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Microfungi on Tamarix
Kasun M. Thambugala
1,2,3
&Dinushani A. Daranagama
2,3,4
&Alan J. L. Phillips
5
&
Timur S. Bulgakov
6
&Darbhe J. Bhat
7,8
&Erio Camporesi
9,10
&Ali H. Bahkali
11
&
Prapassorn D. Eungwanichayapant
3
&Zuo-Yi Liu
1
&Kevin D. Hyde
2,3
Received: 23 June 2016 /Accepted: 16 July 2016
#School of Science 2016
Abstract Tamar ix species are small trees that grow in various
natural habitats and have a wide geographic distribution.
Microfungal species previously found on Ta ma rix and recently
collected in Italy and Russia were identified based on morpho-
logical characters and analyses of gene sequence data. The sex-
ual morph of the coelomycetous genus Homortomyces was col-
lected for the first time and is described and illustrated. A new
family, Homortomycetaceae (Dothideomycetes, families
incertae sedis) is introduced to accommodate Homortomyces.
Two new genera Neomicrosphaeropsis (Didymellaceae)and
Tamaricicola (Pleosporaceae) are introduced in this paper.
Phoma tamaricicola was recollected and is placed in
Neomicrosphaeropsis based on morphology and molecular data.
Ten new species, Cytospora italica,C. unilocularis,Diaporthe
ravennica,Eutypella tamaricis,Neomicrosphaeropsis italica,
N. novorossica,N. rossica,Keissleriella tamaricicola,
Paracamarosporium tamaricis and Tamaricicola muriformis
are introduced, while Alternaria tenuissima,Dothiorella
sarmentorum,Neofusicoccum luteum,Paraepicoccum
amazonense,Pleospora herbarum and Pseudocamarosporium
propinquum are reported for the first time on Tamari x
spp. with descriptions and illustrations. Multi-gene anal-
yses show that Paraepicoccum amazonense should be
placed in Pleosporineae,Pleosporales, where it is close-
ly related to Camarosporium sensu stricto. Several her-
barium specimens were studied to illustrate other fungal
species recorded on Tamarix species. A comprehensive
account of microfungi on Tama ri x is provided, which
includes a list with data from the literature, as well as
those identified in the present study. The taxonomic
placement of most taxa discussed in this study is based
on a modern taxonomic framework based on analysis of
multi-gene sequence data.
Keywords Dothideomycetes .Homortomycetaceae .
Incertae sedis .New species .Phylogeny .Pleosporales .
Sordariomycetes .Tama ri ca ceae .Taxono my
Fungal Diversity
DOI 10.1007/s13225-016-0371-z
*Zuo-Yi Liu
gzliuzuoyi@163.com
1
Guizhou Key Laboratory of Agricultural Biotechnology, Guizhou
Academy of Agricultural Sciences, Guiyang 550006, Guizhou,
People’s Republic of China
2
Centre of Excellence in Fungal Research, Mae Fah Luang University,
Chiang Rai 57100, Thailand
3
School of Science, Mae Fah Luang University, Chiang Rai 57100,
Thailand
4
State Key Laboratory of Mycology, Institute of Microbiology,
Chinese Academy of Sciences, No 3 1st West Beichen Road,
Chaoyang District, Beijing 100101, People’s Republic of China
5
Faculty of Sciences, Biosystems and Integrative Sciences Institute
(BioISI), University of Lisbon, Campo Grande,
1749-016 Lisbon, Portugal
6
Academy of Biology and Biotechnology, Southern Federal
University, Rostov-on-Don, 344090 Rostov Region, Russia
7
Formerly Department of Botany, Goa University, Goa, India
8
Curca, No. 128/1–J, Azad Housing Society, Goa Velha, India
9
A.M.B. Gruppo Micologico ForliveseBAntonio Cicognani^,Via
Roma 18, Forlì, Italy
10
A.M.B. Circolo Micologico BGiovanni Carini^,C.P.314,
Brescia, Italy
11
Department of Botany and Microbiology, College of Science, King
Saud University, P.O. Box: 2455, Riyadh 1145, Saudi Arabia
Table of Contents
Dothideomycetes
Botryosphaeriaceae (Botryosphaeriales)
1. Botryosphaeria tamaricis (Cooke) Theiss. & Syd.
2. Diplodia tamaricina Sacc.
3. Dothiorella sarmentorum (Fr.) A.J.L. Phillips, J. Luque
&A.Alves
4. Neofusicoccum luteum (Pennycook & Samuels) Crous,
Slippers & A.J.L. Phillips
Pleosporaceae (Pleosporales)
5. Alternaria tenuissima (Kunze) Wiltshire
6. Pleospora herbarum (Pers.) Rabenh.
7. Tama ri ci cola Thambugala, E. Camporesi & K.D. Hyde,
gen. nov.
8. Tamaricicola muriformis Thambugala, Camporesi &
K.D. Hyde, sp. nov.
Didymellaceae Gruyter, Aveskamp & Verkley (Pleosporales)
9. Neomicrosphaeropsis Thambugala, Camporesi & K.D.
Hyde, gen. nov.
10. Neomicrosphaeropsis italica Thambugala, Camporesi
& K.D. Hyde, sp. nov.
11. Neomicrosphaeropsis novorossica Thambugala,
Bulgakov & K.D. Hyde, sp.nov.
12. Neomicrosphaeropsis rossica Thambugala, Bulgakov
&K.D.Hyde,sp.nov.
13. Neomicrosphaeropsis tamaricicola Wanasinghe,
Thambugala & K.D. Hyde, comb.nov.
Didymosphaeriaceae Munk (Pleosporales)
14. Pseudocamarosporium propinquum (Sacc.) Wijayaw.,
Camporesi & K.D. Hyde
15. Paracamarosporium tamaricis Thambugala, Camporesi
& K.D. Hyde, sp. nov.
Lentitheciaceae Yin. Zhang, C.L. Schoch, J. Fourn., Crous &
K.D. Hyde (Pleosporales)
16. Keissleriella tamaricicola Thambugala, Camporesi &
K.D. Hyde, sp.nov.
Pleosporales, genera incertae sedis
17. Paraepicoccum amazonense Matsush.
StigmatodiscaceaeVoglmayr & Jaklitsch (Stigmatodiscales)
18. Asterodiscus tamaricis Voglmayr, Gardiennet &
Jaklitsch
Valsariaceae Jaklitsch, K.D. Hyde & Voglmayr (Valsariales)
19. Valsaria tamaricis Mundk. & S. Ahmad
Dothideomycetes, families incertae sedis
Homortomycetaceae Thambugala, A.J.L. Phillips & K.D.
Hyde, fam.nov.
20. Homortomycetaceae Thambugala, A.J.L. Phillips &
K.D. Hyde, fam.nov.
21. Homortomyces tamaricis Wijayaw., Camporesi &
K.D. Hyde
Sordariomycetes
Va l s a c ea e Tul. & C. Tul (Diaporthales)
22. Cytospora tamaricis Brunaud
23. Cytospora italica Thambugala, Camporesi & K.D.
Hyde, sp.nov.
24. Cytospora unilocularis Thambugala, Camporesi &
K.D. Hyde, sp.nov.
Diaporthaceae Höhn. ex Wehm. (Diaporthales)
25. Diaporthe ravennica Thambugala, Camporesi & K.D.
Hyde, sp.nov.
Diatrypaceae Nitschke (Xylariales)
26. Eutypella tamaricis Thambugala, Camporesi & K.D.
Hyde, sp.nov.
Sordariomycetes genera incertae sedis
27. Coryneopsis tamaricis (Cooke) Grove
Introduction
Tamari x is a plant genus of small trees or shrubs belonging to
the family Tam ar ica ce ae and commonly known as saltcedar
or tamarisk. There are about 60 species in the genus. Species
of this genus grow in various natural habitats, such as riparian
regions, deserts, dunes, wetlands and river banks. They are
widely distributed on saline soils in arid and semi-arid regions
of Central, Southern and Eastern Asia, Southern Europe, the
eastern Mediterranean and Africa, and are introduced invasive
species in the southwestern part of North America (Brock
1994; Brotherson and Field 1987;Geetal.2006;Di
Tomaso 1998; Venturella et al. 2007). Tamarix species are
deciduous or evergreen plants and bear small pink or white
bisexual flowers in catkin-like racemes (Brock 1994;Saidana
et al. 2008). They have multiple stems and slender branches,
which are brown to reddish brown or blackish brown. The
leaves are scale-like, overlapping and sessile, with narrow
bases (Brock 1994; Venturella et al. 2007)(Fig.1).
Tamari x species are also cultivated as ornamental plants,
windbreaks and shade trees, especially in artificial forests in
Central Asian countries and China. Some Tamarix species
have antioxidant and antimicrobial activities (Saidana et al.
2008;Zainetal.2012). Although tamarisk is planted as an
Fungal Diversity
ornamental, it can become a weed and causes management
problems in western North American riparian areas
(De Loach 1989;BrothersonandField1987). Their
extensive root system is stable and resistant to erosion
(Brotherson and Field 1987). Tam ari x species are toler-
ant to desiccation in water courses and have assumed
greater dominance in desert flood plains, due to their
superior drought tolerance and ability to produce high
density stands with high leaf areas (Blackburn et al.
1982;Cleverlyetal.1997). Therefore, Tama ri x species,
such as T. aphylla (L.) Karst.,T.nilotica (Ehrenb.)
Bunge, T.ramosissima Ledeb., are widely distributed in desert
areas and dunes (Brock 1994; Rekah et al. 2001; Venturella
et al. 2007). They have been used for medicinal purposes
andastonics(Brock1994; Sultanova et al. 2001; Venturella
et al. 2007).
Although a number of sexual and asexual fungal species
have been reported on Tamari x species (Farr and Rossman
2016; Table 4), only some have good illustrations and gene
sequence data. In this study, we collected microfungi on
Tamari x species inItaly and Russia to establish the microfungi
associated with this host. We also studied herbarium speci-
mens to illustrate other fungal taxa recorded on Tam ar ix spe-
cies. These materials are used to provide a comprehensive
account of microfungi on Tamari x and fresh collections were
used to establish a natural placement of the microfungi in the
Ascomycota, based on sequence data and morphological
traits. The data provided will help establish the numbers of
fungi.
Materials and methods
Sample collection, morphological study and isolation
Fungal species associated with Tama ri x hosts were collected
from Italy (Province of Forlì-Cesena and Ravenna) and Russia
(Rostov Region). Herbarium specimens were obtained from
BPI, MFLU and S. Fungi from fresh material were isolated
by a modified single spore/conidial isolation method
(Manamgoda et al. 2012; Chomnunti et al. 2014). Growth
rate, colony characteristics and asexual morph morphology
were determined from cultures grown on 2 % potato-
dextrose agar (PDA) at room temperature (25 °C) in the
dark. Morphological observations and photomicrographs
were made following the method of Thambugala et al.
(2015). Ex-type or representative cultures are deposited in
Mae Fah Luang University Culture Collection (MFLUCC) with
duplicates in International Collection of Microorganisms from
Plants (ICMP), or Guizhou Culture Collection (GZCC). Freshly
collected specimens are deposited in the Herbarium of Mae Fah
Luang University (MFLU), Thailand; Hirosaki University
(HHUF), Japan and Guizhou Academy of Agricultural
Sciences (GZAAS), China. Taxonomic descriptions are
deposited in the FoF database as described in Jayasiri
et al. (2015) and Index Fungorum numbers were obtained as
detailed in Index Fungorum (2016).
DNA extraction, PCR amplification and sequencing
DNA extraction followed the method of Thambugala et al.
(2015). The PCR amplifications were performed in a total vol-
ume of 25 μL of PCR mixtures containing 8.5–9.5 μL ddH
2
O,
12.5 μL 2 × PCR Master Mix (TIANGEN Co., China), 1–2μL
of DNA template, 1 μL of each primer. The EF1-α,ITS,LSU,
SSU, RPB2 and TUB gene regions were amplified for relevant
strains following the conditions stipulated in Table 1.ThePCR
products were visualized under UV light on 1 % agarose elec-
trophoresis gels stained with ethidium bromide. Purification and
sequencing of PCR products were carried out at Invitrogen
Biotechnology Co., Shanghai, China.
Phylogenetic analyses
Phylogenetic analyses based on selected ITS, LSU, SSU,
EF1-α,β-tubulin and RPB2 sequence data were carried out
to establish the phylogeneticplacement of each isolated taxon.
Sequences were aligned with Bioedit 7.1.3.0 (Hall 1999)
and the consensus sequences were further improved with
MUSCLEimplementedinMEGA5v(Tamuraetal.
2011). Alignments were checked and optimized manually
when necessary.
Phylogenetic analyses were performed based on maximum
likelihood (ML) criterion using RAxML-HPC BlackBox
(8.2.4) (Stamatakis 2006;Stamatakisetal.2008)inthe
CIPRES portal (Miller et al. 2010). The general time re-
versible model of evolution including estimation of invari-
able sites (GTRGAMMA + I) and assuming a discrete
gamma distribution with four rate categories was used
for the ML analysis. Trees were rooted with given
outgroups in each analysis. The best scoring trees were
selected and visualized with MEGA v. 5 (Tamura et al.
2011). ML Bootstrap supports (BS) (greater than or equal
to 50 %) are shown below or above each branch. All the
sequences newly generated in this study are deposited in
GenBank (Table 2). The resulting phylogenetic trees are
presented under each relevant description.
Results
Dothideomycetes
Recent treatments of Dothideomycetes are those Hyde et al.
(2013) and Wijayawardene et al. (2014d) and their treatments
of orders and families are followed here.
Fungal Diversity
Fig. 1 a-cHabitats d Scale-like leaves e-g Inflorescences of different Tamarix species. Tamarix gallica (d), T. smyrn ensis (a), T. ramosissima (b,c,f,g,),
T. la xa (e). Photos by Timur Bulgakov, Erio Camporesi, Anatoly Lisitsin, Yuri Rebriev, Roland Tsandekidis, Tatiana Vinokurova
Fungal Diversity
Botryosphaeriales C.L. Schoch, Crous & Shoemaker, in
Schoch, Shoemaker, Seifert, Hambleton, Spatafora & Crous,
Mycologia 98(6): 1050 (2007) [2006]
Botryosphaeriaceae Theiss. & Syd. [as‘Botryosphaeriacae’],
Annls mycol. 16(1/2): 16 (1918).
The family Botryosphaeriaceae is considered to be one of
the largest families in the class Dothideomycetes. Members of
this family are pathogens, endophytes or saprobes with a cos-
mopolitan distribution (Liu et al. 2012;Hydeetal.2013;
Phillips et al. 2013). Botryosphaeriaceae is characterized by
uni- to multi-loculate ascostromata, sparse hypha-like
pseudoparaphyses, 8-spored, bitunicate, asci and hyaline to
pigmented, aseptate to septate ascospores. The asexual morph
is coelomycetous and produces uni- to multi-locular pycnidial
conidiomata, with hyaline, phialidic conidiogenous cells and
hyaline or pigmented, aseptate or septate conidia, sometimes
with mucoid appendages or sheaths (Liu et al. 2012;Hyde
et al. 2013; Phillips et al. 2013;Thambugalaetal.2014a).
Botryosphaeria Ces. & De Not., Comm. Soc. crittog. Ital.
1(4): 211 (1863)
The genus Botryosphaeria was introduced by Cesati and
De Notaris (1863), while Theissen and Sydow (1918)placed
it in Botryosphaeriaceae. This genus is based on the type
species B.dothidea, which typically has pseudothecial
ascostromata forming a botryose aggregate, hyaline, aseptate
ascospores and stromatic conidiomata with fusiform, hyaline,
aseptate conidia, that can become dark-walled and septate
with age (Phillips et al. 2005,2013; Liu et al. 2012).
Although there are 275 epithets listed in Index Fungorum
(2016), only seven species are currently recognized in
Botryosphaeria (Phillips et al. 2013) according to modern
taxonomic concepts based on morphology plus molecular
data.
Botryosphaeria tamaricis (Cooke) Theiss. & Syd., Annls
mycol. 13(5/6): 663 (1915), Fig. 2
Basionym: Dothidea tamaricis Cooke, Grevillea 11(no.
59): 108 (1883)
Facesoffungi number: FoF02146
Saprobic on branches of Tam ar ix species. Sexual morph:
Ascostromata 300–400 μmhigh×550–650 μmd
iam.
(x=330×613μm, n= 5), immersed to partially erumpent
through the bark, gregarious or solitary, forming a botryose
aggregate, black, multi-loculate, with 3–5 locules; indi-
vidual locules, 175–280 μm high × 145–265 μmdiam.
(x= 212 × 189 μm, n= 10), sometimes with a central
ostiole. Peridium of locules 15–40 μm wide, two-layered,
outer layer composed of dark brown, thick-walled cells
of textura angularis, inner layer composed of lightly
pigmented, thin-walled cells of textura angularis lining
the locule. Hamathecium composed of 1–2μmwide,
hypha-like, septate pseudoparaphyses. Asci 60–80 × 11.5–
17 μm(
x=67.8×13.5μm, n= 10), 8-spored, bitunicate,
fissitunicate, clavate, short pedicellate, apically rounded, with a
small ocular chamber. Ascospores 17.5–23 × 5–8.5 μm
(x= 20.9 × 6.8 μm, n= 20), overlapping, uniseriate at the base,
biseriate at the apex, hyaline, aseptate, fusoid to ovoid, smooth-
walled, with granular contents. Asexual morph: Undetermined.
Material examined: USA, South Carolina, Aiken, on
Tamari x sp. (Tamaricaceae), H.W. Ravenel (S-F49472,
holotype of Dothidea tamaricis).
Notes: This species is morphologically similar to the type
species of Botryosphaeria,B.dothidea (Moug.) Ces. & De
Table 1 Primers used during this study with respective PCR amplification conditions
Locus Primers (Forward and Reverse) PCR amplification Conditions Reference
ITS ITS4 and ITS5 94 °C: 3 min, (94 °C: 30 s, 55 °C: 30 s,
72 °C: 1 min) × 34 cycles; 72 °C: 5 min
White et al. 1990
LSU LROR and LR5 94 °C: 5 min, (94 °C: 1 min, 53 °C: 50 s,
72 °C: 1 min) × 37 cycles, 72 °C: 10 min
Vilgalys and Hester 1990
RNA polymerase II second
largest subunit (RPB2)
fRPB2-5F and fRPB2–7cR 95 °C: 5 min, (95 °C: 15 s, 56 °C: 50 s,
72 °C: 2 min) × 37 cycles, 72 °C: 10 min
Liu et al. 1999
SSU NS1 and NS4 94 °C: 4 min, (94 °C: 50 s, 56 °C: 1 min,
72 °C: 1 min,72 °C: 10 min) × 37 cycles
White et al. 1990
Translation elongation
factor-1α
EF1–728F and EF1–986R, (95 °C: 5 min, 95 °C: 30 s, 58 °C: 50 s,
72 °C: 1 min) × 40 cycles, 72 °C: 10 min
Rehner 2001; Carbone and Kohn 1999
EF1–983F and EF1–2218R
β-tubulin Bt2a and Bt2b 95 °C: 5 min, (95 °C: 30 s, 58 °C: 50 s,
72 °C: 1 min) × 40 cycles; 72 °C: 10 min
Glass and Donaldson 1995
BT2Fw and BT4Rd 95 °C: 5 min, (94 °C: 30 s, 55 °C: 50 s,
72 °C: 1 min) × 40 cycles; 72 °C: 7 min
Woudenberg et al. 2009
Fungal Diversity
Not., except in the slight variation in the asci and ascospore
dimensions (Liu et al. 2012; Phillips et al. 2013). However,
fresh collections and molecular analyses are essential to estab-
lish whether B.tamaricis is a synonym of B.dothidea.
