Biodiversity and Conservation 6, 721±738 (1997)

Biodiversity of bryophilous ascomycetes

È BBELER
PETER DO
Institut fuÈr Systematische Botanik der Ludwig-Maximilians-UniversitaÈt MuÈnchen,
Menzinger Straûe 67, D-80638 MuÈnchen, Germany

Received 1 August 1996; accepted 19 September 1996

Up to now, about 300 species of ascomycetes are known to grow obligately on the gametophytes of
mosses or hepatics; these belong to more than 80 ascomycete genera, some of them unknown
elsewhere, and at least nine orders. They vary greatly in relation to the mode of nutrition
(necrotrophic, biotrophic and gall-inducing parasites), the host organs infected, host speci®city, and
geographical distribution. Diversity of these fungi is illustrated by spore outlines of 36 species.
Hyphae growing super®cially, inter- or intracellularly, and various types of appressoria and
haustoria are illustrated. Adaptations to the unique substratum, such as the formation of minute
reduced ascomata containing gelatinous material, a preference for moist microhabitats such as the
interlamellar spaces of leaves in Polytrichales, and phototropic reactions are discussed. Convergent
evolution leading to similar morphological features is rather common. The colonization of mosses
and hepatics by ascomycetes is a very frequent though generally neglected phenomenon. Some
bryophytes seem to grow always with their speci®c parasites. It is supposed that species diversity
among the bryophilous ascomycetes is as high as among the lichenicolous ones. Numerous new taxa
remain to be described, but only a small fraction of bryophytes has been proved to be infected so far.
Keywords: biodiversity; bryophilous ascomycetes; bryophytes as hosts; spore types;
hyphal characters; adaptive features.

Introduction
Bryophilous fungi grow on gametophytic or sporophytic organs of bryophytes,
muscicolous species infecting mosses (Musci), and hepati(ci)colous species colonizing
liverworts (Hepaticae). These fungi are de®ned by their ecology as are lichenicolous or
coprophilous species, and their systematic positions, modes of nutrition and biological
peculiarities may be quite di€erent. Literature on fungi living on the haploid stage of
bryophytes is reviewed by Felix (1988). Pyrenocarpous ascomycetes as well as mitosporic
fungi present on both generations are compiled by Racovitza (1959) in a monographic
study. For bryophilous lichenized ascomycetes see Faegri (1980), Kalb (1994), Obermayer
and Poelt (1994), and Poelt (1986). Sterile intracellular fungal hyphae frequently form
endomycorrhiza-like associations in numerous hepatics (Paul, 1916; Pocock et al., 1984;
Pocock and Duckett, 1985a, b; Duckett and Renzaglia, 1988; During and Van Tooren,
1990). Symbiotic associations between endophytic hyphae and hepatics are refered to as
`mycothalli' by Boullard (1979, 1988). In this paper, I will concentrate on fruit-body-forming
ascomycetes which develop on the haploid, gametophytic stage of mosses or hepatics.
Bryophytes are unique host plants in numerous ways. One of the most important
features `is a life cycle that involves an alternation of generations with a dominant, free-
living gametophyte' (Wyatt, 1982). Vegetative multiplication by various modes is frequent
and reveals an enormous capacity for regeneration. Very important for the establishment

0960-3115 Ó 1997 Chapman & Hall

722 DoÈbbeler
of interactions with fungi seems to be the small size of mosses and hepatics, which means a
limited availability of the substratum, their inde®nite growth in the form of dense colonies,
mats and cushions without periodic leaf-fall, and an often pronounced poikilohydric
organization. Successful fungal colonization requires morphological and physiological
adaptation to these special conditions.
Bryophytes are generally considered as phylogenetically ancient plants with a slow rate
of evolution over at least 300 million years (Szweykowski, 1984a, b). Miller (1982) regards
them as the oldest extant terrestrial plants which represent the level of evolution associated
with transmigration to the land. Frey (1977) argues that the origin of the large groups of
mosses must date back to before the late Palaeozoic age. Therefore, `fungi growing on
living parts of mosses and ferns are considered to be the most ancient plant parasites'
(von Arx, 1985). Savile (1968; p. 651) points out that `obligate parasitism in the fungi is
not a belated evolutionary bypath, but a fundamental attribute of primitive groups'. This
is supported by the fact that some rather archaic insects are present in bryophytes (Gerson,
1982, p. 320).
Bryophilous fungi do not receive much attention, and are not even mentioned in the
relevant bryological literature (Schuster, 1966, 1983, 1984; Smith, 1982; Scho®eld, 1985).
The situation in mycology is comparable. This is certainly a result of their often extremely
small, sometimes light-coloured or hyaline ascomata hidden between host organs. Even
for a keen and experienced observer it is a time-consuming and troublesome task to detect
them. In addition, the overlap between bryology and mycology is very small in contrast to
the situation in lichenicolous fungi. Lack of known economic importance is also an
important factor. According to the general view, `bryophytes, even herbarium specimens,
are hardly ever attacked by microorganisms' (Ando and Matsuo, 1984, p. 181). In reality,
both mosses and hepatics represent excellent hosts for fungi.
What should be the purpose of studies on bryophilous fungi? Of course, they are part of
the still mostly unknown biological diversity of our planet, which is not only of taxonomic
interest. Analyses of morphological and ecological features of the moss-inhabiting fungi,
their relationship with non-bryophilous fungi, host interactions, speci®city, geographical
distribution, and decomposition enlarge our ideas and concepts of the systematics and
biology of fungi in general, and the parasitic ascomycetes in particular. These are
remarkable and unique interactions between members of two di€erent kingdoms of
organisms. Relatively simple anatomical structures of mosses and hepatics easily allow
host and parasite to be distinguished at the cellular level. A further argument is that these
fungi, when better known, can be expected to contribute to the clari®cation of taxonomic
problems and evolutionary relationships of their hosts, having co-evolved in close
relationships with them.
The goal of this article is to draw attention to a highly diverse, widespread,
unquestionably successful and intriguing group of ascomycetes, which undoubtedly
occupies one of the least explored ecological niches for fungi today.

