Diversity and substrate partitioning of Discomycetes in a cloud forest in Venezuela

Abstract

The fungal diversity of discomycetes was surveyed in a cloud forest at El Avila National Park in Venezuela. A systematic collecting scheme based on transects was used for the sampling methodology. Two sites were visited on two occasions during the rainy season (June to October) with a total of 24 samples collected from each site. For each plot, 24 and 27 species were identified respectively. Chao-Sorensen Similarity Coefficient between sites was 43%. Rarefaction curves based on species richness for both sites and the main types of substrates (wood, leaf, bamboo and soil) indicated that there are more species to be found than collected. Similarly, Incidence-based Coverage Estimator of species richness (ICE) used to calculate species richness showed that only 40-55% of the species present were observed during this study. Discomycete species were distributed in the following orders: Ostropales (1.2%), Lecanorales (2.4%), Pezizales (7.3%), Rhytismatales (13.4%), Orbiliales (15.9%) and Helotiales (59.8%) with a total of 42 species. This work adds ten new records of discomycetes to Venezuela and possibly eight new taxa.

Full text

Mycosphere Doi 10.5943/mycosphere/2/6/2
617
Diversity and substrate partitioning of discomycetes in a cloud forest in
Venezuela
Mardones-Hidalgo M1* and Iturriaga T2
1Departamento de Biología de Organismos, Universidad Simón Bolívar, Venezuela (melissamardones@gmail.com)
2Departamento de Biología de Organismos, Universidad Simón Bolívar, Venezuela (teresitaiturriaga@yahoo.com)
Mardones-Hidalgo M, Iturriaga T. 2011 – Diversity and substrate partitioning of Discomycetes in a
cloud forest in Venezuela. Mycosphere 2(6), 617-625, Doi 10.5943/mycosphere/2/6/2
The fungal diversity of discomycetes was surveyed in a cloud forest at El Avila National Park in
Venezuela. A systematic collecting scheme based on transects was used for the sampling
methodology. Two sites were visited on two occasions during the rainy season (June to October)
with a total of 24 samples collected from each site. For each plot, 24 and 27 species were identified
respectively. Chao-Sorensen Similarity Coefficient between sites was 43%. Rarefaction curves
based on species richness for both sites and the main types of substrates (wood, leaf, bamboo and
soil) indicated that there are more species to be found than collected. Similarly, Incidence-based
Coverage Estimator of species richness (ICE) used to calculate species richness showed that only
40–55% of the species present were observed during this study. Discomycete species were
distributed in the following orders: Ostropales (1.2%), Lecanorales (2.4%), Pezizales (7.3%),
Rhytismatales (13.4%), Orbiliales (15.9%) and Helotiales (59.8%) with a total of 42 species. This
work adds ten new records of discomycetes to Venezuela and possibly eight new taxa.
Key words – biodiversity – ecology – Helotiales – rarefaction curves – South America – species
composition
Article
Received 16 November 2011
Accepted 21 November 2011
Published online 02 December 2011
*Corresponding author: Melissa Mardones – e-mail – melissamardones@gmail.com
Introduction
Studies on fungal diversity have increa-
sed in recent years due to a growing interest in
this area of fungal research. This tendency is
necessary since the determination of species
diversity has been proposed to be one of the 25
topics to be addressed by science in the present
century (Pennisi 2005). Thus far, much of this
interest has focused on estimates of the total
number of species of fungi in the world (e.g.
Hawksworth 2001, Schmit & Mueller 2007).
According to Hawksworth’s estimate of 1.5
million species of fungi worldwide (1991), at
present only 5% of fungal taxa have been
described. Since most of the fungal diversity
studies have been carried out in temperate re-
gions of the world, fungi in neotropical forests
have been historically under-sampled. This bia-
sed approach has been evident in most of the
tropical surveys during the last two decades;
where the great majority of species represent
new records for the regions of study and about
15–30% represent new taxa (May 1996).
Discomycetes are a group of fungi cha-
racterized by the presence of ascohymenial
apothecia with either inoperculate or operculate
unitunicate asci (Kirk et al. 2008). At least five
orders can be included in this group; however
the phylogenetic relationships among these are
still under discussion (see Spatafora et al.
