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Persoonia 44, 2020: 1– 40 ISSN (Online) 1878-9080
www.ingentaconnect.com/content/nhn/pimj https://doi.org/10.3767/persoonia.2020.44.01
RESEARCH ARTICLE
INTRODUCTION
Phylogenetic assignment of non-lichenised pyrenocarpous
ascomycetes forming brown muriform ascospores is a complex
and ongoing task. While fungi with such ascospores are rather
rare in Sordariomycetes, e.g., Dictyoporthe s.lat. (Jaklitsch &
Barr 1997), Stegonsporium (Voglmayr & Jaklitsch 2008, 2014)
in Diaporthales, Strickeria (Xylariales; Jaklitsch et al. 2016a),
Thyronectria (Hypocreales; Jaklitsch & Voglmayr 2014, Vogl-
mayr et al. 2016a) or Thyridium (Spatafora et al. 2006), they
are common in many families of Dothideomycetes, particularly
in several of the Pleosporales (Jaklitsch et al. 2016b). The
Cucurbitariaceae is one of these families. In contrast to genera
like, e.g., Thyronectria (Hypocreales; Jaklitsch & Voglmayr
2014, Voglmayr et al. 2016a) or Teichospora (Pleosporales;
Jaklitsch et al. 2016c), where both phragmospores and dictyo-
spores cluster in the same genus, all sexual morphs of the
Cucurbitariaceae (Pleosporales) have dictyospores (Jaklitsch
et al. 2018). Other characters shared by all representatives
of this family are the presence of a subiculum and phoma- or
pyrenochaeta-like asexual morphs, although these characters
may occur in several other families, too (Jaklitsch et al. 2018,
Valenzuela-Lopez et al. 2018). Several species of Cucurbitaria
with no or other asexual morphs have been recently removed to
different families of Pleosporales, e.g., Coniothyriaceae (Cucur-
bitaria varians; Crous & Groenewald 2017), Camarosporidiella-
ceae (most cucurbitaria-like species on fabaceous hosts;
Wanasinghe et al. 2017a), Melanommataceae (e.g., C. obdu-
cens; Jaklitsch & Voglmayr 2017) or Nectriaceae (C. bicolor in
Thyronectria; Checa et al. 2015). In a foregoing publication,
Jaklitsch et al. (2018) redefined the scope of the Cucurbitaria-
ceae and included the generic type of Fenestella, F. fenestrata,
by redescription, illustration, lecto- and epitypification and DNA
data. Other fenestella-like species were included in that work
as the new genera Cucitella, Parafenestella, Protofenestella
and Seltsamia.
After the original publication of Fenestella by Tulasne & Tulasne
(1863), who recognised three species in the genus including
F. princeps, a synonym of F. fenestrata (see Jaklitsch et al.
2018), 52 additional species names were created in the genus.
Eleven names including Fenestella bipapillata (Jaklitsch &
Barr 1997) and Fenestella frit (see Jaklitsch et al. 2018) have
been removed to other genera or they, among others, are not
interpretable, because no type material exists (for more data
see notes to species and Discussion). Barr (1990) recognised
eight species in Fenestella occurring in North America, which
she keyed out and described morphologically. She also gave
a detailed diagnosis of the genus Fenestella recognising its
fungicolous habit. However, she subsumed American fungi
under European Fenestella names without having seen type
material of most of them. As a result, several of her taxonomic
interpretations and conclusions are either erratic or too broad.
A definition of what fenestelloid fungi are is difficult, particularly
when compared to other members of the Cucurbitariaceae. The
main character apart from a more marked tendency to form val-
soid groups or pseudostromatic pustules, are the ascospores,
whose septa are variable in number and often difficult to count
due to incompleteness, dense insertion and apparent oblique
or shifted superposition in sectional view. This character is
Fenestelloid clades of the Cucurbitariaceae
W.M. Jaklitsch1,2, H. Voglmayr1,2
Key words
Cucurbitaria
Dothideomycetes
multigene phylogenetic analysis
new taxa
Phoma
Pleosporales
Pyrenochaeta
1 Institute of Forest Entomology, Forest Pathology and Forest Protection,
Department of Forest and Soil Sciences, BOKU-University of Natural
Resources and Life Sciences, Franz Schwackhöfer Haus, Peter-Jordan-
Straße 82/ I, 1190 Vienna, Austria;
corresponding author e-mail: walter.jaklitsch@univie.ac.at.
2 Division of Systematic and Evolutionary Botany, Department of Botany
and Biodiversity Research, University of Vienna, Rennweg 14, 1030 Wien,
Austria.
Abstract Fresh collections and their ascospore and conidial isolates backed up by type studies and molecular
phylogenetic analyses of a multigene matrix of partial nuSSU-, complete ITS, partial LSU rDNA, rpb2, tef1 and tub2
sequences were used to evaluate the boundaries and species composition of Fenestella and related genera of the
Cucurbitariaceae. Eight species, of which five are new, are recognised in Fenestella s.str., 13 in Parafenestella with
eight new species and two in the new genus Synfenestella with one new species. Cucurbitaria crataegi is combined
in Fenestella, C. sorbi in Synfenestella, Fenestella faberi and Thyridium salicis in Parafenestella. Cucurbitaria
subcaespitosa is distinct from C. sorbi and combined in Neocucurbitaria. Fenestella minor is a synonym of Valsa
tetratrupha, which is combined in Parafenestella. Cucurbitaria marchica is synonymous with Parafenestella salicis,
Fenestella bavarica with S. sorbi, F. macrospora with F. media, and P. mackenziei is synonymous with P. faberi, and
the latter is lectotypified. Cucurbitaria sorbi, C. subcaespitosa and Fenestella macrospora are lecto- and epitypified,
Cucurbitaria crataegi, Fenestella media, F. minor and Valsa tetratrupha are epitypified in order to stabilise the
names in their phylogenetic positions. A neotype is proposed for Thyridium salicis. A determinative key to species
is given. Asexual morphs of fenestelloid fungi are phoma-like and do not differ from those of other representatives
of the Cucurbitariaceae. The phylogenetic structure of the fenestelloid clades is complex and can only be resolved
at the species level by protein-coding genes, such as rpb2, tef1 and tub2. All fungal species studied here occur,
as far as has been possible to determine, on members of Diaporthales, most frequently on asexual and sexual
morphs of Cytospora.
Article info Received: 10 December 2018; Accepted: 13 February 2019; Published: 27 May 2019.
2Persoonia – Volume 44, 2020
Allocucurbitaria botulispora Superficial tissue CBS 142452 holo LT592932 LN907416 – LT593070 – LT593001
Astragalicola amorpha Astragalus angustifolius CBS 142999 = C227a holo MF795753 MF795753 – MF795795 MF795842 MF795883
Cucitella opali Acer opalus CBS 142405 = FV holo MF795754 MF795754 MF795837 MF795796 MF795843 MF795884
Cucurbitaria berberidis Berberis vulgaris ssp. atropurpurea C39 MF795755 MF795755 – MF795797 MF795844 MF795885
Berberis vulgaris CB MF795757 MF795757 – MF795799 MF795846 MF795887
Berberis vulgaris CBS 130007 = CB1 epi MF795758 MF795758 – MF795800 – –
Berberis sp. CBS 142401 = C241 MF795756 MF795756 – MF795798 MF795845 MF795886
Cucurbitaria oromediterranea Berberis aetnensis C265 MF795762 MF795762 – MF795804 MF795850 MF795891
Berberis hispanica C29 MF795759 MF795759 – MF795801 MF795847 MF795888
Berberis hispanica C86 MF795760 MF795760 – MF795802 MF795848 MF795889
Berberis cretica CB2 MF795763 MF795763 – MF795805 MF795851 MF795892
Berberis hispanica CB3 MF795764 MF795764 – MF795806 MF795852 –
Berberis cretica CBS 142399 = C229 holo MF795761 MF795761 – MF795803 MF795849 MF795890
Fenestella crataegi Crataegus monogyna C287 MK356281 MK356281 – – MK357554 MK357598
Crataegus monogyna CBS 144857 = C314 epi MK356282 MK356282 – MK357512 MK357555 MK357599
Fenestella fenestrata Alnus glutinosa CBS 143001 = FP9 epi MF795765 MF795765 – MF795807 MF795853 MF795893
Fenestella gardiennetii Acer saccharum CBS 144859 = FM holo MK356283 MK356283 – MK357513 MK357556 MK357600
Fenestella granatensis Acer granatense CBS 144854 = C279 holo MK356284 MK356284 – MK357514 MK357557 MK357601
Fenestella media Corylus avellana CBS 144860 = FP epi MK356285 MK356285 MK356326 MK357515 MK357558 MK357602
Carpinus orientalis FCO MK356286 MK356286 – MK357516 MK357559 –
Corylus avellana FP1 MK356287 MK356287 – MK357517 MK357560 MK357603
Acer pseudoplatanus FP3 MK356288 MK356288 – MK357518 MK357561 MK357604
Castanea sativa FP7 MK356289 MK356289 – MK357519 MK357562 MK357605
Tilia cordata FP10 MK356290 MK356290 – MK357520 MK357563 MK357606
Fenestella parafenestrata Quercus robur CBS 144856 = C306 holo MK356291 MK356291 – MK357521 MK357564 MK357607
Salix sp. C317 MK356292 MK356292 – MK357522 MK357565 MK357608
Fenestella subsymmetrica Acer campestre CBS 144861 = FP6 holo MK356297 MK356297 – MK357525 MK357569 MK357610
Juglans regia C285 MK356293 MK356293 – MK357523 MK357566 –
Juglans regia C286 MK356294 MK356294 – – MK357567 –
Juglans regia C286x MK356295 MK356295 – – – –
Corylus avellana FP4 MK356296 MK356296 – MK357524 MK357568 MK357609
Salix caprea FP8 MK356298 MK356298 – MK357526 MK357570 MK357611
Fenestella viburni Viburnum lantana CBS 144863 = FVL holo MK356300 MK356300 – MK357528 MK357572 MK357613
Viburnum lantana FP2 MK356299 MK356299 – MK357527 MK357571 MK357612
Neocucurbitaria acanthocladae Genista acanthoclada CBS 142398 = C225 holo MF795766 MF795766 – MF795808 MF795854 MF795894
Neocucurbitaria acerina Acer pseudoplatanus C26a MF795767 MF795767 – MF795809 MF795855 MF795895
Acer pseudoplatanus CBS 142403 = C255 MF795768 MF795768 – MF795810 MF795856 MF795896
Neocucurbitaria aetnensis Genista aetnensis CBS 142404 = C261 holo MF795769 MF795769 – MF795811 MF795857 MF795897
Genista aetnensis C270 MF795770 MF795770 – MF795812 MF795858 MF795898
Neocucurbitaria aquatica Sea water CBS 297.74 holo LT623221 EU754177 – LT623278 – LT623238
Neocucurbitaria cava Unknown CBS 115979 AY853248 EU754198 – LT623273 – LT623234
Wheat-field soil CBS 257.68 epi JF740260 EU754199 – LT717681 – KT389844
Neocucurbitaria cinereae Genista cinerea CBS 142406 = KU9 holo MF795771 MF795771 – MF795813 MF795859 MF795899
Neocucurbitaria cisticola Cistus monspeliensis CBS 142402 = C244 holo MF795772 MF795772 – MF795814 MF795860 MF795900
Neocucurbitaria hakeae Hakea sp. CBS 142109 = CPC 28920 holo KY173436 KY173526 – KY173593 – KY173613
Neocucurbitaria irregularis Subcutaneous tissue CBS 142791 holo LT592916 LN907372 – LT593054 – LT592985
Neocucurbitaria juglandicola Quercus rubra C316 MK356301 MK356301 – MK357529 MK357573 MK357614
Juglans regia CBS 142390 = BW6 holo MF795773 MF795773 – MF795815 MF795861 MF795901
Neocucurbitaria keratinophila Man corneal scrapings CBS 121759 holo EU885415 LT623215 – LT623275 – LT623236
Neocucurbitaria populi Populus sp. CBS 142393 = C28 holo MF795774 MF795774 – MF795816 MF795862 MF795902
Neocucurbitaria quercina Quercus robur CBS 115095 neo LT623220 GQ387619 GQ387558 LT623277 – LT623237
Table 1 Isolates and accession numbers used in the phylogenetic analyses. Isolates/sequences in bold were isolated/sequenced in the present study.
GenBank accession numbers
Taxon Host/ substrate Strain Ex-type status ITS LSU SSU rpb2 tef1 tub2
3
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Neocucurbitaria rhamni Rhamnus frangula CBS 142391 = C1 epi MF795775 MF795775 MF795838 MF795817 MF795863 –
Rhamnus frangula C112 MF795776 MF795776 – MF795818 MF795864 MF795903
Rhamnus frangula C133 MF795777 MF795777 – MF795819 MF795865 MF795904
Rhamnus frangula C190 MF795778 MF795778 – MF795820 MF795866 –
Rhamnus saxatilis C277 MF795779 MF795779 – MF795821 MF795867 MF795905
Neocucurbitaria rhamnicola Rhamnus lycioides CBS 142396 = C185 holo MF795780 MF795780 – MF795822 MF795868 MF795906
Rhamnus alaternus KRx MF795781 MF795781 – MF795823 MF795869 MF795907
Neocucurbitaria rhamnioides Rhamnus saxatilis ssp. prunifolius C222 MF795783 MF795783 MF795839 MF795825 MF795871 MF795909
Rhamnus saxatilis ssp. prunifolius C223 MF795784 MF795784 – MF795826 MF795872 MF795910
Rhamnus myrtifolius CBS 142395 = C118 holo MF795782 MF795782 – MF795824 MF795870 MF795908
Neocucurbitaria ribicola Ribes rubrum CBS 142394 = C55 holo MF795785 MF795785 MF795840 MF795827 MF795873 MF795911
Ribes rubrum C155 MF795786 MF795786 – MF795828 MF795874 MF795912
Neocucurbitaria unguis-hominis Agapornis sp. Lung CBS 111112 LT623222 GQ387623 – LT623279 – LT623239
Neocucurbitaria vachelliae Vachellia gummifera CBS 142397 = C192 holo MF795787 MF795787 – MF795829 MF795875 MF795913
Paracucurbitaria italica Olea europaea CBS 234.92 holo LT623219 EU754176 EU754077 LT623274 – LT623235
Paracucurbitaria riggenbachii Fraxinus excelsior with bacterial canker CBS 248.79 holo LT903672 GQ387608 – LT903673 – LT900365
Parafenestella alpina Cotoneaster integerrimus CBS 145263 = C198 holo MK356302 MK356302 – MK357530 MK357574 MK357615
Salix appendiculata C249 MK356303 MK356303 – MK357531 MK357575 MK357616
Parafenestella austriaca Rosa canina CBS 145262 = C152 holo MK356304 MK356304 – MK357532 MK357576 MK357617
Parafenestella germanica Corylus avellana CBS 145267 = C307 holo MK356305 MK356305 – MK357533 MK357577 MK357618
Parafenestella parasalicum Salix cinerea CBS 145271 = C318 holo MK356306 MK356306 – MK357534 MK357578 MK357619
Parafenestella pseudoplatani Acer pseudoplatanus CBS 142392 = C26 holo MF795788 MF795788 – MF795830 MF795876 MF795914
Parafenestella pseudosalicis Salix cf. alba CBS 145264 = C301 holo MK356307 MK356307 – MK357535 MK357579 MK357620
Parafenestella rosacearum Pyracantha coccinea CBS 145268 = C309 holo MK356311 MK356311 – MK357539 MK357583 MK357624
Pyrus communis C203 MK356308 MK356308 – MK357536 MK357580 MK357621
Crataegus monogyna C269 MK356309 MK356309 – MK357537 MK357581 MK357622
Pyrus communis C283 MK356310 MK356310 – MK357538 MK357582 MK357623
Rosa canina C315 MK356312 MK356312 – MK357540 MK357584 MK357625
Sorbus aria C320 MK356315 MK356315 – MK357543 MK357587 –
Prunus domestica CBS 145272 = FP11 MK356314 MK356314 – MK357542 MK357586 MK357627
Rosa canina FM1 MK356313 MK356313 MK356327 MK357541 MK357585 MK357626
Parafenestella salicis Salix alba CBS 145270 = C313 neo MK356317 MK356317 – MK357545 MK357589 MK357629
Salix alba C303 MK356316 MK356316 – MK357544 MK357588 MK357628
Parafenestella salicum Salix alba CBS 145269 = C311 holo MK356318 MK356318 – MK357546 MK357590 MK357630
Parafenestella tetratrupha Alnus glutinosa CBS 145266 = C304 epi MK356319 MK356319 – MK357547 MK357591 MK357631
Parafenestella vindobonensis Salix babylonica CBS 145265 = C302 holo MK356320 MK356320 – MK357548 MK357592 MK357632
Protofenestella ulmi Ulmus minor CBS 143000 = FP5 holo MF795791 MF795791 – MF795833 MF795879 MF795915
Pyrenochaeta nobilis Laurus nobilis leaves CBS 407.76 = AFTOL-ID 1856 neo MF795792 MF795792 DQ898287 MF795834 MF795880 MF795916
Pyrenochaetopsis americana Unknown UTHSC DI16-225 holo LT592912 LN907368 – LT593050 – LT592981
Pyrenochaetopsis botulispora Respiratory tract CBS 142458 holo LT592946 LN907441 – LT593085 – LT593015
Pyrenochaetopsis confluens Deep tissue /fluids CBS 142459 holo LT592950 LN907446 – LT593089 – LT593019
Pyrenochaetopsis globosa Superficial tissue CBS 143034 holo LT592934 LN907418 – LT593072 – LT593003
Pyrenochaetopsis leptospora Secale cereale CBS 101635 epi MF795793 MF795793 MF795841 MF795835 MF795881 MF795917
Pyrenochaetopsis uberiformis Superficial tissue CBS 142461 holo LT592935 LN907420 – LT593074 – LT593004
Seltsamia ulmi Ulmus glabra CBS 143002 = L150 holo MF795794 MF795794 MF795794 MF795836 MF795882 MF795918
Synfenestella pyri Pyrus communis CBS 144855 = C297 holo MK356321 MK356321 – MK357549 MK357593 MK357633
Synfenestella sorbi Sorbus aucuparia C298 MK356325 MK356325 – MK357553 MK357597 MK357636
Sorbus aucuparia CBS 144858 = C196 MK356324 MK356324 – MK357552 MK357596 MK357635
Sorbus aucuparia CBS 144862 = FR epi MK356322 MK356322 MK356328 MK357550 MK357594 MK357634
Sorbus aucuparia FRa MK356323 MK356323 – MK357551 MK357595 –
Table 1 (cont.)
GenBank accession numbers
Taxon Host/ substrate Strain Ex-type status ITS LSU SSU rpb2 tef1 tub2
4Persoonia – Volume 44, 2020
shared with the morphologically rather pleomassariaceous
genus Seltsamia (Jaklitsch et al. 2018), whose ascospores have
an indefinitely swelling, bipartite sheath. A similar situation is
found in Fenestella as shown below for F. granatensis, where
the ascospore sheath swells however in a limited manner.
Other unrelated, non-lichenised pyrenocarpous fungi on or in
wood and bark having ascospores with many transverse and
longitudinal eusepta in more or less cylindrical, fissitunicate asci
are Aigialus, differing from fenestelloid fungi, e.g., in different
ecology, as ascomata are immersed in submerged wood of
mangroves in marine environments (Kohlmeyer & Schatz 1985),
Decaisnella and Karstenula in the very wide concept of Barr
(1990), which, e.g., lack a subiculum and are not associated
with Diaporthales, or Ostreichnion, which produces conchate,
superficial ascomata on wood (Boehm et al. 2009).
Here we take a detailed look into the taxonomy and phylo-
genetic structure of fenestelloid fungi described from Europe
on woody hosts, from which fresh material was available for
study. These fungi include several species originally described
in Fenestella, Cucurbitaria or Thyridium, and cluster in three
clades representing the three genera Fenestella, Parafenestella
and Synfenestella.
MATERIALS AND METHODS
Isolates and specimens
All isolates used in this study originated from ascospores or
conidia (where noted) of fresh specimens. Numbers of strains
including NCBI GenBank accession numbers of gene sequen-
ces used to compute the phylogenetic trees are listed in Table
1. Strain acronyms other than those of official culture collections
are used here primarily as strain identifiers throughout the work.
Representative isolates have been deposited at the Westerdijk
Fungal Biodiversity Centre, Utrecht, The Netherlands (CBS
culture collection). Details of the specimens used for morpho-
logical investigations are listed in the Taxonomy section under
the respective descriptions. Herbarium acronyms are accord-
ing to Thiers (2018). Freshly collected specimens have been
deposited in the Fungarium of the Department of Botany and
Biodiversity Research, University of Vienna (WU).
Culture preparation and phenotype analysis
Cultures were prepared and maintained as described previously
(Jaklitsch 2009) except that CMD (CMA; Sigma, St Louis, Mis-
souri; supplemented with 2 % (w/v) D(+)-glucose-monohydrate)
or 2 % malt extract agar (MEA; 2 % w/v malt extract, 2 % w/v
agar-agar; Merck, Darmstadt, Germany) was used as the iso-
lation medium. Cultures used for the study of asexual morph
micro-morphology were grown on CMD or 2 % MEA at 22 ± 3 °C
in darkness. Microscopic observations were made in tap water
except where noted. Morphological analyses of microscopic
characters were carried out as described by Jaklitsch (2009).
Methods of microscopy included stereomicroscopy using a
Nikon SMZ 1500 and Nomarski differential interference contrast
(DIC) using the compound microscopes Nikon Eclipse E600 or
Zeiss Axio Imager.A1 equipped with a Zeiss Axiocam 506 colour
digital camera. Images and data were gathered using a Nikon
Coolpix 4500 or a Nikon DS-U2 digital camera and measured
by using the NIS-Elements D v. 3.0 or 3.22.15 or Zeiss ZEN
Blue Edition software packages. Some images obtained by
using the Nikon interference contrast may be slightly too dark.
For certain images of ascomata the stacking software Zerene
Stacker v. 1.04 (Zerene Systems LLC, Richland, WA, USA)
was used. Measurements are reported as maxima and minima
in parentheses and the mean plus and minus the standard
deviation of a number of measurements given in parentheses.
DNA extraction and sequencing methods
The extraction of genomic DNA was performed as reported pre-
viously (Voglmayr & Jaklitsch 2011, Jaklitsch et al. 2012) using
the DNeasy Plant Mini Kit (QIAgen GmbH, Hilden, Germany).
