Taxonomy of Ochroconis, genus including opportunistic pathogens on humans and animals

Abstract

The genus Ochroconis (Sympoventuriaceae, Venturiales) is revised and currently contains 13 species for which the phylogenetic position has been determined using multilocus sequencing. The older generic name Scoleco-basidium is considered to be of doubtful identity because the type specimen is ambiguous. Within the Ochroconis lineage, phylogenetic distances of all markers analyzed are exception-ally large, both between and within species. A new genus Verruconis is proposed for the neurotropic opportunist Ochroconis gallopava. Species accepted within the lineages are keyed out on the basis of phenotypic characters. Main ecological traits within each species are discussed. Verruconis species are thermophilic and one of them is an important agent of infection in the brain, while Ochroconis is mesophilic, several species causing infec-tions in cold-blooded animals.

Full text

Taxonomy of Ochroconis, genus including opportunistic
pathogens on humans and animals
K. Samerpitak &E. Van der Linde &H.-J. Choi &
A. H. G. Gerrits van den Ende &M. Machouart &
C. Gueidan &G. S. de Hoog
Received: 18 March 2013 /Accepted: 25 June 2013
#Mushroom Research Foundation 2013
Abstract The genus Ochroconis (Sympoventuriaceae,
Venturiales) is revised and currently contains 13 species for
which the phylogenetic position has been determined using
multilocus sequencing. The older generic name Scoleco-
basidium is considered to be of doubtful identity because the
type specimen is ambiguous. Within the Ochroconis lineage,
phylogenetic distances of all markers analyzed are exception-
ally large, both between and within species. A new genus
Verruconis is proposed for the neurotropic opportunist
Ochroconis gallopava. Species accepted within the lineages
are keyed out on the basis of phenotypic characters. Main
ecological traits within each species are discussed.
Verruconis species are thermophilic and one of them is
an important agent of infection in the brain, while
Ochroconis is mesophilic, several species causing infec-
tions in cold-blooded animals.
Keywords Taxonomy .Ecology .Thermophilic fungi .
Human infection .Avian pathogen .Fish pathogen .
Neurotropism
Taxonomic novelties: Ochroconis cordanae Samerpitak, Crous & de
Hoog, Ochroconis mirabilis Samerpitak & de Hoog, Ochroconis
minima (Fassat.) Samerpitak & de Hoog, Ochroconis longiphorum
(Matsush.) Samerpitak & de Hoog, Ochroconis sexualis Samerpitak,
Van der Linde & de Hoog, Ochroconis verrucosa (Zachariah, Sankaran
& Leelav.) Samerpitak & de Hoog, Verruconis Samerpitak & de Hoog,
Verruconis calidifluminalis (Yarita, Sano, de Hoog & Nishim.)
Samerpitak & de Hoog, Verruconis gallopava (W.B. Cooke)
Samerpitak & de Hoog, Verruconis verruculosa (R.Y. Roy, R.S.
Dwivedi & R.R. Mishra) Samerpitak & de Hoog
K. Samerpitak :A. H. G. Gerrits van den Ende :G. S. de Hoog
CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands
K. Samerpitak
Department of Microbiology, Faculty of Medicine,
Khon Kaen University, Khon Kaen, Thailand
K. Samerpitak :G. S. de Hoog (*)
Institute for Biodiversity and Ecosystem Dynamics,
University of Amsterdam, Amsterdam, The Netherlands
e-mail: de.hoog@cbs.knaw.nl
E. Van der Linde
Mycology Unit, Biosystematics Programme, Plant Protection
Research Institute, Agricultural Research Council,
Pretoria, South Africa
H.<J. Choi
Evangelisches und Johanniter Klinikum Niederrhein,
Duisburg, Germany
M. Machouart
Service de Parasitologie-Mycologie, CHU de Brabois,
Vandœuvre-lès-Nancy, France
C. Gueidan
Department Life Sciences, Division of Microbial & Genomic
Diversity, Natural History Museum, London, UK
G. S. de Hoog
Peking University Health Science Center, Research Center for
Medical Mycology, Beijing, China; Sun-Yat-sen Memorial
Hospital, Sun Yat-sen University, Guangzhou, China
G. S. de Hoog
Second Medical Military University, Shanghai, China
Fungal Diversity
DOI 10.1007/s13225-013-0253-6

Introduction
In 1927 Abbott introduced a group of filamentous fungi with
rather characteristic cultural and morphological features. All
strains had rust-brown to olivaceous colonies and produced
small, brownish conidiophores bearing small numbers of
dark, septate, rough-walled conidia. Later the process of
conidium liberation was described to be rhexolytic (Ellis
1971), a rare phenomenon in filamentous fungi. The group
was described as Scolecobasidium, with S.terreum Abbott as
type species. The type strain, CBS 203.27, which is now
sterile, had conidia that were Y-shaped. Matsushima (1975)
added several species with elaborately branched conidia,
which led de Hoog and von Arx (1973) to erect another
genus, Ochroconis, typified by O.constricta (Abbott) de
Hoog & v. Arx for the species with unbranched, subspherical
to cylindrical or clavate conidia. The original isolate of
Scolecobasidium constrictum Abbott, CBS 202.27, present-
ly also fails to produce conidia. The number of species in the
generic complex has increased significantly over the years
(de Hoog 1985).
Ochroconis was introduced into medical and veterinary
mycology after the report of a kidney mycosis in Chinook
salmon (Oncorhynchus tshawytschae)byO.tshawytschae
(Doty & Slater) Kirilenko & Al-Achmed (Doty and Slater
1946). The species was originally described as Heterosporium
tshawytschae Doty & Slater, based on type strain ATCC 9915
(= CBS 100438). Later a visceral infection in cultured Masu
salmon (Oncorhynchus masou) was described (Hatai and
Kubota 1989), caused by an Ochroconis species closely relat-
ed to O.tshawytschae.
Ochroconis humicola (Barron & Busch) de Hoog & v. Arx
has also repeatedly been reported as an etiologic agent of
infections in cold-blooded (thermal-conforming) animals. Like
O.tshawytschae, this species has been reported from infections
in salmon. A kidney infection in Coho salmon (Oncorhynchus
kisutch) was reported by Ross and Yasutake (1973)anda
muscular black spot disease was described in Atlantic salmon
(Salmo salar) by Schaumann and Priebe (1994). Further dis-
eases in cold-blooded animals ascribed to this species were an
outbreak in rainbow trout (Ajello et al. 1977), an infection in a
tortoise (Weitzman et al. 1983) and infections in marine cul-
tured fishes such as a devil stinger (Inimicus japonicus)(Wada
et al. 1995), a red sea bream (Pagus major)andaarbled
rockfish (Sebastiscus marmoratus)(Wadaetal.2005).
Judging from these numerous case reports, Ochroconis-
like fungi are consistent etiologic agents of disease in cold-
blooded animals. It is unfortunate that only very few strains
from these reports have been preserved, which makes it
impossible to ascertain which species were involved. Only
a single infection was noted in a warm-blooded animal: a
subcutaneous lesion in a cat (VanSteenhouse et al. 1988).
Ochroconis species morphologically similar to O.humicola
have been isolated from wet areas in the domestic environ-
ment, such as bathrooms (Lian and de Hoog 2010).
Additional animal infections by Ochroconis species concern
poultry and wild birds, which have relatively high body temper-
atures; an overview of published animal cases is given in Table 1.
Ochroconis gallopava (W.B. Cooke) de Hoog was the etiologic
agent of avian brain infections. Several large epidemics occurred
in the U.S.A. (Georg et al. 1964; Blalock et al. 1973; Ranck et al.
1973; Randall and Owen 1981; Shane et al. 1985; Karesh et al.
1987; Salkin et al. 1990), with one report from Australia
(Connole 1967) and India (Mohapatra 1993). In addition to avian
infections, systemic mycoses in a cat (Padhye et al. 1994)anda
dog (Singh et al. 2006) were reported, also from the U.S.A.
The first human case was also reported from the U.S.A.;
this concerned a lung infection (Dixon and Salkin 1986). To
date, 35 human cases of systemic infection by O.gallopava
had been reported from many countries worldwide (Horré and
de Hoog 1999). Lung and brain were the main target organs,
and most patients were immunocompromised as a result of
immunosuppressive drugs administered after organ transplan-
tation (Mancini and McGinnis 1992; Prevost-Smith et al.
1993;Vukmiretal.1994; Kralovic and Rhodes 1995;
Rossmann et al. 1996;Bonhametal.1996; Singh et al.
1997;Jenneyetal.1998; Horré and de Hoog 1999; Burns
et al. 2000; Malani et al. 2001;Wangetal.2003; Ohori et al.
2006). Other underlying conditions of patients were leukemia
(Terreni et al. 1990; Sides et al. 1991; Fukushima et al. 2005),
diabetes mellitus (Horré and de Hoog 1999), chronic pemphi-
gus autoimmune disorder (Zhao et al. 2002), and occupational
dust-related pneumoconiosis (Ohori et al. 2006). Subcutane-
ous infections were noted in a leukemic patient (Fukushiro
et al. 1986) and in a lung-transplant recipient with diabetes
(Mazur and Judson 2001). Bowyer et al. (2000)reportedan
eye infection by O.gallopava in a patient with lymphoma. In
addition to cases by O.gallopava in patients with significant
underlying disorders, occasionally infections in otherwise
healthy individuals were noted. These concerned occupational
lung diseases in coal mine workers (Sides et al. 1991)andin
wood pulp workers (Odell et al. 2000). In summary, O.
gallopava may be regarded as a virulent pathogen, with a
significant health risk for imunocompromised patients.
The environmental habitat of Ochroconis gallopava indi-
cates a significant thermophily of the species. It has been
isolated from thermal soils (Evans 1971a,b; Tansey and Brock
1973;Weitzmanetal.1983;Redmanetal.1999), broiler-house
litter (Waldrip et al. 1974; Randall and Owen 1981), hot springs
(Tansey and Brock 1973; Weitzman et al. 1983; Yarita et al.
2007), the warm effluent of a nuclear reaction station (Rippon
et al. 1980), and from self heated coal waste piles (Tansey and
Brock 1973). Recently a sibling species, O.calidifluminalis
Yari t a e t al. ( 2010) was described from the same hot springs
where O.gallopava was isolated, but with significantly lower
virulence.
Fungal Diversity

