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First record of Neolecta vitellina (Bres.) Korf & J. K. Rogers and Sowerbyella fagicola J. Moravec in Serbia

  • Faculty of Sciences and Mathematics University of Niš
  • National Park Fruska Gora

Abstract and Figures

The rare ascomycetes Neolecta vitellina and Sowerbyella fagi-cola have been recently registered in Serbia as new for the country. Morphological description based on found specimens, along with macro-and microscopic characteristics of fruiting bodies and attributes on their habitat, ecology and distribution in Serbia are given. Apstract: Prvi nalaz vrsta Neolecta vitelina (Bres.) Korf. & J. K. Rogers and Sowerbyella fagicola J. Moravec in Serbia Retke askomicete Neolecta vitellina i Sowerbyella fagicola registrovane su po prvi put u Srbiji. U radu je predstavljen morfološki opis zasnovan na makro-i mirokarakteristikama plodonosnih tela zabeleženih primeraka uz opis staništa i podatke o ekologiji i distribuciji navedenih askomiceta u Srbiji.
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10 (1) September 2019: 1-8
DOI: 10.5281/zenodo.3463980
First record of Neolecta
vitellina (Bres.) Korf & J. K.
Rogers and Sowerbyella
fagicola J. Moravec in Serbia
Original Article
Iskra Kajevska
Department of Biology and Ecology, Faculty of
Sciences and Mathematics, University of Niš, Serbia (corresponding author)
Dragiša Savić
National Park Fruška Gora, Zmajev trg 1, 21208
Sremska Kamenica, R. Serbia
Received: August 28, 2018
Revised: August 23, 2019
Accepted: September 10, 2019
The rare ascomycetes Neolecta vitellina and Sowerbyella fagi-
cola have been recently registered in Serbia as new for the coun-
try. Morphological description based on found specimens, along with
macro- and microscopic characteristics of fruiting bodies and at-
tributes on their habitat, ecology and distribution in Serbia are given.
Key words:
Neolecta vitellina, Sowerbyella fagicola, rst record, Serbia
Prvi nalaz vrsta Neolecta vitelina (Bres.) Korf. & J. K. Rogers and
Sowerbyella fagicola J. Moravec in Serbia
Retke askomicete Neolecta vitellina i Sowerbyella fagicola registrovane
su po prvi put u Srbiji. U radu je predstavljen morfološki opis zasnovan na
makro- i mirokarakteristikama plodonosnih tela zabeleženih primeraka uz
opis staništa i podatke o ekologiji i distribuciji navedenih askomiceta u Srbiji.
Ključne reči:
Neolecta vitellina, Sowerbyella fagicola, prvi nalaz, Srbija
During fungal diversity eld studies conducted
from 2016-2018, in selected regions in southern and
central Serbia, the rare ascomycetes Neolecta vitel-
lina and Sowerbyella fagicola have been registered.
These specimens represent the rst documented
records of the species in Serbia. Both taxa are al-
ready taken into consideration as “relevant for as-
sessment” in the European Red List candidates of
endangered fungi which is still under preparation,
and only few countries (Austria, Germany, Norway
and Sweden) have included these species in their
National Red List of fungi (ECCF, c1998-2019).
Fungal conservation is a complex and challeng-
ing issue due to high species diversity, incomplete
knowledge of degree of rarity or risk status, large
number of little known species, poor understanding
of their trophic or habitat requirements (Molina et
al., 2011). Reports of Neolecta vitellina and Sow-
erbyella fagicola from Europe are scarce, therefore
biogeographical studies with data on local popula-
tions and habitat are necessary to better understand
the nature of its distribution and further conservation
approaches. This study aimed to present identica-
tion of these two rare species based on morphologi-
cal description and examination of the present col-
lections, as well as data on their habitats, ecological
notes and distribution in Serbia.