Diplodia Fr., in Montagne, Annls Sci. Nat., Bot., sér. 2 1: 302
(1834)
Diplodia is a well-known genus and is typified by Diplodia
mutila (Fr.: Fr. ) Fr. Diplodia species are pathogens or
saprobes, mainly on woody hosts with a worldwide distribu-
tion (Phillips et al. 2013;Hydeetal.2014;Ariyawansaetal.
2015b). There are 1251 species epithets for Diplodia in Index
Fungorum (2016) and recollection, epitypification and molec-
ular analysis are required to confirm the placement of those
species, as currently only 17 Diplodia species are known from
a molecular basis (Phillips et al. 2013).
Diplodia tamaricina Sacc., Syll. fung. (Abellini) 3: 343
(1884), Fig. 3
Table 2 GenBank and culture collection accession numbers of species generated in the present study
GenBank accession numbers
Species Isolate number LSU ITS SSU RPB2 EF1-αβ-tubulin
Alternaria tenuissima MFLUCC 14–0441, ICMP 20706 KU561876 KU752186 KU870907 _ KU577440 _
Asterodiscus tamaricis MFLUCC 13–0072, ICMP 20692 KU729859 KU900324 KU900314 _ _ _
MFLUCC 14–0994,ICMP20744KU729860_____
Cytospora italica MFLUCC 14–0440,ICMP20691KU900301KU900329____
Cytospora unilocularis MFLUCC 15–0481, ICMP 21252 KU900304 KU900332 _ KX011166 _ KX444199
MFLUCC 15–0482 KU900305 KU900333 _ KX017567 _ KX029120
MFLUCC 15–0483 KU900306KU900334___KX063631
Diaporthe ravennica MFLUCC 15–0479, ICMP 21251 KU900307 KU900335 _ _ KX365197 KX432254
MFLUCC 15–0480 KU900308 KU900336 _ _ KX426703 KX377688
Dothiorella sarmentorum MFLUCC 14–0579, ICMP 20752 KU561877 KU752183 KU870904 _ KU561873 _
Eutypella tamaricis MFLUCC 14–0444,ICMP20705KU900302KU900330____
MFLUCC 14–0445 KU900303 KU900331 _ _ KX001803 KX453302
Homortomyces tamaricis MFLUCC 13–0280, ICMP 20745 KU561874 KU752184 KU870905 _ _ _
MFLUCC 14–0167,ICMP21071KU561875KU934190____
Keissleriella tamaricicola MFLUCC 14–0168,ICMP20704KU900300KU900328____
Neofusicoccum luteum MFLUCC 16–0504, GZCC 15–
0050
KU900291 KU900316 KU870903 _ KU534414 KX453297
Neomicrosphaeropsis italica MFLUCC 15–0485, ICMP 21253 KU729854 KU900318 KU900309 KU674820 _ _
MFLUCC 15–0484 KU729853 KU900319 _ KU695539 KU981057 KX453298
MFLUCC 16–0284, GZCC 15–
0049
KU900296 KU900321 KU900311 KU714604 KU991165 KX453299
Neomicrosphaeropsis novorossica MFLUCC 14–0578, ICMP 20751 KX198710 KX198709 KX198711 _ _ _
Neomicrosphaeropsis rossica MFLUCC 14–0586, ICMP 20753 KU729855 KU752192 KU870914 _ KU960943 _
Neomicrosphaeropsis
tamaricicola
MFLUCC 14–0443, ICMP 20708 KU729851 KU900322 KU900312 _ KU949580 _
MFLUCC 14–0439, ICMP 20743 KU729858 KU900323 KU900313 _ _ _
Paracamarosporium tamaricis MFLUCC 15–0494 KU900298KU900326___KX453300
MFLUCC 15–0
495 KU900299KU900327___KX453301
Paraepicoccum amazonense MFLUCC 15–0493, ICMP 21255 KU900294 KU752190 KU870912 KU820871 KU663647 _
MFLUCC 15–0491 KU900295 KU752191 KU870913 KU820872 KU663648 _
Pleospora herbarum MFLUCC 14–0442, ICMP 20707 KU561878 KU752185 KU870906 _ _ _
Pseudocamarosporium
propinquum
MFLUCC 14–0166, ICMP 20709 KU900297 KU900325 KU900315 _ _ _
Tamaricicola muriformis MFLUCC 15–0488, ICMP 21254 KU561879 KU752187 KU870909 KU820870 KU577441 _
MFLUCC 15–0489 KU729857 KU752188 KU870910 _ KU600013 _
MFLUCC 15–0490 KU729856 KU752189 KU870911 _ KU600014 _
MFLUCC 16–0488 KU900293 KU900317 KU870908 _ _ _
Fungal Diversity
Fig. 2 Botryosphaeria tamaricis (holotype) a Herbarium material b, c Ascostromata on host surface dVertical section through ascostroma eClose-up
of locule fPeridium g, h Mature and immature asci i-l Ascospores. Scale bars: d-f =50μm, g, h =30μm, i-l =10μm
Fungal Diversity
Facesoffungi number: FoF02147
Saprobic on stems of Tamari x ga llica L. Sexual morph:
Undetermined. Asexual morph:Conidiomata 120–320 μm
high × 280–500 μmdiam.(
x= 268 × 418 μm, n=6),
stromatic, solitary to gregarious, immersed to partially
erumpent, black, globose to subglobose with a flattened base,
uniloculate, sometimes with 2 locules, ostiolate. Conidiomatal
wall 35–70 μm(
x=47μm, n= 20) wide, composed of several
layers of dark brown to lightly pigmented cells of textura
angularis, becoming hyaline towards the conidiogenous region.
Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 6–15 × 1–3μm(
x= 10.4 × 1.9 μm, n= 15), hyaline,
smooth, holoblastic forming conidia at their tips. Conidia
13.5–21 × 8–12 μm(
x= 16.8 × 10 μm, n=40),palebrown
to brown, oblong to sub-cylindrical, with broadly rounded ends,
Fig. 3 Diplodia tamaricina (holotype)aHerbarium material b, c Conidiomata on host surface d, e Vertical sections through conidiomata f, g
Conidiomatal wall h-j Conidiogenous cells and developing conidia kConidia. Scale bars: d, e =100μm, f, g,k=25μm, h-j =10μm
Fungal Diversity
Fig. 4 Phylogram resulting from maximum likelihood (RAxML)
analysis of a combined ITS and EF1-αsequence alignment of
Dothiorella species. Maximum likelihood bootstrap support values equal
or greater than 50 % are indicated above or below the nodes. The
new isolate is in blue. The tree is rooted to Spencermartinsia viticola
Fungal Diversity
aseptate when immature, becoming 1-septate when mature,
slightly constricted at septum, smooth-walled.
Material examined: FRANCE, Gallia, Rochefort., on
stems of Tama ri x gal lica L. (syn. T. anglica Webb)
(Tamari ca ce ae), P. Brunaud (S-F 45168, holotype).
Notes:AlthoughDiplodia tamaricina primarily has typical
morphological characters of Diplodia, molecular analysis of
sequence data is essential to confirm the placement in
Botryosphaeriaceae based on modern taxonomic concepts.
Dothiorella Sacc., Michelia 2(6): 5 (1880)
The genus Dothiorella was introduced by Saccardo
(1880a)withD.pyrenophora Berk. ex Sacc. as the type spe-
cies. The sexual morph of Dothiorella species are rarely re-
ported in nature, but asexual species are widely distributed on
many host plants (Phillips et al. 2005,2013;Jamietal.2012;
Dissanayake et al. 2016). More than 375 species names have
been recorded in Dothiorella (Index Fungorum 2016), but
currently only few species have gene sequence data.
DNA phylogeny
Reference sequences used in the phylogenetic analyses were
selected based on the data from Phillips et al. (2013), and
the present phylogenetic analysis based on ITS and EF1-α
sequence data showed 17 subclades, representing 17 dis-
tinct species in Dothiorella.Spencermartinsia viticola
(CBS 117009) was selected as outgroup taxon. Our strain
MFLUCC 14–0579 clustered together with the two strains of
D.sarmentorum including the ex-type strain (IMI 63581b) and
is phylogenetically not distinct (Fig. 4). Therefore, our collec-
tion (MFLUCC 14–0579 /MFLU 14–0604) is treated as a
voucher specimen for the asexual morph of D.sarmentorum.
Dothiorella sarmentorum (Fr.) A.J.L. Phillips, J. Luque & A.
Alves, Mycologia 97: 522 (2005), Fig. 5
Basionym: Sphaeria sarmentorum Fr., K. svenska
Vetensk-Acad. Handl. 39: 107. 1818.
≡Diplodia sarmentorum (Fr.) Fr., Summ. veg. Scand.
(Stockholm) 2: 417. 1849.
=Diplodia pruni Fuckel, Jahrb. Nassauischen Vereins
Naturk., 23–24: 169. 1870 [1869].
=Botryosphaeria sarmentorum A.J.L. Phillips, J. Luque &
A. Alves, Mycologia 97: 522. 2005.
Facesoffungi number: FoF02148
Fig. 5 Dothiorella sarmentorum (MFLU 14–0604) aAppearance of conidiomata on the host bVertical section through conidiomata cConidiomatal
wall dConidia developing on conidiogenous cells eMature dark brown conidia. Scale bars: b=300μm, c, d =50μm, e=10μm
Fig. 6 Phylogram resulting from maximum likelihood (RAxML) analysis
of a combined ITS and EF1-αsequence alignment of Neofusicoccum
species. Maximum likelihood bootstrap support values equal or greater
than 50 % are indicated above or below the nodes. The new isolate is in
blue. The tree is rooted to Dothiorella sarmentorum
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Saprobic on a wide range of hosts. Sexual morph:
Undetermined on Tama ri x host (see Phillips et al. 2013 for a
description). Asexual morph:Conidiomata 300–440 μm
high × 215–300 μmdiam.(
x= 376 × 250 μm, n=5),
stromatic, solitary or scattered in small groups, immersed,
uniloculate, individual or aggregated, black, globose to
subglobose, ostiolate. Conidiomatal wall 19–32(−45) μm
(x=37μm, n= 15), comprising several layers; outer
layers composed of thick-walled, dark brown, somewhat
flattened cells of textura angularis and inner layers of
larger, thin-walled, lightly pigmented or hyaline cells.
Conidiophores reduced to conidiogenous cells. Conidiogenous
cells 6.8–15.2 × 2.4–4.2 μm(
x= 10.1 × 3.3 μm, n= 15), lining
the conidiomatal cavity, holoblastic, hyaline, subcylindrical,
proliferating at the same level giving rise to periclinal thicken-
ings. Conidia 18.2–23.1 × 7.8–10.3 μm(
x=21×9.4μm,
n= 30), ovoid, with a broadly rounded apex and truncate base,
initially hyaline to lightly pigmented and aseptate, becoming
dark brown and 1-septate, slightly constricted at the septum,
smooth-walled.
Culture characteristics: Conidia germinating on PDAwith-
in 18 h and germ tubes produced from one or both cells.
Colonies fast growing on PDA at 25 °C, covering the medium
surface (9 cm Petri-dish) in 4 days, circular, flat, moderately
dense, surface initially white, becoming greenish olivaceous
to greyish within 7 days, smooth surface with entire to slightly
undulate edge.
Fig. 7 Neofusicoccum luteum (MFLU 16–0664) aAppearance of conidiomata on the host bVertical section through conidioma c, d Conidia
developing on conidiogenous cells eConidia. Scale bars: b=100μm, c, e =20μm, d=10μm
Fig. 8 Maximum likelihood (ML) tree from analysis of a combined
LSU, SSU, RPB2 and ITS dataset of Pleosporinae, Pleosporales.
Bootstrap support values equal or greater than 50 % are given above or
below the nodes. The tree is rooted to Halojulella avicenniae (BCC
20173). Newly generated sequences are in blue
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Fig. 8 (continued)
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Material examined: RUSSIA, Rostov Region, Shakhty
City, Central Park, on Tamarix ramosissima Ledeb.
(Tamaricaceae), 16 March 2014, Timur Bulgakov 74–2
(MFLU 14–0604), living culture MFLUCC 14–0579, ICMP
20752.
Notes: Phillips et al. (2005) introduced Dothiorella
sarmentorum based on the asexual morph of Botryosphaeria
sarmentorum. This species has a worldwide distribution and
has been recorded from 34 host species (Phillips et al. 2005;
Phillips et al. 2013). The sexual morph of D. sarmentorum is
characterized by partially erumpent ascomata with papillate
ostioles, 4–6(−8)-spored asci and oblong to ovate, (0–)1-sep-
tate, finely verruculose ascospores, widest in the middle part
(Phillips et al. 2013).
Neofusicoccum Crous, Slippers & A.J.L. Phillips, Stud.
Mycol. 55: 247 (2006)
Neofusicoccum is morphologically similar to
Botryosphaeria and it is difficult to separate the two genera.
Most of the species of the genus had previously been treated
as Fusicoccum and dichomera-like synasexual morphs in
Neofusicoccum have been used to differentiate
Neofusicoccum from Botryosphaeria.However,thisisnota
good character to separate these two genera as dichomera-like
synasexual morphs have been reported for some
Botryosphaeria species and all Neofusicoccum species do
not form dichomera-like synasexual morphs (Crous et al.
2006; Liu et al. 2012; Phillips et al. 2013). However, recent
studies based on multi-gene phylogenetic analyses showed
that Botryosphaeria and Neofusicoccum are distinct genera
in Botryosphaeriaceae (Liu et al. 2012; Phillips et al. 2013;
Thambugala et al. 2014a).
DNA phylogeny
Reference sequences used in the phylogenetic analyses were
selected based on the data from Phillips et al. (2013)and
Dothiorella sarmentorum (IMI 63581b) was selected as
outgroup taxon. Phylogenetic analysis based on combined
ITS and EF1-αsequence data of Neofusicoccum species re-
vealed 19 subclades representing 19 distinct species. Our
strain MFLUCC 16–0504 grouped together with ex-type
strains of N.luteum (sexual and asexual morphs ex-type
strains: CBS 110299 and CBS 562.92). The subclade
representing Neofusicoccum luteum received high bootstrap
support (96 %) in the present phylogenetic analysis (Fig. 6).
Neofusicoccum luteum (Pennycook & Samuels) Crous, Slippers
& A.J.L. Phillips, Stud. Mycol. 55: 248 (2006). Fig. 7
Basionym: Fusicoccum luteum Pennycook & Samuels,
Mycotaxon 24: 456. 1985.
=Botryosphaeria lutea A.J.L. Phillips, Sydowia 54: 70
(2002).
Facesoffungi number: FoF02149
Saprobic or pathogenic on fruits, stems and twigs of woody
plants. Sexual morph: Undetermined on Ta marix host
(see Phillips et al. 2013 for a description). Asexual
morph:Conidiomata 250–310 μm high × 350–525 μm
diam. (x=280×431μm, n= 5), stromatic, solitary to
gregarious, immersed, partially erumpent at maturity, dark
brown to black, uni- to multi-loculate. Locules 55–225 μm
high × 60–170 μmdiam.(
x=113×102μm, n=10),
ostiolate. Peridium up to 120 μm wide, composed of
several layers of small, heavily pigmented, thick-walled
cells of textura angularis, becoming hyaline and smaller
towards the conidiogenous region. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 6.8–24.5 × 1.7–
3.6 μm(
x= 14.9 × 2.5 μm, n= 25), holoblastic, phialidic,
integrated, hyaline, smooth, cylindrical with periclinal
thickening. Conidia 16.5–23(−27) × 3.8–6.2(−7.1) μm
(x= 20.4 × 5.5 μm, n= 35), hyaline, thin-walled,
aseptate, ellipsoidal, widest in the middle or upper third of
the conidium, apex subobtuse, base truncate, smooth-walled.
Culture characteristics:Conidia germinating on PDA with-
in 24 h and germ tubes produced from one or both ends.
Colonies fast growing on PDA at 25 °C, covering the medium
surface (9 cm Petri-dish) in 5 days, circular, flat, dense, surface
white, becoming light iron-grey after 14 days, smooth surface,
with entire to slightly undulate edge.
Material examined: ITALY, Province of Ravenna [RA],
Lido di Dante, on dead branches of Tama rix sp.
(Tamari ca ce ae), 19 March 2015, Erio Camporesi IT 2421–2
(MFLU 16–0664 and GZAAS), living culture MFLUCC 16–
0504, GZCC 15–0050.
Notes:Neofusicoccum luteum has been reported on various
woody hosts and is an important plant pathogen (Pennycook
and Samuels 1985; Phillips et al. 2002,2013; Sergeeva et al.
2009). This is the first record of N.luteum on Tama ri x species
(Phillips et al. 2013). The sexual morph of N.luteum is char-
acterized by uni- or multi-loculate ascostromata, cylindrical,
to clavate asci and oval to broadly fusiform, hyaline, aseptate
ascospores (Phillips et al. 2013).
Pleosporales Luttr. ex M.E. Barr, Prodr. Cl. Loculoasc.
(Amherst): 67 (1987)
Pleosporaceae Nitschke, Verh. naturh. Ver. preuss. Rheinl.
26: 74 (1869)
Pleosporaceae is the largest family in the order
Pleosporales and the members of this family are pathogenic
or saprobic on wood and dead herbaceous stems or
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leaves. The family Pleosporaceae is characterized by
immersed to erumpent or nearly superficial ascomata,
septate, cellular pseudoparaphyses, 8-spored, bitunicate
asci and phragmosporous or muriform, brown or pale
brown ascospores with coelomycetous or hyphomy-
cetous asexual morph (Zhang et al. 2012;Ariyawansa
et al. 2015a). Ariyawansa et al. (2015a) revised the
family and accepted 18 genera. In this study Alternaria
tenuissima and Pleospora herbarum are reported for the
first time on Tam ari x spp. and a new genus is introduced
in Pleosporaceae based on morphological traits and
phylogeny.
DNA phylogeny
Blast searches at GenBank were carried out for newly gener-
ated sequences in order to reveal the closest taxa. Sequence
data for Pleosporineae,Pleosporales were selected from
Wijayawardene et al. (2014c) and Ariyawansa et al. (2015a).
Halojulella avicenniae (BBC 20173) was selected as the
outgroup taxon. In the phylogenetic analysis (Fig. 8), 14 gen-
era can be distinguished in the Pleosporaceae clade and
all are supported by moderate to high bootstrap values.
Strain MFLUCC 14–0441 clustered with the putatively
named strain of A. tenuissima (CBS 918.96) in the section
Alternata, while MFLUCC 14–0442 grouped together with
strains of Pleospora herbarum in the genus Pleospora.A
new genus Tam ar icico la grouped in Pleosporaceae as a
sister clade to Comoclathris with good BS support value
(92 %).