Table 1 Strains of Ochroconis and relatives used in phylogenetic analyses
Current/obsolete name Culture/voucher Source Country GenBank accession number
SSU ITS LSU ACT1 BT2 TEF1
Ochroconis
O.anellii/id. CBS 284.64 (T) Stalactite Italy KF156070 FR832477 KF156138 KF155912 KF156184 KF155995
O.anomala/id. CBS 131816 (T) Lascaux Cave France KF156065 HE575201 KF156137 KF155935 KF156194 KF155986
O.constricta/id. CBS 202.27 (T) Soil USA KF156072 AB161063 KF156147 KF155942 KF156161 KF156003
O.constricta/id. CBS 211.53 (T) Soil Canada KF156073 HQ667519 KF156148 KF155941 KF156187 KF156005
O.constricta/O.humicola CBS 106.65 Soil Germany HQ667518
O.constricta/id. CBS 269.61 Peat soil Canada KF156074 KF156024 KF156149 KF155939 KF156163 KF156004
O.constricta/id. CBS 381.64 Soil India HQ667517
O.mirabilis/O.humicola CBS 100486 Devil stinger (marine fish) Japan KF156026
O.constricta/O. sp. CBS 131913, dH 22342 Human, cutaneous mycosis Thailand KF156071 KF156025 KF156146 KF155940 KF156176 KF156006
O.cordanae/O.constricta CBS 475.80 (T) Dead leaf Colombia KF156058 KF156022 KF156122 HQ916976 KF156197 KF155981
O.cordanae/O.humicola CBS 172.74 Dead leaf India KF156057 KF156023 KF156121 KF155906 KF156198 JF440566
O.cordanae/O.constricta CBS 412.51 ND USA KF156056 HQ667540 KF156123 KF155907 KF156200 KF155980
O.cordanae/O.humicola CBS 780.83 Podocarpus litter Japan KF156059 HQ667539 KF156120 KF155905 KF156199 KF155979
O.cordanae/O. sp. CBS 123536 Living leaf South Africa FJ372390 –
O.cordanae/Fungal endophyte 9165 Conifer endophyte USA EF419939 –
O.cordanae/Scopulariopsis humicola ATT236 Ant nest USA HQ607921 –
O.cordanae/Dothideomycete FL0760 Endophyte USA JQ760421 –
O.cordanae/unnamed JIA6-8-1 Living root (Oryza) China FJ752621 –
O.cordanae/S.humicola TR072 Ant nest USA HQ608103 –
O.gamsii/id. CBS 239.78 (T) Plant leaf Sri Lanka KF156088 KF156019 KF156150 KF155936 KF156190 KF155982
O.aff.gamsii/O.gamsii CBS 101179 Dead rachis, Palm tree Puerto Rico KF156091 KF156020 KF156151 KF155937 KF156192
O.aff.gamsii/O.gamsii CBS 102491 Soil Spain KF156092 KF156021 KF156152 KF155938 KF156191 KF155983
O.humicola/id. CBS 116655 (T) Peat soil Canada KF156068 HQ667521 KF156124 KF155904 KF156195 KF155984
O.lascauxensis/id. CBS 131815 (T) Stain in Lascaux Cave France KF156069 FR832474 KF156136 KF155911 KF156183 KF155994
O.lascauxensis/id. LX7.2 Stain in Lascaux Cave France HE575198
O.lascauxensis/id. LX101R-6 Stain in Lascaux Cave France HE575200
O.lascauxensis/id. LXA3 Stain in Lascaux Cave France HE575199
O.lascauxensis/id. LX CH2 Stain in Lascaux Cave France HE575195
O.lascauxensis/id. LX CH3 Stain in Lascaux Cave France HE575196
O.lascauxensis/id. LX CH7 Stain in Lascaux Cave France HE575197
O.lascauxensis/id. LXM8-5 Stain in Lascaux Cave France HE605221
O.longiphorum/S.longiphorum CBS 435.76 Soil Canada KF156060 KF156038 KF156135 KF155908 KF156182 KF155978
O.minima/Humicola minima CBS 510.71 (T) Rhizosphere Nigeria KF156087 HQ667522 KF156134 KF155945 KF156172 KF156007
Fungal Diversity

Table 1 (continued)
Current/obsolete name Culture/voucher Source Country GenBank accession number
SSU ITS LSU ACT1 BT2 TEF1
O.minima/S.terreum CBS 423.64 Rhizosphere Netherlands KF156085 HQ667523 KF156131 KF155943 KF156173 KF156008
O.minima/S.terreum CBS 536.69 Forest soil Canada KF156084 HQ667524 KF156132 KF155944 KF156174 KF156009
O.minima/S.terreum CBS 116645 Sandy soil Canada KF156083 HQ667525
O.minima/S.tereum CBS 119792 Soil India KF156086 KF156027 KF156133 KF155946 KF156175
O.minima/S. sp. ASR 130 Rhizosphere Brazil GU973693
O.minima/S. sp. ASR 131 Rhizosphere Brazil GU973694
O.minima/S.terreum NBRC 9845 Aquatic sediment Japan DQ307328
O.mirabilis/O.humicola CBS 729.95 (T) Regulator of diver Netherlands KF156082 KF156029 KF156144 KF155948 KF156171 KF155999
O.mirabilis/O.humicola dH 20186 Bathroom Netherlands KF156037
O.mirabilis/O.constricta CBS 124.65 Human, skin lesion India HQ667532 KF155951 KF156168 KF155996
O.mirabilis/O.constricta CBS 413.51 ND USA KF156076 HQ667536 KF156140 KF155957 KF156164 KF156001
O.mirabilis/O.constricta CBS 102086 Water (25 °C) Austria HQ667538 KF155956 KF156162
O.mirabilis/O.humicola CBS 102468 Human, nail Netherlands HQ667533 KF155947 KF156000
O.mirabilis/O.humicola CBS 113948 Human, finger nail Netherlands KF156081 HQ667530 KF155949 KF155997
O.mirabilis/O. sp. CBS 116659 ND Germany HQ667537
O.mirabilis/O. cf. humicola CBS 118685 Human, skin lesion Sweden HQ667529
O.mirabilis/O. cf. humicola CBS 121963, dH 18625 Bathroom Netherlands HQ667535
O.mirabilis/O.humicola CBS 123237, dH 16673 Human, finger nail Denmark KF156032
O.mirabilis/O.humicola CBS 123268 Human, toenail Denmark HQ667526
O.mirabilis/O.humicola CBS 124178 Bathroom Germany GQ426961
O.mirabilis/O.humicola CBS 124179 Bathroom Netherlands GQ426963
O.mirabilis/O.humicola CBS 124191 Bathroom Gemany GQ426978
O.mirabilis/O. cf. humicola CBS 124210, dH 17059 Human, foot Denmark KF156028
O.mirabilis/O.humicola dH 14815, KMBP H-14670 Fish, brain Germany KF156079 KF156036 KF156145 KF155954 KF156170 KF155998
O.mirabilis/O.humicola dH 16677 Human, toenail Denmark KF156031
O.mirabilis/O. sp. dH 22275 Human, toenail Thailand KF156077 KF156033 KF156141 KF155950 KF156169
O.mirabilis/S. sp. ATT130 Ant nest USA HQ607847
O.mirabilis/O. sp. DPMD19 Dead leaf Thailand KF156075 KF156030 KF156139 KF155955 KF156165
O.mirabilis/S. sp. F69 Sea-Fan Thailand EU714392
O.mirabilis/S. sp. GS2012 Banana, sooty blotch and flyspeck China JQ364738
O.mirabilis/S.dendroides HSAUP074025 ND China FJ914704
O.mirabilis/O. sp. JIM2 Balcony floor Thailand KF156078 KF156034 KF156143 KF155952 KF156166
O.mirabilis/S. sp. KH00281 Seagrass Thailand GU017502
O.mirabilis/S. sp. KH00283 Seagrass Thailand GU017504
Fungal Diversity

Table 1 (continued)
Current/obsolete name Culture/voucher Source Country GenBank accession number
SSU ITS LSU ACT1 BT2 TEF1
O.mirabilis/O. sp. KS4.1 Bathroom Thailand KF156080 KF156035 KF156142 KF155953 KF156167
O.mirabilis/S. sp. MY5-4 Sea sponge China JQ697530
O.mirabilis/S. sp. OUCMBIII 101045 Sea sand China HQ914903
O.mirabilis/ascomycete sp. s068 Rhizospheric soil Spain HQ650002
O.sexualis CBS 135765, PPRI 12991 (T) Domestic South Africa KF156089 KF156018 KF156118 KF155902 KF156189 KF155976
O.sexualis dH 22953 Ant Brazil KF156090 KF156017 KF156119 KF155903 KF156188 KF155977
O.tshawytschae/id. CBS 100438 (T) Fish USA KF156062 HQ667562 KF156126 KF155918 KF156180 KF155990
O.tshawytschae/id. CBS 130.65 Polder soil Netherlands KF156061 HQ667566 KF156127 KF155916 KF156178 KF155989
O.tshawytschae/id. CBS 228.66 Peat-bog soil Ireland KF156064 KF156016 KF156128 KF155915 KF156179 KF155992
O.tshawytschae/id. CBS 454.77 Agricultural soil Netherlands HQ667565 KF155913 KF156181 KF155991
O.tshawytschae/O.anellii CBS 850.73 Air Germany KF156063 KF156125 KF155914 KF156177 KF155988
O.tshawytschae/id. CBS 852.68 Polder soil Netherlands HQ667563 KF155917 KF155993
O.tshawytschae/id. CBS 129970 Human, subcutaneous mycosis China JN974456
O.tshawytschae/Scolecobasidium sp. E000535873 Grapevine soil and plant Spain JN578640
O.tshawytschae/S.tshawytschae HSAUP063055 ND China FJ914694
O.tshawytschae/S.variabile NBRC 32268 Soil China DQ307334
O.verrucosa/Septonema verrucosa CBS 383.81 (T) Soil India KF156067 KF156015 KF156129 KF155910 KF156185
O.verrucosa/O.tshawytschae CBS 225.77 Plant leaf Myanmar KF156066 HQ667564 KF156130 KF155909 KF156186 KF155985
Verruconis
V.calidifluminalis/O.calidifluminalis CBS 125818 (T) Hot sping river Japan KF156046 AB385698 KF156108 KF155901 KF156202 KF155959
V.calidifluminalis/O.calidifluminalis CBS 125817 Hot spring river Japan KF156045 AB385699 KF156107 KF155900 KF156201 KF155958
V.gallopava/O.gallopava CBS 437.64 (T) Turkey, brain USA KF156053 HQ667553 KF156112 HQ916989 KF156203 KF155968
V.gallopava/O.gallopava CBS 118.91 Human, lymphoma USA KF156047 HQ667551 KF156110 KF155932 HQ877643 JF440539
V.gallopava/O.gallopava CBS 166.85 ND France HQ667554 KF155923 KF156212 KF155960
V.gallopava/O.gallopava CBS 265.97 Chicken, brain Australia HQ667555 KF155931 KF156207
V.gallopava/O.gallopava CBS 547.81 ND New Zealand HQ667560 KF156109 KF155922 KF155973
V.gallopava/O.gallopava CBS 863.95 Human, HIV & transplantation USA KF156052 HQ667548 KF156114 KF155925 KF156208 KF155962
V.gallopava/O.gallopava CBS 864.95 Cat, brain USA KF156054 HQ667552 KF156116 KF155926 KF156209 KF155965
V.gallopava/O.gallopava CBS 865.95 Human, mine worker South Africa HQ667549
V.gallopava/O.gallopava CBS 867.95 Human, CVS disease USA KF156051 HQ667561 KF156117 KF155928 KF156213 KF155972
V.gallopava/O.gallopava CBS 119640 Human, disseminated infection Australia KF156049 HQ667558 KF156111 KF155961
V.gallopava/O.gallopava CBS 100437 Chicken, brain UK KF156050 HQ667556 KF156113 KF155924 KF156204 KF155966
V.gallopava/O.gallopava CBS 116646 Human, sputum Australia HQ667559 KF155930 KF156205 KF155963
V.gallopava/O.gallopava CBS 116660 Human, transplantation USA KF156048 HQ667557 KF156115 KF155929 KF156206 KF155969
Fungal Diversity

