Latitudinal, habitat and substrate distribution patterns of freshwater ascomycetes in the Florida Peninsula

Abstract

Freshwater ascomycetes are important decomposers of dead woody and herbaceous debris in aquatic habitats. Despite evidence of their ecological importance, latitudinal, habitat and substrate distributional patterns of freshwater ascomycetes are poorly understood. In this study, we examined the latitudinal and habitat distributional patterns, and substrate recurrences of freshwater ascomycetes by collecting dead submerged woody and herbaceous debris in lentic and lotic habitats at five selected sites along
a north-central-south, temperate–subtropical latitudinal ecotone in Florida. One hundred and thirty-two fungal taxa were collected during the study. Seventy-four were meiosporic and 56 were mitosporic ascomycetes, while two species were basidiomycetes. Canonical analyses of principal coordinates (CAP) and Sørenson’s similarity index of species based on presence/absence data revealed a high turnover in species composition between the northern and southern sites, indicating a change in species composition along the temperate–subtropical latitudinal ecotone of the Florida Peninsula. Results from the ordination analysis indicated that freshwater ascomycete community composition is not significantly different between lentic and lotic habitats in Florida. The geographically broadly distributed species and species commonly found in Florida occurred in both habitats, whereas a number of new or rare species occurred in either lentic or lotic habitats, but not both. The same freshwater ascomycete species
did not necessarily occur on both woody and herbaceous debris; of the 132 taxa collected, 100 were reported only on woody debris; 14 species occurred exclusively on herbaceous debris; and 18 species were found on both woody and herbaceous debris in lentic or lotic habitats. Implications of data from this study to the conservation and knowledge of biodiversity for freshwater ascomycetes is discussed.

Full text

ORIGINAL PAPER
Latitudinal, habitat and substrate distribution patterns
of freshwater ascomycetes in the Florida Peninsula
Huzefa A. Raja Æ John Paul Schmit Æ Carol A. Shearer
Received: 23 April 2008 / Accepted: 26 September 2008 / Published online: 18 October 2008
Ó Springer Science+Business Media B.V. 2008
Abstract Freshwater ascomycetes are important decomposers of dead woody and her-
baceous debris in aquatic habitats. Despite evidence of their ecological importance,
latitudinal, habitat and substrate distributional patterns of freshwater ascomycetes are
poorly understood. In this study, we examined the latitudinal and habitat distributional
patterns, and substrate recurrences of freshwater ascomycetes by collecting dead sub-
merged woody and herbaceous debris in lentic and lotic habitats at five selected sites along
a north-central-south, temperate–subtropical latitudinal ecotone in Florida. One hundred
and thirty-two fungal taxa were collected during the study. Seventy-four were meiosporic
and 56 were mitosporic ascomycetes, while two species were basidiomycetes. Canonical
analyses of principal coordinates (CAP) and Sørenson’s similarity index of species based
on presence/absence data revealed a high turnover in species composition between the
northern and southern sites, indicating a change in species composition along the tem-
perate–subtropical latitudinal ecotone of the Florida Peninsula. Results from the ordination
analysis indicated that freshwater ascomycete community composition is not significantly
different between lentic and lotic habitats in Florida. The geographically broadly distrib-
uted species and species commonly found in Florida occurred in both habitats, whereas a
number of new or rare species occurred in either lentic or lotic habitats, but not both. The
same freshwater ascomycete species did not necessarily occur on both woody and her-
baceous debris; of the 132 taxa collected, 100 were reported only on woody debris; 14
species occurred exclusively on herbaceous debris; and 18 species were found on both
woody and herbaceous debris in lentic or lotic habitats. Implications of data from this study
to the conservation and knowledge of biodiversity for freshwater ascomycetes is discussed.
Keywords Aquatic habitats  Biogeography  Freshwater fungi
H. A. Raja (&)  C. A. Shearer
Department of Plant Biology, University of Illinois at Urbana – Champaign, 265 Morrill Hall,
505 South Goodwin Ave., Urbana, IL 61801, USA
e-mail: raja@illinois.edu
J. P. Schmit
Smithsonian Institution, Smithsonian Environmental Research Center, 647 Countess Wharf Rd.,
P.O. Box 28, Edgewater, MD 21037, USA
123
Biodivers Conserv (2009) 18:419–455
DOI 10.1007/s10531-008-9500-7

Introduction
Freshwater ascomycetes are an ecological assemblage of fungi that occur on submerged or
partially submerged substrates in aquatic habitats (Shearer 1993, 2001; Vijaykrishna et al.
2006). Beginning with the pioneering studies by Ingold (1951, 1954, 1955, 1959, 1961,
1966, 1968) and Ingold and Chapman (1952), this group of fungi has been studied only in
the last 50 years (Dudka 1963, 1985; Shearer 1993; Hyde et al. 1997; Shearer 2001; Goh
and Hyde 1996; Tsui and Hyde 2003; Cai et al. 2006a, b; Vijaykrishna and Hyde 2006;
Shearer et al. 2007). Most other groups of organisms, including other fungal groups, have a
much longer taxonomic and distributional history. Since the pioneering studies by
C.T. Ingold, 123 new genera and about 261 species of freshwater ascomycetes have been
described and currently, about 548 ascomycete species have been reported from freshwater
habitats located mostly in Europe, North America, and South East Asia (for additional
literature see http://www.life.uiuc.edu/fungi/). Although these studies have revealed the
existence of a distinct freshwater ascomycota, there is a lack of knowledge about the
geographical and habitat distribution patterns and substrate specificities for individual
species (Shearer 1993, 2001; Cai et al. 2003, 2006a, b; Shearer et al. 2007).
Although latitude is known to influence the geographical distribution patterns of plants
and animals (Rosenzweig 1995), similar information for most fungi (Arnolds 2007) and
specifically the freshwater ascomycetes is lacking (Shearer 1993, 2001). In contrast, the
marine ascomycetes are relatively well-studied with respect to their biogeography (Hughes
1974; Kohlmeyer and Kohlmeyer 1979; Hyde and Lee 1995; Kis-Papo 2005). Hughes
(1974) compiled geographical information on higher marine fungi using range maps of
species’ distributions. He concluded that the most important environmental parameter in
their latitudinal distribution is temperature. Booth and Kenkel (1986) used ordination
methods and suggested that both temperature and salinity play an important role in the
distribution of lignicolous marine fungi.
The geographical occurrences of Ingoldian mitosporic ascomycetes, viz (=anamorph or
asexual fungi) are relatively well-studied compared to those of the freshwater meiosporic
ascomycetes. The Ingoldian mitosporic fungi most commonly occur on autumn shed leaves
in streams and rivers (Webster and Descals 1981;Ba
¨
rlocher 1992
) and a subset of these
fungi are also known to occur on wood (Willoughby and Archer 1973; Sanders and
Anderson 1979;Re
´
vay and Go
¨
nczo
¨
l 1990; Shearer and Webster 1991; Shearer 1992). These
fungi have conidia that are mostly branched, tetraradiate or long, narrow and sigmoidal;
morphologies considered to be adapted for life in flowing water (Ingold 1953, 1954, 1966,
1975, Read 1990; Goh and Hyde 1996; Wong et al. 1998). Wood-Eggenschwiler and
Ba
¨
rlocher (1985) used distribution data for over 150 species of Ingoldian mitosporic
ascomycetes obtained from the literature (Webster and Descals 1981) and computed
Sørenson’s similarity index based on species presence or absence data from various geo-
graphical locations. They found that there was a higher similarity in species composition of
Ingoldian fungi between geographically distinct tropical locations (South America, West
Africa) than between tropical and temperate regions that were located on the same conti-
nent, either African or North and South American. From their study, Wood-Eggenschwiler
and Ba
¨
rlocher (1985) concluded, ‘‘on a worldwide scale, temperature together with its
influence on vegetation in different climatic regions is the major factor in determining
distribution patterns of Ingoldian mitosporic fungi’’. Since a number of freshwater Ingoldian
mitosporic fungi are asexual states of freshwater ascomycetes (Shearer 1993, 2001; Sivichai
and Jones 2003; http://www.life.uiuc.edu/fungi/), one might expect distribution patterns of
freshwater ascomycetes to be similar to those of Ingoldian mitosporic fungi.
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The geographical distribution patterns of freshwater ascomycetes, at present, are linked
largely to the geographical distribution of the few mycologists who study this ecological
group and the places they have sampled (Shearer 1993; Hyde et al. 1997; Sivichai 1999;
Shearer 2001; Vijaykrishna et al. 2006; Shearer et al. 2007). Some freshwater ascomycetes
are cosmopolitan in distribution, while others are reported only from their type localities
(Shearer 1993, 2001; Cai et al. 2006a, b; http://www.life.uiuc.edu/fungi/). About 65% of
the ascomycetes reported from freshwater habitats have been reported only once (Shearer
et al. 2007). The fact that they are reported only once cannot be interpreted as being absent
from other geographical areas, unless those areas have been intensively sampled. Ho et al.
(2001) compared lignicolous freshwater ascomycete communities in lentic (e.g., lakes,
ponds, and swamps) and lotic (e.g., streams and rivers) habitats in Australia, Brunei, Hong
Kong, Malaysia, Seychelles, South Africa, and the UK. The authors used multivariate
analysis to visualize the fungal community and found that there were distinct fungal
communities between the temperate site (UK), subtropical sites (Hong Kong, South
Africa), and tropical sites (Australia, Brunei, Malaysia, and Seychelles). These compari-
sons were made from a single collection site each in Australia, Brunei, Malaysia,
Seychelles, and the UK and from six collection sites in Hong Kong. Since the comparisons
did not differentiate between lentic and lotic habitats in their analyses, the differences in
species composition might be due in part to the effect of habitat type as well as geography.
In addition, the study by Ho et al. (2001) was a literature study using published data from
studies that were conducted at different times by different investigators.
Due to the lack of comprehensive, directly comparable studies from different geo-
graphical locations, little can be said about the broad geographical distribution patterns of
freshwater ascomycetes. At present, there are no large or medium spatial scale freshwater
mycogeographic studies in a single geographical area that have examined the effects of
latitudinal change to determine how this factor influences geographical distribution pat-
terns and community composition of freshwater ascomycetes. Understanding changes in
species composition along environmental gradients such as latitude can reveal important
information regarding species distribution patterns.
To better understand the distribution of organisms, it is also important to recognize and
characterize their associations with different habitat types. Variation in habitat type is often
known to affect the movement, settlement and community composition of plants and
animals (Morin 1999), but the influence of habitat type on freshwater ascomycete com-
munity composition is unexplored and poorly understood. A number of studies on
freshwater ascomycetes have been conducted at single, spatially limited sites that were
either lentic (Shearer and Crane 1986; Hyde and Goh 1998a; Goh and Hyde 1999; Cai
et al. 2002; Luo et al. 2004; see Wong et al. 1998) or lotic habitats (Lamore and Goos
1978; Shearer and Von Bodman 1983; Hyde and Goh 1997; Hyde et al. 1998; Hyde and
Goh 1998b; Hyde and Goh 1999; Ho et al. 2002; Sivichai et al. 2000, 2002; Tsui et al.
2001, 2003; Tsui and Hyde 2004; Kane et al. 2002; Cai et al. 2003; Fryar et al. 2004;
Gonz
´
alez and Chavarr
´
ia 2005; Vijaykrishna et al. 2006; see Wong et al. 1998). However,
thus far, no studies have examined distribution of freshwater ascomycetes simultaneously
in both lotic and lentic habitats to compare species composition between habitat types.
Most collections of freshwater ascomycetes have been made from submerged woody
debris (Dudka 1963; Shearer 1972; Willoughby and Archer 1973; Lamore and Goos 1978;
Minoura and Muroi 1978; Shearer and von Bodman 1983; Ho et al. 2001; Kane et al. 2002;
Tsui et al. 2000, 2001; Raja et al. 2003, 2005; Raja and Shearer 2008; Vijaykrishna and
Hyde 2006; also see http://fungi.life.uiuc.edu/). In addition to woody debris, freshwater
ascomycetes have also been collected from a variety of emergent aquatic macrophytes,
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123

