to the ascus (Jones and Hyde, 1992). mangrove wood....................................S. paucispora
1. Asci with 8 ascospores............................................. 2
Ten species have been assigned to the genus,
from freshwater or marine habitats, with S. 2. Ascospores appendaged, 26-35 × 11-16 µm, on wood
appendiculata and S. melanospora found on ........................................................S. appendiculata
driftwood associated with sand (Hyde and 2. Ascospores not appendaged ..................................... 3
Jones, 1992b; Abdel-Wahab and Jones, 2000)
3. Ascospores wider than 15 µm, 32-45 × 15-18 µm, on
(Fig. 57). wood associated with sand................S. melanospora
The genus has been referred to various 3. Ascospores narrower than 15 µm, on wood............. 4
higher taxa: Ascomycetes incertae sedis
(Kohlmeyer, 1986b), Tripterosporaceae or 4. Ascospores 24-36 × 8-12 µm...................S. lignicola
Lasiosphaeriaceae in the Sordariales (Jones and 4. Ascospores 33.5-46.5 × 7.5-12 µm.......S. longispora
Hyde, 1992), Halosphaeriales (Barr, 1990a)
and Hypocreales incertae sedis (Cai et al.,
2006). Savoryella superficially resembles
Ascotaiwania both with versicoloured
ascospores and their occurrence in aquatic
habitats, but differ in the elaborate apical ascus
apparatus of Ascotaiwania, and its
Monotosporella and Helicoon farinosum
anamorphs (Chang et al., 1998; Sivichai et al.,
1998; Cai et al., 2006). A molecular study has
failed to resolve the taxonomic position of
Ascotaiwania (Ranghoo et al., 1999) with Cai
et al. (2006) referring it to the Hypocreales
incertae sedis. Their aquatic or intertidal Fig. 57. Savoryella melanospora. Ascospore. Bar = 10
μm.
habitat may have resulted in adaptation to these
environments with consequent reduction of
2. CHAETOSPHAERIALES
features of phyletic significance that may have
indicated their natural affinities. Therefore,
Chaetosphaeriaceae
assignment of Savoryella species to a higher
Chaetosphaeria Select. fung. carpol. (Paris) 2:
taxon on morphological criteria is difficult and
252, 1863...................................................... (1)
warrants molecular studies to resolve its
C. chaetosa Kohlm., Nova Hedw. 6: 307, 1963.
taxonomic position (Read et al., 1993b).
Ascomata subglobose or pyriform,
A recent molecular study of Savoryella
immersed or partly immersed, ostiolate,
and Ascotaiwania indicates they are closely
paplliate, subcoriaceous, dark brown to black,
related, but are not monophyletic. They do not
solitary or gregarious, necks periphysate,
group with any of the major taxa (Boonyuen,
paraphyses septate, ramose, asci cylindrical to
pers. comm.), but form an un-named clade
clavate, long pedunculate, unitunicate, thin-
distantly placed from the Halosphaeriales,
walled, persistent, apical apparatus present,
Sordariales, Xylariales and Hypocreales
ascospores fusiform or elongate-fusiform, 3-
incertae sedis. It is best referred to
septate, constricted at the septa, hyaline, with
Hypocreomycetidae incertae sedis, Sordario-
115
polar and equatorial appendages formed by Key to xylariaceous and xylariaceous
fragmentation of a sheath. The only marine incertae sedis species:
species of a genus with 20 or more terrestrial 1. Ascospores hyaline .................................................. 2
species growing on wood, bark and leaves, and 1. Ascospores brown or black ...................................... 9
with a variety of anamorphs (Kirk et al., 2001; 2. Ascospores apiosporous............ Apioclypea nypicola
Réblová and Seifert, 2008). 2. Ascospores not apiosporous..................................... 3
Chaetosphaeria chaetosa differs in a
number of respects from other species in the 3. Ascospores 0-septate................................................ 4
genus, primarily in ascospores with both polar 3. Ascospores septate ................................................... 6
and equatorial appendages, formed by 4. Ascospores with a single or bipolar caps or
fragmentation of an exosporic sheath, while no appendages............................................................... 5
anamorph have been reported for it (Jones et 4. Ascospores surrounded by a mucilaginous sheath... 7
al., 1983c). Thus it differs from other
Chaetosphaeria species which possess a wide 5. Ascospores filamentous, ascomata superficial under a
stroma ..................................................... Linocarpon
range of phialidic anamorphs. Ascospores bear 5. Ascospores ellipsoidal, ascomata beneath host
a resemblance to Corollospora species, but surface....................................... Fasciatispora nypae
differ in lacking polar spines (Jones et al.,
1983a). Although the genus has been 6. Ascospores cymbiform, 34-44 × 7-15 µm, on
extensively studied at the morphological and Phragmites .............................Phragmitensis marina
6. Ascospores ellipsoid, 17-26 × 5-7 µm on Nypa
molecular level, Ch. chaetosa has not been ........................................... Frondicola tunitricuspis
examined (Réblova, 1999; Huhndorf et al.,
2004; Réblová et al., 1999; Réblová and Seifert, 7. Ascospores 0-1-septate, filliform, on Nypa.............. 8
2008). Its phylogenetic position needs to be 7. Ascospores 1-septate elongate, 24-36 × 3-4.5 µm, on
confirmed by sequence data. mangrove leaves ................... Lanceispora amphibia

8. Ascospores 70-120 × 2-2.8 µm, ascomata deeply

OPHIOSTOMATALES immersed in Nypa ..............................Neolinocarpon
8. Ascospores 74-113 × 3-6 µm, ascomata not deeply
Lanspora K.D. Hyde & E.B.G. Jones, Can. J. immersed in Nypa ......................................Oxydothis
Bot. 64: 1581, 1986...................................... (1)
9. Ascospores 0-septate.............................................. 10
◙ L. coronata K.D. Hyde & E.B.G. Jones, Can. 9. Ascospores 1-septate.............................................. 17
J. Bot. 64: 1581, 1986 (Type species). 9. Ascospores 13-17-septate .....................Pedumispora
Ascomata globose to subglobose,
coriaceous, immersed to partly immersed, 10. Ascospores with appendages............................... 11
solitary or gregarious, neck short, cylindrical, 10. Ascospores with a sheath .................................... 12
10. Ascospores lacking appendages or a sheath........ 13
periphysate, paraphyses and catenophyses
absent, asci cylindrical to oblong-ventricose, 11. Ascospores 20-33 × 8-12 µm, single polar
unitunicate, thin-walled, short pedunculate, appendage appendage, on mangrove substrata........
ascospores ellipsoidal, unicellular, hyaline, .................................................Adomia avicenniae
with longitudinal wall striations, with a crown 11. Ascospores 6.5-16 × 2-8 µm, single or bipolar
of appendages at both ends. Appendages appendages ...................................... Phomatospora
crown-like, radiating, delicate, subgelatinous
12. Ascospores 16-20 × 4-6 µm, prominent sheath
and formed by fragmentation of an exo- drawn out to form polar appendages ......... Nipicola
sporium. A genus tentatively assigned by Hyde 12. Ascospores occasionally with a sheath ...................
and Jones (1986b) to the Halosphaeriales, ........................................................ Anthostomella
however, preliminary molecular data suggests
an affinitiy to the Ophiostomatales (Schoch, 13. Ascospores lacking germ slits ............................. 14
13. Ascospores with germ slits.................................. 15
pers. comm.).
14. Ascospores 6.5-8 × 1.2-16 µm, hyaline to pale
XYLARIOMYCETIDAE yellow, ascomata immersed in a stroma, on
mangrove wood.............................Eutypella naqsii
1. XYLARIALES 14. Ascospores 6-12 × 2.28 µm, immersed in
mangrove wood.....................Eutypa bathurstensis

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 Fungal Diversity

15. Ascomata immersed in a stroma, ascospores pale surrounded by a wide mucilaginous sheath.
brown to golden yellow to reddish-brown Arecophila nypae differs from other species in
............................................................... Astrocystis
15. Ascomata immersed in wood, ascospores 10-18 × having ascospores with rounded cells and
4-7.5 µm, with or without a sheath, or basal cell .... occurs on the brackish water palm Nypa
......................................................... Anthostomella fruticans. Arecophila is similar to another
15. Ascomata superficial on mangrove wood ........... 16 marine ascomycete, Atrotorquata but differs in
lacking an ascospore germ slit (Hyde, 1996).
16. Ascospores 18-28 × 7.5-13 µm, on various
mangrove wood................Halorosellinia oceanica Atrotorquata Kohlm. & Volkm.-Kohlm., Syst.
16. Ascospores 9-12 × 5-6 µm, on Kandelia candel
wood......................................... Nemania maritima
Ascomycetum 12: 8, 1993 ........................... (1)
A. lineata Kohlm. & Volkm.-Kohlm., Syst.
17. Ascospores with a sheath .................................... 18 Ascomycetum 12: 8, 1993 (Type species).
17. Ascospores lacking a sheath, 10-16 × 5-7 µm Ascomata subglobose, immersed,
....................................... Crytosphaeria mangrovei ostiolate, papillate with a long periphysate neck
18. Ascospores 17-23 × 7-9 µm, lacerate germ pore at
(140-200 µm), coriaceous, dark brown, solitary,
each pole, on Juncus ..... Ommatomyces coronatus paraphyses, simple or branched, septate, asci
18. Ascospores 19-26 × 7-8 µm, on Nypa..................... cylindrical, short pedunculate, unitunicate,
.................................................... Arecophila nypae thin-walled, persistent, with an apical apparatus,
18. Ascospores 23-31 × 7.4-10.5 µm on Juncus J+, ascospores ellipsoidal, 1-septate, slightly
............................................................Atrotorquata
constricted at the septum, brown, striate and
with 5-7 germ slits at each apex, with a
1. Cainiaceae
gelatinous sheath. A common and host-specific
Arecophila K.D. Hyde, Nova Hedw. 63: 82,
ascomycete on Juncus roemerianus, but may
1996.............................................................. (1)
not be obligately marine but occurs with other
A. nypae K.D. Hyde, Nova Hedw. 63: 95, 1996.
marine organisms (Kohlmeyer and Volkmann-
Ascomata immersed, subglobose,
Kohlmeyer, 1993c). It was referred to the
ostiolate, papillate, coriaceous, brown to black,
Amphis-phaeriaceae, but we prefer its
solitary to gregarious, necks periphysate,
assignment in the Cainiaceae (Kang et al.,
paraphyses hypha-like, filiform, septate and
1999a, b).
branched, asci cylindrical to clavate, short
pedunculate, unitunicate, thin-walled, J+ 2. Clypeosphaeriaceae
amyloid ring, persistent, ascospores ellipsoidal, Apioclypea K.D. Hyde, J. Linn. Soc., Bot. 116:
1-septate, brown, slightly constricted at the 316, 1994...................................................... (1)
septum, with a mucilaginous sheath. This is a A. nypicola K.D. Hyde, J. Fröhlich & Joanne E.
genus introduced by Hyde (1996) to Taylor, Sydowia 50: 36, 1998.
accommodate ascomycetes collected on palms, Ascomata subglobose, immersed
while others are known from dead culms of underneath a weakly developed clypeus,
bamboo (Umali et al., 1999). membranous, ostiolate, brown, solitary or
Kang et al. (1999a) placed the genus in gregarious, periphysate, paraphyses hypha-like,
the Cainiaceae based on molecular evidence, septate, numerous embedded in a gelatinous
formerly refered to the Amphisphaeriaceae. matrix, asci cylindrical, pedunculate, persistent,
Arecophila differs from Amphisphaeria in with a J+ apical apparatus, ascospores
having deeply immersed, subglobose to ellipsoidal, hyaline, apiosporus, with a
lenticular ascomata, with a peridium of angular mucilaginous sheath. A genus erected to
cells with a small or lacking a clypeus. accommodate unitunicate ascomycetes with
Arecophila species develop within the host ascomata immersed under a clypeus, appearing
tissue beneath a clypeus (or lacking), ascomata as minute blackened ostiolar dots (Hyde et al.,
often lenticular, paraphyses are hyphal-like, 1998). Ascospores form orange ooze around
asci unitunicate, cylindrical with an apical J+ ostioles. Referred to the Hyponectriaceae
ring and brown bicellular ascospores that are (Hyde, 1994b) and Clypeosphaeriaceae (Hyde
verrucose or striated. Ascospores are et al., 1999a), sequence data confirms its

117
position in the latter family (Kang et al., Ascomata globose to subglobose,
1999c,d). immersed, coriaceous, ostiolate, papillate,
brown to black, neck periphysate, thin
Ommatomyces Kohlm., Volkm.-Kohlm. & O.E. entostroma under the periderm, paraphyses,
Erikss., Mycologia 87: 538, 1995 ................ (1) wide, branched, hypha-like, asci cylindrical,
O. coronatus Kohlm., Volkm.-Kohlm. & O.E. unitunicate, thin-walled, long pedunculate,
Erikss., Mycologia 87: 538, 1995 (Type with an apical apparatus, J- ring, ascospores
species). allantoid, light brown, unicellular, no sheath or
Ascomata broadly ovate, immersed, appendages. An ascomycete saprobic on
ostiolate, subcarbonaceous, black, short papilla, intertidal wood of Rhizophora apiculata, with
periphysate, paraphyses, septate, unbranched, large ascomata (325-520 µm high, 650-910 µm
thin-walled, asci cylindrical, short pedunculate, wide), immersed singly within the bark and
unitunicate, with an apical apparatus consisting covered by the periderm. Molecular data
of several rings, J+, ascospores ellipsoidal, 1- required to determine if this species would be
septate, slightly constricted at the septum, better assigned to the genus Eutypa.
brown, thick-walled, with an a lacerate germ
pore at each end, with subglobose gelatinous Eutypa Tul. & C. Tul., Select. fung. carpol. 2:
and evanescent caps and a thin sheath around 52, 1863........................................................ (1)
the spore. Kohlmeyer et al. (1995c) assigned E. bathurstensis K.D. Hyde & Rappaz, Mycol.
the genus to the Amphisphaeriaceae, but Kirk Res. 97: 861, 1993.
et al. (2001) place it in Xylariaceae. It differs Anamorph: coelomycetous
from the previous family by its broadly Ascomata deeply immersed in the wood,
ellipsoidal ascospores with elaborate lacerate entostroma poorly developed, spherical to
germ pores, gelatinous spore caps and sheath. flattened, ostiole poorly developed, neck 120-
The species has a complex ascospore wall 200 µm periphysate, black, paraphyses
comprising a hyaline outer-wall and brown numerous, persistent, asci clavate, long
inner wall (which the authors refer to as pedunculate, apical apparatus J-, ascospores
perispore and endospore respectively, terms allantoid, olive-brown, no sheath or
that are not defined) and a characteristic apical appendages. This species differs from other
lacerate pore at each end. Ascospores have species in having an ascomatal wall that is
apical caps at each pole and a narrow three layered, the central one thick and hyaline
gelatinous sheath. (Hyde and Rappaz, 1993). Anamorph produced
in nature and in culture, the conidia produced
3. Diatrypaceae holoblastically, filiform, curved, hyaline with a
flattened base, 10-33 × 1.5-2 µm.
None of the marine Diatrypaceae has
Eutypella (Nitschke) Sacc., Atti Soc. Venet.-
been subject to a molecular study, but
Trento. Sc. Nat., Padova 4: 80, 1875 ........... (1)
terrestrial species have been examined by
E. naqsii K.D. Hyde, Mycol. Res. 99: 1462,
Acero et al. (2004) and Carmarán et al. (2006)
1995.
and includes 10 genera. The genera Eutypa,
Ascomata formed beneath a raised,
Cryptosphaeria, Diatrype, Eutypella were
blackened crust-like stroma on the host surface,
considered polyphyletic suggesting a revision
ascomata globose to subglobose, membranous,
of the family was required (Acero et al., 2004),
black, ostiolate, neck periphysate, paraphyses
while polysporous asci in the family appears to
hypha-like, sparse, septate, hyaline, and
have evolved several times (Cannon and Kirk,
tapering distally, asci cylindrical-clavate, long
2007).
pedunculate, unituicate, thin-walled, apical
apparatus J-, ascospores allantoid, unicellular,
Cryptosphaeria Ces. & De Not., Comm. Soc.
straight or curved, hyaline to pale brown, no
crittog. Ital. 1, 231, 1863.............................. (1)
sheath or appendages. The only known
C. mangrovei K.D. Hyde, Trans. Mycol. Soc.
Eutypella species on intertidal marine
Jpn. 34: 311, 1993.
mangrove wood often growing on decorticated

118
 Fungal Diversity

Avicennia species, with ascomata developing F. tunitricuspis K.D. Hyde, J. Linn. Soc. Bot.
under blackened stroma, with necks 110: 102, 1992 (Type species).
collectively erumpent through pustulate discs Ascomata immersed under a small
(Hyde, 1995a). clypeus beneath the host epidermis, lenticular,
ostiolar canal periphysate, dark brown to black,
Pedumispora K.D. Hyde & E.B.G. Jones, paraphyses simple, septate, tapering towards
Mycol. Res. 96: 78, 1992 ............................. (1) their tips, asci cylindrical, short pedunculate,
◙ P. rhizophorae K.D. Hyde & E.B.G. Jones, unitunicate, thin-walled, with an apical
Mycol. Res. 96: 78, 1992 (Type species). apparatus, J-, ascospores ellipsoidal, 1-septate,
Pseudostromatic tissue, comprising host not constricted at the septum, with a
cells and fungal hyphae, through which occur pronounced 3-4 lobed mucilaginous sheath. A
numerous darkened raised pustules with 1-4 monotypic genus occurring on the palm Nypa
immersed ascomata and 1-4 short necks. fruticans, and infrequently collected. Referred
Ascomata large, subglobose, coriaceous, by Hyde (1992a) to the Clypeosphaeriaceae it
brown-black, ostiolate, papillate, necks is a genus that needs evaluation at the
collectively erumpent, periphysate, paraphyses molecular level.
filiform, simple, asci irregularly fusiform,
pedunculate, unitunicate, thin-walled, apically Phragmitensis M.K.M. Wong, Poon & K.D.
truncate without an apical apparatus, Hyde, Bot. Mar. 41: 379, 1998 .................... (1)
ascospores filiform, tapering towards both ends, P. marina M.K.M. Wong, Poon & K.D. Hyde,
curved characteristically, hook-shaped at one Bot. Mar. 41: 379, 1998 (Type species).
end, 13-17-septate, not constricted at the septa, Pseudostroma long (0.7-1.9 mm long)
7-9 longitudinal striations running the length of immersed formed from host and fungal tissue,
the spore, tips lacking cytoplasm, yellow to black, with 3-6 ascomata, which are globose to
brown (Fig. 58). A characteristic feature of this subglobose, necks periphysate, paraphyses
species is the filliform, 13-17- septate, hyaline hyphal-like, straight, flexuous, septate,
to yellow to yellowish-brown, ascospores constricted at the septa, tapering, asci clavate,
tapering at both ends, curved and hook-shaped unitunicate, thin-walled, short pedunculate, J-,
at one end. Asci are unitunicate with no apical lacking an apical apparatus, ascospores
apparatus and rupturing to release the unicellular, cymbiform, basal end more tapered,
ascospores or deliquescing at maturity (Hyde hyaline, smooth-walled, with a mucilaginous
and Jones, 1992a). Assigned tentatively to the sheath. An infrequent, saprobic, intertidal
Diaporthales (Hyde and Jones, 1992a), but species on decaying culms of the marsh grass
LSU rDNA sequences positions it in the Phragmites australis (Wong et al., 1998).
Xylariales with the genera Libertella and Although assigned here to the Xylariales, the
Cryptosphaeria as sister groups (Sakayaroj, asci lack an apical apparatus.
2005). We consider it well placed in the
Diatrypaceae as it groups with the genera 5. Xylariaceae
Cryptosphaeria, Diatrype, Eutypa and
Eutypella. Anthostomella Sacc., Atti Soc. Veneto-Trent.
Sci. Nat., Padova 4: 84, 1875....................... (6)
4. Hyponectriaceae A. nypae K.D. Hyde, B.S. Lu & Alias, Mycol.
Res. 103: 1409, 1999.
A family sometimes referred to the A. nypensis K.D. Hyde, Alias & B.S. Lu,
Phyllachorales, we follow Barr (1994) and Mycol. Res. 103: 1410, 1999.
Cannon and Kirk (2007) in placing it in the A. nypicola K.D. Hyde, Alias & B.S. Lu,
Xylariales. Mycol. Res. 103: 1411, 1999.
Frondicola K.D. Hyde, J. Linn. Soc. Bot. 110: A. poecila Kohlm., Volkm.-Kohlm. & O.E.
100, 1992...................................................... (1) Erikss., Bot. Mar. 38: 175, 1995.

119
A. spissitecta Kohlm. & Volkm.-Kohlm., 4. Ascospores shorter than 15 µm, and narrower than
Mycol. Res. 106: 369, 2002. 8.5 µm...................................................................... 5
◙ A. torosa Kohlm. & Volkm.-Kohlm., Mycol.
5. Ascospores 10-15 × 4-7 µm, on Juncus roemerianus
Res. 106: 365, 2002. ...................................................................A. poecila
Ascomata globose to subglobose, 5. Ascospores 13-15 × 5.5-7 µm, with a well developed
immersed in the host, pale brown to dark gelatinous appendage on J. roemerianus .... A. torosa
brown to black, ostiolate, epapillate, clypeate,
coriaceous, periphysate, solitary or gregarious, Astrocystis Berk. & Broome, J. Linn. Soc. Bot.
paraphyses numerous, simple, septate, asci 14: 123, 1873 ............................................... (2)
cylindrical, short pediculate, unitunicate, thick- A. nypae G.J.D. Sm. & K.D. Hyde, Fungal
walled with a J+ apical apparatus, ascospores Diver. 7: 93, 2001.
ellipsoidal, unicellular, with germ-slits A. selangorensis G.J.D. Sm. & K.D. Hyde,
extending over the whole length of the spore, Fungal Diver. 7: 104, 2001.
hyaline with gelatinous caps or sheaths (Fig. Ascomata erumpent, mammiform,
59). Most of these species have been collected globose to subglobose, ostiolate, minutely
on Juncus roemerianus, Nypa fruticans, papillate, black, carbonaceous, solitary or
Spartina spp. and Pandanus sp., but many also gregarious, paraphyses hypha-like, septate, in a
occur on mangrove wood (Jones and Abdel- gelatinous matrix, asci cylindrical, short
Wahab, unpublished data from the Bahamas), pedunculate, apical apparatus J+, ascospores
and need a more detailed examination (Lu and ellipsoidal, unicellular, golden brown to dark
Hyde, 2000; Kohlmeyer and Volkmann- brown, germ slits run more or less the length of
Kohlmeyer, 2002). Anthostomella semitecta, the spore, no sheaths or appendages to the
has also been reported as a halotolerant species marine species (Smith and Hyde, 2001) (Fig.
on J. roemerianus with ascospores measuring 60). Astrocystis is similar to Rosellinia, but
15.9-18.8 × 6.6-8.1 µm, 1-septate, a large differs in ascus and ascospore morphology,
brown and a small hyaline basal cell, and a asci in Astrocystis have short stipes and ascus
prominent gelatinous sheath (Kohlmeyer et al., apparatus is small, while in Rosellinia the ascus
1995a). Lu and Hyde (2000) rejected the stipe is long with a massive barrel shaped ascus
species A. poecila and A. semitecta, regarding apparatus. A number of Astrocystis species
them as synonyms of A. tenacis and A. have been described from terrestrial palms
sepelibilis, respectively. Molecular data place (Smith and Hyde, 2001).
A. torosa, the only marine species sequenced, The two species listed here have been
in the Xylariaceae. described from the brackish water palm, Nypa
fruticans in Kuala Selangor, Malaysia.
1. On Nypa fruticans .................................................... 2
1. On Juncus roemerianus and other plants ................. 4
Although there is no molecular data for the
marine Astrocystis species, Astrocystis
2. Ascospores lacking a sheath, with a hyaline basal eleiodoxae (a peat swamp species) is well
dwarf cell, ascospores 12.5-16.5 × 5-7.5 µm ............. placed in the Xylariaceae (Pinnoi et al.,
................................................................ A. nypicola unpublished data).
2. Ascospores with a sheath, without a basal dwarf cell.
................................................................................. 3
1. Stromata splitting stellately, ascospores 8-10 × 3-5
µm, golden brown, germ slit 4/5 of total length.........
3. Ascospores 11.5-17 × 4.5-5.5 µm, inequilaterally .....................................................................A. nypae
ellipsoidal, thin sheath, full length germ slit. ............. 1. Stromata not splitting stellately, ascospores 15-19 ×
.....................................................................A. nypae 7.5-9 µm, light brown to reddish brown, germ slit
3. Ascospores 12.18 × 5.5-7.5 µm, inequilaterally whole length of the spore................ A. selangorensis
broadly ellipsoidal, thin sheath, full length germ slit
................................................................ A. nypensis
Fasciatispora K.D. Hyde, Trans. Mycol. Soc.
4. Ascospores longer than 15µm, inequilaterally Jpn. 32: 265, 1991........................................ (2)
broadly ellipsoidal, thin sheath, full length germ slit, F. lignicola Alias, E.B.G. Jones & Kuthub.,
16.5-23 × 13.5-15.5 µm, on Spartina alterniflora .....
..............................................................A. spissitecta
Mycotaxon 52: 78, 1994.

120
 Fungal Diversity

F. nypae K.D. Hyde, Trans. Mycol. Soc. Jpn. good support and a sister clade which includes
32: 267, 1991 (Type species). X. hypoxylon (Pinnoi et al., unpublished data)
Stroma dark coloured beneath the host Halorosellinia oceanica produces a wide range
epidermis. Ascomata immersed beneath the of bioactive compounds, including cytocha-
stroma, globose, ostiole periphysate, para- lasins, which is consistent with a taxonomic
physes persistent, septate, rarely branched, asci affinity to Rosellinia and Xylaria (Whalley and
cylindrical, unitunicate, thin-walled, with a J+ Edwards, 1987; Schlinghgam et al., 1998, Li et
apical apparatus, ascospores unicellular, al., 2001). It is world wide in its distribution in
ellipsoidal, yellow brown to brown, with a tropical mangrove habitats. A Nemania species
central pallid band and polar gelatinous occurs frequently in mangrove habitats and this
appendages. Fasciatispora was assigned to the can be confused with H. oceanica, but differs
Amphisphaeriaceae (Hyde, 1991c, 1995b), but in the smaller size of the ascospores and a
molecular data place F. petraki in the pronounced decolourization of the wood
Xylariaceae. around the ascomata.

