Chaetothiersia vernalis, a new genus and species of Pyronemataceae (Ascomycota, Pezizales) from California

Abstract

Chaetothiersia vernalis, a new genus and species of Pyronemataceae (Ascomycota, Pezizales) from California. Fungal Diversity 28: 65-72. Chaetothiersia vernalis, collected from the northern High Sierra Nevada of California, is described as a new genus and species. This fungus is characterized by stiff, superficial, brown excipular hairs, smooth, eguttulate ascospores, and a thin ectal excipulum composed of globose to angular-globose cells. Phylogenetic analyses of nLSU rDNA sequence data support the recognition of Chaetothiersia as a distinct genus, and suggest a close relationship to the genus Paratrichophaea.

Full text

Fungal Diversity
65
Chaetothiersia vernalis, a new genus and species of Pyronemataceae (Ascomycota,
Pezizales) from California
Perry, B.A.1* and Pfister, D.H.1
Department of Organismic and Evolutionary Biology, Harvard University, 22 Divinity Ave., Cambridge, MA 02138,
USA
Perry, B.A. and Pfister, D.H. (2008). Chaetothiersia vernalis, a new genus and species of Pyronemataceae
(Ascomycota, Pezizales) from California. Fungal Diversity 28: 65-72.
Chaetothiersia vernalis, collected from the northern High Sierra Nevada of California, is described as a new genus and
species. This fungus is characterized by stiff, superficial, brown excipular hairs, smooth, eguttulate ascospores, and a
thin ectal excipulum composed of globose to angular-globose cells. Phylogenetic analyses of nLSU rDNA sequence
data support the recognition of Chaetothiersia as a distinct genus, and suggest a close relationship to the genus
Paratrichophaea.
Keywords: discomycetes, molecular phylogenetics, nLSU rDNA, Sierra Nevada fungi, snow bank fungi, systematics
Article Information
Received 31 January 2007
Accepted 19 December 2007
Published online 31 January 2008
*Corresponding author: B.A. Perry; e-mail: brian_perry@post.harvard.edu
Introduction
During the course of our recent investi-
gation of the phylogenetic relationships of
Pyronemataceae (Perry et al., 2007), we
encountered several collections of an appa-
rently undescribed, operculate discomycete
from the northern High Sierra Nevada of
California. Macroscopically, the taxon repre-
sented by these collections might be referred to
genera of the Pyronemataceae such as
Trichophaea Boud., Trichophaeopsis Korf &
Erb, Humaria Fuckel, Geopora Harkn.,
Tricharina Eckblad, or Wilcoxina Chin S.
Yang & Korf. These genera are characterized
by medium- to large-sized apothecia with often
pale hymenia and erect, pigmented marginal
and superficial excipular hairs. Traditionally,
these genera have been distinguished based
upon differences in ascospore ornamentation,
guttulation and reaction to various stains and
media, as well as differences in the size and
nature of the apothecial hairs and composition
of the excipular tissues. Micromorphological
examination of our collections, however,
indicates that this taxon does not fit well within
the limits of any of the described genera
currently recognized in the family (Eriksson,
2006), and requires the erection of a new
genus. We herein propose the new genus and
species, Chaetothiersia vernalis, to accommo-
date this taxon.
The results of our previous molecular
phylogenetic analyses of the family (Perry et
al., 2007) suggest that this new genus and
species are closely related to Paratrichophaea
Trigaux, but fail to resolve this relationship
with significant bootstrap support. In that
study, which encompassed a broad sampling of
taxa across the order Pezizales, several regions
of the dataset were deemed ambiguous and
therefore excluded from the analyses. In an
attempt to further elucidate the relationships of
this new genus and species within the family,
we have analyzed an nLSU rDNA dataset
focusing on the clade in which this new genus
was previously resolved, and including
additional macromorphologically similar
genera, from which no nucleotide regions have
been excluded (Table 1).

66
Table 1. List of taxa included in this study. Numbers in parentheses indicate multiple collections of a single taxon.
Species Collection Geographic Origin Collector
GenBank
Access. no.