2006). In the traditional sense, the orders of
discomycetes are: Helotiales, Orbiliales, Ostro-
pales, Pezizales and Rhytismatales. In these
groups, the total number of non-lichenized

618
discomycetes is approximately 6650 species
(Kirk et al. 2008). According to estimations
made by Mueller and Schmit (2007), the per-
centage of known species for Helotiales and
Pezizales are 3% and 34%, respectively. From
published and unpublished fungal records com-
piled by Iturriaga and Minter (2006), there are
227 species of discomycetes in Venezuela. The
Helotiales are represented by 108 species,
followed by the Pezizales with 82 taxa. The
Ostropales, Rhytismatales and Orbiliales are
represented by 19, 11 and seven species,
respectively.
Studies in systematic biodiversity that
incorporate standardized methods are important
because they allow comparisons between
places. Among published systematic studies on
tropical microfungi (Bills & Polishook 1994,
Lodge & Cantrell 1995) only one, Cantrell
(2004), includes the discomycetes. Discomy-
cetes are important decomposers of dead
woody and herbaceous debris in tropical forests
but their substrate distributional patterns are
poorly understood. Some species are specifics
on a substrate while others may grow on seve-
ral substrates. Logs and twigs represent major
substrates for discomycetes. Leaves are inten-
sively colonized by some particular species.
Some species, mainly operculate discomycetes,
grow exclusive on soil. Bamboo, if present,
also supports species assemblages.
The main objective of the present study
was the collection of data on the biodiversity
and ecology of discomycetes from a cloud
forest in Venezuela. In order to do this, a (i)
characterization of the species assemblage, an
(ii) estimation of the species diversity and
richness and an (iii) analysis of substrates, were
also carried out.
Methods
The present study was carried out during
the rainy season of 2007 in Venezuela,
northern South America. In all instances, the
morphological concept of species was used and
nomenclature follows the 10th edition of the
Dictionary of the Fungi (Kirk et al. 2008). No
samples were isolated in laboratory conditions
and voucher specimens have been deposited in
the National Herbarium of Venezuela (VEN).
Study sites
Two sites located in a premontane cloud
forest on the south side of the El Avila Natio-
nal Park were selected. The first site is located
near Humboldt Peak (10°32'45''N, 66°52'23''W,
2000 m asl – above sea level) and will be refe-
red to in the present study as Boqueron. The
second site is located approximately 700 m east
of the first site (10°32'42''N, 66°52'01''W, 2250
m asl) and will be referred to as Lagunazo.
The structure of the forest in both sites is
characterized by more than three non-discrete
layers of vegetation. The most abundant cano-
py trees include the dominant Clusia multiflo-
ra, Podocarpus pittieri and P. oleifolius. Other
common species in the canopy are Chimarris
microcarpa, Cordia alliodora, Dictyocaryum
fuscum, Ecclinusa abbreviata, Euterpe preca-
toria, Guapira olfersiana, Inga corucans, I.
marginata, Licania cruegeiana, Ectandra kun-
thiana, Platymiscium polystachyum and Pro-
tium araguense. Three of the most common
species in the intermediate-sized layer are the
palms Catoblastus praemorsus, Ceroxylon
klopstockia and Geonoma pinnatifrons. Tree
ferns of the genus Cyathea are dominant in the
understory. Some open areas or gaps are domi-
nated by Arthrostylidium venezuelae, a species
of bamboo.
Sampling method
The sampling methodology included the
establishment of two study areas per study site.
On each of these areas, two transects were used
to sample the fungi following the method
described by Cantrell (2004). Each transect
consisted of a 60 m long imaginary line with a
series of twelve 1×1 m plots located at 5 m
intervals. These plots were randomly position-
ned on either side of the transect sides.
With this methodology, a total of 24 plots
from each study site were established and
fruiting bodies corresponding to discomycetes
were collected twice during each collecting
period on all plots. Specimens of discomycetes
were collected on three different types of dead
plant material and on the ground (abbreviated
as S for soil). For the former, wood (W),
ground litter (L) and bamboo (B) were exami-
ned. If possible, all samples were collected on
the same day and examined using a compound

Mycosphere Doi 10.5943/mycosphere/2/6/2
619
light microscope following the protocol descri-
bed by Iturriaga and Korf (1990).