Table 1 (continued)
Current/obsolete name Culture/voucher Source Country GenBank accession number
SSU ITS LSU ACT1 BT2 TEF1
V.gallopava/O.gallopava CBS 119641 Human, AIDS UK HQ667547 KF155921 KF155970
V.gallopava/O.gallopava CBS 119642 Human, encephalitis ND HQ667550 KF155920 KF155967
V.gallopava/O.gallopava IFM 41473 Human, subcutaneous mycosis Japan AB125280
V.gallopava/O.gallopava IFM 52602 Human, autoimmune disease Japan AB125281
V.gallopava/O.gallopava IFM 54734 Hot spring bath Japan AB272162
V.gallopava/O.gallopava IFM 54735 Hot spring bath Japan AB272163
V.gallopava/O.gallopava IFM 54736 Hot spring river Japan AB272161
V.verruculosa/S.verruculosum CBS 119775 Plant root, Hevea sp. Malaysia KF156055 KF156014 KF156106 KF155919 KF156193 KF155974
Venturiales
Venturia inaequalis CBS 593.70 Plant Netherlands EU282480
V.inaequalis CBS 594.70 Plant Netherlands KF156093 KF156040 GU301879
V.inaequalis ATCC 60070 Plant USA EF114712
Veronaeopsis simplex CBS 588.66 Leaf litter South Africa KF156095 KF156041 KF156103
Sympoventuria capensis CBS 120136 Leaf litter South Africa KF156094 KF156039 KF156104
Fusicladium sicilianum CBS 105.85 T Rotten plant Italy FN549914 FN398150
S.excentricum CBS 469.95(T) Leaf litter Cuba KF156096 HQ667543 KF156105 KF155934 KF156196 KF155975
Pleosporomycetidae incertae sedis
S.arenarium CBS 181.58 Coastral sand France KF156099 KF156010 KF156157
S.salinum CBS 141.60 Timber in sea water Wales, UK KF156011
S.salinum CBS 142.60 Plant, grass UK KF156098 KF156158
S.salinum CBS 619.92 Seaweed USA KF156097 KF156159
S.salinum CBS 734.96 Cuba KF156100 KF156156
Chaetothyriales incertae sedis
S.fusarioideum CBS 210.95 Leaf litter Cuba KF156043 KF156154
S.cateniphorum CBS 769.83 Podocapus litter Japan KF156044 KF156013 KF156153
Helotiales incertae sedis
S.fusiforme CBS 586.82 Needle litter Netherlands KF156101 KF156012 KF156155 KF156160
Pezizomycotina incertae sedis
S.tropicum CBS 380.87 Indoor soil Canada KF156042 KF156102
id identical, ND no data
Fungal Diversity

In its opportunism to cold- and warm-blooded animals and
frequent association with wet environments, the group of
Ochroconis and Scolecobasidium species is quite exceptional
in the fungal Kingdom. Since the first isolation, the classifi-
cation of these fungi has been problematic, but a recent
phylogenetic analysis revealed that they belonged to the fam-
ily Sympoventuriaceae (Venturiales, Dothideomycetes)
(Machouart et al. 2013). However, the phylogenetic relation-
ship between the sister species have rarely been investigated
by molecular data. To clarify the taxonomy and ecology of
Ochroconis and Scolecobasidium species, especially at the
generic and species levels, sequences of the rDNA operon
and, including their previously physiochemical properties
were investigated. The taxa were delimited on the basis of
these datasets, and selected taxa compared on the basis of
morphology. Barcoding of these organisms with clinical and
veterinary significance was undertaken in view of improved
diagnostics.
Materials and methods
Strains and culture conditions
Ochroconis and Scolecobasidium strains (Table 1)werecul-
tured on oatmeal and malt extract agars (OA, MEA) at 24 °C
for 14 d. Microscopic preparations were performed by a slide
culture technique on MEA, corn meal agar (CMA) and potato
dextrose agar (PDA) using lactic acid and lactic acid-cotton
blue as mounting media. Micrographs were taken by a Nikon
Eclipse 80i microscope and DS Camera Head DS-Fi1/DS-
5 m/DS-2Mv/DS-2MBW using NIS-Element freeware pack-
age (Nikon Europe, Badhoevedorp, The Netherlands).
Molecular data
About 1 cm
2
fungal mass from agar culture, 7–14 d, was
collected and transferred to 2 ml Eppendorf tubes containing
300 μL TES buffer and Silica-mix (Merck 7736). DNA ex-
traction was performed according to Najafzadeh et al. (2009).
Six markers, viz. nuSSU, D1/D2 region of nuLSU, ITS, and
the gene fragments actin (ACT1), β-tubulin (BT2) and trans-
lation elongation factor 1-α(TEF1) were amplified by PCR
using primers as reported earlier (Feng et al. 2013). Final
volumes of PCR reactions were 15 μL containing 1 ng
DNA template, 10 pM of each primer, 4 μL of GoTaq®Green
Master Mix (Promega, Leiden, The Netherlands) and the
conditions for each gene were set. PCR was performed in a
GeneAmp 9700 (Applied Biosystems, Foster City, MA,
U.S.A.). For sequencing, 0.1–0.2 pmol of template was used
in Big Dye terminator cycle sequencing RR mix protocol
(Applied Biosystems), with PCR in 30 cycles of 96 °C 10 s,
50 °C 5 s and 60 °C 4 min. Sequences were edited with the
software SeqMan (Lasergene suite, DNASTAR,Madison,WI,
U.S.A.) and BioNumerics v. 4.61 (Applied Maths, Sint-
Martens-Latem, Belgium) was employed for first iterative
alignments. Sequences of nuLSU, nuSSU and ITS were
aligned with the program MUSCLE (www.ebi.ac.uk/Tools/
msa/muscle), and ACT1,BT2,andTEF1 with the program
MAFFT (http://www.ebi.ac.uk/Tools/msa/mafft), and all
aligned sequences were adjusted using BioEdit v. 7.0.5.2.
Phylogenetic analyses: single-locus datasets
Single-locus analyses were carried out for nuSSU, nuLSU, ITS,
ACT1,BT2,andTEF1 in order to investigate phylogenetic
relationships at generic and species levels. For nuLSU, nuSSU
and ITS, these analyses were done using Bayesian Inference
(BI) with MrBayes v. 3.1.2 on the Cipres portal (http://www.
phylo.org/sub_sections/portal). For nuLSU and nuSSU, two
parallel runs of four chains were run for 10,000,000 generations
and trees were sampled every 1,000 generations. For ITS, the
same analysis was run for 30,000,000 generations. TRACER v. 1.
5 was used to verify that the mean likelihood value, effective
sample size (ESS) and other parameters reached a plateau. For
each run, 10 % of the trees were discarded as burnin. Several
other methods were used to obtain additional support values.
Maximum likelihood (ML) and maximum parsimony (MP)
were carried out with MEGA5 (Tamura et al. 2011). ML analysis
with approximate likelihood ratio-test (aLRT) was also
performed with PhyML (Dereeper et al. 2008). Finally, stan-
dard bootstrapping was also performed with TNT using 1,000
replicates (Goloboff et al. 2008). For ACT1,BT2 and TEF1,
ML and MP were used to reconstruct the phylogenies in
MEGA5. The trees presented for these three protein-coding genes
are the ones obtained with ML. Trees were viewed and edited
with TreeView v. 1.6.6, FIGTREE v. 1.1.2 and MEGA5.
Phylogenetic analysis: multi-locus dataset
For 40 selected strains, sequence datasets of the six gene regions
nuSSU,ITS,nuLSU,ACT1,BT2 and TEF1 were concatenated
with the FASTA alignment joiner in FaBox (1.35). A partition-
homogeneity test (PHT) was carried out in PAUP v. 4.0b10 with
100 replicates. The strains CBS 469.95 and CBS 586.82 were
used as outgroup. The multi-locus analysis was performed using
the ML tool in MEGA5, using Tamura-Nei and GTR+I as the best
model and 1,000 bootstrap replicates. A MP analysis was also
carried out in MEGA5 using 1,000 bootstrap replicates. For the
Bayesian approach with MrBayes, two parallel runs of
20,000,000 generations were done with a sampling frequency
of 1,000 trees. A burnin tree sample of 10 % was discarded.
Finally, a ML analyses was carried out with PHYML.Thetree
presented was obtained with ML in MEGA5. Tree reconstruction,
visualization and editing were done as described above.
Fungal Diversity

Results
Phylogeny at generic and species levels based on nuLSU
and nuSSU (Figs. 1and 2)
The nuLSU dataset included 60 taxa among which are species of
Ochroconis,Scolecobasidium and related members of
Venturiales. This dataset contained 825 characters of which 254
were parsimony-informative. The resulting Bayesian tree is
presented here with posterior probabilities (PP) and support
values obtained with other phylogenetic methods. Our results
show that several Scolecobasidium species are in fact unrelated
to Ochroconis (Fig. 1). All Ochroconis species belongs to two
main lineages, A and B. Lineage A comprised a group with O.
calidifluminalis,O.gallopava and S.verruculosum,whileline-
age B contained the remaining species of Ochroconis.Most
species represented by several strains formed well-supported
monophyletic groups with PPs of 1.0 and bootstrap values higher
than 70 %. The nuSSU dataset included 60 taxa and 1,482
characters, among which 227 were parsimony-informative. As
for the nuLSU analysis, two main lineages of Ochroconis were
recovered and most Scolecobasidium species clustered with the
outgroup (Fig. 2). Most Ochroconis species also form well-
supported monophyletic groups.
Phylogeny at generic and species levels based on ITS (Fig. 3)
For the ITS dataset, a total of 117 strains listed under the generic
names Ochroconis and Scolecobasidium,aswellassomepre-
sumably closely related species were analyzed and compared
with sequences available in GenBank using BLAST.Extreme
differences were noted between groups of strains, both in length
and in nucleotide composition (Table 2). Confident alignment of
ITS sequences was difficult over the entire dataset, even when
apparently unrelated strains were discarded. When unrelated
strains were excluded, the alignment contained 1,073 bp-ITS
sequences with 564 parsimony-informative sites. The overall
mean substitution rate of Ochroconis and Scolecobasidium
strains belonging to the complex was 0.143 substitutions per site.
Barcoding gaps between species were significant, with a mini-
mum of 7.9 % ITS difference between Ochroconis
calidifluminalis and O.gallopava. Heterogeneity within clusters
that were preliminarily judged to represent individual species
ranged between 0 and 22.1 %.
Although the ITS dataset was difficult to align, the informa-
tion in the ITS dataset revealed a tree with a similar topology as
those of nuLSU and nuSSU. This ITS analysis resulted in 11
species (Fig. 3). Among these, nine species were supported with
high support values for the three methods used (PP/aLRT/%
TNT-bootstrap). These species included O.mirabilis nob., O.
constricta,O.tshawytschae,O.verrucosa nob., O.lascauxensis,
O.sexualis nob., O.cordanae nob., and O.gallopava.Two
species did not recover support from all methods: O.gamsii
(0.99 PP/−aLRT/51 % bootstrap), and O.calidifluminalis (1.0
PP/0.60 aLRT/100 % bootstrap). The phylogenetic positions of
S.arenarium (CBS 181.58), S.cateniphorum (CBS 769.83), S.
excentricum (CBS 469.95), S.fusiforme (CBS 586.82), and S.
salinum (CBS 141.60), were in accordance with nuSSU and
nuLSU results, and considered to be only distantly related to
Ochroconis.
Phylogeny at generic and species levels based on ACT1,
BT2, and TEF1 (Fig. 4)
The datasets of three structural genes, ACT1,BT2,andTEF1
showed very high average distances between sequences:
ACT1=0.285 substitutions/site, BT2= 0.158 substitutions/site, and
TEF1=0.376 substitutions/site, which in some cases severely ham-
pered reliable alignment in view of tree reconstruction. For this
reason, each species-cluster of relatively closely related groups were
aligned separately and analyzed with ML and MP (Fig. 4). Species
for which high ML and MP support values were recovered for the
three genes ACT1,BT2,andTEF1 consisted of O.calidifluminalis,
O.cordanae nob., O.gallopava,O.minima nob., O.mirabilis nob.,
O.tshawytschae and O.verrucosa nob. The remaining species were
only partially supported: a clade containing O.gamsii was support-
ed only by TEF1 (98 % ML and/100 % MP bootstrap), while O.
constricta was strongly supported by ACT1 and TEF1 (100 % ML
and MP bootstrap for both genes), but strains did not cluster in the
analysis with BT2. The type strain, CBS 202.27 clustered with the
remaining species in ribosomal gene trees but not with coding gene
fragments.
Phylogeny at generic and species levels based on a 6-gene
analysis (Fig. 5)
A multi-locus tree was reconstructed using a set of six gene
regions (nuSSU, ITS, nuLSU, ACT1,BT2 and TEF1)from40
strains. The length of concatenated sequences was 4,906 bp and
contained 1,571 parsimony informative sites. Eight species
obtained the highest support values for the four methods
(ML/MP/PP/aLRT): O.cordanae nob., V.calidifluminalis nob.,
V.gallopava nob., O.minima nob., O.mirabilis nob., O.
tshawytschae,O.sexualis nob., and O.verrucosa nob.
Ochroconis constricta (94/93/ /0.99) and O.gamsii
(93/86/1.0/1.0) obtained lower values, although still supported
by most of the methods used.
Taxonomy
Based on molecular phylogenetic analyses, and morphological
and ecological comparison, two genera are recognized,
Verruconis and Ochroconis. The original genus Scolecobasidium
is considered of doubtful identity because the type material is
ambiguous (see below).
Fungal Diversity