Chaetothiersia vernalis B.A. Perry & Pfister BAP 492 (HOLOTYPE, FH) USA, California J. Laws DQ220403
Chaetothiersia vernalis HDT 53173 (SFSU) USA, California H.D. Thiers DQ220400
Chaetothiersia vernalis DHP & HDT 5.18.86 (FH) USA, California H.D. Thiers, D.H. Pfister DQ220401
Genea arenaria Harkn. Trappe 17288 (FH, dupl. OSC) USA, California J. Graham DQ220332
Genea harknessii Gilkey Trappe 13313 (FH, dupl. OSC) USA, Washington A. & D. Claridge DQ220334
Geopora arenicola (Lév.) Kers KS-94-173 (C) Denmark K. Hansen, S.K. Sandal DQ220336
Geopora cooperi Harkn. HDT 52489 (SFSU) USA, Wyoming J. Ammarati DQ220341
Humaria hemisphaerica (F.H. Wigg.) Fuckel BAP 320 (FH) China, Tibet B.A. Perry DQ220352
Humaria velenovskyi (Vacek ex Svrček) Korf & Sagara HK 24-IX-1975 (C) Denmark H. Knudsen DQ220354
Lasiobolidium spirale Malloch & Cain CBS 782.70 USA, Wyoming R. F. Cain DQ220363
Melastiza contorta (Massee & Crossl.) Spooner & Y.J. Yao KH.01.06 (C) Sweden B.T. Olsen AY500539
Parascutellinia carneosanguinea (Fuckel) T. Schumach. KH.03.34 (FH) Norway K. Hansen, C. Lange DQ220388
Paratrichophaea boudieri (Grélet) Bronkers BAP 481 (FH) USA, California B.A. Perry DQ220402
Pseudaleuria quinaultiana Lusk NSW 7107 USA, Oregon N. S. Weber DQ220389
Pyronema omphalodes (Bull.) Fuckel TL-11685 (QCNE, C) Ecuador, Carchi K. Hansen et al. DQ220397
Spooneromyces laeticolor (P. Karst.) T. Schumach. & J. Moravec C F-48310/HFG 88.013 (C) Denmark H.F. Gøtzsche DQ220434
Tricharina gilva (Boud. ex Cooke) Eckblad BAP 431 (FH) USA, California B.A. Perry DQ220444
Tricharina ochroleuca (Bres.) Eckblad C F-53062/HD Gr83.107 (C) Greenland H. Dissing DQ220445
Trichophaea abundans (P. Karst.) Boud. CBS 348.76 Finland V. Hintikka DQ220448
Trichophaea hemisphaerioides (Mouton) Graddon KH.97.31 (FH) USA, California K. Hansen DQ220457
Trichophaea hybrida (Sowerby) T. Schumach. (1) DHP 30.VIII.2000 (FH) USA, Vermont D.H. Pfister DQ220453
Trichophaea hybrida (2) AMNH-49682/F-17491 (AMNH) Iceland G. G. Eyjólfsdóltir DQ220455
Trichophaea woolhopeia (Cooke & W. Phillips) Arnould BAP 453 (FH) Norway D.H. Pfister, B.A. Perry DQ220459
Trichophaeopsis bicuspis (Boud.) Korf & Erb NSW 8316 (OCS) USA, Oregon N. S. Weber DQ220461
Trichophaeopsis tetraspora Dissing & M.D. Paulsen C F-47525 (C) Denmark H. Dissing DQ220463
Wilcoxina mikolae (Chin S. Yang & H.E. Wilcox) Chin S. Yang & Korf WS 36 (SFSU) USA, Wyoming W. Stoll DQ220468
Wilcoxina sp. ITS RFLP RPC-10 USA, California — AF156926

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Materials and Methods
Morphological observations
Macroscopic descriptions were compiled
from field notes taken on the characters of
fresh material for collections DHP & HDT
May 1986 (FH) and BAP 492 (FH). Color
terms and notations refer to those of Kornerup
and Wanscher (1981). For micromorphological
investigation, small pieces of dried apothecia
were excised with a double-edged razor blade
and allowed to rehydate in distilled H2O for
approximately 12 hours. Once fully rehydrated,
excised portions were sectioned on a freezing
microtome at a thickness of 20-35 µm, or used
to prepare squash mounts. Sections were
subsequently mounted in H2O, congo red or
cotton blue in lactic acid and viewed on a
compound microscope utilizing both bright
field and diffusion interference contrast (DIC)
optics. All measurements were made from
sections and squash mounts in H2O. To
adequately sample the range of ascospore
dimensions, mature ascospores from each
collection were measured at 1000X and the
values pooled to determine an overall mean
ascospore size. Photomicrographs were taken
with a 35 mm camera body attached to an
Olympus BX 60 microscope.