Methods
Under ®eld conditions it is rather dicult, if not impossible, to detect bryophilous asco-
mycetes, although necrotrophic species often causing distinct decolorized patches within
green colonies of mosses and hepatics can attract attention. However, not all the whitish,
yellowish or brownish spots in bryophytes yield ascomata. Randomly collected samples of
Bryophilous ascomycetes 723
potential hosts must be screened in the laboratory with a stereomicroscope for the
presence of fungi. In most cases, it is recommendable or even indispensable to wet the
plants thoroughly. Bryophytes covered by soil particles and detritus should be cleaned
with ¯owing tap water. Unabsorbed water has to be soaked up to avoid inconvenient light
re¯ections. Ascomata can easily be confused with air bubbles, all kinds of detritus,
colonies of cyanobacteria and algae, pollen grains, parts of the host itself (e.g. brotes,
gemmae) or the cysts and eggs of small animals. It is easier to count the number of
basidiomes of the bracket-fungi in a forest than to know the number of ascomata in a
handful of Hylocomium splendens. I fear some mycologists will even look in vain for fruit-
bodies of certain species in material in which I have ascertained they are indeed present.
A very e€ective method for obtaining access to material from di€erent and remote
geographical regions, is the examination of potentially infected specimens in bryological
collections. Bryophytes present in biosystematic collections (`herbaria') represent a rich
source of accidentally gathered fungal parasites, unknown to their collectors, curators and
more signi®cant, often unknown to science. The same applies to ascomycetes on algae
(Kohlmeyer, 1975) and to fungi growing on lichens (lichenicolous fungi). Nevertheless,
necrotrophic species are apparently underrepresented in bryological herbaria, as collectors
prefer healthy green mosses and hepatics.
Fungal infections in colonies of di€erent species of bryophytes deserve special attention.
These colonies constitute natural infection experiments, by which host speci®city often can
be demonstrated easily and conclusively.
It is useful to have at least a basic knowledge of bryophyte morphology and living
habits. Without knowing the capacity of acrocarpous mosses to innovate from
subterranean rhizoids (Meusel, 1935; Wigglesworth, 1947), the ecology of Octospora
would remain unclear.
Over the last 20 years, I have carefully examined tens of thousands of individual plants
of mosses and hepatics, and thousands of collections in order to obtain an idea of their
fungus biota. However, in almost every case I feel that we still know only a very small
fraction of the total systematic, biologic and geographic diversity of the bryophilous fungi.

Systematics
Up to the present, about 300 species (including ca 50 of Octospora) in more than 80 genera
of ascomycetes have been described as growing on the gametophytes of mosses and
hepatics. Their systematic position is quite diverse since no less than nine orders are
involved: Dothideales, Hypocreales, Leotiales, Pezizales, Verrucariales and, to a more
marginal extent, Arthoniales, Lecanorales, Ostropales and Sordariales. The genera
Acrospermum and Dactylospora remain in families incertae sedis. The heterogenity of the
bryophilous fungi is re¯ected at least partially by the diversity of their ascospore-types,
which vary greatly in size, shape, septation, colour, and other features (Fig. 1).
Several genera are known exclusively from mosses and hepatics (obligate bryophilous
genera), e.g. Bryodiscus, Bryoscyphus, Bryosphaeria, Epibryon, Hypobryon, Octospora and
Potriphila. They apparently represent independent phylogenetic lineages, in which
speciation occurred after attaining the bryophilous habit. However, some non-bryophilous
genera do exist which include obligate parasites of mosses and hepatics, for instance
Dactylospora heimerlii, Muellerella frullaniae and several species of Hymenoscyphus and
Nectria. The `waste basket' genus Epibryon with about 35 species is the largest bryophilous
724 DoÈbbeler
Bryophilous ascomycetes 725
`genus'. Octospora and related genera form a biologically de®ned isolated group within the
terrestrial Pezizales morphologically characterized by elaborate infecting structures. These
fungi should be separated taxonomically as they represent a `class of discomycete similar
to the Erysiphaceae' postulated more than 65 years ago (Corner, 1929).
The appropriate systematic position of a bryophilous ascomycete is often dicult to
ascertain, as the tiny and reduced fruit-bodies are a priori poor in characters and con-
vergent evolution has often led to surprisingly similar morphological features.
Fortunately, hyphal characters o€er a rich source of valuable additional information for
the de®nition of taxa, though this has generally been neglected by other authors.