Materials and methods
Specimens have been collected during the period
2016-2018 in two selected regions in southern (Suva
Planina Mountain - Suva Planina Mt.) and central
(Kopaonik National Park - Kopaonik NP) Serbia
(Fig. 1). A fresh material with substratum was ob-
served for macro- and micromorphological charac-
terisation and exsiccates were deposited in the pri-
vate fungal collection of the Mycological Society
of Niš (M.S.N.). Part specimens were also stored
in the ocially unregistered fungarium of Fruška
Gora National Park. Microscopic characteristics and
measurements were made on multiple preparations
primarily of fresh material, to obtain vital taxonomic
characterisation (Baral, 1992), and for comparison
with dry material, using Optika N-400M and Leica
DM 1000 light microscopes. Examination and meas-
© 2019 Kajevska, Savić. This is an open-access article distributed under the terms of the Creative Commons
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under the same license as the original. 1
uring of approximately 30 spores released from the
ascus and other elements of fresh and dry material
were made rst in water, later in Lugol’s solution
(IKI) and the ornamentation of the cells was ob-
served in Congo red and Cotton blue. Macro- and
microscopic descriptions are based only on the col-
lected specimens. The species names follow Index
Fungorum (Kirk, 2018) and MycoBank (Stalpers &
Cock, 2018) and the following works were used as
resources for identication of the species: Moravec
(1973, 1985, 1988), Redhead (1977, 1979), Roer
(1999), Hansen & Knudsen (2000), Klofac & Her-
rman (2003), Landvik et al. (2003), Benkert (2005),
Van Vooren (2007), Perić (2008), Galán et al. (2010),
Chinan & Hewitt (2012), Friebes (2015), Wieschol-
lek (2015), Ribes et al. (2016).
Neolecta vitellina (Bres.) Korf & J.K. Rogers,
Phytologia 21(4): 204 (1971). Figs. 2 – 4.
Basionym: Geoglossum vitellinum Bres., Revue
Mycologique 4: 212 (1882).
Fig 1. Map of investigated areas: 1) Kopaonik NP, Crvene
Bare nature reserve - Neolecta vitellina site; 2) Suva
Planina Mt., above Gadžin Han town - Sowerbyella fagi-
cola site.
Description of studied specimen:
Ascomata 12-38 x 3-7 mm in diameter, slen-
der, cylindrical, club-shaped or irregularly clavate,
spathuliform or somewhat truncate, elongate or lan-
ceolate, some attened, smooth or wrinkled, often
with longitudinal furrows, bright yellow; esh elas-
tic. Stipe 4-25 x 2-4 mm, central, cylindric, gradu-
ally merges with the fertile part, tomentose, white-
coloured. Asci 50-75(120) x 4-7 μm, cylindrical or
narrowly clavate, 8-spored, mostly uniseriate, apex
obtuse and not staining blue in iodine, aporynchous
ascus base. Paraphyses absent. Ascospores 4-6(-9)
x 2.5-4 μm, smooth, hyaline, thick-walled, ellipti-
cal, subglobose, amygdaliform or reniform, usually
contains small globose oil drops and a nucleus most
visible in Lugol’s solution and Congo red. Excipu-
lum of textura intricata, with septate hyaline hyphae
(5)10-20(30) μm.
Habitat: Fruiting bodies were found in a slope,
scattered among the humid fallen needles of Picea
abies (Fig. 2a, b).
Specimen examined:
Kopaonik NP (Fig. 1), Crvene Bare nature re-
serve, 1600 m, P. abies forest, on litter, among P. abies
fallen needles, attached on roots, very wet place,
BIOLOGICA NYSSANA ● 10 (1) September 2019: 1-8
Fig 2. In situ photographs of Neolecta vitellina; a) as-
comata growing from spruce rootlets, photo I. Kajevska,
b) fruiting bodies at dierent stages of maturity. Photo: M.
Kajevska, Savić ● First record of Neolecta vitellina (Bres.) Korf & J. K.
Rogers and Sowerbyella fagicola J. Moravec in Serbia
BIOLOGICA NYSSANA ● 10 (1) September 2019: 1-8 Kajevska, Savić ● First record of Neolecta vitellina (Bres.) Korf & J. K.
Rogers and Sowerbyella fagicola J. Moravec in Serbia
20/07/2016, I. Kajeveska (legator-leg. A. Trailović),
M.S.N. 21/07/16-168; Kopaonik NP, Crvene Bare,
cc. 1600 m, N 43°17’35.6”, E 20°48’21.81”, P. abies
forest, on litter, among P. abies fallen needles, at-
tached on roots, on a slope, 29/07/2017, I. Kajevska,
M.S.N. 29/07/17-240.
Sowerbyella fagicola J. Moravec, Česká Mykol.,
27 (2), p. 66 (1973). Figs. 5 – 7.