1. Ascospores with polar gelatinous appendages, 11.6- Nemania Gray, Nat. Arr. Brit. Pl. (London) 1:
17 × 5.6-7.1 ..................................................F. nypae 516, 1821, emend. Pouzar, Ceská Mykologie
1. Ascospores lacking polar appendages, 8.4-14.9 × 3.9-
7.5 μm ..................................................... F. lignicola 39: 16, 1985 ................................................. (1)
◙ N. maritima Y.M. Ju & J.D. Rogers, Nova
Halorosellinia Whalley, E.B.G. Jones, K.D. Hedw. 74: 102, 2002.
Hyde & Laessøe, Mycol. Res. 104: 368, 2000. Stromata scattered, subglobose or
...................................................................... (1) globose enclosing 1-many ascomata that are
◙ H. oceanica (S. Schatz) Whalley, E.B.G. carbonaceous, ostiolate, dark brown, asci
Jones, K.D. Hyde & Lassøe, Mycol. Res. 104: cylindrical, short pedunculate, with a J+ apical
370, 2000 (Type species). ring, ascospores ellipsoidal, with broadly
Hypoxylon oceanicum Schatz, Mycotaxon 33: rounded end, unicellular, light brown to brown,
413, 1988. with a straight germ slit the length of the spore
Pseudostroma on decorticated wood, (Fig. 61). A species from Kandelia candel
single or/in clusters of up to 30 uniperitheciate mangrove wood collected in Taiwan (Ju and
pseudostroma, ascomata immersed in Rogers, 2002) and widely collected in
pseudostroma, subglobose to hemispherical, subtropical mangroves (Jones and Abdel-
soft to leathery, ostiolate, papillate, paraphyses Wahab, 2005). This species can be confused
numerous, persistent, asci cylindrical with Halorosellinia oceanica as both grow in
unitunicate, J+ ascus tip, ascospores ellipsoidal, the same habitat, but the ascospores of N.
dark grey-olive to brown, wall smooth, germ maritima are much smaller 9-12 × 5-6 (-6.5)
slit straight, half to three quarters total length µm with rounded ends and a germ slit shorter
of spore, without a sheath or appendages. than the spore length. Unlike H. oceanica this
Halorosellinia oceanica was separated species is reported to produce the teleomorph
from Hypoxylon as it did not meet the criteria in culture but no anamorph has been
for that genus: 1. Nodulisporium anamorph; 2. encountered. Ju and Rogers (2002) believe this
Stromata unipartite, 3. Stromatal tissue below species could be segregated to a new genus, but
the perithecial layer solid and homogenous and did not elaborate on the grounds for this. Tang
4. Stromata not upright (Whalley et al., 2000). et al. (2007b) in ITS-RPB2 sequences show
Molecular data indicate a relationships with that N. maritima is placed in the Xylarioideae
Xylaria hypoxylon (18S, 28S sequences and clade and Nemania subclade, being basal to
combined 18S and 28S data set) rather than the other species. Xylaria hypoxylon is nested in
genus Hypoxylon (Smith et al., 2003). the other subclade with X. grammica and a
However, ITS-5.8S-ITS2 sequences support a Xylaria sp. Pinnoi, et al. (unpublished data) in
relationship with Astrocystis eleiodoxae with an ITS dataset also shows that N. maritima is

121
 Fig. 58. Morphological features of Pedumispora rhizophorae. a. Exposed ascomata on wood surface. b, d. Filiform
ascospores tapering towards both ends. c. Longitudinal striations on the ascospore cell wall. Bars a = 1 mm; b-d = 20
µm.

a b

Fig. 59. Ascospores: a. Anthostomella torosa. b. A. spissitecta. Bars = 5 µm.

122
 Fungal Diversity

well placed in the Nemania clade with high Anthostomella in lacking a J+, subapical ring
bootstrap support. However, it groups with N. and in its black ascospores (Hyde and Taylor,
confluens with high support, in a sister clade to 1996).
other Nemania species.
This species is also known from Hong 1. Ascospores 17-20 × 4-6 µm, appendages are part of
an elaborate sheath drawn out at each end, clypeus
Kong and Malaysia and is possibly widespread poorly developed, ascus ring J-........... N. carbospora
in its distribution. The fungus was collected at 1. Ascospores 16-18 × 5-6 µm, germ slit along the
four sites in Hong Kong: Three Fathom Cove, entire length of the spore, elaborate mucilaginous
Ting Kok, Ho Chung and Mai Po. It was sheath, clypeus well-developed, ascal ring J+ ...........
recorded as the most common species at these ........................................................ N. selangorensis
four sites colonizing 14% of the total number
XYLARIALES incertae sedis
of samples (Abdel-Wahab and Jones,
unpublished data). The Hong Kong material Adomia S. Schatz, Trans. Br. Mycol. Soc. 84:
had asci that measure 82-108 × 7.5-10 µm ( x = 555, 1985...................................................... (1)
93.5 × 7.7 µm, n = 20), with a subapical ring A. avicenniae S. Schatz, Trans. Br. Mycol. Soc.
and inverted cap in shape, ascospores 9-10 × 5 84: 555, 1985 (Type species).
µm ( x = 9.6 × 5 µm, n = 40). Figure 61, Ascomata ampulliform to subglobose,
illustrates material collected in Hong Kong. immersed, brown, ostiolate, clypeate,
periphysate, paraphyses broad, asci cylindrical,
Nipicola K.D. Hyde, Cryptog. Bot. 2: 330, unitunicate, thin-walled, deliquescent with a
1992.............................................................. (2) persistent apical cap, ascospore ellipsoidal,
N. carbospora K.D. Hyde, Cryptog. Bot. 2: brown, unicellular, with an appendage at one
330, 1992 (Type species). end of the spore. This poorly known species
N. selangorensis K.D. Hyde, Sydowia 46: 262, was described from material collected in
1994. Australia and the Red Sea, Egypt, on Avicennia
marina, and referred to the Phyllachorales
a b (Schatz, 1985). It has recently been collected in
New Zealand and Thailand (Lintott and Lintott,
2002; Jones et al., 2005). Collections in
Thailand were on Avicennia marina
pneumatophores.

Lanceispora Nakagiri, Okane, Tad. Ito &

Katum., Mycoscience 38: 208, 1997 ........... (1)
L. amphibia Nakagiri, Okane, Tad. Ito &
Fig. 60. Ascospores of a. Astrocystis nypae. b. A. Katum., Mycoscience 38: 208, 1997 (Type
selangorensis. Bars = 5 μm. species).
Ascomata globose to subglobose,
Ascomata formed under the host surface immersed, olivaceous to olivaceous black,
under slightly raised areas, or under a reduced ostiolate, papillate, necks lacking periphyses,
blackened clypei, globose to subglobose, paraphyses filiform, branched, septate, hyaline,
ostiolate, paraphyses amphisphaeriaceous, asci asci cylindrical, unitunicate, long pedunculate,
broadly cylindrical, short pedunculate, apical ring K+, ascospores oblanceolate,
subapical ring J-, ascospores lunate, unicellular, hyaline, rounded apical part, tapering towards
dark brown to black, with or without a germ the lower end, 1-septate, above the spore
slit, with well developed gelatinous sheaths. middle, lacking a sheath or appendages.
The two marine species occur on the intertidal A genus referred by Nakagiri et al. (1997)
palm Nypa fruticans, with black lunate to the Amphisphaeriaceae while Kirk et al.
ascospores, with a layered mucilaginous sheath (2001) are of the opinion it is close to
and appendages at the ends. It differs from Ceriospora-Urosporellopsis. Only one species

123
has been reported from marine habitats while L. structure that encloses microfibrils.
phyllophila has been described from the petiole Appendages are hair-like, wavy, and separate
of an unidentified dicotyledonous species in fibrils are surrounded by a thin membrane,
leaf litter (Sarma and Hyde, 2001). Assignment which is also present in the ascospores of L.
to family is unresolved: originally referred to nypae (Poonyth et al., 2000a). It was referred
the Amphisphaeriaceae because of its affinities to the Hyponectriaceae by Hawksworth et al.
with Leiosphaerella (Nakagiri et al., 1997), (1995), although Wang and Hyde (1999)
while Barr (1994) thought the latter genus excluded it from this family.
should be assigned to the Hyponectriaceae.
Wang and Hyde (1999) rejected this while 1. Ascospores needle-shaped, lacking appendages, 57.5-
Kang et al. (1999b) suggested the 87.5 × 3.5-6 µm ................................. L. angustatum
1. Ascospores filamentous with appendages................ 2
Clypeosphaeriaceae for Leiosphaerella and
Oxydothis. Since Lanceispora has much in 2. Ascospores with appendages at both ends, 90-139 ×
common with Leiosphaerella then this family 2-3 µm...................................................... L. bipolare
should be considered, but further molecular 2. Ascospores with a single appendage........................ 3
studies of this complex is required before final
assessment can be made. 3. Ascospores C-shaped or sigmoid, one end rounded,
one end tapering with mucilage, 124-140 × 2.5-3 µm
Linocarpon Syd. & P. Syd., Ann. Mycol. 15: .......................................................... L. longisporum
3. Ascospores not markedly curved ............................. 4
210, 1917...................................................... (6)
L. angustatum K.D. Hyde & Alias, 4. Ascospores 75-120 × 2-3.5 µm, with thistle-shaped
Mycoscience 40: 145, 1999. appendage, on mangrove wood..................................
◙ L. appendiculatum K.D. Hyde, Trans. Mycol. .................................................... L. appendiculatum
Soc. Jpn. 29: 339, 1989. 4. Ascospores 91-124 × 3.5-4 µm, with a mucilaginous
L. bipolare K. D. Hyde, Sydowia 44: 38, 1992. pad at one end, on mangrove wood.............. L. nypae
L. longisporum K. D. Hyde, Sydowia 44: 44,
1992. Neolinocarpon K.D. Hyde, Bot. J. Linn. Soc.
L. nipae (Henn.) K.D. Hyde, Trans. Mycol. 110: 104, 1992 ............................................. (2)
Soc. Jpn. 29: 346, 1989. ◙ N. globosicarpum K.D. Hyde, J. Linn. Soc.,
Ophiobolus nipae Henn. Hedwigia 49: 297,
Bot. 110: 104, 1992.
1908. N. nypicola K.D. Hyde & Alias, Mycoscience
Pseudostroma black, ascomata circular, 40: 148, 1999.
lenticular, dome-shaped, beneath the Ascomata immersed in the host, oval to
pseudostroma, ostiolate, short papilla may be subglobose, ostiolate, coriaceous, black, neck
present, paraphyses filamentous, smooth, short with periphyses, paraphyses tapering,
septate, hyaline, in a gelatinous matrix, asci septate, simple, longer than the asci, asci long
cylindrical, pedunculate, truncate at the apex, cylindrical, pedunculate, unitunicate, thin-
with J- ring, ascospores filiform, hyaline, walled, ring-like refractive apical apparatus,
slightly curved or straight, apically gelatinous ascospores filiform, straight or curved, hyaline,
caps or pad-like appendages at both ends unicellular, with apical cap-like mucilaginous
(Hyde, 1988; Hyde and Alias, 1999). This appendages. This genus marginally differs
genus is common on Nypa fruticans and from Linocarpon, and molecular data are
mangrove wood in tropical locations (Hyde and required to resolve its taxonomic affinities. A
Alias, 1999; Poonyth et al., 2000a). Ascospores second Neolinocarpon species has been
are filiform to needle-shaped, hyaline, and described from the dead aerial rachids of Nypa
unicellular, often with polar pad-like fruticans (Hyde and Alias, 1999). They differ
appendages or a mucilaginous sheath. as follows: Neolinocarpon has deeply
Ultrastructure studies of L. appendiculatum immersed ascomata forming below a slightly
show that the asci are thin-walled (ca. 70 nm raised or flattened clypeus and with a refractive
thick) with a C-shaped apical ring with globose body below the apical ascus ring,
subapical projections into the epiplasm, while Linocarpon has ascomata that are
ascospores at their tips have a collar-like superficial or slightly immersed and lack a

124
 Fungal Diversity

a b

c d e f

Fig. 61. Nemania maritima. a. Section through ascoma on superficial wood. b. Apical region of the ascoma with asci
and paraphyses. c. Asci and paraphyses. d-f. Asci with eight ascospores and prominent ring. Bars a = 100 µm, b = 10
µm, c = 10 µm, d-f = 4 µm. (Photos by M.A. Abdel-Wahab).

refractive globose body (Yanna et al., 2003). Ascomata beneath the host epidermis,
Ultrastructure studies of selected Linocarpon globose to subglobose, ostiolate, periphysate,
and Neolinocarpon species failed to yield brown to black gregarious, paraphyses
characters for the delineation of the genera numerous, filamentous, hyaline, asci
(Yanna et al., 2003). Currently the genus can cylindrical, pedunculate, unitunicate, thin-
not be assigned to any family within the walled, with an apical ring J+, ascospores
Xylariales. filiform, straight or curved, hyaline, 0-1-septate,
not constricted at the septum, with mucilage at
1. Ascomata formed under a clypeus, solitary, the tip of the spore (Hyde and Nakagiri, 1989).
ascospores 70-119 × 2-3 µm .........N. globosicarpum
1. Ascomata deep within a stroma, ascospores 92-117 × Both species occur on ageing or cut fronds of
2-4 µm..................................................... N. nypicola the palm Nypa fruticans that have fallen into
the water and are well placed in the genus.
Oxydothis Penz. & Sacc., Malpighia 11: 505, Oxydothis nypae appears to be the most
1898.............................................................. (2) common species, recorded from a variety of
O. nypae K.D. Hyde & Nakagiri, Trans. Mycol. collections (Hyde and Alias, 2000;
Soc. Jpn. 30: 70, 1989. Pilantanapak et al., 2005). Kang et al. (1999a)
O. nypicola K.D. Hyde, Sydowia 46: 298, referred the genus to the Clypeosphaeriaceae.
1994.
125
1. Ascospores 1-septate, diameter more than 6 µm, 74- 5. Ascospore appendages indistinct, ascospores large,
92 µm long, fusiform, with apiculate ends................. 11.5-16 × 5.5-8 μm, on mangrove wood....................
................................................................ O. nypicola .............................................................Ph. kandeliae
1. Ascospores 0-septate, diameter less than 6 µm, 82-
113 µm long, ends rounded with mucilage ................ SORDARIOMYCETES incertae sedis
.................................................................... O. nypae
1. Ascomata perithecial, non-stromatic, ostiolate,
Phomatospora Sacc., Grevillea 4: 22, 1875.(6) papillate with long necks, coriaceous to carbonaceous,
P. acrostichi K.D. Hyde, Trans. Br. Mycol. paraphyses, asci variable, thin-walled, deliquescing
Soc. 90: 135, 1988. early, ascospores variable, fusiform, ellipsoidal,
filiform, generally with end-chambers containing
◙ P. bellaminuta Kohlm., Volkm.-Kohlm. & mucilage, anamorphs generally dermatiaceous
O.E. Erikss., Bot. Mar. 38: 181, 1995. hyphomycetes .....................................Lulworthiales
P. kandeliae K.D. Hyde, Trans. Mycol. Soc. 1. Ascomata perithecial, immersed in host tissue,
Jpn. 33: 315, 1992. subiculate or without subiculum, periphysate,
P. nypae K.D. Hyde, Sydowia 45: 200, 1993. paraphysate, asci clavate to fusiform, unitunicate,
deliquescing, ascospores septate, hyaline, thick-
P. nypicola K.D. Hyde & Alias, Mycol. Res. walled, no apical chambers or sheaths
103: 1417, 1999. ......................................................Koralionastetales
P. phragmiticola Poon & K.D. Hyde, Bot. Mar. 1. Ascomata perithecial, immersed in host tissue, often
41: 148, 1998. clypeate, ostioles periphysate, paraphyses wide thin-
Ascomata globose to subglobose, walled, asci cylindrical, persistent with
inconspicuous I-apical ring, ascospores aseptate,
immersed, pale brown to black, membranous, anamorph coelomycetes.................... Phyllachorales
ostiolate, short papillate, solitary, necks 1. Ascomata perithecial, ostiolate, immersed or
periphysate, paraphyses hypha-like, straight, superficial, often staining the substratum,
septate, tapering distally, numerous, asci paraphysate, paraphyses wide at the base tapering
cylindrical, unitunicate, thin-walled, apically, asci cylindrical, unitunicate, ascospores
septate, often filiform, hyaline, yellow to brown
pedunculate, with a J+ refractive ring, ....................................................... Magnaporthales
ascospores unicellular, ellipsoidal, hyaline,
with longitudinal striae and with or without
polar appendages. A genus frequently 1. LULWORTHIALES
encountered on mangrove wood, Nypa
fruticans, Phragmites australis and the fern Key to the genera
Acrostichum speciosum, while other species
1. Ascospores with apical chambers or appendages .... 2
await description (Jones, unpublished data).
1. Ascospores lacking appendages......................Lindra
Cai et al. (2006) refer the genus to
Soradriomycetes incertae sedis, but suggest 2. Ascospores 0-septate................................................ 3
that 18S rDNA sequences indicate a close 2. Ascospores septate ................................................... 7
relationship to the Magnaporthaceae. This
3. Ascospores filiform.................................................. 6
hypothesis requires further study to determine
3. Ascospores fusiform, ellipsoidal.............................. 4
assignment to a family and an order.
4. Ascospore appendages tubular longer than 35µm .....
1. Ascospores lacking an appendage......... Ph. nypicola .......................................................... Kohlmeyeriella
1. Ascospores appendaged ........................................... 2 4. Ascospore appendages apiculate or conical, less than
35 µm long............................................................... 5
2. Appendages at both ends.......................................... 3
2. Appendages at one pole ........................................... 5 5. Ascomata superficial, with sterile hairs, parasitic on
red algae............................................. Spathulospora
3. Ascospore appendages pad-like .......Ph. bellaminuta 5. Ascomata immersed in a gall, lacking sterile hairs, on
3. Ascospore appendages bifurcate .............................. 4 brown algae......................................Haloguignardia

4. Ascospores 6.5-9.5 μm, on Nypa ...............Ph. nypae 6. Ascomata with bell-like structure within the centrum
4. Ascospores 7.5-1.5 μm, on Phragmites ..................... ............................................................... Rostrupiella
..................................................... Ph. phragmitensis 6. Ascomata lacking the bell-like structure..Lulworthia

5. Ascospore appendages prominent, ascospores 6-7 × 7. Ascospores 1-septate..................................Lulwoana

2-3 μm, on a fern................................. Ph. acrostichi 7. Ascospores many times septate.................Lulwoidea

126
 Fungal Diversity

Lulworthiaceae Anamorph: Anguillospora marina

Kohlmeyeriella E.B.G. Jones, R.G. Johnson & Nakagiri & Tubaki
S.T. Moss, Bot. J. Linn. Soc. 87: 210, 1983...(2) ◙ L. thalassiae Orpurt, Meyers, Boral &
◙ K. crassa (Nakagiri) Kohlm., Volkm,.- Simms, Bull. Mar. Sci. Gulf Caribb. 14: 406,
Kohlm., J. Campb., Spatafora & Gräfenhan., 1964.
Mycol. Res. 109: 564, 2005. L. marinera Meyers, Mycologia 61: 488, 1969.
Lulworthia crassa Nakagiri, Trans. Mycol. Soc. Ascomata semiglobose or ellipsoidal,
Jpn. 25: 378, 1984. immersed, carbonaceous to subcarbonaceous,
◙ K. tubulata (Kohlm.) E.B.G. Jones, R.G. black, ostiolate, papillate or epapillate, solitary
Johnson & S.T. Moss, J. Linn. Soc. Bot. 87: or gregarious, asci cylindrical to clavate,
208, 1983 (Type species). unitunicate, thin-walled, deliquescing early,
Corollospora tubulata Kohlm., Ber. Dtsch. Bot. lacking an apical apparatus, ascospores filiform,
Ges. 81: 53, 1968.
many times septate (up to 50), not constricted
Ascomata subglobse, superficial, subicu-
at the septa, with a globose to semiglobose
late, ostiolate, papillate, subcarbonaceous,
gelatinous appendage on wood. Species on
black, solitary often associated with sand, asci
seagrasses with coriaceous ascomata immersed
fusiform to clavate, unitunicate, thin-walled,
in the host, and lacking an apical swelling or
deliquescing early, ascospores unicellular,
appendage. Molecular data indicate that this
fusiform, curved, hyaline, thick-walled, with
genus is distantly related to the Halosphaeriales
long polar appendages, with an apical pore
and Kohlmeyer et al. (2000) have assigned it to
from which a drop of mucilage is released.
the Lulworthiales.
This genus, like Lulworthia, has ascospores
The genus appears to be polyphyletic
with end chambers that are long and filled with
with Lindra obtusa forming a basal group to
mucilage, which is released at maturity as a
the order, while L. crassa and L. marinera are
drop, which aids in spore attachment (Jones,
distantly placed within the Lulworthiales
1994). Initially described as a Corollospora
(Campbell et al., 2005) (Fig. 62). Lindra
species, it was subsequently transferred to the
inflata, the type species, needs to be sequenced
new genus Kohlmeyeriella based on TEM
before further taxonomic changes are
observations (Jones et al., 1983a). Molecular
undertaken. Morphologically L. inflata differs
data shows it has no affinities with the
from L. marinera and L. thalassiae in the
Halosphaeriales and has been transferred to the
deeply embedded carbonaceous ascomata in
Lulworthiales (Campbell et al., 2005).
wood, thick-walled peridium, ascospores 30-
Sequence data shows that Lulworthia crassa
50-septate with a globose swelling gelatinous
groups with K. tubulata and has thus been
apical appendage (Wilson, 1956).
transferred to this genus (Campbell et al., 2005)
(Fig. 62). 1. Ascospores markedly swollen at their tips, 30-51-
septate, 210-415 × 4-6 µm, on wood ..........L. inflata
1. Ascospores narrow, 140-205 × 5-8 µm....... K. crassa 1. Ascsopores not markedly swollen at their tips......... 2
1. Ascospores wider, 137-152 × 17.5-18.5 µm ..............
.................................................................K. tubulata 2. Ascospores more than 8 µm wide, 320-520 µm, 15-
23-septate, on marine angiosperms and algae............
Lindra I.M. Wilson, Trans. Br. Mycol. Soc. 39: .................................................................... L. crassa
411, 1956...................................................... (5) 2. Ascospores less than 8 µm wide, septa fewer than 30
◙ L. crassa (Kohlm.) Kohlm. & Volkm.- ................................................................................. 3
Kohlm., Bot. Mar. 34: 23, 1991. 3. Ascospores longer than 230 µm, 230-390 × 3-6 µm,
L. hawaiiensis Kohlm. & Volkm.-Kohlm., Can. on sea grasses........................................ L. thalassiae
J. Bot. 65: 574, 1987. 3. Ascospores shorter than 230 µm, on other hosts...... 4
L. inflata I.M. Wilson, Trans. Br. Mycol. Soc.
39: 411, 1956 (Type species). 4. Ascospores narrower than 3.5 µm, 182-313 × 2.5-3.5
µm, forming subicula on a hard surface (sand)..........
◙ L. obtusa Nakagiri & Tubaki, Mycologia 75: ....................................................................L. obtusa
488, 1983.

127
4. Ascospores 101-188 × 3.5-5.5 µm, 8-18-septate, in L. calcicola Kohlm. & Volkm.-Kohlm.,
wood................................................... L. hawaiiensis Mycologia 81: 289, 1989.
L. curalii (Kohlm.) Kohlm. & Volkm.-Kohlm.,
Lulwoana Kohlm., Volkm.-Kohlm, J. Campb., Bot. Mar. 34: 24, 1991.
Spatafora & Gräf., Mycol. Res. 109: 62, 2005. Lulworthia kniepii var. curalii Kohlm., Mar. Ecol
...................................................................... (1) (P.S.Z.N.I.) 5: 361, 1984.
◙ L. uniseptata (Nakagiri) Kohlm., Volkm.- ◙ L. fucicola G.K. Sutherl., Trans. Br. Mycol.
Kohlm, J. Campb., Spatafora & Gräfenhan., Soc. 5: 259, 1916 (Type species).
Mycol. Res. 109: 562, 2005 (Type species). Lulworthia medusa var. fucicola (G.K. Sutherl.)
Lulworthia uniseptata Nakagiri. Trans. Mycol. Booth, Can. J. Bot. 61: 500, 1983.
Soc. Jpn. 25: 382, 1984 L. halima (Diehl & Mounce) Cribb & J.W.
Anamorph: Zalerion maritima (Linder) Cribb, Univ. Queensl., Pap. Dept. Bot. 3: 80,
Anastasiou 1955.
Ascomata superficial or immersed, Ophiobolus halimus Diehl & Mounce, Can. J. Res.
globose to subglobose, ostiolate, coriaceous, 11: 242, 1934.
Halophiobolus halimus (Diehl & Mounce) Linder,
dark brown or black, neck short, asci fusiform, Farlowia 1: 419, 1944.
curved in the ascoma, unitunicate, thin-walled, Linocarpon halimum (Diehl & Mounce) Petr., 6:
deliquescing early, ascospores filiform, hyaline, 388, 1952.
1-septate, with conical end chambers L. floridana Meyers, Mycologia 49: 515, 1957.
containing mucilage which is released as a ◙ L. grandispora Meyers, Mycologia 49: 513,
drop when mounted in water and forms an 1957.
adhesive pad for attachment of the ascospores. L. kniepii Kohlm., Nova Hedw. 6: 140, 1963.
A genus erected based on a single Ophiobolus kniepii Ade & Bauch, Publ. Statzione
morphological character of a 1-septate Napoli 15: 389, 1936.
ascospore, a Zalerion maritima anamorph and L. lindroidea Kohlm., Bot. Mar. 23: 537, 1980.
sequence data (Campbell et al., 2005) (Fig. 62). L. longirostris (Linder) Cribb & J.W. Cribb,
Univ. Queensl., Pap. Dept. Bot. 3: 80, 1955.
Halophiobolus longirostris Linder, Farlowia 1:
Lulwoidea Kohlm., Volkm.-Kohlm, J. Campb., 418, 1944.
Spatafora & Gräfenhan., Mycol. Res. 109: 564, L. medusa (Ellis & Everh.) Cribb & J.W. Cribb,
2005.............................................................. (1) Pap. Dept. Bot. Univ. Qd. 3: 80, 1955.
◙ L. lignoarenaria (Jørg. Koch & E.B.G. Jones) Halophiobolus medusa (Ellis & Everh.) Linder,
Kohlm., Volkm.-Kohlm., J. Campb., Farlowia 1: 419, 1944.
Spatatfora & Gräfenhan., Mycol. Res. 109: 564, Linocarpon medusa (Ellis & Everh.) Petr.,
2005 (Type species). Sydowia 6: 388, 1952.
Lulworthia lignoarenaria Jørg. Koch & E.B.G. Ophiobolus medusae Ellis & Everh., Journal of
Jones, Mycotaxon 20: 389, 1984. Mycology 1: 150, 1885.
Ascomata superficial, generally attached ◙ L. purpurea (I.M. Wilson) T.W. Johnson,
to sand grains by subicula, subglobose to Mycologia 50: 154, 1958.
Halophiobolus purpurea I.M. Wilson, Trans Br.
ellipsoidal, carbonaceous, asci clavate, Mycol. Soc. 39: 403, 1956.
pedunculate, unitunicate, thin-walled, deli- Ascomata immersed or superficial,
quescing early, ascospores filiform, hyaline, globose or cylindrical depending on the host
multiseptate, with polar end chambers filled substratum, ostiolate, papillate, solitary or
with mucilage which is released through a pore gregarious, pale brown red to purple to dark
at maturity when in water. Another genus brown depending on species, paraphyses absent,
erected based on sequence data and a single asci cylindrical, clavate to fusiform, unitunicate,
morphological character of ascospores with thin-walled, deliquescing early, ascospores
numerous septa (31-38). A species generally filiform curved, hyaline, with conical end
found on sand grains associated with buried chambers formed from the outgrowth of the
wood (Koch and Jones, 1984). episporium and mesosporium, and filled with
mucilage which is released through a pore at
Lulworthia G.K. Sutherl., Trans. Br. Mycol. the tip of the appendage (Jones et al., 1983a).
Soc. 5: 259, 1916 ....................................... (11) Many Lulworthia species were originally