Infected host organs

Octospora libussae is the only macromycete species known to grow on the algal-like
protonemata of mosses (DoÈbbeler and Itzerott, 1981). However, there is no essential
di€erence between rhizoids and protonematal ®laments (Goode et al., 1992). Several
species of Octospora attack the subterranean vital rhizoids of acrocarpic mosses
(DoÈbbeler, 1979b, 1993; Itzerott, 1983b; Senn-Irlet, 1988), thus being somewhat spatially
independent from the leafy part of the gametophyte. Only few ascomycetes, such as
Bryoscyphus conocephali and B. marchantiae, parasitize thalloid hepatics (Spooner, 1984).
Several species of Riccia are recorded as hosts for di€erent ascomycetes (Srinivasan, 1939;
Stephens, 1939; Bapna, 1962; Bonar, 1965; Singh and Pavgi, 1979). However, the over-
whelming majority of ascomycetes infects leaves or stems of mosses or leafy hepatics.
The hepaticolous Nectria egens develops perithecia mainly on the distal part of the host
leaves, where the photosynthetic activity is highest (Corner, 1935). Microhabitats
protected against rapid moisture loss are often prefered or even exclusively selected for
fruit-body formation, while hyphae also colonize other parts of the host plant. Ascomata
of Bryorella semiimmersa are partly immersed in the stems of Lophoziaceae. Teichospora
jungermannicola and related species form individual fruit-bodies in the leaf-axils of foliose
hepatics, sometimes resulting in a regular zigzag line of ascomata decreasing in size to-
wards the plant apex. However, even from these moisture retaining places ascospore
discharge into the open air still has to be possible. This may be the reason why ascomata
are not developed other than on the generally favoured lower leaf side but at the leaf
margins, as in Nectria praetermissa.
Epibryon hypophyllum on Radula forms fruit-bodies in which the delicate hymenium is
on the protected ventral leaf surface. The leaf blade is perforated by an apical papilla, so

b
Figure 1. Di€erent types of ascospores of bryophilous ascomycetes (all drawn to same scale = 30 lm). Coloured
spores are indicated by stippling. 1, Acrospermum adeanum; 2, `Lizonia' sphagni; 3, Octosporella ornithocephala;
4, Lamprospora retispora; 5, Dawsicola neglecta; 6, Pitya madothecae; 7, Gloeopeziza interlamellaris; 8, Grimmicola
parasiticus; 9, Pseudonectria brongniartii; 10, Muellerella frullaniae; 11, Nectria racomitrii; 12, Epibryon
perrumpens; 13, Protothelenella polytrichi; 14, Bryosphaeria epibrya (spores desintegrating into single cells already
in the asci); 15, Hypobryon bicolor; 16, Bryosphaeria cinclidoti; 17, Bryostroma trichostomi; 18, Belonioscyphella
hypnorum (germinating spores with conidiogenous cells); 19, Bryodiscus hepaticarum; 20, Trichonectria pellucida;
21, Epibryon ®liforme (germinating spores with conidiogenous cells); 22, Epibryon cryptosphaericum; 23, Bryorella
cryptocarpa; 24, Pleosphaeria sp.; 25, Epibryon diaphanum; 26, Bryomyces caudatus; 27, Potriphila navicularis;
28, Dactylospora heimerlii; 29, Dawsophila callichroma; 30, Philobryon anuliferum; 31, Nectria sanramonensis;
32, Nectria lankesteri; 33, Nectria brenesii (spores desintegrating into single cells already in the asci);
34, Bryostroma rhacomitrii; 35, Lizonia emperigonia; 36, Bryostroma halosporum.
726 DoÈbbeler
that the ostiole is situated on the dorsal surface of the leaf. Perforation occurs only in the
distal part of leaves which are not covered by a leaf beneath, allowing unhindered asco-
spore liberation. Several ascomycetes on hepatics with di€erently overlapping leaves
exhibit the same strategy.
Hepaticolous species such as Calonectria fullaniae and Nectria perianthicola are con-
®ned to young sporophytes still enclosed by the perianth. When mature, the apices of the
perithecia perforate the perianth wall to achieve ascospore liberation. Infected
sporophytes do not develop further and decompose.
Bryostroma trichostomi forms ascomata in the leaf axils of pottiaceous mosses, and
Epibryon muscicola on the adaxial leaf side of diverse Tortella species. A similar behaviour
is known from the leotialian Grimmicola parasiticus on Grimmia (DoÈbbeler and Hertel,
1984). Bryorella acrogena selects the leafy shoot apices of pleurocarpous mosses for fruit-
body formation (DoÈbbeler, 1984), as does Bryostroma rhacomitrii on Rhacomitrium
lanuginosum. Lizonia species form densely aggregated ascomata in the antheridial cups of
Polytrichum. Exactly the same niche is occupied by the systematically unrelated Durella
polytrichina (Racovitza, 1940, 1946).
The longitudinal spaces between the photosynthetic leaf-lamellae of Polytrichales
represent one of the most favoured microhabitats. There are more than twenty species of
di€erent systematic positions known to colonize these phylogenetically ancient and stable
niches. They o€er striking examples of convergent evolution, for example the formation of
compressed, sometimes extremely small, and reduced ascomata. Even heavy infections
with hundreds of fruit-bodies in a single leaf of Dawsonia superba do not lead to any visible
symptoms (DoÈbbeler, 1981). Certain species of leaf-inhabiting ascomycetes on poly-
trichaceous mosses show distinct distribution patterns on that host, prefering apical, basal
or lateral parts of the host leaves (DoÈbbeler, 1986).