Description of studied specimen:
Apothecia 5-23 x 3-9 mm in diameter, shallow
cup-shaped, concave to fully extended in maturity;
hymenium smooth, bright yellow or orange, young
specimens somewhat translucent becoming dark or-
ange in age and orange-brown when dry; excipulum
concolorous with the hymenium, nely pubescent
with indistinguishable white hairs; margin entire or
slightly undulate, somewhat with darker color; esh
Fig. 3. Micrographs of Neolecta vitellina, a) ascus with
spores in Congo red; b) asci with spores in IKI; c) as-
cospores in IKI; d) ascus with spores in IKI; e) young as-
cus. Bars: a, b, e = 10 μm; c,d = 5 μm.
Fig. 4. Micrographs of Neolecta vitellina, f) hymenium; g)
asci bases; h) excipulum cells. Bars: f =20 μm; g, h = 10
thick, very soft, light and fragile, lubricous. Stipe
10-27 x 2-4 mm, central, smooth, delicate, concol-
ourous with the cap, often wrinkled-furrowed or
somewhat moniliform, cylindrical but usually with
attenuated base. Asci 200-220(270) x 9-13 μm, cy-
lindrical, 8- spored, uniseriate, apex obtuse or trun-
cate and inamyloid but asci walls somewhat blue
in Melzers reagent, ascus base pleurorynchous.
Paraphyses 200-240 x 2-3 μm, narrowly cylindri-
cal, slender, septate, with yellow to orange pigments
bluish in Melzer’s reagent. Ascospores 16-19.5(-22)
x (6.5-)7.5-8.5(-9) μm, hyaline, thick-walled, fusi-
form, usually contains two globose oil drops (2.5-4.5
μm) accompanied with 1-2 smaller drops (< 2 μm)
on the ends; ornamentation seemingly smooth but
nely verrucose on oil immersion. Medullary excip-
ulum of textura intricata, with hyphae 20-45(65) x
5-7(9) μm, hyaline, with thin walls. Ectal excipulum
of textura subglobulosa to angularis, with hyphae of
20-40(50) x (8)15-30(45) μm. Marginal hairs cylin-
drical, narrow, septate and hyaline to yellowish hairs
of 30-150 x 4-7 μm.
Habitat: Investigated specimens were found in
the base of a gully with a lot of organic matter accu-
mulated in a mixed deciduous forest (Carpinus ori-
entalis, Quercus cerris, Cornus mas, Acer campes-
Fig. 5. In situ photographs of Sowerbyella fagicola; a)
ascomata arising from litter composed of mixed decidu-
ous forest, b) fruiting bodies at dierent stages of maturity.
Photos: I. Kajevska.
BIOLOGICA NYSSANA ● 10 (1) September 2019: 1-8
Fig. 6. Micrographs of Sowerbyella fagicola, a) asci with
spores, paraphyses; b) ascus tip; c) ascus base; d) spores
with ornamentation in Congo red; e) spores in ascus, Con-
go red; f) paraphyses; g) spores in IKI. Bars: a, d, g = 10
Fig. 7. Micrographs of Sowerbyella fagicola, h) cells of
medullary excipulum; i) cells of ectal excipulum; j) cells of
ectal excipulum; k) hairs. Bars: h, i, j, k = 20 μm.
tre, Robinia pseudoacacia, with one Fagus sylvatica
tree for about 20 m distance from the collection site),
on a shaded place and humid for a long time. Fruit-
ing bodies scattered or gregarious, emerging from
the moist forest litter (Fig. 5a, b).
Specimen examined:
Suva planina Mt. (Fig. 1), above Gadžin Han
town (Niš), 347 m, N 43°12’25”, E 22°03’06”,
mixed deciduous forest (C. orientalis, Q. cerris, C.
mas, R. pseudoacacia, A. campestre, F. sylvatica),
on litter, attached on small fallen branches, leafs,
tiny pieces of wood, etc., very wet place, 29/05/2016
and 26/05/2018, D. Wieschollek (leg. I. Kajevska),
M.S.N. 29/05/16-162; M.S.N. 26/05/18-239.
Kajevska, Savić ● First record of Neolecta vitellina (Bres.) Korf & J. K.
Rogers and Sowerbyella fagicola J. Moravec in Serbia
The species Neolecta vitelina was rst described
in Italy as Geoglossum (Microglossum) vitellinum
Bres. (Bresadola, 1882). In North America, it was
reported as Mitrula vitellina (Bres.) Sacc. (Durand,
1908), and lately due to the lack of paraphyses as
an important distinguishing feature Mains (1955) re-
ports it under the name Spragueola vitellina (Bres.)