128
 Fungal Diversity

Fig. 62. One of the MPTs inferered from LSU rDNA sequences of all lulworthialean taxa, generated with maximum
parsimony anlaysis. Bootstrap values higher than 50% are given above branches.
described by Barghoorn and Linder (1944) as order to accommodate them.
Halophiobolus, but subsequently transferred to Since no type material of Lulworthia
the earlier taxon Lulworthia (Sutherland, 1916b) fucicola could be traced, Campbell (2005)
by Cribb and Cribb (1955). Because of the designated a neotype with a Lulworthia
deliquescing asci and ascospores with an end collection from Chile. However this is a
chamber containing mucilage it was assigned lignicolous strain while L. fucicola was
to the Halosphaeriales (Kohlmeyer, 1972a). described from a brown alga. We do not accept
Kohlmeyer et al. (2000) have shown that at the the argument that the significant morphological
molecular level these species have no affinities differences between the algicolous and
with the Halosphaeriales and erected a new lignicolous strains are substrate-induced

129
(Meyers, 1957). Furthermore we reject the Anamorphs of members of the
synonymy of Lulworthia cylindrica with L. Lulworthiales include: Anguillospora marina
fucicola (Cribb and Cribb, 1955). Greater (Lindra obtusa), Cirrenalia pygmea, Cirrenalia
efforts must be made to collect Lulworthia tropicalis, Cumulospora varia and Orbimyces
species from marine algae. spectabilis (all with different Lulworthia spp.)
Nakagiri (1984) reviewed the available and Zalerion maritima (Lulwoana uniseptata)
morphological characters for the separation of (Jones et al., 2008) (Figs. 62, 63).
the genera Lindra and Lulworthia: peridial wall As the result of sequence analysis of 18S
structure, ascospore septation, possession of and 28S a number of transfers have been
end chambers filled with mucilaginous material proposed: Lulworthia crassa to Kohlmeyeriella,
and their anamorph conidial development. For Lulworthia lignoarenaria to Lulwoidea and
their selected examples, only the latter two Lulworthia uniseptata to Lulwoana (Campbell
characters were considered significant at the et al., 2005), but a key to the new genera was
generic level. Peridial wall structure depends not provided. Although there is good molecular
on the habitat of the species: arenicolous support for the separation of Lulwoidea and
species (L. crassa, L. lignoarenaria) have a Lulwoana from Lulworthia, morphological
single outer-layer of thick-walled, large cells, differences are few.
but these were not observed in the lignicolous
species (L. uniseptata). Similar trabeculate 1. Ascospores 9-12-septate, 170-240 × 4 µm................
carbonaceous outer cells have been observed in .............................................................. L. lindroidea
1. Ascospores with no septa......................................... 2
Corollospora besarispora, another arenicolous
fungus (Sundari et al., 1996). This 2. Lignicolous species.................................................. 3
demonstrates that the environment can 2. On other substrata .................................................... 6
markedly influence the phenology of an
ascomycete, as has also been demonstrated for 3. Ascospores over 500 µm long (500-756 × 3-5 µm)
the variability in the length of the necks in tropical ...............................................L. grandispora
Lulworthia and Halosphaeria appendiculata 3. Ascospores shorter than 500 µm.............................. 4
(Jones, 1962).
4. Ascomata deeply embedded in the wood................. 5
The genus has been shown to be 4. Ascomata more superficial, dark brown, 230-266-320
polyphyletic based on 18S and 28S sequences × 3.6-4.2-5.7 µm .................................... L. floridana
analysis and two new genera erected to
accommodate species that do not group within 5. Ascospores 155-299 × 3.3-4 µm......... L. longirostris
the genus Lulworthia sensu stricto (Campbell 5. Ascomata purple in colour, elongate, ascospores 200-
et al., 2005). However, further species need to 350 µm elongate..................................... L. purpurea
be studied at the molecular level to fully
6. Ascomata on coral or coralline algae ....................... 7
resolve the taxonomic position of the various 6. Ascomata on marine grasses or on Fucus ................ 9
species. We include three species not
recognized by Kohlmeyer and Volkmann- 7. Ascospores over 200 µm (200-270 × 2.5-6 µm)........
Kohlmeyer (1991a) and Hyde et al. (2000): L. ................................................................... L. kniepii
floridana with ascospores measuring 230-266 7. Ascospores shorter than 200 µm.............................. 8
× 3.6-5.7 μm and with dark brown ascomata; L.
8. Ascospores 80-150 × 5.5-8.5 µm, on coral rock ........
purpurea with a thin ascomatal wall ................................................................ L. calcicola
comprising only two layers of cells, purplish 8. Ascospores 120-180 × 4-5 µm, coralline red algae,
brown ascomata turning green in lactophenol conch shells, coral reef................................ L. curalii
and median ascospores 185-265 × 3-4 μm and
Rostrupiella danica, a new species with 9. On the brown alga Fucus, ascospores 70-100 (-126)
distinctive inhibition hyphae in the wood and a × 4-6 µm .................................................. L. fucicola
9. On marine grasses .................................................. 10
pad of parenchymatous tissue separating the
centrum from the ostiole in the neck (Koch et 10. On Zostera rhizomes and leaves, ascospores 230-
al., 2007). Lulworthia purpurea differs from 308 × 2-4 µm........................................... L. halima
many Lulworthia species in the thin nature of 10. On culms of Spartina species, ascospores 350-526
the ascomata, which is only two-layered. µm .......................................................... L. medusa

130
 Fungal Diversity

Rostrupiella Jørg. Koch, K.L. Pang & E.B.G. immersed, ostiolate, short papillate, coriaceous,
Jones, Bot. Mar. 50: 294, 2007 .................... (1) hyaline, periphysate, paraphyses absent, asci
R. danica Jørg. Koch, K.L. Pang & E.B.G. ellipsoidal, clavate or cylindrical, thin-walled,
Jones, Bot. Mar. 50: 295, 2007. unitunicate, deliquescing early, no apical
Ascomata ellipsoidal-cylindrical, ostio- apparatus, ascospores ellipsoidal to fusiform,
late, gregarious, deeply embedded, subhyaline, unicellular, hyaline, with a conical acute
cylindrical neck brown, periphysate, lacking chamber at each pole. An unusual genus
paraphyses or catenophyses, asci cylindrical, forming galls on the larger brown algae
short pedunculate, unitunicate, thin-walled, no (Cystoseira, Halidrys, Sargassum) with
apical pore, deliquescing early, ascospores unicellular, hyaline ascospores with polar cap-
hyaline, filiform, tapering towards the poles, 0- like appendages that are divided into a series of
septate, curved or twisted, with an apical chambers containing mucous. Kohlmeyer and
conoid end-chamber from which mucilage is Volkmann-Kohlmeyer (2003b) illustrate H.
released in water. Morphological and irritans with mucous released from the
molecular evidence distinguishes Rostrupiella ascospore end chambers through an apical pore.
from Lulworthia fucicola, the type species of Although they classify the genus in the
the genus (Koch et al., 2007) (Figs. 63, 64). Phyllachoraceae, Phyllachorales, Inderbitzin et
Morphologically R. danica has a bell-like al. (2004) have shown that Haloguignardia
structure that extends from the neck into the irritans clusters within the Lulworthia complex,
centrum and large thin-walled bladder cells, as a separate branch from the L. uniseptata and
prominent pigmented cells around the neck and L. cfr purpurea clade (Fig. 62.) Clearly further
in the host wood tissue. A similar structure has investigations at the ultrastructural and
been noted in Haloguignardia irritans, another molecular level are required to finally resolve
member of the Lulworthiales (Kohlmeyer and the taxonomic position of the genus.
Kohlmeyer, 1979).
1. In the brown algae Cystoseira or Halidrys .............. 2
Haloguignardia Cribb & J.W. Cribb, Univ. 1. In Sargassum species ............................................... 3
Queensl., Pap. Dept. Bot. 3: 97, 1956.......... (5)
H. cystoseirae Kohlm. & Demoulin, Bot. Mar. 2. Ascospores longer than 60 μm............ H. cystoseirae
2. Ascospores shorter than 60 μm ................. H. irritans
24: 9, 1981.
H. decidua Cribb & J.W. Cribb, Univ. 3. Ascospores with persistent appendages .....................
Queensl., Pap. Dept. Bot. 3: 97, 1956. .......................................................... H. tumefaciens
◙ H. irritans (Setch. & Estee) Cribb & J.W. 3. Ascospore appendages deciduous............................ 4
Cribb, Univ. Queensl. Pap. Dept. Bot. 3: 98,
1956. 4. Ascospores shorter than 35 μm ................ H. decidua
Guignardia irritans Setch. & Estee, Univ. Calif., 4. Ascospores longer thsn 35 μm ............... H. oceanica
Berkeley, Publ. Bot. 4: 311, 1913.
H. oceanica (Ferd. & Winge) Kohlm., Mar. Spathulosporaceae
Biol. 8: 344, 1971
Phyllachorella oceanica Ferd. & Winge, Spathulospora A.R. Caval. & T.W. Johnson,
Mycologia 12: 102, 1920. Mycologia 57: 927, 1965 ............................. (5)
H. tumefaciens (Cribb & J.W. Herbert) Cribb ◙ S. adelpha Kohlm., Mycologia 65: 615, 1973.
& J.W. Cribb, Univ. Queensl., Pap. Dept. Bot. ◙ S. antarctica Kohlm., Mycologia 65: 619,
3: 98, 1956. 1973.
Guignardia tumefaciens Cribb & J.W. Herbert,
Univ. Queensl., Pap. Dept. Bot. 3(2): 9, 1954.
S. calva Kohlm., Mycologia 65: 622, 1973.
Haloguignardia longispora Cribb & J.W. Cribb, S. lanata Kohlm., Mycologia 65: 625, 1973.
Univ. Queensl., Pap. Dept. Bot. 3(12): 98, 1954. S. phycophila A.R. Caval. & T.W. Johnson,
Ascomata subglobose or ellipsoidal, Mycologia 57: 927, 1965 (Type species).

131
 Thallus crustose on algal host, though not Pontogeneia, previously not assigned with
always evident, ascomata subglobose, ovoid, confidence in any order (Campbell et al., 2008).
pyriform, subiculate, ostiolate, papillate or These two genera form a monophyletic clade
epapillate, coriaceous or leathery, dark brown, basal to the Lulworthiales clade.
sterile hairs enclosing ascoma, lacking Morphologically they differ in the mature
paraphyses, asci clavate, thin-walled, hamathecium, presence of periphyses, (absent
unitunicate, deliquescing early, ascospores in the Lulworthiales) but present in
fusiform, cylindrical, ellipsoidal, 0-3-septate, Koralionastetales; and the absence of apical
hyaline, with an appendage at each pole. mucous-filled polar appendages in
Antheridia, spermatia and trichogynes reported Koralionastes and Pontogeneia.
(Kohlmeyer and Kohlmeyer, 1979). All species
are parasites of the marine red algal genus Key to the genera
Ballia (B. callitrichia, B. hirsuta, B. scoparia)
often colonizing a single cell of the filamentous 1. On coral rock, associated with crustose sponges .......
thallus. Primarily known from Australia, New ............................................................ Koralionastes
1. On algae .................................................Pontogeneia
Zealand, Antarctic and sub-Antarctic waters,
and more recently from Argentina, Chile and
Koralionastes Kohlm. & Volkm.-Kohlm.,
Kerguelen and Falkland Islands waters
Mycologia 79: 765, 1987 ............................. (5)
(Kohlmeyer and Kohlmeyer, 1975a). Initially
K. angustus Kohlm. & Volkm.-Kohlm.,
Spathulospora was thought to be a primitive
Mycologia 79: 768, 1987.
ascomycete evolved from a red seaweed
K. ellipticus Kohlm. & Volkm.-Kohlm.,
ancestor, with its hyaline, unicellular
Mycologia 79: 765, 1987.
ascospores, conical appendages filled with
K. giganteus Kohlm. & Volkm.-Kohlm., Can. J.
“mucous”, hairs bearing antheridia, simple
Bot. 68: 1554, 1990.
trichogynes and asci that deliquesce early in
K. ovalis Kohlm. & Volkm.-Kohlm.,
development. Sequence data of the genera
Mycologia 79: 765, 1987 (Type species).
Spathulospora and Haloguignardia phylo-
K. violaceus Kohlm. & Volkm.-Kohlm., Can. J.
genetically place them in the Lulworthiales
Bot. 68: 1556, 1990.
although no formal taxonomic transfer has
A unique group of fungi occurring on
been undertaken (Inderbitzin et al., 2004;
coralline-coated rocks and sponges, known
Campbell et al., 2005) (Figs 62, 63).
from the Atlantic Ocean, Belize, Central
1. Ascomata without hairs, ascospores 40-50 × 4-9μm.. America (Kohlmeyer and Volkmann-
.............................................................. S. antarctica Kohlmeyer, 1987b) and Pacific Ocean,
1. Ascomata with hairs................................................. 2 Queensland, Australia (Kohlmeyer and
Volkmann-Kohlmeyer, 1990b). Characterised
2. Ascospores longer than 65 μm ................................. 3
2. Ascospores shorter than 65 μm ................................ 4
by large ascomata (500-1220 μm diam.),
simple septate paraphyses, clavate asci, short
3. Ascospores less than 14 μm wide, spathulate to pedunculate, deliquescing at maturity, no
spoon-shaped at the apices, 80-100 × 10-13 μm ........ apical apparatus, ascospores hyaline, thick-
.............................................................S. phycophila walled and septate near the apices, no
3. Ascospores wider than 14 μm, conical appendages, appendages, with spermatia.
70-104 × 16-23 μm ...................................S. adelpha

4. Ascospore tips spathulate, 39-62 × 8-12 μm, sub- 1. Ascospores ovoid, peridium 3-4 strata..................... 2
polar appendages......................................... S. lanata 1. Ascospores ellipsoidal, peridium 2-3 strata ............. 3
4. Ascospore tips rounded, 45-56 × 14-20 μm,
appendages polar........................................... S. calva 2. Ascospores 4-septate (1-6), 94-131 × 50-77 μm,
peridium 4 strata, 30-60 μm......................... K. ovalis
2. Ascospores 2-septate (2-6), 123-170 × 61-89 μm,
2. KORALIONASTETALES
peridium 3 strata, 125-190 μm ...............K. giganteus
Koralionastetaceae 3. Ascospores 4 septate (3-6), 85-130 × 25-34 μm,
This is a new order erected to peridium 3 strata, 50-70 μm................... K. violaceus
accommodate the genera Koralionastes and 3. Ascospores ellipsoidal, peridium 2 strata................. 4

132
 Fungal Diversity

4. Ascospores 6-septate (1-8), 81-127 × 27-45 μm, 1. Ascospores shorter than 200 µm, on other algal hosts
peridium 40-70 μm..................................K. ellipticus ................................................................................. 3
4. Ascospores 8-septate (6-10), 81-122 × 18-28 μm,
peridium 60-90 μm..................................K. angustus 2. Ascospores 215-325 × 14-21 µm, 10-13-septate .......
.................................................................P. cubensis
Pontogeneia Kohlm., Bot. Jahrb. 96: 200, 1975 2. Ascospores 280-350 × 12-14 µm, 4-5-septate ...........
................................................................. P. enormis
...................................................................... (8) 2. Ascospores 200-275 × 19-26 µm, 9-18-septate on the
P. calospora (Pat.) Kohlm., Bot. Jahrb. 96: 205, alga Microdictyon sp........................... P. microdictyi
1975.
Zignoella calospora Pat., J. Bot., Paris, 11: 242, 3. Ascospores 99-172 × 10-12 µm, 5-9-septate, on
1897. Padina.......................................................P. padinae
P. codiicola (E.Y. Dawson) Kohlm. & E. 3. Ascospores shorter than 100 µm, on other hosts...... 4
Kohlm., Marine Mycology. The Higher Fungi:
4. Ascospores 18-28 × 8-10 µm, 2-septate, with a large
350, 1979. central cell, on Egregia ................................P. erikae
Sphaerulina codiicola E.Y. Dawson, Occas. Pap. 4. Ascospores longer than 50 µm, no large central cell .
Allan Hancock Found. 8: 20, 1949. ................................................................................. 5
P. cubensis (Har. & Pat.) Kohlm., Bot. Jahrb.
96: 207, 1975. 5. Ascospores 54-85 × 8-14 µm, 3-6-septate, on
Zignoella cubensis Har. & Pat., Bull. Soc. Mycol. Castagnea chordariiformis ....................P. calospora
Fr. 20: 65, 1904. 5. Ascospores wider than 20 µm.................................. 6
P. enormis (Pat. & Har.) Kohlm., Bot. Jahrb.
96: 208, 1975. 6. Ascospores 52-79 × 20-28 µm, 3-septate, on Codium
Zignoella enormis Pat. & Har., J. Bot. Paris 17: species..................................................... P. codiicola
228, 1903. 6. Ascospores 72-100 × 21-30 µm, 2-5-septate, on
Valoniopsis pachynema ................... P. valoniopsidis
P. erikae Kohlm., Bot. Mar. 24: 16, 1981.
P. microdictyi Kohlm. & Volkm.-Kohlm., 3. MAGNAPORTHALES see p. 137, 144
Mycol. Res., 2009. DOI 10.1016/j.mycres.
2008.110314 4. PHYLLACHORALES
P. padinae Kohlm., Bot. Jahrb. 96: 201, 1975
(Type species). Key to Phyllachorales and Phyllachorales
P. valoniopsidis (Cribb & J.W. Cribb) Kohlm., incertae sedis
Bot. Jahrb. 96: 209, 1975.
Zignoella valoniopsidis Cribb & J.W. Cribb, Univ. 1. Ascospores unicellular ............................................. 2
Queensl., Pap. Dept. Bot. 4: 41, 1960. 1. Ascospores septate ................................................... 3
Ascomata subglobose to ovoid,
superficial or partly immersed, ostiolate, 2. Ascospores ellipsoidal, 15-18 × 5-6 µm parasitic on
the alga Apophlaea lyallii .... Polystigma apophlaeae
papillate or epapillate, coriaceous, dark brown, 2. Ascospores subglobose to ellipsoidal, 13.5-18.5 × 9-
solitary or gregarious, paraphyses septate, thick, 12 µm, saprobic on Spartina......................................
asci clavate or fusiform, thin-walled, .............................................Phyllachora paludicola
unitunicate, deliquescing early, no apical
apparatus, ascospores filiform, ellipsoidal or 3. Ascospores 1-septate, 18-28 × 6-8 µm, parasitic on
brown algae......................Phycomelaina laminariae
fusoid-ellipsoidal, curved, 3-13-septate, hyaline. 3. Ascospores 3-septate................................................ 4
Parasitic species on a wide range of marine
algae. Initially assigned to Zignoella, but 4. Ascospores 26-31 × 6-8.5 µm, not constricted at the
Kohlmeyer (1975b) established a case for the septa, many oil globules per cell, no sheath
erection of the new genus Pontogeneia because ....................................... Marinosphaera mangrovei
4. Ascospores 25-39 × 12-14 µm, with a sheath
of different morphological features. ........................................... Mangrovispora pemphis
Ascospores are hyaline, which vary greatly in
their morphology, particularly in their length Phyllachoraceae
and degree of septation.
Phyllachora Nitschke ex Fuckel, Jahrb. des
1. Ascospores longer than 200 µm, on the brown alga Nassauischen Vereins Naturk. 23-24: 216,
Halopteris ................................................................ 2 1870.............................................................. (1)
133
Fig. 63. One of the MPTs inferered from LSU rDNA sequences of selected lulworthialean taxa, to show the
phylogenetic relationship of the genus Rostrupiella. Bootstrap values higher than 50% are given above branches.

Ph. paludicola Kohlm. & Volkm.- Kohlm., Polystigma DC., Fl. Fr. 6: 164, 1815........... (1)
Mycologia 95: 120, 2003. P. apophlaeae Kohlm., Bot. Mar. 24: 13, 1981.
Described from dead leaves of Spartina Parasitic on the alga Apophlaea lyallii
alterniflora and collected widely in the USA. causing discolouration of the thallus, and
Well developed, prominent stromata 1.3-4 cm known from a limited number of collections
long, about 0.5 cm wide, with 5-20 ascomata, (Kohlmeyer and Demoulin, 1981). Ascospores
ostiolate, coriaceous, light brown, arranged in 15-18 × 5-6 µm, hyaline, unicellular and
two to four parallel rows, no necks, periphysate; lacking a sheath or appendage. Further
septate paraphyses, apically free; asci collections are required to determine its
cylindrical, short pedunculate, thick-walled, phylogenetic relationship within the
unitunicate, without an apical apparatus, J-; ascomycetes. Some 70 names are listed for
ascospores subglobose to ellipsoidal, Polystigma but many are referred to other
unicellular, hyaline, orange- yellowish in mass, genera, therefore the position of P. apophlaeae
and lacking a sheath or appendages (Fig. 65) remains to be resolved (Index Fungorum).
(Kohlmeyer and Volkmann-Kohlmeyer, 2003a). Polystigma sonneratiae has been described as a
Well placed in the Phyllachoraceae, parasite causing leaf spots of the mangrove tree
Phyllachorales. Sonneratia caseolaris and known from

134
 Fungal Diversity

a b

Fig. 64. Rostrupiella danica. a. Bell-like structure (arrowed) and base of the neck (arrowed N). b. Higher
magnification of the neck region (arrowed N) with bell layer. c. Neck within the woody substratum filled with
pseudoparenchymatous hyaline cells (arrowed). d. Base of the neck at the tip of the centrum. e. Bell-like structure
pseudoparenchymatous cells (arrowed o) and bladder-like cells (arrowed B). Bars = 10 μm. Reproduced with
permission from Botanica Marina.

135
collections in Australia and Philippines (Hyde appendages (Fig. 66). The species is easily
and Cannon, 1992). This species is of identified by its wide, regularly septate
terrestrial origin and may not be correctly paraphyses.
assigned to this genus (Hyde and Cannon, Hyde (1989b) initially placed the genus,
1992). with reservations, in the Phyllachoraceae, but
this has received little support (Kirk et al.,
2001). The ascus tip is thickened, with a small
flattened ocular chamber but no spore
discharge has been observed (Read et al.,
1995). Molecular results, based on SSU and
LSU rDNA analyses, revealed that
Fig. 65. Phyllachora paludicola. Ascospore. Bar = 10
µm.
Marinosphaera does not have any affinities
with the Phyllachorales. Phylogenetically, M.
PHYLLACHORALES incertae sedis mangrovei is located between the
Halosphaeriales and the Microascales, but
Mangrovispora K.D. Hyde & Nakagiri, Syst. without any closely related taxa (Fig. 67).
Ascomycetum 10: 19, 1991 ......................... (1) Therefore its taxonomic position is not fully
M. pemphi K.D. Hyde & Nakagiri, Syst. resolved at this time, suggesting a further group
Ascomycetum 10: 20, 1991 (Type species). of fungi remains to be discovered. This is a
Described from intertidal decayed roots common species, often found as an early
of Pemphis acidula with ascomata that are colonizer of mangrove wood (Alias, 1996).
immersed beneath a thin stroma, globose to
Phycomelaina Kohlm., Phytopathol. Z. 63:
subglobose, hyaline, membranous, ostiolate,
350, 1968...................................................... (1)
papillate, solitary or gregarious, periphysate,
P. laminariae (Rostr.) Kohlm., Phytopathol. Z.
paraphyses numerous, branched, septate in a
63: 350, 1968 (Type species).
gel, asci cylindrical, fissitunicate, pedunculate, Dothidella laminariae Rostr., Bot. Tidsskr. 19:
unitunicate, thin-walled, faint apical tube and 213, 1895.
apical thickening, ascospores ellipsoidal to Endodothella laminariae (Rostr.) Theiss. &
fusiform, hyaline, 3-septate, slightly Sydow, Ann. Mycol. 13: 582, 1915
constricted at the septa, thin-walled and with a Placostroma laminariae (Rostr.) Meyers,
Mycologia 49: 480. 1957.
sheath like structure (Hyde, 1991a). Hypoderma laminariae G.K. Sutherl., New
Hawksworth et al. (1995) refered the genus to Phytol. 14: 190, 1915.
the Hyponectriaceae while Wang and Hyde Pseudostroma black ellipsoidal spots on
(1999) excluded it from this family. Further cortex of algae, ascomata subglobose or
studies are required before this little collected ampulliform, immersed in the pseudostroma,
species can be assigned with confidence to any ostiolate, epapillate, leathery, hyaline to brown,
family. clypeus dark brown, gregarious, papillae with
periphysoids. Originally described by Rostrup
Marinosphaera K.D. Hyde, Can J. Bot. 67: (1894) it has been transferred variously to other
3080, 1989.................................................... (1) genera until Kohlmeyer (1968b) assigned it to
◙ M. mangrovei K.D. Hyde, Can J. Bot. 67: this genus. It occurs on the brown marine algae:
3080, 1989 (Type species). Laminaria species and possibly Alaria
Ascomata ellipsoidal, globose, esculenta. Still not assigned with authority to
subglobose, elongate, immersed, ostiolate, an order and requiring further investigation.
papillate, membranous, light to dark brown,
solitary to gregarious, periphysate, paraphyses Unitunicate ascomycetes family
wide, simple, septate, asci clavate, short incertae sedis
pedunculate, persistent, unitunicate, thin-
walled, J- subapical plate and pore, ascospores Key to unitunicate ascomycetes incertae sedis
broad ellipsoidal to fusiform, initially 0-septate
but becoming distinctly 3-septate, hyaline, 1. In seaweeds .............................................................. 2
smooth-walled and lacking a sheath or 1. In wood and other cellulosic materials..................... 5
136
 Fungal Diversity

2. On green algae ......................................................... 3 cylindrical, brown and thick-walled at the base.