Modes of nutrition
Necrotrophic species, such as Acrospermum adeanum, Belonioscyphella hypnorum,
Bryostroma necans, all species of Lizonia, and several of Nectria, cause necrotic and often
conspicuously discoloured lesions among healthy green plants of the host colony.
Centrifugally expanding and clearly zonated infections are characteristic of Nectria
muscivora destroying acrocarpous mosses on walls in wet winter months. This is one of the
most frequent muscivorous species in the city of Munich. Comparable but larger infections
can form spectacular fairy rings or ring-like systems in arctic and antarctic moss
communities (Hawksworth, 1973; Longton, 1973, 1988; Fenton, 1983; GamundõÂ and
Spinedi, 1988; p. 476); one of the fungi present has been identi®ed as Thyronectria
hyperantarctica (Pegler et al., 1980). The hyphae of all bryophyte-destroying ascomycetes
hitherto analysed grow within their host cells (Fig. 2, number 6). Acrospermum adeanum
and Nectria muscivora attack mosses with an additional super®cial mycelium; this is visible
to the naked eye as a small whitish zone at the periphery of expanding infections.
Most species on bryophytes are biotrophic parasites which do not or just slightly
damage their hosts. Hyphae of these species grow super®cially on the host organs (Fig. 2,
numbers 1 and 2) or within the cell walls irregularly (Fig. 2, number 4) or in a regular
manner (Fig. 2, number 5). Super®cial hyphae prefer to grow along the anticlinal walls,
often following exactly the host cell reticulum. The mycelium of Bryochiton perpusillus
Bryophilous ascomycetes 727
develops subcuticularly or within the outermost part of the host cell wall (Fig. 2, number
3). In general terms, biotrophic parasitism is correlated with a super®cial and/or inter-
cellular mycelium, and necrotrophic parasitism with an intracellular mycelium.
Systemic infections with the fungal mycelium following the, at least theoretically, un-
limited growth of the host colony are frequent. Only the growing points and adjacent
tissues are free from hyphae. Several pyrenocarpous and discocarpous parasites re¯ect this
developmental process by presenting di€erently aged fruit-bodies along the axis of the host
plant. The distal younger parts contain primordia, while the proximal older parts carry
mature ascomata. Vegetative propagules of the host infected by fungal hyphae probably
represent combined distribution units of the host and its parasite. This could be an
e€ective way for an ascomycete to become established on epiphyllous hepatics in
rainforests, which often propagate by large, disc-shaped gemmae (Thiers, 1988). Redhead
(1980) indicates that the hepaticolous agaric Gerronema pseudogrisella is frequently
dispersed as mycelium established on gemmae of the liverwort Blasia pusilla.
The only known gall-inducing species of bryophilous ascomycetes belong to the genus
Octospora. Galls are formed by meristematic regions of the subterranean rhizoids of
mostly acrocarpous mosses. These hypertrophies sometimes resemble rhizoid gemmae,
which are also known from certain acrocarpous mosses (Whitehouse, 1966; Risse, 1987).
Several gall types can be distinguished, belonging to at least six species of Octospora. The
always one-celled, globular galls di€er depending on their position on the rhizoids, and
micromorphological characters such as the number of appressoria produced (DoÈbbeler,
1979b; DoÈbbeler and Itzerott, 1981; Itzerott and DoÈbbeler, 1982; Itzerott, 1983a;
Senn-Irlet, 1988). A careful analysis of the contact zone between Octospora species and
their hosts will certainly reveal further gall-forming species.
Several bryophilous ascomycetes develop preferentially on older, dying parts of the
plant, for example several species of the genus Epibryon in the interlamellar spaces of
Polytrichales. Bates (1979) points out that in Pleurozium schreberi there is no sharp limit
between the dead and vital regions, the ageing parts only showing certain decreased
physiological processes. The occurrence of obligate, true saprophytes on the gametophyte
of mosses and hepatics seems doubtful.