Molecular methods and analyses
DNA extraction, PCR and cycle
sequencing protocols followed those outlined
in Perry et al. (2007). The nLSU rDNA was
symmetrically amplified using primers LROR
and LR5 or LR7 (Moncalvo et al., 2000). In
addition to these, internal primers LR3 and
LR3R (Moncalvo et al., 2000) were also used
for sequencing. Cleaned sequencing reactions
were visualized on an ABI 3100 or 3730
Genetic Analyzer capillary sequencer (Applied
Biosystems). Sequences were edited and
assembled using the software package
Sequencher 3.0 or 4.0 (Gene Codes Corp, Ann
Arbor, MI). Sequences were aligned manually
using MacClade 4 (Maddison and Maddison,
2000), Se-Al v.2.0 (Rambaut, 1996), and the
editor window of PAUP* 4.0b4a (Swofford,
2003). Edited sequences have been deposited in
GenBank (Table 1; http://www.ncbi.nlm.
nih.gov/), and the aligned dataset is available
via TreeBASE (http://www.treebase.org;
S1764, M3216).
Parsimony analyses were conducted
using PAUP* 4.0 (Swofford, 2003). Searches
used a branch-and-bound algorithm with
furthest sequence addition, MulTrees on,
collapse of zero length branches and equal
weighting of all characters. Nodal support of
individual clades was assessed by bootstrap
analyses (Felsenstein, 1985), using 1000
heuristic replicates, each consisting of 10
random addition sequences, with stepwise
addition and tree bisection reconnection (TBR)
branch swapping. Maximum likelihood (ML)
searches were also conducted using PAUP*,
under a GTR + I + G model of sequence
evolution determined with the Akaike
Information Criterion as calculated in the
program Modeltest v.3.7 (Posada & Crandall,
1998). Parameter values were estimated from
an initial tree generated using a simple Jukes-
Cantor model. The resulting parameters were
then fixed, and a search consisting of 500
heuristic replicates (random sequence addition,
TBR branch swapping, collapse of zero length
branches) was performed under the more
complex model of sequence evolution as
determined above. Clade support was assessed
by nonparametric ML bootstrap analyses as
implemented in the program PhyML (Guindon
and Gascuel, 2003), and consisted of 1000
replicates using the same model of sequence
evolution as the ML searches with all
parameters estimated by the program. Bayesian
analyses were performed using Metropolis-
coupled Markov chain Monte Carlo
(MCMCMC) methods as implemented in
MrBayes 3.1.2 (Huelsenbeck and Ronquist,
2001; Ronquist and Huelsenbeck, 2003) using
the same model as the ML analyses. Bayesian
analyses consisted of two parallel searches, run
for 5 million generations and initiated with
random starting trees. Chains were sampled
every 500 generations for a total of 10,000
trees each, sampled from the posterior
distribution. Those trees sampled prior to the
runs reaching a split deviation frequency of
0.01 were discarded from the sample as the
“burn-in,” while the remaining trees were used
to calculate the posterior probabilities of the

68
Fig. 1. Single parsimony tree of Chaetothiersia and other genera of Pyronemataceae inferred from nLSU sequence data
(568 steps, CI = 0.609, RI = 0.666). Numbers separated by / represent parsimony bootstrap proportions, ML non-
parametric bootstrap proportions, and Bayesian posterior probabilities greater than 70%, respectively (- designates a
value below 70%). Numbers isolated below branches refer to Bayesian posterior probabilities, with associated bootstrap
values located above the branch.
individual clades. The default settings were
used in MrBayes to set the incremental heating
scheme (i.e., 3 heated chains to one cold
chain), unconstrained branch lengths [uncon-
strained: exponential (10.0)] and uninformative
topology (uniform) priors.