Nannf. and Imai (1934, 1941) from Japan as Ascoco-
rynium vitellinum (Bres.) S. Ito & Imai. Korf (1971)
transfers M. vitellina into the genus Neolecta Speg.
Redhead (1977) reports N. vitellina in Canada and
nely places the genus Neolecta within Neolectace-
ae Redhead due to its unique morphology. Based on
morphological and molecular analysis, Neolecta has
been placed in a separate order, Neolectales Land-
vik, O. E. Erikss., Gargas & Gustafsson (Landvik et
al., 1993). Subsequently, giving full morphological
and ultrastructural description as well as molecular
data of the genus species, Landvik et al. (1996, 2001,
2003), afterwards Sugiyama et al. (2006) and Healy
et al. (2013) conrms the unique position of Neolecta
(class Neolectomycetes O.E. Erikss. & Winka, sub-
phylum Taphrinomycotina O. E. Erikss. & Winka)
as the earliest lineage within Ascomycota. Neolecta
vitelina so far is the only species of the genus that
has been reported from Europe. Available published
data on this species from Europe come from the fol-
lowing countries: Italy (type specimen-Bresadola,
1882; Carbone et al., 2015), Poland (Bujakiewicz,
1979), Switzerland (Ryman & Holmåsen, 1992),
Germany (Krieglsteiner, 1993), Czech Republic
(Svrček, 1978), Fennoscandia-Norway, Sweden and
Finland (Eckblad, 1963; Ohenoja, 1975; Hansen &
Knudsen 2000), Romania (Chinan & Hewitt, 2012)
and recently has been described and illustrated from
Austria (Friebes, 2015). Mycological eld research
held in summer 2016, at Kopaonik NP, revealed the
rst nding of N. vitellina in Serbia. Specimens have
been preserved. Another location of this species with
numerous fruiting bodies was located the next year
in the same area. Although most reports of N. vitel-
lina are known from northern Europe, presumably,
the precise geographical distribution across Europe
cannot be denite yet. Most recent reports of this
rare ascomycete like those from Romania, Austria
and now from Serbia shift its distribution range to-
wards central and southern Europe. Reports of N.
vitellina are scarce, therefore more focused research,
monitoring and data with insights on local popula-
tions of this species are needed to better understand
the nature of its distribution. Our collection of N.
vitellina originates from a forest slope with fruiting
bodies emerging from the fallen needles in P. abies
(L.) H. Karst. forest at an altitude of cc.1600 m. Nu-
BIOLOGICA NYSSANA ● 10 (1) September 2019: 1-8 Kajevska, Savić ● First record of Neolecta vitellina (Bres.) Korf & J. K.
Rogers and Sowerbyella fagicola J. Moravec in Serbia
merous fruiting bodies have been found at dierent
stages of maturity. Although this rare species has
been found exclusively in spruce needle bogs, many
questions regarding its morphology, phenology and
its trophic anity remain unclear. Readhead (1979)
indicates N. vitellina as parasite on spruce rootlets.
Fruiting bodies of our collections were also attached
to spruce rootlets and while collecting was dicult
to separate them from the substrate. Macro- and mi-
croscopically, the specimens collected from Serbia
correspond well with the descriptions presented in
Redhead (1977, 1979), Hansen & Knudsen (2000),
Landvik et al. (2003), Chinan & Hewitt (2012) and
Friebes (2015). Until now we have only one known
collection site, but it may likely occur on other sites
especially within the Kopaonik NP where spruce
forests prevail. The site where the species is regis-
tered is already within a protected area, as it is situ-
ated in a nature reserve in the National Park, but it
is an area of high recreational use. Accordingly, we
anticipate that all recreational activities on this site
could be an important threat to the survival of this
fungal species.