2. On red seaweeds ...................................................... 4 Not only is the genus different from
Spathulospora, but is sufficiently so to warrant
3. Ascospores elongate ellipsoidal, 8.5-17.5 × 3-5 µm,
on Prasiola..................................Mastodia tessellata erection of a new family, the
3. Ascospores broad ellipsoidal, 8.5-14 × 3.5-7 µm, on Hispidicarpomycetaceae (Nakagiri, 1993a).
Blidingia.................................. Turgidosculum ulvae Diagnostic features are the hyphoid thalli,
spermadochia multiverticilliate and the hispoid
4. Ascospores with polar appendages, 10-18 × 3-6 µm ascoma, which develops by radial growth of
.............................................. Retrostium amphiroae
4. Ascospores lacking appendages, 35-68 × 12-20 µm
hyphae of the ascoma initial (Nakagiri, 1993).
............................ Hispidicarpomyces galaxauricola The affinity of Spathulospora with the
Lulworthiales raises the question as to the
5. Ascospores unicellular ............................................. 6 phylogenetic position of Hispidicarpomyces.
5. Ascospores septate ................................................... 7
Spathulosporaceae
6. Ascospores hyaline .................................................. 7
6. Ascospores yellow to pale brown............................. 8
Retrostium Nakagiri & Tad. Ito, Mycologia 89:
485, 1997...................................................... (1)
7. Ascospores smooth walled, filiform 110-180 × 3-4 R. amphiroae Nakagiri & Tad. Ito, Mycologia
µm ............................. Gaeumannomyces medullaris 89: 485, 1997 (Type species).
7. Ascospores verrucose, broadly ellipsoidal, 14-18 × Ascomata are conical to hemispherical,
7.5-9.5 µm .............................. Papulosa amerospora
superficial, solitary or united, carbonaceous,
8. Ascospores 5-7-septate, 34-44 × 8-10 µm..................
ostiolate, epapillate, and black. Peridium 45-85
.....................................Pseudohalonectria halophila µm thick, but thinner at the base, and two-
8. Ascospores 3-septate, 37-66 × 9.5-14 µm .................. layered. Ostiolar canal forming a tube-like
............................................. Buergenerula spartinae projection into the centrum but lacking
periphyses. Asci subglobose to obpyriform,
Hispidicarpomycetaceae thin-walled, unitunicate early deliquescing and
Hispidicarpomyces Nakagiri, Mycologia 85: without an apical apparatus. Ascospores oblong
639, 1993...................................................... (1) to ellipsoidal, unicellular, hyaline, with polar
H. galaxauricola Nakagiri, Mycologia 85: 639, mucilaginous, subglobose appendages.
1993 (Type species). Penicilliate spermatophores and
An unusual ascomycete with large trichogynes are also present. Nakagiri and Ito
ascomata on the red alga Galaxaura falcata, (1997) are of the opinion that Retrostium
700-1600 µm high, 800-2100 µm diam. represents a phylogenetic link between
Breaking through from the medullary layer of spathulosporalean fungi and other algal-
the host, solitary, ostiolate, epapillate, black inhabiting marine fungi in Spathulospora, the
arising from collapsed spermodochia. Peridium genera Chadefaudia and Haloguignardia. This
composed of three-layers, the outer layer with observation was prophetical in view of the
thick-walled short hyphal projections. recent referral of Haloguignardia to the order
Paraphyses 1-3-septate, branched. Asci are Lulworthiales (Inderbitzin et al., 2004;
obclavate to obpyriform, thin-walled, Campbell et al., 2005).
unitunicate, deliquescing early and lacking an
apical apparatus. Ascospores are ellipsoid to MAGNAPORTHALES
elliptic-fusiform, unicellular, hyaline and Magnaporthaceae
without appendages or sheath. A feature of the
species is the production of verticilliate Since this monograph was accepted for
spermatophores, with closely packed publication, Thongkantha et al. (2008) erected
ampulliform phialides terminating in a column the order Magnaporthales. The genera
and producing hyaline, thin-walled spermatia, Buergenerula, Gaeumannomyces and
inflated at the apex and cupulate at the base. Pseudohalonectria are now included in this
Trichogynes are 2-4-septate, simple, order. See page 144 for further discussion.

137
Buergenerula Syd., Ann. Mycol. 34: 392, hyaline, lacking a sheath or appendages.
1936.............................................................. (1) A species widely reported from Spartina
◙ B. spartinae Kohlm. & R.V. Gessner, Can. J. species, usually growing on senescing or dead
Bot. 54: 1764, 1976. culms. It has been confused with Sphaerulina
Ascomata subglobose to obpyriform, pedicellata, initially described from wood
immersed, ostiolate, papillate, coriaceous, light panels (Johnson, 1956) because of the thick-
to dark brown, solitary or gregarious, necks walled asci when young and thought to be
periphysate, paraphyses filamentous, simple, or bitunicate. Buergenerula spartinae is
branched, septate, asci cylindrical to paraphysate, with unitunicate thick-walled asci
subfusiform, short pedunculate, unitunicate, and producing hyphopodia on the host.
thick-walled, with an apical apparatus, Although a Pyricularia anamorph has been
ascospores clavate, thick at the apex, 3-septate, reported for the genus, no anamorph has been

a d

Fig. 66. Marinosphaera mangrovei. a. Immersed ascomata with long necks on the wood surface. b. Cylindrical asci
with subapical structures. c. Wide-septate, chain-like paraphyses. d. Ornamented ascospores full of oil globules. Bars
a = 200 µm; b = 20 µm; c, d = 10 µm.

138
 Fungal Diversity

Fig. 67. One of two MPTs inferred from combined SSU and LSU rDNA of Marinosphaera mangrovei, generated with
maximum parsimony analysis. Bootstrap values higher than 50% given above branches. Scale bar indicates 10
character state changes.

139
reported for B. spartinae, except for spermatial Mastodiaceae
like structures (Kohlmeyer and Gessner, 1976). Turgidosculum Kohlm. & E. Kohlm., Bot.
Jahrb. Syst. 92: 429, 1972............................ (1)
Gaeumannomyces Arx & D.L. Olivier, Trans. T. ulvae (G.M. Reed) Kohlm. & E. Kohlm.,
Br. Mycol. Soc. 35: 32, 1952 ....................... (1) Bot. Jahrb. Syst. 92: 429, 1972 (Type species).
G. medullaris Kohlm., Volkm.-Kohlm. & O.E. Guignardia ulvae G. M. Reed, Univ. Calif.,
Erikss., Mycologia 87: 540, 1995. Berkley, Publ. Bot. 1: 160, 1902.
Anamorph: Trichocladium medullare
Kohlm. & Volkm.-Kohlm.
Ascomata ellipsoidal, immersed, ostiolate,
coriaceous, dark brown, solitary, necks
cylindrical and periphysate, paraphyses
numerous, unbranched, septate, tapering, asci
fusoid to cylindrical, short pedunculate, thin-
walled, unitunicate, with an apical ring,
persistent, ascospores filiform, slightly tapering
towards the base, without a sheath or apical
Fig. 68. Pseudohalonectria halophila. Ascospore. Bar =
appendages. The only marine species in the 5 μm.
genus and found on the inundated parts of the Ascomata subglobose, immersed in the
culms of Juncus roemerianus. This species algal thallus between the upper and lower algal
differs from other Gaeumannomyces species in cell layers, dark brown, papillate, ostiolate
having a Trichocladium anamorph (Kohlmeyer closed by a gelatinous plug, periphysate, no
et al., 1995c). Cannon (1994) and Cannon and paraphyses, asci clavate developing all around
Kirk (2007) referred the genus to the the inside of the ascomata, pedunculate,
Magnaporthaceae while Thongkantha et al. unitunicate, thick-walled but deliquescing,
(2009) have erected a new order for the family. ascospores ellipsoidal to ovoid, unicellular,
hyaline, lacking a sheath or appendages. A
Pseudohalonectria Minoura & T. Muroi, Trans. parastic ascomycete on Blidingia minima var.
Mycol. Soc. Jpn. 19: 132, 1978.................... (1) vexata, and with a restricted known distribution
P. halophila Kohlm. & Volkm.-Kohlm., Bot. (Schatz, 1980; Kohlmeyer and Volkmann-
Mar. 48: 310, 2005. Kohlmeyer, 2003b).
Ascomata obpyriform, immersed,
ostiolate, coriaceous, light to dark brown, long Mastodia (Hook. f. & Harv.) Hook. f. & Harv.,
neck (130-410 µm) protruding beyond the Fl. Antrac., Part II: 499, 1847 ...................... (1)
substratum, paraphyses thin-walled, septate, M. tessellata (Hook. f. & Harv.) Hook. f. &
unbranched, asci fusiform, short pedunculate, Harv., Fl. Antarc., Part II: 499, 1847.
unitunicate, persistent, with a refractive tip Leptogiopsis complicatula Nyl., Flora, Jena 67:
below the ring, J-, ascospores ellipsoidal to 211, 1884.
Turgidosculum complicatulum (Nyl.) Kohlm. & E.
fusiform, 5-6 (-7)-septate, slightly constricted
Kohlm., Marine Mycology, The Higher Fungi: 361,
at the septa, echinulate, hyaline and pale brown, 1979.
lacking a sheath or appendages (Fig. 68). A Guignardia alaskana G.M. Reed, Univ. Calfornia
frequent inhabitant of fragmented old leaves Publ. Bot. 1: 161, 1902.
and culms of Juncus roemerianus, and the first Laestadia alaskana (G.M. Reed) Sacc. & D.
Sacc., Syll. Fung. 17: 576, 1905.
Pseudohalonectria species from marine waters
Laestadia prasiolae G. Winter, Hedw. 26: 16,
(Kohlmeyer et al., 2005). It most closely 1887.
resembles P. adversaria and P. fuxianii, but Guignardia prasiolae (G. Winter) Lemmerm.,
differs in that it is obligately marine and does Naturwiss. Ver. Bremen, 17: 199, 1901.
not grow on wood. It also had distinctly Laestadiata tessellta G. Winter ex Har., Algues,
in Mission Sci. Cap Horn, 5: 29, 1882-1883. (nomen
broader ascospores than P. adversaria, and
nudum).
more septa than P. fuxianii (Kohlmeyer et al., Physalospora prasiolae Har., J. Bot., Paris 1: 133,
2005). 1887. (nomen nudum).

140
 Fungal Diversity

Kohlmeyera complicatula (Nyl.) S. Schatz, 2. Ascospores septate ................................................... 4

Mycologia 72: 114, 1980.
Plagiostoma prasiolae (Winter) Clauzade, 3. Ascus thick-walled, with an ocular chamber with a
Diederich & Cl. Roux, Bull. Soc. Linn. Provence. Num. net-like apical apparatus, ascospores hyaline, 59-77 ×
spec. 1: 47, 1989. 33-44 µm...................................... Hapsidascus junci
Ascomata subglobose, immersed in the 3. Ascus thin-walled, no apical apparatus, ascospores
algal thallus, ostiolate, epapillate, coriaceous, pale brown, 20-28 × 7-10 µm ..... Rhizophila marina
dark brown, solitary or gregarious, necks 4. Ascospores with more than 3 septa....... Saccardoella
periphysate, paraphyses not observed in mature 4. Ascospores 3-septate................................................ 5
specimens, centrum filled with a gelatinous
matrix, asci clavate to subcylindrical, short 5. Ascus clavate to subcylindrical, with an operculum,
pedunculate, unitunicate, lacking an apical on brown algae............................ Orcadia ascophylli
5. Asci lacking an operculum....................................... 6
apparatus, deliquescing, ascospores elongate-
ellipsoidal to cylindrical, rounded ends, 6. Ascomata sea-green to blue, apical ring cylindrical or
unicellular, hyaline, lacking a sheath or wedge-shaped, ascospores 18-24 µm.........................
appendages. Kohlmeyer and Kohlmeyer (1979) ................................................Aquamarina speciosa
give a detailed history of the placement of this 6. Ascomata pale coloured, ascus ring subapical,
ascospores striate 25.5-40 × 7-10 µm ........................
taxon. A much confused taxon in need of re- ........................................................ Aropsiclus junci
evaluation, especially at the molecular level.
7. Ascospores 0-septate, with bipolar crown of delicate
Papulosaceae appendages....................................................... Eiona
Papulosa Kohlm. & Volkm.-Kohlm., Syst. 7. Ascospores 1-septate................................................ 8
Ascomycetum 11: 96, 1993 ......................... (1)
8. Ascospores with thread-like spirally unfolding
◙ P. amerospora Kohlm. & Volkm.-Kohlm., appendages............................................................... 9
Syst. Ascomycetum 11: 96, 1993 (Type 8. Ascospore appendages not spirally arranged ......... 10
species).
Ascomata subglobose to ellipsoidal, 9. On wood, often associated with sand.... Dryosphaera
immersed, ostiolate, papillate, coriaceous, dark 9. On seaweed, Fucus .................................... Crinigera
brown, solitary, necks periphysate, lateral 10. Ascospores with polar and lateral appendages...........
paraphyses tapering toward the tip, simple or ................................................................ Marisolaris
branched, asci cylindrical, short pedunculate, 10. Ascospores with a mucilaginous sheath...........Biflua
persistent, with an aical ring, J+, ascospores
broadly ellipsoidal, unicellular, hyaline, Aquamarina Kohlm., Volkm.-Kohlm. & O.E.
verruculose, with a thin gelatinous sheath. Erikss., Mycol. Res. 100: 393, 1996 ............ (1)
Saprophyte of senescent culms of Juncus A. speciosa Kohlm., Volkm.-Kohlm. & O.E.
roemerianus (Kohlmeyer and Volkmann- Erikss., Mycol. Res. 100: 393, 1996.
Kohlmeyer, 1993b). Winka and Eriksson (2000) Ascomata subglobose, immersed,
refer P. amerospora to a new family, ostiolate, coriaceous, sea-green to blue, neck
Papulosaceae in the sub-class Sordariomy- long (165-345 µm) within the host, periphysate,
cetidae, and most closely related to the orders paraphyses septate, unbranched in a gelatinous
Ophiostomatales, Diaporthales, Sordariales and matrix, asci cylindrical, short pedunculate,
Phyllachorales. Insufficient unitunicate marine thin-walled, unitunicate with an apical ring J-,
taxa have been sequenced to enable this family ascospores fusiform, 3-septate, not constricted
to be assigned to any order. at the septa, hyaline lacking a sheath or
appendages (Kohlmeyer et al., 1995d).
Untunicate ascomycetes genera Sporulating on the base of culms of Juncus
roemerianus and regarded as an obligately
incertae sedis
marine ascomycete (Kohlmeyer et al., 1995d),
1. Ascomata perithecoid............................................... 2 characterized by sea-green to blue ascomata
1. Ascomata cleistothecial............................................ 7 that are deeply embedded in the substratum.
Kohlmeyer et al. (1995d) were unable to
2. Ascospores 0-septate................................................ 3 suggest a suitable family for its assignment and
141
further evaluation is dependent on fresh have shown the species is based on two taxa,
material for a molecular study. and erected the genus Dryosphaera for the
material on wood.
Aropsiclus Kohlm. & Volkm.-Kohlm., Syst.
Ascomycetum 13: 24, 1994 ......................... (1) Dryosphaera Jørg. Koch & E.B.G. Jones, Can.
A. junci (Kohlm. & Volkm.-Kohlm.) Kohlm. J. Bot. 67: 1184, 1989 .................................. (3)
& Volkm.-Kohlm., Syst. Ascomycetum 13: 24, ◙ D. navigans Jørg. Koch & E.B.G. Jones, Can.
1994. J. Bot. 67: 1185, 1989 (Type species).
Sulcospora junci Kohlm. & Volkm.-Kohlm., Syst.
Ascomycetum 11: 100, 1993.
D. tropicalis Kohlm. & Volkm-Kohlm., Can. J.
Ascomata embedded in the outer small- Bot. 71: 992, 1993.
celled tissue of Juncus roemerianus, cream- D. tenuis Andrienko, Ukr. Bot. J. 58: 244-247,
coloured, necks breaking through the cuticle, 2001.
periphysate, periphysoids present, ascospores Cleistothecia globose, slightly flattened,
hyaline, 3-septate with a striated spore wall superficial, white-light buff coloured,
(Kohlmeyer and Volkmann-Kohlmeyer, 1994). coriaceous, gregarious, attached to wood by
The genus cannot be assigned to any order or thick-walled hyphae branched at their tips,
family of the Ascomycota. ascomata covered with short appendages, each
with a stalk and a terminal crown of repeatedly
Biflua Jørg. Koch & E.B.G. Jones, Can. J. Bot. divided branches, asci numerous, developed on
67: 1187, 1989 ............................................. (1) lobed hymenium, thick-walled when young,
◙ B. physasca Jørg. Koch & E.B.G. Jones, Can. (possibly bitunicate), broadly clavate, short
J. Bot. 67: 1187, 1989 (Type species). pedunculate, persistent, paraphyses absent,
Cleistothecia globose or slightly flattened, ascospores 1-septate, constricted at the septum,
coriaceous, white-light buff colour, gregarious, hyaline and appendaged. Appendages appear as
paraphyses absent, attached to wood by thick- a ball of spirally arranged threads frequently
walled hyphae with broad ascomatal terminating in ball-like structures (Koch and
appendages, asci clavariform to cylindrical, Jones, 1989).
pedunculate, persistent, lacking an apical pore, Dryosphaera tenuis (Fig. 69) has been
ascospores ellipsoidal, 1-septate, constricted at described by Andrienko (2001), from the leaf
the septum, hyaline, with a mucilaginous of a Zostera sp. in the Odessa region of the
sheath. Ukraine. It has larger asci 17.3-38.7-61.9 × 5.7-
Kirk et al. (2001) relegate this genus to 12.2-19 µm, slightly longer ascospores (8.6-
genera incertae sedis. One of three marine 11.8-13.3 × 2.2-3.0-3.9 µm) and significantly
cleistotheical ascomycetes, with sterile appen- longer appendages (9.4-14.2-17.6 × 0.3-0.7-1.1
dages on the peridium and growing on wood in µm) than the previously described species.
association with sand (Koch and Jones, 1989). Papers published by Russian mycologists on
Their taxonomic relationship is unknown, and marine fungi are rarely cited in the general
all require study at the molecular level, but are literature (Kopytina and Andrienko, 1998) and
infrequently collected, possibly they are consequently new taxa remain unknown.
overlooked because of the small size of the
1. On Zostera leaves, ascospores 8.6-13.3 × 2.2-3.9 μm,
ascomata. These cleistothecial ascomycetes are
appendages 9.4-17.6 × 0.3-1.1 μm.............. D. tenuis
generally confined to cold waters and develop 1. On wood associated with sand ................................. 2
on cold incubation of wood samples, and
remain in good condition for 2-3 weeks (Koch, 2. Ascospores 12-15 × 4-5.5 μm................ D. tropicalis
pers. comm.). 2. Ascospores 8-12 × 3-5 μm..................... D. navigans

Crinigera I. Schmidt, Nat. Naturschutz Eiona Kohlm., Ber. Dtsch. Bot. Ges. 81: 58,
Mecklenburg 7: 11, 1969 ............................. (1) 1968.............................................................. (1)
C. maritima I. Schmidt, Nat. Naturschutz E. tunicata Kohlm., Ber. Dtsch. Bot. Ges. 81:
Mecklenburg 7: 11, 1969 (Type species). 58, 1968 (Type species).
A genus of doubtful validity collected on Ascomata cleistothecial, subglobose,
the seaweed Fucus. Koch and Jones (1989) superficial, no ostiole, coriaceous, brown to
142
 Fungal Diversity

Marisolaris Jørg. Koch & E.B.G. Jones, Can. J.

Bot. 67: 1190, 1989...................................... (1)
M. ansata Jørg. Koch & E.B.G. Jones, Can. J.
Bot. 67: 1193, 1989 (Type species).
Ascomata globose, coriaceous, white-
light buff coloured, gregarious, superficial on
wood, and attached to the surface by thick-
walled hyphae, ascomata covered by numerous
simple horn-like appendages, tapering,
paraphyses absent, asci broadly clavate,
Fig. 69. Dryosphaera tenuis. Ascospore. Bar = 5 μm. pedunculate, persistent, bitunicate with an
exotunicate and endotunica visible when
green, gregarious, lacking paraphyses, asci mounted in water, ascsopores 1-septate, not
broadly clavate or ellipsoidal, unitunicate, thin- constricted at the septum, hyaline with single
walled, scattered throughout the ascoma cavity, polar and four equatorial appendages, each
early deliquescing and lacking and apical appendage comprising an inner part which is
apparatus, ascospores ellipsodial, unicellular., looped like a bridge and an outer part a sheath
hyaline, with a crown of appendages at both that envelopes the spore. Another genus, which
ends of the spore (Kohlmeyer and Kohlmeyer, at the present time, cannot be assigned to any
1979). Ascospore appendages formed by taxonomical group and is only known from a
fragmentation of a sheath (Jones and Moss, few collections on wood associated with sand
1978). This species is in need of a molecular (Koch and Jones, 1989).
study to determine its ordinal rank as it may
not belong in the Eurotiaceae. Collected in Orcadia G.K. Sutherl., Trans Br. Mycol. Soc. 5:
abundance on wood associated with sand 151, 1915...................................................... (1)
(Koch and Jones, 1983; Farrant et al., 1985). O. ascophylli G.K. Sutherl., Trans Br. Mycol.
Soc. 5: 151, 1915 (Type species).
Hapsidascus Kohlm. & Volkm.-Kohlm., Syst. Ascomata subglobose or pyriform,
Ascomycetum 10: 113, 1991 ....................... (1) immersed, ostiolate, papillate, coriaceous,
H. hadrus Kohlm. & Volkm.-Kohlm., Syst. subhyaline, gregarious, necks a papilla or long
Ascomycetum 10: 115, 1991 (Type species). (80-180 µm), periphysate, paraphyses? septate,
An unusual ascomycete with large simple or anasotmosing, deliquescing early,
ellipsoidal ascomata (>750 µm), deeply asci clavate, subcylindrical or subfusiform,
embedded in mangrove roots of Rhizophora short pedunculate, thin-walled, unitunicate,
mangle, necks almost as thick as the ascoma, with an apical thickening but no apical
and the ostiolar canal filled with a network of apparatus, operculum present, ascospores
thin hyphae, embedded in a matrix. Peridium cylindrical, slightly curved, 3-septate,
3-layered and a paraphysate hamathecium, the constricted at the septa, hyaline, no sheath or
upper third of the locule filled with thin appendages. Initially referred by Sutherland
anastomosing hyphae in a gelatinous matrix. (1915) to the Hyponectriaceae, Xylariales.
Asci thick-walled, not fissitunicate with an Rossman et al. (1999) excluded the genus from
ocular chamber with a net-like apical apparatus, the Hypocreales because of the reported
J-, and ascospores ellipsoid to ovoid, 59-77 × operculate asci, broad opening of the ascomata,
33-44 µm, unicellular and hyaline (Kohlmeyer and numerous interthecial elements
and Volkmann-Kohlmeyer, 1991b). No known (Kohlmeyer and Kohlmeyer, 1979). A poorly
anamorph and monotypic. It superficially researched species, parasitic/saprophytic on the
resembles Lignincola tropica or Saagaromyces larger brown algae (Ascophyllum, Fucus,
abonnis, but differs in ascoma ontogeny, asci Pelvetia). Rossman et al. (1999) suggested an
with a net-like substructure to the ascus tip and affinity with the Pezizales because of the
ascospores that are unicellular. operculate asci.

