In Press at Mycologia, preliminary version published on October 25, 2012 as doi:10.

3852/12-137

Short title: Phialemoniopsis gen. nov. and new species of Phialemonium and Lecythophora

Phialemoniopsis, a new genus of Sordariomycetes, and new species of Phialemonium and

Lecythophora

Haybrig Perdomo

Dania García

Josepa Gené1

Josep Cano

Mycology Unit, Medical School and IISPV, Universitat Rovira i Virgili, Sant Llorenç 21,

43201-Reus, Spain

Deanna A. Sutton

Fungus Testing Laboratory, Department of Pathology, University of Texas Health Science

Center, 7703 Floyd Curl Drive, San Antonio, Texas 78229-3900

Richard Summerbell

Sporometrics, Inc. 219 Dufferin Street, Suite 20C, Toronto, Ontario, M6K 1Y9, and Dalla

Lana School of Public Health, University of Toronto, 223 College Street, Toronto Ontario,

M5T 1R4

Josep Guarro

Mycology Unit, Medical School and IISPV, Universitat Rovira i Virgili, Sant Llorenç 21,

43201-Reus, Spain

Abstract: In molecular studies involving numerous clinical isolates of the genera

Acremonium, Phialemonium and Lecytophora some of them could not be identified. To

clarify the phylogenetic relationships among these fungi and other related taxa, we performed

a polyphasic study based on a detailed morphological study and on the analysis of sequences

of four loci: the internal transcribed spacer regions, the D1/D2 domains of the 28S rRNA,

actin and β-tubulin genes. The combination of the resulting data let us propose the new genus

Copyright 2012 by The Mycological Society of America.

Phialemoniopsis to accommodate the opportunistic fungi Phialemonium curvatum and

Sarcopodium oculorum and two new species, Phialemoniopsis cornearis and Phialemoniopsis

pluriloculosa. The taxonomy of Phialemoniopsis has not been completely resolved, however,

remaining incertae sedis within the Sordariomycetes. In addition, the new species

Lecythophora luteorubra, Lecythophora cateniformis and Phialemonium globosum are

described and the species Acremonium atrogriseum and Taifanglania inflata are transferred to

the genus Phialemonium. Lecythophora and Phialemonium are currently monophyletic genera

of the families Coniochaetaceae (Coniochaetales) and Cephalothecaceae (Sordariales)

respectively, qaccording to our results. Tables summarizing key morphological features to

distinguish the current species of Lecythophora, Phialemonium and Phialemoniopsis are

provided.

Key words: Cephalotheca, Coniochaeta, Lecythophora, morphology, multilocus

sequencing, Phialemonium, Phialemoniopsis, Taifanglania, taxonomy

INTRODUCTION

Phialemonium W. Gams & McGinnis is an anamorphic genus with species widely distributed

and commonly isolated from air, soil, industrial water and sewage (Gams and McGinnis

1983). They also are involved in human infections, affecting both immunocompromised and

immunocompetent patients (King et al. 1993, Guarro et al. 1999, de Hoog et al. 2000, Proia et

al. 2004, Scott et al. 2004, Rivero et al. 2009). Phialemonium, typified by P. obovatum W.

Gams & McGinnis, was described by Gams and McGinnis (1983) to accommodate

filamentous fungi with morphological features intermediate between Acremonium Link and

Phialophora Medlar. The species of Phialemonium are characterized by moist, flat, never

pink colonies and abundant adelophialides (reduced phialides that are not delimited from

subtending intercalary hyphal cell by a basal septum), although short discrete phialides

without a collarette also are present. The phialides form unicellular, hyaline conidia
aggregated in slimy heads (Gams and McGinnis 1983). Currently, the genus Phialemonium

comprises two species, P. obovatum and P. curvatum W. Gams & W.B. Cooke.

Phialemonium obovatum is characterized by greenish colonies, discrete phialides often

sympodially proliferating and obovate conidia with an apiculate and minutely truncate base

(Gams and McGinnis 1983). Phialemonium curvatum is characterized by ochraceous or

grayish colonies, non-proliferating phialides and cylindrical to allantoid conidia (Gams and

McGinnis 1983, King et al. 1993, de Hoog et al. 2000). Some authors have described the

presence of sporodochium-like conidiomata in P. curvatum, a character not reported in the

original description (Proia et al. 2004, Strahilevitz et al. 2005, Weinberger et al. 2006, Sutton

et al. 2008, Perdomo et al. 2011b).

Yaguchi et al. (2006) described the new species Cephalotheca foveolata Yaguchi,

Nishim. & Udagawa from a patient with a subcutaneous infection, which showed a

Phialemonium-like anamorph. In the same study, analysis of the LSU rDNA sequences

demonstrated a close relationship between P. obovatum and members of the family

Cephalothecaceae (Sordariales), suggesting that the anamorphs of Cephalotheca Fuckel and

Albertiniella Kirschst. could belong to Phialemonium. However, the placement of P.

curvatum into Cephalothecaceae was not supported by the sequence analysis. Similar results

were found by Perdomo et al. (2011b), who demonstrated that Phialemonium is not

monophyletic. Phialemonium curvatum was phylogenetically related to Linocarpon Syd. & P.

Syd. and Neolinocarpon K.D. Hyde, two ascomycetous genera that are incertae sedis within

the Sordariomycetes (Bahl 2006, Huhndorf and Miller 2011).

Studies have demonstrated that Acremonium atrogriseum (Panas.) W. Gams,

characterized by the production of pigmented conidia arranged in chains and slimy heads, is

phylogenetically related to Cephalothecaceae (Perdomo et al. 2011a, Summerbell et al. 2011).

This is a rare opportunistic species described as agent of a human systemic infection and
repeatedly isolated from different clinical sources (de Hoog et al. 2000, Perdomo et al.

2011a). Other fungi show some morphological resemblance and are genetically related to

Phialemonium spp.: Paecilomyces inflatus (Burnside) J.W. Carmich., recently transferred to

the new genus Taifanglania Z.Q. Liang, Y.F. Han, H.L. Chu & R.T.V. Fox (Liang et al. 2009)

and some species of Sarcopodium Ehrenb. and Volutella Tode, such as S. oculorum Gené &

Guarro and V. cinerescens (Ces.) Sacc. The latter two species were described as agents of

ocular infections (de Hoog et al. 2000, Guarro et al. 2002, Weinberger et al. 2006).

Lecythophora spp., reported as anamorphs of Coniochaeta (Sacc.) Cooke

(Coniochateales, Sordariomycetes) (Gams and McGinnis 1983), also are morphologically

similar to Phialemonium. They develop pink-salmon to dark brown colonies and

adelophialides with collarettes, although discrete phialides like those of Acremonium also may

be present (de Hoog et al. 2000, Weber 2002). Lecythophora Nannf. is a cosmopolitan genus,

which comprises six species: L. lignicola Nannf. (type species), L decumbens (J.F.H. Beyma)

E. Weber, Görke & Begerow, L. fasciculata (J.F.H. Beyma) E. Weber, Görke & Begerow, L.

hoffmannii (J.F.H. Beyma) W. Gams & McGinnis, L. luteoviridis (J.F.H. Beyma) E. Weber,

Görke & Begerow, and L. mutabilis (J.F.H. Beyma) W. Gams & McGinnis. The

morphological characters used to differentiate these species are the presence in L. mutabilis

and L. luteoviridis of chlamydospores, which are ovoidal to ellipsoidal and solitary in the

former and globose to oblong or pear-shaped and solitary or in short chains in the latter; the

colony color, which is orange to salmon in L. hoffmannii, brown in L. fasciculata and

olivaceous-brown in L. decumbens. Lecythophora lignicola differs from the other species in

developing ventricose phialides and adelophialides. Lecythophora spp. have been isolated

from wood, water and food (Weber 2002), and some species are also known as human

pathogens. Lecythophora hoffmannii and L. mutabilis have been isolated from keratitis,
subcutanous abscesses, peritonitis, endocarditis and septic shock (de Hoog et al. 2000,

Sakaeyama et al. 2007, Drees et al. 2007, Taniguchi et al. 2009).

In addition Phialemonium, other genera with species morphologically reminiscent of

Lecythophora are Cadophora Lagerb. & Melin, Collophora Damm & Crous, Hormonema

Lagerb. & Melin, Hyphozyma de Hoog & M.T. Sm., Margarinomyces Laxa,

Phaeoacremonium W. Gams, Crous & M.J. Wingf. and Phialophora. These genera belong to

ascomycete orders other than those embracing Lecythophora and Phialemonium (Seifert et al.

2011).

In a recent molecular study of a wide panel of clinical isolates of Phialemonium and

Lecythophora species, analyzing the D1/D2 domains of the 28S rRNA gene sequences, we

found that some isolates could not be properly identified to species and were considered to be

possible cryptic species (Perdomo et al. 2011b). In the present study, to clarify the

phylogenetic relationships among these fungi and other related or morphologically similar

taxa, as well as to describe new species, we performed a multilocus sequence analysis of the

internal transcribed spacer (ITS) regions, the D1/D2 domains of the 28S rRNA (D1/D2), actin

(ACT) and β-tubulin (TUB) genes. Also, we performed an in-depth morphological

characterization of the isolates, re-evaluating the characters seen in the light of the

phylogenetic information obtained in the sequencing study.

MATERIALS AND METHODS

Isolates and morphology.—Twenty-four clinical isolates from the Fungus Testing Laboratory, Department of

Pathology, University of Texas Health Science Center at San Antonio (UTHSC) were included in the present

study. Twenty-two isolates were identified in Perdomo et al. (2011a, b) as Acremonium atrogriseum (n = 1),

Cephalotheca foveolata (n = 4), Phialemonium curvatum (n = 5), Phialemonium obovatum (n = 4),

Phialemonium sp. 1 (n = 5), Phialemonium sp. 2 (n = 1) and Lecythophora sp. 1 (n = 2). Two other clinical

isolates were received as Volutella sp. (UTHSC 04-7 and UTHSC 09-3589). In addition, type or reference strains

of Phialemonium and Lecythophora spp., and species of other morphologically similar genera, such as
Sarcopodium oculorum, S. circinatum Ehrenb. , Volutella ciliata (Alb. & Schwein.) Fr., V. cinerescens,

Cephalotheca sulfurea Fuckel and Taifanglania inflata (Burnside) Z.Q. Liang, Y.F. Han & H.L. Chu, also were

included in the study. (Information on these isolates is detailed in TABLE I.)