Appendages of the mycelium

Hyphae of many bryophilous fungi form sessile or stalked appendages (Fig. 3). Together
with other hyphal features, they are most useful for species description and the de®nition
of higher taxa. Appressoria, haustoria, or a combination of both, are related to attach-
ment and absorption, whereas conidiogenous cells play a role in asexual reproduction or
spermatization.
Nectria hylocomii and related species develop appressoria more or less irregularly on their
host cells (Fig. 3, number 2). The appressoria of N. egens are located at the cell junctions
(Corner, 1935), and the elaborate appressoria of the two species of Dawsomyces exclusively
occupy the periclinal walls of single host cells (Fig. 3, number 1). Complex appressoria-like
structures at the internal apical side of the mucilage cells of Sphagnum are produced by
Discinella schimperi (Fig. 3, number 3; Redhead and Spicer, 1981). The surprising regularity
by which some fungi select particular sites of the host surface for appressorium formation is
evidence of the importance of speci®c microhabitats (Emmett and Parbery, 1975, p. 158).
728 DoÈbbeler
Bryophilous ascomycetes 729
Haustoria originating directly from a super®cial mycelium without the formation of
appressoria may be intraparietal or intracellular. Peg-like haustoria con®ned to the host
cell walls are typical of Bryomyes hemisphaericus and other congeneric species (Fig. 3,
number 5). Potriphila navicularis develops haustoria within individual cells of the hair-like
leaf trichomes of Polytrichum alpinum (Fig. 3, number 4), while Gloeopeziza cuneiformis
infects the cells of the leaf blade of several species of Polytrichum with intracellular
haustoria (DoÈbbeler, 1996a). The combination of appressoria and intraparietal haustoria
characterizes the genus Hypobryon (DoÈbbeler, 1983). Conspicuous appressoria and
intracellular haustoria occur in Octospora and related genera (Fig. 3, number 6; Corner,
1929; Racovitza and Racovitza, 1945; Racovitza, 1946; DoÈbbeler 1979b, 1980b).
In several cases, conidiogenous cells have been observed on the hyphae. Small phialidic
cells are typical for Hypobryon (DoÈbbeler, 1983) and Epibryon ®liforme (DoÈbbeler and
MenjõÂvar, 1992). Similar anamorphs seem to be more frequent, but are dicult to detect.

Adaptations
The colonization of bryophytes by fungi requires biochemical adaptations to the host
metabolism, as in other fungal parasites. In addition, extreme environmental conditions in
ecological niches such as late snow-lie areas in arctic regions or the phyllosphere in tropical
rainforests are re¯ected by speci®c adaptations of both hosts and parasites. Some of the
fungal features related to their bryophilous habit are discussed here. A phenological
synchronization between host and parasite is demonstrated by the Polytrichum-inhabiting
species of Lizonia; the ascomata develop only when uninfected plants produce antheridia.
Perhaps the most striking adaptation of the bryophilous fungi is the small size of their
fruit-bodies. Many ascomata are smaller than 100 lm in diameter. Those of Bryochiton
monascus and B. perpusillus, hardly reach 50 lm in diameter, and, spectacularly, Epibryon
endocarpum has ascomata only 25±35 lm in diameter obligately formed within individual
cells of Plagiochila asplenioides (DoÈbbeler, 1980a). These fungi are among the smallest
fruit-body-forming ascomycetes. The reduced size may be interpreted as a response to the
limited size of the substratum and to the small amount of available nutrients. Redhead
(1984) points out that bryophilous members of Omphalina and other genera are normally
smaller than their non-bryophilous relatives, interpreting the reduced size of the
basidiomata as a response to a limited availability of nutrients.
Mechanisms assuring survival in dry conditions are very important for bryophilous
fungi, and a preference for moist microhabitats on the host plants has been mentioned