such as Carex, Equisetum, Juncus, Phragmites, Scirpus, and Typha, which are common in
lentic habitats, and backwaters of rivers, bogs and swamps (Shearer 1993; Dudka 1985;
Magnes and Hafellner 1991; Fallah and Shearer 2001; Van Ryckegem and Verbeken
2005). Of the 548 species of freshwater ascomycetes reported thus far (http://fungi.life.
uiuc.edu/), 60% are reported only from submerged woody debris and about 30% are
reported only from herbaceous substrates, while only about 9–10% species are reported
from both submerged wood and herbaceous substrates. This suggests that some degree of
substrate specialization occurs. However, few investigators have collected both submerged
woody and herbaceous debris at the same time to compare species composition on different
substrates. Fallah (1999) collected freshwater ascomycetes from submerged woody and
herbaceous debris from seven north temperate LTER (Long Term Ecological Research)
lakes in Wisconsin. He found that Sordariomycetes occurred on wood and Dothideomy-
cetes were more prevalent on herbaceous debris. He also found that species distributions in
the LTER lakes were related to the distribution of the macrophyte species on which they
occurred. Species found on submerged wood were not found on herbaceous debris and vice
versa. Cai et al. (2003) collected both submerged wood and dead decaying bamboo in a
river in the Philippines and found that the community composition of meiosporic and
mitosporic ascomycetes differed between the two substrates. Of the 80 species reported in
their study, only 16 taxa occurred on both submerged wood and bamboo, and fewer taxa
occurred on bamboo than on wood. A high similarity in species composition was found
when Luo et al. (2004) compared occurrences of freshwater fungi on three different
monocotyledon herbaceous substrates. However, Luo’s study revealed differences in
species relative abundance between the fungal communities on different herbaceous sub-
strates. These foregoing studies suggest that substrate type is important in the distribution
of freshwater ascomycetes, but additional comparative studies from different geographical
areas are needed to better understand the role of substrate specificity in the geographical
distribution patterns of freshwater ascomycetes.
Without adequate knowledge about the environmental factors that determine species
distribution patterns, accurately assessing the diversity of freshwater ascomycetes is not
possible. Efforts to develop rational plans to conserve this group of important saprobic
fungi are hindered by this lack of information. To obtain baseline comparative distribution
data, the following study was undertaken.
The objective of this study was to address the following questions about the distribution
patterns of freshwater ascomycetes: (1) Does the species composition of freshwater
ascomycete communities differ among sites at different latitudes along the temperate–
subtropical ecotone of the Florida Peninsula? (2) Does the species composition of fresh-
water ascomycete communities differ between lentic and lotic habitats in Florida? (3) Do
the same freshwater ascomycete species occur on both woody and herbaceous debris?
Methods
Description of study sites
The Florida Peninsula was selected as the location for this study because of its large
number of freshwater lentic and lotic habitats, including 7,800 lakes and 1,700 rivers and
streams (Whitney et al. 2004). In addition, the Florida Peninsula is a biotic transition zone
between the warm temperate and subtropical zones based on climatic and biotic data
(Henry et al. 1948). Because the peninsula is long, climatic conditions vary somewhat
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north to south. Florida lies within the temperate zone but the climate is subtropical in south
Florida (Henry et al. 1948), providing a unique region to investigate hypotheses about
freshwater fungal diversity and distribution. In addition, Florida is a known area of high
biodiversity for other organisms (Whitney et al. 2004). Prior to this study, however, only
three species of freshwater ascomycetes had been reported from Florida (Conway and Barr
1977; Shearer and Crane 1995; Anderson and Shearer 2002).
Five collection sites were chosen in north, central and south Florida based on a lati-
tudinal ecotone along the Florida Peninsula (Fig. 1): (1) Blackwater River State Forest
(BW) (30°N latitude), (2) Apalachicola National Forest (AP) (30°N latitude), (3) Ocala
National Forest (OC) (29°N latitude), (4) Big Cypress National Preserve (BC) (25°N
latitude), and (5) Everglades National Park (EV) (25°N latitude).
According to Platt and Schwartz (1990), the panhandle forests (BW and AP) are
southern hardwood forests, the central peninsula forest (OC) is a temperate broad-leafed
evergreen forest, and south Florida forests (BC and EV) are subtropical forests. Tropical
hardwood hammocks are seen predominately in the BC and EV. These five sites were
chosen because (a) they are located across a temperate and subtropical ecotone along the
Florida Peninsula (Question 1), and (b) they are rich in a variety of lentic and lotic habitats,
so collections can be made from both habitat types and woody and herbaceous debris can
be collected within the same area in each site (Questions 2, 3). In addition, for the region
where the study sites are located, Beaver et al. (1981) found that the lentic habitats differ
on the basis of temperature and categorized these habitats by their thermal properties into
north (warm temperate), central (transitional) and south (subtropical).
Sampling procedure
Submerged, dead woody and herbaceous debris was collected randomly from lentic and
lotic habitats, four times over 2 years. Sampling times included two summer (wet season)
and two winter (dry season) collections at the five sites within the Florida Peninsula from
2004 to 2006. Both woody and herbaceous debris (dead, decaying emergent macrophytes)
Fig. 1 Study area. The solid dots indicate the locations of the five collection sites within the Florida
Peninsula; scale indicates latitudes where sites were located
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were collected. Equal amounts of debris were collected (about 15 pieces of each depending
on availability) from each site. In lotic habitats, such as a stream or river, collections were
made along an imaginary transect 100 meters upstream and downstream at a collection
site. In lentic habitats, collections were made from around the littoral zone from depths of
about 0.5–1.0 m. Efforts were made to identify and collect substrates that had been sub-
merged in water for a considerable time. This was estimated by observation of the degree
of softening by fungal soft-rot and colonization by other aquatic organisms. Samples were
placed in zippered plastic bags containing moist paper towels and transported to the lab in
an insulated cooler containing ice to reduce heat build-up and biological activity. Water
temperature was recorded in the field with a thermometer; pH was measured with an IQ125
miniLab pH meter and Fisher pH strips; and latitude and longitude were measured using a
Garmin global positioning system (GPS). In the lab, substrates were gently rinsed with tap
water and incubated in plastic storage boxes with moistened paper towels at ambient
temperatures (about 24°C) under 12/12 h (light/dark) conditions.
Morphological observations
Samples were examined with a dissecting microscope within one to two weeks of col-
lection and periodically over 6–12 months (Shearer 1993; Shearer et al. 2004). After
sporulating fungi were located on natural substrates or isolation media using the dissecting
microscope, ascomata were opened in a drop of distilled water on a large (25 mm) cover
slip on a slide using fine dissecting needles, and then covered with a second smaller
(18 mm) cover slip. Ascomycetes were identified based on the morphology and anatomy of
the ascomata and the morphology of the hamathecium, asci, and ascospores. Melzer’s
reagent (0.5 g iodine, 1.5 g KI, 20 g chloral hydrate, 20 ml distilled water), and aqueous
cotton blue was used to determine staining reactions of the ascus apical apparatus. India
ink or aqueous nigrosin was added to water mounts to reveal gelatinous sheaths on or
around ascospores as well as gelatinous material surrounding the paraphyses or pseud-
oparaphyses. Mitosporic ascomycetes were identified using the type of conidiogenesis and
conidial morphology. Measurements were made of material mounted in distilled water,
glycerin (100%) or lactic acid containing azure A. Material mounted on slides was pre-
served with glycerin (100%) or lactic acid (85%) using the double cover glass method
(Volkmann-Kohlmeyer and Kohlmeyer 1996). Whenever, possible freshwater ascomycetes
were isolated in pure culture using procedures outlined in Fallah and Shearer (2001) and
Shearer et al. (2004). Slides and specimens of fungi collected during the study are
deposited in the Herbarium of the University of Illinois (ILL).
Data preparation
A site by species matrix based on presence or absence data was constructed. Within this
matrix, sites (aquatic habitats) were the column variables and species were the row variables.
Each cell contained ‘‘1’’ if a particular species was present at that site, and ‘‘0’’ if it was not.
The matrix contained records of all the occurrences of 132 fungal species in 97 sites.
Multivariate statistical analysis
Data were analyzed using canonical analysis of principal coordinates (CAP; Anderson and
Willis 2003). CAP is a multivariate constrained ordination technique in which an initial
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principal coordinates analysis (PCO) is used to reduce the number of axes in the analysis.
A further analysis is then conducted on the matrix produced by the PCO. A canonical
discriminant functional analysis (=discriminant analysis) was used in this study for
questions 1 and 2 to test if the data matrix is structured according to the grouping variables,
representing either latitude (N–C-S) (Question 1) or habitat (lentic vs. lotic) (Question 2) as
detailed below. One of the benefits of using CAP is that it uses permutation tests (trace
statistic, 1st squared canonical correlation) to assign a P-value to the a-priori hypothesis by
determining the probability that the grouping found in the final analysis could arise by
chance alone. The CAP method, therefore, allows the objective evaluation of a hypothesis.
In addition, a Canonical Correlation Analysis (CCA) was employed to test if variation in
community composition is explained by water pH and/or latitude. The CCA analysis was
part of the CAP ordination. Separate analyses were conducted for pH and latitude, fol-
lowed by an analysis of the combined pH and latitude data. For the CCA analysis, pH
values from different collection dates at a single site were transformed to log scale,
averaged and then transformed to antilog and used in the CCA analysis. Canonical analysis
of principal coordinates has been used recently for analyzing the distribution patterns of
aquatic mangrove fungi (Schmit and Shearer 2004) and Ingoldian mitosporic ascomycetes
(Nikolcheva and Ba
¨
rlocher 2005).
All analyses were performed using CAP software, freely available at the Ecological
Society of America’s Ecological Archives (E084-011-S1; http://esapubs.org/archive/ecol/
E084/011/suppl-1.htm). All analyses were performed using ‘‘Gower excluding double
zero’’ (Gower 1971; Lagendre and Legendre 1998) as the distance measure.
Latitudinal distribution
The CAP analysis was used to test the hypothesis that there are differences in the fresh-
water ascomycete fungal communities between sites at different latitudes (north, central
and south) in Florida. A significant result indicates fungal communities differ along the
latitudinal gradient sampled.
In addition to the CAP analysis, fungal similarity among different collections sites
(N-C-S) was calculated using Sørenson’s similarity index (Sørenson 1948; Magurran 1988,
2004).
Habitat distribution
The CAP was used to test the hypothesis that there are consistent differences in freshwater
ascomycete species between lentic and lotic habitats. To test this hypothesis, lentic and
lotic collection sites from north, central, and south Florida were analyzed with CAP. A
significant result indicates that fungal communities differ between habitat types.
Two CAP analyses were performed. The first analysis contained species presence or
absence data from all the 97 habitats sampled along the Florida Peninsula, with 26 lotic
and 71 lentic habitats. Since the results of the CAP analyses containing an unequal
number of habitats could bias the result of the ordination graph, a second analysis was
performed in CAP using habitat data from BW and AP (northern collection sites)
because approximately equal number of habitat types were sampled in these sites. In the
second CAP analyses, presence or absence data of species from 21 lotic and 27 lentic
habitats were included.
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Results
A total of 97 habitats were sampled at the five collection sites in Florida. Appendix 1
shows the latitudes and longitudes, water temperatures and pH measured during the study.
Maximum and minimum ranges of water temperature and pH values from the five col-
lection sites are listed in Appendix 2. In both years of collection, the lowest water
temperatures were recorded in winter at the northern sites, and the highest water tem-
peratures were recorded for the southern sites. The water temperatures recorded at the
central site were intermediate between those of the northern and southern sites. However,
in summer, the water temperature differential along the latitudinal ecotone disappeared,
probably due to the tropical circulation pattern of the Caribbean that influences the climate
of the entire Florida Peninsula (Beaver et al. 1981), and the water temperature along the
entire peninsula becomes generally warmer with an average of about 30°C.
One hundred and thirty-two species of fungi were collected during the study (Table 1).
Seventy-four are meiosporic and 56 are mitosporic ascomycetes, while two species,
Rogersiomyces okefenokeensis and Ingoldiella hamata, are basidiomycetes. Of the 74
meiosporic ascomycetes, 28 are Dothideomycetes, 44 are Sordariomycetes, and two taxa
belong to the Leotiomycetes. Of the 132 taxa collected, 12 species and two genera were
new to science (Raja and Shearer 2006a, b, 2007; Raja et al. 2008, in press; Raja and
Shearer 2008), 26 taxa are new records for North America and 127 taxa are new records
from aquatic habitats in Florida (Table 1). Ayria appendiculata, Falciformispora lignatilis,
and Trematosphaeria lineolatispora are new records for fresh water.
Among the mitosporic ascomycetes collected, six species, Brachiosphaera tropicalis,
Condylospora sp., Dendrospora sp., Ingoldiella hamata, Nawawia filiformis, and Vari-
cosporium sp., are Ingoldian mitosporic ascomycetes; three species, Cancellidium
applanatum, Helicodendron sp., and Helicoon auratum are aeroaquatic fungi, and the
remaining 49 species belong to the miscellaneous mitosporic fungi (Shearer et al. 2007).
All of the mitosporic fungi found belong to the ascomycetes, except I. hamata, which is a
basidiomycete (Shaw 1972).
Distribution patterns along the latitudinal ecotone
Analysis using CAP revealed latitudinal structuring of freshwater fungal communities
along the Florida Peninsula (Fig. 2). North, central and south sites in Florida formed three
clusters on the ordination graph. The northern and central sites are slightly closer to one
another than either is to the southern cluster. The analysis found two canonical axes.
Permutation tests provided statistical support for the hypothesis of latitudinal structuring of
freshwater fungal communities (trace statistic = 1.341, P \ 0.01; 1st squared canonical
correlation = 0.72, P \0.01).
Sørenson’s coefficient of similarity
Species similarity was greatest between the two northern sites (BW and AP 42%) and
between the two southern sites (BC and EV 34%) (Table 2). The northern sites BW and AP
also shared some similarity with the central site OC (31 and 32%, respectively). The least
similarity (13%) occurred between the northernmost site (BW) and the two southern sites
(BC and EV).
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Table 1 Total numbers of fungal species and the number of times they were collected at each of the five
sites along the Florida Peninsula
Species Collection sites
BW AP OC BC EV
Dothideomycetes
Aliquandostipite minuta Raja & Shearer
a
1
Aliquandostipite crystallinus Raja, Ferrer & Shearer
b
1
Aliquandostipite khaoyaiensis Inderb.
c
1
Aliquandostipite siamensiae (Sivichai & E.B.G. Jones)
J. Campbell, Raja, Ferrer, Sivichai & Shearer
c
1
Boerlagiomyces websteri Shearer & J.L. Crane 2 1 6 2
Caryospora obclavata Raja & Shearer
a
1
Caryospora langloisii Ell. & Everh.
b
1
Falciformispora lignatilis K.D. Hyde
b
11
Jahnula aquatica (Plo
¨
ttner & Kirschst.) Kirschst.
b
1
Jahnula australiensis K.D. Hyde
c
1
Jahnula bipileata Raja & Shearer
a
11
Jahnula bipolaris (K.D. Hyde) K.D. Hyde
c
1
Jahnula potamophila K.D. Hyde & S.W. Wong
c
1
Jahnula rostrata Raja & Shearer
a
1211
Jahnula sangamonensis Shearer & Raja
b
11
Kirschsteiniothelia elaterascus Shearer
b
1111
Lepidopterella palustris Shearer & J.L. Crane
b
2
Lepidopterella tangerina Raja & Shearer
a
1
Massarina bipolaris K.D. Hyde
c
12 1
Massarina fronsisubmersa K.D. Hyde
c
1
Massarina ingoldiana Shearer & K.D. Hyde
b
18
Micropeltopsis sp. F115 1
Ophiobolus shoemakeri Raja & Shearer
a
2
Trematosphaeria lineolatispora K.D. Hyde
b
141
Undescribed Massarina sp. F60 2
Undescribed Massarina sp. F65 1
Undescribed Massarina sp. F80 1
Undescribed genus F76 1 1
Leotiomycetes
Aquapoterium pinicola Raja & Shearer
d
312
Gorgoniceps sp. F72 1
Sordariomycetes
Aniptodera aquadulcis (S.Y. Hsieh, H.S. Chang & E.B.G. Jones)
J. Campb., J. Anderson & Shearer
b
1
Aniptodera chaesapeakensis Shearer & Miller
b
1
Aniptodera inflatiascigera K.M. Tsui, K.D. Hyde & I.J. Hodgkiss
c
131
Aniptodera lignatilis K.D. Hyde
c
231
Aniptodera megaloascocarpa Raja & Shearer
a
1
Aniptodera palmicola K.D. Hyde, W.H. Ho & K.M. Tsui
b
261
Biodivers Conserv (2009) 18:419–455 427
123