For morphological characterization, the isolates were examined on oatmeal agar (OA: 30 g oat flakes,

20 g agar, 1L distilled water), potato dextrose agar (PDA, Difco Laboratories, Detroit, Michigan) and potato

carrot agar (PCA: 20 g potatoes; 20 g carrot; 20 g agar; 1 L tap water). Water agar with sterilized plant material

(small pieces of wood, filter paper and leaves of herbaceous plants) also was used to enhance the formation of

ascomata or conidiomata. Cultures were incubated at room temperature (25 C ± 2 C) in the dark, up to 6 mo.

Growth rates of the isolates were determined on PDA plates at 15, 25, 30, 35, 37, 40 and 42 C for 14 d in

darkness. In descriptions color colony codes in parenthesis refer to Kornerup and Wanscher (1978). The

microscopic features were examined and mesured in lactic acid under a light microscope (Olympus CH-2).

Photomicrographs were obtained with a Zeiss AXIO Imager M1 light microscope and a Jeol JSM-6400 scanning

electron microscope.

Molecular study.—DNA was extracted according to Perdomo et al. (2011a). Protocols for the amplification of

the ITS, D1/D2, ACT and TUB sequences follow Cano et al. (2002), Voigt and Wöstemeyer (2000), and

Gilgado et al. (2005). PCR products were purified according to the manufacturer's instructions using a

commercial kit (Illustra GFX™ PCR DNA and Gel Band Purification, General Electric Healthcare,

Buckinghamshire, UK). Sequencing reactions were carried out with the PCR primers using a Taq DyeDeoxy

Terminator Cycle Sequencing Kit (Applied Biosystems, Gouda, the Netherlands), according to the manufacter's

recommendations, and the resulting products were analyzed on a 310 DNA sequencer (Applied Biosystems).

DNA sequences were aligned with the program Clustal X 1.8 (Thompson et al. 1997). In addition to the

sequences of the clinical isolates and those of type and reference strains generated in this study, several

sequences of other related taxa were retrieved from the GenBank for comparison in this analysis.

The relationships of the fungi tested in this study at suprageneric rank were determined first through the

analysis of D1/D2 domains sequences. Then we performed a multilocus sequence analysis of the D1/D2

domains, ITS regions and a fragment of the ACT and TUB genes of those clades obtained in the first analysis

that were considered of interest for the propose of this study.

Maximum likelihood (ML) analysis of the D1–D2 sequences was conducted with MEGA 5.05 (Tamura

et al., 2011), using Tamura Nei substitution model with gamma distribution. Support for internal branches was

assessed by a search of 1000 bootstrapped sets of data. Bayesian analysis was performed with MrBayes3.0
(Huelsenbeck and Ronquist 2001, Ronquist and Huelsenbeck 2003) by running 1 000 000 generations in four

chains, saving the current tree every 100 generations. The last 10 000 trees were used to construct a 50%

majority rule consensus tree and to determine the posterior probabilities of the branches. The tree was conducted

with a sequence of Collophora rubra Damm & Crous (GQ154606) as outgroup.

In the multilocus study, most parsimonious tree (MPT) analyses of the combined dataset of the loci

mentioned above were performed with PAUP* 4.0b10 (Swofford 2002). One hundred heuristic searches were

conducted with random sequence addition and tree bisection reconnection branch-swapping algorithms,

collapsing zero-length branches and saving all minimal-length trees (MulTrees) on different datasets. The gaps

were treated as missing data and support for internal branches was assessed with a heuristic parsimony search of

1000 bootstrapped datasets. The combined dataset was tested for incongruence with the partition homogeneity

test (PHT) as implemented in PAUP*. Tree length, consistency index (CI), retention index (RI) and the

homoplasy index (HI) values also were calculated. In those analyses the sequences of Lecythophora lignicola

(HE610463 of CBS 267.33) or Phialemonium obovatum (FR691997 of CBS 279.76) were used as outgroup.

The sequences generated in this study and the alignments used the phylogenetic analyses were

deposited respectively in GenBank (TABLE I) and in TreeBASE (www.treebase.org, submission numbers

M11953, M11954, M11955 and M11256).

RESULTS

Phylogenetic analyses.—We illustrated the phylogenetic tree inferred from the analysis of the

D1/D2 sequences of the 24 clinical isolates and of other 35 fungi morphologically or

genetically related (FIG. 1). The aligned sequence regions consisted of 469 bp. The sequences

were distributed in three main clades, that is clade I, incertae sedis (99% bs and 1 pp), clade

II, Hypocreales (91% bs and 1 pp) and clade III, Sordariales (clade III, 76% bs and 1 pp),

while that the representatives of Coniochaetales grouped with low bootstrap support (bs).

Clade I embraced 12 clinical isolates, the type and reference strains of P. curvatum, one

reference strain of Volutella cinerescens (CBS 832.71), the type and a reference strain of

Sarcopodium oculorum (IHEM 19077, FMR 7190 respectively) and GenBank sequences of

the ascomycetes Linocarpon livistonae (Henn.) K.D. Hyde, Neolinocarpon globosicarpum

K.D. Hyde and Neolinocarpon enshiense K.D. Hyde, Joanne E. Taylor & J. Fröhl. Because
the type species of the genus Phialemonium (P. obovatum) was distant from clade I and

included in the order Sordariales (clade III), clade I may represent an anamorph genus

different from Phialemonium. This clade is also different from Sarcopodium or Volutella

because reference strains of the type species of these two genera (S. circinatum, V. ciliata

respectively) were placed in the order Hypocreales (Clade II). Clade III included members of

the family Cephalothecaceae related to P. obovatum; that is type strains and clinical isolates

of A. atrogriseum and C. foveolaata, the type strains of Albertiniella polyporicola (Jacz.)

Malloch & Cain, Cryptendoxyla hypophloia Malloch & Cain and P. inflatus, reference strains

of T. inflata and C. sulfurea, in addition to the clinical isolate UTHSC 03-3661, which was

closely related to the latter three fungi and identified in Perdomo et al. (2011b) as

Phialemonium sp. 2. Finally, the type and reference strains of all the currently accepted

species of Lecythophora, some members of the ascomycetous genus Coniochaeta and the

clinical isolates UTHSC 01-20-1 and UTHSC 01-1644 identified in Perdomo et al. (2011b) as

Lecythophora sp. 1 constituted a separate group.

To confirm the phylogenetic relationships in the D1/D2 domains tree and determine

the boundaries of some of the taxa included in the study, we performed separated multilocus

analyses of clades I, III and the Coniochaetales group, including the most representative

members of each clade (FIG. 1).

We illustrated the multilocus analysis of clade I (FIG. 2). Two MPT were produced

from a heuristic search using the combined dataset of 2135 characters from the four loci

tested. The resulting topology was similar to the topologies obtained by analyzing the genes

individually (data not shown). Analyses of the combined partitions support the recognition of

four well supported terminal clades similar to those previously observed (FIG. 1), each

representing a different phylogenetic species.

 The first clade (100% bs) was composed of the type (IHEM 19077), a reference strain

(FMR 7190) of S. oculorum, and three additional isolates (UTHSC 05-2527, UTHSC 07-

3736, UTHSC 09-2358). Sequences variation within these isolates was 0–1 bp/454 total bp

for D1/D2 (99.3–100%), 1–8/476 for ITS (99.2–99.7%), 0–12/713 for ACT (98.2–100%) and

0–2/375 for the TUB (99.8–100%). The second and the third clades were composed of two

clinical isolates each (100% and 96% bs respectively). The fourth clade (100% bs) included

the type and reference strains of P. curvatum, and the strain V. cinerescens CBS 832.71.

Sequence similarities within this group were 99.4–100% for D1/D2, 99.2–100% for ITS,

99.2–100% for ACT and 99.5–100% for the TUB.

The combined multilocus analysis of the Coniocahetales group included eight strains

of Lecythophora spp. and P. obovatum as outgroup (FIG. 3). The maximum parsimony

analysis of the dataset (2173 bp) produced one MPT, with eight terminal branches. Six

corresponded to known species and the other two, UTHSC 01-20-1 and UTHSC 01-1644,

represented two undescribed species related to L. hoffmannii.

The multilocus sequence analysis of members of clade IIIwith L. lignicola as outgroup

are illustrated (FIG. 4). The phylogenetic analysis of the dataset (2175 bp) yielded one MPT,

in which three well supported (100% bs) primary clades, were shown. The first clade was

composed of two sister terminal subclades; one (100% bs) included the type and different

isolates of P. obovatum, and the other (100% bs) grouped the type strain and four clinical

isolates of C. foveolata. The second clade contained the type strain of A. atrogriseum and

three conspecific isolates (Perdomo et al. 2011a). The similarities among the sequences of the

members of this group were 99.8–100% for D1/D2, 98.6–100% for ITS, 98.5–99.9% for ACT

and 98.8–100% for the TUB. The third clade was divided into three terminal subclades; the

first one included a reference strain of T. inflata and the type strain of P. inflatus. The

sequence similarity was 98.9–100% for all the loci studied, which demonstrated that both
species are conspecific. The second subclade included only the clinical isolate UTHSC 03-

3661, and finally the third subclade was constituted by CBS 135.34, a reference strain of C.

sulfurea which is the type species of the genus Cephalotheca.

Morphological study.—The morphological study of the 24 clinical isolates and numerous type

and reference strains of Phialemonium, Lecythophora and related taxa, let us characterize

phenotypically the different clades shown in the trees inferred from the phylogenetic analyses.

The isolates of clade I (FIG. 1) are characterized by cylindrical, straight to slightly

flexuose, hyaline to brown conidiogenous cells (discrete phialides and adelophialides), in all

cases with collarettes. They are solitary, borne directly on undifferentiated hyphae or

aggregated in sporodochium-like or pycnidium-like conidiomata. Intermediate forms between

the conidiomatal types also were present in some cases. Four morphotypes could be

distinguished within theses isolates, which corresponded to the four main phylogenetic clades

observed in the multilocus analysis (FIG. 2). One morphotype included the three clinical

isolates identified as Phialemonium sp. 1 (Perdomo et al. 2011b) and the type and a reference

strain of S. oculorum; they all develop sporodochium- and pycnidium-like conidiomata. The

conidiomatal structures were characterized by a wall with two types of tissue, textura intricata

and textura epidermoidea. They produced ellipsoidal to slightly allantoid conidia.