b
Figure 2. Positions of hyphae in relation to the host cells (scale = 25 lm). 1, Epibryon plagiochilae on Plagiochila
porelloides: super®cial hyphae following the anticlinal walls of the host cells (Germany, Bavaria, Kreuzeck,
24 July, 1980, Poelt & DoÈbbeler 3484, M); 2, The same leaf in section; 3, Bryochiton perpusillus on Polytrichum
piliferum: transverse section through a leaf showing subcuticular hyphae at the abaxial side of the leaf (Austria,
Tyrol, Fimbertal, 28 July, 1979, DoÈbbeler 3548, M); 4, Dactylospora heimerlii on Plagiochila asplenioides s. l.:
transverse section through the outer part of the stem; hyphae grow irregularly within the cell walls (Germany,
Bavaria, Jachenau, 1 November, 1978, DoÈbbeler 3164, M); 5, Punctillum hepaticarum on Plagiochila sp.: trans-
verse section through the stem, hyphae growing in the angles of the cell walls nearly exclusively in the longitudinal
direction of the axis (Tasmania, Mt Barrow, without date, Jarman 1477, M); 6, Nectria muscivora on Didymodon
rigidulus: hyphae within cells of the basal part of the leaf, note the ®ne hyphal pegs perforating the anticlinal cell
walls (Germany, North Rhine-Westphalia, Olpe, 7 January, 1978, DoÈbbeler 2852, M).
730 DoÈbbeler
Bryophilous ascomycetes 731
above. Perithecia or perithecia-like ascomata are more frequently developed than
apothecia-like ones; the latter are less xerophytic than perithecia (Eckblad, 1968, p. 151).
The fruit-bodies of the hepaticolous Octosporella jungermanniarum (Pezizales), which re-
semble perithecia, are interpreted by Corner (1929) as modi®ed apothecia consequent to
depauperation and xerophily.
As in lichen-forming ascomycetes, gelatinous material may be produced in the
hymenium and/or ascomatal wall as a water storing substance (Ingold, 1959; P®ster, 1976).
Mucilage in the hymenium often reacts with Lugol's solution resulting in either a reddish
or a bluish colour after pretreatment with potassium hydroxide (DoÈbbeler, 1978, p. 12).
Baral (1987) terms this conspicuous reaction `hemiamyloid'. It is regarded as one of the
diagnostic characters for the genera Epibryon, Hypobryon and some other bryophilous
taxa.
Discomycetes with reviving ascomata which are closed when dry and open when moist in
order to expose the hymenium (Haines and McKnight, 1977; Sherwood et al., 1980;
Sherwood, 1981; Kropp and Carpenter, 1984) are to be expected among the bryophilous
fungi.
Phototropic movements of di€erent parts of ascomata towards the incident light have
been sporadically observed in di€erent bryophilous ascomycetes. They are typical in those
on Polytrichum sexangulare, which grows in dense carpets (DoÈbbeler, 1987). The
formation of fruit-bodies mainly in the upper halves of leaves and a positive phototropism
facilitates ascospore discharge into the open air. The phototropic reaction is achieved in
di€erent ways: by the whole ascomata, by ascomatal necks (as in Protothelenella
polytrichi), by laterally orientated ostioles (e.g. certain Bryochiton species), or by the ascus
apices (e.g. Gloeopeziza interlamellaris of Leotiales). The phototropic curving of asci has
previously been reported mainly in Pezizales (Ingold, 1971, p. 14).
Species such as Epibryon diaphanum occurring mostly on Hypnaceae, or as parasites of
epiphyllous hepatics like E. deceptor (sp. nov. ined.) on Radula ¯accida show a remarkable
tendency towards reduced fertility. Scattered ascomata contain few asci with large spores,
and anamorphs are unknown. These fungi apparently occupy stable niches in which a high
reproductive potential does not yield a selective advantage. The same phenomenon occurs
in the coprophilous genus Sordaria and in the Laboulbeniales (Raper, 1968). We face the

b
Figure 3. Hyphae with appressoria and/or haustoria (scale = 20 lm). 1, Dawsomyces mirabilis on Dawsonia
superba: appressoria without haustoria covering single periclinal walls of cells of leaf-lamellae; on the left side two
appressoria in surface view, on the right side section through an infected leaf lamella (Tasmania, Castra Road, 24
June, 1891, Burchard, M±holotype); 2, Nectria hylocomii on Hylocomium splendens: irregularly oriented
appressoria without haustoria on the cells of a leaf in surface view (Germany, Bavaria, Spitzingsee, 8 October,
1977, DoÈbbeler 2788, M); 3, Discinella schimperi (sterile infection) on Sphagnum squarrosum: infected
mucilaginous cells of the apical parts of the stem with an intracellular appressorium closely appressed to the
internal cell wall; above in surface view, below optical section (Canada, Ontario, Nipigon, 1 July, 1956, Heinrich,
M); 4, Potriphila navicularis on Polytrichum alpinum: intracellular haustoria in cells of trichomes arising from the
leaf-lamellae, without appressoria (Austria, Tyrol, Fimbertal, 28 July, 1979, DoÈbbeler 6759, M±holotype); 5,
Bryomyces hemisphaericus on Plagiochila porelloides: hyphae grow super®cially and follow strictly the anticlinal
cell walls, develop peg-like haustoria into the cell walls, and lack appressoria; above seen from above, below
section (Germany, Bavaria, Rossstein, 7 October, 1978, DoÈbbeler 3351, M); 6, Octosporella ptilidii on Ptilidium
ciliare: appressoria with haustoria; on the left side two stalked appressoria seen from above, to the right section
through an appressorium with peg-like perforation-tube and intracellular haustorium (Switzerland, Lower
Engadine, Lavin, 1 September, 1984, MuÈller & DoÈbbeler 5925, M±holotype).
732 DoÈbbeler
paradoxical situation that the delicate and short-lived epiphyllous hepatics, the `biological
nomads' of the rainforest (Richards, 1984), represent stable niches for hepaticolous fungi.