Fig. 1 Bayesian tree for a dataset including 60 taxa and the marker
nuLSU. Numbers on the branches are Bayesian posterior probabilities
(PP), bootstrap values obtained with maximum likelihood (ML) in MEGA5
and bootstrap values obtained with maximum parsimony (MP) in MEGA5
(PP/ML/MP). A red dot on a branch indicates a support higher than 0.95
for all methods. Type strains are highlighted by a T
Fungal Diversity

Key to the genera
1a. Conidia subhyaline, never light to dark brown, smooth-
walled to verruculose; mesophilic ....………… Ochroconis
1b. Conidia light to dark brown, verrucose to coarsely
ornamented; thermophilic ………........…… Verruconis
Ochroconis de Hoog & Arx. MB9136.
Colonies restricted, brown or olivaceous. Aerial hyphae
smooth- or somewhat rough-walled, pigmented. Cleistothecia
up to 40 μm in diam, dark brown; peridium wall composed of
textura angularis. Ascomata bearing antler-shaped appendages,
with serrate edges. Asci bitunicate, clavate, 8-spored; ascospores
Fig. 2 Bayesian tree for a dataset
including 60 taxa and the marker
nuSSU. Numbers on the branches
are Bayesian posterior
probabilities (PP), bootstrap values
obtained with maximum
likelihood (ML) in MEGA5 and
bootstrap values obtained with
maximum parsimony (MP) in
MEGA5 (PP/ML/MP). A red dot on
a branch indicates a support higher
than 0.95 for all methods. Type
strains are highlighted by a T
Fungal Diversity

Fig. 3 Bayesian tree for a dataset including 117 strains and the marker
ITS. Numbers on the branches are bayesian posterior probabilities (PP),
PhyML-approximate likelihood ratio-test (aLRT) and bootstrap values
from TNT-maximum parsimony analysis (PP/aLRT/TNT). Stripes
represent strain distribution; sources, geography and climate distribu-
tion. A black dot on a branch indicates a support higher than 0.95 PP.
Type strains are highlighted by a T
Fungal Diversity

Table 2 ITS data of analyzed Ochroconis and Verruconis species
Current name Culture/voucher ITS length (bp) ITS G+ C % Mean distance
O.anelli CBS 284.64 (T) 649 51.16
O.anomala CBS 131816 (T) 754 58.49
O.constricta CBS 211.53 (T) 678 49.71 0.131
O.constricta CBS 106.65 629 52.94
O.constricta CBS 202.27 (T) 641 52.11
O.constricta CBS 269.61 650 50.15
O.constricta CBS 381.64 626 53.04
O.constricta CBS 131913 641 52.11
O.constricta IFM 52654 640 52.19
O.cordanae CBS 475.80 (T) 566 56.54 0.005
O.cordanae CBS 172.74 565 56.46
O.cordanae CBS 412.51 566 56.54
O.cordanae CBS 780.83 565 56.46
O.cordanae CBS 123536 566 56.18
O.cordanae 9165 565 56.28
O.cordanae TR072 565 56.28
O.gamsii CBS 239.78 (T) 678 53.54 0.221
O. aff. gamsii CBS 101179 668 51.20
O. aff. gamsii CBS 102491 668 51.80
O.humicola CBS 116655 (T) 674 54.90
O.lascauxensis CBS 131815 (T) 584 59.93 0.009
O.lascauxensis LX7.2 584 60.10
O.lascauxensis LX101R-6 583 60.55
O.lascauxensis LXA3 584 59.76
O.lascauxensis LX CH2 585 59.15
O.lascauxensis LX CH3 585 59.32
O.lascauxensis LX CH7 585 59.83
O.lascauxensis LXM8-5 584 59.93
O.longiphorum CBS 435.76 624 57.05
O.minima CBS 510.71 (T) 580 59.31 0.098
O.minima CBS 423.64 591 58.71
O.minima CBS 536.69 592 58.61
O.minima CBS 116645 592 58.61
O.minima CBS 119792 580 59.31
O.mirabilis CBS 729.95 (T) 636 54.40 0.042
O.mirabilis CBS 124.65 645 53.95
O.mirabilis CBS 413.51 644 54.81
O.mirabilis CBS 100486 642 54.05
O.mirabilis CBS 102086 633 55.13
O.mirabilis CBS 102468 642 54.05
O.mirabilis CBS 113948 646 54.02
O.mirabilis CBS 118685 643 54.28
O.mirabilis CBS 121963 643 54.43
O.mirabilis CBS 124210 632 55.22
O.mirabilis dH 22275 645 53.95
O.mirabilis ATT130 638 53.45
O.mirabilis DPMD19 636 55.03
O.mirabilis F69 646 54.18
O.mirabilis GS2012 641 54.87
Fungal Diversity

pale brown, verruculose, 1–3-septate. Conidiogenous cells
scattered, arising from undifferentiated hyphae or integrated in
short lateral branches, elongate to cylindrical; conidium-bearing
denticles cylindrical, scattered alongside the conidiogenous
cell, grouped in the apical region, or in distinct rachis. Conidia
1–4-celled, pigmented, verrucose, ellipsoidal, cylindrical, T- or
Y-shaped, or bilobate, usually provided with a distinct hilum
due to rhexolytic liberation. Brown, one-to many-celled, lateral
or intercalary chlamydospores may be present.
Generic type:Scolecobasidium constrictum E.V. Abbott
Ochroconis anellii (Graniti) de Hoog & Arx, Kavaka 1:
57, 1973. MB318845, Fig. 6.
Table 2 (continued)
Current name Culture/voucher ITS length (bp) ITS G+ C % Mean distance
O.mirabilis HSAUP074025 636 54.04
O.mirabilis JIM2 644 54.04
O.mirabilis KMBP H-14670 645 54.11
O.mirabilis KS4.1 644 54.04
O.mirabilis MY5-4 644 54.04
O.mirabilis OUCMBIII 101045 644 54.04
O.mirabilis s068 638 53.45
O.sexualis PPRI 12991 (T) 592 48.82 0
O.sexualis dH 22953 592 48.82
O.tshawytschae CBS 100438 (T) 707 60.96 0.016
O.tshawytschae CBS 130.65 705 60.99
O.tshawytschae CBS 228.66 705 60.99
O.tshawytschae CBS 454.77 706 60.20
O.tshawytschae CBS 852.68 713 60.87
O.tshawytschae CBS 129970 706 60.76
O.tshawytschae E000535873 706 60.91
O.tshawytschae NBRC 32268 706 60.91
O.tshawytschae HSAUP063055 707 60.82
O.verrucosa CBS 383.81 (T) 699 55.22 0.115
O.veerucosa CBS 225.77 698 56.99
V.calidifluminalis CBS 125818 (T) 668 51.80 0.002
V.calidifluminalis CBS 125817 669 51.87
V.gallopava CBS 437.64 (T) 672 52.23 0.014
V.gallopava CBS 118.91 670 52.09
V.gallopava CBS 166.85 669 52.32
V.gallopava CBS 265.97 669 52.32
V.gallopava CBS 547.81 669 52.32
V.gallopava CBS 863.95 667 51.87
V.gallopava CBS 864.95 670 52.09
V.gallopava CBS 865.95 670 52.39
V.gallopava CBS 867.95 670 52.24
V.gallopava CBS 100437 669 52.32
V.gallopava CBS 116646 670 52.39
V.gallopava CBS 116660 669 52.02
V.gallopava CBS 119641 663 51.58
V.gallopava CBS 119642 671 52.01
V.gallopava IFM 41473 669 52.32
V.gallopava IFM 52602 669 52.32
V.gallopava IFM 54734 669 52.32
V.gallopava IFM 54735 669 52.32
V.gallopava IFM 54736 669 52.32
V.verruculosa CBS 119775 597 60.47
Fungal Diversity

≡Scolecobasidium anellii Graniti, Nuovo G. Bot. Ital. 69:
364, 1963.
Description based on CBS 284.64 at 25 °C on OA after
4 weeks in darkness (Ellis 1971).
Colonies effuse, somewhat raised, cottony or velvety,
isabelline to olivaceous. Hyphae subhyaline to rather pale
olive, smooth-walled, occasionally minutely verruculose. Co-
nidiophores straight or flexuose, sometimes clavate, smooth-
walled, pale to mid olive. Conidia straight, oblong, rounded at
the ends or sometimes ellipsoidal, pale to mid olivaceous-
brown, verruculose or minutely echinulate, with 2–5(mostly
3) transverse septa and occasionally a longitudinal septum,
when mature often slightly constricted at the septa.
Type: Italy, Bari, Grotto di Castellani, from stalactite, A.
Graniti, April 1964. Type strain: CBS 284.64= IHEM
4516=IMI 089069=LCP 71.2121 = MUCL 9473.
Fig. 4 MEGA5 maximum likelihood trees, from data set of each species and three genes (ACT1,BT2,TEF1). Numbers on branches are bootstrap
values obtained from maximum likelihood and maximum parsimony analyses. Type strains are highlighted by a T. x=not cluster-member
Fungal Diversity

Fig. 5 MEGA5-maximum likelihood tree from a concatenated dataset
including the 6 gene regions nuSSU, ITS, nuLSU, ACT1,BT2 and
TEF1. Numbers on the branches are bootstrap values for MEGA5-
maximum likelihood (ML), MEGA5-maximum parsimony (MP),
Bayesian posterior probabilities (PP) and PhyML-approximate likeli-
hood ratio-test (aLRT); (ML/MP/PP/aLRT). A red dot on a branch
indicates a support higher than 0.95 for all methods. Type strains are
highlighted by a T
Fungal Diversity

Fig. 6 Ochroconis anellii, CBS 284.64. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–J. Conidial apparatus with rhexolytic conidia
produced from simple conidiophores. K, L. Hyphal coils. M. Anastomosing hyphae. Scale bar = 10 μm
Fungal Diversity