The following loci were amplified and sequenced: the terminal 3’
end of the small subunit nuclear ribosomal DNA (nSSU rDNA),
the complete internally transcribed spacer region (ITS1-5.8S-
ITS2) and a c. 900 bp fragment of the large subunit nuclear
ribosomal DNA (nLSU rDNA), amplified and sequenced as a
single fragment with primers V9G (De Hoog & Gerrits van den
Ende 1998) and LR5 (Vilgalys & Hester 1990); a c. 1.0–1.4
kb fragment at the 5’ end of the nSSU rDNA with primers SL1
(Landvik et al. 1997) and NSSU1088 (Kauff & Lutzoni 2002);
a c. 1.2 kb fragment of the RNA polymerase II subunit 2 (rpb2)
gene with primers fRPB2-5f and fRPB2-7cr (Liu et al. 1999) or
dRPB2-5f and dRPB2-7r (Voglmayr et al. 2016a); a c. 1.3–1.5
kb fragment of the translation elongation factor 1-alpha (tef1)
gene with primers EF1-728F (Carbone & Kohn 1999) and
TEF1LLErev (Jaklitsch et al. 2005) or EF1-2218R (Rehner &
Buckley 2005); and a c. 0.7 kb fragment of the beta tubulin
(tub2) gene with primers T1 (O’Donnell & Cigelnik 1997) or
T1HV (Voglmayr et al. 2016b) and BtHV2r (Voglmayr et al.
2016b, 2017). PCR products were purified using an enzymatic
PCR clean-up (Werle et al. 1994) as described in Voglmayr
& Jaklitsch (2008). DNA was cycle-sequenced using the ABI
PRISM Big Dye Terminator Cycle Sequencing Ready Reac-
tion Kit v. 3.1 (Applied Biosystems, Warrington, UK) with the
same primers as in PCR; in addition, primers ITS4 (White et
al. 1990), LR2R-A (Voglmayr et al. 2012) and LR3 (Vilgalys &
Hester 1990) were used for the ITS-LSU region. In some cases
the tef1 was cycle-sequenced with internal primers TEF1_INTF
(forward; Jaklitsch 2009) and TEF1_INT2 (reverse; Voglmayr &
Jaklitsch 2017). Sequencing was performed on an automated
DNA sequencer (3730xl Genetic Analyzer, Applied Biosystems).
Analysis of sequence data
For the phylogenetic analyses, a combined matrix of nSSU-ITS-
LSU rDNA, rpb2, tef1 and tub2 sequences was produced. The
newly generated sequences were complemented with GenBank
sequences of Cucurbitariaceae from Jaklitsch et al. (2018), and
sequences of six taxa of Pyrenochaetopsis (Pyrenochaetopsi-
daceae) were added as outgroup according the results of the
phylogenetic analyses of Jaklitsch et al. (2018). All alignments
were produced with the server version of MAFFT (www.ebi.
ac.uk/Tools/mafft), checked and refined using BioEdit v. 7.2.6
(Hall 1999). Large insertions sometimes present in the SSU and
at the terminal 3’ end of the SSU of the partial SSU-ITS-LSU
fragment were removed from the alignments.
The combined matrix contained 5 707 nucleotide characters,
represented by 1 688 from the partial SSU-ITS-LSU, 999 from
the SSU, 1 067 from rpb2, 1 258 from tef1 and 695 from tub2.
Maximum parsimony (MP) analysis was performed using a
parsimony ratchet approach. For this, a nexus file was prepared
using PRAP v. 2.0b3 (Müller 2004), implementing 1 000 ratchet
replicates with 25 % of randomly chosen positions upweighted
to 2, which was then run with PAUP v. 4.0a164 (Swofford
2002). The resulting best trees were then loaded in PAUP and
subjected to heuristic search with TBR branch swapping (MUL-
TREES option in effect, steepest descent option not in effect).
Bootstrap analysis with 1 000 replicates was performed using 5
rounds of replicates of heuristic search with random addition of
sequences and subsequent TBR branch swapping (MULTREES
option in effect, steepest descent option not in effect) during
each bootstrap replicate. In all MP analyses molecular char-
acters were unordered and given equal weight; analyses were
performed with gaps treated as missing data; the COLLAPSE
5
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fenestella subsymmetrica FP4
Fenestella subsymmetrica C285
Fenestella subsymmetrica C286x
Fenestella subsymmetrica C286
Fenestella subsymmetrica FP8
Fenestella subsymmetrica CBS 144861T
Fenestella media FP3
Fenestella media CBS 144860T
Fenestella media FCO
Fenestella media FP10
Fenestella media FP7
Fenestella media FP1
Fenestella viburni FP2
Fenestella viburni CBS 144863T
Fenestella parafenestrata CBS 144856T
Fenestella parafenestrata C317
Fenestella gardiennetii CBS 144859
T
Fenestella granatensis CBS 144854T
Fenestella fenestrata CBS 143001T
Fenestella crataegi CBS 144857T
Fenestella crataegi C287
Parafenestella rosacearum FP11
Parafenestella rosacearum C320
Parafenestella rosacearum CBS 145268T
Parafenestella rosacearum C203
Parafenestella rosacearum FM1
Parafenestella rosacearum C315
Parafenestella rosacearum C283
Parafenestella rosacearum C269
Parafenestella pseudoplatani CBS 142392T
Parafenestella germanica CBS 145267T
Parafenestella austriaca CBS 145262T
Parafenestella salicum CBS 145269T
Parafenestella parasalicum CBS 145271T
Parafenestella tetratrupha CBS 145266T
Parafenestella salicis C303
Parafenestella salicis CBS 145270T
Parafenestella pseudosalicis CBS 145264T
Parafenestella vindobonensis CBS 145265T
Parafenestella alpina CBS 145263T
Parafenestella alpina C249
Synfenestella sorbi C196
Synfenestella sorbi C298
Synfenestella sorbi CBS 144862T
Synfenestella sorbi FRa
Synfenestella pyri CBS 144855T
Seltsamia ulmi CBS 143002T
Allocucurbitaria botulispora CBS 142452T
Astragalicola amorpha CBS 142999T
Neocucurbitaria rhamnioides C222
Neocucurbitaria rhamnioides C223
Neocucurbitaria rhamnioides CBS 142395T
Neocucurbitaria rhamnicola CBS 142396T
Neocucurbitaria rhamnicola KRx
Neocucurbitaria rhamni C133
Neocucurbitaria rhamni C190
Neocucurbitaria rhamni C112
Neocucurbitaria rhamni CBS 142391T
Neocucurbitaria rhamni C277
Neocucurbitaria ribicola CBS 142394T
Neocucurbitaria ribicola C155
Neocucurbitaria acerina C26a
Neocucurbitaria acerina C255
Neocucurbitaria irregularis CBS 142791T
Neocucurbitaria keratinophila CBS 121759T
Neocucurbitaria aquatica CBS 297.74
Neocucurbitaria unguis hominis CBS 111112
Neocucurbitaria vachelliae CBS 142397T
Neocucurbitaria juglandicola CBS 142390T
Neocucurbitaria juglandicola C316
Neocucurbitaria populi CBS 142393T
Neocucurbitaria cava CBS 115979
Neocucurbitaria cava CBS 257.68T
Neocucurbitaria cisticola CBS 142402T
Neocucurbitaria hakeae CPC 28920T
Neocucurbitaria aetnensis CBS 142404T
Neocucurbitaria aetnensis C270
Neocucurbitaria cinereae CBS 142406T
Neocucurbitaria acanthocladae CBS 142398T
Neocucurbitaria quercina CBS 115095T
Cucurbitaria oromediterranea C86
Cucurbitaria oromediterranea CB3
Cucurbitaria oromediterranea C29
Cucurbitaria oromediterranea CBS 142399T
Cucurbitaria oromediterranea CB2
Cucurbitaria oromediterranea C265
Cucurbitaria berberidis CB
Cucurbitaria berberidis CBS 130007T
Cucurbitaria berberidis C39
Cucurbitaria berberidis C241
Paracucurbitaria riggenbachii CBS 248.79T
Paracucurbitaria italica CBS 234.92T
Cucitella opali CBS 142405
Protofenestella ulmi CBS 143000
Pyrenochaeta nobilis CBS 407.76
Pyrenochaetopsis uberiformis CBS 142461T
Pyrenochaetopsis globosa CBS 143034T
Pyrenochaetopsis americana UTHSC DI16 225T
Pyrenochaetopsis botulispora CBS 142458T
Pyrenochaetopsis leptospora CBS 101635
Pyrenochaetopsis confluens CBS 142459T
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Pyrenochaetopsi-
daceae
incertae sedis
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*
*
*
*
T
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T
Fig. 1 Phylogram of one of 33 MP trees 6 241 steps long (CI = 0.386, RI = 0.817), obtained by PAUP from an analysis of the combined matrix (SSU-ITS-LSU,
rpb2, tef1, tub2) of Cucurbitariaceae and Pyrenochaetopsidaceae, with the latter selected as outgroup. MP and ML bootstrap support above 50 % are given at
the first and second position, respectively, above or below the branches. Strains formatted in bold were isolated and sequenced in the current study; ex-type
strains are indicated by a superscript T. Nodes collapsed in the strict consensus of the 33 MP trees are marked by an asterisk (*).
6Persoonia – Volume 44, 2020
command was set to minbrlen. Maximum likelihood (ML)
analyses were performed with RAxML (Stamatakis 2006) as
implemented in raxmlGUI 1.5 (Silvestro & Michalak 2012),
using the ML + rapid bootstrap setting and the GTRGAMMA
substitution model with 1 000 bootstrap replicates. The matrix
was partitioned for the individual gene regions, and substitution
model parameters were calculated separately for them. For
evaluation and discussion of bootstrap support, values below
70 % were considered low, between 70 and 90 % medium/
moderate and above 90 % high.
RESULTS
Phylogenetic analyses
Of the 5 707 nucleotide characters of the combined matrix,
1 266 are parsimony informative (283 of ITS-LSU, 5 of SSU,
423 of rpb2, 302 of tef1 and 253 of tub2). Maximum parsimony
analyses revealed 33 MP trees 6 241 steps long, one of which
is shown as Fig. 1. Topologies of the MP trees were identical
except for one backbone node each within the Fenestella and
Parafenestella clades, and two backbone nodes within the
Neocucurbitaria clade (marked by asterisks in Fig. 1).
Like in the previous phylogenetic analyses of Jaklitsch et al.
(2018), many of the deeper nodes within Cucurbitariaceae were
unsupported or received only low support, while the genera
Cucurbitaria and Neocucurbitaria were highly supported. The
fenestelloid clade received maximum support and contained
three highly supported subclades here recognised as three
distinct genera: Fenestella, Parafenestella (both with maximum
support in MP and ML analyses) and Synfenestella (99 % MP
and ML bootstrap support). The sister group relationship of
Fenestella and Parafenestella received only low support (54 %
MP and 51 % ML), while most backbone nodes within the three
genera received high to maximum support (Fig. 1).
Taxonomy
Fenestella Tul. & C. Tul., Select. Fung. Carpol. (Paris): Xylariei-
Valsei-Spaeriei 2: 207. 1863, emend.
Type species. Fenestella fenestrata (Berk. & Broome) J. Schröt.
Fenestella fenestrata (Berk. & Broome) J. Schröt., in Cohn,
Krypt.-Fl. Schlesien (Breslau) 3.2(4): 435. 1897 (1908)
Basionym. Valsa fenestrata Berk. & Broome, Ann. Mag. Nat. Hist., ser.
III 3: 366. 1859.
Synonym. Fenestella princeps Tul. & C. Tul., Select. Fung. Carpol. (Paris)
2: 207. 1863.
Notes — See Jaklitsch et al. (2018) for description and typi-
fication of the genus and its type species. In that work it was
stated that all materials of F. fenestrata available for study
were more or less overmature, which made identification of the
fungal host difficult. The black encasement of ascomata was
interpreted as belonging to a Diaporthe sp. Considering that all
other species of the genus occur on Cytospora spp. (see below),
it appears probable that the stromatic encasement belonged to
a Cytospora sp. having a Leucostoma sexual morph.
The second specimen given in the protologue of F. fenestrata
was examined: England, Leicestershire, Orton Wood, on dead
twigs of Quercus robur, Mar. 1859, A. Bloxam (K(M) 233193;
as Valsa fenestrata). Although the plant host would suggest
Fenestella parafenestrata (see below), the fungus in this speci-
men is morphologically indistinguishable from F. fenestrata:
Pseudostromatic pustules are 1–4 mm diam, brown to black,
outside limited by a black stromatic line, ascomata 400–770 µm
diam, ostiolar areas 180–420 µm diam, asci are cylindrical and
ascospores (35–)43 –61(–74) × (14.5 –)16.5– 22.5(–27) µm, l/w
(2–)2.3 –3.1(– 3.5) (n = 30), ellipsoid to fusoid, very dark brown,
symmetric or asymmetric, with up to 16 distinct transverse and
7 longitudinal septa and up to 4 µm long hyaline apiculi. See
Fig. 6b1 for the illustration of an ascospore.
Fenestella crataegi (Niessl) Jaklitsch & Voglmayr, comb. nov.
— MycoBank MB829741; Fig. 2
Basionym. Cucurbitaria crataegi Niessl, Verh. Naturf. Vereins Brünn 10:
199. 1872.
Holotype. CzeCh RepubliC, near Brno, on dry branches of Crataegus
oxyacanthae (used at that time for C. monogyna), no date given, G. Niessl
(M-0281851). Epitype, here designated: AustRiA, Burgenland, Purbach, Pur-
bacher Heide, on Cytospora sp. on a branch of Crataegus monogyna, soc. Di-
plodia sp., 24 Mar. 2018, H. Voglmayr (WU 36987; MBT385683; ex-epitype
culture CBS 144857 = C314).
Ascomata (330–)368 – 507(–540) µm (n = 12) diam, globose,
subglobose to pyriform, immersed in groups of 0.9–2.4 mm
diam or length containing usually less than 10 individuals or
solitarily above Cytospora (Valsa) ascomata or conidiomata in
a single or two vertical layers, and erumpent from bark; asco-
mata individually surrounded by shiny, pale to dark brown,
thick-walled, 2–6.5 µm wide subicular hyphae connecting
them and sometimes forming pseudostromatic structures.
Ostiolar areas 90–180 µm diam, black, poorly differentiated
from the venter and inconspicuous, less commonly papillate.
Peridium c. 20–75 µm wide, pseudoparenchymatous, consist-
ing of (3.5–)5.5 –10(–14.5) µm (n = 30) wide cells, dark brown
outside, gradually paler, larger and thinner-walled to the inside;
the innermost layer ill-defined, variably comprising hyaline or
pale brown compressed cells. Hamathecium consisting of nu-
merous branched, 1–4 µm wide, apically free paraphyses. Asci
(214–)245 –295(– 317) × (20–)22– 26.7(–28) µm (n = 22), cylin-
drical to oblong or narrowly clavate, bitunicate, fissitunicate, with
an ocular chamber, a short stipe and simple or knob-like base,
containing (4–)8 ascospores in (obliquely) uniseriate, some-
times partly biseriate arrangement; unstable in water, stable
in 3 % KOH. Ascospores (31–) 36.5–45.5 (–54.5) × (11–)15–
19.5(– 23) µm, l/w (2–) 2.2– 2.6(–3) (n = 95), ellipsoid or broadly
fusoid, with 11–14(–16) transverse and 2– 4 longitudinal septa,
constricted at the median or nearly median primary septum with
upper part often wider than lower, thick-walled, first hyaline,
turning yellowish, finally medium to dark brown, in 3 % KOH
blackish brown; terminal part of terminal cells hyaline and
broadly or narrowly rounded, projecting to c. 3.5 µm, becoming
elongate upon germination.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 4 mm after 1 wk,
21 mm after 3 wk; colony circular, thick, dense, first white,
turning greyish olive, dark grey-brown and finally black, slightly
zonate, becoming velvety by aerial hyphae, odour indistinct.
Pycnidia developing on and around the plug submerged in
the agar to superficial, globose, c. 90–300 µm diam, oliva-
ceous to black, aggregating and confluent to large masses;
peridium pseudoparenchymatous, bearing hyaline to brown
hyphae and dark brown thick-walled setae 10–50 × 1.5 –5 µm;
releasing conidia as whitish to olivaceous turbid drops. Phial-
ides (4.5–)5.7– 8.7(–11) × (1.5–)2.2– 3.5(–4.2) µm (n = 30),
lageniform, ampulliform or subcylindrical, sessile or on short
few-celled conidiophores; conidia also formed on lateral pegs.
Conidia (3.2–)3.5 –4.7(–5.6) × (1.2 –)1.4 –1.8(–2.1) µm, l/w
(1.8–)2.2 – 3.2(– 4.3) (n = 51), cylindrical, oblong to ellipsoid,
1-celled, hyaline to pale greyish olivaceous, containing 2 sub-
terminal drops, smooth.
Habitat — On Cytospora sp. (sexual and asexual morphs)
on branches and twigs of Crataegus monogyna.
Distribution — Central Europe (Czech Republic, Austria).
7
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fig. 2 Fenestella crataegi. a– v. Sexual morph. a. Horizontal section through unevenly immersed ascomata; b– c. Immature ascomata immersed at the ostiolar
levels of the Cytospora morph (b) and the Valsa morph (c); d. peridium in vertical section; e. subicular hyphae below vertically sectioned peridium; f– h. asci
(in f young and opening); i. ascus apex surrounded by hamathecial threads; j– v. ascospores (j –n from fresh material; germinating in s – u); w– a1. asexual
morph from CMD at 22 °C; w. pycnidia with conidial drops; x. pycnidial seta; y. phialides; z–a1. conidia (greyish olivaceous in a1) (g– h, v. in 3 % KOH). a– i,
m, o– a1. WU 36987/CBS 144857 (C314); j–l, n. C287. — Scale bars: a– c = 300 µm; d– e, j– v, x = 10 µm; f– i = 25 µm; w = 150 µm; y– z = 5 µm; a1 = 3 µm.
8Persoonia – Volume 44, 2020
Fig. 3 Fenestella gardiennetii (WU 36986, CBS 144859 = FM). a–o. Sexual morph. a. Ascomata in face view; b. vertical section of ascoma on a Cytospora
pseudostroma; c. peridium in vertical section; d. subicular hyphae; e. ascus apex; f –h. asci; i–o. ascospores (i. initial stage); p. conidia from CMD at 22 °C;
(n– o. in 3 % KOH). — Scale bars: a = 300 µm; b = 500 µm; c–d, f– h = 25 µm; e, i–o = 10 µm; p = 5 µm.
Other materials examined. AustRiA, Burgenland, Purbach am See, Pur-
bacher Heide, on branch of Crataegus monogyna, 4 Feb. 2017, H. Voglmayr
(culture C287; specimen lost); Niederösterreich, Wolfsthal, grid square
7868/3, on corticated twigs of Crataegus monogyna, 1 Apr. 2000, W. Jaklitsch
W.J. 1434 (WU 37020).
Notes — There are two specimens of Niessl labelled Cu-
curbitaria crataegi in M. Specimen M-0281852 contains a
Parafenestella with ascospores (25–)26 – 30(–31) × (11.8–)
12.3–14(–14.3) µm, recognised as P. austriaca (see below). It
was collected in Rosenthal near Hütteldorf (Vienna, Austria),
thus it cannot be type material. Specimen M-0281851 was
collected at the type locality and contains a Diplodia (plus its
botryosphaeriaceous sexual morph) in excess and a Massarina
sp. (s.lat.) with bicellular hyaline ascospores. Also present are
a few pycnidia on subiculum, containing rod-like unicellular
hyaline conidia (3–)3.3 –4(– 4.4) × (1.2 –)1.3–1.6(–1.8) µm,
l/ w 2.1–2.9(– 3.7) (n = 30), on lageniform to subcylindrical
phialides. This is presumably the asexual morph of C. crataegi.
As the sexual morph is apparently used up and the speci-
men is depauperate, epitypification is essential. Von Niessl’s
(1872) measurements of ascospores lies at the lower end of
our measurements, but his illustrations strongly suggest that
our material represents this taxon, therefore we stabilise this
name by epitypification rather than describing a new species.
He compared his fungus with C. acervata, of which he had not
seen authentic material.
Fenestella gardiennetii Jaklitsch & Voglmayr, sp. nov. — Myco-
Bank MB829742; Fig. 3
Etymology. Named after its collector Alain Gardiennet.
Holotype. FRAnCe, 21, Longvic, Arboretum, on Cytospora sp. on twigs of
Acer saccharum, soc. Diplodia sp., 27 June 2013, A. Gardiennet (WU 36986;
ex-type culture CBS 144859 = FM).
Ascomata (390–)405 – 565(– 630) µm (n = 12) diam, depress-
ed subglobose to globose, immersed in valsoid groups or in
lines c. 0.7–2 mm long, also solitarily or in pairs, on and sur-
rounded by subiculum on effete Cytospora sp.; ascoma apex
mostly flat, black, sometimes partly covered by brown subicu-
lum. Ostioles 55–150 µm diam, central, papillate to conical,
black, periphysate. Subiculum consisting of subhyaline to dark
brown, thick-walled, 2–6 µm wide hyphae. Peridium 20 –75
µm, apically to 110 µm thick, pseudoparenchymatous, consis-
ting of thick-walled, dark brown cells (5–)7.5–14 (–17) µm
(n = 45) diam becoming gradually lighter towards the interior,
sometimes terminated inside by pale brown compressed cells.
Hamathecium consisting of richly branched 1–3.5 µm wide
threads in a gel matrix. Asci (176–)202 –243(– 263) × (20.5 –)
21–25.5(– 30) µm (n = 25), cylindrical to oblong, bitunicate,
fissitunicate, with an ocular chamber, a usually short stipe and
simple or knob-like base, containing 4–8 ascospores in (obli-
quely) uniseriate, sometimes partly biseriate arrangement. Asco-
spores (34–)36.5 – 45(– 49) × (13.5 –)15 –19(– 22.5) µm, l/ w
(1.9–)2.2 –2.6(–2.8) (n = 50), broadly ellipsoid to clavate, thick-
walled, first hyaline with 1–4 main transverse septa, usually
distinctly asymmetric with submedian primary septum and
smaller lower part, developing additional septa and turning
dark brown, when mature with 11–16 distinct transverse and
9
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
3–5 longitudinal septa, distinctly constricted at the primary
septum; surface verruculose; ends of terminal cells concolor-
ous or hyaline, often narrowed and projecting as 1–2 µm long
apiculi; in 3 % KOH ascospores turning blackish brown when
mature, apiculi remaining hyaline.
Culture characteristics and asexual morph in culture — Asco-
spores germinating simultaneously from many cells. Colony
radius on CMD at 22 °C in the dark 10 mm after 1 wk, 35 mm
after 4 wk; colony circular, thick and dense, first white, turning
dull olivaceous brown to brownish grey from the centre, zonate;
aerial hyphae forming loose greyish mesh; reverse dark grey to
black; odour indistinct. Pycnidia formed after 4 d, mostly remain-
ing submerged in agar and densely aggregated on and around
the inoculation plug, globose, 120–210 µm diam, greenish,
turning black; peridium pseudoparenchymatous, consisting
of isodiametric cells with irregularly distributed dark brown
pigment, surrounded by brown hyphae. Conidia amassing in
pale greyish mucous drops, (3.3–)3.5 –4.5(–5) × (1.1–)1.2–1.5
(–1.7) µm, l/w (2 –)2.5 –3.4(–4.4) (n = 30), cylindrical, straight to
slightly curved, 1-celled, smooth, containing 2 minute guttules.