Host speci®city
The choice of hosts has to be carefully considered because `the host is part of the parasite's
environment and must be part of the species circumscription' (Savile, 1993, p. 270).
Indeed, nearly all bryophilous fungi exhibit a certain degree of host speci®city. Notwith-
standing the fact that many species are known only from the original collection, there are
rather well ± documented examples of species depending on single host species and others
infecting a wide range of hosts. Between these extremes intermediate types occur. Epibryon
metzgeriae is known only from Apometzgeria pubescens (DoÈbbeler, 1985), and
Pseudonectria brongniartii is recorded exclusively from Frullania dilatata (DoÈbbeler, 1984).
The very common Epibryon plagiochilae is speci®c to Plagiochila asplenioides and the
closely related P. porelloides and does not infect Pedinophyllum interruptum (syn.
Plagiochila interrupta), a member of the only additional genus of the Plagiochilaceae in
Europe. Pedinophyllum interruptum is easily confused with Plagiochila porelloides (Grolle,
1969). I checked more than 60, mostly Central European, specimens of Pedinophyllum
interruptum in the Botanische Staatssammlung MuÈnchen (M) without ®nding any
infection. The presence of Epibryon plagiochilae is a useful way of determining Plagiochila
asplenioides s.1.
After studying hundreds of polytrichaceous mosses, I consider Potriphila navicularis to
be speci®c for Polytrichum alpinum in Central Europe. The infection of Pogonatum
urnigerum (syn. Polytrichum urnigerum) by this parasite has never been observed, though
the moss can be confused with Polytrichum alpinum. On the other hand, the extremely
common Epibryon pogonati-urnigeri on Pogonatum urnigerum does not attack Polytrichum
alpinum. These examples show that the Polytrichum-inhabiting ascomycetes are excellent
taxonomists, sometimes knowing their hosts better than bryologists (DoÈbbeler, 1996b).
The four accepted species of Lizonia are known exclusively from polytrichaceous
mosses. European populations of Polytrichum commune, P. formosum and P. sexangulare
are frequently infected. These hosts, and P. piliferum have speci®c parasites, de®ned by host
selection and anatomical characters. Mosses only once recorded as hosts for Lizonia are
Pogonatum sp. from Nepal (DoÈbbeler, 1978), Oligotrichum aligerum from Alaska (Parriat
and Moreau, 1954) and Dendroligotrichum dendroides from Chile (DoÈbbeler, unpubl.).
Dactylospora heimerlii is known to grow on nine di€erent host genera of the
Jungermanniales, most commonly on Plagiochila asplenioides, which has received special
attention as a rich host for unusual fungi. However, until now neither Frullania nor
Radula, which both belong to the same order, are documented as hosts for Dactylospora,
though many specimens of these hepatics were examined (DoÈbbeler and Triebel, 1985).
A very wide host range characterizes Acrospermum adeanum, with most of the infected
bryophytes belonging to the Hypnobryales, growing on bark of trees or on calcareous rocks
(DoÈbbeler, 1979a). Sphagnum supports, unsurprisingly in view of its relationships, a my-
cobiota of its own with several genus- or species-speci®c parasites, for example Epibryon
turfosorum, Discinella schimperi, and diverse species of Lasiosphaeria. Bryochiton perpusillus
is frequently recorded from hepatics such as Ptilidium and the moss Polytrichum (DoÈbbeler,
1978). In this case, conspeci®city should be re-investigated carefully.
Bryophilous ascomycetes 733
Didymosphaeria marchantiae on Marchantia polymorpha is now regarded as a synonym
of the cosmopolitan Phaeodothis winteri, which occurs on an astonishingly wide range of
hosts, including fungi as well as ligni®ed and non-ligni®ed phanerogams (Aptroot,
1995a, b). If the conspeci®city is proved, this is one of the very few examples of a
facultative bryophilous species.
Members of the systematically isolated Polytrichaceae (including Dawsonia) are nearly
always infected by fungi. More than 40 ascomycetes are known to be associated with them
(DoÈbbeler, 1986). These large or even gigantic mosses with long-lived gametophytes are
characterized by a structural complexity in kind or degree which is achieved by no other
group of bryophytes (Smith, 1971, p.76). They provide many di€erent microhabitats. In
addition, the antiquity of Polytrichum and allied genera has probably favoured the
evolution of a large number of parasites restricted to them. The lichen genus Peltigera is
also considered to be particularly old, partly because of the many obligately lichenicolous
fungi known only from this host genus (Hawksworth, 1980, 1982; Hawksworth and
Santesson, 1988).