A second morphotype contained two isolates, UTHSC 06-1820 and UTHSC 06-1465.

It forms unilocular pycnidium-like conidiomata characterized by a wall of textura intricata

and obovoid conidia.

A third morphotype included the isolates UTHSC 04-7 and UTHSC 09-3589. These

isolates have multilocular conidiomata, with a wall of textura epidermoidea, and ellipsoidal

conidia.

The last morphotype included the isolates identified as P. curvatum; three of them,

UTHSC 06-4324, UTHSC R-3447 and UTHSC R-3448, were characterized by the presence
of sporodochium-like conidiomata. These structures also were observed in the conspecific

strain CBS 832.71, which was received from the CBS collection as V. cinerescens.

All the members of clade III (FIG. 1) developed discrete phialides or adelophialides

with a cylindrical or slightly swollen basal region, usually without visible collarettes. These

characters are typical of the genus Phialemonium. Within this clade, we recognized six

morphotypes, which corresponded with the terminal clades shown in the corresponding

multilocus analysis (FIG. 4).

The isolates in the first morphotype, identified as P. obovatum, developed obovate

conidia with a truncate base and oval chlamydospores. Those in the second morphotype,

which corresponded to C. foveolata isolates, showed cylindrical conidia, globose to

ellipsoidal chlamydospores and the characteristic teleomorph. The P. obovatum and C.

foveolata isolates had conidia arranged in slimy heads.

The third morphotype, with corresponded to isolates identified as A. atrogriseum

(Gams 1971, Perdomo et al 2011a), produced flask-shaped phialides and adelophialides,

occasionally with a markedly inflated base, and produced slightly pigmented obovoid conidia

with a weakly apiculate base, arranged in slimy heads or in chains.

A fourth morphotype included the strains of T. inflata and P. inflatus, which showed

discrete phialides with a swollen basal part, tapering into a long neck, and citriform conidia.

The fifth morphotype, consisting of a single clinical isolate (UTHSC 03-3661),

showed phialides similar to those described in strains of A. atrogriseum and P. inflatus, but

globose conidia.

The sixth morphotype, which corresponded to the reference strain of C. sulfurea (CBS

135.34), developed an anamorph with flask-shaped phialides and ovate conidia. The conidia

in strains of the last-mentioned three morphotypes formed long and persistent chains, and they
had connectives on both ends. This character was not described previously for the genus

Phialemonium (Gams and McGinnis 1983).

Most isolates included in the Coniochareales group (FIG. 1) showed the typical

morphological features of Lecythophora. However, two of them, UTHSC 01-20-1 and

UTHSC 01-1644, had characteristics that were distinct from those of the other described

species of Lecythophora. The first isolate, UTHSC 01-20-1, excreted a diffusible orange

pigment and produced discrete phialides and adelophialides with a swollen base, while the

second, UTHSC 01-1644, produced chains of darkly pigmented chlamydospores.

TAXONOMY

The morphological features shown by the Phialemonium species in clade I are very different

from the typical features shown by Phialemonium s. str. and its relatives (clade III). Because

these morphological differences also are reflected in the multilocus sequence analysis, we

therefore consider that the fungi included in clade I represent a new genus, which is proposed

below as Phialemoniopsis. Four species are described in the new genus. Furthermore, three

new species of Phialemonium and two other of Lecytophoraare are described in the present

study.

Phialemoniopsis H. Perdomo, D. García, Gené, Cano & Guarro, gen. nov.

MycoBank MB563874

Colonies growing moderately with aerial mycelium sparse or lacking, brown or grayish.

Vegetative hyphae hyaline to brown, smooth- to rough-walled. Sporulation from

conidiophores solitary, borne on undifferentiated hyphae from the aerial mycelium or grouped

into sporodochium-like or pycnidium-like conidiomata growing on the agar surface.

Conidiogenesis enteroblastic. Conidiophores simple or slightly branched, hyaline or pale

brown. Conidiogenous cells monophialidic, discrete, terminal or lateral, cylindrical to flask-

shaped, straight to slightly flexuose, or intercalary with a cylindrical lateral projection

(adelophialides), with collarettes, hyaline, subhyaline or brown with age. Sporodochium-like

conidiomata superficial, solitary, applanate, pulvinate or cupulate, subhyaline to dark brown.

Pycnidium-like conidiomata superficial or semi-immersed, solitary or aggregated, uni- or

multilocular, globose, subglobose, ellipsoidal or pyriform, surrounded by a thin wall of

several layers; wall of textura intricata or textura epidermoidea or both, with an irregular

dehiscence (lacking apical preformed opening or ostiole), often appearing cupulate when

mature. Setae usually present, forming a frill at the margin of open conidiomata or

interspersed with conidiophores, simple or slightly branched, septate, straight to slight

flexuose, hyaline, subhyaline to brown. Conidia one-celled, ellipsoidal, allantoid to obovoid,

and slightly apiculate at the base, hyaline to subhyaline, smooth-walled, aggregated in slimy

heads. Chlamydospores sometimes present. Teleomorph unknown.

Type species: Phialemoniopsis ocularis (Gené & Guarro) H. Perdomo, D. García,

Gené, Cano & Guarro.

Etymology: Phialemonium, a generic name of hyphomycetes; from the Greek -opsis, a

suffix meaning appearance.

Phialemoniopsis cornearis H. Perdomo, D. García, Gené, Cano & Guarro, sp. nov. FIG. 5

MycoBank MB563875

Colonies on OA at 25 C attained 54–60 mm diam after 14 d, flat, velvety, with whitish soft

cottony aerial mycelium and olive-brown (4F3) submerged hyphae; reverse colorless to dark

brown (6F5) with age. Colonies on PDA at 25 C, 35–40 mm diam after 14 d, fasciculate,

radially folded, initially white to brownish gray (4E2), becoming olive-brown (4E4) to black;

reverse colorless first becoming olive-brown (4E4). The fungus grew slowly at 37 C (up to 9

mm diam after 14 d); no growth at 40 C; optimal temperature for growth was at 25–30 C.

Vegetative hyphae initially hyaline, smooth-walled, becoming brown and verruculose, 2–4

μm wide. Sporulation from conidiophores growing solitary on undifferentiated hyphae of the

aerial mycelium, or aggregated in sporodochium-like and pycnidium-like conidiomata.

Conidiophores from aerial mycelium mostly short, erect, often simple or few branched, 10–30

× 1–2 μm. Phialides and adelophialides straight to slightly flexuose, cylindrical, 2–24 × 1–2

μm, with conspicuous collarette, initially hyaline to subhyaline becoming brown, smooth-

walled. Conidia one-celled, ellipsoidal to slight allantoid, 4–7 × 1–2 μm, slightly apiculate at

the base, biguttulate, hyaline, smooth-walled, in slimy heads. Sporodochium-like conidiomata

superficial, solitary, applanate, cupulate or pulvinate, up to 350 μm diam, hyaline to brown,

setose. Pycnidium-like conidiomata superficial, solitary, unilocular, non-ostiolate, globose to

subglobose, 300–650 × 200–450 μm; wall 3–4 μm thick, with textura intricata, dark brown to

black. These conidiomata open irregularly and often appearing cupulate when mature and

surrounded by subhyaline to brown setae. Setae forming a frill at the margin of the

conidiomata, or interspersed with the conidiophores, usually unbranched, cylindrical, up to 70

μm long, 1–2 μm wide, hyaline, subhyaline, or brown, smooth and slightly thick-walled.

Conidiomatal conidiophores often irregularly branched, with branches bearing a terminal

group of 3–5 discrete phialides. Phialides cylindrical or subcylindrical, 9–25 × 1–1.5 μm, with

a barely visible collarette, hyaline, smooth-walled. Conidia one-celled, obovoid or ellipsoidal,

slightly apiculate at the base, 2–3 × 1–1.5 μm, one-guttulate, smooth-walled, hyaline, pale

yellow in slimy masses. Chlamydospores present on undifferentiated hyphae, terminal or

lateral, solitary or in short chains, globose, oval to pyriform, 5–9 × 4–5 μm, initially hyaline

and thin-walled, becoming brown, thick- and rough-walled. Teleomorph unknown.

Etymology: From the Latin cornearis referred to the clinical specimen from which the

type strain was isolated.

Specimens examined: USA. FLORIDA. From human corneal fluid, D.A. Sutton. (CBS H-20781,

HOLOTYPE; UTHSC 06-1820, CBS 131711, FMR 10701, ex-type cultures). USA. SOUTH CAROLINA. From

shin aspirate, D.A. Sutton (UTHSC 06-1465).

 Phialemoniopsis cornearis is similar to P. ocularis and P. pluriloculosa in the

presence of sporodochium- and pycnidium-like conidiomata and chlamydospores. However,

the pycnidium-like conidiomata of P. cornearis shows a wall of textura intricata and it usually

produces obovoid conidia.

Phialemoniopsis curvata (W. Gams & W. B. Cooke) H. Perdomo, D. García, Gené, Cano &

Guarro, comb. nov.

MycoBank MB563876

≡Phialemonium curvatum W. Gams & W. B. Cooke, in Gams & McGinnis, Mycologia 75:980 (1983).

Colony on OA at 25 C attained 40–70 mm diam after 14 d, smooth or finely floccose, at first

white becoming brown (6E5) to grayish brown (6E3). On PDA at 25 C, 46–48 mm diam after

14 d, cream-colored (4A3), with small areas light brown (6D4). The fungus grew slowly at 37

C (5–11 mm diam in 14 d); no growth at 40 C; optimal temperature for growth was at 25–30

C. Vegetative hyphae hyaline to light brown, 1–2 μm wide, smooth- to slightly rough-walled.

Sporulation on undifferentiated hyphae from the aerial mycelium or from sporodochium-like

conidiomata in some isolates (CBS 832.71, UTHSC 06-4324, UTHSC R-3447, UTHSC R-

3448). Adelophialides abundant, 4–12 × 1–1.5 µm, with barely visible collarettes. Discrete

phialides less common, long, cylindrical, slightly tapering toward the apex, 8–20 × 2–3.5 µm,

with barely visible collarettes. Conidia cylindrical or allantoid, 4–6 × 1–2 µm, hyaline,

smooth-walled, in slimy heads. Sporodochium-like conidiomata usually pulvinate, up to 1.5

mm diam, composed of differentiated, straight or flexuous, irregularly branched

conidiophores, with a marginal frill of setae and with a slimy conidial mass covering the

entire upper surface of the conidioma. Setae unbranched or slightly branched toward the base,

straight or flexuose, septate, cylindrical, with slightly pointed or blunt tip, up to 60 μm long,

3–4 μm wide, hyaline to subhyaline, slightly thick-walled. Conidia hyaline, ellipsoidal to

ovoidal, smooth-walled, 1–2 × 0.5–1 µm. Chlamydospores absent. Teleomorph unknown.