Habitat — On Cytospora sp. (sexual and asexual morphs)
on Acer saccharum.
Distribution — Europe (France), only known from the type
locality; possibly occurring also in North America.
Notes — Fenestella gardiennetii may have travelled on its
host from North America and may thus occur also there. It is
closely related to F. granatensis, which occurs on Acer gra-
natense and differs from the former by more distinctly clavate
ascospores that are surrounded by a mucous sheath.
Fig. 4 Fenestella granatensis. a –r. Sexual morph (WU 36985). a. Cytospora (Valsa) pseudostroma with laterally inserted ascoma; b. vertical section of
3 ascomata sitting on a Cytospora (Valsa) ascoma; c. peridium in vertical section; d. subicular hyphae; e. free end of a paraphysis with sheath; f. apex of im-
mature ascus; g–h. asci (immature in g); i –r. ascospores (i. initial stage; j– l. young; r. compressed); s– x. asexual morph (CBS 144854 (C279) from CMD at
22 °C); s. pycnidia; t– u. phialides; v– x. conidia (d, h, k– l. in 3 % KOH). — Scale bars: a = 500 µm; b = 300 µm; c– d, f, i–r = 10 µm; e, t–u = 5 µm; g–h = 20
µm; s = 150 µm; v– x = 3 µm.
10 Persoonia – Volume 44, 2020
Fenestella granatensis Jaklitsch & Voglmayr, sp. nov. — Myco-
Bank MB829743; Fig. 4
Etymology. Referring to the place of its occurrence, Granada, and its
host Acer granatense.
Holotype. spAin, Andalusia, Granada, La Zubia, Cerro del Trevenque,
near the Jardín Botanico de la Cortijuela, elev. 1600 m, on Cytospora sp.
on twig of Acer granatense, soc. Dictyoporthe sp., a melanommataceous
fungus, and Myriangium durieui, 14 May 2014, W. Jaklitsch & S. Tello (WU
36985; ex-type culture CBS 144854 = C279).
Ascomata (330–)358 – 636(– 900) µm (n = 20) diam, globose,
depressed subglobose to subpyriform, immersed and erumpent
through bark fissures, in valsoid groups of 1–10 typically around
ostiolar necks of Cytospora (Valsa) ascomata, forming pustules
0.9–2 mm diam; individually surrounded by whitish to dark
brown subiculum consisting of thick-walled, 2 –7 µm wide hy-
phae. Ostiolar structures short-cylindrical, hardly noticeable on
the surface or appearing as black dots 75–170(– 210) µm diam;
interior periphysate. Peridium c. 30–65 µm, apically to 100 µm
thick, pseudoparenchymatous, consisting of thick-walled, dark
brown cells (4.5–)6 –11.5(–16) µm (n = 40) diam becoming
gradually lighter towards the interior, sometimes terminated
inside by pale brown compressed cells. Hamathecium consis-
ting of numerous, richly branched, 1–3 µm wide, apically free
paraphyses in a dense matrix. Asci (195–)222 –265(– 284) ×
(28.5–)31– 34(– 34.5) µm (n = 15), cylindrical, oblong or cla-
vate, bitunicate, fissitunicate, with an ocular chamber, a usu-
ally short stipe and simple or knob-like base, containing 4–8
ascospores in (obliquely) uni- to biseriate arrangement. Asco-
spores (36.5–)43 –53.5(– 60) × (13 –)17– 22.5(– 28.5) µm, l/w
(1.9–)2.2 –2.7(–3.4) (n = 100), clavate, fusoid to subellipsoid
with the upper part always wider than the lower, when young
hyaline to yellowish and with 1–7 transverse and 1–2 longitu-
dinal septa, eventually dark brown, with 12–17 transverse and
4–5 longitudinal septa; usually slightly constricted at the nearly
median primary septum; tips of end cells slightly lighter or con-
colorous; cells filled with oil drops, surface warted and appear-
ing fissured; surrounded by a bipartite, 1– 2 µm wide gelatinous
sheath swelling in water and terminally fraying out at as variably
shaped, appendage-like distortions or protuberances; in 3 %
KOH blackish brown, primary septum appearing more distinct,
other septa becoming less distinct, sheath swelling indefinitely.
Pycnidia of the presumed asexual morph with minute rod-like
hyaline unicellular conidia on lageniform to subulate phialides
on filiform conidiophores also present between ostiolar necks
of the Cytospora (Valsa) host.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 10–13 mm after 1 wk,
33 mm after 3 wk; colony circular, thick, dense, first white to
yellowish, later turning dark olivaceous brown to dark grey,
reverse black; aerial hyphae forming a dense brownish mat;
odour indistinct. Pycnidia appearing after 4 d in the colony
centre, numerous, separate or aggregating to larger complexes,
globose, 150 –270 µm diam, first hyaline to greenish, turning
olivaceous to black; conidia becoming released in whitish turbid
drops. Phialides sessile or formed terminally on cylindrical to
globose intercalary cells, (2.7–)5 –9(–11.3) × (1.5–)2.3–4 (–5)
µm (n = 20), lageniform, ampulliform to subglobose with a long
neck. Conidia formed on phialides and lateral pegs, (3–)3.5 –
4.8(– 6) × (1.4 –)1.6 –2.1(–2.7) µm, l/w (1.5 –)2– 2.7(– 3.1)
(n = 25), oblong, allantoid or narrowly ellipsoid, 1-celled, with
usually 2 subterminal drops, smooth.
Habitat — On Cytospora sp. (sexual and asexual morphs)
on Acer granatense.
Distribution — Europe (Spain), only known from the type
locality.
Notes — Fenestella granatensis is easily recognisable by
its clavate pleomassariaceous ascospores having a gelatinous
sheath and by its host, Acer granatense. As with several other
species of the fenestelloid clades, a few pycnidia are present
on the fungal host; always discrete, globose and collapsing
cupulate.
Fenestella media Tul. & C. Tul., Select. Fung. Carpol. (Paris)
2: 208. 1863. — Fig. 5
Synonym. Fenestella macrospora Fuckel, Jahrb. Nassauischen Vereins
Naturk. 25– 26: 313. 1871.
Typification. Holotype of Fenestella media. FRAnCe, Meudon (close to
Versailles; in the protologue: circa Versalias), Mar. 1860, parasitizing the
Cytospora state of Valsa salicis (as Cytospora fagaci (Bull.), salicicola) on
Salix alba (PC 0706651); donated by L.R. Tulasne to PC in 1873. Another
specimen extant in PC (PC 0706650) was collected in 1892, i.e., after its
first description. The material of the holotype contains numerous effete Cyto-
spora pseudostromata in bark and only few are infected by the Fenestella
with small pustules containing c. 1–4 ascomata. Lectotype of Fenestella
macrospora, here designated: GeRmAny, Hessen, Oestrich-Winkel, Reich-
artshausen, on twigs of Corylus avellana, L. Fuckel (G 00127659, from
Herbier Barbey-Boissier, Herbier Fuckel 1894; distributed as Fungi Rhenani
2328; MBT385684). Epitype of Fenestella media and F. macrospora, here
designated: AustRiA, Kärnten, St. Margareten im Rosental, shrubs between
the village and Stariwald, grid square 9452/4, on Cytospora sp. on Corylus
avellana, soc. Fenestella subsymmetrica, 10 Jan. 2011, W. Jaklitsch (WU
31641; MBT385685, MBT385686; ex-epitype culture CBS 144860 = FP).
Pseudostromatic pustules 0.6–3.6 mm diam or long, with circu-
lar, elliptic or oblong outline, lenticular, subglobose or pulvinate,
erumpent from bark and projecting to c. 0.6 mm, sometimes
confluent to rows of up to 10 mm, often compact; surface con-
vex, plane or with sunken centre, typically with a brittle, pale
brown, less commonly yellow- or dark brown disc or crust due
to condensed subiculum. Ascomata (330–)450 –665(–780) µm
(n = 40) diam, globose, subglobose to pyriform or distorted by
mutual pressure, loosely or densely aggregated in one or two
layers, connected by subiculum, also solitary on conidiomata
or ascomata of the Cytospora host. Subiculum dense or scant,
present at bases, sides and/ or surface of ascomata, consist-
ing of hyaline to dark brown, thick-walled, c. 2.5–6 µm wide
hyphae merging into pseudoparenchyma of the outer peridium.
Ostioles (90–)110–210(–270) µm (n = 47) diam, indistinct at
the surface, sometimes discoid or papillate with plane or convex
top and more or less circular outline, dark brown to black, often
only visible upon injury showing the whitish interior; sometimes
mixed with ostioles of the host. Peridium 20–90 (–120) µm thick,
pseudoparenchymatous, consisting of a dark brown narrow
outer and a highly variable glassy hyaline inner layer, the lat-
ter often thickened in upper regions particularly when young;
cells more or less isodiametric, thick-walled, (4.5–)6 –12(–19)
µm (n = 110) diam; outermost layer darkening in 3 % KOH.
Hamathecium consisting of often rather sparse, 1.5–2.5 (–3) µm
wide, branched and anastomosing ?paraphyses. Asci (185–)
207–294(– 328) × (18–)21– 26.5(– 28) µm (n = 32), cylindrical
to oblong, bitunicate, fissitunicate, with an ocular chamber,
a usually short stipe and simple or knob-like base, contain-
ing 8 ascospores in (obliquely) uniseriate, sometimes partly
biseriate arrangement. Ascospores (30–)34.5 –43.5(– 53.5)
× (12–)14–18 (–21) µm, l/w (1.9 –)2.2– 2.7(– 3.3) (n = 304),
ellipsoid or broadly fusoid, thick-walled, first hyaline to yellow-
ish with 1–5 transverse septa, asymmetric with submedian
primary septum, developing additional septa, turning yellowish
brown, when mature with often indistinct, 11–18 transverse
and 3–6 (–7) longitudinal septa, yellow- to golden brown when
fresh, medium to dark brown when dried; surface verruculose;
often upper part wider than lower; terminal cells concolorous or
hyaline, often narrowed and projecting as 1–2 µm long apiculi,
becoming longer (3–4 µm) when old; germinating from apiculi;
11
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fig. 5 Fenestella media. a– m1. Sexual morph. a–c. Pseudostromatic pustules in face view (obliquely sectioned to expose ascomata in c); d– e. ascomata
in vertical section (e. above peripheral Cytospora (Valsa) ascomata); f– h. peridium in vertical section; i. subicular hyphae; j–l. asci (j –k. young); m. apex of
young ascus; n. section of hamathecium; o– m1. ascospores (o– u. initial and young stages; v– x. from fresh material; g1. germinating); n1–o1. asexual morph
from CMD at 22 °C; n1. pycnidia; o1. conidia (f– g, j1– m1 in 3 % KOH). a, j, n, q. WU 36970 (FCO); b, h1. WU 36972 (FP3); c, f, i, l, s, e1, j1. lectotype of
F. macro spora (G 00127659); d, v, w, c1, i1, k1, l1, n1, o1. WU 31641 /CBS 144860 (FP); e, z, d1, g1. WU 36967; g, o, t, f1. holotype of F. media (PC 0706651);
h, p. WU 15513; k, m, u. WU 36974 (FP10); r, m1. WU 36971 (FP1); x. WU 36973 (FP7); y. WU15069; a1. WU 36969; b1. WU 36968. — Scale bars: a –c =
500 µm; d– e = 300 µm; f– i = 20 µm; j– l = 25 µm; m–m1 = 10 µm; n1 = 100 µm; o1 = 5 µm.
12 Persoonia – Volume 44, 2020
in 3 % KOH ascospores turning olivaceous when young and
dark to blackish brown when mature, apiculi remaining hyaline.
Culture characteristics and asexual morph in culture — Asco-
spores germinating simultaneously from many cells. Colony
radius on CMD at 22 °C in the dark 23 mm after 4 wk; colony
thick, dense, aerial hyphae forming dense white to pale grey mat
on pale brown mycelium, sometimes turning citrine and citrine
pigment diffusing into agar, centre turning black by crowded
pycnidia amassing from 4 days, eventually entire colony brown,
reverse grey, slightly zonate; odour indistinct. On MEA colony
soon turning brown, numerous pycnidia formed. Pycnidia when
formed usually covered by aerial hyphae, 120–210 µm diam,
more or less globose, greyish brown to black, discrete, crowded,
later fusing and forming clusters or irregular stromatic masses.
Conidia forming whitish to brown turbid drops, (2.2–)3 –4(– 4.6)
× (1–)1.3–1.6 (–2) µm, l/w (1.4 –)1.8 –2.8(– 3.9) µm (n = 70),
oblong to narrowly ellipsoid, hyaline, 1-(rarely 2-)celled, smooth,
with 1–2 subapical guttules.
Habitat — On Cytospora spp. (sexual and asexual morphs)
on various deciduous trees and shrubs, particularly common
on Corylus avellana.
Distribution — Europe, possibly North America; locally com-
mon in winter.
Other materials examined (all on Cytospora spp. on corticated twigs and
branches): AustRiA, Kärnten, St. Margareten im Rosental, shrubs between
the village and Stariwald, grid square 9452/ 4, on Corylus avellana, partly
overgrown by Exidia sp., 24 Feb. 1992, W. Jaklitsch (WU 15069); ibid., on
stem of Rubus idaeus, soc. ?Neocucurbitaria sp. (possibly on Apioporthe
vepris), 31 Dec. 1994, W. Jaklitsch W.J. 412 (WU 36965); ibid., on Corylus
avellana, 31 Dec. 1994, W. Jaklitsch W.J. 413 (WU 36966); ibid., on Corylus
avellana, 7 Jan. 1994, W. Jaklitsch (WU 15513); ibid., on Corylus avellana,
24 Oct. 1993, W. Jaklitsch (WU 15786); ibid., on Acer pseudoplatanus, soc.
Thyridaria sp. s.lat., 28 Dec. 2013, W. Jaklitsch (WU 36972; culture FP3); St.
Margareten im Rosental, Gupf, grid square 9452/ 4, on Corylus avellana, soc.
Massarina s.lat., 15 Apr. 1995, W. Jaklitsch W.J. 564 (WU 36967); Gupf, grid
square 9452/ 2, on Corylus avellana, soc. Fenestella subsymmetrica, 8 Nov.
2013, W. Jaklitsch (WU 36971; culture FP1); Niederösterreich, Maissau, grid
square 7460/ 2, on Corylus avellana, 26 Oct. 1995, W. Jaklitsch W.J. 764 (WU
36968); Mauerbach, close to the cemetery, grid square 7763/1, on Carpinus
betulus, 28 Sept. 1996, W. Jaklitsch W.J. 964 (WU 36969); Oberösterreich,
Schärding, Raab, Rothmayrberg, grid square 7648/1, on Corylus avellana,
mostly immature, 5 Sept. 2009, H. Voglmayr (WU 32630); Wetzlbach, on Tilia
cordata, 13 Aug. 2017, H. Voglmayr (WU 36971; culture FP10); Steiermark,
Steinberg, on Castanea sativa, 3 Nov. 2015, H. Voglmayr & W. Jaklitsch (WU
36970; culture FP7). – CRoAtiA, Istrija, NE Pula, near Krnica, on Carpinus
orientalis, 25 Sept. 2010, H. Voglmayr (WU 36970; culture FCO).
Notes — This is one of three cryptic species difficult to identi-
fy morphologically. Characteristic for F. media is the asymmetric
ascospore septation. See also notes under F. subsymmetrica
and F. viburni. As the fungus is not specific for the plant host, we
use a specimen on Corylus for epitypification. Fructifications of
F. media can be found particularly in winter, after the Cytospora
has become old. Size and development of pseudo stromata vary
considerably. The largest pseudostromata occur on Corylus and
may be locally very common. In contrast to Barr (1990), who
considered F. macrospora as a synonym of F. fenestrata (as
F. princeps), F. macrospora is clearly a synonym of F. media.
Nonetheless, morphological identification of some older, non-
cultured and non-sequenced specimens here included under
examined specimens is not always easy and therefore at least
in part, tentative.
Fenestella parafenestrata Jaklitsch & Voglmayr, sp. nov. —
MycoBank MB829744; Fig. 6
Etymology. Referring to the close relationship with Fenestella fenestrata.
Holotype. AustRiA, Oberösterreich, Raab, Wetzlbach, on Cytospora
(Valsa) sp. on a branch of Quercus robur, 24 Feb. 2018, H. Voglmayr (WU
36988; ex-type culture CBS 144856 = C306).
Ascomata (420–)460 –610(– 690) µm (n = 15) diam, globose,
subglobose to pyriform, immersed in and erumpent from bark,
solitary or in small groups of usually less than 10 individuals
forming pustules 0.5–2.2 mm diam on and connected by sub-
iculum on or associated with conidiomata or ascomata of Cyto-
spora (Valsa) sp. in the ostiolar region of the latter. Pustule sur-
face brownish by compacted subiculum or blackened by spore
deposits. Ostiolar areas 90– 240(–270) µm diam, dark brown,
flat or convex disc-like or irregular, sometimes slightly papillate.
Subiculum consisting of thick-walled, hyaline to greyish or dark
brown, 2–7 µm wide hyphae merging with the outer peridial
layer. Peridium 20 –75 µm thick, pseudoparenchymatous, con-
sisting of (5–)7.5–14 (–16.5) µm (n = 37) wide cells, dark brown,
gradually paler toward the interior, at the ostiole to 100 µm wide
and paler with (sub-)hyaline cells toward the interior; inside
compressed brownish cells present. Hamathecium consisting
of numerous richly branched, 1–3 µm wide ?paraphyses. Asci
(216–)241– 320(–342) × (21–)23– 27(–28) µm (n = 20), cylin-
drical, bitunicate, fissitunicate, with an ocular chamber, a usually
short stipe and simple or knob-like base, containing 6–8 asco-
spores in (obliquely) uniseriate arrangement; unstable in water;
biseriate rearrangement and long stipes generated by pres-
sure. Ascospores (32–)41.5 – 52.5(–61) × (13–)15 –19(–23)
µm, l/w (2.1–)2.4 –3.1(– 3.6) (n = 100), ellipsoid with upper part
usually broader than lower, constricted at the median to sub-
median primary septum, thick-walled, hyaline to yellowish
and with 3–5 (–8) transverse septa and 1 longitudinal septum
when young, turning yellow-brown to dark brown and with
11–16(–20) transverse and 2– 4 longitudinal septa; terminal
cells concolorous or hyaline at the tips and often with 1–4 µm
long acute apiculi; smooth, containing minute guttules; in 3 %
KOH mature spores turning blackish brown. Pycnidia of the
presumed asexual morph sometimes associated with ascomata
on the natural host.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 8 mm after 1 wk, 26 mm
after 4 wk; colony thick, dense, slightly zonate, white, turning
cream to pale brownish, releasing bright yellow pigment dif-
fusing into surrounding agar, centre turning olivaceous to dark
brown due to pycnidia, surface velvety by a white to pale greyish
or brownish mat of aerial hyphae; reverse yellow-brown, dark
brown in the centre; odour indistinct to pleasant or leathery.
Pycnidia appearing after 4 d below white aerial hyphae, (90 –)
150–330 µm diam, globose, black, first hyaline to greenish,
turning olivaceous and eventually black, surrounded by brown
hyphal appendages, numerous, tightly aggregated and fusing
into stromatic masses to c. 2 mm diam with many ostioles re-
leasing conidia as turbid whitish to olivaceous drops; peridium
thin, pseudoparenchymatous. Phialides (4–)4.5 – 6.3(–6.6) ×
(1.8–)2 –3.5(– 4) µm (n = 12), sessile, subglobose to ampulli-
form to conical. Conidia (2.5–)3 –4(–4.7) × (1–)1.1–1.5(– 2) µm,
l/w (1.9–) 2.4–3.2 (– 3.9) (n = 65), cylindrical, oblong to sub-
allantoid or narrowly ellipsoid, smooth, with 1–2 subterminal gut-
tules.
Habitat — On Cytospora (Valsa) spp. on deciduous trees,
confirmed for Quercus and Salix.
Distribution — Europe.
Other specimens examined. AustRiA, Burgenland, Mattersburg, Starem-
bühl / Rosaliengebirge, on Cytospora (Valsa) sp. on attached twig of Quercus
petraea, 1 Oct. 2001, W. Jaklitsch W.J. 1815 (WU 36990); Oberösterreich,
Raab, Wetzlbach, on Cytospora (Valsa) sp. on a branch of Salix sp., 31 Mar.
2018, H. Voglmayr (WU 36989; culture C317).
Notes — This species is morphologically intermediate be-
tween F. fenestrata and the cryptic species F. media, F. subsym-
metrica and F. viburni. See also notes under F. fenestrata.
13
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fig. 6 Fenestella parafenestrata. a–w. Sexual morph. a –c. Ascomatal groups connected by subiculum erumpent through bark fissures in face view; d. pe-
ridium in vertical section; e– g. asci (e. young, from fresh material); h– i. ascus apices (immature in h); j– w. ascospores (j– k. young; j, l– o. from fresh material);
x– a1. asexual morph from CMD at 22 °C; x. pycnidia; y. phialides and conidia; z–a1. conidia; b1. ascospore of Fenestella cf. fenestrata K(M) 233193 (t, v.
in 3 % KOH). a –b, d– j, l–m, p –t, v, x, a1. WU 36988/CBS 144856 (C306); c, n –o, y– z. WU 36989 (C317); k, u, w. WU 36990. — Scale bars: a = 300 µm;
b– c = 500 µm; d, h– w, y, b1 = 10 µm; e –g = 30 µm; x = 200 µm; z–a1 = 5 µm.
14 Persoonia – Volume 44, 2020
Fig. 7 Fenestella subsymmetrica. a– z. Sexual morph. a. Pseudostromatic pustule in face view; b– c. ascomata in vertical section (b. above Cytospora (Valsa)
ascomata); d. peridium with subicular hyphae in vertical section; e, i– k. ascus apices (e. immature; i. from fresh material); f– h. asci (f. immature; g. from fresh
material); l– z. ascospores (l–n. initial and young stages; o. from fresh material); a1–e1. asexual morph from CMD at 22 °C; a1. pycnidia; b1. phialides; c1–e1.
conidia (e–f, h, j– k, n, z. in 3 % KOH). a, c, o, y. WU 36978 (FP4); b, d– f, h, j, k, m– n, u, x, z– e1. WU 36979 /CBS 144861 (FP6); g, i, p, s –t. WU 36975;
l, r, v. WU 36977 (C286); q. WU 36976 (C285); w. WU 36980 (FP8). — Scale bars: a– b = 500 µm; c, a1 = 200 µm; d, f–h = 25 µm; e, i– k, n– z = 10 µm;
l– m, b1 = 5 µm; c1–e1 = 3 µm.