Alternaria tenuissima (Kunze) Wiltshire, Trans. Br. mycol.
Soc. 18(2): 157 (1933), Fig. 9
Basionym: Helminthosporium tenuissimum Kunze [as
‘Helmisporium’], in Nees & Nees, Nova Acta Phys.-
Med. Acad. Caes. Leop.-Carol. Nat. Cur. 9: 242 (1818)
Facesoffungi number: FoF02152
Pathogenic or saprobic on wide range of hosts. Sexual
morph: Undetermined. Asexual morph:Sporodochia dark,
moderately dense. Conidiophores 60–135 × 4.4–6.7 μm
(x=105×5.4μm, n= 20), simple or branched, straight or
flexuous, pale, olivaceous or golden brown, smooth, septate,
with one or several sympodially arranged conidiogenous
loci. Conidiogenous cells mono- to polytretic, integrated,
terminal, later becoming sympodial. Conidia in vivo: 22–
58 × 8–14(−16) μm(
x=37.5×11.7μm, n=35),simple
or branched chains, obclavate, obpyriform, ovoid or ellip-
soidal, often with a short conical or cylindrical beak, pale
to mid dark brown, with cicatrized black scars at the
beaked end, up to 7 transverse and usually several longi-
tudinal or oblique septa, slightly constricted near some
septa, finely verruculose; in vitro: 13–29.2 × 6.1–
11.8 μm(
x=21.2×8.2μm, n= 30), obclavate,
obpyriform, ovoid or ellipsoidal, sometimes subsphaerical,
pale to dark brown, up to 5 transverse and usually several
longitudinal or oblique septa, slightly constricted near
some septa, finely verruculose.
Culture characteristics: Conidia germinating on PDAwith-
in 18 h and germ tubes produced from one or several septa.
Colonies on PDA reaching 36 mm diam. After 4 days at
25 °C, flat, circular, dense, initially white, becoming light
olivaceous brown, surface smooth, with entire to slightly un-
dulate edge.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte - Forlì, dead branches of Tamarix
gallica L. (Tamari ca ce ae), 28 March 2014, Erio
Camporesi IT1785A (MFLU 14–0600, reference specimen
designated here), living culture MFLUCC 14–0441, ICMP
20706.
Notes:Alternaria tenuissima (Kunze) Wiltshire is a com-
mon pathogen on a wide range of hosts with a worldwide
distribution (Gannibal et al. 2007; Rahman et al. 2002; Luan
et al. 2007). This species belongs to the section Alternata,
which comprises almost 60 species and the molecular varia-
tion within this section is low (Woudenberg et al. 2013). The
type specimen cannot be located hence we designate the cur-
rent specimen as a reference specimen with molecular data
(Ariyawansa et al. 2014b).
Pleospora herbarum (Pers.) Rabenh., Klotzschii Herb. Viv.
Mycol.: no. 547 (1854), Fig. 10
Basionym: Sphaeria herbarum Pers., Syn. meth. Fung.
(Göttingen) 1: 78 (1801)
Saprobic or parasitic on stems and leaves. Sexual morph:
Ascomata 145–230 μmhigh×105–245 μmdiam.
(x=182×198μm, n= 6), scattered to gregarious, immersed
to partially erumpent, black, subglobose, ostiolate. Ostiole
papillate, ostiolar canal filled with hyaline cells. Peridium
30–55 μm wide, broad at sides and thinner at the apex,
usually with two layers, outer layer of heavily pigmented,
thick-walled cells of textura angularis, inner layer com-
posed of hyaline, thin-walled cells of textura angularis.
Hamathecium of 2–3μm wide, cellular, septate, broad,
dense pseudoparaphyses. Asci 100–210 × 23.5–28.2 μm
(x= 145 × 25.7 μm, n= 20), 8-spored, bitunicate,
fissitunicate, cylindrical to clavate, pedicel furcate, apically
rounded, with a minute ocular chamber. Ascospores 26.5–
31 × 11–15 μm(
x=29×13μm, n= 40), uniseriate to
biseriate, ellipsoidal, muriform, brown or pale brown, with a
thick mucilaginous sheath. Asexual morph: Hyphomycetous.
Conidiophores macronematous, mononematous, scattered or
Fungal Diversity
caespitose, unbranched or rarely loosely branched,
straight or flexuous, usually nodose, with a number of
vesicular swellings, pale to mid brown or olivaceous
brown, smooth or in part verruculose. Conidiogenous
cells holoblastic, integrated, terminal, percurrent, at first
clavate or subsphaerical with thin wall at the apex.
Conidia 15–22 × 10.5–15 μm(
x=18×13.7μm,
n= 20), solitary, dry, acrogenous, oblong, rounded at
the ends, ellipsoidal, obclavate or subsphaerical, dark brown
to olivaceous brown, verrucose or echinulate, muriform, often
constricted at one or more of the septa.
Culture characteristics: Ascospores germinating on PDA
within 18 h and germ tubes produced from one or several
cells. Colonies on PDA reaching 30 mm diam. After 4 days
at 25 °C, flat, circular, white, surface smooth, with entire to
slightly undulate edge.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte - Forlì, dead branches of Tama ri x g al lic a
L. (Tamaricaceae), 28 March 2014, Erio Camporesi
IT1785B (MFLU 14–0601), living culture MFLUCC 14–
0442, ICMP 20707.
Fig. 9 Alternaria tenuissima (MFLU 14–0600)aAppearance of sporodochia on host surface b, c, i Conidiophores and developing conidia d-g, j-l
Conidia (i-l from MFLUCC 14–0441) hGerminating conidia. Scale bars: b, c =25μm, i=20μm, j-l =10μm
Fungal Diversity
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Notes:Pleospora herbarum (Stemphylium herbarum)is
the type species of the genus Pleospora and has been reported
on various plant hosts (Ariyawansa et al. 2015a). This species
seems to be a species complex (Ariyawansa et al. 2015a)and
further studies are needed to establish this.
Tama ri cic ol a Thambugala, E. Camporesi & K.D. Hyde,
gen. nov.
Index Fungorum number: IF552087; Facesoffungi
number: FoF 02153
Etymology: From the Latin cola meaningdwellingonand
Tamarici in reference to holotype occurring on Tamarix
species
Saprobic on dead herbaceous stems and branches. Sexual
morph:Ascomata immersed to partially erumpent, solitary or
scattered to gregarious, black, coriaceous, ostiolate. Peridium
comprising several layers, outer layers heavily pigmented,
thick-walled, comprising blackish to dark brown cells of
textura angularis, inner layers composed of lightly
pigmented to hyaline, thin-walled cells of textura angularis.
Hamathecium composed of septate, branched or not, cellu-
lar pseudoparaphyses. Asci (4–)–8 spored, bitunicate,
fissitunicate, cylindrical to cylindric-clavate, pedicellate,
apically rounded, with a distinct ocular chamber.
Ascospores uni- to biseriate, ellipsoidal, muriform, trans-
versely 3-septate, with 1–3 vertical septa, yellowish brown
at maturity, smooth-walled, without a mucilaginous sheath.
Sexual morph: Coelomycetous. Conidiomata pycnidial,
immersed, dark brown to black, subglobose, ostiolate.
Conidiomata comprising few layers of dark brown to hya-
line cells of textura angularis.Conidiophores reduced to
conidiogenous cells. Conidiogenous cells phialidic, hyaline,
smooth, ampulliform. Conidia ellipsoidal or short-cylindri-
cal, hyaline, rounded at both ends, 1-celled, smooth-walled.
Type species:Tamaricicola muriformis Thambugala,
Camporesi & K.D. Hyde
Notes:Tamaricicola is introduced here to accommodate a
novel species from Tama ri x. Molecular phylogenetic analysis
(Fig. 8) shows that Tamaricicola clustered into a distinct clade
in Pleosporaceae.Tamaricicola morphologically resembles
species of Pleosporaceae in having immersed to partially
erumpent ascomata, cylindrical asci, brown or pale brown,
muriform ascospores and a coelomycetous asexual morph
(Ariyawansa et al. 2015a). However, this genus differs from
Pleospora Rabenh. ex Ces. & De Not., the generic type of
Pleosporaceae in having immersed ascomata, transversely 3-
septate ascospores, lacking a mucilaginous sheath and a
Bphoma-like^coelomycetous asexual morph.
Tamaricicola muriformis Thambugala, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552088; Facesoffungi
number: FoF 02,154; Figs.11 and 12
Etymology: Named after its muriform ascospores
Holotype:MFLU16–0676
Saprobic on dead herbaceous stems and branches. Sexual
morph:Ascomata 140–200 μm high × 175–300 μmdiam.
(x=175 × 230 μm, n= 6), immersed to partially erumpent,
solitary or scattered to gregarious, black, coriaceous, ostiolate.
Peridium 20–40 μm wide, comprising several layers, outer
layers heavily pigmented, thick-walled, comprising blackish
to dark brown cells of textura angularis, inner layers com-
posed of lightly pigmented to hyaline, thin-walled cells of
textura angularis.Hamathecium composed of 1–2.5 μm
wide, numerous, septate, branched or not, cellular
pseudoparaphyses. Asci 70–105 × 12–15(−16.7) μm
(x= 86 × 13.6 μm, n=24),(4–)8 spored, bitunicate,
fissitunicate, cylindrical to cylindric-clavate, pedicellate, api-
cally rounded, with a distinct ocular chamber. Ascospores 13–
19 × 5.5–9μm(
x= 15.8 × 7.3 μm, n= 40), partially overlap-
ping, uni- to biseriate, ellipsoidal, muriform, transversely 3-
septate, with 1–3 vertical septa, upper part wider, constricted
at the central septum, initially hyaline, becoming yellowish
brown at maturity, smooth-walled, without a mucilaginous
sheath. Sexual morph: coelomycetous. Conidiomata 90–
160 μmhigh×90–165 μmdiam.(
x= 106 × 124 μm,
n= 10), pycnidial, immersed, dark brown to black,
subglobose, ostiolate. Conidiomatal wall 10–21 μm wide,
comprising few layers of dark brown to hyaline cells
of textura angularis.Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 3–4.5 × 1.8–
3.8 μm(
x= 3.7 × 2.9 μm, n= 10), phialidic, hyaline,
smooth, ampulliform. Conidia 2–4.5 × 1.2–2.3 μm(
x=
3.1 × 1.7 μm, n= 60), ellipsoidal or short-cylindrical,
hyaline, rounded at both ends, 1-celled, smooth-walled.
Culture characteristics: Ascospores germinating on PDA
within 24 h and germ tubes produced from one end or both
ends. Colonies growing on PDA reaching 25 mm diam. After
12 days at 25 °C, circular, umbonate, dense, surface iron-gray,
white at the margin, reverse black, olivaceous brown at the
margin, smooth surface with edge entire to curled.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte- Forlì, on dead branches of Tamarix
Fig. 10 Pleospora herbarum (MFLU 14–0601 and MFLUCC 14–
0442)aAppearance of ascomata on host surface bVertical section
through ascoma cPeridium d, e Mature and immature asci f-h
Ascospores surrounded by thick mucilaginous sheaths iGerminating
ascospore jConidiophores and developing conidia k, l Conidia. Scale
bars: b=100μm, c=25μm, d, e =50μm, f-j =20μm, k-l =10μm
Fungal Diversity
gallica L. (Tamaricaceae), 10 January 2014, Erio Camporesi
IT 917–3 (MFLU 16–0676, holotype), ex-type living culture
MFLUCC 15–0488, ICMP 21254; ibid. (HHUF 30463,
isotype); ibid., 21 January 2015, Erio Camporesi IT 917–4
(MFLU 16–0677), living culture MFLUCC 15–0489, ICMP,
ibid., 22 November 2012, Erio Camporesi IT 917 (MFLU 14–
0582, paratype), living culture MFLUCC 13–0071, ICMP
20690; TALY, ibid., 23 November 2014, Erio Camporesi IT
Fig. 11 Tamaricicola muriformis (holotype). a, b Appearance of
ascomata on host surface c, d Vertical sections through ascomata e
Peridium fPseudoparaphyses gImmature ascus h, i Mature bitunicate
asci jApex of ascus k-m Ascospores. Scale bars: c= 100 μm, d-
e=50μm, f=20μm, g-i=25μm, j=10μm, k-m =5μm
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917–5 (MFLU 16–0678), living culture MFLUCC 15–0490,
ICMP, ibid., 22 November 2012, Erio Camporesi IT 917–2
(MFLU 14–0583), living culture MFLUCC 16–0488.
Pleosporales, genera incertae sedis
Paraepicoccum Matsush., Matsush. Mycol. Mem. 7: 59 (1993)
Facesoffungi number: FoF 02,155
The genus Paraepicoccum was introduced by Matsushima
(1993)toaccommodateP. amazonense and is presently placed
in Pezizomycotina, genera incertae sedis (Index Fungorum
2016). Paraepicoccum amazonense grows on leaf litter and
aquatic plants (Orłowska et al. 2004; Seifert et al. 2011). We
found P.amazonense associated with dead branches of
Tamari x species and phylogenetic analysis (Fig. 16)of
sequence data revealed the placement of this species in
Pleosporineae,Pleosporales anditformedaseparate
clade in Camarosporium sensu stricto, however only
coelomycetous asexual morphs have been reported in
Camarosporium sensu stricto (Wijayawardene et al.
2014c). Therefore, tentatively we retain Paraepicoccum
in Pleosporales,generaincertae sedis. We designate our
collection as epitype (sensu Ariyawansa et al. 2014b)
because the holotype of Paraepicoccum amazonense is
not available for study.
Typ e species:Paraepicoccum amazonense Matsush.,
Matsush. Mycol. Mem. 7: 59 (1993)
Fig. 12 Tamaricicola muriformis (asexual morph)a, b Appearance of
conidiomata on host surface c, g Vertical section through conidiomata d,
e, h Conidiomatal wall, conidiogenous cells and developing conidia f, i
Conidia (g-i from MFLUCC 15–0488). Scale bars: c, d =50μm,
e=15μm, f=10μm
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Paraepicoccum amazonense Matsush., Matsush. Mycol.
Mem. 7: 59 (1993)
Index Fungorum number: IF360894; Facesoffungi
number: FoF 02,156; Fig.13
Saprobic on dead leaves or branches. Sexual morph:
Undetermined. Asexual morph:Sporodochia dark, dense.
Conidiophores up to 12 μmlong,4–5μm diam., simple
or branched, pale, olivaceous or golden brown, smooth, septate,
with one or several apical conidiogenous loci. Conidiogenous
cells monoblastic, pale brown. Conidia in vivo: 9.5–17.4 × 9.6–
17.2 μm(
x=13×12.4μm, n= 35), ellipsoidal, obclavate or
subsphaerical, yellowish brown and 0–1 septate when young,
becoming brown, muriform with several longitudinal or oblique
septa when mature, finely guttulate; in vitro: 12.5–25 × 12–
22.5 μm(
x= 18.5 × 11.7 μm, n= 35), yellowish to golden
brown.
Culture characteristics: Conidia germinating on PDAwithin
24 h and germ tubes produced from one or two cells. Colonies
growing on PDA reaching 25 mm diam. After 21 days at 25 °C,
circular to irregular, umbonate, dense, surface initially white,
becoming dark olivaceous brown, after 2 weeks, reverse light
golden brown, smooth surface with edge entire to curled.
Material examined: ITALY, Province of Ravenna [RA],
Lido di Dante, on dead branches of Tama rix sp.
(Tamari cacea e), 7 December 2014, Erio Camporesi IT
2287–3(MFLU16–0672, epitype designated here), living
culture MFLUCC 15–0493, ICMP 21255; ibid. IT 2287–5
(MFLU 16–0673), living culture MFLUCC 15–0491
Didymellaceae Gruyter, Aveskamp & Verkley, Mycol. Res.
113(4): 516 (2009)
The family Didymellaceae was introduced to accom-
modate Didymella and Phoma and phoma-like genera
(de Gruyter et al. 2009). The family encompasses plant
pathogenic, saprobic and endophytic species associated
with a wide range of hosts and substrates (de Gruyter
et al. 2009;Hydeetal.2013;Chenetal.2015;Liu
et al. 2015). Chen et al. (2015) revised the family based
on multi-gene (ITS, LSU, RPB2 and tub2) phylogenetic
analysis and morphological observations, while introducing
nine new genera and a new family Microsphaeropsidaceae
to accommodate Microsphaeropsis, which is morphologically
distinct from the members of Didymellaceae.
DNA phylogeny
Reference sequences used in the phylogenetic analysis
were selected from strains of Didymellaceae and
Microsphaeropsidaceae (Chen et al. 2015). The present
phylogeny based on LSU, ITS, RPB2 and β-tubulin
sequence data revealed two major clades representing
the families Didymellaceae and Microsphaeropsidaceae,
while 18 subclades represent 18 distinct genera in
Didymellaceae. Most of these sub-clades received moderate
to high bootstrap support (BS) in the analyses. The tree is
rooted to Leptosphaeria doliolum (strain CBS 505.75).
Newly collected microsphaeropsis-like species in this study
appear distinguishable from Microsphaeropsidaceae
(Fig. 14) and these strains clustered in Didymellaceae as a
distinct group between the Phoma and Calophoma clades,
together with the ex-type strain of Phoma tamaricicola
Wanas., Camporesi, E.B.G. Jones & K.D. Hyde (MFLUCC
14–0602). Another three species can be distinguished in this
clade and have moderate to high bootstrap support.
Neomicrosphaeropsis Thambugala, Camporesi & K.D.
Hyde, gen. nov.
Index Fungorum number: IF552089; Facesoffungi
number: FoF02157
Etymology: The generic epithet, neo (Lat., new), refers to
the similarity to Microsphaeropsis
Saprobic or weak pathogens on branches of Tam arix spe-
cies. Sexual morph:Ascomata solitary, scattered, immersed,
slightly erumpent, dark brown to black, ostiolate. Peridium
comprising 6–8 layers, outer layers heavily pigmented,
thick-walled, comprising blackish to dark brown cells of
textura angularis, inner layers composed of lightly pigmented
to hyaline, thin-walled cells of textura angularis.
Hamathecium comprising numerous, filamentous, branched,
septate, pseudoparaphyses. Asci 8-spored, bitunicate,
fissitunicate, cylindrical to cylindric-clavate, pedicellate,
apically rounded, with an ocular chamber. Ascospores
1–2-seriate, partially overlapping, muriform, ellipsoidal,
4–6 transversely septate, with 3–4 vertical septa, con-
stricted at the central septum, yellowish brown, conical
and narrowly rounded at the ends, smooth-walled, with-
out a mucilaginous sheath (description modified from
Crous et al. 2014). Asexual morph:Conidiomata pyc-
nidial, scattered or solitary, immersed, slightly erumpent,
black, globose to subglobose, uni- to multi-loculate,
ostiolate. Conidiomatal wall consisting of light to dark
brown, thick-walled cells of textura angularis,becoming
hyaline towards the conidiogenous region. Conidiogenous
cells enteroblastic, phialidic, hyaline, cylindrical, discrete or
integrated, smooth. Conidia hyaline to light brown, aseptate,
obovoid to ellipsoidal, smooth-walled.