Table 1 continued
Species Collection sites
BW AP OC BC EV
Annulatascus velatisporus K.D. Hyde
b
6172
Arnium gigantosporum Raja & Shearer
a
1
Ascitendus austriacus (Re
´
blova
´
, Winka & Jaklitsch) J. Campb. & Shearer
b
33
Ascosacculus heteroguttulatus (S.W. Wong, K.D. Hyde & E.B.G. Jones)
J. Campb., J.L. Anderson & Shearer
b
221 1
Ascotaiwania hsilio H.S. Chang & S.Y. Hsieh
b
11
Ayria appendiculata Fryar & K.D. Hyde
c
1
Cataractispora bipolaris (K.D. Hyde) K.D. Hyde, S.W. Wong & E.B.G. Jones
c
1
Cyanoannulus petersenii Raja, J.Campb. & Shearer
b
41
Fluviatispora reticulata K.D. Hyde
c
253
Flammispora pulchra Raja & Shearer
a
1
Hanliniomyces hyaloapicalis Raja & Shearer
d
21
Jobellisia luteola (Ellis & Everh.) M.E. Barr
b
33
Lockerbia striata Raja & Shearer
a
11
Luttrellia estuarina Shearer
b
1
Luttrellia sp. F14 1
Lulworthia sp. F54 1
Nais inornata Kohlm. 1 5 3 1
Natantispora retorquens (Shearer & J.L. Crane) J. Campb.,
J.L. Anderson et Shearer
b
21
Ophioceras commune Shearer, J.L. Crane & Chen
b
22
Phaeonectriella lignicola R.A. Eaton & E.B.G. Jones
b
2
Phomatospora triseptata Raja & Shearer
a
1
Pseudoproboscispora caudae-suis (Ingold) J. Campbell & Shearer
b
3
Physalospora limnetica Raja & Shearer
a
1
Savoryella aquatica K.D. Hyde
b
21
Savoryella fusiformis W.H. Ho, K.D. Hyde & I.J. Hodgkiss
c
2
Submersisphaeria aquatica K.D. Hyde
b
15 6 1
Torrentispora fibrosa K.D. Hyde, W.H. Ho, E.B.G. Jones,
K.M. Tsui & S.W. Wong
b
212
Vertexicola caudatus K.D. Hyde, S.W. Wong & V.M. Ranghoo
b
11
Zopfiella latipes (N. Lundq.) Malloch & Cain
b
311
Zopfiella lundqvistii Shearer & J.L. Crane
b
2
Unidentified ascomycete sp. F28 1
Undescribed sp. F41 2
Undescribed genus F44 1
Undescribed genus F50 1
Undescribed sp. F57 2
Undescribed genus F99 1
Undescribed sp. F100 1
Undescribed sp. F107 2
Basidiomycetes
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123