Note: The first strain, CBS 284.64 was isolated from a
cave in Italy. A second strain with morphological features of
S.anellii, CBS 850.73 was recovered from air in Germany
(Table 1). However, after phylogenetic analyses (Figs. 1,2,4)
this strain appeared to be very different and was re-identified as
O.tshawytschae.HenceO.anellii contains only a single strain.
Its nearest neighbor, Ochroconis anomala, had very similar
ecology, originating from a cave in France. Given the wide
intraspecific heterogeneity in the Ochroconis lineage the pos-
sibility remains that a single species is concerned. More mate-
rial is needed to reach a sound conclusion. Ochroconis
lascauxensis from the same cave in France is widely different.
Ochroconis anomala A. Nováková & P.M. Martin-
Sanchez, Fungal Biol. 116: 584, 2012. MB561939.
Type: France, Montignac, Lascaux Cave, on the cave
sediment in the Painted Gallery, F. Bastian, Aug. 2007. Type
strain: CBS 131816.
Note: A full description of this species was given by
Nováková and Martin-Sanchez (2012). The species is charac-
terized by erect conidiophores, producing two types of
conidiogenous cells which lead to conidia in short chains or
on denticles. The conidia are 2–5-celled. Ochroconis anomala
is a sister species of O.anellii.
Ochroconis constricta (E.V. Abbott) de Hoog & Arx,
Kavaka 1: 57, 1973. MB318847, Fig. 7.
≡Scolecobasidium constrictum E.V. Abbott, Mycologia
19: 30, 1927.
≡Dactylaria constricta (E.V. Abbott) D.M. Dixon &
Salkin, J. Clin. Microbiol. 24: 13, 1986.
=Heterosporium terrestre R.G. Atk., Mycologia 44: 813,
1952.
Description based on CBS 211.53 at 25 °C after 4 weeks
in darkness.
On OA, colonies moderately expanding, smooth, dry, flat,
greyish brown to dark brown. On MEA, colonies flat, felty,
with some shallow radial fissures. Hyphae brown, with rather
thick walls. Conidiophores short, mostly flask-shaped to cylin-
drical, sometimes barely differentiated, bearing 1–3 conidia
near the apex. Denticles fragile, scattered. Conidia two-celled,
verrucose, subhyaline to pale brown, verrucose, broadly ellip-
soidal to cylindrical, slightly narrower around the middle.
Conidial secession rhexolytic, frills remaining on denticle and
on conidial base. Cardinal temperatures: growth abilities rang-
ing from 15 to 30 °C, optimal growth at 25–30 °C. Growth with
5%MgCl
2
and with 5 % NaCl (Horré et al. 1999).
Type:U.S.A., Louisiana, from soil, E.V. Abbott 1927.
Type strain: CBS 202.27= MUCL 9471.
Notes: This is one of the commonly mentioned species in
the literature, but most of these reports may have concerned
other ochroconis-like species; only very few strains were
available for molecular verification. The type strain, CBS
202.27 showed limited sporulation with irregularly verru-
cose, rather broadly cylindrical to somewhat dumbbell-
shaped conidia. Considerable intraspecific variability is not-
ed in all markers (Figs. 1,2,3,4, and 5); the species may
indicate hidden diversity, but the number of isolates available
was too small for a genealogical concordance study.
Ochroconis cordanae Samerpitak, Crous & de Hoog, sp.
nov. MB519170, Fig. 8, see also Crous et al. (2008).
Description based on CBS 475.80 at 25 °C after 4 weeks
in darkness.
On OA, colonies attaining about 4 cm diam, flat, with
predominantly submerged mycelium, dark olivaceous-
brown. Hyphae branched, with thin septa, hyaline to pale
brown, smooth-walled, 2.0–2.5 μm wide. Conidiophores
clearly differentiated, arising at right angles from creeping
hyphae, unbranched, with 1–3 very thin septa, straight to
flexuous, brown, thick-walled, 10–50× 2.5–3.5 μm, produc-
ing conidium-bearing denticles that are widely spaced in the
apical region. Denticles acuminate, up to 0.5 μm long,
rhexolytic with narrow unpigmented scars. Conidia solitary,
subhyaline to hazel brown, finely verruculose, thin-walled,
with a thin median septum, obovoidal to broadly fusiform,
(5−)7–9(−10)×(2.5−)3.0–3.5 μm. Teleomorph unknown.
Type:Colombia, Villavicencio, from dead leaf, W.
Gams, Dec. 1979. Type strain: CBS 475.80.
Additional strains examined: see Table 1.
Habitat: The species is repeatedly found inhabiting living
leaves, sometimes as an endophyte, and is also found on
plant litter, worldwide. Two strains originated from ant nests
in subtropical Brazil.
Notes: The species is morphologically similar to
Scolecobasidium crassihumicola Matsush., which has somewhat
larger conidia and is currently not available for sequencing. The
species seems to be fairly common but has frequently been
misidentified. Four strains isolated from living plants,
EF419939 (= isolate 9165, an unnamed fungal endophyte of
Platycladus orientalis), CBS 123536 (= FJ372390, as
Ochroconis sp. from living leaves of Encephalartos lebom-
boensis), and FJ752621 (= JIA 6-8-1), an unnamed endophyte
of wild rice roots (Oryza granulata) and JQ760421 (= isolate
FLO760, an unnamed endophyte), are corrected to be this new
species. Two strains from an ant nest, HQ608103 (= TR072 as S.
humicola) and HQ607921 (= ATT286, as Scopulariopsis
humicola) are also identified as O.cordanae (Fig. 3).
Ochroconis gamsii de Hoog, Stud. Mycol. 26: 52, 1985.
MB104571, Fig. 9.
Type: Sri Lanka, Peradeniya, on leaves of Caryota
plumosa, W. Gams, Jan. 1978. Type strain: CBS 239.78.
Note: A full description of this species was given by de
Hoog (1985). It is characterized by large conidiophores and
characteristic, unilaterally flattened conidia, and took an
isolated position within the genus Ochroconis (Fig. 5). At
present the species contains only a single confirmed strain.
Ochroconis humicola (G.L Barron & L.V. Busch) de
Hoog & Arx, Kavaka 1: 57, 1973. MB318849, Fig. 10.
Fungal Diversity

Fig. 7 Ochroconis constricta, CBS 211.53. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–N. Conidial apparatus with rhexolytic conidia
produced from short simple conidiophores. O, P. Hyphal coils. Scale bar= 10 μm
Fungal Diversity

Fig. 8 Ochroconis cordanae, CBS 475.80. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. Colony on C–I. Conidial apparatus with rhexolytic
conidia produced from simple conidiophores. J. Hyphal coils and anastomosing hyphae. Scale bar = 10 μm
Fungal Diversity

Fig. 9 Ochroconis gamsii, CBS 239.78. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–F. Conidial apparatus with rhexolytic conidia
produced from long, brown conidiophores. G, H. Anastomosing hyphae. I, J. Hyphal coils. K, L. 4-celled conidia. Scale bar=10 μm
Fungal Diversity

Fig. 10 Ochroconis humicola, CBS 116655. A. Colony on OA, 4 wk. B. Colony on MEA, 4 wk. C–L. Conidial apparatus with cylindricall, 2-celled,
rhexolytic conidia produced from long-cylindrical conidiophores. M,N. Chlamydospores. O. Hyphal coil. Scale bar=10 μm
Fungal Diversity

≡Scolecobasidium humicola G.L. Barron & L.V. Busch,
Can. J. Bot. 40: 83, 1962.
Description based on CBS 116655 at 25 °C after 4 weeks
in darkness.
On OA, colonies moderately expanding, smooth, dry, flat,
olivaceous black. On MEA, colonies flat, felty, with some
shallow radial fissures. Hyphae brown, with rather thick walls.
Conidiophores short, mostly flask-shaped to short-cylindrical,
sometimes barely differentiated, bearing 1–3 conidia near the
apex. Denticles fragile, scattered. Conidia two-celled, verru-
cose, subhyaline to pale brown, broadly ellipsoidal to cylin-
drical, slightly narrower around the middle. Conidial seces-
sion rhexolytic, frills remaining on denticle and on conidial
base. Cardinal temperatures: growth abilities ranging from 15
to 30 °C, optimal growth at 25–30 °C. Growth with 10 %
MgCl
2
and with 5 % NaCl (Horré et al. 1999).
Type: Canada, Guelph, from peat soil, G.L. Barron. Type
strain: CBS 116655= IMI 110131 = UAMH 10241.
Note: The name Ochroconis humicola appears commonly
in the literature, but presently only a single confirmed strain is
available, which is located at an isolated position in the all
analysed trees (Figs. 1,2,3,4,and5). The species has a relatively
robust morphology, with erect, dark brown conidiophores.
Ochroconis lascauxensis A. Nováková & P.M. Martin-
Sanchez, Fungal Biol. 116: 580, 2012. MB561938.
Type: France, Montignac, Lascaux Cave, on a visible
black stain on the rock in the Passageway, F. Bastian, Aug.
2008. Type strain: CBS 131815.
Additional strains examined: see Table 1.
Note: A full description of this species was given by
Nováková & Martin-Sanchez (2012). It is characterized by
erect conidiophores which are cylindrical or flexuose and
have no or few septa, while the conidia are 2–3-septate. The
species is a sister taxon of O.tshawytschae (Fig. 5), and is
phylogenetically remote from the other cave-species above,
O.anellii and O.anomala.
Ochroconis longiphorum (Matsush.) Samerpitak & de
Hoog, comb. nov. MB804902, Fig. 11.
≡Scolecobasidium longiphorum Matsush., Icones
Microfungorum a Matsushima lectorum, Kobe, p. 127, 1975.
Description based on CBS 435.76 at 25 °C after 4 weeks
in darkness (see also Matsushima 1975).
Colonies effuse, minutely hairy, brown. Mycelium mainly
immersed; hyphae branched, septate, subhyaline or brown,
thick-walled, 1.0–2.5 μm. Conidiophores arising from re-
pent hyphae, brown, solitary, erect, simple, septate, straight
or curved, 50–600 μm long, thick-walled, elongating
sympodially, with spaced conidium-bearing denticles.
Conidia cylindrical, rounded at both ends, 3-septate, not or
slightly constricted at the septa, verruculose, 12–19 ×3.5–
4.5 μm, crowded, pale brown.
Type: Japan, Kyoto City, from insect dung on dead oak
leaf, T. Matsushima. Type strain: MFC-2451.
Note: The original isolate MFC-2451 was not available
for study. CBS 435.76= UAMH 3972 was isolated from soil
under asphalt paving of a car park in Alberta, Canada and
identified by L. Sigler. This strain was published by Davies
and Westlake (1979)asS.longiphorum, but given the very
different source of isolation identity of the strains remains
uncertain.
Ochroconis minima (Fassat.) Samerpitak & de Hoog,
comb. nov. MB519046, Fig. 12.
≡Humicola minima Fassat., Česká Mykol. 21: 87, 1967
(basionym).
Description based on CBS 510.71 at 25 °C after 4 weeks
in darkness.
On OA, colonies attaining about 3 cm diam, felty, pale
greyish brown. On MEA, colonies attaining max 2 cm diam,
convex, greyish brown; a dark olivaceous brown pigment is
exuded into the agar. Hyphae brown, 1.3–1.7 μm wide.
Conidiogenous cells standing at right angles from undifferen-
tiated hyphae, flask-shaped to clavate, 7.5–12.0 μm long, with
some scattered denticles in the apical region. Conidia smooth-
walled, with a median septum, clavate to somewhat T- or Y-
shaped, 9.0–13.5 μm long, lower cell about 4.5 μm wide,
upper cell up to 8.0 μm wide. Spherical, brown, smooth- and
thick-walled chlamydospores 4–5μm in diam are inserted
alongside the hyphae.
Type: Nigeria; Samaru, Zaria, from rhizosphere of
Gossypium arboretum, M. Dransfield. Type strain: CBS
510.71=ATCC 22631 = IMI 082933.
Additional strains examined: see Table 1.
Note: Because of the somewhat Y- or T-shaped conidia the
species had longtime been listed as a synonym of
Scolecobasidium terreum, but is phylogenetically remote from
several strains with terreum-like morphology in all markers
analyzed. Although O.minima consists of two monophyletic
clades, the similarity of all strains in conidial morphology and
ecology provides a strong support to classify them in a single
species, albeit with high intraspecific heterogeneity as observed
in most members of the Ochroconis lineage. Judging from ITS
data, three GenBank submissions cluster with O.minima
(Fig. 3,Table1). Also DQ307328 (= NBRC 9845, previously
identified as S.terreum) clustered with CBS 119792 as a sister
strain of the type, and is possibly re-identified as O.minima.
Ochroconis mirabilis Samerpitak & de Hoog, sp. nov.
MB519045, Fig. 13.
Description based on CBS 729.95 at 25 °C after 4 weeks
in darkness.
On OA, colonies attaining 5 cm diam, smooth to felty, dry,
flat, greyish brown to dark brown. On MEA, colonies
attaining 4.8 cm diam, smooth, velvety. Hyphae brown, with
rather thick walls. Conidiophores flexible, mostly cylindrical
to acicular, sometimes barely differentiated, 0(−1)-septate,
8–15 μm long, bearing a few conidia near the apex. Denti-
cles fragile, scattered. Conidia two-celled, smooth-walled to
Fungal Diversity