Results
Phylogenetic analyses
The nLSU dataset consists of 834 aligned
positions for 26 ingroup taxa, and contains 155
parsimony-informative positions. The parsi-
mony analysis recovered a single tree of 568
steps (Fig. 1) similar to the tree produced by
the ML analysis (-lnL = 3884.24117, not
shown), differing only in the placement of
Lasiobolidium and Trichophaea hemisphae-
rioides within the larger clade. Bayesian
analyses reached an average standard deviation
of split frequencies below 0.01 after appro-
ximately 250,000 generations, and the first 500
trees were excluded as the “burn-in.” The three
collections of Chaetothiersia vernalis are
highly supported as a monophyletic group, but
weakly supported by bootstrap analyses as the
sister group to Paratrichophaea boudieri.
These taxa are part of a large, weakly

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Figs 2-4. Chaetothiersia vernalis (BAP 492, Holotype, FH). 2. Apothecia on bark of Abies magnifica, × 1.1. 3. Light
microscopic slide of superficial hairs arising from ectal excipulum of the apothecial margin, × 400. 4. Light microscopic
slide of ectal excipulum demonstrating the globose to angular-globose cells, formation of pustules, and the bases of two
excipular hairs arising from this tissue layer, × 200. Photographs: B. A. Perry.
supported clade containing species of genera
macromorphologically similar to Chaetothi-
ersia, including Genea, Trichophaea, Humaria,
Trichophaeopsis and Wilcoxina. The two
species of Genea sampled form a well
supported monophyletic group, as do both
Trichophaeopsis and Wilcoxina. Humaria is
non-monophyletic, with H. hemisphaerica
forming the sister group to Genea, and H.
velenovskyi grouping with Spooneromyces
laeticolor. Taxa from Tricharina and Geopora,
also morphologically similar to Chaetothiersia,
form a separate, moderately supported clade
with Trichophaea abundans and T.
woolhopeia, rendering the later genus non-
monophyletic. The two species of Geopora
sampled, G. arenicola and G. cooperi, form a
well supported monophyletic group, but
Tricharina does not.
Chaetothiersia vernalis B.A. Perry & Pfister,
gen. et spec. nov. (Figs 2-4)
MycoBank: 510434.
Etymology: Chaeto (Latin) indicating the long
hairs of the excipulum, and thiersia in reference to and
in honor of Dr. Harry D. Thiers (1919-2000), professor
of Biology and curator of mycological collections at San
Francisco State University (H.D. Thiers Herbarium), and
one of the first to collect specimens of this new genus
and species; vernalis (Latin) pertaining to Spring; in
reference to the fruiting of this species during the Spring
months in the High Sierra Nevada.
Apothecia cupulatum vel disciforme, pallide
canum ad cretaceum, margo distinctus; externus pilosus,
pilus superficialis, rigidus ad curvus, hyalinus vel plus
saepe fuscus, obtusus. Excipulum medullare crassum,
densum, texturae intricatae; excipulum ectale tenue,
perexiguum, cellulae globosae vel angulara-globosae.
Asci octosporae, non-amyloideus, operculate.
Ascosporae leaves, ellipsoidia, eguttulatae, hyalinae.
Paraphyses simplices, angustae, rectae, septatae.
Gregarium, lignatile.

70
Holotypus: BAP 492 (FH), Sierra Co., California,
USA.
Apothecia 5-30 mm diam, deep cupulate
with a narrowed base at first, expanding to
discoid and tending to become ±convoluted,
larger forms often appressed to the substrate.
Hymenium pale gray (2-3A3) to nearly white,
smooth; margin distinct, covered with erect,
dark brown hairs that are often borne in
fascicles. Receptacle ± concolorous with hyme-
nium, densely to sparsely covered with dark to
pale brown, ± appressed and occasionally
projecting hairs. Ectal excipulum only a few
cells thick, up to 40 µm, extending beyond
hymenium to form distinct margin, composed
of angular-globose to broadly clavate cells 4.8-
16 µm diam, tending to form small irregular
pustules. Medullary excipulum thicker, up to
840 µm in sections 1-2 mm inwards from
margin, composed of densely compacted
textura intricata, cells 1.6-6.4 µm diam.