Fig. 13 Paraepicoccum amazonense (MFLU 16–0672, epitype)a, b
Appearance of sporodochia on host surface c, h, i Conidiophores and
developing conidia d, e, j Conidia fGerminating conidia gColonies on
PDA (h-j from MFLUCC 15–0493). Scale bars: c-f, j =15μm, h, i
=10μm
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Fungal Diversity
Notes:Microsphaeropsis was established by Von Höhnel
(1917), and placed in the family Montagnulaceae.
Microsphaeropsis is morphologically similar to some species
in the genus Coniothyrium and is characterized by phialidic
conidiogenous cells with a periclinal thickening, and pale
greenish brown, 0–1 septate conidia (De Gruyter et al. 2013;
Chen et al. 2015). Barr (1987) classified Microsphaeropsis as
an asexual morph of Phaeosphaeriaceae, while some
Microsphaeropsis species have been reported in
Montagnulaceae (= Didymosphaeriaceae) (Someya et al.
1997; Zhang et al. 2012; Verkley et al. 2014). Recent studies
(De Gruyter et al. 2009,2013;Aveskampetal.2010; Hyde
et al. 2013) however, treated this genus in Didymellaceae based
on the sequences of the type species M.olivacea (Bonord.)
Höhn. Chen et al. (2015) showed in their phylogenetic analysis
Fig. 15 Neomicrosphaeropsis italica (holotype)aAppearance of conidiomata on host surface b, c Vertical sections through conidiomata d
Conidiogenous cells and developing conidia econidia fCulture growing on PDA. Scale bars: b=100μm, c=50μm, d,e=25μm
Fig. 14 Phylogram resulting from Maximum likelihood (RAxML) anal-
ysis of the combined LSU, ITS, RPB2 and β-tubulin sequences of 82
strains representing Didymellaceae and Microsphaeropsidaceae.
Maximum likelihood bootstrap support values equal or greater than
50 % are indicated above or below the nodes. The new isolates are in
blue. The tree is rooted to Leptosphaeria doliolum
Fungal Diversity
basedonLSU,ITS,β-tubulin and RPB2 sequence data,
that Microsphaeropsis grouped basal to the Didymellaceae
and introduced a new family Microsphaeropsidaceae con-
sidering both morphological traits and phylogeny.
Neomicrosphaeropsis represents a species complex which
includes morphologically similar but phylogenetically different
species. Recollecting and sequencing of more coniothyrium-
like species would be helpful to classify their placement in
the genera Microsphaeropsis and Neomicrosphaeropsis.
Type species:Neomicrosphaeropsis italica Thambugala,
Camporesi & K.D. Hyde
Neomicrosphaeropsis italica Thambugala, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552090; Facesoffungi
number: FoF02158; Fig. 15
Etymology: Named after the country (Italy), where the ho-
lotype was collected.
Holotype:MFLU16–0674
Saprobic or weak pathogen on twigs of Tamari x species.
Sexual morph: Undetermined. Asexual morph:
Conidiomata 95–200 μm high × 200–270 μm diam. (x=
142 × 228 μm, n= 8), pycnidial, scattered or solitary, im-
mersed, slightly erumpent, black, globose to subglobose,
uni- to multi-loculate, ostiolate. Conidiomatal wall 18–
40 μm(
x=28μm, n= 15), consisting of light to dark brown,
thick-walled cells of textura angularis, becoming hyaline to-
wards the conidiogenous region. Conidiogenous cells 2.5–
5(−6.4) × 1.8–3.1(−4) μm(
x=4.3×2.5μm, n=15),
enteroblastic, phialidic, hyaline, cylindrical, discrete or
integrated, smooth. Conidia 3.6–6.2 × 2.9–4.6 μm
(x=5.1×3.8μm, n= 60), hyaline to light brown,
aseptate, obovoid to ellipsoidal, smooth-walled.
Culture characteristics: Conidia germinating on PDAwith-
in 18 h. Colonies growing on PDA reaching 14 mm diam.
After 7 days at 25 °C, circular, flat, surface pinkish white,
surface smooth with entire to curled edge.
Material examined: ITALY, Province of Ravenna [RA],
Lido di Dante, on dead branches of Ta ma rix sp.
(Tamari cacea e), 7 December 2014, Erio Camporesi IT
2286–2 (MFLU 16–0674, holotype), ex-type living culture
MFLUCC 15–0485, ICMP 21253; ibid. (GZAAS, isotype);
ibid. 24 December 2014, Erio Camporesi IT 2286 MFLUCC
15–0484; ibid. IT 2287–2(MFLU16–0663, paratype), living
culture MFLUCC 15–0487; Province of Ravenna [RA], Lido
di Dante, on dead branches of Tamari x sp. (Tamaricaceae), 19
March 2015, Erio Camporesi IT 2421–1(MFLU16–0675,
GZAAS), living culture MFLUCC 16–0284, GZCC 15–0049.
Fig. 16 Neomicrosphaeropsis novorossica (holotype)aAppearance of conidiomata on host surface b, c Vertical sections through conidiomata d
Conidiogenous cell and developing conidium econidia. Scale bars: b,c=50μm, d=5μm, e=20μm
Fungal Diversity
Neomicrosphaeropsis novorossica Thambugala, Bulgakov &
K.D. Hyde, sp.nov.Fig. 16
Index Fungorum Number: IF552091; Facesoffungi
number: FoF02159
Etymology: Named after the historical name of region
«Novorossiya» (in Russia), from which the host plant species
had been introduced.
Holotype:MFLU14–0603
Fig. 17 Neomicrosphaeropsis rossica (MFLU 14–0605,holotype)aAppearance of conidiomata on host surface b, c Vertical sections through
conidiomata d, e Conidiogenous cells and developing conidia fConidia gCulture growing on PDA. Scale bars: b,c=50μm, d=20μm, e,f=10μm
Fungal Diversity
Saprobic or weak pathogen on twigs and branches of
Tamari x species. Sexual morph: Undetermined. Asexual
morph:Conidiomata 60–160 μmdiam.×55–130 μmhigh
(x=123×107μm, n= 10), pycnidial, gregarious, scattered or
solitary, immersed slightly erumpent, black, globose to
subglobose, uniloculate, ostiolate. Conidiomatal wall 13–
34 μm(
x=21μm, n= 15), consisting of 3–4layersoflight
brown cells of textura angularis, becoming hyaline towards the
conidiogenous region. Conidiogenous cells 2.6–4.2(−5.8) × 2–
3.4(−4.8) μm(
x=3.7×3μm, n= 15), enteroblastic, phialidic,
hyaline, cylindrical, discrete or integrated, smooth. Conidia
4.3–7.5 × 3.6–5.1 μm(
x= 5.8 × 4.4 μm, n= 40), hyaline when
immature, becoming light brown, aseptate, ellipsoidal, obovoid
or globose, smooth-walled.
Culture characteristics: Conidia germinating on PDAwith-
in 18 h. Colonies growing on PDA reaching 22 mm diam.
after 10 days at 25 °C, circular, flat, moderately dense, surface
olivaceous brown, smooth surface with entire edge.
Material examined: RUSSIA, Rostov Region, Shakhty
City, Central Park, on dead branches of Tamarix ramosissima
Ledeb. (Tamaricaceae), 21 May 2014, Timur Bulgakov 74
(MFLU 14–0603, holotype), ex-type living culture
MFLUCC 14–0578, ICMP 20751; ibid. (GZAAS, isotype).
Neomicrosphaeropsis rossica Thambugala, Bulgakov &
K.D. Hyde, sp.nov.Fig. 17
Index Fungorum number: IF552092; Facesoffungi
number: FoF0 2160
Etymology: Named after the country (Russia), where the
holotype was collected.
Holotype:MFLU14–0605
Saprobic or weak pathogen on twigs of Ta ma rix
ramosissima Ledeb. Sexual morph: Undetermined. Asexual
morph:Conidiomata 85–150 μmhigh×90–150 μmdiam.
(x= 122 × 121 μm, n= 10), pycnidial, gregarious,
scattered or solitary, immersed slightly erumpent, black,
globose to subglobose, uniloculate, ostiolate. Conidiomatal
wall 15–30 μm wide, consisting of 3–5layersofbrown
cells of textura angularis, becoming hyaline towards the
conidiogenous region. Conidiogenous cells (2–)3–4.6 × 2–
4μm(
x=3.8×3μm, n= 15), enteroblastic, phialidic,
hyaline, cylindrical, discrete or integrated, smooth. Conidia
Fig. 18 Neomicrosphaeropsis tamaricicola (MFLU 14–0602, asexual morph)aConidiomata on host surface b, c Vertical sections through
conidiomata d, e Conidiogenous cells and developing conidia fconidia. Scale bars: b=50μm, c=30μm, d-e =5μm, f=10μm
Fig. 19 Phylogram resulting from maximum likelihood (RAxML)
analysis of combined ITS and LSU sequence data of selected strains in
Didymosphaeriaceae. Maximum likelihood bootstrap values equal or
greater than 50 % are indicated above or below the nodes. The new
isolates are in blue. The tree is rooted to Stagonospora paludosa
Fungal Diversity
Fungal Diversity
4.4–5.7(−6.3) × 2.9–3.9 μm(x=5.3×3.4μm, n=35),
hyaline when immature, becoming light brown, aseptate,
ellipsoidal, obovoid or globose, smooth-walled.
Culture characteristics: Conidia germinating on PDAwith-
in 18 h. Colonies growing on PDA reaching 30 mm diam.
After 10 days at 25 °C, circular, flat, moderately dense, surface
dark olivaceous brown, smooth surface with entire to slightly
undulate edge.
Material examined: RUSSIA, Rostov Region, Oktyabrsky
District, Persianovsky Township, arboretum of Don State
Agrarian University, Tamarix ramosissima Ledeb.
(Tamaricaceae), 4 June 2014, Timur Bulgakov 79 (MFLU
14–0605, holotype), ex-type living culture MFLUCC 14–
0586, ICMP 20753.
Notes:Neomicrosphaeropsis rossica is introduced as a
new species and this species mainly differs from N.italica,
the type species of the genus, in having uniloculate
conidiomata which are smaller and recorded from a different
host species in a different country (Russia). In the present
phylogenetic analysis of combined LSU, ITS, RPB2 and β-
tubulin sequence data, N. rossica formed a separate subclade
in the Neomicrosphaeropsis clade in Didymellaceae with
high BS support (96 %). Neomicrosphaeropsis rossica
shares similar morphology with Coniothyrium tamaricis,
but the conidial dimensions are slightly different and the
taxa are recorded from different hosts (Tam ari x gal li ca L.)
and continents (Saccardo 1902). Therefore, C.tamaricis can
be another species that may represent Neomicrosphaeropsis
and needs to be recollected from the same location and
analyzed with gene sequence data to confirm its placement.
Fig. 20 Pseudocamarosporium propinquum (MFLU 16–0669)a, b
Appearance of conidiomata on the host c, d Vertical sections through
conidiomata e, f Conidiogenous cells and developing conidia g, h
Mature and immature conidia i-j Germinating conidia kColony after
2 weeks growth on PDA at 25 °C. Scale bars: c=100μm, d=50μm,
e-h =10μm, i-j =20μm
Fungal Diversity
We observed the type material of Coniothyrium tamaricis
Henn., which is an illegitimate name, and this taxon has
larger conidia (5–8×3.5–4.5 μm) and was reported on
Tamarix gallica in Germany (Kabát and Bubák 1907).
Therefore, re-collection and molecular data analyses
are essential to identify whether this species is distinct.
Fig. 21 Paracamarosporium tamaricis (holotype)aAppearance of
conidiomata on host surface b, c Vertical section through conidiomata d
Conidiomatal wall e-f Conidiogenesis and developing conidia g-h
Conidia iGerminating conidium. Scale bars: b, c =100μm,
d=25μm, e=20μm, f=5μm, g=15μm, h- i =10μm
Fungal Diversity
Neomicrosphaeropsis tamaricicola (Wanas., Camporesi,
E.B.G. Jones & K.D. Hyde) Thambugala, Wanasinghe &
K.D. Hyde, comb.nov.Fig. 18
Basionym: Phoma tamaricicola Wanas., Camporesi,
E.B.G. Jones & K.D. Hyde, in Crous et al., Persoonia,
Mol. Phyl. Evol. Fungi 33: 281 (2014)
Index Fungorum Number: IF552093; Facesoffungi
number: FoF02161
Saprobic or weak pathogen on twigs and branches of
Tamari x species. Sexual morph:Ascomata solitary, scattered,
immersed, slightly erumpent, dark brown to black, ostiolate.
Peridium comprising 6–8 layers, outer layers heavily
pigmented, thick-walled, comprising blackish to dark
brown cells of textura angularis, inner layers composed
of lightly pigmented to hyaline thin-walled cells of textura
angularis.Hamathecium comprising numerous, filamen-
tous, branched, septate pseudoparaphyses. Asci 8-spored,
bitunicate, fissitunicate, cylindrical to cylindric-clavate,
pedicellate, apically rounded, with a minute ocular chamber.
Ascospores 1–2-seriate, partially overlapping, muriform, el-
lipsoidal, 4–6 transversely septate, with 3–4verticalsepta,
constricted at the central septum, initially hyaline, becom-
ing yellowish brown at maturity, conical and narrowly
rounded at the ends, smooth-walled, without a mucilagi-
nous sheath (description modified from Crous et al. 2014).
Asexual morph:Conidiomata 65–155 μmdiam.×75–
120 μmhigh(x
̅=115×97μm, n= 6), scattered to
gregarious, immersed to erumpent, black, subglobose,
uniloculate, ostiolate. Conidiomatal wall 12–22 μmcom-
posed of 3–5 layers of dark brown, thick-walled cells of
textura angularis, fusing at the outside with the host tissues.
Conidiogenous cells 2–4 × 1.6–3.2 μm(x
̅=3.1×2.2μm,
n=15), enteroblastic, phialidic, hyaline, cylindrical, discrete
or integrated, smooth. Conidia 3.5–6.6 × 2.5–3.4 μm
(x
̅=5.3×2.9μm, n= 40), ellipsoidal or obovoid, hyaline
to light brown, straight, rounded at both ends, 1-celled,
smooth-walled.
Culture characteristics: Conidia germinating on PDAwith-
in 18 h. Colonies growing on PDA reaching 30 mm diam.
After 10 days at 25 °C, circular, flat, moderately dense, surface
grey, olivaceous brown to dull green from below, surface
smooth with entire to slightly undulate edge.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte - Forlì, on dead branches of Tamarix
gallica L. (Tamaricaceae), 15 January 2014, Erio Camporesi
(MFLU 14–0333, holotype); ibid. 28 March 2014, Erio
Camporesi IT 1785C (MFLU 14–0602), living culture
MFLUCC 14–0443, ICMP 20708; ibid. 22 November 2012
Erio Camporesi IT 918–1(MFLU14–0591), living culture
MFLUCC 14–0439, ICMP 20743.
Didymosphaeriaceae Munk, Dansk bot. Ark. 15(no. 2): 128
(1953)
The family Didymosphaeriaceae is characterized by im-
mersed ascomata surrounded by hyphae forming an apical
clypeus, 2–8-spored, bitunicate, fissitunicate asci and hya-
line to pigmented, phragmosporous or muriform asco-
spores, with coelomycetous or hyphomycetous asexual
morphs (Hyde et al. 2013; Ariyawansa et al. 2014a;Liu
et al. 2015;Lietal.2016). Ariyawansa et al. (2014a)
accepted 16 genera in Didymosphaeriaceae and subsequently
Austropleospora,Paracamarosporium,Pseudocamarosporium,
Pseudotrichia and Spegazzinia were included (Thambugala
et al. 2014b; Ariyawansa et al. 2015b; Wijayawardene
et al. 2014b).
DNA phylogeny
The present phylogenetic analysis based on ITS and LSU
sequence data and reference sequences used in the
phylogenetic analyses were selected based on the data
from Wijayawardene et al. (2014b)a
ndCrousetal.
(2015). The tree is rooted to Stagonospora paludosa
(CBS 135088). The strain MFLUCC 14–0166 grouped
together with the ex-type strain of Pseudocamarosporium
propinquum, while strains MFLUCC 15–0494 and
MFLUCC 15–0495 clustered as a sister clade to
Paracamarosporium hawaiiense (CBS 120025) (Fig. 19).
Pseudocamarosporium Wijayaw. & K.D. Hyde, in
Wijayawardene et al., Cryptog. Mycol. 35(2): 185 (2014)
Pseudocamarosporium was established by Wijayawardene
et al. (2014b) for some camarosporium-like species grouping
in Didymosphaeriaceae (= Montagnulaceae) and typi-
fied by Pseudocamarosporium propinquum. Later,
Camarosporium brabeji Marincowitz,M.J.Wingf.&
Crous, Paraconiothyrium africanum Damm, Verkley &
Crous and Pseudocamarosporium pini (Westend.)
Phukhamsakda, Camporesi & K.D. Hyde were included
in the genus as new combinations based on their phy-
logenetic affinities (Crous et al. 2015;Lietal.2016).
We collected P.propinquum associated with dead
branches of Tamari x galli ca in Italy and it perfectly
matches with the epitype description (Wijayawardene
et al. 2014b) and clustered together with the ex-type
Fig. 22 Phylogram resulting from maximum likelihood (RAxML)
analysis of combined ITS and LSU sequence data for taxa of
Lentitheciaceae. Maximum likelihood bootstrap values equal or greater
than 50 % are indicated above or below the nodes. The new isolate is in
blue. The tree is rooted with Macrodiplodiopsis desmazieri
Fungal Diversity
Fungal Diversity
Fungal Diversity
strain (Fig. 19). This is the first record of P.propinquum on
Tamari x species.
Type species:Pseudocamarosporium propinquum (Sacc.)
Wijayaw., Camporesi & K.D. Hyde, in Wijayawardene, et al.
Cryptog. Mycol. 35(2): 191 (2014).
Pseudocamarosporium propinquum (Sacc.) Wijayaw.,
Camporesi & K.D. Hyde, in Wijayawardene, et al. Cryptog.
Mycol. 35(2): 191 (2014)
Facesoffungi number: FoF02162; Fig. 20
Saprobic on Tamarix gallica L. Sexual morph:
Undetermined. Asexual morph:Conidiomata pycnidial,
200–400 μmdiam.×120–200 μmhigh(
x=323×168μm,
n= 6), solitary, scattered, black, immersed, unilocular,
subglobose or irregular in shape, frequently associated
with other fungi, ostiolate. Pycnidial wall 14–25 μm
(x=18.9μm, n= 12), comprising 3–4 layers of hya-
line to light brown cells of textura angularis and indis-
tinguishable from the host tissues. Conidiophores re-
duced to conidiogenous cells. Conidiogenous cells 4–
6×5–8μm(
x= 12.6 × 7 μm, n=30),withpercurrent
phialidic development, smooth, short, hyaline, formed
from the inner layer of the pycnidial wall. Conidia
10–15 × 5.5–8μm(
x= 12.6 × 6.5 μm, n=30),ob-
long, straight, initially hyaline, aseptate, becoming
muriform, with 3 transverse septa and 1–2 longitudinal
septa, pale brown to dark brown, smooth-walled.
Culture characteristics: Conidia germinating on PDAwith-
in 24 h and germ tubes produced from one or several septa.