Frequency and geographical distribution

To understand the frequency and the geographical distribution of the fungi on bryophytes,
two facts should be taken into consideration. First, the presence of a fungus is recorded by
the presence of its fertile structures, i.e. ascomata. However, mosses or hepatics growing
under natural conditions are actually never free from hyphae, i.e. non-sporing fungal
infection. Normally, hyphal features do not permit the distinction of taxa at a lower level,
an exception being some fungi with characteristic hyphal appendages. Discinella schimperi,
a highly specialized parasite of Sphagnum squarrosum, is very frequent in the northern
circumpolar region. The presence of sterile mycelia with unique appressoria (Fig. 3,
number 3) indicates that the infection rate may be higher than 50 % (Redhead and Spicer,
1981). From a sterile mycelium with characteristic appressoria and haustoria on the
hepatic Schistochila aligera I can recognize an Octosporella-like fungus from Sri Lanka
(DoÈbbeler, 1978, p. 107). Ascomata of Octosporella, and the recently separated genus
Filicupula (Yao and Spooner, 1996), are until now recorded only from Europe, Costa Rica
and Venezuela.
Second, the presence or absence of fruit-bodies in a host population depends on the size
of the collection. Infections are often randomly distributed in patches within mosses or
hepatics. The investigation of more individual host plants, or larger cushions, increases the
probability of discovering parasites. In general terms, bryophilous fungi are much more
frequent than records indicate.
DoÈbbeler (1987) concluded that about half of the collections of Polytrichum
sexangulare deposited in reference collections are inhabited by at least one of the six
species of ascomycetes which attack this host. Several parasites of the tropical epiphyllous
Radula ¯accida occur regularly in the whole American as well as the African area of the
host (DoÈbbeler, unpubl.). Epibryon pogonati-urnigeri on Pogonatum urnigerum is known
from Europe, North America and Asia (DoÈbbeler, 1985), and has been recorded recently
from Zaire in Africa (DoÈbbeler, unpubl.). Potriphila navicularis has a bipolar distribution,
as does its host Polytrichum alpinum (DoÈbbeler, 1996b). Bryochiton perpusillus is extremely
widespread because it seems to be constantly present on Polytrichum piliferum, which is
734 DoÈbbeler
almost cosmopolitically distributed (Long, 1985) and infects also other polytrichaceous
mosses, including Dawsonia.

The actual situation

Several examples clearly demonstrate that our knowledge of the bryophilous ascomycetes
is in an almost Linnean situation, especially outside Central Europe. Five of the six species
of Hypobryon are known exclusively from their type collections (DoÈbbeler, 1983; DoÈbbeler
and Menyivar, 1992). Four new species of Nectria on epiphyllous hepatics are known only
from Costa Rica (DoÈbbeler and Carranza, 1993). Actually, the genus Frullania is the `best'
host genus of the hepatics, with 15 species of associated ascomycetes which are nearly
exclusively known from F. dilatata and F. tamarisci, though the genus contains perhaps
over 500 species (Schuster, 1988). Ten species are known on Plagiochila, nearly always
recorded from P. asplenioides s. l., yet, Plagiochila is by far the largest genus of the
Hepaticae with hundreds of species (Schuster, 1980). The situation is similar in large
genera like Porella and Radula, in which only very few, mostly European, species have
been demonstrated as substrata for fungi.
A single collection of the epiphyllous hepatic Radula ¯accida from Africa (Tanzania,
Amani, 1911, Braun, M) yielded seven parasitic non-lichenized species, whereas up to now
only four bryophilous ascomycetes are known from the whole continent (DoÈbbeler, unpubl.).
Polytrichaceous mosses are relatively well studied, but the approximately 20 species
recorded as hosts represent only 5 % of the 354 species of the Polytrichidae (Walther,
1983).
Scho®eld (1985) indicates that there are about 18 000 species of bryophytes, 10 000
Musci and 8000 Hepaticae. They all constitute potential hosts, though by no means all of
them may harbour speci®c parasites. Until now only a very small fraction has been proved
to be a substratum for fungi. Remarkably, the number of bryophyte species is comparable
to that of the estimated world total of 17 000 to 20 000 lichens postulated to exist
(Galloway, 1992), although only around 13 500 lichenized fungi are as yet described
(Hawksworth et al., 1995). Mosses and hepatics share several important ecophysiological
characters with lichens, though they are evolutionarily unrelated (Nash and Egan, 1988).
Hawksworth (1991) points out that there may be at least 2000 species of obligately
lichenicolous fungi. I am convinced that species diversity among bryophilous fungi is at
least as high as among the lichenicolous ones.
The ®rst clear and accurate descriptions and illustrations of bryophilous ascomycetes
were published by the bryologist Johann Hedwig in 1789, introducing the generic name
Octospora for several muscicolous species (MuÈller, 1977). More than two hundred years
later we realize that within the bryophilous fungi more remains to be discovered and
described than in many other ecological groups. Bryomycology urgently needs more
attention from both bryologists and mycologists!

Acknowledgements
I wish to thank Eva Hofmann (MuÈnchen) for help with the English text. Meike
Piepenbring (TuÈbingen) gave valuable comments on an early draft of the manuscript. I am
particularly grateful to David Hawksworth (Egham) and Roy Watling (Edinburgh) for
various improvements in this paper.
Bryophilous ascomycetes 735
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