Data Analysis
A number of general analyses were car-
ried out with the obtained dataset. To charac-
terize the assemblages, the number of species
per genus (S/G) was calculated from both
collecting sites and from the total data set, as
recommended by Stephenson et al. (1993). A
low value for S/G implies a higher diversity
than a high value. Similarly, an abundance-
based categorization of observations was con-
structed following the method described by
Stephenson et al. (1993) for the complete
dataset. Four categories were established and
species abundances were assigned to one of
these by means of their relative abundance. In
this way, those species regarded as “rare” were
represented by <2.0% of the total number of
collections, “occasional” between >2.0% and
<3.8% of the total, “common” between >3.8%
and <5.8% and “abundant” species were repre-
sented by >5.8% of the total. Similarly, the
incidence (presence or absence) of the species
was used as a measurement of abundance in
each plot as recommended by Schmit et al.
(1999). The Shannon-Wiener index (SI) was
obtained for both study sites as well as for the
entire assemblage of discomycetes recorded in
this study.
In addition to the latter, a comparison of
species richness values across study sites and
among the main types of substrates, was car-
ried out by constructing rarefaction-based spe-
cies accumulation curves along with their 95%
confidence interval curves using EstimateS,
version 8.2.0 (Colwell 2006). These curves
were based on the values obtained for the Chao
2 estimator, as recommended by Unterseher et
al. (2008). The same program was used for the
Incidence-based Coverage Estimator of species
richness (ICE) for both sites and all types of
substrates, to estimate the sampling efficiency
as recommended by Chao et al. (2000). The
similarity between both sites was determined
using the Chao-Sorensen Similarity Coefficient
(Chao et al. 2005).
Results
A total of 76 specimens representing 42
species of discomycetes were recorded being
Chlorencoelia torta, Hyalorbilia inflatula,
Orbilia delicatula and several species of the
genus Lachnum the most abundant taxa. Out of
the total number of species, 24 were recorded
from Boqueron and 27 from Lagunazo. The
S/G values calculated for each site and for the
total dataset ranged from 1.85 to 2 (see Table
1). The SI value calculated for the total assem-
blage of species was 3.57 (see Table 1). A list
of discomycete species and their abundance
category are given in Table 2.
Of all the species recorded, ten represent
new records for Venezuela, five are new
records for the Neotropics and eight may repre-
sent undescribed species, which for practical
purposes were assigned to the genera Dascycy-
phella (1 sp), Dimorphotricha (1 sp), Chloren-
coelia (1 sp), Lachnum (3 spp) and Strossma-
yeria (2 spp, see Table 2). Overall, the relative
abundance of collections in each order was:
Ostropales (1.%), Lecanorales (2.4%), Pezi-
zales (7.3%), Rhytismatales (13.4%), Orbiliales
(15.9%) and Helotiales (59.8%). The species-
abundance distribution (Table 2) shows that
there were few abundant species (16%) and a
high proportion of species with intermediate
abundance or rare (60%).
The ascomata of most wood-inhabiting
species were collected on logs, twigs and leaf
litter; and correspond to 71% of the total
collections. Similarly, species growing on leaf
litter represented 17% and the rest of the spe-
cies were almost equally divided between bam-
boo-inhabiting species (7%) and soil-inhabiting
species (5%).
Species accumulation curves for both
sites are shown in Fig 1, with effort measured
as the number of individuals collected on both
study sites. Although species richness estimates
for Lagunazo are higher than for Boqueron,
differences are clearly not significant because
the confidence intervals overlap.
The species accumulation curves for the
most common substrates (wood and leaf litter),
appear to indicate that wood-inhabiting species
are more diverse than leaf litter species. This
difference is not significant on the basis of the
confidence intervals (Fig 2), even when the
number of individuals is higher on wood than
on litter.