143
Rhizophila K.D. Hyde & E.B.G. Jones, Saccardoella transylvanica, but Barr (1990b)
Mycotaxon 34: 527, 1989 ............................ (1) does not concur with that view. Geographically
◙ R. marina K.D. Hyde & E.B.G. Jones, the genus is well distributed with terrestrial,
Mycotaxon 34: 528, 1989 (Type species). marine and freshwater species (Hyde, 1992;
Ascomata large (500-965 µm wide, 425- Tsui et al., 1998).
730 µm long), globose to subglobose,
immersed, ostiolate, papillate, periphysate, 1. Ascospores with 7-9 septa, 26-33 × 6-8 µm ..............
coriaceous, dark brown to black, solitary or ..............................................................S. mangrovei
1. Ascospores with fewer than 7 septa ......................... 2
gregarious, paraphyses irregular shaped cells,
septate, asci clavate, pedunculate, unitunicate, 2. Ascospores with (4-) 5- (-6) septa, 19-26 × 6-8 µm ..
thin-walled, with a slight apical thickening, no ........................................................... S. rhizophorae
apical apparatus, ascospores ellipsoidal to 2. Ascospores with 3 septa, 25-31 × 7.5-10 µm.............
fusiform, unicellular, hyaline but becoming .......................................................... S. marinospora
yellowish brown, lacking a sheath or
appendages. Often found deeply embedded in MAGNAPORTHALES
mangrove wood and the droppers of MAGNAPORTHACEAE
Rhizophora species (Hyde and Jones, 1989a,
Jones, unpublished data). Canon and Kirk (2007) accepted the
Similarities with the Phyllachoraceae and Magnaporthaceae in the Sordariomycetes, a
Glomerella were reported by Hyde and Jones small family of unitunicate perithecial
(1989a), but molecular sequences suggest that ascomycetes, comprising 13 genera and 93
it may be better placed in the Microascales or species (Kirk et al., 2008). Gaeumannomyces
Hypocreales (Spatafora and Blackwell, 1994; (2 species) and Magnaporthe (5 species) are
Winka and Eriksson, 2000). the key members of the family as many species
are economically important plant pathogens.
Saccardoella Speg., Michelia 1: 461, 1879…(3) Magnaporthe grisea (anamorph Pyricularia
S. mangrovei K.D. Hyde, Mycologia 84: 803, oryzae) is the cause of rice blast and a wide
1992. variety of plants (Yaegashi and Herbert, 1976).
S. marinospora K.D. Hyde, Mycologia 84: 806, Because few of the Magnaporthaceae have
1992. been sequenced, placement at the ordinal level
S. rhizophorae K.D. Hyde, Mycologia 84: 806, has remained unresolved (Hibbett et al., 2007,
1992. while Lumbsch and Hundorf (2007) and Zhang
Ascomata globose to subglobose, et al. (2006) merely refer the family to the
immersed or erumpent, ostiolate, papillate, Sordariomycetidae along with two other poorly
coriaceous, carbonaceous, solitary or studied families, Annulatascaceae (Wong et al.,
gregarious, dark brown to black, clypeate, no 1998) and Papulosaceae (Winka and Eriksson,
periphyses, paraphyses numerous, filamentous, 2000).
hypha-like, branching and anastomosing in Thongkantha et al. (2008) investigated
gelatinous matrix, asci long-cylindrical, short the molecular phylogeny of selected genera in
pediculate, thin-walled, unitunicate, apically the Magnapothaceae (Buergenerula spartinae,
flattened or flattened with a subapical ring, Gaeumannomyces spp., Magnaporthe spp.,
ascospores fusiform, 3-9-septate, not Mycoleptodiscus coloratus, Ophioceras spp.
constricted at the septa, with acute ends, and Pseudohalonectria spp.), using 18S and
variable septation, hyaline, with a thin to well 28S rDNA phylogeny. These genera formed a
developed gelatinous sheath. A genus referred monophyletic clade, closely related to the
to the Clypeosphaeriaceae and Barr (1994) Diaporthales and Ophiostomatales. However,
later revised and extend the limits of the genus. the Magnaporthaceae could not be
There is confusion about the ascus structure in accommodated in any known fungal order and
Saccardoella as in slide preparations the ascus consequently they erected a new order, the
may appear thickened and behave like a Magnaporthales (Thongkantha et al., 2008).
bitunicate ascus. Mathiassen (1989) was of the Marine representatives of the Magna-
opinion that the asci are bitunicate in porthaceae, Magnaporthales are Buergenerula
144
 Fungal Diversity

spartinae, Gaeumannomyces medullaris and accommodate the little known genera

Pseudohalonectria halophila. Only G. Koralionastes and Pontogeneia, previously not
medullaris has a known anamorph in assigned with authority to any group (Campbell
Trichocladium (T. medullare). The anamorphs et al., 2008). Molecular studies have also
of Gaeumannomyces are generally Phialo- enabled a better understanding of the generic
phora species. relationships of many marine ascomycetes:
Ascomycetes, and their anamorphs Corollospora (Campbell et al. (2002),
comprise the largest marine fungal group (424 Halosarpheia, Lignincola and Nais (Pang et al.,
+ 94 species, respectively) and the most 2003a, b), Antennospora (Pang et al., 2008)
intensively studied. The orders Halosphaeriales and anamorphic Lulworthiales (Jones et al.,
and Lulworthiales have been extensively 2008).
studied at the molecular level, both forming In comparison the bitunicate marine
distinct clades in the Sordariomycetidae ascomycetes have been less well served at the
incertae sedis (Hibbett et al., 2007). These molecular level. Some 40 marine bitunicate
results have shown that the Halosphaeriales are ascomycetes have been sequenced, but
a sister group to the Microascales and are assignment to families remain unresolved. This
secondary invaders of the marine environment reflects the genera lack of resolution within the
(Spatafora et al., 1998). It has been confirmed bitunicate ascomycetes. Sequences of
that the Lulworthiales are morphologically and Manglicola guatemalensis places it in the
phylogenetically distinct from the Halo- Jahnulales.
sphaeriales (Kohlmeyer et al., 2000). Another
significant taxonomic result has been the TOTAL ASCOMYCETES.................... (424)
erection of a new order the Koralionastetales to

145
 Section C: 12. Conidiophores penicilliate or on a terminal ampulla
ANAMORPHIC FUNGI ............................................................................ 13
12. Conidiophores not penicilliate or on a terminal
(COELOMYCETES and ampulla................................................................ 15
HYPHOMYCETES)
13. Conidiogenous cells phialidic ............................. 14
1. Spores produced on hyphae ............... Hyphomycetes 13. Conidiogenous cells anellidic, 5-7 × 4-5 µm, on
1. Spores produced in sporocarps.............Coelomycetes algae .............................Scopulariopsis halophilica

HYPHOMYCETES 14. Phialides born on penicilliate conidiophores,

1. Conidia hyaline ........................................................ 2 conidia variously coloured ................... Penicillium
1. Conidia coloured .................................................... 11 14. Phialides on a terminal ampulla, conidia brown .....
................................................ Aspergillus sydowii
2. Conidia 1-celled ....................................................... 3 14. Phialides verrucose, at tip of undifferentiated
2. Conidia septate......................................................... 7 conidiophore, conidia verrucose, 5-7 × 3-4 μm
....................................... Stachybotrys mangiferae
3. Conidia produced in a sporodochium....................... 4
3. Conidia not produced in a sporodochium ................ 5 15. Conidia basauxic, 10.5-15.3 × 5.8-7.9 µm, on algae
............................................... Arthrinium algicola
4. Sporodochia superficial, on wood from deep waters, 15. Conidia acroblastosporic......................... Periconia
conidia 16-24 × 6.5-9 µm...Allescheriella bathygena
4. Sporodochia cushion-shaped, on salt marsh plants, 16. Conidia with transverse septa.............................. 17
conidia 3-4 × 1.5-2 µm....Tubercularia pulverulenta 16. Conidia muriform................................................ 24
16. Conidia helicoid .................................................. 27
5. Conidia tetraradiate, 7-9 × 6-7 µm, on driftwood ...... 16. Conidia tetraradiate or staurospores.................... 31
............................................... Heliscella stellatacula 16. Chlamydospores only produced .......................... 33
5. Conidia not tetraradiate or staurosporous................. 6
17. Conidia over 120 µm long, on submerged test
6. Conidiophores unbranched or sparingly, collarette panels ................................ Sporidesmium salinum
cylindrical, spore mass pale yellowish-brown, on 17. Conidia shorter than 120 µm............................... 18
diseased shrimp, conidia 3-4 × 1.5-2 µm
..................................... Plectosporium oratosquillae 18. Conidia with a prominent scar ............................ 19
6. Conidiophores simple or branched, on algae or soil 18. Conidia without a basal scar ............................... 20
..............................................................Acremonium
6. Conidiophores subglobose, lateral or apical, conidia 19. Conidiophores macronematous, conidia olive-
2-5 µm, on various substrata ...................................... brown, 8-24 × 4.5-8 µm, with protruberant scars at
....................................... Botryophialophora marina each end, conidia in short chains, on algae, drift
plant material..................... Cladosporium algarum
7. Conidia septate tetraradiate .................Varicosporina 19. Conidiophores macronematous, conidia never in
7. Conidia filamentous, multiseptate............................ 8 chains, slightly constricted at the septa, with short
7. Conidia elongate, obovoidal, ellipsoidal, elongate, basal peg-like denticle, on algae, occasionally wood
clavate, 1-5-septate, predacious on nematodes in .........................................................Dendryphiella
mangrove habitats (mangrove wood and leaves) ..... 9
20. Conidia 1-3-septate ............................................. 22
8. Conidia filiform, 9-13 (-19)-septate, end cells with 20. Conidia with more than 3-septate........................ 21
cytoplasm, 150-255 (-312) × 2.5-4 µm ......................
...............................................Anguillospora marina 21. Conidia narrower than 8 µm, 24-29 × 5.5-6.5 (-8)
8. Conidia sigmoid, curved, variously septate, end cells µm .............................................Amorosia littoralis
lacking cytoplasm ............................. Halosigmoidea 21. Conidia wider than 8 µm.................Trichocladium

9. Conidia apically and on lateral denticles................ 10 22. Conidia spathulate, versicolorous, apical cells
9. Apical conidia only ........................Monacrosporium brown, basal cell hyaline, 35-42 × 9-11 µm, on
mangrove wood......... Phragmospathula phoenicis
10. Trapping nematodes with adhesive networks.......... 22. Conidial cells in straight rows, concolorous,
............................................................ Arthrobotrys constricted at the septa ....................Trichocladium
10. Trapping nematodes with adhesive knobs or non- 22. Conidial pyriform................................................ 23
constricting rings................................. Dactylellina
23. Conidiophores in sporodochia, conidia obpyriform,
11. Conidia coloured, one-celled............................... 12 1-2-septate, not constricted at the septa, 20-33.6 ×
11. Conidia coloured, septate .................................... 16 14.5-20.5 µm......................Bactrodesmium linderi

146
 Fungal Diversity

23. Conidiophores not in sporodochia, conidia Mycelium septate, branched, phialides

pyriform, 21-27 × 10-17.5 µm ................................ erect, simple, solitary or in groups, conidia
.............................. Brachysporium helgolandicum
elliptical, obovoid, broadly-ellipsoidal, short,
24. Conidia flattened in one plane, up to 90 µm long truncate basal hilum, hyaline, forming a ball of
and 120 µm wide, dictyosporous............................. conidia in a mucilaginous matrix (Tubaki,
.............................Mycoenterolobium platysporum 1973b; Zuccaro et al., 2004). Acremonium fuci
24. Conidia not flattened in one plane....................... 25 was isolated from the brown algae Fucus
25. Conidiophores with percurrent proliferation, serratus (Germany) and F. distichus (Canada)
conidia variously shaped ................... Stemphylium with conidia measuring 5-8 (-15) × 3.2-5 (-6.0)
25. Conidia aleuriospores.......................................... 26 µm (Fig. 70). This species groups within a
marine clade of Emericellopsis and
26. Conidia entire, clavate, 15-41 × 12-37 µm.............. Stanjemonium species primarily isolated from
............................................... Monodictys pelagica
26. Conidia formed by rows of cells or branched..........
marine habitats, and Zuccaro et al. (2004)
......................................................... Dictyosporium suggest that further new species remain to be
described from this environment. Acremonium
27. Cells in conidia equal width along the coil, on polyaleurum was recovered from coastal muds
wood..........................................................Zalerion and characterized by its small conidia, the
27. Cells variable in width along the coil.................. 28
formation of aleuriospores and its better growth
28. Apical cell the largest in the coil ............ Cirrenalia on seawater media when compared with
28. Apical cells variable in width.............................. 29 growth on freshwater media. No teleomorphs
have been reported for the two marine species
29. Conidiophores polydenticulate, conidia 15-29 × 12- listed above. Zuccaro et al. (2004) list a
15 µm, on Nypa palm ...... Helicorhoidion nypicola
29. Conidiophores not denticulate............................. 30
number of Acremonium species isolated from
the sea: Acremonium potronii (skin lesion of a
30. Conidia 15-65 × 14-56 µm, cells less than 10 µm dolphin); A. tubakii (Fucus serratus);
wide, spirally contorted into a ball of up to 100 Emericellopsis minima (mangrove soil, Fucus
cells ....................................................Halenospora serratus) and E. stolkiae (mud saline lake).
30. Conidia initially a spiral/coil, cells over 10 µm
wide, forming a ball of cells by division in several
Acremonium species are frequently
planes ................................................Cumulospora encountered on incubated mangrove wood in
the topics and warrant further investigation.
31. Conidia with tetraradiate arms ............................ 32 See section on Emericellopsis, page 72.
31. Conidia unicellular but aggregated into groups up
to 20, 9-29 × 4-9 µm, on various substrata 1. Conidia 5-8 (-15) × 3.2-5 (-6) µm, obovoid-broadly
............................................ Asteromyces cruciatus ellipsoidal, on brown algae .......................... A. fuci
1. Conidia 3-6 × 1.5-2.0 µm, elliptical, from marine
32. Basal cell small, light brown, 6-20 µm, conidia 20- sediments................................................. A. tubakii
70 × 4-6 µm, on various substrata ...........................
..............................................Clavatospora bulbosa
32. Basal cell large, dark brown to black, conidia 24-42
× 18-37 µm, on wood ...........Orbimyces spectabilis

33. Spores hyaline, 35-150 × 8-33 µm, on Cladophora

spp. ....................................Blodgettia confervoides
33. Spores dark brown, on mangrove wood ..................
................................................................Xylomyces

Acremonium Link, Magazin Ges. naturf.

Freunde, Berlin, 3: 15, 1809 ........................ (2)
◙ A. fuci Summerb., Zuccaro & W. Gams,
Stud. Mycol. 50: 288, 2004.
◙ A. tubakii W. Gams, Cephalosporium-artige
Schimmelpilze (Stuttgart): 55, 1971. Fig. 70. Conidiophores and conidia of Acremonium fuci.
Cephalosporium polyaleurum Tubaki, Mycologia Bar = 5 µm.
65: 939, 1973.
147
Allescheriella Henn., Hedw. 36: 244, 1897. (1) A. marina Nakagiri & Tubaki, Mycologia 75:
A. bathygena Kohlm., Revue. Mycol. 41: 199, 488, 1983.
1977. Teleomorph: Lindra obtusa Nakagiri &
Sporodochial superficial, conidiophores Tubaki.
monoblastic, terminal, determinate, cylindrical, Mycelium septate, hyaline, conidiophore
straight, hyaline to pale brown conidia simple or branched, hyaline, conidiogenous
acrogenous, solitary, ellipsoidal to fusiform, cells hyaline, monoblastic, terminal, precurrent,
unicellular, thick-walled, smooth, and pale without a separating cell, conidia solitary,
brown (after Kohlmeyer, 1977). Only known filiform, straight or curved, 9-13 (-19)-septate,
from a collection made on wood from the swollen at both ends, hyaline, no sheath or
Tongue of the Ocean, off the Bahamas Islands appendages.
(Kohlmeyer, 1977).

Alternaria Nees, Syst. Pilze (Würzburg): 72,

1816........................................# Alternaria spp.
It is uncertain if these are truly marine
fungi, but numerous collections have been
listed in the literature. These include
Alternaria maritima on a wide range of
substrata (Sutherland, 1916; Johnson and
Sparrow, 1961), A. radicina (Anastasiou,
1963b) and A. tenuis (Siepmann and Johnson,
1960; Anastasiou, 1963b). Kohlmeyer and Fig. 71. Conidia of Amorosia littoralis (After Mantle et
Volkmann-Kohlmeyer (1991a) do not regard al., 2006). Bar = 5 µm.
any Alternaria species as obligately marine.
Arthrobotrys Corda, Pracht-Fl. Eur.
Amorosia Mantle & D. Hawksw., Mycol. Res. Schimmelbild.: 43, 1839 ............................ (12)
110: 1373, 2006 ........................................... (1) A. arthrobotryoides (Berl.) Lindau, Rabenh.
◙ A. littoralis Mantle & D. Hawksw., Mycol. Krypt. –Fl. (leipzg) 1(8): 371, 1905.
Res. 110: 1373, 2006. Cephalothecium roseum var. arthrobotryoides
Berl., Malpighia 2: 245, 1888.
Teleomorph: may be found in the Didymozoophaga arthrobotryoides (Berl.)
Sporormiaceae, Pleosporales (Mantle et al., Soprunov & Galliulina, Microbiol., Reading 20: 493,
2006). 1951.
A dematiaceous hyphomycete isolated A. brochopaga (Drechsler) S. Schenck, W.B.
from the littoral zone in the Bahamas and Kendr. & Pramer, Can. J. Bot. 55: 982, 1977.
producing the novel axa-anthraquinone Arthrobotrys gracilis (Dudd.) S. Schenck, W.B.
scorpinone and caffeine. Condiophores Kendr. & Pramer, Can. J. Bot. 55: 983, 1977.
Candelabrella brochopaga (Drechsler) Subram.,
micronematous or semi-macronematous, Kavaka 5: 95, 1978.
arising singly sympodially, monoblastic, Dactylaria gracilis Dudd., Trans Br. Mycol. Soc.
determinate, subhyaline to pale brown and 34: 194, 1951.
smooth walled. Conidia arising singly, 3-4- Dactylariopsis brochopaga (Drechsler) Mekht.,
septate, pale brown to brown, smooth-walled Mikol. Fitopatol. 1: 278, 1967.
Dactylariopsis gracilis (Dudd.) Mekht.,
lacking appendages or a sheath, 24-27.5 (-29) × Khishchnye Nematofagovye Griby-Gifomitsety (Baku)
5.5-6.5 (-8) µm (Fig. 71). It most closely 117, 1979.
resembles Trichocladium species: T. asperum, Dactylella brochopaga Drechsler, Mycologia 29:
T. lignicola, T. opacum and T. pyriforme but 517, 1937.
are not phylogenetically related (Mantle et al., Drechslerella brochopaga (Drechsler) M.
Scholler, Hagedorn & A. Rubner, Sydowia 51: 99, 1999.
2006).
A. cladodes var. macroides Dreschler,
Anguillospora Ingold, Trans. Br. Mycol. Soc. Mycologia 36: 144, 1944.
Trichothecium cladodes var macroides (Drechsler)
25: 401, 1942 ............................................... (1) Soprunov, Predacious fungi - Hyphomycetes and their

148
 Fungal Diversity

use in the control of pathogenic nematodes: 113, 1958. Didymozoophaga superba (Corda) Soprunov &
Arthrobotrys macroides (Drechsler) Mekht., Galiulina, Microbiol., Reading 20: 493, 1951.
Khishchnye Nematofagovye Griby–Gifomitsety (Baku) A. vermicola (R.C. Cooke & Satchuth.) Rifai,
89, 1979. Reinwardtia 7: 371, 1968.
A. dactyloides Drechsler, Mycologia 29: 486, Dactylaria vermicola R.C. Cooke & Satchuth.,
1937. Trans. Br,. Mycol. Soc. 49: 27, 1966.
Arthrobotrys anchonia Drechsler, Mycologia 46: Nematophagus vermicola (R.C. Cooke &
762, 1954. Satchuth.) Mekht., Khishchnye Nematofagovye Griby –
Dactylaria dactyloides (Drechsler) Soprunov, Gifomitsety (Baku): 105, 1979.
Predacious fungi–Hyphomycetes and their use in the Teleomorph: in the Orbiliaceae.
control of pathogenic nematodes: 142, 1958.
Dactylariopsis dactyloides (Drechsler) Mekht., Colonies white to cream to yellow, fast
Khishchnye Nematofagovye Griby–Gifomitsety (Baku): growing. Mycelium hyaline, septate and
120, 1979. branched. Conidiophores erect 100-900 µm
Drechslerella anchonia (Drechsler) M. Scholler,
Hagedorn & A.Rubner, Sydowia 51: 99, 1999.
long 1.5-5 µm wide, septate, rarely producing
Drechslerella dactyloides (Drechsler) M. Scholler, side branches, proliferating and bearing
Hagedorn & A. Rubner, Sydowia 51: 99, 1999. numerous conidia. Conidia are hyaline, 1-5-
Nematophagus anchonius (Drechsler) Mekht., septate, variously shaped, elongate, ellipsoidal,
Khishchnye Nematofagovye Griby–Giforitsety (Baku): pyriform, rarely constricted at the septa, up to
108, 1979. 60 µm long (Fig. 72). These fungi from
A. javanica (Rifai & R.C. Cooke) Jarow., Acta adhesive network devices for trapping
Mycologica, Warszawa 6: 373, 1970. nematodes. Johnson and Autery (1961) were
Candelabrella javanica Rifai & R.C. Cooke,
Trans. Br. Mycol. Soc. 49: 162, 1966.
the first to document a predacious fungus from
A. mangrovispora Swe, Jeewon, Pointing & brackish water habitats when they tentatively
K.D. Hyde, Bot. Mar. 51: 332, 2008. identified Arthrobotrys dactyloides growing on
A. musiformis Deschler, Mycologia 29: 481, pine panels (Pinus taeda) submerged for 37
1937. days in the saline zone of the Neuse River
Candelabrella musiformis (Drechsler) Rifai & estuary, North Carolina. Meyers et al. (1963)
R.C.Cooke, Trans. Br. Mycol. Soc. 49: 163, 1966. have documented the relationship between
Dactylella musiformis (Drechsler) Matsush., marine filamentous fungi and nematodes.
Microfungi of the Solomon Islands and Papua-New Generic delineation of predacious fungi varies
Guinea (Osaka) 22, 1971.
between different authors, but generally based
A. oligospora Fresen., Beitr. Mykol. 1: 18,
on the morphology of the nematode trapping
1850.
Arthrobotrys superba var. oligospora (Fresen.)
structures.
Coem.
Didymozoophaga oligospora (Fresen.) Soprunov 1. Conidia predominantly 1-septate ............................. 2
& Galiulina, Microbiol., Reading 20: 493, 1951. 1. Conidia with 2 or more-septate................................ 6
A. polycephala (Drechsler) Rifai, Reinwardtia
2. Conidia with central septum .................................... 3
7: 371, 1968.
2. Conidia with septum below the middle.................... 4
Dactylaria polycephala Drechsler, Mycologia 29:
530, 1937.
Woroninula polycephala (Drechsler) Mekht., 3. Conidiophores up to 850 µm, conidia elongate,
Khishchnye Nematofagovye Griby – Gifomitsety (Baku): obovoidal, ellipsoidal. 13-26 × 7-10 µm. 8-20 conidia
110, 1979. per conidiophore ..................................... A. cladodes
3. Condiophores 100-280 µm, conidia broad-ellipsoidal
A. pyriformis (Juniper) Schenk, W.B. Kendr. 16-22 × 7-9 µm, 5-14 conidia per conidiophore ........
& Pramer, Can. J. Bot. 55: 984, 1977. ................................................... A. arthrobotryoides
Dactylaria pyriformis Juniper, Trans. Br. Mycol. 3. Conidiophores 220-400 µm, conidia obovoid to
Soc. 37: 437, 1954. clavate, 20-37.5 × 7.5-10 µm ...................A. javanica
Dactylariopsis pyriformis (Juniper) Mekht.,
Khishchnye Nematofagovye Griby–Gifomitsety (Baku): 4. Condiophores up to 900 µm..................................... 5
119, 1979. 4. Conidiophore up to 700 µm, conidia elongate-clavate
A. superba Corda, Pracht-Fl. Eur. 24-36 × 6-8 µm ................................... A. dactyloides
Schimmelbid: 43, 1839.

149
4. Conidiophores up to 440 µm, conidia obovoid to This is a species widely reported as
pyriform, 17.5-28.8 × 10-16 µm, 10-15 conidia per causing widespread mortalities of sea fans
conidiophore ........................................ A. oligospora
(Gorgonia ventalina, G. flabellum) in the
5. Conidiophores up to 830 µm, conidia elongate, Bahamas, Caribbean, Costa Rica, Cuba,
obovoidal, ellipsoidal, slightly curved, 25-40 × 8.8- Mexico, USA and Venezuela (Nieves-Rivera,
12.5 µm…………..……………..…….A. musiformis 2002). Its identification as the causal pathogen
5. Conidiophores up to 900 µm, conidia elongate, was determined by a combination of
oboidal, ellipsoidal, 12-23 × 6-10 µm, 12 conidia per
conidiophore ............................................ A. superba morphology, virulence assays, histopathology
and molecular analysis of the 18S of rDNA
6. Conidia narrow, less than 16 µm ............................. 7 (Geiser et al., 1998). Other Aspergillus species
6. Conidia wider, 25-50 × 12-24 µm.A. mangrovispora have been frequently reported from sediments,
but treatment of these must await further
7. Conidiophores 90-270 µm, conidia ellipsoidal 12.5-5
× 3.5-4.5 µm, three dimentional networks .................
studies.
...........................................................A. polycephala
7. Conidiophores 78-580 µm, conidia cylindrical or
elongate-ellipsoidal 20-45 × 5-12.5 µm, trapping
with constricting rings........................ A. brochopaga
7. Conidiophores 150-500 µm, conidia 38-52 × 10-11
µm ........................................................ A. pyriformis

Fig. 73. Sea fan (Annella sp.) showing tissue loss due to
fungal infections (arrowed) (Photo by Sakanan
Plathong).

Fig. 72. Arthrobotrys mangrovispora. Conidiophores

and conidium. Bar = 10 μm. (after Sue et al., 2008).

Arthrinium Kunze, Mykologische Hefte

(Leipzig) 1: 9, 1817...................................... (1)
A. algicola (N.J. Artemczuk) N.J. Artemczuk
Papularia algicola N.J. Artemczuk, Mikol.
Fitopatol. 14: 95, 1980.
Described as a Papularia species it is
referred here to Arthrinium, and isolated from
the brown alga Cystoseira barbata. Colonies
terminal, hyaline, becoming dark, reverse
colony pale brown, conidiophores hyaline,
conidia lentiform, brown 10.5-15.3 × 5.8-7.9
µm and similar to Papularia sphaerosperma.

Aspergillus Link, Observ. Mycol. (Lipstae) 1:

16, 1809........................................................ (1)
A. sydowii (Bainier & Sartory) Thom &
Church, The Aspergilli 147, 1926. Fig. 74. White encrustation spreading over the axial
Sterigmatocystis sydowii Bainier & Sartory, skeleton of sea fan (Annella sp.) (Photo by Sakanan
Annls. Mycol. 11: 25, 1913. Plathong).