Fenestella subsymmetrica Jaklitsch & Voglmayr, sp. nov. —
MycoBank MB829745; Fig. 7
Etymology. Referring to the nearly symmetric shape of ascospores due
to the submedian to median insertion of the primary septum.
Holotype. AustRiA, Vienna, 21st district, at Marchfeldkanalweg near Felix
Slavikstraße, on/ soc. Cytospora holomorph on Acer campestre, soc. Diplodia
sp., Fusarium sp., 8 Nov. 2015, W. Jaklitsch (WU 36979; ex-type culture CBS
144861 = FP6).
Pseudostromatic pustules 0.7–3.4 mm wide or long, with circu-
lar, elliptic or oblong outline, subglobose or pulvinate, erumpent
from bark, sometimes confluent; surface usually ill-defined and
irregular, convex, plane or with sunken centre, often partly co-
vered by bark fibres or brownish condensed subiculum, pale
brown to nearly black. Ascomata (300–)370 –600(–765) µm
(n = 49) diam, subglobose to pyriform or distorted by mutual
pressure, often obliquely oriented and convergent toward the
15
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
pustule centre, loosely or densely aggregated in valsoid or ill-
defined groups of up to c. 20 individuals on and connected by
subiculum, sometimes fusing laterally, also solitary on conidio-
mata or ascomata of the Cytospora host in its ostiolar region.
Subiculum present at bases, sides and/or surface of ascomata,
consisting of mostly pale brown, thick-walled, 2.5–6 (–7) µm
wide hyphae. Ostiolar areas (75–)85 –163(–180) µm (n = 10)
diam, ill-defined, irregular, often only visible by spore deposits,
sometimes roundish and slightly projecting, black. Peridium
20–80 (–90) µm thick, pseudoparenchymatous, consisting of a
dark brown narrow outer and a glassy pale brownish to hyaline
inner layer, the latter often thickened in upper regions particu-
larly when young; cells more or less isodiametric, thick-walled,
(3.5–)5.5 –11(–15) µm (n = 82) diam; in 3 % KOH outermost
layer turning blackish brown. Innermost part of the inner layer
often slightly darker and of distinctly compressed elongate
cells. Hamathecium consisting of numerous 1– 3 µm wide,
branched and anastomosing ?paraphyses. Asci (182–)207–
302(– 345) × (19 –) 21.5–25.5 (–26.5) µm (n = 32), cylindrical
to oblong, bitunicate, fissitunicate, with an ocular chamber, a
usually short stipe and simple or knob-like base, containing
4–8 ascospores in (obliquely) uniseriate, sometimes partly
biseriate arrangement. Ascospores (28–)34.5 –44.5(– 54.5) ×
(13–)15.5–19.5 (–24.5) µm, l/w (1.8–) 2–2.5 (–2.9) (n = 201),
broadly ellipsoid, oblong or broadly fusoid, thick-walled, first
hyaline to yellowish with 1–4 transverse septa, asymmetric
to subsymmetric, with submedian to median primary septum,
developing additional septa, turning pale brown to olivaceous,
when mature with distinct, 11–16(–18) transverse and 3– 6
longitudinal septa and yellow- to golden brown when fresh,
dark brown when dried; surface verruculose; often upper part
wider than lower; terminal cells concolorous or hyaline, often
narrowed and projecting as 1–2 µm long apiculi, becoming
longer (3–5 µm) when old; germinating from apiculi; in 3 %
KOH ascospores turning olivaceous when young and dark to
blackish brown when mature, apiculi remaining hyaline.
Culture characteristics and asexual morph in culture — Asco-
spores germinating simultaneously from many cells. Colony
radius on CMD at 22 °C in the dark 6 mm after 1 wk, c. 20 mm
after 3– 4 wk; colony white, centre turning black by pycnidia
after 4 d, soon entire colony turning grey, brownish grey to
olivaceous, margin often hyaline to white, covered by a white
to pale grey mat of aerial hyphae; odour indistinct; no diffusing
pigment formed. Pycnidia 120–240 µm diam, more or less glo-
bose, first hyaline to greenish, turning green to black, numerous,
often concentrically and very densely arranged, spreading over
entire colony or remaining in the centre; often covered by mats
of aerial hyphae; conidia amassing in whitish to greenish turbid
drops. Phialides 4.5– 8 × 2–4 µm, lageniform to subglobose with
a long neck. Conidia (3.2–)3.5 –4.2(–4.5) × (1.1–)1.3–2 (–2.3)
µm, l/ w (1.7–)1.9–3 (–3.7) (n = 30), cylindrical, oblong to el-
lipsoid, 1-celled, hyaline, with 1–3 drops, smooth.
Habitat — On Cytospora spp. (sexual and asexual morphs)
on various deciduous trees and shrubs.
Distribution — Europe, possibly North America; locally com-
mon in winter; sometimes co-occurring with F. media.
Other materials examined (all on or in pseudostromata of Cytospora
spp. including their Valsa sexual morphs): AustRiA, Kärnten, St. Margareten
im Rosental, Aussicht, grid square 9452/3, on branch of Corylus avellana,
on Valsa morph, soc. Parafenestella sp., 8 Jan. 1994, W. Jaklitsch W.J. 91
(WU 15613); Gupf, grid square 9452/2, on Corylus avellana, soc. Fenestella
media, 8 Nov. 2013, W. Jaklitsch (WU 36975; part of WU 36971); Niederöster-
reich, Bad Vöslau, Grossau, near Haidlhof, on old Cytospora holomorph on
Salix caprea, 22 Feb. 2016, W. Jaklitsch & H. Voglmayr (WU 36980; culture
FP8); Oberösterreich, Schärding, Raab, between Riedlhof and Großrotmayr,
grid square 7647/ 2, on branch of Corylus avellana, 18 Mar. 2015, H. Voglmayr
(WU 36978; culture FP4); Vienna, 22nd district, at AGES, Spargelfeldstraße
191, on Valsa morph on cut branches of Juglans regia; soc. Diaporthe sp., 25
Jan. 2017, R. Moosbeckhofer (WU 36976; culture C285); ibid., other tree of
Juglans regia, 25 Jan. 2017, R. Moosbeckhofer (WU 36977; cultures C286,
C286x).
Notes — Fenestella subsymmetrica is hardly distinguishable
from F. media by morphology alone. Ascospores of F. subsym-
metrica often tend to appear broader, with more distinct septa
and a more median primary septum. However, individual speci-
mens pose serious problems in morphological identification.
For example, culture C286x derived from distinctly asymmetric
asco spores of WU 36977 yielded ITS and LSU sequences,
which are identical with those derived from symmetric asco-
pores. In cultures on CMD no pigment is formed. Mature asci
are very unstable in water, therefore they were mostly measured
and illustrated in 3 % KOH.
Fenestella viburni Jaklitsch & Voglmayr, sp. nov. — MycoBank
MB829746; Fig. 8
Etymology. Owing to its occurrence on Viburnum spp.
Holotype.
AustRiA, Niederösterreich, Wr. Neustadt, Markt Piesting, on the
Hart N Piesting, grid square 8162/ 2, elev. 500 m, on Cytospora (Leucostoma)
sp. on Viburnum lantana, 12 Oct. 2014, H. Voglmayr (WU 36982; ex-type
culture CBS 144863 = FVL).
Ascomata (330–)390 – 600(–720) µm (n = 21) diam, subglo-
bose to subpyriform, immersed singly or in small groups in the
ostiolar region above ascomata or in conidiomata of Cytospora
(Leucostoma) sp., less commonly forming pulvinate pseudo-
stromatic pustules 0.7–2.5 mm diam with circular or oblong
outline, in loose association with the fungal host, erumpent
from bark. Subiculum individually surrounding ascomata and
connecting them, consisting of thick-walled, pale to dark brown,
2–6 µm wide hyphae, sometimes condensed to brown crusts
between ascomata. Ascomatal apices obtuse, brown, mostly
90–180 µm diam; ostioles 70–150 µm diam, usually inconspicu-
ous, rarely papillate, black, sometimes whitish. Peridium 20–60
µm thick, pseudoparenchymatous, consisting of a dark brown
narrow outer and a glassy pale brownish to hyaline inner layer;
cells more or less isodiametric, thick-walled, (5–)7–13.5(–17)
µm (n = 40) diam; innermost part of the inner layer of distinctly
compressed brownish cells. Hamathecium consisting of nu-
merous, 1–3 µm wide, branched and anastomosing ?para-
physes. Asci (247–) 258–295(– 312) × (19–)22 – 26(– 29) µm
(n = 34), cylindrical to oblong, bitunicate, fissitunicate, with an
ocular chamber, a usually short stipe and simple or knob-like
base, containing 8 ascospores in uniseriate arrangement.
Ascospores (29–)38 –46(–49.5) × (12.5–)15–18 (–22) µm, l/w
(2–)2.3 – 2.8(– 3.1) (n = 193), ellipsoid or fusoid, sometimes
distinctly pointed at the ends, asymmetric to subsymmetric, with
submedian to median primary septum, first hyaline to yellowish,
turning olivaceous, when mature yellow- to golden brown when
fresh, dark brown when dried, with distinct 11–16 transverse
and 3–6 longitudinal septa; surface verruculose; upper part
often slightly wider than lower; terminal cells concolorous or
terminally hyaline and projecting as minute, 1–2 µm long apiculi,
becoming slightly longer when old; in 3 % KOH ascospores
turning olivaceous when young and darker to blackish brown
when mature, apiculi remaining hyaline.
Culture characteristics and asexual morph in culture — Asco-
spores germinating simultaneously from many cells. Colony
radius on CMD at 22 °C in the dark 4 mm after 1 wk, 22 mm
after 4 wk; colony thick, dense, white, turning grey or olivaceous
grey with white margin, velvety by a dense whitish to greyish mat
of aerial hyphae; odour indistinct. Pycnidia appearing after 5 d,
globose, 90–250 µm diam, first hyaline to olivaceous, turning
black, immersed to superficial, tightly aggregated or fusing
in large numbers around the inoculation plug or spreading
over the colony, often covered by aerial hyphae, releasing
conidia in whitish turbid drops through ostioles lined by clavate
16 Persoonia – Volume 44, 2020
Fig. 8 Fenestella viburni. a– q. Sexual morph. a. Ascomata in face view (inserted right and left in a Cytospora (Leucostoma) pseudostroma); b. ascomata above
Cytospora (Leucostoma) ascomata in vertical section; c. peridium in vertical section; d– e. ascus apices; f– h. asci (f– g. from fresh material); i– q. ascospores
(i– l. from fresh material; m. young); r– u. asexual morph from CMD at 22 °C; r. pycnidia and conidial drops; s. peridium in surface view; t –u. conidia. a, e, h,
m, o, q. WU 36983 (FP2); b– c, f–g, i–l, r –u. WU 36984/CBS 144863 (FVL); d, n. WU 36982; p. WU 15341. — Scale bars: a –b, r = 300 µm; c–e, i –q = 10
µm; f– h = 25 µm; s = 15 µm; t– u = 3 µm.
hyaline marginal cells; peridium thin, pseudoparenchymatous,
consisting of thin-walled cells (4.5–)6.5 –11(–14) µm (n = 30)
diam. Phialides (3.8 –)4.8–7.5 (–8.2) × (1.7–)2.5 – 4(–4.2) µm
(n = 10), crowded, lageniform to subglobose with long neck or
subulate. Conidia (3.3 –)3.5–5 (–6.3) × (1.4–)1.7– 2.3(– 2.7) µm,
l/ w (1.6–)1.9– 2.5(–3) (n = 32), cylindrical, oblong to ellipsoid,
sometimes pinched, 1-celled, hyaline, with 2 subterminal drops,
smooth.
Habitat — On Cytospora spp. (both morphs; sexual morph
of the Leucostoma type) on Viburnum spp.
Distribution — Europe.
Other materials examined. AustRiA, Kärnten, St. Margareten im Rosental,
Aussicht, grid square 9452/ 3, on Viburnum lantana, 8 Jan. 1994, W. Jaklitsch
(WU 15341); shrubs between the village and Stariwald, grid square 9452/4,
on Cytospora sp. on Viburnum opulus, 24 Dec. 1995, W. Jaklitsch W.J. 814
(WU 36982); Stariwald, grid square 9452/4, on Cytospora sp. on Viburnum
lantana, 10 Jan. 1995, W. Jaklitsch W.J. 454 (WU 36981). – FRAnCe, Aude,
Belcaire, chemin du Traouc, elev. 1050 m, on Cytospora (Leucostoma) sp. on
Viburnum lantana, 25 Oct. 2013, J. Fournier J.F.13212 (WU 36981; culture
FP2).
Notes — Fenestella viburni is one of three cryptic species,
morphologically most closely related to F. subsymmetrica, but
difficult to differentiate. In individual specimens ascospores
tend to be distinctly pointed terminally. Formation of pseudo-
stromatic pustules is less pronounced and asci are more stable
in water than with F. media and F. subsymmetrica. Pustules are
difficult to assess, as they are usually produced basically by
its Leucostoma host. Older, not sequenced specimens from
Viburnum spp. are added tentatively to the list above.
Neocucurbitaria Wanas. et al., Mycosphere 8: 408. 2017,
emended by Jaklitsch & Voglmayr in Jaklitsch et al. (2018)
Type species. Neocucurbitaria unguis-hominis (Punith. & M.P. English)
Wanas. et al.
17
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Notes — Neocucurbitaria juglandicola is not host-specific, as
it has been recently collected on Quercus rubra, too. Cultured
and sequenced material: Austria, Oberösterreich, St. Willibald,
Große Sallet, on a branch of Quercus rubra, 30 Mar. 2018, H. Vogl-
mayr (WU 36984; culture C316).
Neocucurbitaria subcaespitosa (G.H. Otth) Jaklitsch & Vogl-
mayr, comb. nov. — MycoBank MB829747; Fig. 9
Basionym. Cucurbitaria subcaespitosa G.H. Otth, Mitth. Naturf. Ges. Bern
711–744: 103. 1871 ‘1870’.
Synonym. Fenestella subcaespitosa (G.H. Otth) M.E. Barr, Ann. Univ.
Turku., A II 55: 14. 1974.
Lectotype, here designated: switzeRlAnd, near Bern, on twigs of Sorbus
aria, without date, G.H. Otth (B 700016481; transferred from Münster in 1936;
MBT385687). On the label Otth noted that he retained this material as No.
10, a rather bad but perhaps not entirely useless part of No. 90. He might
have sent No. 90 to Nitschke for inspection. No additional material is extant
in B, but according to R. Berndt (pers. comm.) Otths’ material was transferred
from Bern to Z, where a part of the type may be present but is currently not
accessible. For this reason we designate B 700016481 as lectotype.
Ascomata (300 –)354 –550 (–600) µm (n = 20) diam, more or less
globose, immersed-erumpent from bark, loosely aggre gated on
subiculum in valsoid groups or in rows or firmly united by grey -
ish or brown subiculum forming pseudostromatic pustules
0.5–2.6 mm diam of various shapes containing up to c. 10
ascomata; ascomata also solitary and glabrous or individu ally
covered by brown, crust-like subiculum. Ostioles (60–)95 –
186(– 210) µm (n = 21) diam outside, papillate, or cylindrical
and projecting to c. 210 µm, sometimes apically flattened,
circular, angular or substellate in section, shiny black, whitish
inside when injured. Asci cylindrical, bitunicate, containing 8
ascospores in uniseriate arrangement. Asco spores (20–)21.5 –
26(– 29.5) × (8 –)9–11.5(–14) µm, l/w (1.9 –)2.1– 2.6(–2.9)
(n = 81), ellipsoid, with (4–) 5–7(– 9) distinct transverse and 1–2
longitudinal septa, distinctly constricted at the median primary
septum, less distinctly at other septa, pale brown when im-
mature, dark brown when mature, ends rounded, concolorous,
surrounded by a narrow hyaline perispore swelling in KOH to
2 µm.
Habitat — On dead partly corticated twigs of Sorbus aria.
Distribution — Europe.
Other material examined. AustRiA, Kärnten, St. Margareten im Rosental,
Schwarzgupf, grid square 9452/4, on branch of Sorbus aria, 25 May 1997,
W. Jaklitsch W.J. 1072 (WU 36991).
Fig. 9 Neocucurbitaria subcaespitosa. a–e. Ascomata in face view; f –l. ascospores (i– l. in 3 % KOH). a, d– e, i–l. WU 36991; b–c, f –h. Cucurbitaria sub-
caespitosa holotype B 700016481. — Scale bars: a– e = 300 µm; f– l = 5 µm.
18 Persoonia – Volume 44, 2020
Fig. 10 Parafenestella alpina. a– r. Sexual morph. a. Ascomata in face view; b. vertical section of young ascoma above the perithecial host and one ascoma
in horizontal section; c. vertical section of laterally fused ascomata surrounded by subiculum; d– f. asci; g. peridium of laterally fused ascomata in vertical
section; h– i. ascus apices (h. immature); j. paraphysis tip near immature ascus apex; k –r. ascospores (k. young; q, r. aberrant); s– v. asexual morph from
CMD at 22 °C; s. pycnidia and conidial drops; t –u. phialides; v. conidia (h, j, o. in 3 % KOH). a–m, p– v. WU 36997/ CBS 145263 (C198); n– o. WU 36998
(C249). — Scale bars: a– c, s = 200 µm; d– g = 20 µm; h, j, r = 10 µm; i, k– q, v = 7 µm; t–u = 5 µm.
19
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Notes — In both studied specimens ascomata are over-
mature, and only fragments of asci allowing interpretations of
ascus shape and ascospore arrangement. Our material was
neither cultured nor sequenced, therefore relegation to Neocu-
curbitaria is tentative but strongly suggested by the morphology,
particularly based on features of ascomata, ascospores and
ostioles (compare Jaklitsch et al. 2018). Barr (1990) incorrectly
synonymised this species with Cucurbitaria sorbi without ha-
ving seen type material and anticipated occurrence on several
Sorbus spp. in Europe and North America. However, N. sub-
caespitosa seems to occur only on Sorbus aria.
Parafenestella Jaklitsch & Voglmayr, in Jaklitsch et al., Stud.
Mycol. 90: 108. 2018
Type species. Parafenestella pseudoplatani Jaklitsch & Voglmayr.
Parafenestella alpina Jaklitsch & Voglmayr, sp. nov. — Myco-
Bank MB829748; Fig. 10
Etymology. For its occurrence in subalpine to alpine regions.
Holotype.
AustRiA, Osttirol, Prägraten am Großvenediger, Wallhorn, Boden -
alm, elev. c. 2000 m, on dead attached twigs of Cotoneaster integerrimus, soc.
Cytospora (Leucostoma morph) sp., Discostroma sp. (in excess), Mollisia sp.,
cf. Nigrograna sp., cf. Teichospora sp., 18 June 2015, W. Jaklitsch & H. Vogl-
mayr (WU 36997; ex-type culture CBS 145263 = C198).
Ascomata (180–)240 – 375(– 450) µm (n = 22) diam, globose,
subglobose or pyriform, usually tightly aggregated in bark on a
perithecial host fungus in small numbers and connected by sub-
hyaline to dark brown, thick-walled, 2–5 µm wide subicular hy-
phae, dark brown to black; tightly packed ascomata sometimes
covered by a brown to black, densely packed mesh of subicular
hyphae and ejected ascospores. Ostiolar areas (53–)60 –
105(–135) µm (n = 12) diam, slightly papillate, rounded, black.
Peridium 15–70 µm thick, pseudoparenchymatous, consisting
of isodiametric cells (4–)5 –9.5(–12) µm (n = 30) diam, out-
side moderately thick-walled and dark brown, paler to hyaline
in upper regions and thinner-walled to the inside; confluent
with tightly appressed ascomata. Hamathecium consisting of
numerous 0.5 –1 µm (to 2.5 µm in 3 % KOH) wide, branched
paraphyses with free ends. Asci (143–)170– 208(– 227) × (18–)
18.5–21.5 (–24.5) µm (n = 25), cylindrical to oblong, bitunicate,
fissitunicate, with a truncate ocular chamber, a short stipe
and simple or knob-like base, containing 6–8 ascospores in
uniseriate arrangement. Ascospores (19–)24 –30.5(– 35) ×
(10.5–)12–14 (–15.5) µm, l/ w (1.4–)1.8– 2.4(–2.9) (n = 73),
typically ellipsoid to fusoid, often inequilateral, very variable in
shape and size, from subglobose to clavate or lower part elon-
gated fusoid, first with 1–5 main septa, constricted at the more
or less median primary septum, developing (7–)8 –12(–15)
transverse and (2–)3 – 4 longitudinal septa, with upper part
often broader, first hyaline to yellowish, turning medium to dark
brown, blackish brown when old; ends concolorous; in 3 % KOH
turning pale olivaceous when young and dark to blackish brown
when mature or old.
Culture characteristics and asexual morph in culture — Asco-
spores germinating si multaneously from many cells. Colo ny
radius on CMD at 22 °C in the dark 8 mm after 1 wk, 25 mm
after 3 wk, 38 mm after 5 wk; colony dark grey to olivaceous,
centre darker; aerial hyphae long, white, forming an initially
loose later dense mesh above the colony; odour indistinct.
Pycnidia 75–170 µm diam, (sub-)globose, papillate with a
pale opening, numerous, first appearing after 3 d, hyaline,
turning greenish, olivaceous to black, mostly immersed, parti-
ally erumpent, solitary and in firm packs, spreading from the
centre; conidia emitted as whitish turbid drops. Peridium thin,
pseudoparenchymatous, olivaceous, surrounded by subhyaline
submoniliform hyphae. Phia lides 4.8–7.5(– 9.3) × 2– 3.5(– 4.4)
µm (n = 14), sessile, varying from subglobose over ampulliform
and lageniform to subulate. Conidia (3.4–)3.7– 4.3(– 4.6) ×
(1–)1.1–1.4(–1.5) µm, l /w (2.6 –)2.9– 3.7(–3.9) (n = 24), cylin-
drical to allantoid, less commonly narrowly ellipsoid, 1-celled,
hyaline with 2 small drops, smooth.
Habitat — On perithecial fungi on Cotoneaster integerrimus
and Salix appendiculata.
Distribution — Central Europe (Austria).
Other materials examined. AustRiA, Steiermark, Deutschlandsberg, Kor-
alpe, at the parking place of the walking path to the Grünanger- and Bären-
talhütte; N46°49'44" E15°00'56", elev. c. 1540 m; on dead attached twigs of
Salix appendiculata, soc. effete Plagiostoma sp., Plenodomus hendersoniae
(in excess), 6 May 2016, G. Friebes (WU 36998; culture C249).
Notes — Due to the rough climate in (sub)alpine regions,
asci and ascospores are often aberrantly developed. The fungal
host of P. alpina may be Cytospora, but due to the many other
fungi that are present on the specimens, this is uncertain.