Table 1 continued
Species Collection sites
BW AP OC BC EV
Rogersiomyces okefenokeensis J.L. Crane & Schoknecht
b
1
Mitosporic fungi
Acrogenospora sphaerocephala (Berk. and Broome) M. B. Ellis
b
111 1
Ardhachandra cristaspora (Matsush.) Subram. & Sudha
c
1
Bactrodesmium linderi (J.L. Crane & Shearer) M.E. Palm & E.L. Stewart
b
4231
Berkleasmium concinnum Berk. (S. Hughes)
b
1
Brachiosphaera tropicalis Nawawi
c
1
Brachysporium obovatum Kiessl
b
1
Cancellidium applanatum Tubaki
b
781
Canalisporium caribense (Hol.-Jech. & Mercado) Nawawi & Kuthub.
c
1
Canalisporium kenyense Goh, W.H. Ho, K.D. Hyde, S.R. Whitton & T.E. Umali
b
1
Cacumisporium sigmoideum Mercado & R.F. Castan
˜
eda
b
31
Chaetospermum camelliae Agnihothr.
c
112
Coleodictyospora micronesia (Matsush.) Matsush. 1 1
Cordana abramovii Seman & Davydkina var. seychellensis K.D. Hyde & Goh
b
1
Condylospora sp. FH 34 1
Cryptophiale cucullata Kuthub.
b
1
Dactylaria hyalotunicata K.M. Tsui, Goh & K. D. Hyde
c
11
Dactylaria tunicata Goh & K.D. Hyde
b
61
Delortia palmicola Pat.
b
5
Dendrosporium lobatum Plakidas & Edgerton ex J.L. Crane
b
1
Dendrospora sp. FH 87 1
Dictyosporium sp. FH39 1
Dictyosporium digitatum J.L. Chen, C.H. Hwang & S. S. Tzean
c
223
Dictyosporium elegans Corda
b
1
Dictyosporium giganticum Goh & K.D. Hyde
b
1
Dictyosporium heptasporum (Garov.) Damon
b
1
Ellisembia abscendens (Berk.) Subram.
b
12 1
Exserticlava globosa Roa & de Hoog
c
1
Exserticlava triseptata (Matsush.) S. Hughes
c
11
Excerticlava vasiformis (Matsush.) S. Hughes
c
23
Helicodendron sp. FH38 3
Helicoon auratum (Ellis) Morgan
b
1
Helicomyces roseus Link
b
31
Helicosporium aureum (Corda) Linder
b
1
Helicosporium guianense Linder
b
1
Helicosporium gigasporum K.M. Tsui, Goh, K.D. Hyde & Hodgkiss
b
32
Helicosporium lubricopsis Linder
b
1
Humicola asteroidea Udagawa & Y. Horie
b
1
Ingoldiella hamata D.E. Shaw
c
1
Intercalarispora sp. FH88 1
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123

Habitat distribution
Based on the presence or absence data in a site by species matrix, analyses using CAP
provided some separation of freshwater fungal communities based on lentic and lotic sites,
with some lentic habitats forming a separate cluster from the lotic habitats on the ordination
graph (Fig. 3). However, there was not enough evidence to support distinct cluster for-
mation between lentic and lotic habitats. The analysis found one canonical axis.
Permutation tests did not provide statistical support for the hypothesis that fungal com-
munities in lentic and lotic habitats are distinctly different (trace statistic = 0.5108,
P [0.05; 1st squared canonical correlation = 0.5108, P [ 0.05). An additional CAP
analysis between lentic and lotic habitats in northern Florida, where the number of lentic
and lotic sites were more similar, also showed some separation between habitats (Fig. 4),
but again, permutation tests did not provide statistical support. The analysis found a single
Table 1 continued
Species Collection sites
BW AP OC BC EV
Monacrosporium ellipsosporum (Preuss) R.C. Cooke & C.H. Dickinson
b
2
Monotosporella sp. FH20 3 1
Melanocephala australiensis G.W. Beaton & M.B. Ellis
b
1
Nawawia filiformis (Nawawi) Marvanova
´
c
1
Pleurophragmium malaysianum Matsush.
c
922
Pleurothecium recurvatum (Morgan) Ho
¨
hn.
b
1
Septonema hormiscium Sacc.
b
11
Speiropsis sp. FH59 1
Sporidesmiella hyalosperma var. novae-zealandiae
b
(S. Hughes). P.M. Kirk 1
Sporidesmium tropicale M.B. Ellis
b
1
Sporidesmium sp. FH27 1
Sporoschisma saccardoi Mason & S. Hughes
b
11
Tetraploa aristata Berk. & Broome 1
Thozetella sp. 1
Vargamyces sp. FH86 1
Varicosporium sp. FH22 1
Wiesneriomyces larinus (Tassi) P.M. Kirk
b
11
Xylomyces chlamydosporus Goos, R.D. Brooks & Lamore
b
133
Total number of samples collected
e
52 41 49 25 13
Species richness 41 59 63 33 19
BW Blackwater River State Forest, AP Apalachicola National Forest, OC Ocala National Forest, BC Big
Cypress National Preserve, EV Everglades National Park
a
New species described in this study
b
New records for Florida
c
New records for North America and Florida
d
New genus described in this study
e
Each collection consisted of about 20–30 samples of wood and herbaceous debris depending on
availability
430 Biodivers Conserv (2009) 18:419–455
123

canonical axis (trace statistic = 0.4643, P [ 0.05; 1st squared canonical correlation =
0.4643, P [ 0.05).
Seventy-five species were collected only from lentic habitats, 16 species were collected
only from lotic habitats, and 41 species were collected from both lentic and lotic habitats
(Appendix 3). Numerous species of freshwater ascomycetes, including Boerlagiomyces
websteri, Jahnula rostrata, J. sangamonensis, Lepidopterella palustris, Trematosphaeria
lineolatispora, Undescribed Massarina sp. F60, Aquapoterium pinicola, Aniptodera ligna-
tilis, A. palmicola, Annulatascus velatisporus, Ascitendus austriacus, Ascosacculus
heteroguttulatus, Cyanoannulus petersenii, Fluviatispora reticulata, Hanliniomyces hyalo-
apicalis, Nais inornata, Natantispora retorquens, Ophioceras commune, Submersisphaeria
aquatica, Torrentispora fibrosa, Vertexicola caudatus, Zopfiella latipes, and Z. lundqvistii
occurred in both lentic and lotic habitats. A number of these species also occurred frequently
and were reported from more than one collection site along the Florida Peninsula (Table 1;
Appendix 3).
Aliquandostipite minuta, Arnium gigantosporum, Caryospora obclavata, Flammis-
pora pulchra, Lockerbia striata, Ophiobolus shoemakeri, Phomatospora triseptata,
-0.2
-0.1
0
0.1
0.2
0.3
0.30.20.10-1.0-0.2
Axis 1
North
Central
South
Fig. 2 Canonical analysis of principal coordinates (CAP) of fungal communities in freshwater habitats
based on latitudinal ecotone along north, central and south Florida (n = 97)
Table 2 Index of Sørenson’s similarity among five collection sites
Sites BW AP OC BC EV
BW – 0.42 0.31 0.13 0.13
AP – 0.32 0.17 0.20
OC – 0.25 0.21
BC – 0.34
EV –
Biodivers Conserv (2009) 18:419–455 431
123

Physalospora limnetica, undescribed Massarina sp. F65, F80, undescribed sp. F41, F44,
F99, F100, and F107 were collected only from lentic habitats. Whereas, two undescribed
genera, F76 and F50, were collected exclusively from lotic habitats (Appendix 3).
-0.2
-0.1
0
0.1
0.2
0.3
Axis 1
Lo
Le
Lo = Lotic habitats
Le = Lentic habitats
Fig. 3 Canonical analysis of principal coordinates (CAP) of fungal species in freshwater habitats based on
lotic and lentic habitat type (n = 97). Lo = Lotic habitats, Le = Lentic habitats
-0.3
-0.2
-0.1
0
0.1
0.2
0.3
Axis 1
Lo
Le
Lo = Lotic habitats
Le = Lentic habitats
Fig. 4 Canonical analysis of principal coordinates (CAP) of fungal species in freshwater habitats based on
lotic and lentic habitat type in north Florida (n = 48). Lo = Lotic habitats, Le = Lentic habitats
432 Biodivers Conserv (2009) 18:419–455
123

Substrate distribution
One hundred of the 132 species collected from Florida were recorded only on woody
debris, 14 species only on herbaceous debris, and 18 species occurred on both herbaceous
and woody debris (Appendix 3). Freshwater ascomycetes, such as O. shoemakeri and
P. limnetica, were collected exclusively from herbaceous debris in lentic habitats. Aqu-
apoterium pinicola, collected only from pine needles, was found in both lentic and lotic
habitats. Other taxa that occurred only on herbaceous debris include some freshwater
mitosporic ascomycetes such as Ardhachandra cristaspora, Coleodictyospora micronesia,
Cryptophiale cucullata, Dendrosporium lobatum, Sporidesmium sp., and Tetraploa
aristata.
Eighteen species occurred on both herbaceous and woody debris and may be substrate
generalists. Among these, Annulatascus velatisporus, Ascitendus austriacus, Ascosacculus
heteroguttulatus, Bactrodesmium linderi and Cancellidium applanatum were the most
commonly occurring species during the study and were found in more than two collection
sites in both lentic and lotic habitats along the Florida Peninsula (Appendix 3). Species that
occurred on both woody and herbaceous debris usually occurred on herbaceous debris in
lentic habitats. Two exceptions are A. austriacus and a new undescribed fungus (F76) that
occurred on herbaceous debris in lotic habitats.
Relationship of freshwater ascomycete community composition to pH and latitude
When community composition was examined as a function of latitude and pH separately
(Figs. 5, 6), both these variables remained significant as determinants of variation in
community composition. Canonical correspondence analyses with pH as the variable found
one canonical axis (trace statistic = 0.462131, P \0.05; 1st squared canonical correla-
tion = 0.462131, P \ 0.05) and pH explained about 40% of the variation (Fig. 5).
Canonical correspondence analyses with latitude also found one canonical axis (trace
R
2
= 0.4564
-0.2
-0.1
0
0.1
0.2
0.3
987654
pH
Fig. 5 Canonical correlation analysis using CAP to explain the relationship of freshwater fungal species
with pH (n = 97)
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123