None of the species accumulation curves
(for both sites and substrates) appear to be

620
Table 1 Summary of discomycete species diversity recorded in the present study.
Parameter
Study sites
Total
Boqueron
Lagunazo
Total number of collection
43
38
81
Number of genera
13
14
21
Number of species
24
27
42
S/G value
1.85
1.93
2.00
SI value
2.93
3.22
3.57
leveling off (Figs 1 and 2). These results are
comparable with the ICE values for the maxi-
mum number of expected species in both sites
and in different substrates (Fig 3). According to
the latter estimations the studied plots contain-
ed about 40% of the species present in Boque-
ron, 55% in Lagunazo, 40% of wood-inhabi-
ting species, 75% of species on leaf litter, 75%
of species on bamboo and 50% of species
growing on soil.
The Chao-Sorensen similarity coefficient
between sites was 0.43. Six species were found
in both areas: Chlorociboria aeruginosa, Chlo-
rencoelia torta, Hyalorbilia inflatula, Lachnum
brasiliensis, Lachnum pseudoesclerotii and
Lachnum sp.1. The low number of common
species found on both sites seems to indicate
that species composition in each site is mode-
rately different.
Discussion
The study sites surveyed herein yielded a
total of 24 and 27 species of Discomycetes. In
a previous study carried out in the tropical
forests of Puerto Rico and Dominican Republic
(Cantrell 2004) using the same sampling me-
thod, the number of species in two sites per
country were 31 and 30 (Puerto Rico) and 25
and 26 (Dominican Republic). The number of
total records was higher for Puerto Rico (180
and 127 for each site). Differences in the num-
ber and frequency of sampling periods may
account for some of the discrepancies in the
number of records. While only two visits were
made to each collecting site in the present
study, Cantrell (2004) made twelve visits to
each site. Although there are differences in the
number of records, the number of species is
similar. The mean number of species per genus
for Boqueron and Lagunazo was 1.85 and 1.93,
respectively. The values obtained for the data
set from each site in Puerto Rico and Domi-
nican Republic was 2.2, 2.8, 2.1 and 1.86,
respectively. It seems clear that the values ob-
tained in the present study are lower than those
obtained in the study by Cantrell (2004).
Our results show that the four main types
of substrates support assemblages of discomy-
cetes. The most productive substrate, both in
terms of records and species present, was
decaying wood. Leaf litter occupied the second
place, followed by bamboo and soil. In general,
members of the Orbiliales and Helotiales tend
to be commonly associated with woody sub-
strates, while members of the Rhytismatales
were almost always collected from leaf litter,
specifically on leaves of Clusia multiflora.
Several authors have recognized specificity of
some species of Rhytismatales with this host in
the tropics (Sherwood 1980, Johnston 1989a,
b). Bamboo-inhabiting discomycetes form a
distinct assemblage of a few but highly specia-
lized species. Some of the taxa that have been
previously reported growing only on bamboo
include species such as Lachnum pseudoescle-
rotii and Strossmayeria jamaicensis. Interes-
tingly, all the species present on soil (Fig. 3),
belong to the order Pezizales. This is not sur-
prising, considering that this group seems to be
more abundant in temperate forests (Lodge et
al. 1995). Furthermore, members of this order
often fruit in habitats with high pH and low
content of organic matter (Hansen & Pfister
2006). According to Ataroff (2001) soils in El
Avila National Park are characterized by
having low pH and high content of organic
matter.
In both of the species accumulation
curves the asymptote is far from being reached.
This implies that there are more species to be
found. According to the ICE values, the
number of expected species for Boqueron and
Lagunazo were 62 and 49 respectively. These
values indicate that between 40–55 % of the
expected total number of species are yet to be
found. Despite the similar number of species

Mycosphere Doi 10.5943/mycosphere/2/6/2
621
Table 2 Species of discomycetes recorded in the present study and their occurrence on the studied substrates.