Parafenestella austriaca Jaklitsch & Voglmayr, sp. nov. —
MycoBank MB829749; Fig. 11
Etymology. For its occurrence in Austria.
Holotype.
AustRiA, Oberösterreich, Schärding, St. Willibald, Geitzedt, grid
square 7648/1, on branch of Rosa canina, 19 Mar. 2015, H. Voglmayr (WU
37014; ex-type culture CBS 145262 = C152).
Ascomata (270–)295 – 412(– 450) µm (n = 10) diam, subglo-
bose to pyriform, immersed-erumpent from bark, scattered or
aggregated in small valsoid groups, often on an effete peri-
thecial fungus, laterally collapsing from above, basally and
laterally surrounded by subhyaline to dark brown, thick-walled,
2–5 µm wide, smooth to verruculose subicular hyphae turn-
ing olivaceous in 3 % KOH. Ostiolar area (60–)75–128 (–150)
µm (n = 17) diam, convex or papillate with rounded opening,
black. Peridium 15– 85 µm thick, pseudoparenchymatous, con-
sisting of isodiametric cells (3.5–)4 –10(–14.5) µm (n = 40)
diam, outside thick-walled and dark brown, paler to hyaline
in upper regions and thinner-walled to the inside; darkening
in 3 % KOH. Hamathecium consisting of numerous 1–2.5
µm wide, branched ?paraphyses in a gel matrix. Asci (150–)
159–205 (–237) × 16–19.5(–24.2) µm (n = 33), cylindrical, bi-
tunicate, fissitunicate, with a distinct ocular chamber, a short
stipe and simple or knob-like base, containing 4–8 ascospores
in uniseriate arrangement. Ascospores (25–)27– 32.5(–38.2) ×
(12–)13–15 (–16.5) µm, l/w (1.9 –)2 –2.3(– 2.6) (n = 87), broadly
ellipsoid with usually broadly rounded ends and upper part often
wider, constricted at the median or slightly supra- or submedian
primary septum, with 9–13(–14) distantly spaced transverse
including v-septa in end cells and 3–4 (–5) longitudinal septa,
first hyaline to yellowish, with 1–3 main septa, turning yellow-
ish brown, finally medium to dark brown or dark reddish brown
with concolorous, sometimes paler to hyaline ends; surface
appearing verruculose; in 3 % KOH wall appearing smooth,
interior containing numerous minute droplets, turning green-
ish to yellow-green when immature, dark to blackish brown
when mature.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 7–8 mm after 1 wk, 16
mm after 2 wk, 26–27 mm after 23 d; colony first white, soon
turning greyish olivaceous to greyish brown; aerial hyphae form-
ing a dense mesh above the colony hiding pycnidia; reverse
dark grey to black; odour indistinct. Pycnidia c. 100–200 µm
diam, subglobose with papillate to cylindrical ostioles, first ap-
pearing after 3 d, hyaline, turning greenish, olivaceous to black,
mostly immersed, scattered to densely aggregated, covered
by aerial hyphae; conidia emitted as whitish turbid drops.
20 Persoonia – Volume 44, 2020
Fig. 11 Parafenestella austriaca. a– y. Sexual morph. a. Ascomata in face view and in vertical section; b. subicular hyphae; c. peridium in vertical section;
d– f. asci (d. young, 4-spored); g. periphysis; h –i. ascus apices; j –y. ascospores (j –n. immature; y. germinating); z– d1. asexual morph from CMD at 22 °C;
z. phialide; a1–d1. conidia. (d– f, h–i, l, n –o, s, u–x, z. in 3 % KOH). a –l, n–q, s –v, y–d1. WU 37014 /CBS 145262 (C152); m, r, w, x. M 0281852. — Scale
bars: a = 250 µm; b, g– h, x– y = 10 µm; c– f = 20 µm; i–w = 7 µm; z –b1 = 5 µm; c1– d1 = 3 µm.
Peridium thin, pseudo parenchymatous, olivaceous. Phialides
(3.5–)4.2 –7(– 8.2) × (1.5–)2– 3.2(– 3.8) µm (n = 16), sessile or
formed on short, 1–2 celled conidiophores, varying from sub-
globose over ampulli form and lageniform to subulate. Conidia
(3–)3.5 –4.2(– 5) × 0.9 –1.5(–2) µm, l/w (2.1–)2.6 –3.7(–4.2)
(n = 50), cylindrical to allantoid, less commonly narrowly ellip-
soid, 1-celled, hyaline, with 2 small drops, smooth; produced on
phialides and pegs. After a few transfers no pycnidia formed.
Habitat — Associated with perithecial fungi on Rosa canina
and possibly Crataegus monogyna.
Distribution — Central Europe (Austria).
Notes — Von Niessl’s Austrian collection from Crataegus in
Rosenthal bei Hütteldorf, Vienna (M-0281852; as Cucurbitaria
crataegi Niessl) is morphologically indistinguishable and there-
fore apparently this species. As we have not seen type material
of Cucurbitaria rosae G. Winter & Sacc. (none is present in W),
21
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
its concept is unclear, but its protologue suggests that it may be
a synonym of C. acervata. However, a fungus matching the de-
scription of that species in the sense of Mirza (1968) was found
not belong to the Cucurbitariaceae. Another species described
from Rosa is Cucurbitaria occulta Fuckel, with ascospores 16
× 8 µm having 4–5 transverse and 1 longitudinal septa. Its
type material (Germany, Hessen, Oestrich, Oestricher Wald,
erumpent on Rosa canina in the spring, Fuckel (G 00266382;
Fungi rhenani 1279, from Herbier Boissier, labelled Agyrium
nitidum Lib.) contains a drawing with a cylindrical ascus 102
× 13 µm with 8 uniseriate ascospores, one ellipsoid, with 5/1
septa, 16 × 8 µm and bark fragments containing the asexual
Fig. 12 Parafenestella faberi. a– b. Ascomata in face view (with superficial subiculum in b); c– e. asci; f. ascoma above Cytospora (Valsa) ascomata in vertical
section; g– h. peridium in vertical section (g. from side; h. from top); i– j. ascus apices; k– z. ascospores (k. immature; l, q. surface view) (e, q –w, y–z. in 3 %
KOH; h, o. in lactoglycerol (from slides prepared by R. Phookamsak). a, c–d, f, h –i, o, q– y. lectotype GZU; b, e, g, j–n, p, z. WU 37022. — Scale bars: a–b,
f = 200 µm; c– e, g– h = 15 µm; i, q, u– w, y– z = 10 µm; j–p, r –t, x = 7 µm.
22 Persoonia – Volume 44, 2020
fungus Agyriella nitida as black gelatinous drops producing
masses of cylindrical to allantoid, 1-celled hyaline conidia on
ampulliform phialides, a Cytospora (Valsa morph) sp. and a
Diplodia sp. No sexual morph matching C. occulta was found.
Fuckel (1870) interpreted the Agyriella as asexual morph of his
C. occulta. For comparison with other Parafenestella spp. on
Rosaceae see notes under P. rosacearum.
Parafenestella faberi (J. Kunze) Jaklitsch & Voglmayr, comb.
nov. — MB829750; Fig. 12
Basionym. Fenestella faberi J. Kunze, Fung. Sel. Exs., Cent. 3: no. 263.
1879.
Synonyms. Fenestella mackenziei Wanas. et al., Mycosphere 8: 407.
2017b.
Parafenestella mackenziei (Wanas. et al.) Jaklitsch & Voglmayr, in Ja-
klitsch et al., Stud. Mycol. 90: 109. 2018.
Lectotype of Fenestella faberi, here designated. GeRmAny, Sachsen-
Anhalt, Mansfeld-Südharz, Eisleben, Oberrißdorf, on dead corticated sticks of
Rosa canina, Sept. 1878, J. Kunze (GZU, Inv No. 226, Digi Bota ID 365656;
Joannes Kunze, Fungi Selecti exsiccati, ex museo botanico berolinensi; as
Thyridium faberi J. Kunze, nom. nud.; MBT385688). No type material is extant
in B, therefore the material in GZU, which was originally received from B,
may be the only available type specimen.
Ascomata (240–)300 – 450(– 480) µm (n = 14) diam, subglo-
bose to pyriform or subconical, black, tightly or loosely aggre-
gated in small numbers in more or less valsoid configuration
below blackened epidermis on inner bark or among ostiolar
necks of Cytospora (Valsa) sp., partly erumpent through bark
fissures, surrounded and connected by hyaline to brown, thick-
walled, 2–4.5 µm wide subicular hyphae; sometimes subiculum
also forming brown discs c. 0.2–1 mm diam at the bark surface.
Ostiolar areas 50–180 µm diam, inconspicuous, appearing as
black dots or blunt black papillae. Peridium 15–70 µm thick,
thickest around the ostiole, pseudoparenchymatous, consisting
of (4–)5.5 –10.5(–14.5) µm (n = 54) wide cells, outside thick-
walled and very dark brown textura angularis with encrusted
pigment, gradually paler to hyaline and thinner-walled to the
inside, partly terminated at the inner side by a compressed layer
of pale brown longish cells; cells more isodiametric at upper
levels and sometimes vertically elongated at the sides; in 3 %
KOH turning dark olivaceous to dark brown. Hamathecium con-
sisting of numerous 1–3 µm wide, branched ?paraphyses. Asci
(110–)135 –180(– 200) × (15.5 –)18.5–23.5(– 26) µm (n = 22),
cylindrical to oblong or narrowly clavate, bitunicate, fissitunicate,
with a distinct ocular chamber, a short stipe and simple or knob-
like base, containing 4–8 ascospores in uniseriate to partly
biseriate arrangement. Ascospores (23.5–)28.5 – 36(–42) ×
(11–)12.5–16(–17.5) µm, l/ w (1.9–)2.1– 2.5(–2.8) (n = 103),
ellipsoid, sometimes fusoid, with upper part broader than lower,
first hyaline, with 1–4 main septa, turning pale or yellowish
brown, eventually dark brown, reddish brown in herbarium ma-
terial, with 7–12(–14) transverse and 1– 3(–5) longitudinal
septa, slightly constricted at the median to submedian primary
septum; end cells broadly or narrowly rounded, concolorous
except for a truncate to convex hyaline terminal part of their
walls; in 3 % KOH turning olivaceous to grey-brown; surface
slightly verruculose.
Other material examined. AustRiA, Osttirol, Prägraten am Großvenediger,
Umbalfälle, grid square 8939/4, on Cytospora (Valsa) sp. on a branch of Rosa
canina, soc. Diplodia sp., 28 Aug. 2000, W. Jaklitsch W.J. 1539 (WU 37022).
Notes — Although ascospore size of Parafenestella faberi is
in the range of P. austriaca and P. rosacearum (see below), its
ascospores are unique due to the hyaline terminal wall of the
terminal cells and a uniform shape. Ascospore septa appear
rather distant in surface view, but dense in sectional view due to
strong superposition. Ascospore size varies slightly among as-
comata and specimens. We found the largest ascospores in the
isotype, and they were even up to 34.7 × 12.4 µm in the slides
prepared by R. Phookamsak. We synonymise P. mackenziei
with P. faberi here, as we do not see a difference between them.
The authors did not compare their new species with Fenestella
faberi, although the latter was redescribed by Phookamsak &
Hyde (2015); in illustrations of Wanasinghe et al. (2017b) up to
12 transverse septa including incomplete ones are discernible.
We give only a morphological account of this species here, as
our material was not cultured and sequenced.
Parafenestella germanica Jaklitsch & Voglmayr, sp. nov. —
MycoBank MB829751; Fig. 13
Etymology. For its occurrence in Germany.
Holotype. GeRmAny, Baden-Württemberg, Hornberg, Am Rubersbach, on
Diaporthe decedens on a branch of Corylus avellana, soc. ?Cosmospora sp.
and a dothideomycete with minute muriform spores, 18 Feb. 2018, K. Pätzold,
comm. B. Wergen (WU 37017; ex-type culture CBS 145267 = C307).
Ascomata (195–)230 – 450(–570) µm (n = 12) diam, globose
to subglobose, laterally collapsing from above, black, solitary
or in small groups on inner bark or on the ostiolar level of old
Diaporthe decedens, individually surrounded and connected by
hyaline to dark brown, thick-walled, 1.5–6.5 µm wide subicular
hyphae with often swollen and sometimes forked attachment
cells, near the ostiole often short with rounded ends (blunt
setae). Ostiolar areas c. 60–160 µm diam, indistinct or slightly
papillate, rounded, black, sometimes convergent in clustered
ascomata. Peridium 10– 60 µm thick, pseudoparenchymatous,
consisting of (4–)5.5 –10.5(–13.5) µm (n = 32) wide cells, out-
side thick-walled and dark brown, paler to hyaline and thinner-
walled to the inside. Hamathecium consisting of numerous 1–3
µm wide, branched ?paraphyses; in 3 % KOH swelling to c. 6 µm
when old. Asci (128–)140–173 (–193) × (17–)17.5–22 (–24.5)
µm (n = 25), cylindrical to oblong, bitunicate, fissitunicate, with
a distinct ocular chamber, a short stipe and simple or knob-like
base, containing 2–8 ascospores in (obliquely or overlapping)
uniseriate arrangement. Ascospores (25.5–)29 – 39.5(– 47)
× (11–)13–16.5(–19) µm, l/w (1.9–) 2.1– 2.5(– 2.7) (n = 70),
ellipsoid to broadly fusoid, symmetric, with the upper part
often broader, first hyaline, with 1–3 main septa, more or less
constricted at the median or slightly eccentric primary septum,
turning yellow to yellow-brown and finally dark brown when
mature, with often broadly rounded, paler to hyaline end cells,
sometimes larger terminal parts paler than the middle, with (8–)
9–13(–15) transverse and 3– 6 longitudinal septa; in 3 % KOH
turning dark to blackish brown, end cells remaining hyaline.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark up to 10 mm after
1 wk, 18 mm after 2 wk, 25 mm after 3 wk; colony first hyaline
to whitish, dense, turning olivaceous, later greyish brownish
due to a thick mat of aerial hyphae, surface and reverse be-
coming zonate; reverse dull grey to dark greyish olivaceous;
odour indistinct. Pycnidia appearing after 4 d, immersed to
erumpent, c. 70 –200 µm diam, more or less globose, first
hyaline, turning olivaceous to black, with whitish conidial drops.
Peridium thin, pseudoparenchymatous, olivaceous. Phialides
(3.2–)4.7–7.5(–9) × (1.7–)2 –3(– 4) µm (n = 30), sessile, sub-
globose to lageniform. Conidia (3.2–)3.5 – 4.3(– 4.7) × (1.1–)
1.2–1.4(–1.6) µm, l/w (2.5 –)2.7– 3.4 (–3.8) (n = 50), oblong
to ellipsoid, 1-celled, hyaline, with 1–2 minute drops, smooth;
also produced on pegs present below phialides. After a few
transfers no pycnidia formed.
Habitat — On Diaporthe decedens on Corylus avellana.
Distribution — Central Europe (Germany), only known from
the type locality.
Notes — Parafenestella germanica is phylogenetically clo-
sely related to P. pseudoplatani but has distinctly larger asco-
spores than the latter species.
23
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fig. 13 Parafenestella germanica. a– v. Sexual morph (WU 37017). a. Ascomata and subiculum in face view; b. ascoma above Diaporthe ascomata in vertical
section; c– f. asci (2-, 3-, 4- and 8-spored); g. peridium in vertical section; h. seta-like subicular hypha; i. ascus apex; j–v. ascospores (j – k. immature); w– z.
asexual morph in culture (CBS 145267 (C307) from CMD at 22 °C); w. pycnidia; x– y. phialides; z. conidia (d, h –i, u–v. in 3 % KOH). — Scale bars: a–b,
w = 150 µm; c– g = 15 µm; h– i, o– v = 10 µm; j –n = 7 µm; x –z = 5 µm.
Parafenestella parasalicum Jaklitsch & Voglmayr, sp. nov. —
MycoBank MB829752, Fig. 14
Etymology. Para = at, near; the epithet refers to the close phylogenetic
relationship with P. salicum.
Holotype.
AustRiA, Niederösterreich, Marchauen, Drösing, village area, on
branch of Salix cinerea, on/soc. Cytospora (Valsa) sp., 7 Apr. 2018, H. Vogl-
mayr (WU 37006; ex-type culture CBS 145271 = C318).
Ascomata 270–400 µm diam, globose, subglobose or pyriform,
black, immersed in bark in the ostiolar region of Cytospora (Val-
sa morph) or on inner bark, scattered, in valsoid configuration
or in rows in small numbers, forming groups 0.5–1.7 mm diam,
individually surrounded or connected by pale to dark brown,
thick-walled, 1.5–6 µm wide subicular hyphae, the latter
widened up to 10 µm at the connection to the peridium; incon-
24 Persoonia – Volume 44, 2020
Fig. 14 Parafenestella parasalicum. a–x. Sexual morph (WU 37006). a. Ascomata in face view and in obliquely vertical section; b. peridium and subiculum
hyphae in vertical section; c. hamathecium; d, i. ascus apices; e– h. asci (f. partially developed); j –x. ascospores (j, m. immature); y – c1. asexual morph in
culture (CBS 145271 (C318) from CMD at 22 °C); y. pycnidia and conidial drops; z –c1. conidia; d– e, j–l. from fresh material; x. in 3 % KOH. — Scale bars:
a = 300 µm; b, e– h = 20 µm; c– d, i– x = 10 µm; y = 100 µm; z–c1 = 3 µm.
spicuous at the bark surface, becoming visible in fissures. Osti-
olar areas (70–)75–138 (–160) µm (n = 10) diam, flattened, con-
vex or slightly papillate, shiny black, often mixed with minute
pycnidia. Peridium 15 –50 µm thick, pseudoparenchymatous,
consisting of isodiametric cells (3.5–)5.5 –11(–14) µm (n = 30)
diam, outside thick-walled and very dark brown, paler brown
to hyaline and thinner-walled toward the inner side and there
also with some brownish compressed elongate cells; darken-
ing in 3 % KOH. Hamathecium consisting of numerous, 1– 4
wide, branched ?paraphyses. Asci (176–)185– 219 (–239) ×
(20–)22 –27(– 30) µm (n = 12), cylindrical to oblong, bitunicate,
fissitunicate, with an ocular chamber, a short stipe and simple
or knob-like base, containing 4–8 ascospores in (overlapping,
obliquely) uniseriate to partly biseriate arrangement. Asco-
25
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fig. 15 Parafenestella pseudosalicis (WU 37016/CBS 145264 (C301)). a. Ascomata in bark fissures; b. apically depressed ascoma in oblique face view
and in vertical section; c. ascoma in vertical section; d. peridium in vertical section; e– g. asci (e. 4-spored, spore part); h–i. ascus apices (h. immature); j– v.
ascospores (j –l. immature; m. immature and mature mixed) (e–i, k– l, t –v. in 3 % KOH). — Scale bars: a = 300 µm; b = 200 µm; c = 100 µm; d, i = 10 µm;
e– g = 20 µm; h, j– v = 7 µm.
spores (33.5–)36 – 44(– 49.5) × (14.5 –)15.8–19.3(– 22.2) µm,
l/w (2–)2.1–2.4 (–2.6) (n = 44), fusoid or ellipsoid, first hyaline
and 2-celled, developing 2 additional main septa, turning yel-
lowish to yellow-brown and finally dark brown and eventually
with 11–16 distinct and densely inserted transverse and 3– 5
longitudinal septa; constricted at the primary septum, with the
upper part usually broader and often longer than the lower; ends
usually broadly rounded, concolorous, sometimes papillate and
paler to hyaline, smooth, containing numerous minute droplets;
in 3 % KOH turning dark to blackish brown.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 7– 8 mm after 1 wk,
22 mm after 22 d; colony first pale to medium grey, turning
dark olivaceous grey, centre black; aerial hyphae long, white,
forming a loose, later dense and thick mesh above the colony;
reverse dark grey to black; odour indistinct. Pycnidia c. 45–100
µm diam, (sub-)globose, papillate, first appearing after 3– 4 d,
hyaline, turning dark olivaceous, mostly immersed, spreading
from the centre, solitary or in firm small packs, with whitish turbid
conidial drops. Peridium thin, pseudoparenchymatous, oliva-
ceous, surround ed by olivaceous, partly submoniliform hyphae.
Phialides (4–)4.2 – 6.2(–7) × (1.8–) 2.2–3.5(– 4) µm (n = 16),
sessile, varying from subglobose over ampulliform or lageniform
to subulate. Conidia (3.3–)3.6 – 4.4(– 4.8) × (1–)1.1–1.5 (–1.8)
µm, l/ w (2.2–)2.6 – 3.4(– 4.1) (n = 24), cylindrical to allantoid,
less commonly narrowly ellipsoid, 1-celled, hyaline with 2 small
drops, smooth.
Habitat — On both morphs of a Cytospora (Valsa) sp. on
Salix cinerea.
Distribution — Central Europe (Austria), only known from
the type locality.
Notes — Parafenestella parasalicum is phylogenetically
and morphologically close to P. salicum, but differs from the
latter by larger ascospores.
26 Persoonia – Volume 44, 2020
Parafenestella pseudosalicis Jaklitsch & Voglmayr, sp. nov.
— MycoBank MB829753; Fig. 15
Etymology. Pseudo = false; the epithet denotes phylogenetic distinctness
from P. salicis despite morphological similarity.
Holotype. ukRAine, Ivano-Frankivsk region, Kosiv district, National Nature
Park ‘Hutsulshchyna’ (Carpathians), on twigs of Salix cf. alba, soc. Cytospora
sp., cf. Keissleriella sp., 5 Aug. 2017, A. Akulov (WU 37016; ex-type culture
CBS 145264 = C301).
Ascomata (270–)300 – 400(–420) µm (n = 11) diam, subglo-
bose to subpyriform, solitary or loosely or tightly aggregated
in small numbers, immersed in bark or on ascomata of an ef-
fete perithecial fungus, often with concave apex, individually
surrounded or connected by rather scant subhyaline to dark
brown, thick-walled, 2–5.5 µm wide subicular hyphae. Ostiolar
areas (53–)60 –116(–142) µm (n = 10) diam, indistinct and in-
conspicuous, concave or papillate, black. Peridium 15–55 µm
thick, pseudoparenchymatous, consisting of isodiametric cells
(3.5–)4 –8 (–12) µm (n = 32) diam, outside thick-walled and dark
brown, slightly paler and thinner-walled toward the inner side;
darkening in 3 % KOH. Hamathecium consisting of numerous,
1–4 wide, branched ?paraphyses. Asci (180–)186– 215(– 220)
× (17–)17.5 –19(–19.5) µm (n = 10), cylindrical to oblong, bi-
tunicate, fissitunicate, with an ocular chamber, a short stipe and
simple or knob-like base, containing 4–8 ascospores in uni-
seriate arrangement. Ascospores (23–)25 – 29(– 32) × (11–)
12–14(–15) µm, l/w (1.9–) 2–2.3 (–2.7) (n = 41), ellipsoid, first
hyaline to yellowish, with 1–3 main septa, turning yellow-brown
to dark brown, developing 7–10(–11) transverse and 2–4 longi-
tudinal septa, distinct in surface view, in section difficult to count
due to oblique superposition; constricted at the median primary
septum, upper part often wider, ends concolorous, wall smooth,
contents with minute guttules; in 3 % KOH turning very dark to
blackish brown, yellow-green to greyish brown when immature.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 6–7 mm after 1 wk,
15–16 mm after 2 wk; colony first hyaline to whitish, thick,
dense, turning olivaceous from the centre, becoming grey to
greyish brown due to a dense whitish to greyish villose mat of
aerial hyphae; reverse dark grey to black; odour indistinct. No
asexual morph detected.