Colonies growing on PDA reaching 47 mm diam. After
14 days at 25 °C, circular, flat, moderately dense, surface
greenish olivaceous, white at the margin, becoming dark olive
Fig. 23 Keissleriella tamaricicola (holotype-sexual morph) a, b
Appearance of ascomata on the host surface c, d Vertical sections
through ascomata ePeridium fPseudoparaphyses g-j Mature and
immature bitunicate asci k-n Ascospores. Scale bars: c, d =100μm,
e=50μm, f=10μm, g-j =25μm, k- n =5μm
Fig. 24 Keissleriella tamaricicola (asexual morph from ex-type living culture) a Appearance of conidioma on PDA bVertical section through
conidioma cConidiomatal wall dConidiophores and conidiogenous cells producing conidia eConidia. Scale bars: b=100μm, c-e =10μm
Fungal Diversity
to greyish after 2 weeks, reverse black, smooth surface with
entire to slightly undulate edge.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte - Forlì, Tamarix gallica L.
(Tamaricaceae), 10 January 2014, Erio Camporesi IT 917-
CD100 (MFLU 16–0669), living culture MFLUCC 14–
0166, ICMP 20709.
Paracamarosporium Wijayaw. & K.D. Hyde, in
Wijayawardene et al., Cryptog. Mycol. 35(2): 183 (2014)
Paracamarosporium was introduced by Wijayawardene
et al. (2014b) to accommodate Camarosporium psoraleae
Crous & M.J. Wingf. and subsequently, Crous et al. (2015)
added Microdiplodia hawaiiensis Crous (Paracamarosporium
hawaiiense (Crous) Crous), Camarosporium leucadendri
Marinc. et al. (Paracamarosporium leucadendri (Marinc.
et al.) Crous) and Paracamarosporium fagi Crous & R.K.
Schumach. to the genus. However, it is difficult to morpholog-
ically distinguish the taxa in this genus as both camarosporium
and coniothyrium-like species grouped in this genus (Crous
et al. 2015).
Paracamarosporium tamaricis Thambugala, Camporesi &
K.D. Hyde, sp. nov.
Index Fungorum number: IF552094, Facesoffungi
number: FoF02163, Fig. 21
Etymology: The species epithet Btamaricis^refers to the
host genus Tam ar ix on which the holotype was collected.
Holotype:MFLU16–0661
Saprobic on Tamarix gallica L. Sexual morph:
Undetermined. Asexual morph:Conidiomata 280–350 μm
diam. × 160–320 μmhigh(
x=311×237μm, n= 6), pyc-
nidial, single to solitary, or scattered, immersed, brown to dark
brown, globose to subglobose, ostiolate. Conidiomatal wall
15–35 μm wide, comprising several layers of light brown cells
of textura angularis, becoming hyaline towards the
conidiogenous region. Conidiophores reduced to
conidiogenous cells. Conidiogenous cells 3–9.2 × 2.2–
4.4 μm(
x=6.5×3.2μm, n= 15), hyaline, smooth, holoblas-
tic forming conidia at their tips. Conidia 8.9–12.1 × 4.5–
6.4 μm(
x= 10.2 × 5.2 μm, n= 40), initially hyaline, becom-
ing pale brown to brown, oblong to sub-cylindrical, with
rounded ends, 1-septate, smooth-walled.
Culture characteristics: Conidia germinating on PDAwith-
in 24 h and germ tubes produced from one or both ends.
Colonies growing on PDA 15 mm diam. After 14 days at
25 °C, circular, flat, moderately dense, surface white, becom-
ing pale brown when aged, smooth surface, with entire to
slightly undulate edge.
Material examined: ITALY, Province of Forlì-Cesena,
Converselle - Castrocaro Terme e Terra del Sole, dead
branches of Tamarix gallica L. (Tamaricaceae), 24
December 2014, Erio Camporesi IT 933 (MFLU 16–0661,
holotype), ex-type living culture MFLUCC 15–0494,
ICMP; ibid. (MFLU 16–0662, isotype); ibid., 22 January
2015, Erio Camporesi IT 933–2 (MFLU 16–0662, paratype),
living culture MFLUCC 15–0495.
Notes:Paracamarosporium tamaricis is introduced to
accommodate a new paraconiothyrium-like species,
based on phylogeny and new taxonomic concepts of
Paracamarosporium (Crous et al. 2015). In our phylo-
genetic analysis, P.tamaricis (strains MFLU 16–0661
and 16–0662) clustered in a separate clade in
Paracamarosporium and is phylogenetically distinct
from Paracamarosporium hawaiiense and P.fagi
(Fig. 19), the other paraconiothyrium-like species in
Paracamarosporium. This is the first report of
Paracamarosporium species on Tam ar ix species.
Lentitheciaceae Yin. Zhang, C.L. Schoch, J. Fourn., Crous &
K.D. Hyde, Studies in Mycology 64: 93
The family Lentitheciaceae was established by Zhang
et al. (2009) and members of this family occur on stems
and twigs of herbaceous and woody plants in terrestrial
or aquatic habitats as endophytes and saprobes (Hyde
et al. 2013; Wanasinghe et al. 2014;Knappetal.
2015;Ariyawansaetal.2015b;Singtripopetal.2015;
Tanaka et al. 2015). Hyde et al. (2013)acceptedfour
genera, while Tanaka et al. (2015) included ten genera
Darksidea,Katumotoa,Keissleriella,Lentithecium,
Murilentithecium,Neoophiosphaerella,
Phragmocamarosporium,Poaceascoma,Setoseptoria
and Tingoldiago in Lentitheciaceae.
DNA phylogeny
Phylogenetic analysis is based on combined ITS and LSU
sequence data and reference strains of the family
Lentitheciaceae were selected from Ariyawansa et al.
(2015b); Singtripop et al. (2015) and Li et al. (2016). The tree
is rooted with Macrodiplodiopsis desmazieri (CBS 140062).
The analysis separates the 10 genera currently recognized in
this genus (Fig. 22) and our strain (MFLUCC 14–0168)
Fig. 25 Phylogram resulting from maximum likelihood (RAxML)
analysis of LSU sequences alignment showing the placement of
Homortomyces and Asterodiscus (Stigmatodiscales). Maximum
likelihood bootstrap support values equal or greater than 50 % are
indicated above or below the nodes. The new isolates are in blue. The
tree is rooted to Megalotremis verrucosa
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Fungal Diversity
clustered together with other Keissleriella species in
Lentitheciaceae, close to K.linearis E. Müll. ex Dennis
(IFRD 2008).
Keissleriella Höhn., Sber. Akad. Wiss. Wien, Math.-naturw.
Kl., Abt. 1128: 582 (1919)
Keissleriella was introduced by Von Höhnel (1919)andis
characterized by ascomata with an ostiolar neck covered by
short dark setae (Liu et al. 2015; Ariyawansa et al. 2015b;
Singtripop et al. 2015). Although sequence data for the type
species K.aesculi (Höhn.) Höhn. are unavailable, many au-
thors accept Keissleriella in Lentitheciaceae (Zhang et al.
2009;Hydeetal.2013;Ariyawansaetal.2015b;Singtripop
et al. 2015), based on available sequence data and morpholo-
gy. Our specimen (MFLU14–0597) shows typical character-
istics of Keissleriella, such as short brown setae around the
apex of ascomatal ostiole and cylindro-clavate asci with hya-
line, 1-septate ascospores (Barr 1990; Zhang et al. 2012), but
differs from other Keissleriella species in having immersed
ascomata under a blackened clypeus. Hence, we introduce a
new species K.tamaricicola considering both morphology
and phylogeny.
Keissleriella tamaricicola Thambugala, Camporesi & K.D.
Hyde, sp.nov.
Index Fungorum number: IF552095, Facesoffungi
number: FoF02164, Figs. 23 and 24
Etymology: The species epithet Btamaricicola^refers to the
host genus Tam ar ix on which the holotype occurs.
Holotype:MFLU14–0597
Saprobic on branches of Tam ar ix species. Sexual morph:
Ascomata 160–300 μmd
iam.×130–240 μmhigh(x=
210 × 167 μm, n= 7), solitary or in small groups, scattered,
immersed under a blackened clypeus, globose or subglobose,
dark brown to black, coriaceous, fused to the host tis-
sue, ostiolate. Ostiole papillate, slightly protruding, with
apical dark brown, short setae. Peridium 8–24 μmwide,
composed with dark brown to black, cells of textura
angularis, cells towards the inside lighter and somewhat
flattened, at the outside, darker, fusing with host tissues.
Hamathecium comprising 1–2.5 μm wide, branched,
septate, cellular pseudoparaphyses. Asci 78–100 × 8–
11.6 μm(
x= 88.5 × 9.8 μm, n= 20), 8-spored,
bitunicate, cylindric-clavate, slightly curved, with short
bulbose pedicel, apically rounded, with an ocular chamber.
Ascospores 12.5–15.2 × 3.8–5.6 μm(
x= 13.7 × 4.7 μm,
n= 20), hyaline, broadly fusiform, 1-septate, cells of unequal
size, deeply constricted at the septum, widest at the middle and
tapering towards the narrow ends, straight or slightly curved,
with small guttules smooth-walled. Asexual morph:
Conidiomata 200–430 μmdiam.×190–410 μmhigh
(x=326×306μm, n= 5), pycnidial, scattered or clustered,
gregarious, semi-immersed to superficial on PDA, globose to
subglobose, surrounded by vegetative hyphae. Conidiomatal
Fig. 26 Phylogram resulting
from maximum likelihood
(RAxML) analysis of LSU
sequences alignment showing the
placement of Asterodiscus
tamaricis in Stigmatodiscaceae.
Maximum likelihood bootstrap
support values equal or greater
than 50 % are indicated above or
below the nodes. The new isolates
are in blue. The tree is rooted to
Botryosphaeria dothidea
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wall 15–32 μm wide, composed of several layers of black
to hyaline pseudoparenchymatous cells of textura
angularis.Conidiophores aseptate, hyaline, mostly reduced
to conidiogenous cells. Conidiogenous cells 3.5–
7.6 × 1.2–3.4 μm(
x=5.7×1.9μm, n=20),
enteroblastic, phialidic, single, discrete, determinate, doliiform
to cylindrical or ampulliform, hyaline, arising from basal stra-
tum. Conidia 2.5–4.8 × 1.7–2.9 μm(
x=2.5×2.1μm, n= 60),
one-celled, ellipsoidal, with rounded to obtuse ends, hyaline,
smooth-walled.
Culture characteristics: Ascospores germinating on PDA
within 24 h and germ tubes produced from one or both ends.
Colonies growing on PDA reaching 17 mm diam. After
16 days at 25 °C, circular, flat, moderately dense, surface
white, smooth surface with entire to slightly undulate edge,
asexual morph produced after 2 weeks.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte - Forlì, on dead branches of Tamarix
gallica L. (Tamaricaceae), 15 January 2014, E. Camporesi
IT 1458 (MFLU 14–0597, holotype), ex-type living culture
MFLUCC 14–0168, ICMP 20704; ibid. (GZAAS, isotype).
Stigmatodiscales Voglmayr & Jaklitsch, in Voglmayr,
Gardiennet & Jaklitsch, Fungal Diversity: 10.1007/s13225–
016-0356-y, [5] (2016)
Voglmayr et al. (2016) established the order Stigmatodiscales
to accommodate the family Stigmatodiscaceae.
Fig. 27 Asterodiscus tamaricis (MFLU 14–0593)a, b Appearance of ascomata on host surface cVertical section through ascoma dPeridium e-g Asci
surrounded by pseudoparaphyses i-k Ascospores lAscospore stained in Indian ink. Scale bars: c=100μm, d-g =50μm, i-l =20μm
Fungal Diversity
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Stigmatodiscaceae Voglmayr & Jaklitsch, in Voglmayr,
Gardiennet & Jaklitsch, Fungal Diversity: 10.1007/s13225–
016-0356-y, [5] (2016)
The family Stigmatodiscaceae was introduced by
Voglmayr et al. (2016) to accommodate Asterodiscus and
Stigmatodiscus. Species of this family are characterized by
apothecioid ascomata, broadly fusoid to saccate, with thin
ecto- and thick endotunica asci, hyaline to brown ascospores
and coelomycetous asexual morphs with hyaline, irregularly
sinuously curved to falcate conidia (Voglmayr et al. 2016).
DNA phylogeny
Based on the closest matches of GenBank Nucleotide BLAST
search of strain MFLUCC 13–0072, 64 isolates belonging to
six orders (Botryosphaeriales,Dothideales,Hysteriales,
Pleosporales,Stigmatodiscales and Tubeufiales) were se-
lected. Megalotremis verrucosa (strain number: 104) was
selected as the outgroup taxon (Fig. 25). Stigmatodiscaceae
(Stigmatodiscales) deviated from all the orders included in the
phylogenetic analysis. In the second phylogenetic analysis
(Fig. 26) 11 isolates belonging to Stigmatodiscaceae were
selected with Botryosphaeria dothidea (CMW 8000) as the
outgroup taxon.
Asterodiscus tamaricis Voglmayr, Gardiennet & Jaklitsch,
Fungal Diversity: 10.1007/s13225–016-0356-y, [6] (2016),
Fig. 27
Saprobic on Tamarix species. Sexual morph:Ascomata
140–220 μm high × 170–310 μmdiam.(
x=170×262μm,
n= 10), apothecioid, solitary or scattered, immersed, slightly
erumpent through the bark, forming under black, irregularly
shaped, shallow depressions on the host surface, pyriform to
subglobose, with a wide irregular opening, ostiolate, ostiolar
area black. Peridium 10–25 μm wide, broad at the apex, com-
prising several layers of lightly pigmented to brown cells of
textura angularis, somewhat flattened at the base, broad and
black at the apex. Hamathecium comprising 2–3μmwide,
numerous, septate, cellular pseudoparaphyses, with swollen
free apical ends. Asci 75–105 × 30–48 μm(
x=90×40μm,
n= 15), 8-spored, bitunicate, broadly clavate, broadly fusoid
to saccate, short pedicellate, apically rounded, with an ocular
chamber. Ascospores 35–46 × 10.8–17 μm(
x= 39 × 13.8 μm,
n= 30), bi- to tri-seriate, hyaline, becoming brown to dark
brown when mature, ellipsoidal, 3-septate, third cell from the
base larger, constricted at the septa, deeply constricted at the
central septum, smooth-walled, with a large mucilaginous
sheath, which is constricted near the central septum. Sexual
morph: see Voglmayr et al. (2016).
Culture characteristics: Ascospores germinating on PDA
within 36 h and germ tubes produced from one or both ends.
Colonies growing on PDA slowly, 25 mm diam. After 14 days
at 25 °C, circular to slightly irregular, flat, surface white, be-
coming pale brown, when aged, smooth surface, with entire to
slightly undulate edge.
Notes:Asterodiscus was introduced by Voglmayr et al.
(2016) as a monotypic genus to accommodate A.tamaricis.
Asterodiscus tamaricis is another Tama ri x inhabitant species
reported in Austria and Croatia. We collected A.tamaricis
from Italy associated with Tamarix gallica.Massarina
mauritiana Poonyth, K.D. Hyde, Aptroot & Peerally could
be another species in Asterodiscus and is saprobic on
Bruguiera gymnorrhiza (L.) Lam. (Rhizophoraceae).
Asterodiscus closely resembles Massarina mauritiana in mor-
phology, but can be distinguished by its smaller ascospores
(26–34 × 8–9.5 μm), with a gelatinous sheath, which swells
strongly in water and appears granular at the poles and refrac-
tive adjacent to the septa in M.mauritiana (Poonyth et al.
1999). However, M.mauritiana requires molecular phyloge-
netic studies to confirm its placement in Asterodiscus.
Material examined: ITALY, Province of Forlì-Cesena [FC],
Ravaldino in Monte - Forlì, on dead branches of Tamarix
gallica L. (Tamarica ce ae ), 22 November 2012, Erio
Camporesi IT 920 (MFLU 14–0593), living culture
MFLUCC 13–0072, ICMP 20692; ibid. (HHUF 30464); ibid.
(Tamari ca ce ae), 8 October 2013, Erio Camporesi IT 920–2
(MFLU 14–059 = HHUF 30465), living culture MFLUCC
14–0994, ICMP 20744; ITALY, ibid. 15 January 2014, Erio
Camporesi IT 920–3(MFLU16–0667); ITALY, Province of
Forlì-Cesena, Fiumana, Predappio, 30 January 2014, Erio
Camporesi IT 920–4 (MFLU 16–0668)
Valsariales Jaklitsch, K.D. Hyde & Voglmayr, in Jaklitsch,
Fournier, Dai, Hyde & Voglmayr, Fungal Diversity
73(1):167 (2015)
Jaklitsch et al. (2015) established the order Va ls a r i a l e s to ac-
commodate the family Val s a r i a c e a e .
Va l s a r ia c e a e Jaklitsch, K.D. Hyde & Voglmayr, in Jaklitsch,
Fournier, Dai, Hyde & Voglmayr, Fungal Diversity 73(1):167
(2015)
The family Valsariaceae was introduced by Jaklitsch
et al. (2015)toaccommodateVa l s a r i a,Bambusaria and
Fig. 28 Valsaria tamaricis (BPI 619574)aHerbarium material b, c
Appearance of ascostromata on host surface dVertical section through
ascostroma eClose up of a locule f-g Peridium hPseudoparaphyses i-k
Asci l-o Ascospores. Scale bars: d=500μm, e-g =50μm, h=10μm, i-
k=25μm, l-o =5μm
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Myrmaecium. Species of this family are saprobic in bark or on
culms of bamboo and characterized by perithecioid ascomata,
usually monostichous in valsoid or diatrypoid configuration,
(4–)6–8-spored asci, ellipsoid to subfusiform, dark brown, 2-
Fig. 30 Homortomyces tamaricis (sexual morph, MFLU 14–0595)a,
bAppearance of ascomata on host surface cVertical section through
ascoma dPeridium ePseudoparaphyses f-i Mature bitunicate asci j-l
Ascospores mGerminating ascospore.Scale bars: c= 100 μm,
d=50μm, e, j-m =10μm, f- i =20μm
Fig. 29 Phylogram resulting from maximum likelihood (RAxML)
analysis of LSU sequence data showing the placement of Homortomyces.
Maximum likelihood bootstrap support values equal or greater than 50 %
are indicated above or below the nodes. The new isolates are in blue. The
tree is rooted to Schismatomma decolorans
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Fungal Diversity
celled ascospores and hyphomycetous or coelomycetous
asexual morphs with 1-celled, hyaline conidia (Jaklitsch
et al. 2015).
Valsaria tamaricis Mundk. & S. Ahmad, Mycol. Pap. 18: 7
(1946), Fig. 28
Facesoffungi number: FoF02165
Saprobic on dead wood. Sexual morph:Ascostromata
900–1100 μm diam. × 540–660 μmhigh(
x=
994 × 587 μm, n= 5), erumpent from bark, solitary, scattered
or gregarious, black, broadly conical or subglobose with flat-
tened base, coriaceous, multiloculate, peridium of
ascostromata 75–180 μm wide composed of brown to dark
brown cells of textura angularis.Locules 110–240 μmdiam.
×165–210 μmhigh,(
x=180×234μm, n= 10), subglobose
to conical or irregular. Ostiole inconspicuous at the surface,
papillate. Peridium of locules 13–20 μm, comprising few
layers of lightly pigmented to dark brown, flattened cells of
textura angularis, becoming hyaline towards the inner region.