Fig. 11 Ochroconis longiphorum, CBS 435.76. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–H. Conidial apparatus with rhexolytic 4-celled
conidia produced from conidiophores. I. Anastomosing hyphae. J. Hyphal coil. Scale bar = 10 μm
Fungal Diversity

Fig. 12 Ochroconis minima, CBS 510.71. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–G. Conidial apparatus with rhexolytic conidia
produced from brown, stout conidiophores. H, I. Conidia and anastomosing hyphae. J. Chlamydospores and anastomosing hyphae. Scale bar= 10 μm
Fungal Diversity

Fig. 13 Ochroconis mirabilis, CBS 729.95. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–K. Rhexolytic conidia narrowed at the septum,
produced from conical to cylindrical conidiophores. L, M. Anastomosing hyphae. Scale bar = 10 μm
Fungal Diversity

verruculose, subhyaline to pale brown, cylindrical, constricted
at the septum, 9.0–13.5×4.8–6.7 μm. Conidial secession
rhexolytic, frills remaining on denticle and on conidial base.
Cardinal temperatures: growth abilities ranging from 15 to
30 °C, optimal growth at 25–30 °C. Growth in 10 % MgCl
2
and with 5 % NaCl (Horré et al. 1999).
Type: The Netherlands, Haarlem, Streeklaboratorium
voor Volksgezondheid, from regulator of diver, Nov. 1995,
CBS-H 20527. Type strain: CBS 729.95.
Additional strains examined: see Table 1.
Habitat: waterborne, opportunist in fish; also indoor wet
cells such as bathrooms and causing mild cutaneous infections
in humans; occasionally isolated from soil and plant material.
Notes: Judging from the number of strains available, this is
one of the relatively common species of the Scolecobasidium/
Ochroconis complex. It is a waterborne species which is found in
relatively warm environments, and can commonly be isolated
from moist localizations in bathrooms, e.g. bath edges, sinks or
shower cabins. Lian and de Hoog (2010) suggested that
bathroom-associated fungi, among which are many black yeast
species, are picked up by human skin and in nails when the skin
barrier is weakened during bathing. Strain CBS 729.95 originat-
ed from a regulator of a diver after a near-drowning accident, the
same species having been isolated from the diver’s pulmonary
aspirate. Also waterborne cold-blooded animals are prone to
infection, as described by Schaumann and Priebe (1994)where
the fungus caused muscular black spot disease in Atlantic
salmon.
The ITS region contains about 641 bp, and has 54 % mole
G+ C. A cluster of strains was found to be consistently differ-
ent in all genes analyzed, at a mean ITS distance of 4.2 %
(Table 2). This group contained a GenBank deposition
FJ914704 under the name Scolecobasidium dendroides Piroz.
& Hodges (Pirozynski and Hodges 1973). This species was
originally described as a hyperparasite on a Circinotrichum
species and has not been cultured. FJ914704 came from putrid
plant material and is thus unlikely to represent the same
species. The ‘dendroides’-cluster shows the same combina-
tion of habitats as the main cluster, i.e., comprising isolates
from bathrooms and from human skin, and for this reason we
provisionally leave the group within O.mirabilis. Other
GenBank depositions, which all originate from or are
otherwise associated with sea water, such as HQ914903
(= OUCMBIII 101045, as Scolecobasidium sp.) from sea
sand, EU714392 (= F69 as Scolecobasidium sp.) from a sea-
fan, JQ697530 (= MY54 as Scolecobasidium sp.) from a sea
sponge, and two Scolecobasidium sp. from seagrass,
GU017502 and GU017504 (= KH00281, KH00283 respec-
tively), are re-identified here as O.mirabilis.However,some
GenBank accessions matching with O.mirabilis were isolated
from soil (HQ650002=s068), from a lesion of a diseased plant
(JQ364738=GS2012), and DPMD19 from a dead leaf, were
also proven to belong to O.mirabilis.
Ochroconis sexualis Samerpitak, Van der Linde & de
Hoog, sp. nov. MB803666, Fig. 14.
Description based on PPRI 12991at 25 °C after 4 weeks in
darkness.
On OA, colonies growing slowly, velvety, dark grey to
olivaceous-brown; reverse dark grey to almost black. Myce-
lium pale brown, smooth- and thick-walled. Conidiophores
with conidia produced on denticles, with rhexolytic conidium
secession, pale brown, verrucose, cylindrical with rounded
ends, 1–3-septate, 15–22 μm× 3.5–5.0 μm. Cleistothecia dark
brown, spherical, up to 40 μm diam astomate; peridium wall
composed of textura angularis. Appendages golden brown,
varying in number and branching, with serrate edges. Asci
bitunicate, clavate, 8-spored, 15–20×7–9μm. Ascospores
pale brown, verruculose, 1- or 3-septate when mature, 8–
10× 2.5 3.5 μm.
Type: South Africa, Durban, obtained from quality con-
trol swabs in a laboratory providing medical supplies, pre-
served in National Collections of Fungi, South Africa (PPRI).
Type strain=PPRI 12991; PREM 60690.
Notes. The teleomorph found in a single strain, PPRI
12991, is the first ascosporulating strain in the Ochroconis
lineage. The ascomata bear characteristic antler-shaped ap-
pendages, which are unique in the Venturiales. The strain
produced conidia similar to those of O.tshawytschae in low
abundance. A second strain of O.sexualis, dH 22953, iden-
tified as such by identical multilocus sequence data, was
completely sterile, the culture merely consisting of hyphae.
Ochroconis tshawytschae (Doty & D.W. Slater)
Kirilenko & Al-Achmed, Mykrobiol. Zhurn. 39: 305, 1977.
MB318851, Fig. 15.
≡Heterosporium tshawytschae Doty & D.W. Slater, Am.
Midl. Nat. 36: 663, 1946.
≡Scolecobasidium tshawytschae (Doty & D.W. Slater)
McGinnis & Ajello, Trans. Br. Mycol. Soc. 63: 202, 1974.
Description based on CBS 100438 at 25 °C after 4 weeks
in darkness.
On OA, colonies attaining 4.3 cm diam, with flat, felty,
brown aerial mycelium and dark brown submerged myceli-
um. On MEA, colonies attaining 3.5 cm diam, woolly,
domed, with dense dark hairs at the centre and grey, loose
mycelium near the edge. Hyphae subhyaline to pale oliva-
ceous brown, smooth- and thick-walled, forming a compact
mycelium. Conidiophores erect, straight, cylindrical, often
somewhat inflated or flexuose, up to 30 μm long, 2.0–
3.5 μm wide; conidia produced on denticles in the apical
region. Conidia (2−)4-celled, verrucose, pale brown, cylin-
drical or slightly clavate, with round ends, 12–25 ×3.5–
5.5 μm. Cardinal temperatures: growth abilities ranging
from 15 to 30 °C, with optimal growth at 25–30 °C. Growth
with 5 % MgCl
2
and with 5 % NaCl (Horré et al. 1999).
Type: U.S.A., California, from infected kidney of young
Oncorhynchus tshawytscha (chinook salmon), Anderson.
Fungal Diversity

Fig. 14 Ochroconis sexualis, CBS 135765. A. Colony on MEA 2 wk.
B. Colony on OA, 2 wk. C, I–P. Rhexolytic conidia narrowed at the
septum, produced from denticles to cylindrical conidiophores. D, E.
Cleistothecial ascomata. F, G. Asci and ascospors. H, Q. Anastomosing
hyphae. Scale bar= 10 μm
Fungal Diversity

Fig. 15 Ochroconis tshawytschae, CBS 100438. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–L. Conidial apparatus with rhexolytic, 4-
celled conidia, produced from short to long cylindrical conidiophores. M–O. Chlamydospores. P. Hyphal coil. Scale bar= 10 μm
Fungal Diversity