Habitat — Associated with a perithecial fungus (probably
Cytospora, Valsa morph) on Salix cf. alba.
Distribution — Europe, only known from the type locality in
Ukraine.
Notes — Parafenestella pseudosalicis is morphologically
similar to P. salicis, but differs from that species by a more regu-
lar, symmetric ascospore shape and more longitudinal septa
being visible in surface view. Also, colonies on CMD tend to be
more distinctly brown than with other species from Salix, which
are more grey, and no asexual morph was produced in culture.
However, as the species is based on a single species, further
material would be necessary to evaluate its natural variation.
Parafenestella rosacearum Jaklitsch & Voglmayr, sp. nov. —
MycoBank MB829754; Fig. 16
Etymology. For its occurrence on various species of Rosaceae.
Holotype.
AustRiA, Vienna, 22nd district, Spargelfeldstraße, on ?Diaporthe
sp. on a branch of Pyracantha coccinea, soc. Diplodia sp., 25 Jan. 2018,
R. Moosbeckhofer, comm. B. Wergen (WU 37010; ex-type culture CBS
145268 = C309).
Ascomata (270–)285 – 432(– 510) µm (n = 42) diam, globose,
subglobose to subpyriform, dark brown to black, immersed on
often blackened inner bark or at the ostiolar level of Cytospora
spp. or other perithecial fungi, scattered or in small groups,
erumpent through bark fissures, laterally and basally loosely or
tightly connected by hyaline to pale brown, less commonly dark
brown, thick-walled, 2–5 µm wide subicular hyphae. Ostiolar
areas (53–)73–125 (–150) µm (n = 47) diam, inconspicuous
or papillate to short-cylindrical, sometimes depressed when
immature, sometimes mixed with slender ostiolar necks of
the host or often by pycnidia of its presumed asexual morph.
Peridium 15–50 µm thick, pseudoparenchymatous, consisting
of isodiametric cells (2.7–)4 –8(–12) µm (n = 30) diam, outside
thick-walled and dark brown, paler to hyaline and thinner-walled
to the inside; darkening in 3 % KOH. Hamathecium consisting
of numerous 1–3, basally to 5 µm wide branched paraphyses
with free ends. Asci (150–)181–240 (–290) × (17–)19– 22(–24)
µm (n = 59), cylindrical to oblong, bitunicate, fissitunicate,
with a distinct ocular chamber, a short, sometimes contorted
stipe and simple or knob-like base, containing (2–)4 –8 asco-
spores in uniseriate, rarely partly biseriate arrangement. Asco-
spores (23.3–)28 – 35(– 44.5) × (11–)13.5–16.5 (–19.5) µm,
l/w (1.7–)1.9– 2.2(–2.6) (n = 230), ellipsoid, symmetric to in-
equilateral, constricted at the more or less median primary
septum, initially hyaline to yellowish, with 1–3 main septa,
developing (7–)9 –13(–15) transverse and 2 – 4(– 5) longitu-
dinal septa and turning yellow-brown, pale-, medium- to dark
brown, ends broadly rounded and concolorous, less commonly
narrowly rounded and slightly paler, surface appearing slightly
verruculose; in 3 % KOH turning olivaceous when young, black-
ish brown when mature.
Culture characteristics and asexual morph in culture — Asco-
spores germinating simultaneously from many cells. Colony
radius on CMD at 22 °C in the dark 7–9 mm after 1 wk, up to
18 mm after 2 wk, 35 mm after 5 wk; colony first hyaline, turning
greyish oliva ceous, dense, usually becoming thick and pale grey
by a dense mesh of white aerial hyphae, long remaining pale
greyish brownish olivaceous, finally darkening; reverse dark
grey to black; odour indistinct. Pycnidia appearing after 4 d,
c. 80–200 µm diam, more or less globose, olivaceous, often
numerous, aggregated, immersed in the agar and usually com-
pletely covered by a dense mat of aerial hyphae, with whitish
conidial drops. Peridium thin, pseudoparenchymatous, brown
to olivaceous, surrounded by submoniliform hyphae. Phialides
(2.7–)4.7– 8.2(–9.7) × (2–) 2.5 –4(– 5.2) µm (n = 25), sessile,
varying from subglobose over ampulliform or lageniform to
subulate; conidia also formed on lateral pegs below phialides.
Conidia (3–)3.5 –4.2(–4.8) × (1.1–)1.2 –1.5(–1.7) µm, l/w (2 –)
2.5–3.3 (–4.1) (n = 55), cylindrical or oblong, less commonly
ellipsoid, 1-celled, hyaline, with 1–3 drops, smooth.
Habitat — Associated with Cytospora spp. and other peri-
thecial fungi on various species of Rosaceae, recorded from
Crataegus, Prunus, Pyracantha, Pyrus, Rosa and Sorbus aria.
Distribution — Central Europe (Austria).
Other materials examined. AustRiA, Burgenland, Purbach, Purbacher Heide,
on Cytospora sp. on a branch of Rosa canina, 24 Mar. 2018, H. Voglmayr
(WU 37011; culture C315); Siegendorf, Königschüssel, on effete perithecia
of ?Diaporthe sp. on Crataegus monogyna, 18 Sept. 2016, W. Jaklitsch &
H. Voglmayr (WU 37008; culture C269); Niederösterreich, Ulrichschlag, on
Cytospora sp. on an attached branch of Pyrus communis, soc. Diplodia sp., 11
July 2015, W. Jaklitsch & H. Voglmayr (WU 37007; culture C203); Oberöster-
reich, Schärding, Raab, Wetzlbach, on a branch of Prunus domestica, soc.
diverse fungi including a Botryosphaeria sp., 2 Dec. 2017, H. Voglmayr (WU
37013; culture CBS 145272 = FP11); Steiermark, N Rein, Kaschsteig, on a
branch of Sorbus aria, 30 June 2018, G. Friebes (WU 37023; culture C320);
Ratten, Kirchenviertel, vlg. Kirchenberger, beim ‘Dörrofen’, on a branch of
Pyrus communis, soc. Acanthostigma sp., ‘Cucurbitaria acervata’, a perithe-
cioid fungus with yellow muriform ascospores and diverse pycnidia, 6 Jan.
2017, R. Moosbeckhofer (WU 37009; culture C283); Vienna, 21st district, at
Marchfeldkanalweg near Felix Slavikstraße, on Cytospora sp. on a branch of
Rosa canina, soc. Calosphaeria sp., Massarina sp., 8 Nov. 2015, W. Jaklitsch
(WU 37012; culture FM1).
Notes — As deduced from the protologue, P. rosacearum
may be what Saccardo (1884) termed Cucurbitaria delitescens
27
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fig. 16 Parafenestella rosacearum. a– y. Sexual morph. a–b. Ascomata in face view; c. ascoma in vertical section; d– h. asci (d– e. from fresh material; f. im-
mature); i. basal peridium in vertical section; j. immature ascus apex and apically free paraphysis; k– l. ascus apices (k. from fresh material); m– y. ascospores
(m– o. immature; p– s. from fresh material); z– b1. asexual morph from CMD at 22 °C; z– a1. phialides; b1. conidia (j, n. in 3 % KOH). a, c, f, z– b1. WU 37010/
CBS 145268 (C309); b, d– e, i–k, p– s, y. WU 37013 (FP11); g, t, w. WU 37008 (C269); h, v. WU 37009 (C283); j, n. WU 37007 (C203); l, m, u, x. WU 37012
(FM1); o. WU 37011 (C315). — Scale bars: a–b = 250 µm; c = 150 µm; d– g = 25 µm; h–i = 15 µm; j, q –r, u–y = 10 µm; k– p, s–t = 7 µm; z –b1 = 5 µm.
*prunorum. However, we have not seen type material and a
varietal name is not binding. Parafenestella rosacearum is a
complex species. In spite of splitting into two or three groups in
multigene analyses (see Fig. 1), we recognise a single species,
because there are no morphological differences among those
groups and particularly due to the following observations: tef1
sequences of C203 and C283 are identical and tef1 of C309 is
nearly identical with them, whereas rpb2 sequences of C203,
C315, FM1 and FP11 are virtually identical, while those of C269
and C283, which are identical, differ from the first group by c.
20 nucleotides. This finding was verified by repetition of DNA
amplifications and sequencing. Species of Parafenestella on
Rosaceae, particularly on Rosa spp., are difficult to distinguish
morphologically; ascospores of P. faberi are characteristic due
28 Persoonia – Volume 44, 2020
to their often truncate hyaline terminus of the end cells, and
P. austriaca differs from P. rosacearum by a rather invariable
ascospore shape. In both of the latter species ascospore end
cells are rounded, slightly projecting and may be only diffusely
paler to hyaline.
Parafenestella salicis (Rehm) Jaklitsch & Voglmayr, comb.
nov. — MycoBank MB829755; Fig. 17
Basionym. Thyridium salicis Rehm, in Thümen, Beiträge zur Pilzflora
Sibiriens IV, Bull. Soc. Imp. Naturalistes Moscou 55 (1–4): 218. 1880.
Synonyms. Fenestella salicis (Rehm) Sacc., Syll. Fung. (Abellini) 2: 330.
1883.
Cucurbitaria marchica Kirschst., Ann. Mycol. 34: 186. 1936.
Fig. 17 Parafenestella salicis. a– v. Sexual morph. a. Ascomata in face view; b. ascoma in oblique view and in vertical section; c– g. asci (c. from fresh mate-
rial); h. peridium in vertical section; i– j. ascus apices (i. young); k –v. ascospores (k –l. immature / young; m –p. from fresh material); w– a1. asexual morph
from CMD at 22 °C; w. pycnidia; x–y. phialides and short conidiophores; z–a1. conidia (g, v. in 3 % KOH). a– f, h, j– k, s, w–a1. WU 37002/CBS 145270
(C313); g, r. K(M) 251618; i, l–n. WU 37003 (C311); o– p, u. WU 37001 (C303); q. part of WU 37015; t, v. C. marchica holotype B 700016516. — Scale bars:
a– b, w = 250 µm; c– h = 20 µm; i, k– v = 7 µm; j = 10 µm; x–a1 = 5 µm.
29
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Typification. According to the herbarium curator of LE, Dr. Olga Morozova,
comm. Dr. Eugene Popov, there is only a younger specimen of the original
collector, N. Martianov, but not the type, extant in LE. Also, no type material
has been found in other herbaria. Therefore we here propose a neotype
for Thyridium salicis: AustRiA, Niederösterreich, Marchegg, on Cytospora
(Valsa) sp. on branchlets of Salix alba, soc. a coniothyrium-like fungus, 17
Mar. 2018, H. Voglmayr (WU 37002; MBT385689; ex-neotype culture CBS
145270 = C313).
Ascomata (255–)275 – 442(–510) µm (n = 20) diam, globose,
subglobose to pyriform or subconical, black, immersed below
the epidermis on inner bark or in the ostiolar region of Cytospora
(Valsa) sp., scattered or tightly or loosely aggregated in valsoid
or irregular configuration, partly erumpent through bark fissures,
surrounded laterally and basally and connected by subhyaline
to medium brown, thick-walled, 2–5 µm wide subicular hyphae.
Ostiolar areas (35–)55 –118(–180) µm (n = 22) diam, usually
indistinct, sometimes papillate, rounded or apically flattened or
compressed and furrowed, black. Peridium 20–80 µm thick,
pseudoparenchymatous, consisting of (4–)6 –13(–18) µm
(n = 30) wide cells, outside thick-walled and very dark brown,
gradually paler and thinner-walled to the inside. Hamathecium
consisting of numerous, 1–3.5 µm wide, branched ?paraphy-
ses; when old widened up to c. 6 µm and submoniliform at their
bases. Asci (131–)141–188(–220) × (14.8–)16–19 (–21.5) µm
(n = 33), cylindrical to oblong, bitunicate, fissitunicate, with a
distinct ocular chamber, a short stipe and simple or knob-like
base, containing (1–)4 –8 ascospores in (obliquely) uniseriate to
partly biseriate arrangement. Ascospores (20.5–)23 –29(– 34)
× (10–)11–13.5(–15) µm, l/w (1.5 –)1.9–2.3 (–2.7) (n = 152),
ellipsoid to fusoid, slightly to distinctly constricted at the more
or less median primary septum, often also at other septa, sym-
metric to inequilateral, first hyaline to yellowish, with 1–3 main
septa, turning bright or golden yellow to yellow-brown (when
fresh), eventually dark brown, with (5–6 –)7–10(–11) transverse
and (1–)2 – 3 longitudinal septa, the former often with oblique
superposition in section, the latter often forming parallel lines;
ends broadly rounded and concolorous, less commonly nar-
rowly rounded and only rarely hyaline immediately before
germination, smooth. Often tightly accompanied by the pre-
sumed asexual morph forming pycnidia with minute cylindrical
to allantoid, 1-celled, hyaline conidia.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 9 –11 mm after 1 wk,
22 mm after 2 wk; colony dense, first hyaline, turning olivaceous
and later pale to dark grey owing to aerial hyphae; reverse
dark grey to black; odour indistinct to sour. Pycnidia appearing
after 3 d, c. 80– 200 µm diam, numerous, immersed, partially
erumpent, scattered or tightly aggregated to confluent in small
numbers, globose or subglobose, papillate, olivaceous to
black, with whitish to greyish conidial drops, partly covered
by aerial hyphae, surrounded by olivaceous to brown hyphae.
Peridium thin, pseudo parenchymatous, olivaceous. Phialides
(4.3–)5.3 –9(–10.5) × (1.8 –)2 – 3(– 3.8) µm (n = 22), subglo-
bose to subcylindrical, sessile or formed on short conidiophores;
conidia also formed on pegs. Conidia (3.2–)3.7– 4.5(–5.2) ×
(1–)1.1–1.5(–1.7) µm, l /w (2.1–)2.7– 3.6(– 4.2) (n = 64), cy-
lindrical to allantoid, sometimes narrowly ellipsoid, 1-celled,
hyaline, with 1–2 subterminal drops, smooth.
Habitat — On both morphs of Cytospora (Valsa) sp(p). on
Salix spp.
Distribution — Europe (Austria, Germany, UK), Russia, etc.
Other materials examined. AustRiA, Niederösterreich, Orth, on Cytospora
(Valsa) sp. on branchlets of Salix alba, soc. Diplodia sp., 3 Feb. 2018, H. Vogl-
mayr (WU 37001; culture C303); same area, on Cytospora sp. on branches
of Salix alba, soc. Coniothyrium sp., Parafenestella salicum (removed as WU
37004), 10 Mar. 2018, W. Jaklitsch & H. Voglmayr (WU 37003). – GeRmAny,
Westhavelland, Quermathen bei Großbehnitz, Löffelpfuhl, on branches of
Salix cinerea, 4 Nov. 1935, W. Kirschstein (B 70 0016516; holotype of Cucur-
bitaria marchica). – uk, England, Kew, on twigs of Salix cinerea, apparently
on Plagiostoma sp., May 1887, no collector given (K-M 251618).
Notes — Judging on the specimens we have seen, P. salicis
is the most common species of the genus on Salix spp. and oc-
curs often together with other Parafenestella species. Counting
of ascospore septa in sectional view is particularly difficult in
this species, due to conspicuous superposition; they differ from
those of P. salicum and P. parasalicum in less and more widely
spaced septa and from the latter also by size. Ascospores are
often inequilateral and often slightly constricted at other than
the primary septum, traits not seen with P. pseudosalicis. Like
with other species, mature ascospores are often dark reddish
brown in herbarium material.
Other species/ names described on Salix: Cucurbitaria salicina
was described from Salix fragilis and S. triandra with asco-
spores measuring 22 × 9 µm having 4–5 transverse septa
and 1 longitudinal septum. The holotype (Germany, Oestrich,
L. Fuckel, G 00127757, ex Herbier Barbey-Boissier, Nassau’s
Flora. 6) does not contain a sexual morph but numerous
pycnidia with camarosporium-like conidia and a Diplodia sp.
Cucurbitaria cinerea, described from Salix aurita, is Disco-
stroma corticola, according to Brockmann (1976). Cucurbitaria
rubefaciens, described from Salix caprea, does not belong to
the Cucurbitariaceae (unpublished results).
Parafenestella salicum Jaklitsch & Voglmayr, sp. nov. — Myco-
Bank MB829756; Fig. 18
Etymology. For its occurrence on species of Salix.
Holotype.
AustRiA, Niederösterreich, Orth, on thin branches of Salix alba,
soc. effete Cytospora sp., Massarina sp., Parafenestella salicis (removed as
WU 37003) and effete Plagiostoma sp., 10 Mar. 2018, W. Jaklitsch & H. Vogl-
mayr (WU 37004; ex-type culture CBS 145269 = C311).
Ascomata (270–)280 – 375(– 420) µm (n = 11) diam, globose,
subglobose or pyriform, black, laterally collapsing, immersed
in bark over a perithecial host fungus or firmly connected to the
inner bark layers, scattered or aggregated in small numbers
in valsoid groups, connected by subhyaline to dark brown,
thick-walled, 1.5–5 µm wide subicular hyphae. Ostiolar areas
85–200 µm diam, inconspicuous at the bark surface, roundish
to longish, black, often apically flattened, sometimes concave
or papillate; sometimes accompanied by pycnidia of the puta-
tive asexual morph. Peridium 15–65 µm thick, pseudoparen-
chymatous, consisting of (3.5–)6 –10(–12.5) µm (n = 40) wide
cells, outside thick-walled and very dark brown and forming a
narrow layer, gradually paler to hyaline and thinner-walled to the
inside, terminated by a narrow layer of brownish compressed
cells. Hamathecium consisting of numerous 1– 3.5(–4) µm wide,
branched ?paraphyses. Asci (172–)181–228 (–246) × (17.5–)
19.5–24(–28) µm (n = 20), cylindrical, bitunicate, fissitunicate,
with a distinct ocular chamber, a short stipe and simple or
knob-like base, containing 6–8 ascospores in (overlapping)
uniseriate arrangement. Ascospores (22.5–)27– 33(– 40) ×
(11–)12.5–16(–18) µm, l/w (1.7–)1.9– 2.3 (–2.5) (n = 110),
broadly ellipsoid to broadly fusoid with broadly rounded ends
or one end narrowly rounded, upper part often slightly wider,
not or slightly constricted at the median primary septum, first
2-celled and hyaline, turning golden yellow to golden brown,
eventually dark brown or dark reddish brown, ends concolor-
ous, only rarely slightly paler and sometimes slightly projecting,
with 9–13(–14) transverse and 3 –4 longitudinal septa; smooth.
Presumed asexual morph forming pycnidia at the apices or
sides of ascomata, c. 70–150 µm diam, more or less globose,
with masses of oblong, allantoid to narrowly ellipsoid, 1-celled,
hyaline conidia typically containing two drops.
Culture characteristics and asexual morph in culture — Asco-
spores germinating simultaneously from many cells. Colony
radius on CMD at 22 °C in the dark 5–7 mm after 1 wk, 16 mm
after 2 wk, 18 mm after 27 d; colony pale brownish to pale grey-
30 Persoonia – Volume 44, 2020
Fig. 18 Parafenestella salicum. a– t. Sexual morph. a. Ascomata in face view; b, g. ascomata in oblique view and in vertical section (detached in g); c–f. asci
(c. from fresh material; d. young; e. early stage on the left side); h. peridium in vertical section; i. subicular hyphae; j– k. ascus apices; l–t. ascospores (l– m.
from fresh material); u– y. asexual morph from CMD at 22 °C; u. pycnidia; v. submoniliform hyphae; w. peridium in face view; x. phialides; y. conidia. a– g, i–r,
u– y. WU 37004 /CBS 145269 (C311); h, s–t. WU 37005. — Scale bars: a –b, g = 150 µm; c–f, h = 20 µm; i –k, p–q, v –w = 10 µm; l– o, r–t = 7 µm; u = 300
µm; x– y = 5 µm.
ish brown, covered by a dense and thick mat of aerial hyphae;
reverse dark grey; odour indistinct to unpleasant. Pycnidia
appearing after 3 d, numerous, immersed, partially erumpent,
80–180 µm diam, solitary, in lines or tightly aggregated in
small numbers, globose, subglobose or vertically elongated,
papillate, greenish, olivaceous to black, with narrow periphy-
ses in the ostiole; conidial drops whitish to greyish. Peridium
thin, pseudoparenchymatous, an olivaceous textura angularis
of (6.5–)8.5 –14.5(–19) µm (n = 30) wide cells; surrounded
by submoniliform hyphae. Phia lides (4–)4.5 – 6.2(–7.3) ×
(1.5–)1.8– 3.5(–4.7) µm (n = 20), sessile, subglobose, ampul-
liform, lageniform to subcylindrical. Conidia (3.3–)3.5 –4(–4.5)
31
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
× (0.9–)1.1–1.6(– 2) µm, l/w (1.9–)2.4 –3.4(– 4.1) (n = 42),
cylindrical to allantoid, sometimes nar-rowly ellipsoid, 1-celled,
hyaline, with 2 subterminal drops, smooth.
Habitat — On thin branches of Salix alba, presumably on
both morphs of a Cytospora (Valsa) sp.
Distribution — Europe (Austria).
Other material examined. AustRiA, Vienna, 22nd district, Lobau, Panoz-
zalacke, grid square 7865/1, on thin branches of Salix alba, soc. effete
Cytospora sp., Keissleriella holmiorum, 8 Feb. 1997, W. Jaklitsch W.J. 1015
(WU 37005).
Notes — Parafenestella salicum differs from P. salicis by
larger asci, larger, particularly broader ascospores with a larger
number of septa. While phylogenetically closely related to
Fig. 19 Parafenestella tetratrupha. a– y. Sexual morph. a. Ascomata and subiculum in face view; b. cluster of ascomata in conidioma of Melanconis alni (grey
patches at bottom and left are host conidia); c. young ascomata in conidioma of Melanconis alni; d– h. asci (f. upper part, partially developed, apically opening);
i. ascomatal setae; j. peridium in vertical section (showing host conidia); k–m. ascus apices; n–y. ascospores (n. immature); z–c1. asexual morph from CMD
at 22 °C; z. pycnidia and conidial drops; a1. phialide and basal peg; b1–c1. conidia (d –e, i– j, m, r, x– y. in 3 % KOH). a, d– e, i, l–m, w, y. WU 37000; b, g–h,
s– v, x. Valsa tetratrupha holotype K-M 251617; c, f, j, n– o. Fenestella minor holotype PC 0706652; k, p– r, z– c1. WU 36999/CBS 145266 (C304). — Scale
bars: a– c, z = 200 µm; d– e, g– j = 20 µm; f, k–m, o –q, t– v, y = 10 µm; n, r– s, w– x = 7 µm; a1–c1 = 5 µm.