Hamathecium 1.5–3.5 μm wide, composed of hyaline, sep-
tate, cellular pseudoparaphyses. Asci 70–88 × 7.5–11 μm
(x= 82.4 × 9.5 μm, n= 15), 8-spored, bitunicate, fissitunicate,
cylindrical, with a short pedicel, apically rounded with an
ocular chamber. Ascospores 10.6–14.2(−15.7) × 5.3–6.9 μm
(x= 12.8 × 6.2 μm, n= 25), uniseriate, partially overlapping,
yellowish brown when immature, becoming dark brown when
mature, (0–)1-septate, not distinctly constricted at the septum,
ellipsoid with broadly obtuse ends, finely striate. Asexual
morph: Undetermined.
Material examined: PAKISTAN, Rahim Yar Khan, on a log
of Tamarix articulata Vahl., 25 August 1964 (BPI 619574).
Notes: The genus Va l s a r ia was established by Cesati and
De Notaris (1863) and typified by V.insitiva (Tode) Ces. & De
Not. Kirk et al. (2008) categorized the genus in Diaporthales,
Sordariomycetes, while Jaklitsch et al. (2015) placed the ge-
nus Valsaria in a newly introduced family Valsariaceae,
Valsariales within the class Dothideomycetes based on
morphology and molecular phylogeny. Valsaria tamaricis
was introduced by Mundkur and Ahmad (1946) but no se-
quence data are available for this species. However, the mor-
phology fits with the genus Va ls a r i a (Jaklitsch et al. 2015)and
recollection and molecular analyses are needed to establish if
this is a distinct species.
Dothideomycetes, families incertae sedis
Homortomycetaceae Thambugala, A.J.L. Phillips & K.D.
Hyde, fam.nov.
DNA phylogeny
The phylogeny of the Homortomyces strains is shown with
two analyses based on selected LSU sequence data used in
Crous et al. (2012), Slippers et al. (2013) and Wijayawardene
et al. (2014a). The first phylogenetic analysis was carried out
using sequence data of 40 isolates belonging to four different
orders: Botryosphaeriales,Hysteriales,Tubeufiales and
Umbilicariales (Fig. 29)withSchismatomma decolorans
(DUKE 0047570 Ertz 5003) as the outgroup taxon. The
second phylogenetic analysis was done using 62 isolates
of six orders: Botryosphaeriales,Dothideales,Hysteriales,
Pleosporales,Stigmatodiscales and Tubeufiales.
Megalotremis verrucosa (strain number: 104) was selected
as the outgroup taxon (Fig. 25). In the first phylogenetic
analysis (Fig. 29), Homortomyces strains grouped in
Botryosphaeriales as a basal clade with a good BS sup-
port value (78 %), while in the second analysis (Fig. 25),
Homortomyces clustered in Botryosphaeriales as a distinct
clade separated from neighboring families.
Homortomycetaceae Thambugala, A.J.L. Phillips & K.D.
Hyde, fam.nov.
Index Fungorum Number: IF552086; Facesoffungi
number: FoF02150
Foliicolous, associated with leaf spots or saprobic on dead
twigs or branches. Sexual morph:Ascomata scattered, im-
mersed to partially erumpent, black, globose to subglobose,
ostiolate. Peridium comprising few layers of brown to dark
brown, thick-walled cells of textura angularis.Hamathecium
composed of septate, cellular pseudoparaphyses. Asci 2–6-
spored, bitunicate, fissitunicate, cylindrical, apically rounded.
Ascospores uni to bi-seriate, fusiform with broadly to narrowly
rounded ends, yellowish brown to brown, 3-septate, smooth-
walled. Asexual morph: Coelomycetous. Conidiomata pyc-
nidial, solitary or gregarious, immersed or slightly erumpent,
uniloculate or multi-loculate, globose to subglobose, ostiolate.
Conidiomatal wall comprising a few layers, of lightly
pigmented to hyaline cells of textura angularis.Paraphyses
intermingled among conidiogenous cells, extending above co-
nidia, hyaline, cylindrical, smooth, flexuous, apex obtuse,
sparingly septate. Conidiophores reduced to conidiogenous
cells or one supporting cell. Conidiogenous cell with
supporting cell, hyaline, percurrently proliferating at the tip
of the supporting cells. Conidia ellipsoid to subcylindrical,
straight to slightly curved, golden brown to dark brown,
3(−4)-euseptate, apex obtuse, base truncate with a visible scar.
Fig. 31 Homortomyces tamaricis (asexual morph MFLU 14–0596)a,
bAppearance of conidiomata on host m, n Appearance of conidiomata
on PDA c, d,oVertical sections through conidiomata e, f, p Conidiomatal
wall g-i, q–sDeveloping conidia and paraphyses j, k Conidia l
Germinating conidia. Scale bars: c-f =50μm, g=20μm, h-l,
q-s =10μm, o, p =100μm
Fungal Diversity
Notes: The genus Homortomyces was introduced by Crous
et al. (2012) to accommodate H.combreti Crous & M.J.
Wingf., which was associated with leaf spots on Combretum
erythrophyllum (Burch.) Sond. and placed in Dothideomycetes
genera incertae sedis. Wijayawardene et al. (2014a)described
another Homortomyces species, H.tamaricis Wijayaw.,
Camporesi & K.D. Hyde from dead branches of Tamarix
gallica. In the present study, we found the sexual morph of
H.tamaricis associated with the same host and this is con-
firmed by the phylogenetic analyses (Fig. 29). Crous et al.
(2012) showed Homortomyces to have phylogenetic affinities
in Botryosphaeriales. In our preliminary analysis using
LSU sequence data, H.combreti and H.tamaricis formed
aseparatecladeinBotryosphaeriales that was distinct from
all other families in this order (Fig. 25). Homortomyces is
morphologically similar to Stilbospora, which is currently
classified in Stilbosporaceae,Diaporthales,
Sordariomycetes (Crous et al. 2012; Wijayawardene et al.
2014a; Maharachchikumbura et al. 2016), while Crous
et al. (2012) pointed out four genera (Endocoryneum,
Hendersoniopsis,Angiopomopsis and Ceratopycnis) that
have conidia similar to Homortomyces. However, all these
genera have been classified in incertae sedis and lack se-
quence data. Morphologically, Homortomyces shares some
characteristics of Botryosphaeriales such as stromatic
conidiomata, that develop within a preformed stroma, with
septate, pigmented, thick-walled, diplodia-like conidia, and
clavate, bitunicate, fissitunicate asci with a thick endotunica
and a well-developed apical chamber (Phillips et al. 2013).
Homortomyces can be distinguished from other members
of Botryosphaeriales in having uniloculate, thin-walled
ascomata and 3-septate, pigmented ascospores. Based on
our phylogenetic analyses (Figs. 25 and 29) and its unique
morphology, we recognize Homortomyces as a distinct
family (Homortomycetaceae), but its phylogenetic place-
ment is uncertain. Therefore, tentatively we place
Homortomycetaceae in Dothideomycetes,families incertae
sedis, rather than assigning to Botryosphaeriales, until
more fresh collections are made and multi-gene phyloge-
netic analyses carried out.
Type genus:Homortomyces Crous & M.J. Wingf., in
Crous, Groenewald, Lombard & Wingfield, IMA Fungus
3(2): 110 (2012)
Foliicolous, associated with leaf spots or saprobic on dead
twigs or branches. Sexual morph:Ascomata scattered, im-
mersed to partially erumpent, black, globose to subglobose,
ostiolate. Peridium comprising 4–6 layers of brown to dark
brown, thick-walled cells of textura angularis.Hamathecium
composed of septate, cellular pseudoparaphyses. Asci 2–6-
spored, bitunicate, fissitunicate, cylindrical, apically rounded,
with a distinct ocular chamber. Ascospores uni- to bi-seriate,
fusiform with broadly to narrowly rounded ends, yellowish
brown to brown, 3-septate, slightly curved, smooth-walled.
Asexual morph:Conidiomata pycnidial, solitary or gregari-
ous, immersed to slightly erumpent, uniloculate to multi-
loculate, globose to subglobose, ostiolate. Conidiomatal wall
comprising a few layers, of lightly pigmented to hyaline cells
of textura angularis.Paraphyses intermingled among
conidiogenous cells, extending above conidia, cylindrical,
smooth, flexuous, apex obtuse, sparingly septate.
Conidiophores reduced to conidiogenous cells or one
supporting cell, hyaline, cylindrical, with 1–4 inconspicuous,
percurrent proliferations at the apex. Conidiogenous cell with
supporting cell, hyaline, percurrent proliferation at the tip
of the supporting cells. Conidia ellipsoidtosubcylindrical,
straight to slightly curved, golden brown to dark brown,
3(−4)-euseptate, apex obtuse, base truncate with a visible
scar, verruculose or smooth-walled (Crous et al. 2012;
Wijayawardene et al. 2014a).
Type species:Homortomyces combreti Crous & M.J.
Wingf., in Crous, Groenewald, Lombard & Wingfield, IMA
Fungus 3(2): 113 (2012)
Table 3 Synopsis of Cytospora species on Tam ar ix
Species Conidia (μm) Host Locality References
C.tamaricella Syd. & P. Syd. 7–9×1–1.5 Ta ma ri x ga ll ic a Germany Sydow and Sydow (1904)
C.tamaricis Brunaud 5 × 2 Tamar ix g al li ca Germany Saccardo and Sydow (1899)
C.tamaricis Brunaud 4.2–6.5 × 1–1.6 Tamarix parviflora USA BPI 365752
C.amaricophila Maire & Sacc. 3 × 1 Tamarix africana Corsica Saccardo (1906)
C.italica Thambugala, Camporesi
& K.D. Hyde
3.7–5.2 × 1–1.3 Ta ma rix gallica Italy Present study
Fig. 32 Phylogram resulting from maximum likelihood (RAxML)
analysis of ITS sequence data for Cytospora species. Maximum
likelihood bootstrap support values equal or greater than 50 % are
indicated above or below the nodes. The new isolates are in blue. The
tree is rooted to Phomopsis vaccinii
Fungal Diversity
Fungal Diversity
Homortomyces tamaricis Wijayaw., Camporesi & K.D.
Hyde, in Wijayawardene et al., Phytotaxa 176(1): 160
(2014). Figs. 30 and 31
Facesoffungi number: FoF02151
Saprobic on Tamari x ga llica L. Sexual morph:Ascomata
180–275 μm high × 270–340 μmdiam.(
x=234×310μm,
n= 6), scattered, immersed to partially erumpent, black, glo-
bose to subglobose, ostiolate. Peridium 20–45 μm wide, com-
prising 4–6 layers of brown to dark brown, thick-walled cells
of textura angularis.Hamathecium composed of 2–4μm
wide, septate, cellular pseudoparaphyses. Asci 75–98 × 18–
22 μm(
x= 85 × 20.6 μm, n= 8), mostly (1–)2–6-spored,
bitunicate, fissitunicate, cylindrical, short pedicellate, apically
rounded, with a distinct ocular chamber. Ascospores 23–
29 × 8.5–12 μm(
x= 26 × 10.1 μm, n= 25), uni- to bi-seriate,
fusiform with broadly to narrowly rounded ends, yellowish
brown to brown, 3-septate, slightly curved, smooth-walled,
without a mucilaginous sheath. Asexual morph:
Conidiomata 150–240 μm high × 300–360 μmdiam.(
x=
203 × 332 μm, n= 6), solitary or gregarious, immersed to
slightly erumpent, pycnidial to irregular, uniloculate to
multi-loculate, subglobose, ostiolate. Conidiomatal wall 9–
23 μm(
x= 14.3 μm, n= 15), comprising a few layers of
Fig. 33 Cytospora tamaricis (BPI 365752)aHerbarium material bAppearance of conidiomata on the host surface c-e Sections through conidiomata f
Ostiole gConidiomatal wall hConidiogenous cells and developing conidia iConidia. Scale bars: c, d =200μm, e-g =50μm, h=20μm, i=10μm
Fungal Diversity
lightly pigmented to hyaline cells of textura angularis.
Paraphyses numerous, aseptate, cylindrical. Conidiophores
reduced to conidiogenous cells. Conidiogenous cell 6–
27 × 2.5–5.5 μm(
x= 14 × 3.5 μm, n=20),withsupporting
cell, hyaline, percurrently proliferating at the tip of the
supporting cells. Conidia ellipsoid to subcylindrical, straight
to slightly curved, in vivo; 22–27.5 × 9–12.2 μm(
x=
24.6 × 11 μm, n= 30), golden brown to dark brown, becoming
coloured before release from conidiogenous cells, smooth-
walled, 3-septate, apex obtuse, base truncate, smooth-walled;
in vitro; 20–31 × 8.5–12.5 μm(
x= 24.5 × 10.5 μm, n=30),
light brown, aseptate or 1-septate, guttulate.
Culture characteristics: Ascospores germinating on PDA
within 18 h and germ tubes produced from one or both ends.
Colonies growing on PDA at 25 °C reaching 30 mm diam.
After 14 days at 25 °C, irregular, flat, moderately dense, sur-
face white, with slightly undulate edge, producing conidia
after 14 days frequently at the margin.
Material examined: ITALY, Province of Forlì-Cesena
[FC]), Ravaldino in Monte Forlì, on dead branches of
Tamari x ga lli ca L. (Tamaricaceae), 22 November 2012,
Erio Camporesi IT 922 (MFLU 14–0595, sexual morph),
ex-type living culture of sexual morph MFLUCC 13–
0280, ICMP 20745; Fiumana –Predappio, on dead
branches of Tamarix gallica L. (Tamaricaceae), 25
September 2013, Erio Camporesi IT 922–2(MFLU14–
0596, asexual morph), living culture MFLUCC 14–0167,
ICMP 21071.
Fig. 34 Cytospora italica (holotype) a Appearance of conidiomata on
the host surface bTransverse sections through conidiomata c-d
Longitudinal sections through conidiomata eConidiomatal wall f-g
Conidiogenous cells and developing conidia hConidia. Scale bars:
c=200μm, d=50μm, e, h =10μm, f-g =15μm
Fungal Diversity
Sordariomycetes
The class Sordariomycetes has been treated by
Maharachchikumbura et al. (2016) and the taxonomic ar-
rangements used by these authors are followed here.
Fig. 35 Cytospora unilocularis (holotype) a Appearance of ascostromata on host surface bVertical section through ascostroma cPeridium dAscus e-f
Ascospores gUpper view of the colony on PDA. Scale bars: b=100μm, c,e=50μm, d=10μm, f=20μm
Fig. 36 Phylogram resulting from maximum likelihood (RAxML)
analysis of combined ITS and β-tubulin sequence data of selected
strains of Eutypella species in Diatrypaceae. Maximum likelihood
bootstrap values equal or greater than 50 % are indicated above or
below the nodes. The new isolates are in blue. The tree is rooted to
Cryptovalsa ampelina
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Fungal Diversity
Va l s a c ea e Tul. & C. Tul. [as ‘Va ls a rum ’], Select. fung. carpol.
(Paris) 1: 180 (1861)
The family Valsaceae was introduced by Tulasne and
Tulasne (1861) and is characterized by immersed to erumpent
ascomata bearing inoperculate, unitunicate, 8-spored asci with
shallow, refractive, J–apical ring and hyaline or yellowish
brown ascospores. Members of Vals a cea e possess a
pseudostromata usually with a clypeus surrounding the
ostiolar neck. The asexual morph of the Valsaceae are
acervular or pycnidial, with phialidic conidiogenous cells
(Adams et al. 2005). Maharachchikumbura et al. (2016)ac-
cepted 13 genera in the family.
Cytospora Ehrenb., Sylv. mycol. berol. (Berlin): 2 (1818)
Cytospora species are important phytopathogens causing
severe canker disease on a broad range of hosts (Adams et al.
2006;Ariyawansaetal.2015b; Fan et al. 2015a,b;Lietal.
2016)andCytospora canker is a common disease in
Eucalyptus in both commercial plantations and natural forests
(Fraser and Davison 1985). Val s a species are known as the
sexual morph of Cytospora species and Va ls a species are treat-
ed as synonyms of Cytospora (Fan et al. 2015a,b). Although
577 Cytospora epithets are recorded in Index Fungorum
(2016), ex-type sequence data is available for only a few spe-
cies. In this study we collected both sexual and asexual
morphs of two different Cytospora species associated with
Tamari x in Italy. Three different Cytospora species (asexual
morph) have been reported from Tam arix species (Table 3),
but no sequence data are available for them in GenBank.
Our asexual morph collection (MFLU 14–0592) differs
from these three Cytospora species mainly in conidial dimen-
sions. Our collections of the sexual morph have uniloculate
ascomata and show fungicolous characters that are unique and
no Va l s a species (which are considered as sexual morph of
Cytospora)havebeenreportedonTama ri x hosts.
DNA phylogeny
Reference sequences used in the phylogenetic analyses were
selected based on the data from Ariyawansa et al. (2015b)a
nd
Fan et al. (2015a,b). The present phylogenetic analysis based
on ITS and β-tubulin sequence data showed 53 subclades
representing 53 distinct species in Cytospora. The two strains
clustered in separate lineages from other Cytospora species.
Therefore, two new species, Cytospora italica and C.
unilocularis are introduced in this study. Phomopsis vaccinii
(ATCC 18451) was selected as the outgroup taxon (Fig. 32).
Cytospora tamaricis Brunaud, Act. Soc. linn. Bordeaux,
Trois. sér. 53: 13 [repr.] (1898), Fig. 33
Saprobic on dead branches of Tamarix parviflora DC.
Sexual morph: Undetermined. Asexual morph:Stromata
immersed in host. Conidiomata 440–800 μmdiam.×240–
450 μmhigh(
x=537×340μm, n= 8), scattered, or some-
times in small groups, immersed to partially erumpent, black,
multi-loculate, ostiolate. Locules 65–330 μmdiam.×150–
332 μmhigh(
x= 200 × 230 μm, n= 10), single,
monostichous, subglobose to ovoid or obpyriform.
Conidiomatal wall 7–26 μm(
x=15.8μm), comprising a few
layers of dark brown to lightly pigmented, thick-walled cells of
textura angularis.Conidiophores indistinct. Conidiogenous cells
9–18 × 0.6–1μm(
x= 13.4 × 0.7 μm, n= 30), enteroblastic,
phialidic, hyaline, smooth. Conidia 4.2–6.5 × 1–1.6 μm(
x=
5.4 × 1.3 μm, n= 50), hyaline, allantoid, aseptate, thin-walled.
Material examined: USA, Oregon, Portland, on Tamar ix
parviflora DC., 1931 April 15, J.R. Hansbrough (BPI
365752)
Notes: Due to the lack of molecular data, the phylogenetic
distinctiveness of Cytospora tamaricis has not been con-
firmed. The taxonomic classification of this species has not
been treated in detail.
Cytospora italica Thambugala, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF 552098, Facesoffungi
number: FoF02168, Fig. 34
Etymology: Named after the country (Italy), where the ho-
lotype was collected.
Holotype:MFLU14–0592
Saprobic or necrotrophic on twigs and branches of Tamar ix
gallica L. Sexual morph: Undetermined. Asexual morph:
Stromata immersed in bark. Conidiomata 580–730 μmdiam.