Species
Abundance
Substrates
Species
Abundance
Substrates
Wood
Litter
Bamboo
Soil
Wood
Litter
Bamboo
Soil
Arachnopeziza colachna b
Rare
X
Lachnum undescribed 2
Rare
X
Bisporella discedens
Rare
X
Lachnum undescribed 3
Occasional
X
c.f. arachnopeziza
Rare
X
Lophodermium platyplacum
Rare
X
Cerion leucophaeum
Rare
X
Lophodermium sp.1
Rare
X
Chlorencoelia undescribed
Rare
X
Mollisia sp.1
Rare
X
Chlorencoelia torta
Common
X
Mollisia sp.2
Rare
X
Chlorociboria aeruginosa
Common
X
Niptera c.f. stictoidea
Rare
X
Coccomyces limitatus
Occasional
X
Niptera sp.1
Rare
X
Coccomyces radiatus
Rare
X
Niptera sp.2
Rare
X
Dactylospora stygia var. striata a
Rare
X
Niptera trichophoricola b
Common
X
X
Dactylospora stygia var. tenuispora
Rare
X
Orbilia delicatula
Abundant
X
Dascycyphella undescribed b
Rare
X
Orbilia gaillardi
Occasional
X
Dimorphotricha undescribed b
Rare
X
Plectania melastomaa
Occasional
X
Hyalorbilia inflatula
Abundant
Plectania rhytidia
Rare
X
Lachnum abnorme
Rare
X
Poculum c.f. bambusae
Rare
X
Lachnum brasiliensis
Abundant
X
Pseudoplectania nigrella a
Occasional
X
Lachnum calosporum
Rare
X
Pulvinula orichalcea
Rare
X
Lachnum correae b
Common
X
Stictis radiata
Rare
X
Lachnum pseudoesclerotii b
Abundant
X
X
Strossmayeria jamaicensis a
Rare
X
Lachnum sclerotii
Occasional
X
Strossmayeria undescribed 1
Rare
X
Lachnum undescribed 1
Rare
X
Strossmayeria undescribed 2
Rare
X
aNew record for Venezuela
bNew record for the Neotropics

622
Fig. 1 – Species accumulation curves for discomycetes from the study sites in the present study.
Dashed lines indicate 95% confidence intervals.
Fig. 2 – Species accumulation curves for discomycetes associated with wood and leaf litter in the
present study. Species richness values (solid lines) were calculated using the Chao 2 estimator with
95% confidence intervals (dash lines).
found on both sites, the structure of the data
suggests that the Boqueron site has higher
species richness. The ICE values for the maxi-
mum number of species indicate that the sur-
vey was more complete for woody substrates
than for leaf litter, bamboo and soil. These
estimates however, may represent underestima-
tions simply because all of the different sub-
strates were not sampled with equal intensity.
The ICE values for expected species
conform with the results obtained in the curve
analyses. Both results suggest that a greater
sampling effort is required to find the remain-
ing species. In both cases, the nonparametric
estimators magnified the differences in species
richness.
The Chao-Sorensen Similarity Coefficient
between sites was 0.43. This value suggests
low similarity between the assemblages of
discomycetes at both study sites, which may
indicate that both sites were established at
appropriate distances as to represent indepen-
dent samples of the same community (Lodge &
Cantrell 1995). This value may also be an
indicative of the high diversity of species in El
Avila National Park, since only six species of

Mycosphere Doi 10.5943/mycosphere/2/6/2
623
Fig. 3 – Observed versus expected species richness values for both sites and all type of substrates,
using the Incidence-based Coverage Estimator (ICE). Numbers over columns indicate the actual
value.
the total of 45, were shared between sites. Si-
milar results were obtained by Cantrell (2004)
for Puerto Rico and Dominican Republic (0.5
and 0.40 respectively).
Records of new taxa – The information
on the diversity of discomycetes in tropical
ecosystems is scarce. The present study repre-
sents a useful set of data to further understand
the diversity and biogeography of discomy-
cetes in tropical areas. Our study has increased
the number of species known from Venezuela
to 244. Two genera are recorded for the first
time for the Neotropics: Dascycyphella sp. and
Dimorphotricha sp, as well as three species:
Arachnopeziza colachna, Lachnum pseudoes-
clerotii and Lachnum correae. Dascycyphella
occurs predominantly in temperate areas, and
none of its 22 described species have been
reported for tropical areas (Raitviir 2002).