150
 Fungal Diversity

As a result of the 2004 tsunami, many Trichocladium species, especially T. nypae

gorgonian sea fans of the genus Annella at Mu which has smaller conidia 15-20 × 10-15 µm,
Ko Similan National Park, Andaman Sea, 2-septate, both occurring in marine habitats
Thailand were destroyed or became infected by (Goh and Hyde, 1999). No teleomorph reported
unknown microbes. These infections were for B. linderi, but other species show affinities
characterized by white encrustations spreading with Stuartella (Dothideales, incertae sedis).
rapidly over the axial skeleton causing tissue
loss and finally death of the colony (Figs. 73, Blodgettia Harv., Smithson. Contrib. bot. 10:
74). Phongpaichit et al. (2006) in a preliminary 42, 46, 1858.................................................. (1)
report noted an Aspergillus sp. and other fungal B. confervoides Harv., Smithson. Contrib. bot.
strains had infected the sea fan based on 10: 48, 1858.
morphological and molecular data. B. bornetii E.P. Wright, Trans. R. Ir. Acad. 28: 25,
1881.
Asteromyces Moreau & M. Moreau, Revue A confusing taxon, symbiotic
Mycol., Paris 6: 79, 1941 ............................. (1) (mycophycobioses) with the green algae
A. cruciatus Moreau & M. Moreau ex. Cladophora caespitosa and C. fuliginosa and
Hennebert, Can. J. Bot. 40: 1213, 1962 (Type collected from various localities (Kohlmeyer
species). and Kohlmeyer, 1979). Mycelium growing
Colony on agar, greenish-brown, within the inner and outer wall layer of the host,
becoming pale brown, mycelium septate, septate, branched, occurring as chlamydospores
branched, hyaline to brown, conidiogenous 35-150 × 8-33 µm, 1-7-septate terminal or
cells cylindrical, subglobose to subclavate, intercalary chains, constricted at the septa,
hyaline, from which arise up to 12-14 conidia, hyaline, yellow to yellow-brown (after
conidia ovoid to obpyriform, thin-walled, Kohlmeyer and Kohlmeyer, 1979).
brown, develop singly, first terminal, others
laterally, with a short stalk or denticles, Botryophialophora Linder, Farlowia 1: 403,
conidial mass released in a cluster. A common 1944.............................................................. (1)
species isolated from sand with unicellular B. marina Linder, Farlowia 1: 404, 1944 (Type
conidia forming cruciate arrangement of cells. species).
When grown on different media and under Hyphae septate, branched, hyaline to pale
different light regimes the conidia can become brown, conidiophores subglobose, lateral or
septate (Jones and Ward, 1973). apical, conidiogenous cells phialidic, flask-
shaped, hyaline, conidia globose, unicellular,
Bactrodesmium Cooke, Grevillea 12: 35, 1883. hyaline, no sheath or appendage. Reported
B. linderi ( J.L. Crane & Shearer) M.E. Palm & from a wide range of substrata, wood, test
E.L. Stewart, Mycotaxon 15: 319, 1982 ...... (1) panels, sand, sediments, but Kohlmeyer and
Trichocladium linderi J.L. Crane & Shearer., Kohlmeyer (1979) question its marine status.
Mycologia 70: 866, 1978.
Colonies sporodochial, hyaline becoming Brachysporium Sacc., Syll. fung. (Abellini) 4:
black, condiophores micronematous, smooth, 423, 1886 (emend Mason & Hughes) .......... (1)
thin-walled, hyaline or thick-walled brown, B. helgolandicum Schaumann, Helgoländer
conidiogenous cells holoblastic, integrated, wiss. Meeresunters. 25: 26, 1973.
terminal or intercalary, smooth, cylindrical, Mycelium superficial on substratum,
determinate, conidia solitary, subglobose to septate, pale brown, conidiophores simple,
obpyriform, 1-2-septate, not constricted at the erect, septate, pale brown, conidia broadly
septa, dark brown to black 20-33.6 × 14.5-20.5 pyriform, 21-27 × 10-17.5 μm, 2-3-septate, pale
µm. Moved to Bactrodesmium because of its brown, and attached to the conidiophores by
compact sporodochia, not a feature of long spirally coiled pedicels. No sheath or
Trichocladium which has mononematous and appendages. This is the only species known
scattered conidiophores. Extremely common from marine habitats and is most similar to the
on mangrove wood and often confused with genera Trichocladium and Cirrenalia.
151
Collected by Schaumann (1973a) on drift bark 3. Apical cells over 16 µm long ................................... 4
in the sea at Heligoland, Germany, however 3. Apical cells shorter than 16 µm ............................... 6
Kohlmeyer and Kohlmeyer (1979) do not list it
as a marine species. 4. Apical cell large, 16-23 µm, conidia 3-4-septate,
hooked appearance, black to fuscous ........C. pygmea
4. Apical cell not hooked ............................................. 5
Cirrenalia Meyers & R.T. Moore, Am. J. Bot.
47: 346, 1960 ............................................... (7) 5. Apical cell 11-22 µm, 2-4-septate, brown to dark
C. adarca Kohlm., Volkm.-Kohlm. & O.E. brown ............................................................C. fusca
Erikss., Bot. Mar. 40: 292, 1997. 5. Apical cell 16-20 µm, 3-6-septate, fuscous to grey-
brown ..................................C. pseudomacrocephala
Teleomorph: Juncigena adarca Kohlm.,
Volkm.-Kohlm. & O.E. Erikss. 6. Apical cell 9-16 µm, 3-5-septate, pale brown ............
C. basiminuta Raghuk. & Zainal, Mycotaxon ............................................................ C. basiminuta
31: 163, 1988. 6. Apical cell 6-14 (17) µm, 2-6-septate, reddish-brown
C. fusca I. Schmidt, Mycotaxon. 24: 419, 1985. ...................................................... C. macrocephala
C. macrocephala (Kohlm.) Meyers & R.T.
Moore, Am. J. Bot. 47: 347. 1960 (Type Cladosporium Link Mag. Ges. naturf. Feunde,
species). Berlin 7: 37, 1816 ........................................ (1)
Helicoma macrocephala Kohlm., Ber. dtsch. bot. C. algarum Cooke & Massee, Grevillea 16: 80,
Ges. 71: 99, 1958. 1888.
Teleomorph: in the Halosphaeriales. Hyphae septate, branched, pale brown,
C. pseudomacrocephala Kohlm., Mycologia conidiophores macronematous, cylindrical,
60: 266, 1968. septate, simple, straight, smooth, brown,
◙ C. pygmea Kohlm., Ber. dtsch. bot. Ges. 79: conidiogenous cells polyblastic, integrated,
35, 1966. terminal with prominent scars, conidia
Teleomorph: in the Lulworthiales. ellipsoidal, with a basal scar, 0-3-septate, not
◙ C. tropicalis Kohlm., Mycologia 60: 267, constricted at the septa, smooth, thick-walled,
1968. olive-brown, no sheath or appendages.
Teleomorph: in the Lulworthiales. Reported from drift brown seaweeds and sea
Conidiophores present or lacking, grasses. Cladosporium species are widely
cylindrical, septate or lacking septa, hyaline to reported from drift material but no
pale brown, conidiogenous cells monoblastic, investigation has been carried out on the
integrated, terminal, determinate, conidia marine taxa.
solitary, helicoid, 2-12-septate, generally
constricted at the septa, brown, the apical cell Clavatospora Sv. Nilsson ex Marvanová & Sv.
generally the largest. The morphology and Nilsson, Trans. Br. Mycol. Soc. 57: 531, 1971
dimensions of the apical cell determine the ...................................................................... (1)
species. A number of terrestrial species have ◙ C. bulbosa (Anastasiou) Nakagiri & Tubaki,
been described but in our opinion these are not Bot. Mar. 28: 489, 1985 (Type species).
well placed in the genus (Raghukumar et al., Clavariopsis bulbosa Anastasiou, Mycologia 53:
11, 1962.
1988). The genus is not monophyletic (See Fig.
62) with teleomorphs in different ascomycete Teleomorph: Corollospora pulchella
genera while new anamorphic genera need to Kohlm., I. Schmidt & N.B. Nair.
be described (Jones, personal observation). Conidia dark brown, composed of a basal
Reported from a wide range of substrata in the bulbous cell and 1-3 radiating arms, 3-6-
sea. septate, hyaline to pale brown, cells constricted
at the septa, in culture the bulbous basal cell
1. Conidia 6-septate or more ........................................ 2 may be absent and conidia develop as a single
1. Conidia 5-septate or less .......................................... 3 row of brown cells. Kohlmeyer and Kohlmeyer
(1979) refer to these as chlamydospores. A
2. Apical cell 6-12 µm, 4-9-celled, hyaline to pale common species on various substrata,
brown, on Juncus ....................................... C. adarca
particularly wood, when incubated with a layer
2. Apical cell 10-20 µm, 6-12-septate, reddish-brown,
on wood.................................................. C. tropicalis of water over the surface.

152
 Fungal Diversity

Cumulospora I. Schmidt, Mycotaxon 24: 420, Dactylellina M. Morelet, Bull. Soc. Sci.
1985.............................................................. (2) naturelles Archeologie, Toulonet et du Var 178:
C. marina I. Schmidt, Mycotaxon 24: 421, 6, 1968.......................................................... (2)
1985 (Type species). D. huisuniana (J.L. Chen, T.L. Huang &
Vesicularia marina I. Schmidt, Natur Naturschutz Tzean) M. Scholler, Hagedorn & A. Rubner,
Mecklenberg 12: 117, 1974. Sydowia 51: 111, 1999.
Basramyces marinus (I. Schmidt) Abdullah, Dactylella huisuniana J.L. Chen, T.L. Huang &
Abdulk. & Goos, Intern. J. Mycol. and Lichenol. 4: 183, Tzean, Mycol. Res. 102: 1269, 1998.
1989.
D. lysipaga (Drechsler) Scholler, Hagedorn &
◙ C. varia Chatmala & Somrith., Fungal Diver.
A. Rubner, Sydowia 51: 111, 1999.
17: 3, 2004. Dactylella lysipaga Drechsler, Mycologia 29:503,
Teleomorph: in the Lulworthiales. 1937
Mycelium septate, branched, superficial Teleomorph: in Orbilia.
or immersed, pale brown, conidiophores absent,
conidiogenous cells holoblastic, integrated, Colonies light yellow, mycelium
terminal, determinate, conidia initially spiral, branched, septate. Conidiophores up to 250 µm
cell division in several planes leads to a tangled erect. First conidium apical, lateral conidia on
knot of cells, numbering up to 40 or more, dark branches, condia spindle-shaped, 3-4-septate,
brown to black and constricted at the septa (Fig. 20-49 × 6-10 µm and hyaline. Trapping device,
75). Cumulospora varia is well placed in the adhesive stalked knobs.
Lulworthiales (100% bootstrap value and 1.00
posterior probability). The two isolates of 1. Conidiophores up to 250 µm, conidia spindle shaped,
Cumulospora varia, isolated from Mu Ko 3-4 septate, 20-49 × 6-10 µm...................D. lysipaga
Chang Island, Thailand, from different 1. Conidiophores 86-264 µm, conidia fuisiform, 3-
septate, 25-52.5 × 4-6 µm ...................D. huisuniana
collections, are monophyletic, and form a sister
group to Lulwoana uniseptata and its
anamorph Zalerion maritima (Jones et al., Dendryphiella Bubák & Ranoj., Ann. Mycol.
2008). 12: 417, 1914 ............................................... (2)
◙ D. arenaria Nicot, Revue Mycol., Paris 23:
1. Conidia 52-91 × 40-71 µm, up to 20 cells, cells up to 93, 1958.
30 µm ....................................................... C. marina Scolecobasidium arenarium (Nicot) M.B. Ellis,
1. Conidia 24-87 × 21-51 µm, up to 40 cells, cells up to More Dematiaceous Hyphomycetes (Kew): 194, 1976.
20 µm ............................................................ C. varia
◙ D. salina (G.K. Sutherl.) G.J.F. Pugh &
Nicot, Trans. Br. Mycol. Soc. 47, 266, 1964.
Cercospora salina G.K. Sutherl., New Phythol.
15: 43, 1916.
Teleomorph: in the Pleosporales,
Pleosporaceae.
Hyphae septate, branched, pale brown,
conidiophores macronematous, cylindrical, 1-
3-septate, simple, apically swollen, pale brown
to olive-brown, scars visible when conidia shed,
conidia ellipsoidal, cylindrical, 1-11-septate,
constricted at the septa, pale brown to
olivaceous, solitary, with short stalks but not
denticulate. Molecular sequence places these
species in the Pleosporales (Jones et al., 2008).
However the taxonomic position of the species
needs needs resolution. Ellis (1976) referred
the species to Scolecobasidium, however in the
Fig. 75. Cumulospora varia. Conidium. Bar =10 µm.
marine Dendryphiella species conidiogenous
cells are enteroblastic and denticles are absent.
153
Ellis (1976) described pegs on the 1. Conidia with 5 rows of cells, (44-) 50-80 × 24-31(-37)
conidiogenous cells but these may be confused µm, golden to reddish-brown ...................D. elegans
1. Conidia irregular in shape and number of cells,
with extensions from the conidia as seen in appears muriform, 12-66 × 9-28 (-36) µm, dark
SEM micrographs (Fig. 76). A new genus may brown to black ..................................... D. pelagicum
be warranted for the marine species.
Fusarium sp.
1. Conidia 1-3-septate, 9-20 × 4-6 µm ........ D. arenaria A number of Fusarium species have also
1. Conidia 2-9-septate, 14-75 × 6-10 µm ........ D. salina been isolated from sediments and sand dunes
Dictyosporium Corda, Beitr. Gesammten including Fusarium oxysporum (Nicot, 1958)
Natur-Heilwiss. 1: 87, 1836 ......................... (2) and F. solani growing on recovered submerged
D. pelagicum (Linder) G.C. Hughes ex T.W. twigs of Tamarix aphylla (Anastasiou, 1963b),
Johnson & Sparrow, In: Fungi in Oceans and but none of these have been considered to be
Estuaries 391, 1961. marine.
Speira pelagica Linder, Farlowia 1: 407, 1944.
Speira littoralis Höhnk, Veröeff. Inst.
Halenospora E.B.G. Jones gen. nov............ (1)
Meeresforsch. Bremerhaven 3: 221, 1955. Mycobank 513077
Holotytypus: Helanospora varia (Anastasiou)
E.B.G. Jones [≡ Zalerion varium Anastasiou]
Etymology: From halen = welsh for salt in
reference to its marine habitat and spora = spore.
Fungus leotiacearum mitosporarum. Mycelio
hyalino, septato, ramoso; conidiophoris hyalinis,
simplicibus, interdum nullis; conidiis fuscis vel nigris,
conidiarum celluis catenatis tandem tortuosis denique
dictyosporiformibus terminalibus. Pneumocandin nullis.
Anamorphic Ascomycota (Leotiaceae).
Conidiophores simple or absent, conidia dark brown to
black, cells formed in a chain, becoming tortous and
appearing as terminal dictyospores. Pneumocandin
negative. Phylogenetically groups with Glarea lozyensis
in the Leotiaceae.
◙ H. varia (Anastasiou) E.B.G. Jones, comb.
nov. Fig. 77, Mycobank 513086.
Fig. 76. Scanning electron micrograph of Dendryphiella Zalerion varium Anastasiou, Can. J. Bot. 41:
salina, conidiogenous cells and conidia. Peg-like 1136, 1963.
structures on the conidia (white arrows), black arrow Teleomorph: In the Leotiaceae,
indicates pore at the tip of the conidiogenous cell. Bar =
10 μm. No denticles found on the conidiogenous cells.
Leotatiales.
Zalerion varia (as Z. varium) is not
D. elegans Corda, Weitenweber’s Beitrage zur congeneric with Z. maritima (as Z. maritimum)
Nat.: 87, 1837. (Bills et al., 1999) the type species. Based on
Conidiophores micronematous, ITS sequences, H. varia is positioned in the
mononematous, hyaline to light brown, Leotiaceae in a clade comprising Galrea
conidiogenous cells integrated, determinate, lozoyensis, Hymenocyphus monotropae and
conidia solitary, holoblastic, branched, cheiroid, Pezicula carpinea (Bills et al., 1999), while Z.
dark brown to black. Both species occur on maritima is a member of the Lulworthiales
submerged wood, D. pelagicum a frequently (Campbell et al., 2005; Jones et al., 2008).
occurring species, while D. elegans has been Worldwide in its distribution occurring
reported on Rhizophora stylosa wood in the on intertidal wood, submerge leaves, seedling
Shiira River, Japan (Nakagiri, 1993b). Sutton of Rhizophora mangle. No teleomorph in the
(1985) considered D. pelagicum better placed Leotiaceae has been identified for the species.
in Monodictys, a view shared by Goh et al. Although the conidia of both
(1999), however, neither proposed formal Halenospora and Zalerion initially are simple
designation. Monodictys species rarely have coil, they differ in that in H. varia they
branched conidia, thus we keep D. pelagicum “produce a lateral rather than a terminal spiral
in Dictyosporium. as in Z. maritima” (Anastasiou, 1963a). The

154
 Fungal Diversity

individual cells in H. varia are narrower than conidia devoid of contents when mature (Fig.
those of Z. maritima, and form knot-like 78).
structures (Fig. 77) but they are not Halosigmoidea differs from Sigmoidea in
phylogenetically related. A number of fungi the following respects: (1) Conidia markedly
with similar morphology have been observed in coiled, rarely sigmoid. (2) Conidial cells
tropical locations and await further slightly swollen. (3) Conidial ends cells lacking
identification and description (Jones, pers. contents. (4) Phylogenetically placed in the
comm.). Halosphaeriales and (5) Marine species found
predominantly on decaying seaweeds,
especially members of the Fucales (Haythorn et
al., 1980; Zuccaro and Mitchell, 2005). Mature
conidia are generally not constricted at the
septa, but before germination each conidial cell
becomes rounded and separate into individual
cells or several cell clusters, from which
hyphae germinate.
1. Conidia less than 100 µm, 4-5 µm wide near the
middle, with indentation at some septa .... H. parvula
1. Conidia longer than 100 µm .................................... 2

Fig. 77. Halenospora varia. Highly convoluted, knot- 2. Conidia 110-180 × 4.5-10 µm near the middle,
like conidia. Bars = 10 µm. colony pale................................................ H. marina
2. Conidia 108-222 × 4.5-7.5 µm near the middle,
Halosigmoidea Nakagiri, K.L. Pang & E.B.G. colony bright yellow ..................................H. luteola
Jones, Bot. Mar. (in press) ........................... (3) Helicorhoidion S. Hughes, Can. J. Bot. 36: 773,
H. luteola (Nakagiri & Tubaki) Nakagiri K.L. 1958.............................................................. (1)
Pang & E.B.G. Jones Bot. Mar. (in press) H. nypicola K.D. Hyde & Goh, Mycol. Res.
(Type species). 103: 1420, 1999.
Sigmoidea luteola Nakagiri & Tubaki., Trans. Mycol.
Soc. Japan 23: 102. 1982.
Colonies on wood black and glistening,
Teleomorph: Corollospora luteola conidiophores macronematous, unbranched,
Nakagiri & Tubaki.
H. marina (Haythorn & E.B.G. Jones).
Nakagiri, K.L. Pang & E.B.G. Jones, Bot. Mar.
(in press).
Sigmoidea marina Haythorn & E.B.G. Jones.
Trans. Brit. Mycol. Soc. 74: 620. 1980.
Teleomorph: In Corollospora close to C.
luteola.
H. parvula Zuccaro, J.I. Mitch. & Nakagiri,
Bot. Mar. (in press).
Teleomorph: In Corollospora.
Mycelial hyphae branched, septate,
hyaline. Conidiophore hyaline pleurogenous on
the mycelium initially short and simple then
becoming longer and septate. Conidiogenous
cells holoblastic, terminal, sympodial or Fig. 78. Halosigmoidea parvula. End cells (arrowed)
devoid of cytoplasm. Conidia 4-5-septate. Bar = 25 μm.
irregularly sympodial and denticulate with
(Photo by Akira Nakagiri).
schizolytic secession. Conidia aleuriospores, C
to U-shaped, rarely sigmoid, solitary, septate, pale brown, smooth, 1-4-septate, condiogenous
hyaline, terminal and basal cells of mature cells indeterminate, terminal, polydenticulate,

155
conidia ellipsoidal or globose, irregularly Teleomorph: in the Orbiliaceae,
helicoid, very tightly coiled, constricted at the Orbiliales.
septa, 15-20 × 12-15 µm, holoblastic produced
1. Conidia 30-60 × 13-24 μm, broad at the apex,
on denticles by schizolysis, multiseptate, chlamydospores present ................... M. thaumasium
smooth and brown, occurring on Nypa 1. Conidia wider, end cells pointed, no chlamydospores
fruticans (Hyde et al., 1999b). May be .........................................................M. eudermatum
confused with Zalerion varia, but differs in the
production of denticles on the conidiophores Monodictys S. Hughes, Can. J. Bot. 36: 785,
which are 1-4-septate, and long (30-70 × 2.5-4 1958.............................................................. (1)
μm). ◙ M. pelagica (T.W. Johnson) E.B.G. Jones,
Trans. Br. Mycol. Soc. 46: 138, 1963.
Piricauda pelagica T.W. Johnson, J. Elisha
Heliscella Marvanová, Trans. Br. Mycol. Soc. Mitchell sci. Soc. 74: 42, 1958.
75: 224, 1980 ............................................... (1) Piricauda arcticoceanorum R.T. Moore, Rhodora
61: 95, 1959.
H. stellatacula (P.W. Kirk ex Marvanová & Sv.
Nilsson) Marvanová, Trans. Br. Mycol. Soc 75: Teleomorph: Nereiospora cristata
224, 1980. (Kohlm.) E.B.G. Jones, R.G. Johnson & S.T.
Clavatospora stellatacula P.W. Kirk, Mycologia Moss.
61: 178, 1969 A cosmopolitan species occurring on a
Hyphae septate, branched, hyaline, wide range of substrata, largely with a
conidiophores phialidic, lageniform, simple, temperate distribution. Conidiogenous cells
hyaline, conidia hyaline, 7-9 × 6-7 µm, monoblastic, integrated, terminal, determinate,
enteroblastic-phialidic, stellate, unicellular, cylindrical, doliiform or subspherical; conidia
comprising a main axis and 3-4 subconical solitary, dry, acrogenous, simple, oblong
processes. Occurring on wood (pine test-blocks) rounded ends, pyriform, clavate, ellipsoidal,
and rarely collected since described by Kirk subspherical or irregular, muriform, brown to
(1969). black, smooth or verrucose, basal cell
sometimes inflated, paler and thinner walled
than the other cells. Free sporulating in culture,
Monacrosporium Oudem., Ned. kruidk. Archf,
conidial measurements vary between different
2 sér. 4: 250, 1885........................................ (2)
collections. Teleomorph connection established
M. eudermatum (Drechsler) Subram., J. Indian
by culture techniques (Mouzouras and Jones,
bot. Soc. 42: 293, 1964.
Arthrobotrys eudermata (Drechsler) M. Scholler,
1985).
Hagedorn & A. Rubner, Sydowia 51: 102, 1999.
Dactylaria eudermata Drechsler, Mycologia 42:
Mycoenterolobium Goos, Mycologia, 62: 171,
40, 1950. 1970.............................................................. (1)
Dactylella eudermata (Drechsler) Seifert & W.B M. platysporum Goos, Mycologia 62: 172,
Kendr., Univ. Waterloo Biol. Ser. 27: 30, 1983. 1970.
Genicularia eudermata (Drechsler) Rifai, Conidiophores micronematous, monone-
Reinwardtia 7: 367, 1968.
Geniculifera eudermata (Drechsler) Rifai,
matous, short, hyaline, conidiogenous cells
Mycotaxon 2: 216, 1975. integrated, determinate, conidia solitary,
Golovinia eudermata (Drechsler) Mekht., holoblastic, dictyosporus, flattened in one
Khishchnye Nematofagovye Griby – Giforitsety (Baku): plane, variable shape, fan shaped, and dark
153, 1979. brown (Nakagiri, 1993b). Numerous
M. thaumasium (Drechsler) de Hoog & collections from submerged Rhizophora stylosa
Oorschot, Stud. Mycol. 26: 120, 1985. wood collected in the Shiira River, Japan.
Arthrobotrys thaumasia (Drechsler) S. Schenck,
W.B. Kendr. & Pramer, Can. J. Bot. 55: 984, 1977.
Orbimyces Linder, Farlowia 1: 404, 1944 ... (1)
Candelabrella thaumasia (Drechsler) Rifai,
Reinwardtia 7: 369, 1968. ◙ O. spectabilis Linder, Farlowia 1: 404, 1944
Dactylaria thaumasia Drechsler Mycologia 29: (Type species).
522, 1937. Teleomorph: In the Lulworthiales.
Golovinia thaumasia (Drechsler) Mekht., Mikol. Conidiophores short arising from the
Fitopatol. 1: 276, 1967.
mycelium, hyaline to pale brown, conidia with

156
 Fungal Diversity

a large black basal cell, subglobose to ovoid, 1. Colonies yellowish-green, becoming deep red,
thick-walled, glistening, with 1-2 branches conidia oval, 2.4-2 × 1.8-2 µm.. P. dimorphosporum
1. Colonies pale, with no pigment production, irregular
giving rise to a crown of radiating appendages, biverticillate, conidia globose to subglobose, 2.8-3.3
generally one apical and 4 latter arms, each × 2.5-3 µm ...............................................P. limosum
one 2-4 septa, slightly constricted at the septa,
and pale brown. Sporulates in culture with a
putative teleomorph in the Lulworthiales
(Jones et al., 2008). Orbimyces spectabilis, an
infrequently collected anamorphic fungus
isolated from intertidal wood in Denmark, was
basal to the Lulwoidea clade, but with weak
support (Fig. 62). Reported from driftwood,
test panels, and geographically widely
distributed, but uncommon.

Penicillium Link, Magazin Ges. Naturf.

Freunde, Berlin 3: 16, 1809 ......................... (3)
P. dimorphosporum H.J. Swart, Trans. Br.
Fig. 79. Penicillium dravuni. Conidiophore, phialides
Mycol. Soc. 55: 310, 1970. and conidia. Bar = 5 µm.
◙ P. dravuni Janso, Mycologia 97: 445, 2005.
P. limosum S. Udea, Mycoscience 36: 451, Periconia Tode, Fung. mecklenb. sel.
1995. (Lüneburg) 2: 2-3, 1791............................... (2)
Teleomorph: Eupenicillium limosum S. P. abyssa Kohlm., Rev. Mycol. 41: 202, 1977.
Udea. ◙ P. prolifica Anastasiou, Nova Hedw. 6: 260,
Penicillium limosum and P. dimor- 1963.
phosporum were described from marine Teleomorph: Okeanomyces cucullatus
sediments and P. dravuni collected on a (Kohlm.) K.L. Pang & E.B.G. Jones.
submerged alga Dictyosphaeria versluysii by Hyphae pale brown, septate, branching,
scuba diving off the coast of Fiji (Janso et al., conidiophores monoblastic, ellipsoidal,
2005). Penicillium dimorphosporum was cylindrical to clavate, septate, hyaline to pale
isolated from a mangrove swamp in Australia brown, conidia unicellular, subglobose or
and has yellowish-green colonies, eventually ovoid, smooth, thick-walled, pale brown to
turning deep reddish brown, conidiophores dark brown, catenulate, conidia dispersing, no
short, smooth, unbranched, phialides variable, sheath or appendages. Periconia prolifica is
conidia oval, smooth, later with prominent very common on tropical wood (Vrijmoed et
tubercles with no known teleomorph (Swart, al., 1984) occurring on a wide range of
1970). Penicillium dravuni is monoverticillate, substrata, sporulating readily in culture.
conidia spherical to subglobose, smooth-walled Periconia abyssa is known from recovered
to finely roughened, colonies yellow-grey and wood at 3975-5315 m in the Gulf of Angola
belongs in the P. thomii series, subseries P. and Iberian deep sea (Kohlmeyer, 1977).
turbatum and most closely resembles P.
turbatum (Fig. 79). Demonstrated to produce 1. Conidia 6-13 (-20) µm diameter, light to dark brown,
the secondary metabolites dictyosphaeric acids littoral species .........................................P. prolifica
1. Conidia 16-20 µm diameter, brown, deep sea species
A and B and carviolin (Janso et al., 2005). No ....................................................................P. abyssa
known teleomorphs for P. dravuni and P.
dimorphosporum. Phragmospathula Subram. & N.G. Nair,
1. Colonies light brown to white, reverse amber to
Antonie van Leeuwenhoek 32: 384, 1966 ... (1)
yellow at the margins, monoverticilliate, conidia P. phoenicis Subram. & N.G. Nair, Antonie
spherical, 2.5-3 µm long, 2 µm wide ........P. dravuni van Leeuwenhoek 32: 384, 1966.

157
 Conidiophores macronematous, mono- Mycelium septate, branched, hyaline,
ematous, arising from hyphae on the wood, annellophores borne on hyphae, hyaline, 3-18
conidiogenous cells integrated, percurrent, µm, base cylindrical or slightly constricted,
conidia solitary, holoblastic, spathulate, 3- conidia thallic, subglobose to ovate, truncate
septate, apical cell hyaline, middle cells brown, base, rounded apex, conidial mass white,
basal cell hyaline, no sheath or appendage. catenulate in a short chain (Tubaki, 1973a).
Collected on intertidal wood of Rhizophora Isolated from the seaweed Undaria pinnatifida
stylosa in the Shiira River, Shi-Ya-O, Japan (salted) and requires seawater for growth.
(Nakagiri, 1993b). This species may be more
common than reported in the literature (Hyde, Stachybotrys Corda, Icon. fung. (Abellini) 1:
1988). 21, 1837........................................................ (2)
S. atra Corda, Icon. Fung. (Prague) 1: 21, 1837.
Plectosporium M.E. Palm, W. Gams & S. mangiferae P.C. Misra & S.K. Srivast.,
Nirenberg, Mycologia, 87: 398, 1995 .......... (1) Trans. Br. Mycol. Soc. 78: 556, 1982.
P. oratosquillae Duc, Yaguchi & Udagawa, Conidiophores, macronematous,
Mycopth. Mycol. Appl. (In press) branched, septate, hyaline to pale brown,
Conidiophores superficial, solitary, verrucose, conidiogenous cells phialidic, in
unbranched to sparingly branched, smooth groups of 6-8 at the tip of the conidiophore,
walled; conidiogenous cells phialidic, hyaline, clavate, pale brown, conidia, ovoid to ellipsoid,
cylindrical or subulate; conidia unicellular, verrucose, brown. Stachybotrys atra was
hyaline, pale yellowish brown in mass, reported from submerged twigs of Tamarix
ellipsoidal, cylindrical to obovoid, sometimes aphylla in the Salton Sea (Anastasious, 1963b)
curved, 3-10 × 2-4 µm, smooth-walled, while S. mangiferae was reported on
guttulate, in slimy heads at the tips of the submerged wood of Rhizophora stylosa from
phialides (Fig. 80). The fungus is known from the Shiira River, Japan by Nakagiri (1993b).
diseased mantis shrimp (Oratosquilla oratoria) Stachybotrys has been reported by others from
collected in Yamaguchi Pref., Japan (Duc et al., marine habitats (e.g. Meyers and Reynolds,
in press). 1959) and is included here so that its
occurrence in the sea can be documented and
its ecological role determined.