statistic = 0.620395, P \ 0.01; 1st squared canonical correlation = 0.620395, P \ 0.01)
and latitude explained about 60% of the variation (Fig. 6). Ordination analysis also pro-
vided some evidence of interaction of both pH and latitude as explanatory variables in
determining variation in community composition. The analysis found two canonical axes
with significant statistical support (trace statistic = 1.123, P \ 0.01; 1st squared canonical
correlation = 0.664, P \ 0.01). The freshwater fungal communities showed some sepa-
ration based on both latitude (vertical axis) and pH (horizontal axis) on the ordination
graph (Fig. 7). The southern sites formed a cluster distinct from the northern and central
sites on axis 2 (vertical axis) and colors representing low pH values \ 6.5 are at the top,
the middle pH values are mostly in the middle and the high pH values are almost all at the
bottom of the ordination graph (Fig. 7).
Discussion
Latitudinal distribution patterns along the temperate–subtropical ecotone in Florida
This study demonstrates that species composition of freshwater ascomycete communities
differs along the temperate–subtropical ecotone in the Florida Peninsula. The differences in
species composition are resolved in the ordination graph as three different clusters shown
as north, central and south (Fig. 2). Species composition in the north and central sites are
more similar to one another than either is to the southern sites. High turnover in species
composition is indicated by the lower similarity values between the northern and southern
sites, than between the northern and central sites (Table 2). The results of the canonical
correspondence analysis for latitude (Fig. 7) are somewhat consistent with the results of
discriminant function analysis (Fig. 2), which shows the freshwater fungal communities in
north and central Florida are clustered closer together than to those in south Florida. For
R
2
= 0.6623
-0.3
-0.2
-0.1
0
0.1
0.2
20 22 24 26 28 30 32
Latitude in Degrees North
Fig. 6 Canonical correlation analysis using CAP to explain the relationship of freshwater fungal species
with latitude (n = 97)
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example, freshwater ascomycetes, Lepidopterella palustris, L. tangerina, A. austriacus,
Jobellisia luteloa, Lockerbia striata, Pseudoproboscispora caudae-suis, F41, and F57
occurred only in north Florida sites, but were absent from south Florida. On the other hand,
Aliquandostipite minuta, A. khaoyaiensis, A. siamensiae, Jahnula bipolaris, and J. aus-
traliensis, and mitosporic ascomycetes, Coleodictyospora micronesia, Canalisporium
caribense, Brachiosphaera tropicalis, Dictyosporium heptasporum, and Nawawia filiformis
occurred in the southern sites, but were completely absent from the northern sites in
Florida (Table 1). The above species recorded from south Florida have been reported
previously from the paleotropics and subtropics as well as from freshwater habitats in the
neotropics (http://fungi.life.uiuc.edu/).
Previous studies on aquatic fungi have also revealed differences in species composition
along latitudinal zones that differ in climatic conditions. Miura (1974) conducted a study of
freshwater Ingoldian mitosporic ascomycetes in Japan along a group of islands that were
located along a latitudinal gradient from 45°N, 37°N, and 25°N and found that the species
composition showed a pronounced change along the subtropical collection sites. He also
found that the species composition of Ingoldian fungi in streams sampled at the northern
latitudes were more similar to each other than either of them was to the southern sub-
tropical region in Japan. Shearer and Burgos (1987) found a change in species composition
of lignicolous marine fungi along collection sites between north, central and south Chile.
They found that some species occurring in northern subtropical Chile did not occur in the
anti-boreal southern part of Chile. For marine fungi studied along a wider latitudinal
gradient, Hughes (1974) and Booth and Kenkel (1986) also found that temperature was an
important environmental parameter in explaining the latitudinal distribution patterns of
marine fungi. Similarly, Taylor et al. (2000) and Fro
¨
hlich and Hyde (2000) found the palm
saprobes from sites of the same latitude/longitude were more similar than sites from
different latitudes.
-0.3
-0.2
-0.1
0
0.1
0.2
-0.3 -0.2 -0.1 0 0.1 0.2
Axis 1
N <6.5
N 6.5–7.49
N >7.5
C <6.5
C 6.5–7.49
S <6.5
S 6.5–7.49
S >7.5
Fig. 7 Canonical correlation analysis (CCA) to explain the relationship of freshwater fungal species with
pH and latitude (n = 97)
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Differences in species composition observed in this study are also similar to patterns
observed for other groups of organisms sampled along the temperate–subtropical ecotone
of the Florida Peninsula. The northern and southern distributional limits of several ver-
tebrate taxa in Florida (Whitaker 1968) show distinct patterns similar to those observed for
the freshwater ascomycetes. A number of vertebrate taxa found commonly in eastern North
America apparently do not occur in central and south Florida (Sprunt 1954; Whitaker
1968). Humphrey (1975) found that the common bat species Lasiurus cinerus and
L. borealis are absent south of 29° latitude in Florida. In another study, Christman (1975)
examined the distribution of snake species in Florida and concluded that seasonal tem-
perature extremes were mainly responsible for a pronounced north–south replacement
series along the Florida Peninsula. In a study conducted at the beginning of the 20th
century, Brown (1909) delineated three forest regions for peninsular Florida as north
Florida (29.5° latitude), transitional pine (27.5–29.5° latitude), and south Florida or
Antillean (27.5° latitude) based on the distribution of tree species.
Our results support a previous study that showed greater similarity in species compo-
sition among sites located at similar latitudes than between species on the same continent
found at different temperate and tropical latitudes (Wood-Eggenschwiler and Ba
¨
rlocher
1985). For example, in our study several freshwater ascomycete species, such as A. kha-
oyaiensis, J. australiensis, Massarina bipolaris, Aniptodera inflatiascigera, A. palmicola,
Ayria appendiculata, Fluviatispora reticulata, Torrentispora fibrosa, and Vertexicola
caudatus, and mitosporic ascomycetes, P. malaysianum and D. tunicata, have been
reported previously from southeastern Asian tropical and subtropical regions (Fryar and
Hyde 2004; Hyde 1993, 1994, 1995; Hyde et al. 1999, 2000; Inderbitzin et al. 2001;
Ranghoo et al. 2000; Tsui et al. 1997), but not from other states in USA north of Florida
(Shearer 1993, 2001). Speculating on the latitudinal distribution of aquatic hyphomycetes,
Ingold (1966) stated: ‘‘I think I could tell my latitude with an error of 15° by examining the
aquatic spores in a sample of stream foam.’’
An increase in species richness from higher latitudes (polar zones) towards lower latitudes
(equatorial zones) is the oldest and most fundamental pattern concerning the distribution of
life on earth and a widely documented macroecological pattern (see Rosenzweig 1995;
Brown and Lomolino 1998; Gaston and Blackburn 2000). The latitudinal gradient in
diversity and/or richness has been well documented for several groups of organisms in
marine (Gray 2001), freshwater (France 1992), and terrestrial habitats (Stevens 1989;
Blackburn and Gaston 1996; Ruggiero 1999). For freshwater ascomycetes sampled along the
Florida Peninsula, however, there was no prominent richness gradient (Table 1), despite
changes in species composition along the peninsula as discussed above. A total of 75 species
were collected from north Florida (BW ? AP) from 48 habitats sampled, 63 species were
collected from central Florida (OC) from 24 habitats sampled, while only 43 species were
collected from south Florida (BC ? EV) from 25 habitats sampled. Since species richness is
dependent, in part, on sampling effort (Magurran 2004), which to some extent was uneven
among the collection sites in this study, further work along the Florida ecotone is needed to
validate this pattern.
One reason for lower richness in the southern sites in this study might be due to the
relatively homogenous vegetational composition at the BC site. Although extensive
freshwater cypress swamps are present in BC, and EV is an extensive freshwater wetland
(Kushlan 1990), the number of easily accessible discrete freshwater habitats such as lakes
and streams are fewer in BC and EV compared to the other northern and central sites in the
study. In addition, EV, is referred to as the ‘‘river of grass’’ (Douglas 1947), and one of the
most abundant sedge species in the EV is a Cladium sp., commonly called saw grass. This
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substrate is highly depauperate of fungal growth and we have found only a few fungal
species colonizing submerged saw grass (H.A. Raja and C.A. Shearer, personal observa-
tion). Sampling of fungi in other freshwater habitats not colonized by saw grass, but still
located at lower latitudes should be carried out in future studies to further resolve species
richness patterns with respect to latitudinal distribution in the Florida Peninsula.
Reverse richness gradients have been observed previously in peninsulas and this pattern
is so called the ‘‘peninsula effect’’ (Simpson 1964; and see Brown and Lomolino 1998).
The peninsula effect is a pattern of diversity documented thus far mostly in terrestrial
environments where species richness decreases along peninsulas with increasing distance
from their continental attachments. The peninsula effect has been shown along Florida for
herpetofauna (Means and Simberloff 1987) and for avian species (Robertson 1955; Rob-
ertson and Kushlan 1974). The above mentioned studies suggest that the decline in species
richness occurs due to a combination of historical effects and decline in habitat for species
to colonize the lower parts of the peninsula.
Data from this study suggest that the compression of latitudinal thermoclines along the
temperate–subtropical ecotone in peninsular Florida may be an important driving force in
shaping the species composition of freshwater ascomycete communities.
Habitat distribution
Although CAP showed some clustering of species from lentic habitats separated from lotic
habitats based on comparisons of species collected in all lentic and lotic habitat types in
Florida (Fig. 3), analyses showed that habitat was not a significant factor (P [ 0.05) in
explaining the distribution patterns of freshwater ascomycete species. The analyses com-
paring all lotic and lentic habitats might be biased from uneven sampling due to the larger
number of lentic habitats sampled (n = 71) compared to lotic habitats (n = 26) in the
study. Therefore, another CAP analysis was employed for lentic and lotic habitats in north
Florida where the number of lotic (n = 21) and lentic (n = 27) habitats sampled were
geographically closer to each other. The second CAP analysis (Fig. 4) revealed similar
results to the first and showed that habitat was not a significant factor (P [ 0.05). These
data suggest that freshwater ascomycetes communities are not significantly different
although some species appeared to be exclusive to one or the other habitat type.
Numerous species of freshwater ascomycetes, including Boerlagiomyces websteri,
Jahnula rostrata, J. sangamonensis, L. palustris, Trematosphaeria lineolatispora, Unde-
scribed Massarina sp. F60, Aquapoterium pinicola, Aniptodera lignatilis, A. palmicola,
Annulatascus velatisporus, Ascitendus austriacus, Ascosacculus heteroguttulatus, Cyano-
annulus petersenii, F. reticulata, Hanliniomyces hyaloapicalis, Nais inornata, Natantispora
retorquens, Ophioceras commune, Submersisphaeria aquatica, T. fibrosa, V. caudatus,
Zopfiella latipes and Z. lundqvistii occurred in both lentic and lotic habitats (Appendix 3). A
number of these species also occurred frequently and were reported from more than one
collection site along the Florida Peninsula and may be habitat generalists (Appendix 3).
Fourteen new taxa were collected only from lentic habitats, whereas only two new taxa
were collected exclusively in lotic habitats (Appendix 3). Since lentic habitats are often
spatially discrete, which can restrict gene flow between their respective fungal populations;
the populations may be subjected to different selective pressures. This characteristic could
lead to higher rates of speciation than non-discrete habitats such as streams and rivers. The
above hypothesis might be one of several possible reasons why more new species were
found in lentic habitats than in lotic habitats. Additional comparative collections of
freshwater ascomycetes from lentic and lotic habitats and population level studies of lentic
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and lotic species are needed to determine the importance of isolation as a mechanism of
speciation in aquatic ascomycetes.
Comparison of the freshwater ascomycetes from Florida lakes to those collected by
Fallah (1999) from north temperate lakes in Wisconsin showed that P. caudae-suis and
M. ingoldiana were the only two species common to collections made in lentic habitats
from Wisconsin and Florida. These two species are widely distributed in both lentic and
lotic habitats in North America (Fallah and Shearer 2001; Shearer 2001; Campbell et al.
2003). Because there is a greater difference in the water temperature regimes between
Wisconsin and Florida, differences in species composition between lakes in these two
regions is not unexpected. Another factor that might contribute to the striking difference in
fungal species composition between Wisconsin and Florida might be due to geological
ages of lakes in the two regions. Lakes in the northern latitudes are geologically younger
than those in Florida due to the most recent glaciation, which did not reach Florida
(Brenner et al. 1990).
We also compared the species composition of freshwater fungi from Florida Lakes to
that of a tropical lake in Australia (Hyde and Goh 1998a), and to subtropical lakes in China
(Cai et al. 2002; Luo et al. 2004). About 10% of the freshwater meiosporic and mitosporic
fungal species were similar with Lake Barrine in Australia, and approximately 10–12%
species were similar to the lakes from China. Fungal taxa that showed overlap among the
lakes in the three geographical areas are common species that are distributed worldwide,
such as Aniptodera chesapeakensis, Kirschsteiniothelia elaterascus, Nais inornata,
Sporoschisma saccardoi, and Zopfiella latipes (see http://fungi.life.uiuc.edu/).
Water pH was shown to be significant in explaining the variation in community com-
position of freshwater fungi in Florida as fungal communities were divided into three more
or less distinct groups based on pH values (Fig. 7). Water chemistry plays an important
role in the distribution of freshwater organisms (Hynes 1970; Wetzel 2001). Previous
studies of freshwater fungi have also shown that pH is an important factor in the distri-
bution of freshwater ascomycetes (Fallah 1999), as well as freshwater Ingoldian mitosporic
fungi (Wood-Eggenschwiler and Ba
¨
rlocher 1983; Shearer and Webster 1985;Ba
¨
rlocher
1987;Ba
¨
rlocher and Rosset 1981). In a review of the effects of water chemistry on the
distribution of freshwater Ingoldian mitosporic ascomycetes, Chamier (1992) concluded
that pH may have an indirect effect on the distribution of freshwater mitosporic fungi by
affecting the solubility of Al or other metals in freshwater habitats.
Substrate distribution
Eighteen of 132 taxa occurred on both herbaceous and woody debris and may be substrate
and habitat generalists. Among these, Annulatascus velatisporus, A. austriacus, Ascosac-
culus heteroguttulatus, Bactrodesmium linderi and Cancellidium applanatum were the
most commonly occurring species and were found in more than two collection sites in both
lentic and lotic habitats along the Florida Peninsula (Table 1; Appendix 3). Species that
occurred on both woody and herbaceous debris usually occurred on herbaceous debris in
lentic habitats. Two exceptions are A. austriacus and a new undescribed fungus (F76) that
occurred on herbaceous debris in lotic habitats.
Freshwater ascomycetes that occur on herbaceous substrates are reported more fre-
quently from lentic habitats probably because herbaceous macrophytes are much more
common in lentic habitats than lotic habitats (Shearer 1993; Shearer
2001). Fallah (1999)
found a number of freshwater ascomycetes exclusively on herbaceous material in lentic
habitats. Ascovaginospora stellipala Fallah, Shearer & J.L. Crane, Aquadulciospora
438 Biodivers Conserv (2009) 18:419–455
123