Dimorphotricha is a monotypic genus reported
only for Australia (Spooner 1987) and this is
the second report in the world for this genus.
Similarly Lachnum correae has also been
reported for Australia (Spooner 1987). Twenty-
eight percent of the total number of species
collected represent new reports for Venezuela.
This is an interesting fact since the study of
Discomycetes in this country is relatively
ahead of the situation in other countries of the
region.
In summary, the data obtained in the
present study indicate that the assemblages of
discomycetes present in the study sites conform
to similar patterns reported for other areas in
the Neotropics. Although more research is
needed to determine a good estimate of the true
number of discomycetes species, we conclude
that this study adds to the small amount of data
previously known of discomycetes diversity in
the Neotropics. However, the lack of fungal
studies with the group and the disparities in
methodology difficult comparisons among stu-
dies and large-scale patterns of tropical fungal
diversity remain difficult to make reliable
conclusions. We envision that more research,
based on consistent methods, would help deter-
mine the degree of gamma diversity present in
tropical areas. At the same time, studies as-
sessing substrates preference help elucidate
ecological patterns of these fungi. At present,
we continue collecting data in the study sites
described in the present study with the goal of
obtaining more accurate information to esti-
mate the number of species present in these
sites over a longer period of time.
Acknowledgements
We thank to the Decanato de Estudios de
Postgrado of Simón Bolívar University for
financial support. We also thank Dr. Carlos
Rojas and MSc. Alejandro Muñoz for a critical
and linguistic review of the manuscript and
additional statistical support. We would like to
acknowledge the support of the El Avila
National Park and INPARQUES for facilities
during the work fields and to Jesús Hernández,

624
Roberto Fernández and Carlos Rondón for field
assistance. M. Mardones would like to acknow-
ledge to Ministerio de Ciencia y Tecnología
from Costa Rica, for the financial support du-
ring her Master Degree in Venezuela, which
led to this paper. T. Iturriaga thanks FONACIT
(Fondo Nacional de Ciencia, Tecnología e In-
novacion) Caracas-Venezuela for funding pro-
ject NºS1-2001000663: “Sistemática Molecular
y Morfológica de Hongos Ascomycetes y Basi-
diomycetes presentes en las Sucesiones Fúngi-
cas Descomponedoras de Madera en Bosques
Tropicales” under the direction of TI which
helped with partially funding this research
References
Ataroff M. 2001 – Venezuela. In: Bosque Un-
blados del Neotrópico (eds M Kappelle,
AD Brown). INBio. Costa Rica 397–442.
Bills GF, Polishook JD. 1994 – Abundance and
diversity of microfungi in leaf litter of a
lowland rain forest in Costa Rica. Myco-
logia 86, 187–198.
Cantrell S. 2004 – A Comparison of two sam-
pling strategies to assess Discomycete
diversity in the wet tropical forest. Carib-
bean Journal of Science 40, 8–16.
Chao A, Chazdon RL, Colwell RK, Shen TJ.
2005 – A new statistical approach for as-
sessing compositional similarity based on
incidence and abundance data. Ecology
Letters 8, 148–159.
Chao A, Hwang WH, Chen YC, Kuo CY. 2000
– Estimating the number of shared spe-
cies in two communities. Statistica Sinica
10, 227–246.
Colwell RK. 2006 – Estimates: Statistical esti-
mation of species richness and shared
species from samples. Version 8.0. User's
Guide and application published at:
http://purl.oclc.org/estimates
Hansen K, Pfister DH. 2006 – Systematics of
the Pezizomycetes–the operculate disco-
mycetes. Mycologia 98, 1029–1040.
Hawksworth DL. 2001 – The Magnitude of
fungal diversity: the 1.5 million species
estimate revisited. Mycological Research
105, 1422–1432.
Iturriaga T, Korf RP. 1990 – A monograph of
the discomycete genus Strossmayeria
(Leotiaceae), with comments on it’s ana-
morph Pseudospiropes (Dematiaceae).
Mycotaxon 36, 383–454.