32 Persoonia – Volume 44, 2020
P. parasalicum, P. salicum differs from that species by smaller
ascospores, although some old aberrant ascospores may
approach those of P. parasalicum in size. The fungal host of
P. salicum is not unequivocally clear. As is usual in the Cucur-
bitariaceae, ascospores are more reddish brown in herbarium
material.
Parafenestella tetratrupha (Berk. & Broome) Jaklitsch &
Voglmayr, comb. nov. — MycoBank MB829757; Fig. 19
Basionym. Valsa tetratrupha Berk. & Broome, Ann. Mag. Nat. Hist., ser.
III, 3: 366. 1859.
Synonyms. Fenestella tetratrupha (Berk. & Broome) Sacc., Syll. Fung.
(Abellini) 2: 326. 1883.
Fenestella minor Tul. & C. Tul., Select. Fung. Carp. 2: 207. 1863.
Holotype. uk, England, North Somerset, Batheaston, on conidiomata
of Melanconis alni on twigs of Alnus sp. (probably A. glutinosa), Feb. 1852,
C.E. Broome (K-M 251617, as Fenestella minor written on the outer, Spha-
eria tetratrupha on the inner label). Epitype of Fenestella minor and Valsa
tetratrupha, here designated: AustRiA, Kärnten, St. Margareten im Rosental,
village area, at the brook Tumpfi, on conidiomata of Melanconis alni on a
branch of Alnus glutinosa, partly overgrown by Fusarium/Cosmospora (s.lat.)
sp., 4 Feb. 2018, W. Jaklitsch (WU 36999; MBT385691, MBT385690; ex-
epitype culture CBS 145266 = C304).
Ascomata (240–)300 –500(– 630) µm (n = 31) diam, globose,
subglobose or pyriform, immersed in the ostiolar region of Me-
lanconis alni conidiomata singly or in more or less valsoid
groups with often convergent ostiolar necks, tightly or loosely
aggregated in whitish to dark brown subiculum, forming pustules
c. 0.5–2.3 mm depending on the size of the host, projecting
up to 0.6 mm from the bark surface. Pustules pulvinate, caus-
ing small bumps in elevated bark, erumpent through fissures,
outline circular, elliptic or elongate, surface obtuse, dark brown
to black, often partly covered by brown subiculum consisting
of subhyaline to dark brown, thick-walled, 1.5–6 µm wide hy-
phae, mixed with dark brown, pointed, thick-walled, up to 60
µm long and 2–6 µm wide setae on the ascomatal surface.
Ostiolar areas 70–186(– 270) µm (n = 17) diam, inconspicuous
or papillate, sometimes eccentric and convergent, rounded or
flat and angular, brown to black, whitish inside, often mixed
with shiny black ostiolar necks of the host. Peridium 15–100
µm thick, pseudoparenchymatous, consisting of isodiametric
cells (4–)6 –11(–13.5) µm (n = 30) diam, outside moderately
thick-walled and dark brown, paler and thinner-walled to the
inside. Hamathecium consisting of numerous 0.5– 3.5 µm wide
branched ?paraphyses. Asci (141–)154–229 (–294) × (16.8–)
18.5–22.2 (–26) µm (n = 50), cylindrical to oblong, bitunicate,
fissitunicate, with a truncate ocular chamber, a short stipe
and simple or knob-like base, containing 2–8 ascospores in
uniseriate arrangement. Ascospores (19–)26.5 – 33.5(–39) ×
(10–)13–16.5 (–18) µm, l/w (1.7–)1.9 – 2.3(–2.8) (n = 120),
ellipsoid with broadly rounded ends or upper end narrowly
rounded, sometimes broadly fusoid, first hyaline, with 1–3 main
septa, turning yellow to yellow- or pale brown and finally red-
dish brown to dark brown with concolorous, rarely paler ends
and 8–14(–17) distinct transverse and 2 –4 longitudinal septa;
constricted at the median primary septum and upper part often
slightly wider; surface finely verruculose; in 3 % KOH becoming
dark olivaceous to blackish brown.
Culture characteristics and asexual morph in culture — Asco-
spores germinating simultaneously from many cells. Colony
radius on CMD at 22 °C in the dark 9 mm after 1 wk, 27 mm
after 24 d; colony olivaceous, but in face view appearing pale
grey to greyish brown due to a thick and dense mat of whitish
aerial hyphae; reverse dark grey; odour indistinct. Pycnidia
c. 100 –200 µm diam, subglobose, olivaceous, completely co-
vered by aerial hyphae, scattered, immersed, partly erumpent,
with whitish conidial drops; spreading from the centre. Peridium
thin, pseudoparenchymatous, olivaceous. Phialides (3.5–)4.5 –
6.7(–7.5) × (1.7–)2.2 – 4(– 4.2) µm (n = 21), sessile, varying
from subglobose over ampulliform and lageniform to subulate;
conidia also formed on lateral pegs. Conidia (3.2–)3.5 – 4.3(–6)
× (0.9–)1–1.2(–1.4) µm, l/w (2.8 –)3.2–4.1(– 4.7) (n = 50), cy-
lindrical to allantoid, rarely narrowly ellipsoid, 1-celled, hyaline,
with 2 small drops, smooth.
Habitat — On or in conidiomata of Melanconis alni on Alnus
glutinosa.
Distribution — Europe (Austria, France, UK).
Other materials examined. AustRiA, Kärnten, St. Margareten im Rosental,
village area, at the brook Tumpfi, grid square 9452 /4, on conidiomata of
Melanconis alni on a branch of Alnus glutinosa, 17 Dec. 1994, W. Jaklitsch
W.J. 377 (WU 37000). – FRAnCe, Chaville, on conidiomata of Melanconis
alni on Alnus glutinosa, soc. Cytospora sp., Apr. 1860, L.-R. Tulasne (PC
0706652, holotype of Fenestella minor; donated to PC in 1873 as Sphaeria
fenestrata deminuta in herb).
Notes — The fungal host of P. tetratrupha is clearly Melan-
conis alni, usually present as asexual morph and easily iden-
tifiable by the white central stromatic column and the brown
conidia having a pale median band. Conidia of the host are
present directly below ascomata in all studied specimens and
usually adhere in masses to the peridium. The holotype of
Fenestella minor has scant and mostly immature material,
whereas the holotype of Valsa tetratrupha has well-developed
ascomata with mostly 8-spored asci, although 4-spored asci are
given in the protologue. The number of ascospores in the ascus
depends much of the developmental condition of ascomata.
Parafenestella vindobonensis Jaklitsch & Voglmayr, sp. nov.
— MycoBank MB829758; Fig. 20
Etymology. For its occurrence in Vienna, Austria.
Holotype.
AustRiA, Vienna, 21st district, Wasserpark, on Cytospora sp. on
Salix babylonica, soc. Massarina sp., Parafenestella salicis and a pycnidial
fungus with fusoid, hyaline, 1-celled conidia, 27 Jan. 2018, W. Jaklitsch (WU
37015; ex-type culture CBS 145265 = C302).
Ascomata (240–)308 – 425(– 450) µm (n = 13) diam, globose,
subglobose or pyriform, black, immersed in bark, partially
erumpent, forming more or less valsoid groups in usually loose
connection by hyaline to dark brown, thick-walled, 2–5 µm
wide subicular hyphae or entirely covered by subiculum, some-
times tightly aggregated in small groups on inner bark and on
conidiomata and pseudostromata of a Cytospora sp., partly
blackening inner bark. Ostiolar areas (70–)75–145 (–160) µm
(n = 11) diam, indistinct or papillate, black, rounded or angular
in section. Peridium 15–65 (–80) µm thick, pseudoparenchy-
matous, consisting of (4–)5 –10(–12) µm (n = 40) wide cells,
outside thick-walled and dark brown, paler and slightly thinner-
walled to the inside. Hamathecium consisting of numerous
1–3 µm wide, branched paraphyses with free ends. Asci
(162–)179– 214(–228) × (12.5–)13.5 –15.5(–16) µm (n = 25),
cylindrical, bitunicate, fissitunicate, with a distinct ocular cham-
ber, a short stipe and simple or knob-like base, containing 4– 8
ascospores in uniseriate arrangement. Ascospores (20–)24.5 –
30.5(– 37.5) × (8.5 –) 9.5–11(–13) µm, l/w (2 –)2.4 – 2.9(– 3.4)
(n = 85), oblong, fusoid or narrowly ellipsoid, first hyaline, with
1–6 main septa, turning yellowish, pale, yellow- to golden
or medium brown with concolorous or slightly paler ends,
thick-walled, when mature with 7–9(–11) thick transverse and
1–2(– 3) septa, constricted at the median or supramedian pri-
mary septum, smooth, containing minute droplets; in 3 % KOH
turning greenish to yellow-brown when young, dark olivaceous
when mature.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 7–8 mm after 1 wk,
17–18 mm after 2 wk, 28 mm after 25 d; colony first hyaline to
33
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fig. 20 Parafenestella vindobonensis. a– e1. Sexual morph (WU 37015). a– b. Ascomata in face view; c– d. ascomata in vertical section (with subicular disc
in c; on Valsa ascomata in d); e– g. asci (note paraphysis in e); h. peridium in vertical section; i. aged hamathecium; j–m. ascus apices (immature in j –l);
n– e1. ascospores (n–o. immature); f1–l1. asexual morph in culture (CBS 145265 (C302) from CMD at 22 °C); f1. pycnidia; g1–h1. phialides; i1 –l1. conidia
(e– g, j–n, y–e1. in 3 % KOH). — Scale bars: a–d = 250 µm; e –h = 20 µm; i, q, s–v, e1 = 10 µm; j–p, r, w, x–d1, g1– h1 = 7 µm; f1 = 150 µm; i1–l1 = 3 µm.
34 Persoonia – Volume 44, 2020
whitish, thick, dense, turning olivaceous, later greyish brown
from the centre, covered by a velvety mat of whitish to brown-
ish aerial hyphae; reverse dark grey to black; odour indistinct.
Pycnidia appearing after 4 d, mostly solitary, greenish; forma-
tion of pycnidia not reproducible on CMD. Colony on MEA grey,
aerial hyphae forming a loose mesh over numerous pycnidia
aggregating in dense masses. Pycnidia c. 60–130 µm diam,
hyaline, turning olivaceous from the base, globose, with grey-
ish olivaceous conidial drops. Peridium thin, pseudoparen-
chymatous, olivaceous, surrounded by submoniliform hyphae.
Phialides (3.8–)5–7.5(–9.5) × (2 –)2.5 – 4(–4.8) µm (n = 39),
sessile, varying from subglobose to lageniform, often with a
long neck. Conidia (2.8–)3.5 –4(–4.7) × (1.1–)1.3–2 (–2.7) µm,
l/ w (1.4–)1.9–2.7(– 3) (n = 42), oblong to ellipsoid, 1-celled,
hyaline, with two or more minute drops, smooth.
Fig. 21 Synfenestella pyri. a –t. Sexual morph (WU 36996). a. Ascomata in face view; b– c. ascomata in vertical section (c. in a Cytospora (Leucostoma)
pseudostroma); d. hamathecial threads; e– f. ascus apices; g –i. asci (g. immature; note sheaths around ascospores); j. peridium and subiculum in vertical
section; k– t. ascospores (k. showing size variation); u–w. asexual morph in culture (CBS 144855 (C297) from CMD at 22 °C); u. pycnidia and conidial drops;
v– w. conidia (f, i, t. in 3 % KOH). — Scale bars: a– c, u = 250 µm; d–f, l– t = 10 µm; g–i = 25 µm; j –k = 15 µm; v– w = 3 µm.
35
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Habitat — On both morphs of a Cytospora (Valsa) sp. on
Salix babylonica.
Distribution — Central Europe (Austria), only known from
the type locality in Vienna.
Notes — This species is well characterised by its narrow
ascospores.
Synfenestella Jaklitsch & Voglmayr, gen. nov. — MycoBank
MB829759
Etymology. Syn = together with, for its close phylogenetic relationship
with Fenestella.
Type species. Synfenestella sorbi Jaklitsch & Voglmayr.
Ascomata 300–1000 µm diam, globose, subglobose or pyri-
form, ostiolate, dark brown to black, immersed below bark
epidermis, scattered, forming inconspicuous valsoid groups
or conspicuous pseudostromatic pustules on pseudostromata
or conidiomata of Diaporthales, surrounded and connected by
thick-walled subicular hyphae, the latter sometimes short and
subsetose in the ostiolar region. Peridium c. 15–130 µm thick,
thicker and whitish inside in the apical region, pseudoparen-
chymatous, dark brown and thick-walled outside and pale to
hyaline and thin-walled inside. Hamathecium consisting of
numerous 1–3.5 µm wide, branched and anastomosing para-
physes with free ends in a matrix. Asci cylindrical to oblong,
bitunicate, fissitunicate, with an ocular chamber, a short or long
undulating stipe and simple or knob-like base, containing 4–8
ascospores in (obliquely) uniseriate arrangement. Ascospores
ellipsoid, oblong to fusoid, sometimes subglobose, symmetric
to slightly curved, with upper part often wider, initially 2-celled,
hyaline and surrounded by a swelling sheath, developing
additional transverse and longitudinal septa, turning yellow
to golden brown (when fresh), finally dark brown with con-
colorous rounded ends, ends sometimes paler or apiculate
upon germination, usually strongly constricted at the primary
septum; in 3 % KOH surface smooth; turning dark olivaceous
to blackish brown.
Culture characteristics and asexual morph in culture — Co-
lony on CMD at 22 °C in the dark typically producing a diffusing
yellow pigment. Pycnidia more or less globose, green to black,
densely aggregating and fusing. Phialides lageniform, ampul-
liform, subglobose or subconical, sessile or produced on short
simple conidiophores. Conidia cylindrical, oblong to allantoid,
less commonly ellipsoid, 1-celled, hyaline, with 2 drops, smooth;
pro duced on phialides and pegs.
Habitat — On or in pseudostromata or conidiomata of Dia-
porthales or in loose association with them on Rosaceae.
Distribution — Europe, North America.
Synfenestella pyri Jaklitsch & Voglmayr, sp. nov. — MycoBank
MB829760; Fig. 21
Etymology. Referring to its plant host, Pyrus communis.
Holotype. AustRiA, Niederösterreich, Ottenschlag, on Cytospora (Leuco-
stoma) sp. on a branch of Pyrus communis, 19 Mar. 2017, W. Jaklitsch (WU
36996; ex-type culture CBS 144855 = C297).
Ascomata (330–)370 – 620(–720) µm (n = 21) diam, subglo-
bose to globose, immersed in bark singly or in small groups
c. 0.6 –1.6 mm diam, on or in loose association with both morphs
of a Cytospora (Leucostoma type) sp., on and connected by
subhyaline to dark brown, thick-walled, 2–5 µm wide subicular
hyphae, often only visible on the surface due to spore deposits
in bark fissures or slightly erumpent, rarely projecting up to 350
µm above the bark surface. Ostiolar areas 90–270 µm diam,
concealed by spore deposits or short-papillate, black, shiny.
Peridium 15–70 µm thick, thicker and whitish inside in the osti-
olar region, pseudoparenchymatous, consisting of isodiametric
cells (3.8–)5 –9 (–11) µm (n = 40) diam, outside thick-walled and
dark brown, paler to hyaline and thinner-walled to the inside.
Hamathecium consisting of numerous 1–3.5 µm wide, branched
and anastomosing paraphyses with free ends in a matrix. Asci
(206–)228 – 268(–296) × (21–) 21.8 –23.7(–25) µm (n = 14),
cylindrical to oblong, bitunicate, fissitunicate, with an ocular
chamber, a short to long stipe and simple or knob-like base,
containing 4–8 ascospores in (obliquely) uniseriate arrange-
ment; unstable in water. Ascospores (27–)31.5–42 (–51.5) ×
(13–)15–19 (–22) µm, l/w (1.9–) 2 –2.4(– 3.1) (n = 50), very
variable, typically ellipsoid to fusoid, sometimes subglobose,
symmetric or inequilateral or slightly curved, initially 2-celled,
hyaline and surrounded by a sheath, developing additional
septa, turning yellow to golden brown (when fresh), finally dark
brown with concolorous ends, with 8–14 distinct transverse
and 3–5 longitudinal septa, usually strongly constricted at the
primary septum; surface smooth; in 3 % KOH turning blackish
brown when mature.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 13 mm after 1 wk,
27 mm after 4 wk; colony circular, thick by a dense whitish
to greyish mat of aerial hyphae, first white, turning dull yel-
low and later becoming concentrically zonate with medium
or greyish brown and dark brown zones and a white margin;
producing a yellow diffusing pigment; odour indistinct. After 5 d
pycnidia densely disposed, appearing on and around the plug
or in rings, green, turning black, 120–300 µm diam, becoming
fused to larger complexes in aged cultures, emitting conidia in
white to olivaceous turbid drops. Phialides (4–)4.5 –7(–7.8) ×
(2–)2.3 – 3.3(– 4) µm (n = 19), crowded, sessile or on interca-
lary cells, mostly lageniform, also subconical, ampulliform to
subglobose. Conidia (3.5 –)4–4.8 (–5.5) × (1.2 –)1.3 – 2(–2.5)
µm, l/w (2.2–) 2.4–3.2 (– 3.8) (n = 32), cylindrical, oblong or
allantoid, 1-celled, smooth, with 2 guttules.
Habitat — On Cytospora sp. (both morphs; sexual morph of
the Leucostoma type) on Pyrus communis.
Distribution — Europe (Austria), only known from the type
locality.
Notes — Synfenestella pyri is an inconspicuous fungus. It
differs from Parafenestella spp. occurring on Rosaceae by the
swelling sheath visible in immature ascopores within asci.
Synfenestella sorbi (P. Karst.) Jaklitsch & Voglmayr, comb.
nov. — MycoBank MB829761; Fig. 22
Basionym. Cucurbitaria sorbi P. Karst., Bidrag Kannedom Finlands Natur
Folk 23(2): 62. 1873.
Synonyms. Gibberidea sorbi (P. Karst.) Kuntze, Revis. Gen. Pl. (Leipzig)
3(2): 481. 1898.
Fenestella bavarica Kirschst., Ann. Mycol. 34(3): 193. 1936.
Lectotype of Cucurbitaria sorbi, here designated. FinlAnd, Aboe, on
branches of Sorbus aucuparia, soc. Cytospora (Leucostoma) sp., 8 Apr. 1861,
P.A. Karsten (H 3686, from the herbarium Karsten; MBT385692). Epitype of
Cucurbitaria sorbi, here designated: AustRiA, Niederösterreich, Reichenau,
Rax, near Seehütte, on twig of Sorbus aucuparia, on/ soc. Cytospora sp.
(Leucostoma type; holomorph), 21 Sept. 2014, W. Jaklitsch (WU 36992;
MBT385693; ex-epitype culture from ascospores CBS 144862 = FR; culture
from conidia = FRa).
Pseudostromatic pustules when present 0.9–4.7 mm diam,
erumpent from bark and projecting up to 1.5 mm beyond the
bark surface, subglobose to pulvinate, roundish to longish in
outline; surface brown (compacted subiculum) to dark grey or
black (ascomatal apices), sometimes containing also narrow
cylindrical ostiolar necks of the Diaporthe or Leucostoma host.
Ascomata (330–)425 –745(–990) µm (n = 47) diam, globose,
subglobose or pyriform, immersed in bark in rows or valsoid
configuration, often laterally in the ostiolar region of the fungal
36 Persoonia – Volume 44, 2020
host or inserted in the pustules on a single or two levels upright
or obliquely and convergent, singly or up to c. 20 individuals
loosely or tightly aggregated and surrounded by usually ample
dark brown subiculum, partly directly on the fungal host; subicu-
lum also present below epidermis. Ascomatal apices containing
inconspicuous ostiolar openings 120–300 (–600) µm (n = 46)
diam, black, whitish when injured, often flattened. Subiculum
consisting of hyaline, pale to dark brown, thick-walled, 2–7 µm
(widened up to 9.5 µm at points connecting to the peridium) wide
hyphae, darker brown, particularly thick-walled, often short and
subsetose (‘hyphal appendages’) near the ascomatal apices.
Peridium 20–65 µm thick, up to 130 µm at the ostiolar level,
pseudoparenchymatous, consisting of cells (4–)7–14(– 23) µm
(n = 63) diam, outside very dark brown, thick-walled, to the
inside gradually paler and thinner-walled and terminated by a
narrow layer of pale brownish compressed cells. Hamathecium
Fig. 22 Synfenestella sorbi. a– w. Sexual morph. a– b. Ascomata/ pseudostromata in face view; c. Diaporthe impulsa pseudostroma with a small S. sorbi
ascoma inserted laterally; d. subsetose subicular hyphae at ascoma apex; e. ascomata above Diaporthe ascomata in vertical section; f. peridium and subicu-
lum in vertical section; g– i. ascus apices (g. immature; note sheath around ascospores); j–n. asci; o– w. ascospores (o. immature; p. young); x. pycnidia on
the natural host; y– c1. asexual morph from CMD at 22 °C; y. pycnidia and conidial drops; z. phialides; a1–c1. conidia (h, j, n–o, t –w. in 3 % KOH). a, k, u– v.
Cucurbitaria sorbi lectotype H 3686; b, d–f, l, q–r, z–b1. WU 36994/ CBS 144858 (C196); c, i, p, w. WU 36995 (C298); g, m, o, s– t, x– y, c1. WU 36992/CBS
144862 (FR); h, j. Fenestella bavarica holotype B 700016482; n. WU 36993. — Scale bars: a –c = 1 mm; d, f, j –n = 25 µm; e, x –y = 300 µm; g, q–w = 10
µm; h– i, o– p, z = 7 µm; a1–c1 = 3 µm.