 Specimens examined: USA. GEORGIA: Atlanta. From skin lesion, W.B. Cooke (CBS 490.82, ex-type

culture). USA. CALIFORNIA: From chaparral soil, W.B. Cooke (CBS 491.82). USA. MINNESOTA: From

human sinus, D.A. Sutton (UTHSC 04-956). USA. TEXAS: From canine pleural fluid, D.A. Sutton (UTHSC 06-

4324). USA. UTAH. From human blood, D.A. Sutton (UTHSC 08-2292). ISRAEL. From human eye, D.A.

Sutton (UTHSC R-3447); from human eye, D.A. Sutton (UTHSC R-3448). As Volutella cinerescens: from

keratomycosis, R. Zapater (CBS 832.71).

Phialemoniopsis curvata is mainly distinguished from the other species of the genus

by the absence of pycnidium-like conidiomata and of chlamydospores.

Phialemoniopsis ocularis (Gené & Guarro) H. Perdomo, D. García, Gené, Cano & Guarro,

comb. nov. FIG. 6

MycoBank MB563877

≡ Sarcopodium oculorum Gené & Guarro, in Guarro et al., J. Clin. Microbiol. 40:3074 (2002).

Colonies on OA at 25 C attained 47–48 mm diam after 14 d, granulose, smooth toward the

periphery, with whitish, soft cottony aerial mycelium and brownish gray (4E2) submerged

hyphae. On PDA at 25 C, the colonies reaching 39–50 mm diam after 14 d, cream-colored

(4A3) to brownish gray (4E2), radially folded and granulose due to the abundant production

of conidiomata. The fungus grew slowly at 37 C (9–14 mm diam after 14 d); no growth at 40

C; optimal temperature for growth was 25 C. Sporulation from conidiophores solitary on the

aerial mycelium or aggregated into sporodochium-like and pycnidium-like conidiomata.

Phialides and adelophialides from aerial mycelium straight to slightly flexuose, cylindrical, 3–

19(–36) × 1–2 μm, with collarettes, initially hyaline to subhyaline becoming brown, smooth-

walled. Conidia ellipsoidal to slightly allantoid, 3–5 × 1–1.5 μm, slightly appiculate at the

base, biguttulate, hyaline, smooth-walled, arranged in slimy heads. Sporodochium-like

conidiomata superficial, applanate to cupulate or pulvinate, up to 400 μm diam, with setae

forming a frill at the margin of the conidiomata and interspersed with the conidiophores.

Pycnidium-like conidiomata superficial, solitary, unilocular, globose to subglobose, 300–500

× 120 – 600 μm; wall 6–7 μm thick, dark brown to black, with two types of textura intricata

and epidermoidea. These conidiomata open by an irregular rupture and often appear cup-

shaped when mature and surrounded by setae. Setae erect, usually unbranched, straight or

flexuose, septate, cylindrical, with slightly pointed or blunt tips, up to 65 μm long, 1.5–2.5

μm wide, subhyaline to dark brown, smooth-walled, thin- to slightly thick-walled.

Conidiomatal conidiophores hyaline, straight, simple or irregularly branched, with branches

bearing a group of 2–4 phialides terminally. Phialides usually with a slightly swollen basal

part and slightly tapering toward the tip, 5–8 × 1–1.5 μm, hyaline. Conidia one-celled,

ellipsoidal, some lightly allantoid, 2–3 × 1–1.5 μm, with a slightly apiculate base, one-

guttulate, hyaline, smooth-walled. Chlamydospores present, intercalary, solitary or in chains,

globose to pyriform, 5–9 × 3–6 μm, initially hyaline becoming brown, thick- and rough-

walled. Teleomorph unknown.

Specimens examined: BRAZIL. Sao Paulo: From human keratitis, P. Godoy (IHEM 19077, ex-type

culture). BRAZIL. Sao Paulo: From human keratitis, P. Godoy (FMR 7190). USA PENNSYLVANIA: From

peritoneal dialysis catheter, D.A. Sutton (UTHSC 05-2527). USA. FLORIDA: Left hand, D.A. Sutton (UTHSC

07-3736). USA. MASSACHUSETTS: From aspirate cellulitis, D.A. Sutton (UTHSC 09-2358).

A description of this species was given by Guarro et al. (2002). However, pycnidium-

like conidiomata or chlamydospores were not described in the protolog. In P. ocularis, as in

P. cornearis and P. pluriloculosa, the sporulation is associated with conidiophores arising

from the aerial mycelium and with the sporodochium- and pycnidium-like conidiomata. The

main distinctive feature of P. ocularis is that the pycnidium-like conidiomata have a wall

made of both textura intricata and textura epidermoidea, and the conidia from these structures

are usually ellipsoidal. In addition, it differs from the other species by its growth at 37 C.

Phialemoniopsis pluriloculosa H. Perdomo, D. García, Gené, Cano & Guarro, sp. nov. FIG. 7

MycoBank MB563878
Colonies on OA 25 C attained 54–55 mm diam after 14 d, velvety and yellowish white (4A2),

but soon become olive-brown (4E4) to black; reverse colorless to black with age. On PDA at

25 C the colonies reaching 40–43 mm diam after 14 d, mucous, radially folded, yellowish

white (4A2); reverse pale yellow (4A3). The fungus grew slowly at 35 C (up to 8 mm diam in

14 d); no growth at 37 C; optimal temperature for growth was 25 C. Vegetative hyphae

initially hyaline, smooth-walled, becoming brown and slightly verruculose, 1–2 μm wide.

Sporulation from conidiophores growing on the aerial mycelium, and from sporodochium-

and pycnidium-like conidiomata. Phialides and adelophialides born on aerial mycelium

straight to slightly flexuose, cylindrical, 6–20 × 1–2 μm, with short collarettes, initially

hyaline to subhyaline, but turning brown, smooth-walled. Conidia one-celled, ellipsoidal to

cylindrical, some slightly curved, 3–5 × 1–2.5 μm, with a slightly apiculate base, biguttulate,

hyaline, smooth-walled, in slimy heads. Sporodochium-like conidiomata superficial, solitary,

pulvinate, up to 450 μm diam, subhyaline to brown, setose. Pycnidium-like conidiomata

superficial or semi-immersed, solitary or confluent, multilocular, globose to subglobose, 150–

450 × 150–350 μm; wall 10–15 μm thick, with textura epidermoidea, dark brown to black,

non-ostiolate, opening irregularly and often appearing cupulate and with marginal setae. Setae

unbranched or slightly branched toward the base, straight or flexuose, septate, cylindrical,

with slightly pointed or blunt tips, up to 60 μm long, 2.5–3 μm wide, subhyaline to brown,

slightly thick-walled. Conidiomatal conidiophores straight, compact, filiform, simple or few

branched, and often proliferate sympodially showing conidiogenous cells as lateral cylindrical

projections, hyaline. Phialides subcylindrical, slightly tapering toward the tip, 6–12 × 1–1.5

μm, hyaline. Conidia one-celled, ellipsoidal, 2–3 × 1–1.5 μm, with a slightly apiculate base,

1–2 guttulate, smooth-walled, hyaline, pale yellow in mass. Chlamydospores present,

unicellular, intercalary, usually in chains, subglobose, oval to pyriform, 5–9 × 4–6 μm,

hyaline becoming brown, thick- and rough-walled. Teleomorph unknown.

 Etymology: From the Latin pluriloculosa referred to the multilocular conidiomata.

Specimens examined: USA. NEVADA: From human toe nail, D.A. Sutton (CBS H-20782,

HOLOTYPE; UTHSC 04–7, CBS 131712, FMR 11070, ex-type cultures). USA. MINNESOTA: From human

synovial fluid, D.A. Sutton (UTHSC 09-3589).

Phialemoniopsis pluriloculosa differs from the other species of the genus by the

presence of multilocular pycnidium-like conidiomata with textura epidermoidea and by the

production of filiform conidiophores that usually proliferate sympodially. The key

morphological features to distinguishing the Phialemoniopsis species are summarized (TABLE

II).

Phialemonium W. Gams & McGinnis, Mycologia 75:978, 1983

Emended description.

Colonies spreading, flat, velvety, plane, or slightly floccose, white, yellow-gray, ochraceous

or dark olivaceous gray. Conidiophores undifferentiated, simple or occasionally branched.

Phialides often inflated at base, with or without barely visible collarettes. Adelophialides

often present, short cylindrical. Conidia hyaline or pigmented, one-celled, cylindrical,

obovate, globose to citriform, mostly smooth-walled or inconspicuously rugose, arranged in

slimy heads or in long chains, with or without connectives at both ends. Chlamydospores

present or absent. Teleomorphs: Albertiniella and Cephalotheca (Cephalothecaceae,

Sordariales).

Type species: Phialemonium obovatum W. Gams & McGinnis

The concept of Phialemonium is extended to include species with phialides inflated at

base and more or less conspicuous collarettes, and conidia arranged in long chains that may

have connectives at both ends.

Phialemonium atrogriseum (Panas.) D. García, H. Perdomo, Gené, Cano & Guarro, comb.

nov. FIG. 8A–G

MycoBank MB563879

≡ Phaeoscopulariopsis atrogrisea Panas., Mycologia 56:60 (1964).

≡ Acremonium atrogriseum (Panas.) W. Gams, Cephalosporium-artige Schimmelpilze (Stuttgart) 89

(1971).

Colony on OA at 25 C attaining a diameter of 28–40 mm after 14 d, powdery, at first pale

ochraceous, becoming olive-brown (4E5) to black. No growth at 35 C; optimal temperature

for growth was at 25 C. Conidiophores absent or poorly differentiated, often single lateral

phialides born directly from aerial hyphae , or consisting of whorls of 2–4 phialides on top of

lateral supporting cells. Phialides flask-shaped with more or less inflated basal part and a

long, tapering neck bearing a barely visible collarete, 8–20 × 2–3.5 µm wide at base, mostly

subhyaline and smooth-walled, pale yellowish-brown and slightly rough-walled in some

isolates. Adelophialides may be also present, 7–11 × 1.5–2 µm. Conidia 1-celled, ellipsoidal

or obovoid, 3–5 × 2–3 µm, with an apiculate and minutely truncate base, subhyaline to

grayish brown, smooth-walled, in slimy heads and in long chains, without connectives.