Excipular hairs 60-720 × 4-14.4 µm, brown to
nearly hyaline, multiseptate, thick-walled (up
to 2.4 µm), apically rounded, arising from
angular-globose cells of the ectal excipulum,
those near the margin typically straight and
projecting, those of the receptacle often flexous
and quite narrowed apically (~2 µm),
interspersed with shorter, broader forms.
Hymenium 300-400 µm thick. Asci 8-spored,
304-380 × 8-16 µm, operculate, cylindrical, J–,
arising from a pleurorhynchous base.
Ascospores 16.8-18.4(-19.2) × 10-11.2 (-12)
µm (
x
= 17.8 × 10.6 µm, n = 60 spores),
ellipsoid, smooth, eguttulate, hyaline, not
refractive in media containing lactic acid, with
granular cytoplasmic contents, often collapsing
in media other than H2O. Paraphyses narrow,
1.6-2.4 µm diam, equal to barely expanding
apically, straight, multiseptate, unbranched,
hyaline, exceeding asci by up to 10.4 µm.
Habit and habitat: Gregarious to caespi-
tose on decaying wood and bark of conifers
(Abies magnifica A. Murray), and on woody
debris in soil.
Known distribution: Northern High
Sierra Nevada mountains, California, U.S.A.,
April to June.
Material examined: USA. California: El Dorado
County, Crystal Basin Recreation Area, caespitose on
woody debris in soil under conifers, 4 May 1974, HDT
32218, coll. by Harry D. Thiers (SFSU). El Dorado
County, Silver Fork Road, near China Flat Campground,
elev. ca. 1525 m, on woody debris in soil and rotting
wood, 26 April 1986, HS 3068, coll. by Herb Saylor
(SFSU). Sierra County, Highway 49, 3 miles east of
Yuba Pass, solitary on woody debris in soil, 15 June
1983, HDT 45925, coll. by Harry D. Thiers (SFSU).
Sierra County, Highway 49, “Steele Gulch,” elev. ca.
1770 m, on conifer log, 18 May 1986, coll. by Donald H.
Pfister & Harry D. Thiers (FH, paratype). Sierra County,
Highway 49, Yuba Pass, elev. ca. 2400 m, on woody
debris in soil under conifer bark, 6 June 1990, HDT
53173, coll. by Harry D. Thiers (SFSU, paratype). Sierra
County, Highway 49, Yuba Pass, elev. ca. 2400 m, on
backside of bark on decaying stump of Abies magnifica,
2 June 2003, BAP 492, coll. by Jack Laws (FH,
holotype). Tuolumne County, Pinecrest Lake, on woody
debris near dead logs, 1 April 1972, HDT 28743, coll. by
Harry D. Thiers (SFSU). Tuolumne County, Pinecrest
Lake, on woody debris near melting snow, 1 April 1972,
HDT 28744, coll. by Harry D. Thiers (SFSU).
Discussion
Chaetothiersia vernalis is characterized
by the following combination of features:
smooth, eguttulate, ellipsoid ascospores,
brown, non-rooting, stiff superficial excipular
and marginal hairs with obtuse apices; a dense
medullary excipulum composed of textura
intricata; a very thin ectal excipulum of
globose to angular-globose cells; medium to
rather large apothecia (relative to other
members of Pyronemataceae); and growth on
wood and woody debris associated with
moisture produced by the Spring snowmelt in
the High Sierra Nevada. Our morphological
observations, as well as the results of the
phylogenetic analyses, indicate a close
relationship between Chaetothiersia and
Paratrichophaea. These genera are quite
similar in the presence of smooth, ellipsoid
ascospores and long, projecting excipular hairs.
However, the hairs of Paratrichophaea are
typically rooting, arising from the medullary
excipulum, are apically acuminate, and often
bifurcating basally. The hairs of Chaetothiersia
are superficial in nature, arising from the cells
of the thin ectal excipulum, obtuse apically,
and have not been observed to branch basally.