rhomboidia Fallah & Shearer, Phaeosphaeria barriae Fallah & Shearer, Phaeosphaeria
vilasensis Fallah, Shearer & Leuchtm. and Phomatospora muskellungensis Fallah &
Shearer occurred only on herbaceous debris collected from lakes in Wisconsin (Fallah
1999). The above taxa have not been reported from wood in lentic or lotic habitats from
other geographical locations thus far (http://fungi.life.uiuc.edu/) and may be specialists on
herbaceous material. Whether or not these taxa might have adapted to freshwater with their
plant hosts is an interesting question (Shearer 1993, 2001).
Aquapoterium pinicola and Ophiobolus shoemakeri, two species of freshwater mei-
osporic ascomycetes reported only from herbaceous debris in this study, were tested for the
production of extracellular enzymes in vitro and found positive for cellulase, endoglu-
canase, beta-glucosidase, xylanase and amylase (Simonis et al. 2008), enzymes important
in decomposing herbaceous substrates. Although positive for xylanase, A. pinicola and O.
shoemakeri were both negative for production of the lignin modifying enzymes, peroxidase
and tyrosinase, and they did not cause soft-rot in balsa wood. It is plausible that lack of
lignin modifying enzymes and inability to form soft-rot cavities may prevent species such
as A. pinicola and O. shoemakeri from being competitive on woody substrates and thereby
limit their occurrences to herbaceous debris.
Studies that have examined substrate specificity of mitosporic Ingoldian fungi (Gulis
2001; Nikolcheva and Ba
¨
rlocher 2005) suggest that most aquatic hyphomycetes can col-
onize and grow on a wide range of substrate types (Webster and Descals 1981;Ba
¨
rlocher
1992; Suberkropp 1992). However, relative frequencies of individual species may be
influenced by the substrate (Ba
¨
rlocher 2005). For example, Ba
¨
rlocher (1982) found that the
dominant species on conifer needles differ from the dominant species on deciduous leaf
litter. Ba
¨
rlocher and Grac¸a (2002) showed that fungal communities of streams running
through eucalyptus stands are more similar to each other than those running through mixed
deciduous forest. Gulis (2001) conducted a study on Ingoldian mitosporic ascomycetes to
test preferences among various substrate types such as leaf litter, woody debris and her-
baceous debris (grass blades) and found that wood and grass blades had a distinct fungal
assemblage clearly different from those supported by deciduous leaf litter. Ferreira et al.
(2006) reported a clear preference of aquatic hyphomycete species towards either leaves or
wood, they found that the fungal assemblage on leaves was different from those colonizing
balsa wood veneers. On the other hand, Nikolcheva and Ba
¨
rlocher (2005) did not find strict
substrate preferences for Ingoldian mitosporic ascomycetes using both traditional and
molecular techniques, and they concluded that strict exclusion of fungi by substrate type
was rare, and that presence of different species or phylotypes was governed by seasonality
in their study.
In our study, some differences were found in the freshwater ascomycete species colo-
nizing herbaceous debris versus those found on woody debris, but, in general, except for
Aquapoterium pinicola, Ayria appendiculata, O. shoemakeri and Physalospora limnetica,
and other mitotic ascomycetes (Appendix 3) most species occurred on woody debris. Of the
548 species of freshwater meiosporic ascomycetes reported in the literature, 375 were col-
lected from wood and 173 were collected from herbaceous debris (http://fungi.life.uiuc.edu/)
while in the present study, of the 74 freshwater meiosporic ascomycetes, 61 species occurred
only on woody debris, eight occurred on both substrate types, while only four occurred
exclusively on herbaceous debris (Appendix 3). The occurrences of few meiosporic asco-
mycetes on herbaceous debris might be due to some features of wood not characteristic of
herbaceous debris, such as persistence over time (Shearer 1992) or nutrient content (higher
carbon: nitrogen ratio) which may stimulate the production of sexual states.
Biodivers Conserv (2009) 18:419–455 439
123

Conclusions
1. Results from CAP analyses, and Sørenson’s similarity index indicate that the fresh-
water ascomycete community composition differs between northern and southern sites
along the temperate–subtropical latitudinal ecotone of the Florida Peninsula. Canon-
ical correspondence analyses revealed that both latitude and pH are important factors
in explaining the distribution patterns of freshwater ascomycetes. A number of
freshwater ascomycete species from central and south Florida occur in the paleotropics
as well as neotropics but have not been reported thus far from north Florida as well as
other states north of Florida.
2. Ordination analysis of freshwater ascomycete communities indicated that lentic and
lotic communities were not significantly different. Some geographically broadly
distributed species and species commonly found in Florida occurred in both habitats
whereas a number of rare species were collected either in lentic or lotic habitats.
3. Of the 132 taxa of meiosporic and mitosporic freshwater ascomycetes reported from
Florida, 100 species were reported on woody debris, while only 14 species occurred
exclusively on herbaceous debris. Eighteen species, of which some were commonly
occurring taxa, were found on both woody and herbaceous debris. Most of the taxa
reported from herbaceous debris were reported from lentic habitats. Species occurring
on woody debris in lotic habitats were also found on wood in lentic habitats, but
species found exclusively on herbaceous debris in lentic habitats were seldom found
on wood or herbaceous debris in lotic habitats. This may reflect, to some degree, the
absence of herbaceous substrates in many lotic habitats.
4. With respect to the conservation of freshwater ascomycetes, a broad latitudinal
approach must be taken. Given the large number of new species discovered in this
single study, additional geographically broad studies are warranted to fully understand
the biodiversity of this group.
Acknowledgments We thank Dr. Andrew N. Miller, Christopher Brown, and Dr. J.L. Crane for their
assistance with collecting. Appreciation is expressed to the rangers at Blackwater River State Forest,
Apalachicola National Forest and Ocala National Forest, for permission to collect within the forests. We are
grateful to the Superintendent of Big Cypress National Preserve and Everglades National Park for providing
permits to collect aquatic fungi. Financial support of this study by the National Science Foundation (NSF
Grant No. DEB 03-16496) and the National Institutes of Health (NIH Grant No. R01GM-60600), Clark
Research Grant from Integrative Biology, UIUC, and The Mycological Society of America Graduate Fel-
lowship Award are gratefully acknowledged. Any opinions, findings, and conclusions or recommendations
expressed in this publication are those of the authors and do not necessarily reflect the views of the National
Science Foundation and National Institutes of Health. This work represents a portion of a thesis in partial
fulfillment of the requirements for the doctoral degree at the Graduate College of the University of Illinois at
Urbana-Champaign.
Appendices
Appendix 1 List of habitats sampled with their water temperature and pH ranges as measured at the time
of sample collection at five sites within the Florida Peninsula
Collection site Latitude/longitude Temperature (°C) pH
Blackwater River State Forest
Blackwater River 30°56
0
01
00
N, 86°44
0
07
00
W 11–27 5.5–7
Bone Creek 30°44
0
10
00
N, 86°43
0
56
00
W 25–27 5.5–6.7
440 Biodivers Conserv (2009) 18:419–455
123