Chlamydospores absent. Teleomorph unknown.

Specimens examined: UKRAINE. Kharkiv: From noodles, V.T. Panasenko (CBS 604.67, ex-type

culture). GERMANY. München; From human urine, A. Haas (CBS 774.97). USA. DELAWARE: Dover. From

a lower lobe lung, A.A. Padhye (CBS 306.85). USA. PENNSYLVANIA: From BAL aplastic anemia, D.A.

Sutton (UTHSC 03-986).

Phialemonium atrogriseum differs from other Phialemonium species forming conidial

chains by its slightly pigmented conidia without connectives. Although P. atrogriseum forms

conidia similar to those of P. obovatum, in the latter they are not pigmented and are arranged

only in slimy heads. Also, P. obovatum produces chlamydospores and P. atrogriseum does

not.

Phialemonium globosum D. García, H. Perdomo, Gené, Cano & Guarro, sp. nov. FIG. 8H–L

MycoBank MB563880
Colonies on OA and PDA at 25 C attained up to 30 mm diam after 14 d, flat, velvety to

powdery, sometimes fasciculate in the center, white. The fungus grew slowly at 37 C (up to

12 mm diam in 14 d); no growth at 40 C; optimal temperature for growth was 25 C.

Conidiophores absent or poorly differentiated, often single lateral phialides borne directly

from aerial hyphae, occasionally 1–2 phialides borne on top of a lateral supporting cell.

Phialides terminal or lateral, sometimes sympodially proliferating from the supporting cell

without basal septa, cylindrical or with a slightly swollen basal part, sometimes slightly

constricted at midpoint, and a long tapering neck, (7–)9–24(–36) × 1.5–3 µm near the base, 1–

1.5 µm wide at apex, with inconspicuous collarettes, hyaline, smooth-walled. Adelophialides

scarce, cylindrical, up to 15 × 3 µm near the base. Conidia one-celled, globose to subglobose,

4–6 × 3.5–5 µm, with distinct connectives on both ends, hyaline, smooth-walled, forming

persistent chains. Chamydospores absent. Teleomorph unknown.

Etymology: From the Latin globosum referred to the conidial shape.

Specimen examined: USA. ILLINOIS: From human sinus, D.A. Sutton (CBS H-20783, HOLOTYPE;

UTHSC 03–3661, CBS 131713, FMR 10390, ex-type cultures).

Phialemonium globosum is morphologically similar to the anamorph of C. sulfurea

and Phialemonium inflatum (see below). They all share phialides with a more or less swollen

basal portion and form long conidial chains with connectives at both ends. However, these

three species can be distinguished by their conidial morphology: P. globosum has globose

conidia up to 5 µm diam, the anamorph of C. sulfurea has ovoid conidia up to 3 µm diam, and

P. inflatum has citriform conidia up to 3 µm diam.

Phialemonium inflatum (Burnside) D. García, H. Perdomo, Gené, Cano & Guarro, comb.

nov. FIG. 8M–Q

MycoBank MB563881

≡ Myceliophthora inflata Burnside, Pap. Mich. Acad. Sci. 8:82 (1928)

 ≡ Paecilomyces inflatus (Burnside) Carmichael, Can. J. Bot. 40:1148 (1962)

≡ Taifanglania inflata (Burnside) Z.Q. Liang, Y.F. Han & H.L. Chu, Fungal Diversity 34:72 (2009)

Colony on OA at 25 C attaining 35 mm diam after 14 d, velvety to granular, white to pale

yellow (4A3). No growth at 35 C; optimal temperature 25 C. Conidiophores absent or poorly

differentiated, mostly consisting of single phialides borne irregularly on aerial hyphae,

occasionally 2–3 phialides borne atop lateral supporting cells. Phialides lateral or terminal,

flask-shaped, with a swollen basal part and tapering abruptly into a long neck, 7–15(–19) ×

(1.5–)2–3 µm, hyaline, smooth-walled. Conidia one-celled, citriform, 4–5 × 2–3 µm, with

distinct connectives on both ends, hyaline, smooth- to slightly rough-walled, and arranged in

chains. Adelophialides and chlamydospores absent. Teleomorph unknown.

Specimens examined: USA. WASHINGTON: From Apis mellifera, J.A. Stevenson (CBS 259.39, ex-

type culture). As Taifanglania inflata: RUSSIA. Ust-Omchug. From cabbage field soil, T. Ito and T. Yokoyama

(NBRC 31965).

The key morphological features, based on OA cultures, useful for distinguishing the

Phialemonium species included in the present study, are summarized (TABLE III).

Lecythophora cateniformis H. Perdomo, D. García, Gené, Cano & Guarro, sp. nov. FIG.

9A–I

MycoBank MB563407

Colonies on OA at 25 C attained 35 mm diam after 14 d, velvety, brownish gray (8F2) at the

center and white at the periphery; reverse brownish gray (8F2). On PDA at 25 C, the colonies

reaching 35 mm diam after 14 d, flat, velvety, moist, orange-white (6A2); reverse pale yellow

(4A3). The fungus grew moderately at 37 C (up to 32 mm diam in 14 d); no growth at 40 C;

optimal temperature was 25 C. Vegetative hyphae 1–2 μm wide, hyaline or subhyaline,

smooth-walled. Adelophialides predominant, consisting of lateral collarettes up to 1.5 μm

long, growing directly on the undifferentiated hyphae, or as cylindrical lateral projections up

to 3 μm long. Phialides rare. Conidia one-celled, obovoidal to ellipsoidal, 3–4(–6) × 2–3(–4)

μm, with slightly protruding and truncate base, one-guttulate, hyaline, smooth-walled,

aggregated in slimy heads. Chlamydospores solitary or in short chains, subglobose, oval to

pyriform, 4–7(–9) × 4–5(–6) μm, pale brown to brown, smooth and slightly thick-walled.

Teleomorph unknown.

Etymology: From the Latin cateniformis referred to the chains of chlamydospores

produced by the fungus.

Specimen examined: USA. ALABAMA: From canine bone marrow aspirate, D.A. Sutton (CBS H-

20779, HOLOTYPE; UTHSC 01-1644, CBS 131709, FMR 10743, ex-type cultures).

Lecythophora cateniformis is similar to L. luteoviridis, but the latter species produces

hyaline or faintly brown chlamydospores and its conidia are ellipsoidal to cylindrical up to 2

μm wide.

Lecythophora luteorubra H. Perdomo, D. García, Gené, Cano & Guarro, sp. nov. FIG. 9J–P

MycoBank MB563408

Colonies on OA at 25 C attained 25 mm diam after 14 d, flat, slightly fasciculate at the center,

orange (6B8); reverse orange (6A7). Colonies on PDA at 25 C attained 25 mm diam after 14

d, downy, orange (6B8); reverse orange (6B7). Diffusible pigment orange-red to brownish red

(10D8) was produced on both media at 25 C, but it is absent at 30 C. The fungus grew slowly

at 37 C (up to 11 mm diam in 14 d); no growth at 40 C; optimal temperature was at 30 C.

Vegetative hyphae hyaline, 2–4 μm wide. Phialides predominant, terminal or lateral,

cylindrical or ampulliform, 5–10 × 1–3 μm, with distinct cylindrical collarettes, hyaline,

smooth-walled. Adelophialides rare, some reduced to lateral collarettes on undifferentiated

hyphae, 1–1.5 × 1 μm. Conidia one-celled, ellipsoidal or cylindrical, 3–6(–7) × 1–2 μm, with

a slightly apiculate base, biguttulate, hyaline, smooth-walled, aggregated in slimy heads.

Chlamydospores absent. Teleomorph unknown.

Etymology: From the Latin luteorubra, referred to the orange-red diffusible pigment.
 Specimen examined: USA. MASSACHUSETTS: From leg wound, D.A. Sutton (CBS H-20780,

HOLOTYPE; UTHSC 01-20-1, CBS 131710, FMR 10721, ex-type cultures).

Lecythophora luteorubra is morphologically similar to L. hoffmannii and the

anamorph of Coniochaeta ligniaria (Grev.) Massee. However, L. hoffmannii does not produce

diffusible pigment, and its conidia are slightly smaller (3.5–5 × 1.2–2 μm) and often

somewhat more curved than those of L. luteorubra. In the anamorph of C. ligniaria, the

conidia often are curved; the adelophialides are predominant; although some cylindrical

phialides may be present, they are never ampulliform; and the teleomorph is usually present.

The key morphological features useful to distinguish current Lecythophora species are

summarized (TABLE IV).

DISCUSSION

The genus Phialemonium was proposed to accommodate three species with morphological

features intermediate between Acremonium and Phialophora (Gams and McGinnis 1983).

Yaguchi et al. (2006) and Perdomo et al. (2011b) demonstrated that, as originally conceived,

the genus was polyphyletic. They suggested that P. curvatum should be accommodated in

different genus. Based on combined morphological and additional molecular data, we propose

the new anamorphic genus Phialemoniopsis to accommodate P. curvatum and three additional

species, Phialemoniopsis cornearis and P. pluriloculosa proposed here as new and P. ocularis

previously included in Sarcopodium (Guarro et al. 2002).

Phialemoniopsis can be distinguished from Phialemonium by having phialides and

adelophialides both with collarettes and by the development of sporodochium- or pycnidium-

like conidiomata, which are absent in Phialemonium s. str.

Other anamorphic genera that have the same conidial ontogeny and develop

conidiomata similar to those of Phialemoniopsis are Chaetomella Fuckel and Cornucopiella

Höhn., both traditionally placed among the coelomycetes (Sutton 1980), and the

hyphomycetous genera Collophora, Sarcopodium and Volutella.

Chaetomella is mainly distinguished by its pycnidial conidiomata with a wall of

textura angularis and by the production of fusiform or falcate conidia. Molecular data placed

the members of Chaetomella within the Leotiomycetes (Helotiales) (Rossman et al. 2004).