Iturriaga T, Minter D. 2006 – Hongos de Vene-
zuela. www.cybertruffle.org.uk/venefung
/esp.
Johnston PR. 1989a – Lophodermium (Rhytis-
mataceae) on Clusia. Sydowia 41, 170–
179.
Johnston PR. 1989b – Rhytismataceae in New
Zealand 2. The genus Lophodermium on
indigenous plants. New Zealand Journal
of Botany 27, 243–274.
Kirk PM, Cannon PF, Minter DW, Stalpers JA.
2008 – Dictionary of the Fungi. 10th Edi-
tion. Cromwell Press. UK.
Lodge DJ, Cantrell SA. 1995 – Diversity of
litter agarics at Cuyabeno, Ecuador: cali-
brating sampling efforts in tropical rain-
forest. The Mycologist 9, 149–151.
Lodge DJ, Chapela I, Samuels G, Uecker FA,
Desjardin D, Horak E, Miller OK,
Hennebert GL, Decock CA, Ammirati J,
Burdsall HH, Kirk PM, Minter DW,
Hailing R, Laessøe T, Mueller G,
Huhndorf S, Oberwinkler F, Pegler DN,
Spooner B, Petersen RH, Rogers JD,
Ryvarden L, Watling R, Turnbull E,
Whalley AJS. 1995 – A Survey of
Patterns of Diversity in Non-Lichenized
Fungi. Mitt. Eidgenöss. Forsch. anst.
Wald Schnee Landsch 70, 157–173.
May RM. 1996 – Conceptual aspects of the
quantification of the extend of biological
diversity. In: Biodiversity, Measurement
and estimation (ed DL Hawksworth).
Chapman & Hall, London 101–118.
Mueller GM, SchmiT JP. 2007 – Fungal biodi-
versity: what do we know? What can we
predict. Biodiversity and Consevation 16,
1–5.
Pennisi E. 2005 – What determines species
diversity? Science 309, 90.
Raitviir A. 2002 – A Revision of the Genus
Dasiscyphella (Hyaloscyphaceae, Helo-
tiales). Polish Botanical Journal 47, 227–
241.
Schmit JP, Mueller GM. 2007 – An estimate of
the lower limit of global fungal diversity.
Biodiversity and Conservation 16, 99–
111.
Schmit JP, Murphy JF, Mueller GM. 1999 –
Macrofungal diversity of a temperate oak

Mycosphere Doi 10.5943/mycosphere/2/6/2
625
forest: a test of species richness estima-
tors. Canadian Journal of Botany 77,
1014–1027.
Sherwood MA. 1980 – Taxonomic studies in
the Phacidiales: The genus Coccomyces
(Rhytismataceae). Occasional Papers of
the Farlow Herbarium of Cryptogamic
Botany 15, 1–120.
Spatafora JW, Sung GH, Johnson D, Hesse C,
O’Rourke B, Serdani M, Spotts R,
Lutzoni F, Hofstetter V, Miadlikowska J,
Reeb V, Gueidan C, Fraker E, Lumbsch
T, Lücking R, Schmitt I, Hosaka K,
Aptroot A, Roux C, Miller AN, Geiser
DM, Hafellner J, Hestmark G, Arnold
AE, Büdel B, Rauhut A, Hewitt D,
Untereiner WA, Cole MS, Scheidegger
C, Schultz M, Sipman H, Schoch CL.
2006 – A five-gene phylogeny of Pezizo-
mycotina. Mycologia 98, 1018–1028.
Spooner BM. 1987 – Helotiales of Australasia:
Geoglossaceae, Orbiliaceae, Sclerotinia-
ceae, Hyaloscyphaceae. Bibliotheca My-
cologica Band 116, 1–711.
Stephenson SL, Kalyanasundaram, I, Lakhan-
pal T. 1993 – A comparative biogeogra-
phical study of Myxomycetes in the mid-
Appalachians of eastern North America
and two regions of India. Journal of Bio-
geography 20, 645–657.
Unterseher M, Schnittler M, Dormann C,
Sickert A. 2008 – Application of species
richness estimators for the assessmentof
fungal diversity FEMS Microbiological
Letters 282, 205–213.