1. Conidia 6.1-10.4 × 3.3-3.6 μm........................ S. atra

1. Conidia 5-7 × 3-4 μm.......................... S. mangiferae

Sporidesmium Link, Mag. Ges. naturf.

Freunde, Berlin 3: 41, 1809 ......................... (1)
S. salinum E.B.G. Jones, Trans Br. Mycol. Soc.
46: 135, 1963.
Mycelium septate, branched, brown to
dark brown. This species has only been
collected on submerged test blocks at Port Erin
Fig. 80. Plectosporium oratosquillae. Conidiophore and Marine Station, Isle of Man, when nine
conidium. Bar = 5 µm. collections were made on beech and Scots pine
submerged test panels (Jones, 1963). Does not
Scopulariopsis Bainier, Bull. Soc. mycol. Fr. correspond to typical Sporidesmium species
23: 98, 1907 ................................................. (1) and further collections are required to resolve
S. halophilica Tubaki, Trans. Mycol. Soc. Jpn. its taxonomic position. Typically has a large
14: 367, 1973. apical conidial cell circa 75 µm long.
Basipetospora halophila (J.F.H. Beyma) Pitt &
A.D. Hocking, Mycotaxon 22: 198, 1985.
Oospora halophila J.F.H. Beyma, Zentbl. Bakt.
Stemphylium Wallr., Flora crypt. Germ,
ParasitKde, Abt. II 88: 134, 1933. (Nürnberg) 2: 300, 1833 .............................. (3)

158
 Fungal Diversity

S. gracilariae E.G. Simmons, Mem. N.Y. bot. Teleomorph: Gaeumannomyces medul-

Gdn. 49: 305, 1989. laris Kohlm., Volkm-Kohlm. & O.E. Erikss.
S. maritimum T.W. Johnson, Mycologia 48: T. melhae E.B.G. Jones, Abdel-Wahab &
844, 1957. Vrijmoed, Fungal Diver. 7: 50, 2001.
S. triglochinicola B. Sutton & Piroz., Trans Br. ◙ T. nypae K.D. Hyde & Goh, Mycol. Res. 103:
Mycol. Soc. 46: 519, 1963. 1420, 1999.
Conidiophores macronematous, mono- Mycelium septate, branching, pale brown
nematous, septate, smooth or verrucose, pale to brown, conidiophores poorly differentiated,
brown to brown or olivaceous, solitary or in short pedicels, 0-3-septate, simple, straight,
groups, conidiogenous cells monoblastic, smooth, conidia produced at the apex, hyaline
integrated, terminal, percurrent, clavate to or pale brown, conidiogenous cells monoblastic
subglobose, thin-walled, conidia oblong, or polyblastic, integrated, terminal, solitary,
ellipsoidal, obclavate or subglobose, muriform, clavate, obovoid, pyriform or cylindrical, 1-4-
constricted at the septa or not, smooth, septate, thick-walled, smooth or verrucose, pale
verrucose or echinulate, scar at the base, pale to dark brown to black (Jones et al., 2001) (Fig.
brown, brown or olivaceous, solitary 81). A number of marine species have been
acrogenous, no sheath or appendages. Marine assigned to this genus, but they are not
species collected on various substrata: algae, monophyletic, as the teleomorph connections
salt marsh plants and on submerged wood referred to above indicates.
panels in the sea (Johnson, 1957). It is doubtful Goh and Hyde (1999) referred Tricho-
if any of these belong in Stemphylium. cladium linderi to Bactrodesmium linderi.
Trichocladium nypae differs only slightly from
1. Conidia trigonous, conidia 40-82 × 18-46 µm, up to
12 trans-septate, 1-4-longi-septate, on the salt marsh T. linderi. de Bertoldi et al. (1972) and Lepidi
plant Triglochin..............................S. triglochinicola et al. (1977) pointed out that Humicola
1. Conidia ellipsoidal, on other substrata ..................... 2 alopallonella was incorrectly referred to
Humicola because it lacked one-celled conidia,
2. Conidia 22-31 × 13-20 µm, 3-4-trans-septate, 1-2- thus Kohlmeyer and Volkmann-Kohlmeyer
longi-septa, on red seaweeds................ S. gracilariae
2. Condia 19-32 × 17-29 µm, 3-4-trans-septa, 2-4- (1995) transferred it to Trichocladium.
longi-septa, on submerged wood panels ....................
.............................................................S. maritimum 1. Conidia 1-2 (3)-septate ........................................... 2
1. Conidia with more than 3-septate ............................ 3
Trichocladium Harz., Bull. Soc. Imp. nat. 2. Conidia 10-22 (-38) × 8-18 µm, apical cell 8.5-15.5 ×
Moscou 44: 125, 1871 ................................. (7) 7-12 µm fuscous............................ T. alopallonellum
T. achrasporum (Meyers & R.T. Moore) 2. Conidia 6.5-14 × 3.5-9 µm, apical cell 6.5-13 × 4-9
Dixon, Trans Br. Mycol. Soc. 51: 163, 1968. µm, dark brown..........................................T. melhae
Culcitalna achraspora Meyers & R.T. Moore,
Am. J. Bot. 47: 349, 1960. 3. Conidia with 2-3-septa, 17-26 × 7-10 µm, distal cell
Teleomorph: Halosphaeriopsis medi- fuscous, elongate ellipsoid, on Juncus .......................
.............................................................. T. medullare
osetigera (Cribb & J.W. Cribb) T.W. Johnson. 3. Conidia with 2-3-septate, 15-20 × 10-13 (-15),
T. alopallonellum (Meyers & R.T. Moore) curved, dark-brown on Nypa........................ T. nypae
Kohlm. & Volkm.-Kohlm., Mycotaxon 53: 352, 3. Conidia with more than 3-septate ............................ 4
1995.
Humicola alopallonella Meyers & R.T. Moore, 4. Conidia with 2-5 (-6)-septate, 25-32 × 12-17 µm,
Am. J. Bot. 47: 346, 1960. light brown, slightly constricted.............. T. lignicola
◙ T. constrictum I. Schmidt, Nat. Naturschutz 4. Conidia with 2-4-septate .......................................... 5
Mecklenberg 12: 114, 1974. 5. Conidia 25-47 × 8-20 µm, reddish-brown,
T. lignicola I. Schmidt, Mycotaxon 24: 420, subglobose, markedly constricted at the septa, 2-4-
1985. septate ................................................ T. constrictum
T. medullare Kohlm. & Volkm.-Kohlm., 5. Conidia (15-) 20-34 (45) × (8-) 10-24 µm, often
Mycotaxon 53: 349, 1995. sporodochial, dark brown, compressed, less
constricted at the septa, 2-4-septate .T. achrasporum

159
Fig. 81. Trichocladium melhae. Conidium. Bar = 10 µm.
Fig. 82. Conidia of Varicosporina anglusa. Bars = 15
Tubercularia Tode, Fung. mecklenb. sel. µm. (Photo by Mohamed Abdel-Wahab).
(Lüneburg): 1: 18, 1790 ............................... (1)
T. pulverulenta Speg., An Soc. Cient. Argent. Xylomyces Goos, R.D. Brooks & Lamore,
13: 32, 1882 Mycologia 69: 282, 1977 ............................. (1)
X. rhizophorae Kohlm. & Volkm.-Kohlm.,
Sporodochia cushion-shaped, sessile or Fungal Diversity. 1: 160, 1998.
short stalk, erumpent, conidiophores, filiform, Xylomyces produces only chlamydo-
simple, conidia produced at the apex, hyaline, spores of variable form, with 5 species
conidia unicellular, ellipsoidal, hyaline to pink described from freshwater and one on
in a mass, smooth-walled (after Kohlmeyer and mangrove wood (Goos et al., 1977; Goh et al.,
Kohlmeyer, 1979). Saprophytic on various 1997; Kohlmeyer and Volkmann-Kohlmeyer,
Salicornia species and poorly known. 1998d). Chlamydospores in Xylomyces
rhizophorae mostly apical, single or in chains,
Varicosporina Meyers & Kohlm., Can. J. Bot. rarely branching, filamentous, straight or
43: 916, 1965 ............................................... (3) curved, mostly widest at the tips, tapering
V. prolifera Nakagiri, Trans Mycol. Soc. Jpn. towards the base, dark brown, 11-34-septate
27: 198, 1986. and constricted at the septa (Kohlmeyer and
Teleomorph: In the genus Corollospora Volkmann-Kohlmeyer, 1998d). They also
◙ V. ramulosa Meyers & Kohlm., Can. J. Bot. reported and illustrated Xylomyces
43: 916, 1965. chlamydosporus from a saline location in Bay
Teleomorph: In the genus Corollospora. Minette, Alabama. We also include this species
◙ V. anglusa Abdel-Wahab & Nagah., in the key as it may be encountered in marine
Mycoscience (in press). habitats.
Teleomorph: Corollospora anglusa
1. Chlamydospores with 11-34 (-64)-septa, 95-370 (-
Abdel-Wahab & Nagah. 500) × 8-16 µm ..................................X. rhizophorae
Mycelium septate, branched, hyaline to 1 Chlamydospores with 5-19-septate, 95-420 × 26-42
pale brown, conidiophores cylindrical, septate, µm...............................................X. chlamydosporus
simple, conidiogenous cells monoblastic,
integrated, generally terminal, conidia Zalerion R. T. Moore & Meyers, Can. J.
acrogenous, solitary, branched, septate, hyaline, Microbiol. 8: 408, 1962 ............................... (1)
main axis from which arise 2-3 side branches, ◙ Z. maritima (Linder) Anastasiou, Can. J. Bot.
typically tetra-radiate, and sporulating 41: 1136, 1963 (Type species).
profusely on wood in a layer of seawater (Fig. Helicoma maritimum Linder, Farlowia 1: 405,
1944.
82). Helicoma salinum Linder, Farlowia 1: 406, 1944.
Zalerion nepura R.T. Moore & Meyers, Can. J.
1. Conidia 1-4 (-7)-septate in the main axis V. prolifera Microbiol. 8: 413, 1962.
1. Conidia 1-2 (-4)-septate in the main axisV. ramulosa Zalerion eistla R.T. Moore & Meyers, Can. J.
1. Conidia 1-2-septate in the main axis ........ V. anglusa Microbiol. 8: 413, 1962.

160
 Fungal Diversity

◙ Zalerion xylestrix R.T. Moore & Meyers, Can. 5. Conidia one-celled ................................................... 6
J. Microbiol. 8: 414, 1962. 5. Conidia 3-7-septate, on various substrata ..................
Zalerion raptor R.T. Moore & Meyers Can. J. .............................................................Stagonospora
Microbiol. 8: 415, 1962. 5. Conidia with appendages ....................................... 11
Teleomorph: Lulwoana uniseptata.
Conidia pluricellular, monacrogenous, 6. Conidia filiform ....................................................... 7
6. Conidia spherical or ellipsoidal................................ 8
single or branched, on simple conidiophores,
filaments variable in length and septation, 7. Conidia 10-16 × 0.5 µm, on wood .............................
coiled irregularly, producing a balled .......................................... Halonectria milfordensis
appearance, and subhyaline to fuscous. Three 7. Conidia 9-12 × 1.5-2 µm, saprobic on Avicennia
Zalerion species were accepted (Kirk et al., wood. ................................ Rhabdospora avicenniae
2001), two marine and one of terrestrial origin
8. Occurring on Nypa fruticans palm........................... 9
(Buczacki, 1973), while four species described
8. Occurring on wood and other substrata ................. 10
by Moore and Meyers (1962), based largely on
their different enzyme activity, have been 9. Pycnidium superficial with a large reddish-brown
reduced to synonymy with Z. maritima. neck, conidia 3.5-5 × 2-2.5 µm ..................................
Isolates of Z. arboricola (a terrestrial species) ............................................ Pleurophomopsis nypae
are not congeneric with Z. maritima and isolate 9. Pycnidium on a subiculum, conidia 2.4-4 × 1.8-2.4
µm..............................................Nypaella frondicola
ATCC 20868 was transferred to a new genus
Glarea lozoyensis, based on DNA 10. On wood, conidia 2.5-4.5 × 2.5-2 µm
fingerprinting (Bills et al., 1999). Zalerion ...................................Phialophorophoma litoralis
varia likewise is not congeneric with Z. 10. On Rhizophora, conidia 11-18 × 3-4 μm.
maritima, but groups with G. lozoyensis in the ............................................Phomopsis mangrovei
10. On various substrata, pycnidia flask-shaped
Leotiaceae, Leotiales, and is therefore assigned
superficial or submerged .........................................
to the new genus Halenospora (see page 154). .......................... Phoma and Macrophoma species
Zalerion varia isolates differ greatly in their
morphology and collections made may not be 11. Conidia 1-2-septate, with three polar appendages,
monophyletic. A strain of Z. xylestrix clusters on various substrata............................................. 12
with Z. maritima and is considered by 11. Conidia unicellular, cylindrical, 39-49 × 6.5-8.5 μm,
with 3-4 apical appendages formed by
Campbell et al. (2005) to be conspecific with it, fragmentation of a sheath, on Juncus roemerianus
supporting the findings of Anastasiou (1963a) ........................................... Tiarosporella halmyra

COELOMYCETES 12. Conidiogenesis phialidic, conidia 16-23 × 2.5-4 µm,

on Phragmites communis ........................................
1. Conidia in an acervulus............................................ 2 ................................. Pseudorobillarda phragmitis
1. Conidioma eustromatic ............................................ 3 12. Conidiogenesis holoblastic, conidia 10-14 × 3-4.5
1. Conidioma pycnidial ................................................ 4 µm, on Rhizophora mangle, and other Rhizophora
species ...............................Robillarda rhizophorae
2. Conidia with polar spines, 11-15 × 2.5-3.5 µm, on
drift wood........................ Dinemasporium marinum 13. Conidia one-celled .............................................. 14
2. Conidia with no polar spines, hyperparasite of 13. Conidia one transverse septum............................ 15
Haloguignardia species, conidia 2-5 × 2-2.5 µm 13. Conidia muriform................................................ 17
.............................................. Gloeosporidina cecidii
14 Conidia with funnel-shaped polar appendages,
3. On Rhizophora wood, roots and seedlings, conidia 3- fusiform or naviculate .......................Koorchaloma
6 × 1-1.5 µm..........................Cytospora rhizophorae 14. Conidia ellipsoidal, dark brown, thick-walled, with
3. On Phragmites, conidia 17-5-75 × 2.5-5 µm 5-9 tentacle-like appendages ...................................
.............................Cytoplacosphaeria phragmiticola .......................................... Octopodotus stupendus
3. On Nypa fruticans, conidia 3-4.5 × 1.2-1.6 µm 14. Conidia with no appendages, 4-8 × 4-6 μm
............................................... Plectophomella nypae ............................................Coniothyrium obiones

4. Conidia hyaline ........................................................ 5 15. Saprobic on salt marsh plants.............................. 16

4. Conidia coloured .................................................... 13 15. Saprobic on driftwood, conidia yellowish-brown, 6-
8 × 3.5-4.5 μm........................ Diplodia orae-maris

161
16. On Halimione, conidia yellow-brown, 9-12 × 4-5 Ascochyta salicorniae var. salicorniae-patulae
μm .............................................Ascochyta obiones Trotter, Annls. Mycol. 3: 30, 1905.
16. On Salicornia, conidia olive to light brown, 4-8 × Ascochyta salicorniae-patulae (Trotter) Melnik,
4-6 μm ................................. Ascochyta salicorniae Nov. Sist. Niz. Rast., 12: 205, 1975.
Ascochyta salicorniae Trotter, Annls. Mycol. 2:
17. Conidia with no sheath, 3-transverse septate, 1- 536, 1904, nom. illegit
longi-septate, 10-20 × 7-13 μm ............................... Diplodia salicorniae Jaap, Verh. Bot. Ver. Prov.
................................. Camarosporium roumeguerii Brandenburg 49: 16, 1907.
17. Conidia with a polar cap-like appendage at each Pycnidia immersed, erumpent, ovate,
end, 22-42 × 10-17 μm, on Ammophila subglobose, ellipsoidal or pyriform, ostiolate,
...........................Amarenographium metableticum epapillate to papillate, coriaceous, olive brown,
17. Conidia with a pronounced mucilaginous sheath, 5-
transverse septate, 1-longiseptate, 20-34 × 9-20 μm
brown to black, solitary or gregarious,
......................................Camarosporium palliatum conidiogenous cells phialidic, flask-shaped to
pyriform, hyaline, conidia blastic, ellipsoidal,
Amarenographium O.E. Erikss., Mycotaxon obovate or cylindrical, 1-septate, slightly
15: 199, 1982 ............................................... (1) constricted at the septum, smooth-walled,
A. metableticum (Trail) O.E. Erikss., hyaline to yellowish to pale brown. Parasitic or
Mycotaxon 15: 199, 1982 (Type species). saprophytic on the salt marsh plants Halimione
Camarosporium metableticum (Trail) Grove, portulacoides, Salicornia europaea, S.
British Stem and Leaf Fungi, Cambridge 2: 108, 1937. herbacea and S. patula.
Camarosporium graminicola Ellis and Everh.,
Proc. Acad. Nat. Sci. Philadelphia, 1893: 161, 1893. 1. Ascospore with a sheath, hyaline to yellowish or light
Camarosporium metableticum Trail, Scottish brown, 10-12 × 4-7 µm, ...................... A. salicorniae
Naturalist 8: 267, 1886. 1. Ascospores without a sheath, 9-12 × 4-5 µm, pale
Diplodina ammophilae Trail, Scott. Natural. 10: yellowish to brown....................................A. obiones
76, 1889.
Leptosphaeria ammophila (Lasch) Rehm, Asc.: Camarosporium Schulzer, Verh. zool.-bot. Ges.
69, 1882.
Wien 20: 649, 1870...................................... (2)
Teleomorph: Amarenomyces ammophilae
C. palliatum Kohlm. & E. Kohlm., Marine
(Lasch) O.E. Erikss.
Mycology, The Higher Fungi: 519, 1979.
Pycnidia subglobose, immersed, ostiolate,
C. roumeguerii Sacc., Michelia 2: 112, 1880.
papillate, clypeate?, coriaceous, black, solitary, Camarosporium obiones Jaap, bot. Ver. Prov.
conidiogenous cells cylindrical, conidia Brandenburg 47: 97, 1905.
ellipsoidal to trapezoidal, muriform, 3-9 Pycnidia immersed or erumpent,
transverse septate, 1-2 longitudinal septate, subglobose or ellipsoidal, ostiolate, epapillate
slightly constricted at the septa, fuscous, apical or papillate, dark brown, solitary or gregarious,
cells lighter, bearing cap-like, gelatinous, paraphyses present, simple, hyaline, filiform,
striate appendages at either end of the nonseptate, conidiogenous cells phialidic,
conidium. Reported from bark, maritime flask-shaped, simple, hyaline, conidia
grasses and salt marsh plants. subglobose, ellipsoidal or oblong, enteroblastic,
monophialidic, muriform, 1-6 transverse septa,
Ascochyta Lib., Pl. Crypt. Arduenna 1 (Praef.):
1-4 longitudinal septa, slightly constricted at
8, 1830.......................................................... (2)
the septa, smooth, gold, yellowish-brown,
A. obiones (Jaap) P.K. Buchanan, Mycol. Pap.
olive-brown to brown, with a gelatinous sheath,
156: 28, 1987.
Ascochytula obiones (Jaap) Died., Annls mycol.
lacking appendages. Both species occur on the
10: 141, 1912. salt marsh plants Halimione portulacoides, and
Diplodia obiones Jaap, Verh. Bot. Ver. Prov. various Salicornia species.
Brandenburg 47: 96, 1905.
A. salicorniae Magnus, in Jaap, Schr. 1. Conidia 5-septate, 20-34 × 9-20 µm, with a
Naturwiss. Ver. Schleswig-Holstein 12: 345, pronounced gelatinous sheath, on Salicornia species
...............................................................C. palliatum
1902. 1. Conidia 3-septate, 10-20 × 7-13 µm, lacking a sheath,
Stagonosporopsis salicorniae (Magnus) Died., on stems and leaves of salt marsh plants....................
Annls. Mycol. 10: 141-142, 1912. .......................................................... C. roumeguerii

162
 Fungal Diversity

Coniothyrium Corda, Icon. Fung. (Abellini) 4: scattered over the surface, conidiophores
38, 1840........................................................ (1) simple, septate, elongate-cylindrical, hyaline,
◙ C. obiones Jaap, Schr. naturw. Ver. conidiogenous cells cylindrical, smooth,
Schlesing-Holstein 14: 29, 1907. hyaline, producing conidia at their tips, conidia
Pycnidia subglobose, immersed to 11-15 × 2.5-3.5 µm, fusoid-allantoid,
erumpent, ostiolate, short papillate, coriaceous, unicellular, smooth, hyaline, with a single hair-
brown, gregarious, conidiogenous cells like appendage at each end. Reported from
phialidic, flask-shaped, unicellular, hyaline, driftwood and rarely collected since its original
conidia 4-8 × 3.5-6 µm, ellipsoidal, ovoid, description.
subglobose, unicellular, smooth-walled,
olivaceous to pale brown, lacking a sheath or Diplodia Fr., Annls. Sci Nat. Bot., Sér. 2, 1:
appendages. Occurs on the salt marsh plant 302, 1834...................................................... (2)
Halimione portulacoides. D. orae-maris Linder, Farlowia 1: 403. 1944.
D. thalassia N.J. Artemczuk, Mikol. Fitopatol.
Cytoplacosphaeria Petr., Annls mycol. 17: 79, 14: 95, 1980.
1920.............................................................. (1) Pycnidia immersed to erumpent,
C. phragmiticola Poon & K.D. Hyde, Bot. Mar. subglobose, ostiolate, papillate, pale to dark
41: 148, 1998. brown, glaborous, membranous or coriaceous,
Pycnidia large (circa 800 μm), loosely solitary or gregarious, conidiophores short
aggregated into a stroma with 1-5 locules, cylindrical, hyaline, conidia 1-3-septate,
immersed in the substratum (Phragmites slightly to markedly constricted at the septa,
communis), ostiole indistinct, brown, ovoid, ellipsoidal, cylindrical, yellow to brown,
ellipsoidal to lenticular, conidiogenous cells yellow in mass. Collections of D. orae-maris
enteroblastic, phialidic, pronounced collarette, on driftwood while D. thalassia was isolated
conidia 17.5- 75 × 2.5-5 μm, straight or slightly from marine sediments (Fig. 83, Artemczuk,
curved, thin-walled, 0-5 septate, not constricted 1980).
at the septa, hyaline (Poon and Hyde, 1998).
1. Conidia 6-8.5 × 3.5-7.5 µm, on wood. D. orae-maris
Cytospora Ehrenb., Sylv. mycol. berol. (Berlin): 1. Conidia larger 8.5-12 × 5.5- 6 µm, in sediments .......
................................................................D. thalassia
28, 1818........................................................ (1)
C. rhizophorae Kohlm. & E. Kohlm.,
Mycologia 63: 847, 1971.
Pycnidia immersed, composed of several
locules, irregular morphology, ostiolate,
epapillate, coriaceous, brown, solitary or
gregarious, conidiophores cylindrical, filiform,
simple, septate producing conidia apically,
hyaline, conidia 3-6 × 1.1-1.5 µm, allantoid or
ellipsoidal-cylindrical, unicellular, hyaline,
Fig. 83. Diplodia thalassia. Bar = 5 µm.
lackng a sheath or appendages, conidia released
in a mass. Frequently collected on mangrove Gloeosporidina Petr., Annls. mycol. 19: 214,
trees, particularly roots and seedlings of 1921.............................................................. (1)
Rhizophora species (Kohlmeyer and G. cecidii (Kohlm.) B. Sutton, The
Kohlmeyer, 1971). Coelomycetes (Kew): 517. 1980.
Sphaceloma cecidii Kohlm., J. Elisha Mitchell
Dinemasporium Lév., Annls. Sci. Nat. Bot., Scient. Soc. 88: 255, 1972.
Sér. 35: 274, 1846. ....................................... (1) Acervuli discoid, olive coloured, later
D. marinum Sv. Nilsson, Bot. Not. 110: 321, black, conidiogenous cells phialidic, cylindrical
1957. or slightly attenuated, simple with funnel-
Acervuli cupulate, superficial, greyish- shaped mouths, conidia unicellular, hyaline,
black, sessile, setae needle-shaped, brown smooth-walled, ellipsoidal, 3-4.5 × 1.8-2.5 µm,
163
truncate at the base, with a cylindrical basal a b
appendage. Growing on the galls of
Haloguignardia species on Sargassum natans.