Cornucopiella comprises two species, C. mirabilis Höhn., type species, and. C.

fusispora (P. Karst.) Seifert. Their conidiomata are usually cylindrical, sessile and non-setose,

with a wall of textura porrecta (Sutton 1980, Seifert 1985). The taxonomical placement of

Cornucopiella is still uncertain among ascomycetes. Cultures of the first species are not

available for study; however, we examined and sequenced the D1/D2 and ITS regions of a

reference strain of C. fusispora (CBS 113544). That strain did not sporulate in any agar media

tested. The BLAST query of the ITS and D1-D2 of it showed similarities below 91% with

uncultured Leotiomycetes (JF449892.1) and 97% with Collophora pallida (GQ154612.1)

respectively.

Collophora is a genus incertae sedis in the Leotiomycetes (Helotiales). It produces

white, cream or reddish colonies and pycnidium-like conidiomata with walls of textura

epidermoidea (Damm et al. 2010). Sporodochial conidiomata is not present in this genus. The

species of Collophora are known only as agents of wood necroses of Prunus trees (Damm et

al. 2010).

Members of Sarcopodium and Volutella develop only sporodochial conidiomata; in

Sarcopodium they are brown pinkish or purple, with brown setae (Sutton 1981, Domsch et al.

2007), and in Volutella they are white to pale with hyaline setae around the margin (Maud and

Duke 1928, Gräfenhan et al. 2011). Most species of these two genera are saprotrophic or

pathogenic on different plants, but two of them, V. cinerescens and S. oculorum, have been

described as human opportunists of ocular infections (de Hoog et al. 2000, Guarro et al.
2002). Our molecular analysis shows that type species of Sarcopodium (S. circinatum) and

Volutella (V. ciliata) are distantly related to Phialemoniopsis in the Hypocreales. The isolates

of the respective opportunistic species wrongly assigned to those genera were clearly related

to P. curvata (FIG. 1). That is the case of V. cinerescens CBS 832.71, isolated from a case of

endophthalmitis, whose affinity to P. curvata was suggested by Proia et al. (2004). That

isolate, as well as other clinical isolates included in the Proia et al. (2004) study, developed

the typical sporodochium-like conidiomata described for P. curvata in the present study. It is

noteworthy, however, that such structures were not observed in the type strain and in two

other reference strains of P. curvata in any of the media tested in this study. These are

relatively old strains and we conjectured that they have degenerated, thereby losing the ability

to produce conidiomata.

The description of S. oculorum was based on two isolates obtained from a corneal

ulcer of a human patient (Guarro et al. 2002). These isolates were initially identified as

Phoma sp., but the development of sporodochial conidiomata favored re-identification as a

new species of Sarcopodium. Examination of additional clinical isolates of S. oculorum

showed sporodochial conidiomata intermixed with pycnidia-like conidiomata in all culture

media tested, a character shared with P. curvata. Therefore, the presence of conidiomata and

the phylogenetic relationships with the latter fungus support inclusion of S. oculorum in

Phialemoniopsis (FIG. 2).

In the D1/D2 analysis (FIG. 1), Phialemoniopsis was phylogenetically related to some

species of the genera Linocarpon (L. livistonae) and Neolinocarpon (N. globosicarpum and N.

enshiense). Those genera show anamorphs producing phialides with collarettes similar to

those present in Phialemoniopsis (Hyde et al. 1998, Bahl 2006). However, Huhndorf and

Miller (2011) demonstrated that Linocarpon is polyphyletic and the species of that genus and

of Neolinocarpon, particularly related with Phialemoniopsis, were not associated with any
known family. We could not include Neolinocarpon species in our combined analysis of

DNA phylogenies because strains of members of that genus are not available in any public

culture collection

Our results confirmed the anamorph-teleomorph association between Phialemonium

and Cephalotheca, and let us enlarge the circumscription of the genus Phialemonium to

include species producing conidia in slimy heads or in chains. We found a close phylogenetic

relationships and morphological similarities among P. obovatum, P.globosum, Taifanglania

inflata, Acremonium atrogriseum (FIGS. 1, 4). The genus Taifanglania was erected by Liang

et. al. (2009) to accommodate thermotolerant fungi, as well as some Paecilomyces species

showing single phialides on undifferentiated hyphae like Paecilomyces inflatus. The ex-type

strain of P. inflatus, isolated from a worker bee in USA, and the strain used by Liang et al.

(2009) to propose T. inflata, isolated from soil in Russia, were studied. They proved to be

conspecific and related to the type species of Phialemonium. It is worth mentioning that T.

inflata is the only Taifanglania species that is not thermotolerant. According to our

phylogenetic analysis, Phialemonium belongs to the Cephalothecaceae, while the other

members of Taifanglania are placed within the Chaetomiaceae (Liang et al. 2009). These

facts support the transference of T. inflata to Phialemonium, as P. inflatum. Phialemonium

inflatum is closely related to C. sulfurea and P. globosum (FIG. 4). Morphologically, they all

develop discrete phialides with more or less swollen bases, usually without visible collarettes,

and produce long persistent chains of conidia with connectives (Chester 1935, Onions and

Barron 1967, Liang et al. 2009). Differences among these species are summarized (TABLE

III). Our polyphasic taxonomic approach also supports the accommodation of A. atrogriseum

in the genus Phialemonium. This rare opportunistic fungus differs morphologically from the

other Phialemonium species by the lack of chlamydospores and by producing grayish brown

conidia without connectives, arranged in chains and slimy heads.

 Other anamorphic genera that have morphological similarities to Phialemonium are

Sagenomella W. Gams, Phialosimplex Sigler, Deanna A. Sutton, Gibas, Summerb. & Iwen,

Monocillium S.B. Saksena and Septofusidium W. Gams. However, all these fungi produce

only catenulate conidia. These genera are phylogenetically unrelated to Phialemonium; the

first two genera belong to the Trichocomaceae (Eurotiales), while Monocillium is placed in

the Niessliaceae and Septofusidium in the Nectriaceae, both latter within Hypocreales

(Luangsa-ard et al. 2004, Castlebury et al. 2004, Sigler et al. 2010).

Perdomo et al. (2011b) showed that Lecythophora and Coniochaeta constituted a well

defined group with several unidentified clinical isolates. Here, we have described two new

species closely related to L. hoffmannii (FIG. 3); however, other putative new species (i.e.

Lecythophora sp. 2–4) were not formally proposed because they could not be clearly

distinguished morphologically from the currently accepted Lecythophora species. The novel

species, L. luteorubra and L. cateniformis, are distinguished from each other mainly by the

presence of a characteristic diffusible pigment and the production chlamydospores,

respectively (TABLE IV). It is important to note that descriptions of those two species are

based on single isolate each. Therefore, phenotypic identification characters are susceptible to

be re-evaluated with the finding of new isolates.

ACKNOWLEDGMENTS

The authors are indebted to the curators of the Centraalbureau voor Schimmelcultures (Utrecht, the Netherlands),

BCCM™/IHEM Biomedical Fungi and Yeasts Collection (Brussels, Belgium) and NITE Biological Resource

Center (Chiba, Japan) for supplying many of the strains used in the study. This work was supported by the

Spanish Ministerio de Ciencia e Innovación grants CGL 2009-08698/BOS and CGL 2011-27185/BOS.

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LEGENDS

FIG. 1. Maximum likelihood (ML) tree obtained from D1/D2 domains of the 28S rDNA sequences of isolates

(TABLE I). Numbers on the branches are bootstrap ML values above 55%, followed by Bayesian posterior

probabilities (Bpp) above 0.56. Branch lengths are proportional to distance. Sequences not generated in this

study and obtained from the GenBank database are indicated in parentheses. Type strains of the different species

are indicated in boldface and with T.

FIG. 2. One of the two most parsimonious trees inferred from heuristic search of a combined alignment of the

domains D1/D2, ITS, ACT and TUB of the isolates included in clade I (FIG. 1). Bootstrap support values above

70% are indicated at the nodes. Lecythophora lignicola (CBS 267.33) was chosen as outgroup. GenBank

accession numbers for sequences included in this analysis are listed (TABLE I). T Type strain. New taxa are in

boldface.

FIG. 3. One of the two most parsimonious trees inferred from heuristic search of a combined alignment of the

domains D1/D2, ITS, ACT and TUB of the isolates included in the clade III (FIG. 1). Bootstrap support values

above 70% are indicated at the nodes. Phialemonium obovatum (CBS 279.76) was chosen as outgroup. GenBank

accession numbers for sequences included in this analysis are provided (TABLE I). The news species are

indicated in boldface. T Type strain. New taxa are in boldface.

FIG. 4. The most parsimonious tree inferred from heuristic search of a combined alignment of the domains

D1/D2, ITS, ACT and TUB of the isolates included in clade IV (FIG 1). Bootstrap support values above 70% are

indicated at the nodes. Lecythophora lignicola (CBS 267.33) was chosen as outgroup. GenBank accession

numbers for sequences included in this analysis are provided (TABLE I). T Type strain. New taxa are in boldface.
FIG. 5. Phialemoniopsis cornearis, UTHSC 06-1820T. A. Colony surface on OA after 14 d at 25 C. B. Colony

surface on OA after 6 mo incubation at 25 C with conidiomata. C. Conidiophore from aerial mycelium. D–L.

Phialides, adelophialides (arrows) and conidia from aerial mycelium. M. Chlamydospore. N, O. Sporodochium-

like conidiomata. P. Conidiophores lining the inner wall of sporodochium-like conidiomata. Q, R. Detail of

branched conidiophoes and conidia from sporodochium-like conidiomata. S, T. Pycnidium-like conidiomata

with wall of textura intricata. U, V. Detail of branched conidiophores from pycnidium-like conidiomata and

conidia. Bars: C–I, M, Q = 10 μm; J–L, R, V = 5 μm; N, O, S = 100 μm; T = 50 μm; P, U = 20 μm.

FIG. 6. Phialemoniopsis ocularis, IHEM 19077T. A–E. Phialides, adelophialides and conidia from the aerial

mycelium. F, G. Pycnidium-like conidiomata. H–J. Conidiophores lining the inner wall of a sporodochium-like

conidioma and conidia. K, L. Chlamydospores. Conidiophores from a pycnidium-like conidiomata and conidia.

Bars: A, B, D, E, H, K = 10 μm; C, J–L = 5 μm; F, G = 50 μm.