37
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
consisting of numerous tightly packed, richly branched, 1–3 µm
wide ?paraphyses. Asci (199–)228 – 297(–377) × (15 –)18–
21(–23) µm (n = 64), cylindrical to oblong, bitunicate, fissi-
tunicate, with an ocular chamber, a usually long undulating
stipe and simple or knob-like base, containing 4–8 ascospores
in (obliquely) uniseriate arrangement. Ascospores (22–)28 –
35(– 43.5) × (10 –)13 –15.5(–18.5) µm, l /w (1.6 –) 2–2.5 (– 3.2)
(n = 171), very variable in shape, ellipsoid, oblong to fusoid, first
hyaline, 2-celled and with a swelling sheath, developing 3–5
main thick septa and later additional thin septa, eventually with
7–13, rarely 15, distinct transverse and (1–)2 – 4 longitudinal
septa, turning golden yellow, yellow brown, pale or reddish
brown and finally dark brown; often strongly constricted at the
primary septum, upper part often wider, ends usually broadly
rounded or upper end narrowly and lower broadly rounded,
concolorous, only paler or apiculate upon germination; smooth,
containing small droplets, often smaller in the ascus apex, often
several in the ascus aborted or aberrant; in 3 % KOH turning
dark olivaceous to blackish brown. Sexual morph sometimes
accompanied by black, shiny, globose, non-papillate, apically
collapsing-cupulate pycnidia 90– 300 µm diam seated on scant
to ample dark brown subiculum on innermost bark layers around
old Leucostoma pseudostroma (black encasement) and on
?Diaporthe ascomata.
Culture characteristics and asexual morph in culture — Co-
lony radius on CMD at 22 °C in the dark 8 mm after 1 wk,
21 mm after 3 wk; colony circular, thick, dense, first white to
bright yellow, aerial hyphae forming a thick white mat, reverse
yellow, yellow pigment diffusing into the agar, later colony
turning greyish brown from the centre, diffusing pigment turn-
ing dull yellow; odour indistinct. After 4 d pycnidia numerous,
mostly covered by a white mat of aerial hyphae, 120–330 µm
diam, more or less globose, hyaline to greenish olivaceous,
turning black, densely aggregating and forming stromatic
masses; conidia appearing in whitish turbid drops from many
ostioles; peridium pseudoparenchymatous, yellow to dark
olivaceous, surrounded by olivaceous to brown hyphae. Phia-
lides (4.8–)5 –8(–11.5) × 2– 4(–5.7) µm (n = 15), very variable,
subglobose to ampulliform to lageniform, on short simple
conidiophores, also with lateral pegs or on hyaline intercalary
cells. Conidia (3.3–) 4 –4.5(– 5) × (1.2 –)1.5 – 2(– 2.8) µm, l/w
(1.5–)2.1– 3.1(– 3.9) (n = 64), cylindrical, oblong to allantoid,
less commonly ellipsoid, 1-celled, hyaline, with 2 drops, smooth.
Habitat — On both morphs of Cytospora sp. of the Leuco-
stoma type, in pseudostromata on effete ascomata of Diaporthe
impulsa or in loose association with it on Sorbus aucuparia.
Distribution — Europe, North America fide Barr (1990; under
Fenestella subcaespitosa) and Mirza (1968; under Cucurbitaria
sorbi).
Other materials examined. AustRiA, Osttirol, Prägraten am Großvenediger,
Hinterbichl, Umbalfälle, on cf. Diaporthe impulsa on a dead attached branch
of Sorbus aucuparia, soc. cf. Karstenula sp., Mollisia caespiticea, 17 June
2015, H. Voglmayr & W. Jaklitsch (WU 36994; culture CBS 144858 = C196);
Steiermark, Deutschlandsberg, Freiländer Alm, W Freiländer Almhütte;
E15°02'57" N46°54'47", elev. 1410 m, on Diaporthe impulsa on a branch
of Sorbus aucuparia on the ground, 3 May 2017, G. Friebes (WU 36995;
culture C298); highway parking place close to the Pack tunnel, on a branch
of Sorbus aucuparia attached to the tree, soc. Diaporthe impulsa, 17 Feb.
1995, W. Jaklitsch W.J. 499 (WU 36993). – GeRmAny, Bavaria, Bayerisch
Häusl bei Eisenstein, on Cytospora (Leucostoma) on branches of Sorbus
aucuparia, June 1935, W. Kirschstein (B 700016482, holotype of Fenestella
bavarica).
Notes — In H two syntypes of Cucurbitaria sorbi are extant;
No. 3687 (Finland, Vaasa, on Sorbus, 12 Aug. 1867, P.A.
Karsten) contains mostly Dothiora pyrenophora, a fungus with
3-septate ascospores (possibly Nigrograna sp.), an immature
nectriaceous fungus and only little and old S. sorbi (ascospore
measurements given on the label: 25–30 × 12–14). In contrast,
No. 3686 contains good material and is thus selected as lecto-
type. The latter was also examined by M.E. Barr according to
the annotation slip. Synfenestella sorbi has been found in direct
association with both Cytospora sp. (Leucostoma morph) and
Diaporthe impulsa. Also other ascomycetes are associated
with the fungus, thus parasitism on other fungi may be possi-
ble. Fenestella bavarica is clearly a synonym of S. sorbi. In its
holotype, S. sorbi grew on Cytospora (Leucostoma morph), has
typical long-pedicellate asci with often aberrantly developed
ascospores, many aborted or distorted or even globose. At the
pustule surface cylindrical Leucostoma ostioles peek through
brown compacted subiculum of the Synfenestella; also a Tym -
panis is present on thin twigs.
Synfenestella sorbi is extremely variable in appearance, rang-
ing from solitary ascomata (on Diaporthe) to large conspicuous
pseudostromata. Hyphal appendages sensu Barr (1990; sub
Fenestella subcaespitosa) are subicular hyphae and occur
variably in the entire family. In S. sorbi they are often but not
always differentiated from other hyphae around the ascomatal
apices by having a nearly setose appearance, darker colour
and slightly thicker walls. The long-pedicellate asci were already
described by Mirza (1968; under Cucurbitaria sorbi). Cucurbi-
taria subcaespitosa from S. aria is clearly a different species
and here combined in Neocucurbitaria.
There are some sequence differences between isolates derived
from specimens containing the fungal host Cytospora (Leuco-
stoma) and those growing on Diaporthe impulsa. However, as
these differences are not convincing and the specimens are
morphologically indistinguishable, we do not recognise two
separate species. In the lectotype H 3686 S. sorbi is associated
with Cytospora, therefore we epitypify it with material containing
the same fungal host.
KEY TO FENESTELLOID SPECIES
Fungal hosts are only given in the key when different from or
additional to Cytospora spp. Parafenestella ostryae from Ostrya
carpinifolia (Wanasinghe et al. 2017b) is not included, as we
have not seen material of this species. Some species, particu-
larly Fenestella media vs F. subsymmetrica, Parafenestella
salicis vs P. pseudosalicis, or P. austriaca vs P. rosacearum
cannot be safely distinguished morphologically.
1. Ascospores with a gelatinous sheath ............... 2
1. Ascospores lacking a sheath ......................5
2. Sheath persistent during ascospore development...... 3
2. Sheath only present initially in the ascus ............. 4
3. Ascospores (52 –)58 – 67(–72) × (16–)19–23 (–26) μm, l/w
2.5–3.7, fusoid to subclavate, with c. 17– 25 transverse and
3–7 longitudinal septa, surrounded by a sheath around each
hemisphere, quickly swelling and losing contours in water;
on Ulmus glabra .................... Seltsamia ulmi
3. Ascospores (36 –)43– 54(–60) × (13 –)17– 22(–28) µm, l /w
2–3.4, clavate to fusoid, with 12 –17 transverse and 4 – 5
longitudinal septa; on Acer granatense ...............
............................Fenestella granatensis
4. Ascomatal groups inconspicuous, ascospores (27–)32 –42
(–51) × (13–)15 –19(–22) µm, l /w 2 – 3, typically ellipsoid to
fusoid, ends concolorous, with 8–14 distinct transverse and
3–5 longitudinal septa; on Pyrus .....Synfenestella pyri
4. Ascomata often forming conspicuous pseudostromatic pus-
tules, ascospores (22–)28 –35 (–44) × (10–)13 –16(–19) µm,
l/w 1.5– 3.2, ellipsoid, oblong to fusoid, with 7–13(–15) trans-
verse and (1–)2 – 4 longitudinal septa, ends concolorous,
only paler or apiculate upon germination; on Sorbus aucu-
paria ...........................Synfenestella sorbi
38 Persoonia – Volume 44, 2020
5. Ascomata immersed evenly or randomly in bark, as-
cospores (32–)38 – 52(–63) × (14–)16– 21(– 25) μm, l/w
1.8–3.3, with 12 –18(– 20) transverse and 3–7 longitudinal
septa, with wall distinctly thicker than septa; ends some-
times apiculate; on Ulmus spp. .... Protofenestella ulmi
5. Ascomata forming inconspicuous groups or well-defined
pseudostromatic pustules ....................... 6
6. Ascospores (36 –)49– 65(–73) × (14 –)18– 25 (–31) µm, most-
ly fusoid, with 13–20 transverse and 4 –6(–7) longitudinal
septa, often with hyaline, 2–7 μm long terminal apiculi; on
Alnus glutinosa and possibly Quercus ..............
............................ Fenestella fenestrata
6. Ascospores smaller ............................ 7
7. Ascospores (32 –) 41–53(– 61) × (13–)15 –19(–23) µm, l/w
2–3.6, with 11–16(– 20) transverse and 2 –4 longitudinal
septa; on Quercus and Salix . . Fenestella parafenestrata
7. Ascospores smaller ............................ 8
8. Ascomata forming more or less compact pustules, asco-
spores (28–)32 – 46(– 55) µm long ................ 9
8. Ascomata forming inconspicuous groups, ascospores
short er . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
9. Ascospores (30 –)32–41(– 47) × (14–)15 –19(–24) μm, l/w
1.8–2.6, with (9 –)10 –14(–16) transverse and 3– 5 longi-
tudinal septa and pale non-apiculate ends; on Acer opalus
..................................Cucitella opali
9. Ascopores with apiculate ends ..................10
10. Ascospores (31–) 36–46 (– 55) × (11–)15– 20(– 23) µm,
l/w 2– 3, with 11–14 (–16) transverse and 2–4 longitudinal
septa and usually broad, often hyaline terminal cells some-
times transformed into an ampulliform cellular appendage;
on Crataegus monogyna ..........Fenestella crataegi
10. Apiculi not ampulliform, up to 2 µm long, to 5 µm when
old ........................................ 11
11. Ascospores with distinctly submedian primary septum,
clavate to ellipsoid, (34–)36 –45(–49) × (13 –)15–19(– 22)
µm, l/ w 2–2.8, with 11–16 distinct transverse and 3–5
longitudinal septa; on Acer saccharum ..............
...........................Fenestella gardiennetii
11. Ascospores ellipsoid or fusoid, with submedian to median
primary septum .............................. 12
12. On Viburnum spp.; ascospores (29–) 38 –46(– 49) × (12 –)
15–18(– 22) µm, l/w 2 –3, with 11–16 transverse and 3–6
longitudinal septa, sometimes distinctly pointed at ends
...............................Fenestella viburni
12. On diverse woody hosts .......................13
13. Ascospores (30–) 34– 44(–53) × (12–)14–18 (–21) µm, l/w
2–3.3, mostly asymmetric, with indistinct 11–18 transverse
and 3–6 (–7) longitudinal septa ...... Fenestella media
13. Ascospores (28–) 34–44 (–55) × (13–)15–20 (–25) µm, l/w
1.8–3, asymmetric to subsymmetric, with distinct 11–16
(–18) transverse and 3–6 longitudinal septa .........
........................ Fenestella subsymmetrica
14. In (sub)alpine regions on Cotoneaster and Salix; asco-
spores (19–)24 – 30.5(–35) × (10.5 –)12 –14(–15.5) µm,
l/ w 1.4–2.9, with (7–)8 –12(–15) transverse and (2 –)3–4
longitudinal septa and concolorous ends . . . . . . . . . . . .
............................ Parafenestella alpina
14. Not exclusively in (sub)alpine regions ............. 15
15. On Melanconis alni on Alnus; ascospores (19–)26 –34(– 39)
× (10 –)13–17(–18) µm, l/w 1.7–2.8, with 8 –14 (–17) dis-
tinct transverse and 2–4 longitudinal septa and concolor-
ous, rarely paler ends ...... Parafenestella tetratrupha
15. On other hosts ..............................16
16. On species of Diaporthe .......................17
16. Not known to occur on Diaporthe ................18
17. Ascospores (22 –)24 –30(–32) × (11–)12–14(–16) μm, l/w
1.6–2.5, with (7–) 8–12(–13) transverse and 3 – 4 longitu-
dinal septa and often subacute concolorous, sometimes
lighter ends; on Acer pseudoplatanus ...............
......................Parafenestella pseudoplatani
17. Ascospores (26 –)29 –40(–47) × (11–)13–17(–19) µm, l /w
1.9–2.7, with (8 –)9 –13(–15) transverse and 3–6 longitudi-
nal septa and often broadly rounded, paler to hyaline end
cells; on Corylus .......... Parafenestella germanica
18. On Salix spp. ................................19
18. On Rosaceae................................ 23
19. Ascospores (20 –) 24–31(–38) × (8.5 –) 9.5–11(–13) µm,
l/ w 2–3.4, with 7–9 (–11) thick transverse and 1–2(–3)
longitudinal septa and concolorous or slightly paler ends
..................... Parafenestella vindobonensis
19. Ascospores wider ............................20
20. Ascospores (33–) 36–44 (– 50) × (14–)16 –19(–22) µm, l/w
2–2.6, with 11–16 distinct transverse and 3–5 longitudinal
septa and mostly concolorous ends ................
....................... Parafenestella parasalicum
20. Ascospores smaller ........................... 21
21. Ascospores (22 –)27– 33(–40) × (11–)13 –16(–18) µm, l/ w
1.7–2.5, with 9–13 (–14) transverse and 3–4 longitudinal
septa and concolorous, rarely slightly paler ends ......
...........................Parafenestella salicum
21. Ascospores with fewer septa ...................22
22. Ascospores (20–)23–29 (–34) × (10 –)11–14(–15) µm, l/w
1.5–2.7, often inequilateral, with often indistinct (5 – 6 –)
7–10(–11) transverse and (1–)2 –3 longitudinal septa, the
latter often forming parallel lines; ends concolorous, rarely
hyaline ...................... Parafenestella salicis
22. Ascospores (23 –)25 –29(–32) × (11–)12–14(–15) µm, l/w
1.9–2.7, symmetric, with distinct 7–10 (–11) transverse and
2–4 longitudinal septa; ends concolorous ...........
...................... Parafenestella pseudosalicis
23. Ascospores (23–)28 –36 (–42) × (11–)12–16(–18) µm, l/w
1.9–2.8, with 7–12 (–14) transverse and 1–3(– 5) longitu-
dinal septa; end cells concolorous except for a truncate to
slightly convex hyaline terminal part of their walls .....
............................ Parafenestella faberi
23. Ascospore end cells rounded and concolorous or diffusely
to entirely paler to hyaline ...................... 24
24. Ascospores (25–) 27–33(– 38) × (12–)13 –15(–16) µm, l/w
1.9–2.6, ellipsoid, symmetric, with 9 –13 (–14) distantly
spaced transverse and 3–4 (– 5) longitudinal septa . . . .
..........................Parafenestella austriaca
24. Ascospores (23 –)28 –35(–45) × (11–)13–17(–19) µm, l /w
1.7–2.6, symmetric to inequilateral, with (7–)9 –13(–15)
transverse and 2–4 (–5) longitudinal septa ...........
........................Parafenestella rosacearum
DISCUSSION
Other species of Fenestella (and Cucurbitaria)
See notes under the species described in the Taxonomy section
above and in Jaklitsch et al. (2018), where many names in
Cucurbitaria are commented on. In this work we studied parti-
cularly European species of Fenestella s.lat. with available
fresh material. We did not study, e.g., F. betulae nor F. parvula,
because no fresh material from Betula was available. No type
material of Fenestella cydoniae (basionym Pleospora cydoniae)
could be located. The protologue may suggest affiliation with
F. media or a similar species. Fenestella microspora was de-
scribed from Corylus in France with ascospores 10–11 × 6 –7
µm, thus it is apparently not a member of Cucurbitariaceae.
39
W.M. Jaklitsch & H. Voglmayr: Fenestelloid clades of the Cucurbitariaceae
Fenestella phaeospora is not a member of the Cucurbitaria-
ceae phylogenetically (unpublished results). Fenestella pru-
nastri from Prunus spinosa in Luxemburg was described with
asco spores having 3–4 transverse and 1 longitudinal septa,
20–22 × 8 –9 µm, pale yellow, as similar to F. lycii, i.e., not a
species of Fenestella, but possibly rather belonging to Camaro-
sporidiellaceae or Coniothyriaceae. Species with muriform asco-
spores on Lycium spp. (Cucurbitaria varians, Fenestella lycii)
are referable to Camarosporium. Fenestella subvestita from
Alnus glutinosa in Lyngby, Denmark, was described with asco-
spores 15–19 × 7.5–8.5 µm having 3 transverse and 1 longi-
tudinal septa, i.e., not a species of Cucurbitariaceae. The genera
Teichospora and Thyridium may house additional cucurbi-
tariaceous species, but the task to examine all available type
materials of these genera is beyond this work.
Molecular phylogeny
As shown in Fig.1, there is significant infraspecific variation in
taxa of the genera Parafenestella and Synfenestella. On the
other hand, in those Fenestella species, where several isolates
were available, F. media, F. subsymmetrica and F. viburni, there
is no or very little variation in DNA sequences of all markers.
This is particularly interesting, as most species of all genera
grow on Cytospora spp., i.e., they share the same ecology.
The reason for the infraspecific variability is unclear but may
indicate that these lineages are in the process of ongoing
speciation, probably triggered by host specificity on different
Cytospora hosts. This, however, is difficult to assess, as the
identification of the effete Cytospora hosts is usually impossible
when ascomata are produced.
Ecology, hosts and development
Fenestelloid fungi produce their ascomata and conidiomata on
or in tight association with effete ascomata and conidiomata of
mostly if not always Diaporthales, in temperate zones mostly
in the cold season (late autumn to early spring). This means
that climatic conditions have first to be favourable enough to
promote development of the fungal host and secondly, the fol-
lowing period must be long and favourable enough to support
ascomatal development and maturation. In the cold season
such periods are often short and irregular, as temperatures
may variably decrease below 0 °C often terminating fungal de-
velopment, and this may influence development of ascospores
dramatically. Moisture is usually not a criterion limiting fungal de-
velopment in the cold season, but in recent years and decades,
even this has changed in many temperate regions, where little
precipitation occurs even in winter. At high elevations (montane
to alpine levels) impact on development by climatic variation is
even more drastic. One example is Synfenestella sorbi, whose
plant host Sorbus aucuparia occurs from lowlands to subalpine
levels. As a result, ascospores often vary tremendously with
regard to size and shape, even within a single ascus, in this
but also other species. Often only one to four ascospores in an
ascus are mature and others are aborted. The resulting drastic
infraspecific variation of ascospore characters makes species
recognition and identification using morphological traits alone
extremely difficult.
Also, size and development of ascomatal groups or pseudo-
stromata vary considerably, as they depend on the size and
development of the Cytospora (and other) hosts, thickness of
twigs or branches, and again on (micro-)climatic conditions on
twigs and branches determining the magnitude of infection of
the present host pseudostromata or conidiomata. As the latter
are mostly effete when colonised by the Fenestella, we have
no information on host identity on the species level, which is
difficult also due to the unsettled taxonomy of Cytospora and
the fact that several Cytospora species may occur on a single
host plant. Specificity toward the fungal host is unknown, as,
e.g., Fenestella media occurs on several plant hosts, which
may be substrates of different Cytospora spp. Thus, F. media
either has a wide specificity regarding its Cytospora host spe-
cies or alternatively, it grows on a single Cytospora sp., which
occurs on many different plants. As species of Cytospora are
numerous and common, we expect that many more species
of fenestelloid fungi will be described in future, at least from
temperate zones worldwide.
Morphological characters
Ascomata of fenestelloid fungi are always immersed in to erum-
pent from bark and have a marked tendency to form groups,
besides forming solitary ascomata in a colony, too. Numbers
of ascomata in a group is often less than 10 in Parafenestella,
but may be much higher in Fenestella and Synfenestella. A few
species of the latter genera may produce conspicuous pseudo-
stromatic pustules. Morphological variation in Synfenestella is
remarkable, as fructifications of S. pyri are very inconspicuous
and consist of only few ascomata in a group, and asci are mostly
short-stipitate, while in contrast S. sorbi often forms large con-
spicuous pseudostromata and asci are long-stipitate. However,
the swelling sheath in young ascospores within asci unites the
two species. It should be borne in mind that elongated ascus
stipes and partly biseriate arrangement of ascospores in micro-
scopic mounts may also be consequences of exerting pressure
on the cover slip in order to free asci from the hymenium. The
peridium of Fenestella was characterised as 3-layered by Barr
(1990; sub Fenestellaceae). However, the innermost narrow
layer of brownish compressed and elongated cells is not always
present and the outer two layers (dark brown, of thick-walled
cells outside, followed by thinner-walled and paler brown to
hyaline cells) are, after consideration of all species, rather a
single layer with internal variation. The outermost region can
be narrow and then more easily be interpreted as a layer of
its own, particularly when the transition to lighter colour and
thin walls, which basically is always gradual, takes place at a
short distance. This is especially obvious in F. fenestrata and
F. parafenestrata and in the three sibling species F. media,
F. subsymmetrica and F. viburni. The hamathecium has been
identified as apically free paraphyses present among immature
asci in several species, thus this may be a character common
to the entire family Cucurbitariaceae (see also Jaklitsch et al.
2018). Morphology and illustration of several species in this
work was done using dried material; thus images of fresh vital
ascospores are missing. However, ascospores of fenestelloid
fungi are mostly yellow-brown or golden-brown when fresh, but
turn dark brown upon drying and may often be dark reddish
brown in herbarium material. They turn greenish olivaceous
when immature to blackish brown when mature in 3 % KOH.
Verruculose ornamentation has been seen in many species.
This might however be an effect of drying of the perispore.
Asexual morphs in the Cucurbitariaceae
As pointed out by Jaklitsch et al. (2018), asexual morphs of
the Cucurbitariaceae do not offer sufficient traits for reliable
distinction at the species or generic level. As we have seen
here, asexual morphs of all fenestelloid fungi are morphologi-
cally similar and may be termed phoma-like.
Acknowledgments We thank the fungarium curators A. Bond (K), B. Buyck
(PC), P. Clerc (G), M. Hyvarinen and O. Miettinen (H), A. Igersheim (W), R.
Lücking (B), O. Morozova (LE), C. Scheuer (GZU), W. Till (WU) and D. Triebel
(M) for sending and managing collections, R. Berndt (Z) and A. Igersheim
(W) also for information on specimens, A. Akulov, J. Fournier, G. Friebes,
A. Gardiennet, I. Greilhuber, R. Moosbeckhofer, K. Pätzold, E. Popov, S. Tello
and B. Wergen for fresh material or data. The financial support by the Aus-
trian Science Fund (FWF; project P25870-B16) is gratefully acknowledged.
40 Persoonia – Volume 44, 2020
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