Appendix 1 continued
Collection site Latitude/longitude Temperature (°C) pH
Penny Creek 30°45
0
05
00
N, 86°46
0
54
00
W 23 6.6
Alligator Creek 30°44
0
39
00
N, 86°52
0
54
00
W 7–24 5.5–5.7
Juniper Creek 30°50
0
01
00
N, 86°54
0
11
00
W 11–24 5.5–6.3
Sweetwater Creek 30°51
0
21
00
N, 86°51
0
03
00
W 13–26 5.5–6.8
Coldwater Creek 30°52
0
59
00
N, 86°57
0
28
00
W 11–25 5.5–6.4
Pringle Branch 30°54
0
33
00
N, 86°58
0
04
00
W 11–23 5.5–6.4
Hurricane Creek 30°56
0
36
00
N, 86°44
0
50
00
W30 8
Maria Branch 30°46
0
39
00
N, 86°54
0
42
00
W 7 5.5
Hawkins Creek 30°58
0
11
00
N, 86°59
0
43
00
W 30 5–5.5
Dixon Branch 30°54
0
50
00
N, 86°57
0
52
00
W 23 6.7
Horns Creek Swamp 30°46
0
31
00
N, 86°54
0
43
00
W30 6
Pitman Creek Swamp 30°48
0
57
00
N, 86°55
0
31
00
W 27 5.7
Deaton Bridge Swamp 30°43
0
49
00
N, 86°52
0
29
00
W 36 6.3
Bear Lake 30°51
0
43
00
N, 86°49
0
56
00
W 11–34 6.5–7.6
Hurricane Lake 30°56
0
19
00
N, 86°45
0
12
00
W 11–36 5.5–8.7
Karick Lake 30°53
0
45
00
N, 86°38
0
30
00
W 13 5.5
Kennedy Branch Swamp 30°56
0
37
00
N, 86°44
0
49
00
W 11–12 5.5
Blackwater River Shingles Branch Swamp 30°43
0
30
00
N, 86°7
0
40
00
W 13 5.5
Calloway Swamp 30°53
0
28
00
N, 86°58
0
03
00
W 12 5.5
Apalachicola National Forest
Big Gulley Creek 30°15
0
42
00
N, 85°00
0
46
00
W 27 6.6
Kennedy Creek 30°06
0
31
00
N, 85°03
0
37
00
W 27–28 7
Ochlockonee River 30°05
0
37
00
N, 84°37
0
42
00
W 24 5.5
Fisher Creek 30°18
0
51
00
N, 84°23
0
57
00
W 25–39 5.5–7.5
Syfreet Creek 30°05
0
02
00
N, 84°34
0
54
00
W 26 5.5–7.4
Little Gully Creek 30°15
0
39
00
N, 85°00
0
46
00
W 9–26 5.5
Rowletts Creek 30°51
0
30
00
N, 85°01
0
10
00
W 9–35 5.5
New Crossing Branch 30°13
0
08
00
N, 84°59
0
21
00
W 8 5.5
Apalachicola River 29°56
0
24
00
N, 85°00
0
41
00
W9 6
Field Branch Creek 30°03
0
24
00
N, 85°03
0
36
00
W30 7
Apalachicola River Swamp 29°56
0
20
00
N, 85°00
0
45
00
W 30 7.5
Owls Creek Swamp 30°03
0
31
00
N, 85°01
0
11
00
W 30 7.5
Leon Sinks 30°18
0
35
00
N, 84°20
0
46
00
W21 8
Dog Pond 30°20
0
55
00
N, 84°24
0
41
00
W 30 5.5
Unnamed Pond 29°54
0
07
00
N, 84°20
0
36
00
W30 6
Unnamed Swamp 1 30°17
0
02
00
N, 84°50
0
25
00
W 30 6.5
Andrew Lake 30°24
0
09
00
N, 30°24
0
09
00
N 33 6.5
Unnamed Lake 30°21
0
55
00
N, 84°22
0
54
00
W 30 6.5
Wood Lake 30°01
0
34
00
N, 84°33
0
57
00
W 30 6–8.5
Unnamed Swamp 2 29°49
0
42
00
N, 84°58
0
28
00
W 8 5.5
Unnamed Swamp 3 30°15
0
57
00
N, 84°48
0
53
00
W9 6
Whitehead Lake 30°09
0
54
00
N, 84°40
0
30
00
W 12–29 6–7.5
Hitchcock Lake 30°04
0
54
00
N, 84°39
0
05
00
W30 6
Biodivers Conserv (2009) 18:419–455 441
123

Appendix 1 continued
Collection site Latitude/longitude Temperature (°C) pH
Wright Lake 30°00
0
02
00
N, 85°00
0
08
00
W– 6
Long Bay Swamp 30°19
0
23
00
N, 84°37
0
23
00
W21 5
Swamp at Fort Gadsden 29°55
0
07
00
N, 84°58
0
38
00
W 29 7.5
Camel Pond 30°16
0
36
00
N, 84°59
0
20
00
W 33 5.5
Ocala National Forest
Alexander Springs 29°04
0
52
00
N, 81°33
0
57
00
W 18–27 7
Ocklawaha River 29°22
0
19
00
N, 81°53
0
56
00
W 19–25 7
Juniper Springs 29°10
0
59
00
N, 81°42
0
46
00
W 22–27 6–6.5
Redwater Lake 29°11
0
49
00
N, 81°53
0
28
00
W 35–36 5.5–6.4
Doe Lake 29°02
0
14
00
N, 81°49
0
09
00
W 30–34 5.5–6.3
Mary Lake 29°04
0
23
00
N, 81°49
0
57
00
W 30–37 5–5.7
Lake Dorr 29°00
0
50
00
N, 81°38
0
07
00
W 33–36 5–5.9
Bock Lake 29°05
0
52
00
N, 81°39
0
11
00
W 37 6.5
Faries Prairie 29°06
0
15
00
N, 81°40
0
27
00
W 39 6.5
South Grasshopper Lake 29°08
0
04
00
N, 81°37
0
11
00
W 27–37 6.5
Chain-O-Lake 29°07
0
57
00
N, 81°38
0
36
00
W 17–39 5.5–6.3
Beakman Lake 29°07
0
34
00
N, 81°37
0
14
00
W 36 6.5
Wildcat Lake 29°10
0
14
00
N, 81°37
0
40
00
W35 6
Fore Lake 29°16
0
17
00
N, 81°55
0
03
00
W 18–37 5.5–7
Lake Eaton 29°15
0
18
00
N, 81°51
0
55
00
W 18–36 5.5–7
Half moon Lake 29°09
0
32
00
N, 81°49
0
17
00
W 19–34 5.5–6.9
Mill Dam Lake 29°10
0
42
00
N, 81°50
0
03
00
W 17–34 5.5–6.7
Lake Kerr 29°21
0
19
00
N, 81°48
0
45
00
W 21–39 6.7
Quarry Pond 29°12
0
45
00
N, 81°53
0
45
00
W21 7
Little Lake Kerr 29°20
0
57
00
N, 81°43
0
49
00
W 21–32 7
Unnamed Pond 1 29°07
0
46
00
N, 81°37
0
34
00
W 15 5.5
Lake George 29°12
0
06
00
N, 81°34
0
40
00
W38 7
Unnamed Pond 2 29°06
0
03
00
N, 81°32
0
52
00
W18 7
Clearwater Lake – 35 5.5
Big Cypress National Preserve
Turner River Canal 25°54
0
21
00
N, 81°15
0
43
00
W30 7
Pine Crest well head swamp 25°23
0
27
00
N, 80°47
0
56
00
W27 7
Cypress Swamp 1 25°45
0
37
00
N, 81°00
0
50
00
W30 7
Cypress Swamp 2 25°45
0
36
00
N, 81°02
0
07
00
W25 7
Cypress Swamp 3 25°45
0
37
00
N, 81°03
0
18
00
W25 7
Cypress Swamp 4 25°47
0
01
00
N, 81°05
0
39
00
W25 7
Monument Lake 25°52
0
14
00
N, 81°06
0
51
00
W 20–30 6
Burns Lake 25°53
0
44
00
N, 81°13
0
49
00
W 21–33 7
Roadside Swamp 26°10
0
21
00
N, 81°16
0
00
00
W26 8
East Hanson Swamp 26°11
0
06
00
N, 81°16
0
02
00
W27 7
Tamiami Canal Swamp 1 25°52
0
36
00
N, 81°13
0
40
00
W29 7
Tamiami Canal Swamp 2 – –7
Cypress Swamp 5 25°44
0
51
00
N, 80°57
0
24
00
W18 7
442 Biodivers Conserv (2009) 18:419–455
123

Appendix 1 continued
Collection site Latitude/longitude Temperature (°C) pH
Cypress Swamp 6 25°45
0
37
00
N, 81°02
0
09
00
W– 7
Cypress Swamp 7 25°46
0
53
00
N, 81°05
0
31
00
W20 7
Cypress Swamp 8 25°45
0
03
00
N, 80°58
0
02
00
W25 7
Cypress Swamp 9 25°45
0
16
00
N, 80°58
0
42
00
W 25 6.5
Cypress Swamp 10 25°45
0
37
00
N, 81°02
0
54
00
W 27 6.5
Cypress Swamp 11 25°45
0
37
00
N, 81°03
0
18
00
W 27 6.5
Cypress Swamp 12 25°45
0
45
00
N, 80°55
0
09
00
W30 6
Everglades National Park
Mahagony Hammock 25°19
0
16
00
N, 80°49
0
33
00
W 20–35 7.5
Sisal Pond 25°23
0
28
00
N, 80°47
0
58
00
W 20–25 7
Paurotis Pond 25°16
0
56
00
N, 80°47
0
59
00
W25 7
Long Key Pine Pond 25°24
0
02
00
N, 80°39
0
32
00
W 20–32 6–7
Royal Palm Pond 25°22
0
58
00
N, 80°36
0
36
00
W 20–30 7
Appendix 3 Fungal species found on submerged woody or herbaceous debris in lentic or lotic habitats at
each of the five sites along the Florida Peninsula
Species Collection sites
BW AP OC BC EV
Dothideomycetes
Aliquandostipite minuta Raja & Shearer Le
w
Aliquandostipite crystallinus Raja, Ferrer &
Shearer
Lo
w
Aliquandostipite khaoyaiensis Inderb. Le
w
Aliquandostipite siamensiae (Sivichai &
E.B.G. Jones) J. Campbell, Raja, Ferrer,
Sivichai & Shearer
Le
w
Appendix 2 Ranges in water temperature and pH of all the freshwater habitats sampled during different
seasons at the five collection sites
BW AP OC BC EV
WS WS WS WS WS
Temp 7–14 23–36 8–12 21–35 18–21 25–39 21–30 27–33 20–35 27–35
pH 5–8.5 5–8.5 5–6.5 6–8 7–8
W winter; S summer; Temperature measured in degrees Celsius
BW Blackwater River State Forest, AP Apalachicola National Forest, OC Ocala National Forest, BC Big
Cypress National Preserve, EV Everglades National Park
Biodivers Conserv (2009) 18:419–455 443
123