FIG. 7. Phialemoniopsis pluriloculosa, UTHSC 04-7T. A. Colony surface on OA after 14 d at 25 C. B. Colony

surface on OA after 6 mo incubation at 25 C with dark pigmented conidiomata (arrow). C. Chlamydospores.

D–F. Phialides, adelophialides (arrow) and conidia from the aerial mycelium. G, H, J. Multilocular pycnidium-

like conidiomata. I. Conidiophores lining the inner wall of a sporodochium-like conidiomata. K, L.

Conidiophores and conidia from a pycnidium-like conidiomata. Bars: G, H, J = 100 μm; C, D, I, K = 10 μm; E,

F, L = 5 μm.

FIG. 8. A–G. Phialemonium atrogriseum, CBS 604.67T. A, B. Colony surface and reverse on OA after 14 d at

25 C. C–G. Phialides, some with a visible collarette (arrow) and conidia. H–L. Phialemonium globosum,

UTHSC 03-3661T. H. Colony surface on OA after 14 d at 25 C. I–L phialides and conidia in chain. M–Q.

Phialemonium inflatum, CBS 259.39T. M. Colony surface on OA after 14 d at 25 C. N. Conidiophores and

conidia. O. Conidia in chains. P, Q. Phialide and conidia. Bars: C, I, J, N = 10 μm; D–F, G, K, L, O–Q = 5

μm.

FIG. 9. A–I. Lecythophora cateniformis, UTHSC 01-1644T. A, B. Colony surface on PDA and OA respectively

after 14 d at 25 C. C, D. Phialides without basal septum. E, F. Conidia. G, H, I. Chlamydospores. J–P.

Lecythophora luteorubra, UTHSC 01-20-1T. J, K. Colony surface and reverse on PDA after 14 d at 25 C. L.

Colony surface and diffusible pigment on PDA after 2 mo incubation at 25 C. M. Phialides and conidia in slimy

heads. N, O. Phialides without basal septum. P. Conidia. Bars: G–I, M–O = 10 μm; C–F, P = 5 μm.

FOOTNOTES
Submitted 19 Apr 2012; accepted for publication 11 Sep 2012.
1
Corresponding author. E-mail: josepa.gene@urv.cat
TABLE I. Phialemonium and Lecythophora clinical isolates and type or reference strains of related species included in the study

Isolate Origin Species GenBank accession number

(Original identification) D1/D2 rDNA ACT ITS TUB
T
UTHSC 01-20-1 (Lecythophora sp. 1) Leg wound, USA Lecythophora luteorubra HE610328 HE610338 HE610330 HE610346
UTHSC 01-317 (Phialemonium obovatum) Graft tissue, USA Phialemonium obovatum FR745951 HE599317 HE610355 HE599336
UTHSC 01-1399 (Phialemonium obovatum) Abscess, USA Phialemonium obovatum FR745945 HE599318 HE610354 HE599337
T HE610339 HE610331 HE610347
UTHSC 01-1644 (Lecythophora sp. 1) Canine bone marrow, USA Lecythophora cateniformis HE610329
UTHSC 03-986 (Acremonium atrogriseum) BAL, USA Phialemonium atrogriseum FR691979 HE599324 HE610359 HE599343
UTHSC 03-2258 (Phialemonium obovatum) Arm, USA Phialemonium obovatum FR745950 HE599313 HE610356 HE599332
UTHSC 03-2653 (Cephalotheca foveolata) Endocarditis, Singapure Cephalotheca foveolata FR745941 HE599323 HE610357 HE599342
UTHSC 03-3661 T (Phialemonium sp. 2) Sinus, USA Phialemonium globosum HE610461 HE599330 HE610358 HE599349
UTHSC 04-7 T (Volutella sp.) Toe nail, USA Phialemonipsis pluriloculosa HE599271 HE599254 HE599286 HE599303
UTHSC 04-616 (Phialemonium obovatum) Arm, USA Phialemonium obovatum FR745944 HE599314 HE610360 HE599333
UTHSC 04-956 (Phialemonium curvatum) Sinus, USA Phialemoniopsis curvata HE599278 HE599263 HE599295 HE599312
UTHSC 05-2527 (Phialemonium sp. 1) Peritoneal dialysis catheter, USA Phialemoniopsis ocularis HE599266 HE599249 HE599281 HE599298
UTHSC 06-733 (Cephalotheca foveolata) Lymph node Bx, USA Cephalotheca foveolata FR745943 HE599320 HE610361 HE599339
UTHSC 06-1465 (Phialemonium sp. 1) Shin aspirate, USA Phialemoniopsis cornearis HE599270 HE599253 HE599285 HE599302
UTHSC 06-1820 T (Phialemonium sp. 1) Corneal fluid, USA Phialemoniopsis cornearis HE599269 HE599252 HE599284 HE599301
UTHSC 06-4324 (Phialemonium curvatum) Canine pleural fluid, USA Phialemoniopsis curvata HE599273 HE599256 HE599288 HE599305
UTHSC 07-1556 (Cephalotheca foveolata) Bronchial, USA Cephalotheca foveolata FR691996 HE599322 HE610362 HE599341
UTHSC 07-3736 (Phialemonium sp. 1) Left hand, USA Phialemoniopsis ocularis HE599268 HE599251 HE599283 HE599300
UTHSC 08-2292 (Phialemonium curvatum) Blood, USA Phialemoniopsis curvata HE599277 HE599262 HE599294 HE599311
UTHSC 08-2766 (Cephalotheca foveolata) Eye capsule, USA Cephalotheca foveolata FR691995 HE599319 HE610363 HE599338
UTHSC 09-2358 (Phialemonium sp. 1) Aspirate cellulitis, USA Phialemoniopsis ocularis HE599267 HE599250 HE599282 HE599299
UTHSC 09-3589 (Volutella sp.) Synovial fluid, USA Phialemoniopsis pluriloculosa HE599272 HE599255 HE599287 HE599304
UTHSC R-3447 (Phialemonium curvatum) Eye, Israel Phialemoniopsis curvata HE599274 HE599259 HE599291 HE599308
UTHSC R-3448 (Phialemonium curvatum) Eye, Israel Phialemoniopsis curvata HE599275 HE599260 HE599292 HE599309
CBS 135.34 Garden cane, United Kingdom Cephalotheca sulfurea HE610462 HE599331 HE610372 HE599350
T HE610340 HE610337 HE610348
CBS 153.42 Fruit, the Netherlands Lecythophora decumbens HE610463
T HE610341 HE610334 HE610349
CBS 157.44 River water, Germany Lecythophora mutabilis FR691990
T HE610342 HE610336 HE610350
CBS 205.38 Butter, Switzerland Lecythophora fasciculata FR691988
T HE610343 HE610333 HE610351
CBS 206.38 Butter, Switzerland Lecythophora luteoviridis FR691987
T HE610344 HE610332 HE610352
CBS 245.38 Butter, Switzerland Lecythophora hoffmannii FR691982
TABLE I (cont.)
Isolate Origin Species GenBank accession number
(Original identification) D1/D2 rDNA ACT ITS TUB
T HE599328 HE610364 HE599347
CBS 259.39 (Paecilomyces inflatus) Apis mellifera, USA Phialemonium inflatum HE610464
T HE610345 HE610335 HE610353
CBS 267.33 Unknown, Sweden Lecythophora lignicola FR691986
CBS 279.76T Systemic infection, USA Phialemonium obovatum FR691997 HE599315 HE610365 HE599334
CBS 298.64 Air, the Netherlands Volutella ciliata HE610465
CBS 306.85 (Acremonium atrogriseum) Lung, USA Phialemonium atrogriseum HE610466 HE599326 HE610366 HE599345
CBS 360.35 Soil, the Netherlands Volutella ciliata HE610467
CBS 376.81 Dead stem, the Netherlands Sarcopodium circinatum HE610468
CBS 408.80 Dead stem, the Netherlands Sarcopodium circinatum HE610469
CBS 457.88 Ganoderma applanatum, Germany Albertiniella polyporicola AF096185a
CBS 490.82T (Phialemonium curvatum) Skin lesion, USA Phialemoniopsis curvata FR691977 HE599258 HE599290 HE599307
CBS 491.82T (Phialemonium dimorphosporum) Soil, USA Phialemoniopsis curvata FR691976 HE599257 HE599289 HE599306
CBS 508.70 T Wood, Canada Cryptendoxyla hypophloia AB191032b
CBS 551.75 Wood, Norway Coniochaeta subcorticalis AF353593c
CBS 604.67T (Acremonium atrogriseum) Noodles, Ukraine Phialemonium atrogriseum HE610470 HE599327 HE610367 HE599346
CBS 730.97 Peritoneal dialysis fluid, USA Phialemonium obovatum HE610471 HE599316 HE610368 HE599335
CBS 774.97 (Acremonium atrogriseum) Urine, Germany Phialemonium atrogriseum HE610472 HE599325 HE610369 HE599344
CBS 832.71 (Volutella cinerescens) Keratomycosis, Unknown Phialemoniopsis curvata HE599276 HE599261 HE599293 HE599310
d
CBS 109872 Wood, Argentina Coniochaeta rhopalochaeta GQ351561
CBS 110467 Wood, Germany Coniochaeta ligniaria AF353583c
CBS 120873T Wood, South Africa Collophora rubra GQ154606e
CBS 120875 Prunus armeniaca, South Africa Coniochaeta prunicola GQ154602e
E. W. 93.262 Wood, Germany Coniochaeta velutina AF353594c
E. W. 95.605 Wood, Germany Coniochaeta ligniaria AF353584c
IHEM 19077 T (Sarcopodium oculorum) Keratomycosis, Brazil Phialemoniopsis ocularis HE599264 HE599247 HE599279 HE599296
FMR 7190 (Sarcopodium oculorum) Keratomycosis, Brazil Phialemoniopsis ocularis HE599265 HE599248 HE599280 HE599297
FMR 8227 Nematode eggs, Spain Volutella ciliata HE610475
HKUCC 2984 Neolinocarpon globosicarpum DQ810224f
HKUCC 2954 Linocarpon livistonae DQ810206f
HKUCC 2983 Neolinocarpon enshiense DQ810221f
NBRC 100905T Subcutaneous infection, Korea Cephalotheca foveolata HE610473 HE599321 HE610370 HE599340
NBRC 31965 (Taifanglania inflata) Cabbage field soil, Russia Phialemonium inflatum HE610474 HE599329 HE610371 HE599348
T
Type strains; CBS, Centraalbureau voor Schimmelcultures, Utrecht, the Netherlands; FMR, Facultat de Medicina i Ciencies de la Salut, Reus, Spain; HKUCC, University
of Hong Kong Culture Collection; IHEM, BCCM/IHEM Biomedical Fungi and Yeasts Collection, Brussels, Belgium; NBRC, NITE Biological Resource Center, Chiba,
Japan; UTHSC, Fungus Testing Laboratory, University of Texas Health Science Center at San Antonio, USA; Accession number sequences retrieved from GenBank
database are in bold face; a Suh and Blackwell (1999); b Yaguchi et al. (2006); c Weber et al. (2002); d Heerden et al. (2011); e Damm et al. (2010); f Bahl (2006).
TABLE II. Distinctive morphological features of Phialemoniopsis species based on OA cultures at 25 C after 14 d
Species Colonies Sporodochium-like conidiomata Pycnidium-like conidiomata Other distinctive features
P. cornearis Whitish to olive-brown Solitary, superficial, hyaline. Conidia Solitary, superficial, globose to subglobose, dark Chlamydospores present
hyaline, obovoid to ellipsoidal brown to black, unilocular, textura intricata.
Conidia hyaline, obovoid, slightly apiculate at
the base
P. curvata White, brown to grayish Solitary, superficial, hyaline to subhyaline. Absent Chlamydospores absent
Conidia hyaline, ellipsoid to ovoid