×350–730 μmhigh(
x= 690 × 582 μm, n= 6), erumpent
through the surface of bark, flat to discoid, with multi-locules.
Locules 150–645 μm diam. × 180–400 μmhigh(
x=
357 × 260 μm, n=8).Conidiophores indistinct.
Conidiogenous cells 10–32 μm×(0.7–)1–2.4 μm(
x=
16 × 1.6 μm, n= 35), enteroblastic, phialidic, formed from
the inner most layer of pycnidial wall, hyaline, smooth.
Conidia 3.7–5.2 μm×1–1.3 μm(
x=4.3×1.1μm, n=50),
hyaline, elongate-allantoid, aseptate, eguttulate.
Culture characteristics: Conidia germinating on PDAwith-
in 18 h. Colonies on PDA fast growing, reaching 32 mm diam.
Fig. 37 Eutypella tamaricis (holotype)aAppearance of stromata on
host bLongitudinal section through ascostroma immersed in host
surface cVertical section through locule dOstiole ePeridium f
Paraphyses g-i Asci jAscospores kGerminating ascospores. Scale
bars: c=200μm, d=100μm, e=30μm, f=10μm, g- i =50μm,
j=25μm, k=20μm
Fungal Diversity
Fungal Diversity
After 3 days at 25 °C, flat, circular to irregular, dense, white,
reverse light olivaceous green, surface smooth, with entire to
slightly undulate edge.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte - Forlì, on dead branches of Tamarix
gallica L. (Tam ar icace ae ), 28 March 2014, Erio Camporesi
IT 918D (MFLU 14–0592, holotype), ex-type living culture
MFLUCC 14–0440, ICMP 20691.
Notes: Unfortunately, there is no sequence data for those
Cytospora species previously reported on Tam ar ix. However,
conidial dimensions of C.italic are different (Table 3) and the
new species is described based on morphological features and
molecular data are provided for future studies.
Cytospora unilocularis Thambugala, Camporesi & K.D.
Hyde, sp. nov.
Index Fungorum number: IF552099, Facesoffungi
number: FoF02169, Fig. 35
Etymology: In reference to uniloculate ascostromata
Holotype:MFLU16–670
Saprobic on twigs and branches of Tamari x spp. or
sometimes forming fruiting bodies in old ascostromata.
Sexual morph:Ascostromata 175–300 μmdiam.×
140–240 μmhigh(
x= 238 × 185 μm, n=8),im-
mersed in old ascomata, slightly erumpent through the
surface of bark, subglobose to irregular, uniloculate.
Peridium up to 50 μm wide, composed of several layers
of dark brown to black, flattened, thick-walled cells of
textura angularis, becoming hyaline towards the inner
layers. Asci 61–73 × 7.6–9.7 μm(
x= 65.9 × 8.8 μm,
n= 15), 8-spored, unitunicate, clavate to elongate-obovoid,
with a non-amyloid apical ring. Ascospores 9.6–13.9 × 2.3–
3.7 μm(
x= 11.8 × 2.9 μm, n= 50), overlapping biseriate,
elongate allantoid, hyaline, lacking guttules, aseptate, smooth-
walled. Asexual morph: Undetermined.
Culture characteristics: Ascospores germinated on PDA
within 18 h. Colonies on PDA fast growing, reaching
35 mm diam. After 6 days at 25 °C, flat or effused, circular
to irregular, dense, white, surface smooth, with entire to
slightly undulate edge.
Material examined: ITALY, Province of Ravenna, Lido di
Dante, on dead branches of Tama ri x sp. (Tamaricaceae), 4
December 2014, Erio Camporesi IT 2280 (MFLU 16–670,
holotype), ex-type living culture MFLUCC 15–0481, ICMP
21252; ibid., IT 2280–2(MFLU16–0671, paratype), living
culture MFLUCC 15–0482; ibid., 10 March 2015, Erio
Camporesi IT 2280–3 (MFLU 15–1028, paratype), living
culture MFLUCC 15–0483.
Notes:Cytospora unilocularis is introduced here as a new
species based on molecular evidence and morphological fea-
tures. No sexual morphs of Cytospora species have been re-
ported on Tam arix species and the fungicolous character and
uniloculate ascostromata of this species are unique. Cytospora
unilocularis formed a well-supported sister clade (98 % BS) to
C.acacia Oudem.
Diatrypaceae Nitschke [as ‘Diatrypeae’], Verh. naturh. Ver.
preuss. Rheinl. 26: 73 (1869)
The family Diatrypaceae was introduced by Nitschke (1869)
and currently comprises 14 genera (Maharachchikumbura et al.
2016). Recent papers (Carmarán et al. 2006, Liu et al. 2015;
Ariyawansa et al. 2015b;Lietal.2016) have contributed to new
sequence data and provided backbone trees for the family.
Senanayake et al. (2015) provided a description and reference
specimen for the generic type, Diatrype disciformis (Hoffm.) Fr.
Several genera in the Diatrypaceae (Cryptovalsa, Diatrype,
Diatrypella, Eutypa,andEutypella) are known to occur on
grapevines worldwide (Luque et al. 2006).
Eutypella (Nitschke) Sacc., Atti Soc. Veneto-Trent. Sci. Nat.,
Padova, Sér. 4 4: 80 (1875)
Eutypella species are found on a wide range of hosts, es-
pecially associated with canker diseases in Vitis vinifera
(Vasilyeva and Stephenson 2006; Trouillas et al. 2011;
Luque et al. 2012). Both Maharachchikumbura et al. (2016)
and Senanayake et al. (2015) included Eutypella in
Diatrypaceae. Currently 247 species epithets are listed in
Index Fungorum (2016), but only a few Eutypella species
have sequence data. Until more species are collected,
epitypified and sequenced, understanding of the genus
Eutypella will remain incomplete.
DNA phylogeny
Phylogenetic analysis (Fig. 36) using combined ITS and β-
tubulin sequence data was carried out using strains selected
from Trouillas et al. (2011) and GenBank BLAST searches.
Cryptovalsa ampelina (B10-16 A) was selected as the
outgroup taxon. Strains MFLUCC 14–0444 and MFLUCC
14–0445 clustered together with putatively named strains
of C.rabenhorstii (CBS 128338 and CBS 128339) in the
genus Eutypella.
Fig. 38 Phylogram resulting from maximum likelihood (RAxML)
analysis of a combined ITS, EF1-αand β-tubulin sequence data of
selected strains of Diaporthe species in Diaporthaceae.Maximum
likelihood bootstrap values equal or greater than 50 % are indicated above
or below the nodes. The new isolates are in blue. The tree is rooted to
Diaporthe helianthi
Fungal Diversity
Fungal Diversity
Eutypella tamaricis Thambugala, Camporesi & K.D. Hyde,
sp. nov.
Etymology: The species epithet Btamaricis^refers to the
host genus Tam ar ix on which the holotype was collected.
Index Fungorum Number: IF552235; Facesoffungi
number: FoF02170, Fig. 37
Holotype:MFLU14–0598
Saprobic on Tamari x gallica L. Sexual morph:Stromata
up to 2.6 mm wide, scattered on host, immersed to partially
erumpent, somewhat convex, arising through cracks in bark
epidermis, with numerous perithecia immersed in a single
stroma, ostioles opening in the central region and appearing
as black circular spots. Ascomata 425–620 μmdiam.,825–
925 μmhigh(x=508×868μm, n= 8), perithecial, immersed
in stromatic tissues, aggregated, globose to subglobose,
narrowing towards the apex, dark brown, thin-walled,
ostiolate. Ostiole short, compressed, apex wider than base,
periphysate, ostiolar opening composed of dark brown
to black cells. Peridium 30–40 μmwideatthebase,
comprising several layers of dark brown to lightly
pigmented, thick-walled cells of textura angularis,be-
coming flattened and hyaline towards the inner region,
up to 80 μm wide at the apex, comprising several
layers of dark brown to lightly pigmented, thick-walled
cells of textura epidermoidea,becomingtextura
angularis to textura prismatica and hyaline towards
the inner region. Hamathecium comprising 2–3μmwide,
septate, cellular paraphyses. Asci 74–115(−132) × 16.2–
22.6 μm(
x= 94 × 18.4 μm, n= 15), polysporous, unitunicate,
very long pedicellate, with cylindrical, thick-walled,
swollen upper portion, apex flat, with J-, cylindrical,
Fig. 39 Diaporthe ravennica (holotype)aAppearance of conidiomata on host surface b-d Vertical sections through conidiomata eConidiomatal wall f
Conidiophores and developing conidia gAlpha conidia hBeta conidia. Scale bars: b=100μm, c, d =50μm, e, f =20μm, g, h =10μm
Fungal Diversity
conspicuous, apical ring. Ascospores 9–14.7 × 2.8–4μm
(x= 11.9 × 3.4 μm, n= 50), overlapping, crowded, yellowish
to light-brown, allantoid to oblong, unicellular, thin-walled,
smooth-walled. Asexual morph: Undetermined.
Material examined: ITALY, Province of Forlì-Cesena,
Ravaldino in Monte, on branches of Tamarix gallica L., 25
March 2014, Erio Camporesi IT 1763 (MFLU 14–0598,
holotype), ex-type living culture MFLUCC 14–0444, ICMP
20705; ibid., IT 1763–1(MFLU14–0599), MFLUCC 14–0445.
Notes:Cryptovalsa rabenhorstii has characters typical of
Cryptovalsa (Saccardo 1882; Trouillas et al. 2011), but recent
studies showed that putatively named strains (CBS 128338
and CBS 128339) of C.rabenhorstii clustered with
Eutypella species (Trouillas et al. 2011). However, the holo-
type specimen or ex-type culture of C.rabenhorstii cannot be
located for study. We compared our new collections (MFLU
14–0598 and MFLU 14–0599) with the descriptions of
Saccardo (1882)andTrouillasetal.(2011). The dimensions
of asci and ascospores in those studies differed slightly. In our
Fig. 40 Coryneopsis tamaricis (BPI 374366)aHerbarium material bConidiomata immersed in the host surface cVertical section through conidioma d
Conidioma wall e-j Conidia attach to conidiogenous cells. Scale bars: c=150μm, d=50μm, e=100μm, f-j =10μm
Fungal Diversity
phylogenetic analysis our strains (MFLUCC 14–0444 and
MFLUCC 14–0445) clustered together with the putatively
named strains of C.rabenhorstii which grouped with other
Eutypella species. However, neither Cryptovalsa nor
Eutypella species have been recorded on Tamarix species.
The putatively named strains of C.rabenhorstii have been
obtained from Vitis vinifera in Australia. It is difficult to distin-
guished Cryptovalsa and Eutypella species by considering the
number of ascospores in the asci, as both 8-spored and
polysporous asci are typical of Eutypella species (Trouillas
et al. 2011). We therefore introduce a new species, E.tamaricis.
Diaporthaceae Höhn. ex Wehm., Am. J. Bot. 13: 638 (1926)
The genera within Diaporthaceae are pathogens, endo-
phytes or saprobes and have a cosmopolitan distribution
(Gomes et al. 2013;Udayangaetal.2014). Phylogenetic anal-
yses by Maharachchikumbura et al. (2016) showed that the
family Diaporthaceae clustered as a monophyletic clade sister
to Va l s a c ea e in Diaporthales and currently comprises eleven
genera (Maharachchikumbura et al. 2016).
Diaporthe Nitschke, Pyrenomyc. Germ. 2: 240 (1870)
Nitschke (1870) established the genus Diaporthe typified
by Diaporthe eres Nitschke and this species has been studied
in detail by Udayanga et al. (2015). Species of this genus are
plant pathogens, endophytes or saprobes, commonly isolated
from a wide range of hosts (Gomes et al. 2013; Udayanga
et al. 2014; Hyde et al. 2014). Phomopsis species are the
asexual morphs of Diaporthe and are now considered a syn-
onym. The phylogeny and taxonomy of the genus has been
addressed in several recent studies (Udayanga et al. 2014,
2015; Liu et al. 2015; Gomes et al. 2013;Lombard et al.
2014; Dissanayake et al. 2015).
DNA phylogeny
Reference sequences used in the phylogenetic analyses were
obtained from recent studies (Udayanga et al. 2014;Lombard
et al. 2014) and downloaded from GenBank. Diaporthe
helianthi (CBS 592.81) was selected as the outgroup taxon.
The phylogenetic analysis based on a combined ITS, EF1-α
and β-tubulin sequences, revealed strains MFLUCC 15–0479
and MFLUCC 15–0480 grouped in the genus Diaporthe and
are phylogenetically allied to D.foeniculina and D.baccae,
but in a different subclade (Fig. 38).
Diaporthe ravennica Thambugala, Camporesi & K.D. Hyde,
sp. nov.
Index Fungorum number: IF552100, Facesoffungi
number: FoF02171, Fig. 39
Etymology: Refers to the name of the province in Italy
(Province of Ravenna), where the holotype was collected.
Holotype:MFLU16–0665
Saprobic on dead branches of Tamarix species. Sexual
morph: Undetermined. Asexual morph:Stromata up to
500 μm diam. Scattered, immersed to slightly erumpent,
multi-loculate, black. Locules 140–275 μmdiam.×65–
175 μm high, (x=190×108μm, n= 5), subglobose or
irregular, with or without an ostiole. Conidiomatal wall com-
prising several layers of lightly pigmented to dark brown cells
of textura angularis, becoming hyaline towards the inner re-
gion. Paraphyses not observed. Conidiophores hyaline,
smooth, usually unbranched, cylindrical, straight to sinuous,
12–23 × 1.2–1.8 μm(
x= 16.4 × 1.5 μm, n=40).
Conidiogenous cells phialidic, cylindrical, terminal, slightly ta-
pering towards the apex. Alpha conidia (6.4–)7.2–10.5 × 1.7–
2.8 μm(
x=8.6×2.3μm, n= 40), aseptate, hyaline, ellipsoidal
or fusiform, with no, two or many guttules, smooth-walled.
Beta conidia 21–32(−38) × (1–)1.2–1.7 μm(
x=
26.4 × 1.3 μm, n= 30), aseptate, filiform, hyaline, eguttulate,
slightly curved, abundant, base subtruncate, acute apex.
Material examined: ITALY, Province of Ravenna, Lido di
Dante, on dead branches of Tama ri x sp. (Tamaricaceae), 4
December 2014, Erio Camporesi IT 2273 (MFLU 16–0665,
holotype), ex-type living culture MFLUCC 15–0479, ICMP
21251; ibid., (GZAAS, isotype); ibid., IT 2273–2 (MFLU 16–
0666, paratype), living culture MFLUCC 15–0480.
Notes:Diaporthe ravennica is reported in its asexual form
on Tama ri x and is introducedhere as a new species. Diaporthe
tamaricina Sacc. & Flageolet has also been recorded on
Tamari x, but is a sexual morph (Saccardo 1905). Phomopsis
tamaricaria (Sacc.) Grove, is an asexual morph reported on
Tamari x afric an a from France and has a very brief description
(Saccardo 1884) and, unfortunately the type material in PAD
cannot be borrowed to assess its characters.
Sordariomycetes genera incertae sedis
Coryneopsis tamaricis (Cooke) Grove, J. Bot., Lond. 70: 34
(1933) [1932]
Basionym: Hendersonia tamaricis Cooke, Grevillea
14(no. 69): 5 (1885)
Facesoffungi number: FoF02172; Fig. 40
Saprobic on Tamarix africana Poir. Sexual morph:
Undetermined. Asexual morph:Conidiomata 330–455 μm
diam. × 210–270 μmhigh(
x=384×246μm, n= 5), solitary
or gregarious, immersed, pycnidial to irregular, uniloculate,
globose to subglobose, ostiolate. Pycnidium wall 10–
14.5 μm(
x=12.3μm, n= 15), comprising a few layers, of
lightly pigmented to hyaline cells of textura angularis.
Fungal Diversity
Table 4 List of fungi occuring on Ta ma rix species
Species Host Locality References
Alternaria tamaricis T.Y. Zhang Tamarix chinensis China Zhang 2003, Zhuang 2005
Alternaria tenuissima (Kunze)
Wiltshire
Tamarix g al lica Italy This study
Aplosporella tamaricis (Hollós) Moesz Tamarix g al lica Hungaria MycoBank 2016
Asterodiscus tamaricis Vo g l ma y r ,
Gardiennet & Jaklitsch
Tamarix g al lica,Tamarix
tetrandra
Austria, France, Italy, Croatia Voglmayr et al. 2016,Thisstudy
Ascochyta tamaricis Golovin Tam ar ix sp. China, Turkmenistan Koshkelova and Frolov 1973,
Chen 2002,Zhuang2005
Brunneiperidium gracilentum
Daranagama, Camporesi & K.D.
Hyde
Tamarix g al lica Italy Daranagama et al. 2015
Botryosphaeria tamaricis (Cooke)
Theiss. & Syd.
Tamarix articulate,Ta ma ri x
dioica,Tamarix g al li ca ,
Tamarix sp.
Argentina, India, Pakistan, U.S.A Sydow et al. 1911, Anonymous
1960, Spaulding 1961,Ahmad
1978, Kanaujia 1978,Ahmad
et al. 1997
Botrytis cinerea Pers. Tamarix sp. U.S.A Gilman 1932
Camarosporium sp. Tam ar ix sp. China Zhuang 2005
Calospora tamaricis Maubl. Tamarix sp. France Saccardo 1913
Maublanc 1906
Camarosporium potebniae Sacc. Tamarix sp. Ukraine Morochkovsky et al. 1971
Camarosporium tamaricis Hollós Tamarix ramosissima,Ta mari x sp. Russia, Turkmenistan, Ukraine,
Uzbekistan
Potebnia 1907, Gaponenko 1965,
Morochkovsky et al. 1971,
Koshkelova and Frolov 1973
Camarosporium tamaricum Mekht. Tamarix smyrnensis Azerbaijan Mekhtijeva 1962
Camarosporium tauriae GucevičTamarix ramosissima Russia, Ukraine Gucevičh1960
Chrysothrix septemseptata Jagadeesh
et al.
Tamarix g al lica India Singh and Sinha 2010
Cilioplea coronata (Niessl) Munk Ta ma ri x sp. Ukraine Dudka et al. 2004
Clavariopsis bulbosa Anastasiou Ta ma ri x aphyll a California Anastasiou 1961
Clasterosporium tamaricinum Maire Tamarix a fr ic an a France Ahmad 1967
Colletotrichum dematium (Pers.) Grove Ta ma ri x ga ll ic a Morocco Rieuf 1970
Coniochaeta pulveracea (Ehrh.) Munk Ta ma ri x sp. China Zhuang 2005
Coniothecium tamariscinum Thüm. Tamarix mannifera Egypt Mouchacca 2009
Coniothyrium caespitulosum Sacc. Tamarix hohenackeri,Tamarix
smyrnensis,Tam ar ix sp.,
Georgia, Greece Pantidou 1973, Nakhutsrishvili
1986
Coniothyrium globisporum Bubák &
Serebrian.
Tamarix sp. Turkmenistan Koshkelova and Frolov 1973
Coniothyrium henningsii Gonz. Frag. Tamarix gallica Spain Gonzalez Fragoso 1917
Coniothyrium tamaricis Oudem. Tam ar ix g al li ca ,Tamarix hispida,
Tamarix sp.