Type strain: CBS 100438= ATCC 16027= CDC 45-492-62 -
=MUCL 6683=IFM 52605.
Additional strains examined: see Table 1.
Notes: The species was originally isolated as the etiologic
agent of a kidney mycosis in chinook salmon (Oncorhynchus
tshawytscha) smolts (Doty and Slater 1946). First site of
infection were the mesonephric tubules, extending to the
renal corpuscle. The disease was observed in a fisheries
station while handling smolts returning upstream for hatch-
ing in less than 1 % of the yearlings and attempts to repro-
duce the disease in vitro remained unsuccessful. Hence a low
degree of virulence was supposed. Additional strains of the
species all originated from soil, corroborating the supposi-
tion that its infective ability in fish is purely opportunistic.
The etiologic agent of a visceral mycosis in Masu salmon
and reported to be closely akin to Ochroconis tshawytschae
(Hatai and Kubota 1989) was not available for study and thus
could not be confirmed. A GenBank accession DQ307334
(= NBRC 32268, as S.variabile) from soil, JN578640
(= E000535873, as Scolecobasidium sp.) from grapevine
environment, and FJ914694 (= HSAUP063055, as S.tshawyt-
schae), could now be corrected as O.tshawytschae.
Ochroconis verrucosa (Zachariah, Sankaran & Leelav.)
Samerpitak & de Hoog, comb. nov. MB803668, Fig. 16.
≡Septonema verrucosa Zachariah, Sankaran & Leelav.,
Mycologia. 73: 208, 1981 (basionym).
Description based on CBS 383.81 at 25 °C after 4 weeks
in darkness (Zachariah et al. 1981).
On OA, colonies dark olivaceous brown, with flat, felty,
brown aerial mycelium and dark brown submerged mycelium.
On MEA, colonies attaining 3.5 cm in diam, woolly. Hyphae
subhyaline to pale olivaceous brown, smooth- and thick-
walled, forming a compact mycelium. Conidiophores light
brown, variable in length, non-septate or septate, unbranched
or rarely branched, straight or slightly flexuous, smooth-
walled, 6.3–22.5×2.5–3μm. Conidiogenous cells integrated,
terminal, bearing one or two denticles. Conidia single or in
acropetal, branched or unbranched chains, light brown, cylin-
drical or fusiform, slightly tapered at both ends, mostly 3-
septate (rarely 4-septate), moderately thick-walled, slightly
constricted at septa, verrucose, 7.5–20.0×2.5–4.0 μm.
Type: India, from soil, Calicut University campus. Type
strain CBS 383.81=IMI 211655.
Note: Because of the four-celled conidia, CBS 383.81 had
been misidentified as O.tshawytschae for a long time. Our
phylogenetic analyses clearly showed a large phylogenetic
distance between the two species. Ochroconis verrucosa is
one of a series of relatively closely related species with four-
celled conidia, viz. O.tshawytschae,O.lascauxensis,O.
anellii, and O.verrucosa.
Verruconis Samerpitak & de Hoog, gen. nov. MB519169.
Colonies slow growing, olivacous brown. Conidiophores
absent. Conidiogenous cells erect, pale brown, sparsely septate,
in the apical part with scattered denticles. Conidia produced
sympodially, two-celled, verrucose to almost smooth-walled,
clavate to cylindrical, liberated rhexolytically. Teleomorph
unknown.
Generic type: Diplorhinotrichum gallopavum W.B.
Cooke
Verruconis calidifluminalis (Yarita, A. Sano, de Hoog &
Nishim.) Samerpitak & de Hoog, comb. nov. MB519172.
≡Ochroconis calidifluminalis Yarita, A. Sano, de Hoog &
Nishim., Mycopathologia 170: 29, 2010 (basionym).
On OA, colonies floccose, pale-olivaceous-green, with
radial folds or somewhat crateriform; reverse dark brown,
with a reddish pigment exuded into the agar. Hyphae brown,
with somewhat thickened walls. Conidiogenous cells short,
brown, straight or flexuous, with 1–5 conidia on pronounced
denticles, liberating rhexolytically. Conidia 1-septate, pale to
medium brown, cylindrical to clavate, with or without con-
strictions at the septa, 9.5–20.5 × 2.5–5.0 μm. Cardinal tem-
peratures: good growth at 37 °C, optimum at 42 °C, maxi-
mum at 47 °C.
Type: Japan, Hakone, Kanagawa Prefecture, from water
of a hot stream. Type strain: CBS 125818= IFM 54738.
Habitat: oligotrophic hot water.
Notes: Yarita et al. (2010) described the species, which
was isolated concomitantly with V.gallopava from hot
spring water, as morphologically identical to that species,
but noticed a significant difference at experimental subcuta-
neous inoculation into hydrocortisone-treated mice: the spe-
cies showed low virulence and affected the kidneys, whereas
V.gallopava was a highly virulent neurotroph. The two
species are clearly separate from each other in all gene
fragments analyzed.
Verruconis gallopava (W.B. Cooke) Samerpitak & de
Hoog, comb. nov. MB519171, Fig. 17.
≡Diplorhinotrichum gallopavum W.B. Cooke, in Georg,
Bierer & Cooke, Sabouraudia 3: 241, 1964 (basionym).
≡Dactylaria gallopava (W.B. Cooke) G.C. Bhatt & W.B.
Kendr., Can. J. Bot. 46: 1257, 1968.
≡Ochroconis gallopava (W.B. Cooke) de Hoog, in Howard,
Fung. Path. Hum. Anim. B-II: 181, 1983.
≡Dactylaria constricta (E.V. Abbott) D.M. Dixon &
Salkin var. gallopava (W.B. Cooke) Salkin & D.M. Dixon,
Mycotaxon 29: 379, 1987.
Description based on CBS 437.64 at 25 °C after 4 weeks
in darkness.
On OA, colonies attaining 5.5 cm, smooth to felty, dry,
flat, brown to reddish brown; a pink pigment is exuded into
the agar. On MEA, colony attaining 3.2 cm, smooth to
velvety, dark grey at the center and paler near the edge; a
pink pigment is exuded into the agar. Hyphae brown, with
rather thick walls. Conidiophores flexible, mostly cylindrical
to acicular, with 0(−1) thin septa, poorly differentiated, bear-
ing a few conidia near the apex on fragile denticles. Conidia
Fungal Diversity

Fig. 16 Ochroconis verrucosa, CBS 383.81. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–P. Conidial apparatus with rhexolytic, 4-celled
conidia, produced from short to long cylindrical conidiophores. Q. Conidia and anastomosing hyphae. R. Hyplal coil. Scale bar =10 μm
Fungal Diversity

Fig. 17 Verruconis gallopava, CBS 437.64. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–G, J. Conidial apparatus with rhexolytic conidia.
H. Hyphal coils. I. Anastomosing hyphae. Scale bar=10 μm
Fungal Diversity

two-celled, verruculose to nearly smooth-walled, subhyaline
to pale brown, clavate, constricted at the septum, 11–18× 2.5–
4.5 μm; apical cell wider than basal cell. Conidial secession
rhexolytic, frills remaining on denticle and on conidial base.
Cardinal temperatures: growth abilities ranging from 15 to
50 °C, with optimal growth at 35 °C; growth with 5 % MgCl
2
and 5 % NaCl (Horré et al. 1999).
Type: U.S.A., South Carolina, Bishopville, from brain
abscess of Meleagris gallopavo (turkey), 1964, G. Bierer.
Type strain CBS 437.64= ATCC 16027= CDC 45-492-62 = -
MUCL 6683=IFM 52605.
Additional strains examined: see Table 1.
Habitat: in self-heated environments, hot springs and
warm effluents. Neurotropic opportunist in bird brain tissue,
occasionally in severely immunocompromised humans.
Notes: This species is an obvious thermophile. It has been
encountered in diverse types of hot environments, among
which are thermal soils (Evans 1971a,b;TanseyandBrock
1973;Weitzmanetal.1983;Redmanetal.1999), self-heated
coal waste piles (Tansey and Brock 1973), hot springs (Tansey
and Brock 1973;Weitzmanetal.1983;Yaritaetal.2007), and
warm effluents of a nuclear reaction station (Rippon et al.
1980). Yarita et al. (2007) found the species associated in a
hot spring in Japan with the morphologically similar but mo-
lecularly clearly different species O.calidifluminalis.Ver r u -
conis gallopava was also isolated from broiler-house litter
(Waldrip et al. 1974; Randall and Owen 1981), which was
supposed to be the source of several zoonoses in broiler house
chicken (Georg et al. 1964; Blalock et al. 1973) and trumpeters
(Karesh et al. 1987). Horré and de Hoog (1999) listed 13 brain
infections in young birds and one in a cat brain. The species is
also observed as a neurotrope in humans but then nearly exclu-
sively in severely immunocompromised individuals; for a re-
view is referred to Horré and de Hoog (1999). The species’
neurotropism is exceptional among thermophilic fungi.
The rDNA ITS region is about 670 bp in length, with
52 % mole G+C. The nearest species is V.calidifluminalis,at
a distance of 7.9 % in ITS and thus unambiguously belong-
ing to another taxon. It is remarkable that two sister species
share the same habitat and nevertheless have diverged so
much at the molecular level. Not all environmental strains of
V.gallopava listed in the literature have been confirmed by
sequencing, and thus confusion with V.calidifluminalis is
not excluded.
Verruconis verruculosa (R.Y. Roy, R.S. Dwivedi & R.R.
Mishra) Samerpitak & de Hoog, comb. nov. MB803669,
Fig. 18.
≡Scolecobasidium verruculosum R.Y. Roy, R.S. Dwivedi
& R.R. Mishra, Lloydia 25:164, 1962.
Description based on CBS 119775 (Roy et al. 1962; Ellis
1971).
On OA, colonies effuse, usually slow-growing, hairy, hyaline,
becoming tinged brown to olivaceous brown. Mycelium
immersed and superficial. Conidiophores poorly differentiated,
straight or flexuous, unbranched, brown to olivaceous-brown,
smooth, sparingly septate, 5–45 μm high, 2–3μm wide.
Conidiogenous cells integrated, terminal, sympodial, denticulate,
5–15 μm long, 2–3μm wide. Conidia sparse, sympodial, oblong
with rounded ends, 1-septate, usually strongly constricted at the
septum, pale brown to olivaceous-brown, (6−)7.5(−9) μm long,
(3.5−)4(−5) μm wide, echinulate with prominent spines.
Type: India, grassland soil, New Delhi, India. The type
strain was deposited in the Indian Type Culture Collection,
I.A.R.I., New Delhi.
Discussion
Members of the Ochroconis and Scolecobasidium, are mel-
anized filamentous fungi which are morphologically excep-
tional by having olivaceous colonies often excuding rust
brown to yellowish brown, occasionally reddish pigments
into the medium, and producing roughened, septate conidia
by sympodial conidiogenesis which liberate by rhexolytic
dehiscence. The first teleomorph has been observed in a
member of this group, Ochroconis sexualis. Spherical
cleistothecia are formed bearing thick-walled, antler-shaped
or serrate spines. The bitunicate asci contain eight bicellular,
slightly rough-walled ascospores. The ascocarp, bitunicate
asci and brown, bicellular ascospores are reminiscent of
those of Venturia and related genera. This morphological
resemblance matches with the phylogenetic position of the
Ochroconis/Ve r r u c o n is clade, paraphyletic to the Venturiales,
Sympoventuriaceae. The antler-like spines on the ascocarps
are unknown in the Venturiales (Fig. 14), but using muti-gene
analysis; nuLSU, nuSSU, mtSSU and RPB2 reported by
Machouart et al. (2013), strongly confirmed that the position
of Ochroconis is inside the Sympoventuriaceae.
Nearly all members of Venturiales are plant pathogens,
often with a restricted host range. The ecology of Ochroconis
and Scolecobasidium is enigmatic. A recurrent trait in the
group is oligotrophism, species being found in moist environ-
ments poor in nutrients. Some species are thermophilic, and a
cluster of species is repeatedly found on moist rock in caves.
Opportunistic pathogens are found in the group, mostly on
cold-blooded vertebrates. Some thermophilic species may in-
fect warm-blooded vertebrates, and in the latter case a signif-
icant neurotropic tendency is apparent similarity to the black
yeast-like fungi in the Chaetothyriales (de Hoog et al. 2011).
The Index Fungorum lists 48 names in Scolecobasidium,
and 14 in Ochroconis. Eighteen of these have been redescribed
in the present study, and three new species were added. Au-
thentic material of the majority of the species described as
Scolecobasidium could not be located or/and have never been
available in culture. Seven species, Scolecobasidium arenarium
(Nicot) M.B. Ellis (Ellis 1976; MB323274, CBS 181.58), S.
Fungal Diversity