P. ocularis Whitish to brownish Solitary, superficial, hyaline to subhyaline. Solitary, superficial, globose to subglobose, dark Chlamydospores present
gray Conidia subhyaline, ellipsoid, navicular or brown to black, unilocular, textura intricata and
slightly allantoid epidermoidea. Conidia hyaline, ellipsoidal to
lightly allantoid, slightly apiculate at the base

P. pluriloculosa Yellowish white, olive- Solitary, superficial, multilocular, Solitary or in groups, superficial or semi- Chlamydospores present
brown to black subhyaline to brown. Conidia hyaline, immersed, globose to subglobose, dark brown to
ellipsoidal black, multilocular, textura epidermoidea.
Conidia hyaline, ellipsoidal, slightly apiculate at
the base
TABLE III. Distinctive morphological features of Phialemonium species based on OA cultures at 25 C after 14 d
Species Colonies Conidia Other distinctive features
P. atrogriseum Pale ocher-brown to Subhyaline to dark gray, ellipsoidal or Sometimes, phialides with a short pale
brownish black obovoid with an apiculate and minutely collarette. Ascomata and chlamydospores
truncate base, in slimy heads and in chains absent
without connectives
P. globosum White Hyaline, globose to subglobose, connective Chlamydospores and ascomata absent
on both ends, in chains
P. inflatum White to pale yellow Hyaline, citriform, connective on both Chlamydospores and ascomata absent
ends, in chains
P. obovatum Pale ochraceous to Hyaline, obovoid with a truncate base, in Chlamydospores hyaline, oval and thin-
greenish slimy heads walled. Some isolates develop globose and
dark structures suggestive of immature
ascomata
Phialemonium White to brownish Hyaline, cylindrical, in slimy heads Chlamydospores hyaline, ellipsoidal, thick-
anamorph of walled. Ascomata present, covered with
Cephalotheca sulfurous to glabrous hyphae; ascospora
foveolata elliptic-ovate, foveolate-walled
Phialemonium White to pale yellow Hyaline, ovoid, connective on both ends, in Chlamydospores absent. Ascomata present,
anamorph of chains covered with light yellow to brown hyphal
Cephalotheca sulfurea hairs; ascospores broadly reniform, smooth-
walled
TABLE IV. Distinctive morphological features of Lecythophora species, based on OA cultures at 25 C after 14 d
Species Colonies Conidia Chlamydospores Other distinctive features
L. cateniformis Brownish gray Obovoid to ellipsoidal, with a Pale brown to brown, subglobose, Adelophialides are
protruding and slightly truncate oval to pyriform, solitary or in predominating
base short chains
L. decumbens Olivaceous to Ellipsoidal to cylindrical, often Absent Presence of conidia swollen up
brown somewhat curved to 13 × 3–4.5 μm
L. fasciculata Gray-brown to Ovoid to ellipsoidal, often Absent Adelophialides are
black somewhat curved predominating
L. hoffmannii Orange to salmon Ellipsoidal to cylindrical, often Absent More common short
somewhat curved adelophialides than ventricose
phialides
L. luteorubra Orange Ellipsoidal to cylindrical, slightly Absent Colonies producing a diffusible
apiculate at the base pigment orange-red to
brownish-red
L. luteoviridis Olivaceous to Ellipsoidal to cylindrical, often Hyaline or faintly brown, globose Darkening of the colonies due
brown somewhat curved to oblong or pear-shaped, singly to development of pigmented
or short chains chlamydospores
L. lignicola Brown Ovoid to ellipsoidal Absent Discrete ventricose phialides
abundant
L. mutabilis Brown Ellipsoidal, rarely curved Olivaceous-brown, ovoid to Discrete ventricose phialides
ellipsoidal, more rarely globose, abundant
singly
 CBS 490.82T Phialemonium curvatum
CBS 491.82
UTHSC 06-4324
UTHSC R-3447
99/1 UTHSC R-3448
CBS 832.71
UTHSC 08-2292
99/1 UTHSC 04-956
UTHSC 09-3589
UTHSC 06-1465
Clade I
UTHSC 06-1820 Incertae sedis
UTHSC 09-2358 Incertae sedis
87/0,9
UTHSC 07-3736
IHEM 19077T Sarcopodium oculorum
FMR 7190
UTHSC 05-2527

UTHSC 04-7
Linocarpon livistonae (DQ810206)
94/0,98 Neolinocarpon globosicarpum (DQ810224)
Neolinocarpon enshiense (DQ810221)

90/0,96 CBS 408.80 Sarcopodium circinatum

Incertae sedis
91/1 CBS 376.81 Sarcopodium circinatum Clade II
99/1 CBS360.35 Volutella ciliata Hypocreales
CBS298.64 Volutella ciliata
Nectriaceae
74/0,56 CBS 267.33T Lecythophora lignicola
CBS 206.38T Lecythophora luteoviridis
E. W. 93.262 Coniochaeta velutina (AF353594)
CBS 205.38T Lecythophora fasciculata
92/1
CBS 110467 Coniochaeta ligniaria (AF353583)
89/0,97 E. W. 95.605 Coniochaeta ligniaria (AF353584)
CBS 157.44T Lecythophora mutabilis
77/0,99
CBS 120875T Coniochaeta prunicola (GQ1546031)
Coniochaetaceae Coniochaetales
UTHSC 01-1644
55/0,88
UTHSC 01-20-1
CBS 245.38T Lecythophora hoffmannii
CBS 551.75 Coniochaeta subcorticalis (AF353593)
98/1 CBS 109872T Coniochaeta rhopalochaeta (GQ351561)
CBS 153.42T Lecythophora decumbens
CBS 508.70T Cryptendoxyla hypophoia (AB191032)
78/0,96
CBS 259.39T Paecilomyces inflatus
NBRC 31965 Taifanglania inflata
80/1
CBS 135.34 Cephalotheca sulfurea
76/1 UTHSC 03– 3661
70/0,88 CBS 604.67T Acremonium atrogriseum
CBS 306.85
99/1
CBS 774.97
UTHSC 03–986
CBS 457.88T (AF096185)
Albertiniella polyporicola Clade III
UTHSC 06–733 Cephalothecaceae
UTHSC 07–1556 Sordariales
97/1 UTHSC 08–2766

NBRC 100905T Cephalotheca foveolata

UTHSC 03–2653
--/0,99 CBS 730.97
CBS 279.76T Phialemonium obovatum
69/0,98
UTHSC 01–1399
71/0,86 UTHSC 04–616
UTHSC 03–2258
UTHSC 01–317
CBS120873 Collophora rubra (GQ154606) Helotiales
0.05
 D1-D2, ITS, ACT, TUB
2135 bp 100 IHEM 19077 T Sarcopodium oculorum
FMR 7190
Tree length= 222 87
CI=0.77 Phialemoniopsis
100 UTHSC 05–2527 ocularis
HI=0.23
RI=0.92 UTHSC 09–2358
81 86
UTHSC 07–3736

UTHSC 06–1820 T
100 100 Phialemoniopsis cornearis
UTHSC 06–1465

UTHSC 04–7 T
96 Phialemoniopsis pluriloculosa
UTHSC 09–3589

UTHSC 06–4324
83 CBS 491.82

CBS 490.82 T Phialemonium curvatum

94
UTHSC R–3447
Phialemoniopsis curvata
UTHSC R–3448
100 CBS 832.71 Volutella cinerescens

UTHSC 08–2292
98 UTHSC 04–956

CBS 267.33 T Lecythophora lignicola

1
D1-D2, ITS, ACT, TUB
2173 bp CBS 205.38 T Lecythophora fasciculata
Tree length= 514 77
CI=0.65 100 CBS 267.33 T Lecythophora lignicola
HI=0.35
RI=0.55< 98 CBS 206.38 T Lecythophora luteoviridis

CBS 157.44 T Lecythophora mutabilis

100
UTHSC 01–20-1T Lecythophora luteorubra

UTHSC 01–1644 T Lecythophora cateniformis

CBS 245.38 T Lecythophora hoffmannii

CBS 153.42 T Lecythophora decumbens

CBS 279.76 T Phialemonium obovatum

10
 D1-D2, ITS, ACT, TUB UTHSC 03–2258
2175 bp UTHSC 04–616
CI=0.79
CBS 279.76 T
HI=0.20 Phialemonium obovatum
RI=0.89 CBS 730.97
100
UTHSC 01–317

UTHSC 01–1399

100 UTHSC 08–2766

UTHSC 06–733

100 NBRC 100905 T Cephalotheca foveolata

UTHSC 07–1556

UTHSC 03–2653

100
UTHSC 03–986
CBS 774.97
100
Phialemonium atrogriseum
CBS 306.85
90 CBS 604.67 T Acremonium atrogriseum

100 CBS 259.39 T Paecilomyces inflatus

Phialemonium inflatum
NBRC 31965 Taifanglania inflata
100 UTHSC 03–3661T Phialemonium globosum

CBS 135.34 Cephalotheca sulfurea

CBS 267.33T Lecythophora lignicola

10