Portugal, Turkmenistan , Ukraine,
Uzbekistan, Uzbekistan
Moskovets 1933, Gaponenko
1965, Koshkelova and Frolov
1973, de Sousa Dias and Lucas
1975
Coniothyrium tamarisci (Mont.) Petr. Tamarix gracilios,Tamarix
ramosissima,Tam ar ix
smyrnensis,Tam ar ix sp.,
Tamarix tetrandra
Europe, Georgia, Russia, Ukraine Saccardo 1884, Nakhutsrishvili
1986, Isikov 1989, Dudka et al.
2004, Rebriev et al. 2012
Corollospora pulchella Kohlm., I.
Schmidt & N.B. Nair
Tamarix a ph ylla California Shearer and Crane 1971
Cryptosphaeria tamaricis Grev. Ta ma ri x ge rmanica UK Greville 1823
Cucurbitaria berberidis Fuckel Tamarix tetrandra Ukraine Dudka et al. 2004
Cucurbitaria tamaricina Smyk Tamarix ramosissima Ukraine Smyk 1970
Cytospora chrysosperma (Pers.) Fr. Tamarix ramosissima,Ta mari x sp. Iran, Russia Fotouhifar et al. 2010,Rebriev
et al. 2012
Cytospora italica Thambugala,
Camporesi & K.D. Hyde
Tamarix g al lica Italy This study
Fungal Diversity
Table 4 (continued)
Species Host Locality References
Cytospora tamaricella Syd. & P. Syd. Tamarix chinensis,Tamarix
gallica,Tama ri x sp.
China, Germany, Turkmenistan Sydow and Sydow 1904,
Koshkelova and Frolov 1973,
Chen 2002,Zhuang2005
Cytospora tamaricis Brunaud Tam ar ix g al li ca ,Tam arix sp. Germany, Poland, Portugal Saccardo and Sydow 1899,de
Sousa Dias and Lucas 1975,
Mulenko et al. 2008
Cytospora tamaricophila Maire &
Sacc.
Tamarix a fr ic ana,Ta marix gall ica Canary Islands, Corsica Saccardo 1906, de Urries 1957
Diaporthe ravennica Thambugala,
Camporesi & K.D. Hyde
Tamarix sp. Italy This study
Diaporthe tamaricina Sacc. &
Flageolet
Tamarix a fr ic ana,Ta marix gall ica France, U.K., Morocco Saccardo 1905,Cannonetal.1985
Didymosphaeria futilis (Berk. &
Broome) Rehm
Tamarix ramosissima Georgia Nakhutsrishvili 1986
Didymosphaeria tamaricis Koshk. &
Frolov
Tamarix sp. Turkmenistan Koshkelova and Frolov 1973
Diplodia tamaricina Sacc. Tamarix sp. France , U. S. A Anonymous 1960, Saccardo 1884
Diplodia tamaricis Rabenh. Tamarix ramosissima,Tamarix sp. Germany, Russia Rebriev et al. 2012,Index
Fungorum 2016
Diplodia taxi (Sowerby) De Not. Tamarix ramosissima Ukraine Merezhko 1980
Diplodiella tamaricina S. Ahmad Tamarix articulata Pakistan Ahmad 1969, Ahmad et al. 1997
Dothidotthia ramulicola (Peck) M.E.
Barr
Tamarix sp. Turkmenistan Aptroot 1995
Dothiora tamaricis Dennis & Spooner Ta ma ri x ga ll ic a Azores Dennis et al. 1977
Dothiorella tamaricicola M.T. Lucas
& Sousa da Câmara
Tamarix sp. Portugal Lucas and Sousa da Câmara 1955
Dothiorella sarmentorum (Fr.) A.J.L.
Phillips, J. Luque & A. Alves
Tamarix ramosissima Russia This study
Duportella tristicula (Berk. &
Broome) Reinking
Tamarix indica India Kanaujia 1978
Eutypa lata (Pers.) Tul. & C. Tul. Tamarix sp. Australia Cook and Dubé 1989, Carter 1991
Eutypa ludibunda Sacc. Tam ar ix a fr ic an a Argentina Saccardo 1913
Eutypella tamaricis Thambugala,
Camporesi & K.D. Hyde
Tamarix g al lica Italy This study
Fomes platincola Speg. Tamarix africana Argentina Farr 1973
Fusarium oxysporum f. sp. radicis-
lycopersici Jarvis & Shoemaker
Tamarix nilotica Israel Rekah et al. 2001
Hendersonia tamaricis Cooke =
Coryneopsis tamaricis(Cooke)
Grove
Tamarix a fr ic ana,Ta marix gall ica Ukraine, Yugoslavia Saccardo (1885–1886),
Moskovets 1933
Hendersonia tamaricicola Brunaud Tamarix gallica France Saccardo and Sydow 1899
Homortomyces tamaricis Wijayaw.,
Camporesi & K.D. Hyde
Tamarix g al lica Italy Wijayawardene et al. 2014a, This
study
Hyphodontia sambuci (Pers.) J. Erikss. Tamarix sp. Spain Telleria et al. 1997
Inocutis rheades (Pers.) Fiasson &
Niemelä
Tamarix sp. China Tai 1979, Chen 2002
Inonotus tamaricis (Pat.) Maire Tamarix canariensis,Tama rix
ramosissima,Tam ar ix sp.
Canary Islands, Georgia,
Kazakhstan, Ukraine
Cannon et al. 1985,
Nakhutsrishvili 1986,Beltran
Tejera and Rodríguez-Armas
1999
Inonotus calcuttensis (Bose) S. Ahmad Tamarix articulata Pakistan Ahmad et al. 1997
Inonotus hispidus (Bull.) P. Karst. Ta ma rix aphyll a,Tamarix gallica,
Tamarix ramosissima,Tamarix
smyrnensis,Tam ar ix sp.,
Tamarix tetrandra
China, Georgia, Greece, Italy,
Pakistan, Russia, Spain,
Ukraine
Nakhutsrishvili 1986, Ahmad et al.
1997, Desjardin et al. 1997,
Telleria et al. 1997, Zervakis
et al. 1998, de Simone et al.
2011,Chen2002, Dudka et al.
2004, Rebriev et al. 2012
Fungal Diversity
Table 4 (continued)
Species Host Locality References
Keissleriella tamaricicola
Thambugala, Camporesi & K.D.
Hyde
Tamarix g al lica Italy This study
Leptosphaeria crozalsiana Maire Tam ar ix sp. Algeria Crane and Shearer 1991
Leptosphaeria hollosii Moesz Ta ma ri x ga ll ic a Hungary Crane and Shearer 1991
Leptosphaeria meridionalis D. Sacc. Tamarix g al lica Italy Crane and Shearer 1991
Leptosphaeria tamaricis (Grev.) Sacc. Tamarix g al lica,Tamarix
germanica,Tamarix hispida,
Tamarix sp.
England, Hungary, Pennsylvania,
U. K., U.S.A.
Ellis and Everhart 1892,Preston
1947, Anonymous 1960,
Cannon et al. 1985, Crane and
Shearer 1991
Macrophoma tamaricis Pass. Tamarix smyrnensis Georgia Nakhutsrishvili 1986
Massarina dubia (Wehm. & S.
Ahmad) S. Ahmad
Tamarix articulata Pakistan Ahmad 1969, Ahmad et al. 1997
Melanomma medium Sacc. & Speg. Tam ar ix sp. U.K. Cannon et al. 1985
Microdiplodia microsporella Allesch. Tam ar ix sp. Russia Rebriev et al. 2012
Mollisia ligni (Desm.) P. Karst. Ta ma ri x sp. Morocco, U.S.A. Anonymous 1960, Rieuf 1969
Monochaetia unicornis (Cooke &
Ellis) Sacc. & D. Sacc.
Tamarix g al lica Italy Guba 1961
Nectria cinnabarina (Tode) Fr. Tamarix ramosissima Russia Rebriev et al. 2012
Neofusicoccum luteum (Pennycook &
Samuels) Crous, Slippers & A.J.L.
Phillips
Tamarix sp. Italy This study
Neomicrosphaeropsis italica
Thambugala, Camporesi & K.D.
Hyde
Tamarix sp. Italy This study
Neomicrosphaeropsis novorossica
Thambugala, Bulgakov & K.D.
Hyde
Tamarix ramosissima Russia This study
Neomicrosphaeropsis rossica
Thambugala, Bulgakov & K.D.
Hyde
Tamarix ramosissima Russia This study
Neomicrosphaeropsis tamaricicola
(Wanas., Camporesi, E.B.G. Jones
& K.D. Hyde) Thambugala,
Wanasinghe & K.D. Hyde
Tamarix g al lica Italy Crous et al. 2014,Thisstudy
Paraepicoccum amazonense Matsush. Ta ma ri x sp. Italy This study
Peniophora meridionalis Boidin Tamarix canariensis Canary Islands Beltran Tejera and Rodríguez-
Armas 1999
Peniophora tamaricicola Boidin &
Malençon
Tamarix g al lica,Tamarix sp. Morocco, Portugal, Spain Rieuf 1970, Melo 1989, Telleria
1991,Telleriaetal.1997
Phaeosphaerella tamaricis (Hollós)
Tomilin
Tamarix a fric ana Hungaria Saccardo 1913
Phanerochaete martelliana (Bres.)
Parmasto
Tamarix sp. Spain Telleria et al. 1997
Phellinus torulosus (Petr.) Bourdot &
Galzin
Tamatix tetrandra Russia Isikov and Kuznetsov 1990
Phlebia subochracea (Alb. &
Schwein.) J. Erikss. & Ryvarden
Tamarix sp. Spain Telleria et al. 1997
Phoma africana Speg. Tam ar ix sp. Turkmenistan Koshkelova and Frolov 1973
Phoma rimincola Sacc. Tam arix sp. Georgia Nakhutsrishvili 1986
Phoma tamaricella Sacc. Tamarix sp. France, Turkmenistan, Ukraine Saccardo 1884, Koshkelova and
Frolov 1973,Helutaetal.1992,
Dudka et al. 2004
Phoma tamaricina Thüm. Tam ar ix g al li ca ,Tam ar ix sp. Austria, Turkmenistan, Ukraine Saccardo 1884, Moskovets 1933,
Koshkelova and Frolov 1973
Phoma tamaricis Sacc. Tamarix g al lica _ Saccardo 1880b
Phomopsis tamaricaria (Sacc.) Grove Ta marix africana France Saccardo 1884
Fungal Diversity
Table 4 (continued)
Species Host Locality References
Phymatotrichum omnivorum Duggar Tamarix sp. U.S.A. Anonymous 1960, French 1989
Pilidiella tamaricina S. Ahmad Tamarix articulata Pakistan Ahmad 1969, Ahmad et al. 1997
Platystomum compressum (Pers.)
Trevis
Tamarix sp. Morocco Rieuf 1970
Pleosphaeria echinata (Ellis & Everh.)
J.H. Mill.
Tamarix ramosissima Georgia Nakhutsrishvili 1986
Pleospora turkestanica Rehm Tam arix sp. Turkmenistan Koshkelova and Frolov 1973
Polyporus sulphureus (Bull.) Fr. Tamarix sp. Maryland Anonymous 1960
Polyporus tamaricis (Pat.) Sacc. & D.
Sacc.
Tamarix sp. Greece Pantidou 1973
Poria dictyopora (Cooke) Sacc. Ta ma ri x gallic a New Zealand Cunningham 1965
Paracamarosporium tamaricis
Thambugala, Camporesi & K.D.
Hyde
Tamarix g al lica Italy This study
Pseudocamarosporium propinquum
(Sacc.) Wijayaw., Camporesi &
K.D. Hyde
Tamarix g al lica Italy This study
Pyrenopeziza tamaricis (Roum.) Sacc. Tam ar ix sp. New York Anonymous 1960
Radulomyces confluens (Fr.) M.P.
Christ.
Tamarix sp. Spain Telleria et al. 1997
Roccella podocarpa Vai n . Tam ar ix sp. Southern Africa Doidge 1950
Rosellinia amblystoma Berl. & F. Sacc. Tamarix g al lica Spain Unamuno 1941
Rosellinia aquila (Fr.) Ces. & De Not. Tamarix gallic a var. canariensis Canary Islands de Urries 1957
Rosellinia byssiseda (Tode) J. Schröt. Tamarix g al lica Spain Gonzalez Fragoso 1916
Rosellinia pulveracea (Ehrh.) Fuckel Tamarix sp. Turkmenistan Koshkelova and Frolov 1973
Rosellinia rimincola Rehm Tamarix smyrnensis Georgia Dekanoidze 1984
Sinomyces fusoideus Yong Wang bis &
X.G. Zhang
Tamarix ramosissima China Wang et al. 2011
Sirococcus tamaricis (Syd.) S. Ahmad Tamarix articulata Pakistan Ahmad et al. 1997
Sphaerotheca humuli (DC.) Burrill Tama ri x sp. India Anonymous 1960
Splanchnonema loricatum (C. Tul. &
Tul.) M.E. Barr
Tamarix smyrnensis Georgia Dekanoidze 1984
Strickeria deflectens (P. Karst.)
Cooke
Tamarix sp. Turkmenistan Koshkelova and Frolov 1973
Strickeria sarmenticia (Sacc. & Speg.)
Cooke
Tamarix hispida Uzbekistan Gaponenko 1965
Teichospora obducens (Schumach.)
Fuckel
Tamarix articulata Pakistan Ahmad 1969, Ahmad et al. 1997
Tamaricicola muriformis Thambugala,
Camporesi & K.D. Hyde
Tamarix g al lica Italy This study
Teichospora pomiformis P. Karst. Tamarix articulata Pakistan Ahmad 1978
Trematosphaeria britzelmayriana
(Rehm) Sacc.
Tamarix articulata Pakistan Ahmad et al. 1997
Trichosporum fuscum (Link) Sacc. Tamarix gallica Spain Gonzalez Fragoso 1916
Triposporium echeveriae Tass i Tam ar ix sp. Morocco Rieuf 1970
Valsaria insitiva (Tode) Ces. & De Not. Tamarix sp. Morocco Rieuf 1970
Valsaria tamaricis Mundk. & S.
Ahmad
Tamarix articulata India, Pakistan Mundkur and Ahmad 1946,
Ahmad 1978, Ahmad et al.
1997
Vuilleminia macrospora (Bres.)
Hjortstam
Tamarix sp. Spain Telleria et al. 1997
Xanthochrous rheades (Pers.) Pat. Tamarix sp. China Teng 1996
*The records are taken from the literatures and thus may not be correct and the same taxon could be listed more than once. It would be
necessary to re-examine all collections if available to confirm their identities. Even the molecular data may be needed to establish their
correct names
Fungal Diversity
Paraphyses 1–2μm wide, numerous, aseptate, cylindrical.
Conidiophores reduced to conidiogenous cells, 10–48 × 1.7–
3μm(
x= 25 × 2.4 μm, n= 30), with supporting cells, hyaline,
percurrently proliferating at the apex of supporting cell.
Conidia 19–25.4 × 8.2–10.6 μm(
x= 22.7 × 9.7 μm,
n= 40), ellipsoid to subcylindrical, straight to slightly curved,
initially hyaline, after maturity golden brown to dark brown,
smooth-walled, 3-euseptate, apex obtuse.
Material examined: YUGOSLAVIA, Dalmatia, Rab
(Arbe) Island, on Tamarix africana Poir. (Tamaricaceae),
Jaap Otto, 3 May1912 (BPI 374366).
Notes:Coryneopsis tamaricis was introduced by Grove
(1932) to accommodate Hendersonia tamaricis. However,
currently Coryneopsis is considered as a synonym of
Seimatosporium Corda and the generic name Hendersonia
Sacc. is illegitimate (Index Fungorum 2016), and sequence
data are not available for any species in either Coryneopsis
or Hendersonia. Index Fungorum (2016) lists 1057
Hendersonia names and many have been transferred to
Stagonospora (including the type species Hendersonia
elegans Berk. ≡Stagonospora elegans (Berk.) Sacc. &
Traverso), and several have been transferred to other genera,
but a large number of species have not been adequately char-
acterized and have never been allocated to any other genus.
Coryneopsis tamaricis shows similar morphological traits to
Homortomyces and may represent another species of the ge-
nus Homortomyces.However,C.tamaricis differs from H.
tamaricis in having immersed, unilocular conidiomata and
longer conidiogenous cells. The species needs to be recollect-
ed and sequenced to confirm possible placement in the genus
Homortomyces.
Discussion
In this study, we describe a new family, two new genera and
ten new species. The family Homortomycetaceae is intro-
duced to accommodate Homortomyces with morphological
and molecular evidence. Sequence data of ribosomal RNA
genes (ITS, SSU and LSU) are available for Homortomyces
species. Protein coding gene sequences of Homortomyces spe-
cies and multigene phylogenetic analyses are required to estab-
lish a better phylogeny. Tamaricicola (Pleosporaceae)isintro-
duced as new monotypic genus, while Neomicrosphaeropsis is
established with four species and this genus may represent a
complex of cryptic species, which includes morphologically
similar, but phylogenetically different species. Paraepicoccum
amazonense, the type species of Paraepicoccum is reported for
the first time on Tam ari x spp. There are no sequence data
available for this genus in GenBank. Multi-gene phyloge-
netic analyses of newly generated sequence data in this
study show Paraepicoccum amazonense should be placed
in Pleosporineae,Pleosporales where it is closely related
to Camarosporium sensu stricto.
Tamari x species are distributed in various habitats in many
countries and are colonized by many fungal species (Table 4).
In this study we collected micro-fungal species associated
with different Tamarix species in Italy (Province of Forlì-
Cesena and Ravenna) and Russia (Rostov Region). More than
one hundred micro- and macro-fungal taxa reported on
Tamari x species are listed, although the actual number of fun-
gal taxa associated with Tamarix hosts may be higher. The
number of taxa that may be unique to Tam arix species is more
than 50, among them there are possibly, approximately 5–10
unique fungal species for each Ta ma ri x species. If this study
was expanded to other countries and Tamar ix hosts then many
additional fungi would probably be found. Presently many
fungal species reported from Tamarix species do not have
sequence data. Therefore, recollecting and sequencing these
taxa are essential to establish if the fungal records are correct.
Acknowledgments We are grateful to the Directors and Curators of BPI
and S for making specimens available for examination. The Mushroom
Research Foundation, Chiang Rai, Thailand and Guizhou Key Laboratory
of Agricultural Biotechnology, Guizhou Academy of Agricultural
Sciences, Guiyang, Guizhou Province, People’s Republic of China are
acknowledged for financial support and providing postgraduate scholar-
ship support to Kasun M. Thambugala. Kevin D. Hyde thanks the Chinese
Academy of Sciences (project number 2013T2S0030) for the award of
Visiting Professorship for Senior International Scientists at Kunming
Institute of Botany. The authors extend their sincere appreciations to the
Deanship of Scientific Research at King Saud University for its funding
this Prolific Research Group (PRG-1436-09). The authors would like to
thank the featured microbial resources and diversity investigation in
Southwest Karst area (2014FY120100). Kasun M. Thambugala thanks
Shaun Pennycook, Dhanushka Udayanga, Hiran A. Ariyawansa and
Kazuaki Tanaka for helpful comments and advice on the manuscript.
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