Fig. 18 Verruconis verruculosa, CBS 119775. A. Colony on MEA 4 wk. B. Colony on OA, 4 wk. C–L. Conidial apparatus with rhexolytic conidia,
produced from cylindrical conidiophores. M, N. Hyphal coil. Scale bar = 10 μm
Fungal Diversity

cateniphorum Matsush. (Matsushima 1975; MB323275, CBS
769.83), S.fusarioideum Matsush. (Matsushima 1993;
MB360912, CBS 210.95), S.excentricum R.F. Castañeda
et al. (1997) (MB443160, type, CBS 469.95), S.fusiforme
(CBS 586.82), S.salinum (G.K. Sutherl.) M.B. Ellis (Ellis
1976; MB323288, CBS 141.60, CBS 619.92, CBS 734.96)
and S.tropicum (CBS 360.87) were proven to be unrelated to
the Ochroconis and Verr u co n is when cultures were studied. In
addition, when their LSU and SSU sequences were analyzed
with Ascomycota members (data not shown), S.cateniphorum
(CBS 769.83) and S.fusarioideum (CBS 210.95) were judged
to be related to the order Chaetothyriales (black yeasts and
allies). S.arenarium (CBS 181.58) and S.salinum (CBS
141.60, CBS 142.60, CBS 619.92, CBS 734.76) were consid-
ered to be members of Pleosporales. Scolecobasidium
excentricum strain CBS 469.95 had resemblance to members
of Sympoventuriaceae basal to the Ochroconis lineage.
Scolecobasidium fusiforme Matsush. (Matsushima 1971;
MB323283, CBS 586.82) belonged to the Helotiales, and
Scolecobasidium tropicum Matsush. (Matsushima 1983;
MB124386, CBS 380.87) clustered in the Pezizomycotina.
Phylogenetic species recognition by genealogical concor-
dance was performed in this study with species where sufficient
numbers of strains were available, viz. Ochroconis mirabilis,O.
constricta,O.minima,O.tshawytschae,O.cordanae,and
Verruconis gallopava. Of remaining species only a single or just
a few isolates were available, and were confirmed as separate
taxa by their remote positions in concordant single-locus gene-
alogies. These species include O.anellii,O.anomala,O.gamsii,
O.humicola,O.lascauxensis,O.longiphorum,O.sexualis,O.
verrucosa,V.calidifluminalis,andV.verruculosa.Furthercom-
plicating factors are high substitution rates and orthology of
protein coding genes. In some cases the gene fragments of
ACT1,BT2 and TEF1 yielded conflicting topologies, which
might be attributed to the occurrence of these factors (Fig. 4),
especially to the high substitution rates of these genes compared
to those of ribosomal genes, which may decrease phylogenetic
accuracy (Barker and Lutzoni 2002). The type of O.constricta,
CBS 202.27 clustered with strains of O.mirabilis in three genes
analyses, but with remaining strains interpreted to be O.
constricta with ribosomal trees. More markers should be inves-
tigated to clarify these ambiguous data.
The characteristics of morphology, habitat and biochem-
ical properties (Tables 1and 2) mostly support the results of
molecular phylogeny and genealogical concordance. The
phenotypic properties also played an important role in de-
cisions to maintain several taxa such as O.constricta and O.
minima as a unity despite considerable intraspecific hetero-
geneity. In the case of O.gamsii, the paraphyletic species
clusters in three single analyses (SSU,BT2 and TEF1) and
the high distances among member strains in all analyses,
especially 22.1 % in ITS, strongly support the existence of
a cryptic species next to O.gamsii.
Topologies of gene-trees were concordant in main traits,
despite considerable variation. Within the group of study
strains, two main clusters were found. Combining phenotypic
characters such as morphology, temperature relations, and
pathogenicity, we judged that the group should comprise at
least two genera, Ochroconis and Ver r u c o n i s .Ve rr u c o n i s con-
tains species bearing two-celled conidia which are mostly
waterborne, thermophilic oligotrophs, eventually pathogenic
to warm-blooded vertebrates. Neurotropism in V.gallopava
has primarily been expressed in (wild) birds; it is likely that
occurrence in this habitat is significantly under-reported. The
genus contains V.verruculosa, isolated from plant root in
Malaysia, as an ancestral species. Ochroconis represents spe-
cies with two- to four-celled conidia which are soil- or water-
borne, mesophilic saprobes, and which occasionally infect
cold-blooded vertebrates.
Partition Homogeneity Test (PHT) detected conflicts in the
combined data set of ACT1,BT2,andTEF1, but there was no
conflict between nuSSU, ITS, and nuLSU. The conflict in
PHT was probably due to the conflicting data of O.constricta
and O.gamsii (Fig. 4). The multi-locus tree yielded high
support for all species, confirming the topology of most single
gene analyses (Fig. 5), which would suggest that the signifi-
cant incongruence in the dataset might not be sufficient to
reduce the phylogenetic accuracy, as mentioned by Cunning-
ham (1997). Three main clusters are recognizable in the
multilocus tree of Ochroconis.Ochroconis cordanae is a
saprobe in soil and on plant material, possibly also occurring
as an endophyte. It is a sister species of O.humicola. The large
cluster at the top of the tree comprises species mostly
exhibiting two-celled conidia: O.mirabilis/O.constricta,
and O.gamsii/O.sexualis are two clusters of sister species,
while O.minima is paraphyletic to all of them. A third,
unresolved median cluster contains species such as O.
tshawytschae,O.verrucosa,O.lascauxensis,O.anellii,O.
anomala and O.longiphorum, with rough-walled conidia and
preponderantly oligotrophic life style.
Scolecobasidium terreum, with CBS 203.27 as type strain
of the oldest species described in Scolecobasidium, was found
to be remote from all remaining members in all analyses, and
always far away from other strains with identical Y-shaped
conidia as described for S.terreum, namely CBS 423.67, CBS
536.69, CBS 116645, and CBS 119792 (data not shown).
These strains clustered with CBS 510.71, the type of
Humicola minima, which also produces bilobate conidia.
Horré et al. (1999) noted that CBS 203.27 had lost its ability
of conidium production, only sterile, melanized hyphae
remaining, which, judging from the distant sequencing results,
might belong to a contaminant. Consequently, S.terreum as
based on CBS 203.27 is now regarded as a dubious species.
LSU and ITS analyses proved to have comparable taxo-
nomic power to resolve both generic and species levels,
although ITS was difficult to align over the entire data set.
Fungal Diversity

Tree topologies of both genes were similar to those resulting
from multilocus analysis. For determination of species
boundaries, the partial protein coding genes ACT1,BT2,
and TEF1 were used; variation of individual gene fragments
were too large to determine the generic topology.
Multilocus analysis was possible only when sets of closely
related strains were compared. The unique heterogeneity be-
tween isolates in Ochroconis and Ver r u c o n i s renders all taxo-
nomic decisions difficult. Given the fact that phenotypic char-
acters vary much less than molecular data, it seems that these
fungi have gone through a process of accelerated evolution.
The paraphyletic position with respect to Sympoventuriales
(Machouart et al. 2013), with significantly longer branches in
the Ochroconis lineages underlines this supposition. To inves-
tigate this hypothesis, distances among data sets were inves-
tigated and compared. The variability of concatenated se-
quences in multilocus analysis was 0.094 substitutions per
site, higher than those of SSU (0.017) and LSU (0.037), but
lower than ITS (0.143). The values for individual protein-
coding gene fragments were for ACT1 (0.285), BT2 (0.158),
and TEF1 (0.376). Especially the latter gene fragment showed
high variability, suggesting diversity within species such as O.
constricta and O.mirabilis which is not supported by any
phenotypic or ecological feature. Lewis et al. (2011)sug-
gested that the gene with the highest mean distance, compared
to the average of all barcoding candidates, would qualify best
as a barcoding marker. From our study, it seems that higher
substitution rates in Ochroconis may lead to over-estimation
of the phylogenetic value of barcoding markers. Because of
the length of the sequence, sufficient phylogenetic informa-
tion content, and ease of amplification, ITS is judged to be the
optimal marker for identification of species of Ochroconis and
Ve r r u c on i s . The discriminatory power of ITS for species
identification also reveals distinguishedly in their length and
G+ C% (Table 2).
The fungi in these genera are particularly known through
Verruconis gallopava, a remarkably thermophilic species of
hot environments such as hot springs, broiler house litter or
self heated coal piles (Redman et al. 1999; Waldrip et al.
1974; Randall and Owen 1981; Yarita et al. 2007). Its sib-
ling, V.calidifluminalis (Yarita et al. 2010) was isolated from
a hot spring together with V.gallopava. The two species
resemble each other in their environmental ecology as they
both can be found as thermophilic oligotrophs in hot water,
and are morphologically and culturally very similar. How-
ever, they have diverged considerably in their ITS region and
also in other genes sequenced, and thus seem to have
maintained coexistence over long periods of time while
occupying the same or a very similar ecological slot. They
differ in their pathogenic potential to vertebrates (Yarita et al.
2010). Verruconis gallopava is a neurotropic invader in birds
and in humans. In cultivated animals such as poultry the
infection may take epidemic proportions (e.g., Georg et al.
1964; Blalock et al. 1973; Randall and Owen 1981; Shane
et al. 1985). In humans usually individuals with an underly-
ing immune disorder are affected, leading to brain infections
that have been categorized as primary encephalitis, because
first symptoms are mostly of a neurological nature (Horré
and de Hoog 1999). The route of infection of cerebral infec-
tion a disease which is particularly prevalent in black yeasts
of the order Chaetothyriales is unknown, neither is the exact
ecological background of this infective ability.
Other species, in the literature attributed to O.humicola and
O tshawytschae, have repeatedly been observed in infections
of fish and other cold-blooded vertebrates. Reidentification of
available isolates in the present study (Table 1) suggested that
particularly O.mirabilis seems to be concerned. Nearly all
strains of this species were derived from water or from the
vertebrates thriving in this water. Ocean water was concerned,
with isolates from a sea-fan, a sponge, sea-grass (Sakayaroj
et al. 2010). One remarkable finding was a human pulmonary
infection in a near-drowned sea diver; CBS 729.96 was isolat-
ed from the air regulator of the diver who aspirated sea water
(Table 1). Additional isolates had prevalently been isolated
from moist environments such as bathing facilities, and from
human skin and nails (G.S. de Hoog, unpublished data). Lian
and de Hoog (2010) suggested that particular, predisposed
groups of melanized waterborne oligotrophs (Satow et al.
2008) may infect moisturized human skin after showering; O.
mirabilis seems to have a similar ecology. A single proven fish-
infection outside O.mirabilis was caused by O.tshawytshae
(Doty and Slater 1946), a species otherwise only known from
soil. Both species are unable to grow at 37 °C, having an
optimum between 15 and 30 °C, and show some halotolerance
ingrowingwith5%NaCland10%MgCl
2
(Horré et al. 1999).
However, the first human infection, subcutaneous mycosis, by
O.tshawytscahe,wasfinallyreportedbyYietal.(2012).
Several other species seem to be soilborne saprobes, judg-
ing from current sources of isolation (Table 1). An exception
may be Ochroconis cordanae which has been repeatedly iso-
lated from living plant tissue: CBS 123536 (= FJ372390), from
living leaves of Encephalartos lebomboensis in South Africa
published by Crous et al. (2008)asOchroconis sp., and two
GenBank submissions: EF419939 (= Endophyte isolate 9165),
isolated as an unnamed fungal endophyte of Platycladus
orientalis by Arnold and Hoffman (2008), FJ752621 (= JIA
6-8-1), isolated by Yuan et al. (2010) as an unnamed endophyte
of wild rice roots (Oryza granulata), and JQ760421 (=isolate
FLO760), an endophytic dothideomycetous fungus by U’Ren
et al. (2012). On the other hand, several strains of O.cordanae,
HQ608103 (= TR072 as S.humicola) and HQ607921
(= ATT286, as Scopulariopsis humicola), which were isolated
from an ant nest (Rodrigues et al. 2011). The first isolation
concerned a nest of Trachymyrmex septentrionalis,andthe
second a nest of Atta texana. The common factor between
these habitats, plant and insect, should be further investigated.
Fungal Diversity

Another interesting habitat was revealed by two recently
described species, O.lascauxensis and O.anomala,whichwere
isolated from the Lascaux Cave in France (Martin-Sanchez et al.
2012). Ochroconis lascauxensis produced black stains on ancient
pictures on the wall, while O.anomala originated from the cave
sediment. This niche is quite similar to that of the sister species,
O.anellii, from a stalactite. This habitat matches with the
oligotropic property of some waterborne Ochroconis species.
Acknowledgments The authors are indebted to K. Yarita, P.M. Martin-
Sanchez, C. Saiz-Jimenez, D. Attili-Angelis, A.P.M Duarte and P. Crous for
strains and information. T. Chaowasku, M.J. Najafzadeh, C. Labuschagne
and K. Khayhan are acknowledged for technical assistance. R. Horré is
thanked for constructive discussion, and S.B.J. Menken for critical reading
of the text.
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