Appendix 3 continued
Species Collection sites
BW AP OC BC EV
Boerlagiomyces websteri Shearer &
J.L. Crane
Lo
w
Le
w
Le
w
Le
h
Caryospora obclavata Raja & Shearer Le
w
Caryospora langliosii Ell. & Everh. Le
w
Falciformispora lignatilis K.D. Hyde Le
w
Le
w
Jahnula aquatica (Plo
¨
ttner & Kirschst.)
Kirschst.
Lo
w
Jahnula australiensis K.D. Hyde Le
w
Jahnula bipileata Raja & Shearer Le
w
Le
w
Jahnula bipolaris (K.D. Hyde) K.D. Hyde Le
w
Jahnula potamophila K.D. Hyde &
S.W. Wong
Le
w
Jahnula rostrata Raja & Shearer Lo
w
Le
Lo
w
Le
w
Le
w
Jahnula sangamonensis Shearer & Raja Le
w
Lo
w
Kirschsteiniothelia elaterascus Shearer Le
w
Le
w
Le
w
Le
w
Lepidopterella palustris Shearer &
J.L. Crane
Le
Lo
w
Lepidopterella tangerina Raja & Shearer Lo
w
Massarina bipolaris K.D. Hyde Le
w
Le
w
Le
w
Massarina fronsisubmersa K. D. Hyde Lo
w
Massarina ingoldiana Shearer & K.D. Hyde Le
w
Le
Lo
w
Micropeltopsis sp. F115 Le
w
Ophiobolus shoemakeri Raja & Shearer Le
h
Trematosphaeria lineolatispora K.D. Hyde Lo
w
Lo
Le
w
Le
h
Undescribed Massarina sp. F60 Le
Lo
w
444 Biodivers Conserv (2009) 18:419–455
123

Appendix 3 continued
Species Collection sites
BW AP OC BC EV
Undescribed Massarina sp. F65 Le
w
Undescribed Massarina sp. F80 Le
w
Undescribed genus F76 Lo
w
Lo
h
Leotiomycetes
Aquapoterium pinicola Raja & Shearer Le
Lo
h
Le
h
Le
h
Gorgoniceps sp. F72 Le
w
Sordariomycetes
Aniptodera aquadulcis (S.Y. Hsieh,
H.S. Chang & E.B.G. Jones) J. Campb.,
J. Anderson & Shearer
Le
w
Aniptodera chesapeakensis Shearer & Miller Le
w
Aniptodera inflatiascigera K.M. Tsui,
K.D. Hyde & I.J. Hodgkiss
Le
w
Le
h
Le
w
Aniptodera lignatilis K.D. Hyde Lo
w
Le
w
Lo
w
Aniptodera megaloascocarpa Raja &
Shearer
Le
w
Aniptodera palmicola K.D. Hyde, W.H. Ho
& K.M. Tsui
Le
Lo
w
Le
h
w
Le
w
Annulatascus velatisporus K.D. Hyde Le
Lo
w
Le
w
Le
h
Le
Lo
w
Arnium gigantosporum Raja & Shearer Le
w
Ascitendus austriacus (Re
´
blova
´
, Winka &
Jaklitsch) J. Campb. & Shearer
Lo
Le
h
w
Le
w
Ascosacculus heteroguttulatus (S.W. Wong,
K.D. Hyde & E.B.G. Jones) J. Campb.,
J.L. Anderson & Shearer
Le
w
Le
Lo
w
Le
w
Le
h
Ascotaiwania hsilio H.S. Chang &
S.Y. Hsieh
Lo
w
Le
w
Ayria appendiculata Fryar & K.D. Hyde Le
h
Cataractispora bipolaris (K.D. Hyde)
K.D. Hyde, S.W. Wong & E.B.G. Jones
Le
w
Cyanoannulus petersenii Raja, J.Campb. &
Shearer
Le
Lo
w
Lo
w
Biodivers Conserv (2009) 18:419–455 445
123

Appendix 3 continued
Species Collection sites
BW AP OC BC EV
Fluviatispora reticulata K.D. Hyde Le
Lo
w
Le
Lo
w
Le
w
Flammispora pulchra Raja & Shearer Le
w
Hanliniomyces hyaloapicalis Raja &
Shearer gen. nov
Lo
w
Le
w
Jobellisia luteola (Ellis & Everh.) M.E. Barr Le
Lo
w
Le
Lo
w
Lockerbia striata Raja & Shearer Le
w
Le
h
Luttrellia estuarina Shearer Lo
w
Luttrellia sp. F14 Le
w
Lulworthia sp. F54 Le
w
Nais inornata Kohlm. Le
w
Le
Lo
w
Le
Lo
w
Le
w
Natantispora retorquens (Shearer &
J.L. Crane) J. Campb.,
J.L. Anderson et Shearer
Lo
Le
w
Le
w
Ophioceras commune Shearer, J.L. Crane &
Chen
Le
w
Le
Lo
w
Phaeonectriella lignicola R.A. Eaton &
E.B.G. Jones
Lo
w
Phomatospora triseptata Raja & Shearer Le
w
Pseudoproboscispora caudae-suis (Ingold)
J. Campbell & Shearer
Lo
w
Physalospora limnetica Raja & Shearer Le
h
Savoryella aquatica K.D. Hyde Le
w
Le
w
Savoryella fusiformis W.H. Ho, K.D. Hyde
& I.J. Hodgkiss
Le
w
Submersisphaeria aquatica K.D. Hyde Le
Lo
w
Le
Lo
w
Le
w
Torrentispora fibrosa K.D. Hyde, W.H. Ho,
E.B.G. Jones, K.M. Tsui & S.W. Wong
Lo
w
Lo
w
Le
w
Vertexicola caudatus K.D. Hyde,
S.W. Wong & V.M. Ranghoo
Lo
w
Le
w
446 Biodivers Conserv (2009) 18:419–455
123

Appendix 3 continued
Species Collection sites
BW AP OC BC EV
Zopfiella latipes (N. Lundq.)
Malloch & Cain
Le
Lo
w
Le
w
Le
w
Zopfiella lundqvistii Shearer & J.L. Crane Le
Lo
w
Unidentified ascomycete sp. F28 Le
w
Undescribed sp. F41 Le
w
Undescribed gen. F44 Le
w
Undescribed gen. F50 Lo
w
Undescribed sp. F57 Lo
Le
w
Undescribed gen. F99 Le
w
Undescribed sp. F100 Le
w
Undescribed sp. F107 Le
w
BASIDIOMYCETES
Rogersiomyces okefenokeensis J.L. Crane &
Schoknecht
Le
h
Mitosporic fungi
Acrogenospora sphaerocephala (Berk. and
Broome) M. B. Ellis
Lo
w
Le
w
Lo
w
Le
w
Ardhachandra cristaspora (Matsush.)
Subram. & Sudha
Le
h
Bactrodesmium linderi (J.L. Crane &
Shearer) M.E. Palm & E.L. Stewart
Le
Lo
w
Le
w
Le
w
Le
h
Berkleasmium concinnum Berk. (S. Hughes) Lo
w
Brachiosphaera tropicalis Nawawi Le
w
Brachysporium obovatum Kiessl Le
w
Cancellidium applanatum Tubaki Le
Lo
h
w
Le
Lo
w
Le
w
Canalisporium caribense (Hol.-Jech. &
Mercado) Nawawi & Kuthub.
Le
w
Biodivers Conserv (2009) 18:419–455 447
123

Appendix 3 continued
Species Collection sites
BW AP OC BC EV
Canalisporium kenyense Goh, W. H. Ho,
K. D. Hyde, S. R. Whitton & T.E. Umali
Le
w
Cacumisporium sigmoideum Mercado &
R. F. Castan
˜
eda
Lo
w
Le
w
Chaetospermum camelliae Agnihothr. Le
h
Le
h
Le
h
w
Coleodictyospora micronesia (Matsush.)
Matsush.
Le
h
Le
h
Cordana abramovii Seman & Davydkina
var. seychellensis K. D. Hyde & Goh
Le
w
Condylospora sp. FH34 Le
w
Cryptophiale cucullata Kuthub. Le
h
Dactylaria hyalotunicata K.M. Tsui, Goh &
K.D. Hyde
Lo
h
w
Le
w
Dactylaria tunicata Goh & K.D. Hyde Le
Lo
w
Le
w
Delortia palmicola Pat. Le
h
w
Dendrosporium lobatum Plakidas &
Edgerton ex J.L. Crane
Le
h
Dendrospora sp. FH87 Le
w
Dictyosporium sp. FH39 Le
w
Dictyosporium digitatum J.L. Chen,
C.H. Hwang & S.S. Tzean
Le
h
w
Le
h
Le
h
Dictyosporium elegans Corda Lo
w
Dictyosporium giganticum Goh &
K.D. Hyde
Le
h
Dictyosporium heptasporum (Garov.)
Damon
Le
w
Ellisembia abscendens (Berk.) Subram. Lo
w
Le
Lo
w
Le
w
Exserticlava globosa Roa de Hoog Le
w
Exserticlava triseptata (Matsush.) S. Hughes Le
w
Le
w
448 Biodivers Conserv (2009) 18:419–455
123

Appendix 3 continued
Species Collection sites
BW AP OC BC EV
Exserticlava vasiformis (Matsush.)
S. Hughes
Lo
Le
w
Le
w
Helicodendron sp. FH38 Le
Lo
w
Helicoon auratum (Ellis) Morgan Le
w
Helicomyces roseus Link Le
Lo
w
Le
w
Helicosporium aureum (Corda) Linder Le
w
Helicosporium guianense Linder Le
w
Helicosporium gigasporum K.M. Tsui, Goh,
K.D. Hyde & Hodgkiss
Lo
w
Le
Lo
w
Helicosporium lubricopsis Linder Le
w
Humicola asteroidea Udagawa & Y. Horie Lo
w
Ingoldiella hamata D.E. Shaw Lo
h
Intercalarispora sp. FH88 Lo
w
Monacrosporium ellipsosporum (Preuss)
R.C. Cooke & C.H. Dickinson
Le
h
w
Monotosporella setosa
(Berk. & M.A. Curtis) S. Hughes
Lo
Le
w
Le
w
Melanocephala australiensis
G.W. Beaton & M.B. Ellis
Lo
w
Nawawia filiformis (Nawawi) Marvanova
´
Le
w
Pleurophragmium malaysianum Matsush. Le
Lo
w
Le
w
Le
w
Pleurothecium recurvatum (Morgan) Ho
¨
hn. Le
w
Septonema hormiscium Sacc. Le
h
Le
w
Speiropsis sp. FH59 Le
h
Sporidesmiella hyalosperma var.
novae-zealandiae (S. Hughes) P.M. Kirk
Le
w
Biodivers Conserv (2009) 18:419–455 449
123

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