Koorchaloma Subram., J. Indian bot. Soc. 32:

124, 1953...................................................... (2)
K. galateae Kohlm. & Volkm.-Kohlm., Bot.
Mar. 44: 147, 2001.
K. spartinicola V.V. Sarma, S.Y. Newell &
K.D. Hyde, Bot. Mar. 44: 321, 2001.
Conidiomata stromatic, sporodochial,
punctiform, scattered, superficial, salmon to
orange, with dark brown setae that are 3-4-
septate, conidiophores erect, branched, thin-
walled, smooth, hyaline, conidiogenous cells
monoblastic, terminal, subcylindrical with
collarettes, smooth, hyaline, conidia blastic- Fig. 84. a. Koorchaloma galateae.b. K. spartinicola,
phialidic, solitary, fusiform or naviculate, with conidia with polar appendages. Bars: a = 5 µm, b = 10
µm.
gelatinous appendages at both ends, smooth,
Conidiomata pycnidial, formed on a
hyaline, appendages formed by fragmentation
subiculum, superficial, apricot to pale brown
of a sheath?, becoming inverted, funnel-shaped,
with central ostiole; conidiophores branched at
then splitting into thin, radiating filaments,
base of the conidiomata; conidiogenous cells
conidial secession schizolytic and released in a
phialidic; conidia holoblastic, hyaline, smooth
slimy mass (Figs. 84a, b) (Sarma et al., 2001).
walled, ellipsoidal, unicellular, 2.4-4 × 1.8-2.4
Koorchaloma galateae was described from
µm, lacking a sheath or appendages (Hyde and
Juncus roemerianus (Kohlmeyer and
Sutton, 1992). Saprotrophic on Nypa fruticans.
Volkmann-Kohlmeyer, 2001b) occurring
between 5-25 cm above the rhizome while K.
Octopodotus Kohlm. & Volkm.-Kohlm.,
spartinicola occurs throughout the intertidal
Mycol. Res. 95: 117, 2003 ........................... (1)
range of its host Spartina alterniflora, with a
O. stupendus Kohlm. & Volkm.-Kohlm.,
salinity range of 14 to 29 ‰ (Sarma et al.,
Mycol. Res. 95: 117. 2003 (Type species).
2001).
Conidiomata pycnidial, immersed in the
1. Conidiomata setae 40-65 × 6.5-10 µm, conidia 15.5- mesophyll between vascular bundles of leaves
21.5 × 4.5-6.5 µm, on Juncus roemerianus................ of Spartina alterniflora, light brown, ostiolate
.................................................................K. galateae with a short papilla, peridium 6-12 µm thick;
1. Conidiomata setae 45-80 × 3-5 µm, conidia 14.5- conidiophores reduced to conidiogenous cells
20.5 × 4.5-6.8 µm, on Spartina alterniflora............... that are discrete, conoid or irregular polygonal
........................................................... K. spartinicola
all around the peridial wall; macroconidia
Macrophoma (Sacc.) Berl. & Voglino., Atti aseptate, ellipsoidal, dark brown, thick-walled,
Soc. Veneto-Trentina Sci. Nat. (Padova) 10: verruculose, surrounded by a gelatinous sheath
172, 1886.............................# Macrophoma sp. and with 5-9 tentacle-like appendages. Hyaline
A wide range of Macrophoma strains microconidia also produced on the peridium
have been isolated from intertidal wood, near the ostiolar canal (Kohlmeyer and
mangrove roots, marsh plants, seawater and Volkmann-Kohlmeyer, 2003a).
sediments. However few have been fully
identified. Phialophorophoma Linder, Farlowia 1: 403,
1944.............................................................. (1)
Nypaella K.D. Hyde & B. Sutton, Mycol. Res. P. litoralis Linder, Farlowia 1: 402, 1944.
96: 210, 1992 ............................................... (1) Pycnidia immersed, subglobose or
N. frondicola K.D. Hyde & B. Sutton, Mycol. ellipsoidal, ostiolate, papillate or epapillate,
Res. 96: 210, 1992. subcarbonaceous, brown to black, glaborous,

164
 Fungal Diversity

solitary, conidiophores cylindrical, branched, flask-shaped, conidia ellipsoidal to subglobose,

septate, with terminal phialides, conidiogenous unicellular, hyaline, smooth walled, with no
cells monophialidic, enteroblastic, conidia sheath or appendages. Species marked * have
ellipsoidal, obovoid or clavate, unicellular, all been isolated from marine soils (Pawar et
smooth-walled, hyaline, no sheath or al., 1967), and thus may not be regarded as
appendages (Kohlmeyer and Kohlmeyer, 1979). obligately marine.
Occurring on a variety of substrata, drift and Phoma species are ubiquitous,
intertidal wood, bark and dead roots of geographically widespread, occurring in
Avicennia marina. variety of environments and habitat niches, as
saprophytes, endophytes, as biodeteriogens and
Phoma Sacc., Michelia 2: 4, 1880 ............... (8) parasitic on plants and animals. Because of
*P. capitulum V.H. Panwar, P.N. Mathur & difficulties in their identification, most are
Thirum., Trans. Br. Mycol. Soc. 50: 261, 1967. simply referred to as Phoma species. Some
P. glomerata (Corda) Wollenw. & Hochapfel, 3,000 Phoma epithets have been recorded in
Z. Parasitkde 3: 592, 1936. MycoBank (Crous et al., 2004), but Borerema
Aposphaeria fibricola (Berk.) Sacc., Syll. fung. et al. (2004) only accept circa 223 species.
(Abellini) 3: 176, 1884. Aveskamp et al. (2008) questioned the validity
Aposphaeria glomerata (Corda) Sacc., Syll. fung.
(Abellini) 3: 175, 1884. of some of these and proposed avenues for
Coniothyrium glomeratum Corda, Icon. fung. future research, including DNA-barcoding
(Prague) 4: 39, 1840. (Herbert et al., 2002).
Peyronellaea alternariaceum (F.T. Brooks & We have not attempted a key to Phoma
Searle) Goid., Annali Sper. agr., n. s. 6: 92, 1952. species recovered from marine habitats as the
Peyronellaea fibricola (Berk.) Goid., Annali Sper.
agr., n. s. 6: 92, 1952. only way to resolve their taxonomy is to
Peyronellaea glomerata (Corda) Goid., Atti examine cultures and undertake molecular
Accad. Naz. Lincei, Rendiconti Adunanze Solenni 1: sequencing of their DNA.
455, 1946.
Peyronellaea glomerata (Corda) Goid., ex Phomopsis (Sacc.) Bubák, Öst. bot. Z. 55: 78,
Togliani, Annali Sper. agr. n.s. 6: 93, 1952.
Phoma alternariaceum F.T. Brooks & Searle,
1905.............................................................. (1)
Trans. Br. Mycol. Soc. 7: 192, 1921 (1920). P. mangrovei K.D. Hyde, Mycol. Res. 95:
Phoma fibricola Berk., 1853. 1149, 1991.
* P. hibernica Grimes, M. O’Connor & Teleomorph: In the Diaporthales?
Cummins, Trans. Br. Mycol. Soc. 17: 100, Pycnidia immersed, ellipsoid or
1933. subglobose, ostiolate, solitary or gregarious,
P. laminariae Cooke & Massee, Grevillea 18: black, conidiophores branched, stout to filiform,
53, 1889. septate, hyaline, conidiogenous cells phialidic,
* P. multispora V.H. Pawar, P.N. Mathur. & determinate, integrated, hyaline, cylindrical
Thirum., Trans Br. Mycol. Soc. 50: 260, 1967. with a termial collarette, conidia holoblastic,
* P. navium Woron. Arbeit. Biol. Wolga- fusiform or ellipsoidal, 11-18 × 3-4 µm,
Station, 8: 61 1925. straight, unicellular, hyaline, rounded apically,
* P. ostiolata V.H. Pawar, P.N. Mathur. & truncate at the base, no sheath or appendages
Thirum., Trans Br. Mycol. Soc. 50: 262, 1967. (Hyde, 1991d). Intertidal on prop roots of
P. suaedae Jaap, Schr. Naturw. Ver. Rhizophora species, no teleomorph reported.
Schleswig-Holstein 14: 27, 1907.
# Phoma spp. Plectophomella Moesz, Magyar Bot. Lapok.
Teleomorph: one Phoma sp. has 21: 13, 1922 ................................................. (1)
Tremateia halophila Kohlm., Volkm.-Kohlm. P. nypae K.D. Hyde & B. Sutton, Mycol. Res.
& O.E. Erikss., as its teleomorph. 96: 211, 1992.
Pycnidia ellipsoidal to subglobose, Conidiomata pycnidial, pale, immersed,
immersed or erumpent, ostiolate, papillate or ostiolate; conidiophores hyaline, 1-septate;
epapillate, coriaceous, brown to black, conidiogenous cells phialidic, conidia
conidiogenous cells conical, hyaline, phialidic, holoblastic, hyaline, unicellular, smooth,
165
cylindrical 3-4.5 × 1.2-1.6 µm. Collected on Rhabdospora (Durieu & Mont.) Mont., Syll.
Nypa fruticans. Gen. Spec.: 277, 1856 .................................. (1)
R. avicenniae Kohlm. & E. Kohlm.,
Pleurophomopsis Petr., Annls. mycol. 22: 156, Mycologia 63: 851, 1971.
1924.............................................................. (1) Pycnidia immersed, subglobose,
P. nypae K.D. Hyde & B. Sutton, Mycol. Res. unilocular, ostiolate, papillate or epapillate,
96: 213, 1992. coriaceous, dark brown to black, solitary or
Conidiomata pycnidial, superficial, gregarious, conidiophores cylindrical or
reddish brown, central orange ostiole; attenuate, simple, conidia produced at their
conidiophores at the base and sides of apex, hyaline, conidia filiform, 9-12.5 × 1.5-2
conidiomata; conidiogenous cells phialidic; µm, unicellular, straight mostly curved, hyaline,
conidia holoblastic, hyaline, aseptate ellipsoid lacking appendages or a sheath (Kohlmeyer
to fusiform, smooth 3.5-5 × 2-2.5 µm and and Kohlmeyer, 1979). Occurring on bark and
saprotrophic on Nypa fruticans. Hyde and pneumatophores of Avicennia and proproots
Sutton (1992) indicate that the assignment of and trunks of Rhizophora trees, releasing
this species to “Pleurophomopsis is debatable, conidia in a cirus.
but there appears to be no better choice at
present”. Cultures of these three coelomycetes Robillarda Sacc., Michelia 2: 8, 1880 ......... (1)
(Nypaella, Plectophomella, Pleurophomopsis) R. rhizophorae Kohlm., Can. J. Bot. 47: 1483,
are required to enable determination of their 1969.
phylogenetic relationship. Teleomorph : in the Amphisphaeriaceae,
Xylariales (Rungjindamai et al., unpublished
Pseudorobillarda M. Morelet, Bull. Soc. Sci. data).
nat. Archéol. Toulon Var 175, 1968 ............ (1) Pycnidia immersed or erumpent,
P. phragmitis (Cunnell) M. Morelet, Bull. Soc. ellipsoidal, unilocular, epapillate, subco-
Sci. nat. Archéol. Toulon Var 175: 6, 1968. riaceous, black, glabrous, solitary or gregarious,
Teleomorph: in the Dothideomycetes, conidiogenous cells conical or cylindrical,
(Rungjindamai et al., unpublished data). hyaline, conidia ellipsoidal, 1-septate, slightly
Pycnidia immersed, scattered, globose, dark constricted at the septum, smooth-walled,
brown, paraphyses present, conidiogenous cells hyaline, with 3-4 apical radiating appendages.
subcylindrical, lageniform, colourless, smooth, All collections of R. rhizophorae appear to be
conidia 1-septate, fusiform, both ends rounded, from Rhizophora wood.
hyaline, smooth, eguttulate, 16-23 × 2.5 µm,
with 2-4 appendages, growing on grasses Stagonospora (Sacc.) Sacc., Syll. fung.
(Plaingam et al., 2005). Appendages in (Abellini) 3: 445, 1884................................. (1)
Pseudorobillarda sojae are non-cellular and S. haliclysta Kohlm., Bot. Mar. 16: 213, 1973.
arise from the outer wall layer of the conidium Stagonospora spp.
as an outgrowth (Plaingam, 2002). Pycnidia immersed to superficial,
Pseudorobillarda phragmitis (Fig. 85) was subglobose, ostiolate, epapillate, coriaceous,
collected on pine and yellow poplar panels dark brown, solitary or gregarious, conidio-
from estuarine waters (salinity 3-16‰) by genous cells conoidal, simple, hyaline, conidia
Johnson and Hughes (1960) but originally formed at the apex, conidia holoblastic, 20-
known from Phragmites communis (Cunnell, 27.5 × 3.5-4.5 µm, fusiform with rounded ends,
1958). 3-septate, slightly constricted at the septa,
Although Pseudorobillarda and hyaline, smooth-walled, with an mucilaginous
Robillarda morphologically share some cap on the upper cell. Occurring on the brown
common features (pycnidial, septate hyaline seaweed Pelvetia canaliculata (Kohlmeyer,
conidia with 3-4 polar appendages), 1973a) and known only from the type
phylogenetically they are distantly related collection. Many Stagonospora species have
(Rungjindamai et al., unpublished data). been reported from salt marsh plants,

166
 Fungal Diversity

in the ecology of mangrove ecosystems,

recycling leaf and ligno-cellulose, creating
particulate material and dissolved organic
matter for other organisms in the food web. Yet
much needs to done to quantify this activity,
especially for filamentous fungi.
Marine ascomycetes account for the
greatest diversity of filamentous fungi, also
occurring on diverse substrata (Kis-Popa,
2005). However despite intense studies at the
molecular level, many genera cannot be
assigned to families, and some do not fit into
any orders that have been described to date.
Often new orders have to be erected to
Fig. 85. Pseudorobillarda phragmitis. Conidium with accommodate them: Jahnulales (Pang et al.,
three polar appendages. Bar = 20 µm. 2002), Lulworthiales (Kohlmeyer et al., 2000)
and Koralionastetales (Campbell et al., 2008).
especially Spartina, Phragmites, Juncus and Despite an earlier view that some marine
Carex and warrant further study (Jones, 1963; ascomycetes had originated in the marine
Jones and Oliver, 1964; Hughes, 1969; environment, we now have documented
Henningson, 1974; Gessner and Kohlmeyer, evidence that they are secondary invaders of
1976). the marine milieu. There is no information as
to when this event (s) occurred, but the fact that
Tiarosporella Höhn., Mitteil. bot. Ins. tech.
they are worldwide in their distribution
Hochsch. Wien 1: 82, 1924.......................... (1)
suggests an early migration, even before the
T. halmyra Kohlm. & Volkm.-Kohlm.,
separation of land masses.
Mycotaxon 59: 79, 1996.
Currently we can distinguish several
Pycnidia ellipsoidal, obpyriform,
marine lineages; for example
immersed, ostiolate, papillate, dark brown,
solitary, conidiogenous cells cylindrical to
Basidiomycota:
elongate conical, conidia cylindrical,
1. Physalacriaceae clade in the eugarics
unicellular, hyaline, smooth, with 3-4 apical
2. Nia clade, eugarics
appendages that are tentaculiform, undulate,
3. Peniophoraceae clade
gelatinous, tapering to a thin tip, formed by
4. Ustilaginomycetes clade
fragmentation and eversion of a sheath
5. Tremellomycetes, Cystofilobasidiales
(Kohlmeyer and Volkmann-Kohlmeyer, 1996).
clade
Found on senescent culms of Juncus
6. Agaricostilbomycetes, Agaricostilbales
roemerianus, 7 to 63 cm above the rhizome
group
and considered to be facultative species.
7. Microbotryomycetes, Sporidiobolales
TOTAL 8. Leucosporidiales group
ANAMORPHIC ASCOMYCETES ..... (103)
Ascomycota:
Concluding remarks 9. Saccharomycetales group
10. Halosphaeriales clade
The past 50 years has seen a remarkable 11. Lulworthiales clade
advance in our knowledge of marine fungi, a 12. Hypocreales clade
group that some might regard as a minor 13. Koralionastetales clade
assemblage of fungi of little general interest. 14. Torpedospora/Swampomyces clade
However, marine fungi play a vital role in the 15. Jahnulales clade
recycling of organic matter in coastal and 16. Verrucariales clade
oceanic waters. They are particularly important 17. Cleistothecial bitunicate ascomycetes.
167
 Thus there has been multi-transitions studies or pathological studies of those
from terrestrial to marine habitats often infecting commercial fish (Duc et al., 2009).
resulting in a reduced fruiting body, probably Most studies have been directed at the
and adaptation to the aquatic habitat (Binder et filamentous fungi, in particular the lignicolous
al., 2006). Undoubtedly this transition was ascomycetes, while anamorphic fungi present
from terrestrial to freshwater and possibly in mangrove mud’s, ocean sediments are
mangrove ecosystems to completely oceanic neglected and not considered part of the marine
conditions. There is an overlap in the genera fungal community. There needs to be a new
found in terrestrial/freshwater/mangrove direction set for many taxa remain to be
ecosystems, e.g. Leptosphaeria, Phaeosphaeria discovered in these habitats. Too rigid
and Massarina species, occur in all these delineation of what is marine must be
ecosystems and are also found in fully saline overcome, the important fact is they are
waters. However, few occur under fully repeatedly isolated from marine habitats, what
submerged coniditions. An interesting group is their role in nature? Sam Meyers in the early
has been members of the Jahnulales, initially 1960’s proposed the term thalassiomycetes for
described from freshwater and terrestrial those fungi recovered from marine habitats.
habitats, but also in a peat swamp and with Perhaps we would have made greater
Manglicola a marine genus (Pinruan et al., advancement in the study of fungi in the sea,
2002; Suetrong et al., 2009). had such a term been adopted.
So what of the future of marine
mycology? While some geographical areas New species included in this volume, and
have been widely surveyed (temperate coastal published since Hyde et al. (2000).
lignicolous fungi, tropical mangrove fungi)
vast areas have not been studied (South Basidiomycota
America, Africa) and cold water areas. Haloaleurodiscus mangrovei N. Maek., Suhara
Similarly, while some substrata have been & K. Kinjo
intensively studied (cellulosic and ligno-
cellulose, sand dwelling species, fungi on sea- Ascomycota
grasses), there is much to be done to sample Bitunicate
seaweeds, root inhabiting species, parasites of Caryospora australiensis Abdel-Wahab &
marine animals, endophytes of marine plants E.B.G. Jones
and animals (Kis-Papo, 2005; Raghukumar, Decaisnella formosa Abdel-Wahab & E.B.G.
2008). Jones
Taxonomic studies of marine fungi have Leptosphaerulina mangrovei Inderb. & E.B.G.
made considerable progress in the last 20 years, Jones
especially with advent of molecular techniques. Mauritiana rhizophorae Poonyth, K.D. Hyde,
Our understanding of the physiology and Aptroot & Peerally
biochemistry of marine fungi still remains Platystomum scabridisporum Abdel-Wahab &
patchy, with most studies confined to the effect E.B.G. Jones
of salinity on growth, movement of ions into Trematosphaeria malaysiana Alias, McKeown,
mycelium, their ability to degrade cellulose and S.T. Moss & E.B.G. Jones
lignin (Mouzouras et al., 1988; Pointing et al.,
1998). Another area that has attracted research Unitunicate
interest is the source of new chemical Alisea longicola J. Dupont & E.B.G. Jones
structures and bioactive compounds from Anthostomella spissitecta Kohlm. & Volkm.-
marine fungi. However, this has been confined Kohlm.
to meet the needs of the pharmaceutical A. torosa Kohlm. & Volkm.-Kohlm.
industry (Höller et al., 2000; Pan et al., 2008; Astrocystis nypae G.J.D. Smith & K.D. Hyde
Jones, 2008; Jones et al., 2008), rather to A. selangorensis G.J.D. Smith & K.D. Hyde
explore their role in the ecology of marine Corollospora anglusa Abdel-Wahab & Nagah.
fungi. Few have undertaken broad enzymatic C. baravispora Steinke & E.B.G. Jones sp. nov.

168
 Fungal Diversity

C. indica Pasannarai, K. Ananda & K.R. Trichocladium melhae E.B.G. Jones, Abdel-
Sridhar Wahab & Vrijmoed
C. portsaidica Abdel-Wahab & Nagah.
Dryosphaera tennuis Andrienko Coelomycetes
Halosarpheia unicellularis Abdel-Wahab & Koorchaloma galateae Kohlm. & Volkm.
E.B.G. Jones Kohlm.
Havispora longyearbyenensis K.L. Pang & Koorchaloma spartinicola V.V. Sarma, S.Y.
Vrijmoed Newell & K.D. Hyde
Nemania maritima Y.M. Ju & J.D. Rogers
Phyllachora paludicola Kohlm. & Volkm.- Acknowledgements
Kohlm.
Pontogeneia microdictyi Kohlm. & Volkm.- We are grateful to many friends and colleagues
for their generosity with time, friendship, ideas, critical
Kohlm. comments and helpful guidance and advice: Kevin
Pseudohalonectria halophila Kohlm. & Hyde for reading sections of the manuscript and for
Volkm.-Kohlm. endless discussions; David Hibbett and Mark Binder for
Pseudolignincola siamensis Chatmala & E.B.G. their comments on the text and phylogeny of marine
Jones basidiomycetes; H. Harada for assistance with lichen
literature; Anthony Fletcher and Patrick M McCarthy,
Remispora minuta E.B.G. Jones, K.L. Pang & for reading drafts of the manuscript and the supply of
Vrijmoed photographs; Amy Rossman for invaluable comments
Remispora spitsbergenensis K.L. Pang & on the taxonomy of the nectrioid marine taxa; Jack Fell
Vrijmoed for his assistance with data on marine yeasts; Timothy
Rostrupiella danica Jørg. Koch, K.L. Pang & James for his comments on the origin of the
Chytridiomycota; Mohamed Abdel-Wahab for his
E.B.G. Jones comments on mangrove fungi, Jorgen Koch for valuable
Sablecola chinensis E.B.G. Jones, K.L. Pang & discussions and unfailing assistance with finding
Vrijmoed difficult fungi for our phylogenetic studies; Akira
Savoryella melanospora Abdel-Wahab & Nakagiri for the supply of cultures, photographs and
E.B.G. Jones valuable assistance in so many ways; Teck Koon Tan
for endless discussion and continued assistance with
Swampomyces aegyptiacus Abdel-Wahab, El- field work; Jan and Brigit Kohlmeyer who have assisted
Shar. & E.B.G. Jones in so many ways with the identification and taxonomy
Swampomyces clavatispora Abdel-Wahab, El- of various fungi; Jinx Campbell for supplying us with a
Shar. & E.B.G. Jones prepublication manuscript of their paper on the new
Thalespora appendiculata Chatmala & E.B.G. order Koralionastetales; the late Sam Meyers whose
wisdom was so generously given; Lilian Vrijmoed for
Jones loyal support and assistance with field work; the late
Tirispora mandoviana V.V. Sarma & K.D. Steve Moss who did so much to advance our knowledge
Hyde of the ultrastructure of marine ascomycetes and for all
those discussions when sitting at the electron
Hyphomycetes microscope; various Ph.D. students, (R.A. Eaton, P.J.
Byrne, D.J. Miller, G. Rees, V .Cuomo, R. Mouzouras,
Acremonium fuci Summerb., Zuccaro & W. C. Panebianco, S. Sivichai, S. Stanley, R.G. Johnson,
Gams C.A. Farrant, T.A. Baker, D.W.T. Au., T.A. McKeown,
Amorosia littorlais Mantle & D. Hawksw. S.A. Alias, S.Y. Hsieh, A. Pilantanapak, R. Sadaba and
Arthrobotrys mangrovispora Swe, Jeewon, Y. Musa), who have assisted with many of the projects
Pointing & K.D. Hyde that led to this monograph. Siew Moi Phang,
Vikineswary, Deen Kuthubuttheen, Tony Whalley, Gill
Cumulospora varia Chatmala & Somrith. Hughes, Trevor Steinke, Aisyah Alias, Apiradee
Halenospora varia (Anastasiou) E.B.G. Jones. Pilantanapak, Sung-Yuan Hsieh and H.S. Chang, Alun
Halosigmoidea parvula Zuccaro, J.I. Mitch. & Jones and Jonathan Batten, Howard and Liz Lintott,
Nakagiri Director and staff Friday Harbour Marine Station, USA,
Penicillium dravuni J.E. Janso Somsak Sivichai, Aom Pinnoi, Umpava Pinruan,
Souwalak Phongpaichit, Rattaket Choeyklin, Anupong
Plectosporium oratosquillae Duc, Yaguchi & Klaysuban, Orathai Supaphon, Sita Preedanon for all
Udagawa their help with field work and logistic support. Various

169
herbaria for the loan of material and supply of cultures: Canadian Journal of Botany 81: 598-600.
Kew, IMI, ATCC, CBS, Khun Wanchern Potacharoen, Abdel-Wahab, M.A., Nagahama, T. and Abdel-Aziz,
and BCC. F.A. (2008). Two new Corollospora species and
This monograph would not have been possible one anamorph based on morphological and
without the financial support of BRT, Thailand (BRT: molecular data. Mycoscience (Accepted for
R_245002, R_248002, R_249001, R_251006, publication, in press).
R_149017) and BIOTEC; The Croucher Foundation,
Hong Kong; BIOTEC, Thailand and industrial support; Acero, F.J., González, V., Sánchez-Ballesteros, J., Rubio,
TOTAL E & P Thailand, TOTAL FOUNDATION. V., Checa, J., Bills, G.F., Salazar, O., Plata, G.
Ka-Lai Pang would like to thank the National and Peláez, F. (2004). Molecular phylogenetic
Science Council of Taiwan (Project No. NSC 97-2621- studies on the Diatrypaceae based on rDNA-ITS
3-019-002) for financial support. sequences. Mycologia 96: 249-259.
Satinee Suetrong would like to thank Graduate Alias, S.A. (1996). Ecological and taxonomic studies of
School Prince of Songkla University for financial lignicolous marine fungi in Malaysian mangroves.
support. PhD. Thesis, University of Portsmouth.
We thank Umpava Pinruan and Nattawut Alias, S.A., Hyde, K.D. and Jones, E.B.G. (1996).
Rungjindamai for assistance with the literature search Pyrenographa xylographoides from Malaysia and
and indexing. Particular thanks to Paul Kirk and Index Australia mangroves. Mycological Research 100:
Fungorum for assistance with the taxonomy and search 580-582.
for literature, and Lesley Ragab for library assistance. Alias, S.A., E.B.G. Jones and J. Torres. (1999). Intertidal
We thank various colleagues for allowing us to fungi from the Philippines, with a description
use their photographs (acknowledged in the text), and Acrocordiopsis sphaerica sp. nov. (Ascomycota).
Figures 63, 64 reprinted with permission from Jørgen Fungal Diversity 2: 35-41.
Koch, Ka-lai Pang and E.B. Gareth Jones. 2007. Alias, S.A., Moss, S.T. and Jones, E.B.G. (2001).
Rostrupiella danica gen et sp. nov., a Lulworthia-like Cucullosporella mangrovei, ultrastructure of
marine lignicolous species from Denmark and the USA. ascospores and their appendages. Mycoscience 42:
Bot Mar 50: 294-301. 405-411.
We are particularly gratefull to Dr. Shaun Alker, A.P., Smith, G.W. and Kim, K. (2001).
Pennycook for checking and correcting the fungal Characterization of Aspergillus sydowii (Thom &
authorities and references. Church), a fungal pathogen of Caribbean sea fan
We would like to record our profound thanks to corals. Hydrobiologia 460: 105-111.
Professor Morakot Tanticharoen, Dr. Kanyawim Ananda, K. and Sridhar, K.R. (2001). Aniptodera indica,
Kirtikara and Dr. Lily Eurwilaichitr, BIOTEC, Thailand a new species of mangrove inhabiting ascomycete
for endless support and encouragement without which from west coast of India. Journal Environmental
this monograph would not have seen the light of day. Biology 22: 283-286.
E.B.G.J. would like to thank Marion Jones, for Anastasiou, C.J. (1963a). The genus Zalerion Moore et
dedicated support, unfailing understanding, and for Meyers. Canadian Journal of Botany 41: 1135-
faithfully following him around the world in search of 1139.
marine fungi, and to whom this monograph is dedicated. Anastasiou, C.J. (1963b). Fungi from salt lakes II.
Ascomycetes and Fungi Imperfecti from the
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