Additionally, the apothecia of the Chaetothi-
ersia attain sizes quite a bit larger than those
typically reported for Paratrichophaea (1-4
mm). Paratrichophaea appears to be a rather
rare or easily overlooked genus, and we have
sampled only a single species in our phylo-
genetic analyses. Aside from the original

Fungal Diversity
71
description of the genus by Trigaux (1985),
three species of Paratrichophaea have been
treated from North America by Pfister (1988),
and more recently Bronkers (2003) has treated
two of the same species from Europe. An
additional taxon, Cheilymenia albescens
Dissing et Raitv., was transferred to Paratri-
chophaea by Schumacher (1988) due to the
obvious affinities this species has with the
other members of the genus.
Additional genera that may superficially
resemble Chaetothiersia include Trichophaea,
Trichophaeopsis, Tricharina, Geopora,
Humaria and Wilcoxina. Of these genera, both
Humaria and some Trichophaea species can be
distinguished from Chaetothiersia by the
presence of ornamented, guttulate spores. The
smooth-spored species of Trichophaea, such as
T. woolhopeia and T. abundans, are also
characterized by guttulate spores. Similarly,
Geopora is also characterized by ascospores
containing 1-2 large guttules, and produces
generally subhypogeous apothecia. Wilcoxina
species have eguttulate spores, hair and
excipular structure similar to Chaetothiersia,
but can be distinguished by generally smaller
ascospores (up to 9.5 × 16), which are yellow-
refractive when mounted in media containing
lactic acid. Additionally, the apothecia of
Wilcoxina are often orange to ochraceous, and
with the exception of W. sequoia (W. Phillips)
T. Schumach., which is reported to grow on
bark, foliage and decayed wood, the apothecia
of most species occur on soil. The eguttulate
ascospores of Tricharina are also often yellow-
refractive in media containing lactic acid, and
unlike Chaetothiersia, typically occupy more
than half to nearly the entire length of the
ascus. The excipular hairs of Tricharina are
also generally restricted to the margin, and the
apothecia are not known to occur on woody
substrates. Trichophaeopsis species generally
produce smaller apothecia (up to 4 mm diam.)
than those of Chaetothiersia, and can be
distinguished by the presence of ectal cells
arranged in distinct vertical rows, and often
forked excipular hairs which tend to have a
long branch pointing upwards, relative to the
orientation of the apothecium, and a shorter
branch pointing down.
Phylogenetic analyses of the nLSU data
support the recognition of Chaetothiersia as a
distinct genus in the Pyronemataceae, and
confirm, with weak bootstrap support, our
morphological observations suggesting a sister
relationship between this genus and
Paratrichophaea. The relationships of Chaeto-
thiersia to other morphologically similar
genera of the family, however, are not resolved
with high support. Chaetothiersia and
Paratrichophaea are part of a clade containing
several genera with morphologically similar
features, but this clade is not well supported.
With the exception of Trichophaea, Humaria
and Tricharina, the other genera sampled by
more than one taxon all form well supported
monophyletic groups. In agreement with our
previous investigation (Perry et al., 2007), the
species of Trichophaea sampled are nested in
two independent clades. Trichophaea
woolhopeia, the type of the genus, and T.
abundans, which are both smooth-spored
species, are isolated from the rough-spored
species T. hybrida and T. hemisphaerioides.
Humaria displays a similar pattern, with the
brightly pigmented H. velenovskyi forming the
sister group to Spooneromyces laeticolor.
These results suggest that the generic
placement of both H. velenovskyi and the
rough-spored species of Trichophaea are in
need of reconsideration (Perry et al., 2007).
Also in agreement with our previous analyses,
the phylogenetic delimitation of Tricharina
from Geopora remains unclear, indicating that
the limits of these genera need to be re-
examined.
Acknowledgements
We would like to thank Dennis E. Desjardin,
curator of the herbarium at SFSU, for the loan of
specimens HDT 28743, 28744, 32218, 45925, 53173,
and HS 3068, used in this study. We would also like to
thank Genevieve Lewis-Gentry for her invaluable
assistance with the Latin diagnosis. This research was
funded in part by NSF Grant DEB-0315940 to DHP and
Karen Hansen, and a Harvard University research grant
to BAP.
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