Sowerbyella fagicola so far is reported only from
Europe in the following countries: Czech Republic
(type specimen-Moravec, 1973; 1988), Germany
(Krieglsteiner, 1979, 1993; Häner, 1989, 1993;
Hohmeyer et al., 1989, Benkert, 2005; Wieschollek,
2015), Austria (Rücker & Wittmann, 1995; Klofac
& Voglmayr, 2003), Switzerland (Roer, 1999),
Poland (Chmiel & Ronikier, 2007), France (Van
Vooren, 2007; Moyne & Petit, 2010); Montenegro
(Perić, 2008); Spain (Galán et al., 2010) and re-
cently from Spain (Ribes et al., 2016), Bosnia and
Herzegovina (Jukić & Omerović, 2017) and Bul-
garia (Assyov & Slavova, 2017). Recent reports of
S. fagicola from the abovementioned three Balkan
countries have predicted an expected presence of
this rare species also on Serbian territory. Thus, in
May 2016 and subsequently in 2018, S. fagicola has
been registered during mycological eld research
held in the southern part of Serbia, precisely on
Suva planina Mt., above Gadžin Han town. There-
fore, data on S. fagicola published in this paper is
new for the country. Sowerbyella fagicola was rst
described by Moravec (1973). The spore ornamen-
tation and macroscopically it reminds S. imperialis
(Peck) Korf, but this species has smaller spores, and
it is found in coniferous habitats (Moravec, 1973,
1985, 1988). Our collection contains fruiting bodies
at dierent stages of maturity and macro- and mi-
croscopically, the specimens collected from Serbia
correspond well with the descriptions presented in
Moravec (1973, 1985, 1988), Roer (1999), Klo-
fac & Herrman (2003), Benkert (2005), Van Vooren
(2007), Perić (2008), Galán et al. (2010), Wieschol-
lek (2015), Ribes et al. (2016) except for slightly
dierences in dimensions of the excipulum cells.
The dimensions of the textura intricata cells in Perić
(2008) are 23.6-51.7 x 8.6-15.7 μm and are slightly
wider than those from our collection (5-7(9)). Also
cells of the ectal excipulum from our collection (20-
40(50) x (8)15-30(45)) μm) are longer than those
(11-23 x 10-22) noted by Van Vooren (2007). Inves-
tigated specimens from our collection comprised of
small amount of fruiting bodies (cca. 15) have been
found in a shaded place, humid for a long time at the
base of a gully where lot of organic matter is accu-
mulated. The collection site was situated in a mixed
deciduous forest comprised of C. orientalis Mill., Q.
cerris L., C. mas L., R. pseudoacacia L., A. camp-
estre L. and with only one F. sylvatica L. tree for
about 20 m distance from the collection site. Speci-
mens were found scattered, some gregarious (with
max. 3-4 fruiting bodies), barely visible among the
moist layer of the forest litter. Stipe base has been
attached to fallen leaves, twigs, branches and other
small-sized decaying organic matter. We managed to
establish only leaf litter from C. orientalis, Q. cer-
ris, R. pseudoacacia and C. mas but without Fagus
leaf litter. It might also be worth to mention that the
only herbaceous plant species on the collection site
were Galium trifolium Michx. and Hedera helix L.
but poorly developed due to little sunlight unable to
reach the ground in the dense forest. Regarding the
ecology of S. fagicola, Moravec (1988) points out
the presence of ammonia in the substrate as a pos-
sible condition for developing its fruiting bodies. So
far this has not been conrmed. And although it is
frequently related with Fagus it has also been found
occurring in other habitats. In our case and in reports
from many countries Fagus has not been always the
predominant tree in the habitat but the most uncom-
mon are the cases from France found in an oak forest
(Moyne & Petit, 2010) and Spain found in Mediter-
ranean oak forest (Galán et al., 2010) and from Spain
found on hazel litter (Ribes et al., 2016). Thus, the
real ecology and substrate preferences of S. fagicola,
whether it is or not bound to Fagus, remains unclear,
but it is most likely a saprotrophic species growing
on ne litter in its initial phase of decomposition. We
also agree with the conviction noted by Wieschol-
lek (2015) that S. fagicola grows on a ne litter ac-
cumulated predominantly by deciduous trees. Also,
although Suva Planina Mt., is also a special nature
reserve and a protected area, during our continuous
eld studies we established various negative anthro-
pogenic activities in this region. In 2017, on this site,
we couldn’t found developed fruiting bodies of S.
fagicola. Namely, for the past few years, there were
many actions of illegal logging and an increase in
illegal landlls. These and similar activities soon
BIOLOGICA NYSSANA ● 10 (1) September 2019: 1-8
may be possible threat for the persistence of this and
other rare species encountered in this small but re-
garding mycodiversity rich area. Therefore, as soon
as possible, this region should be an issue of concern
and to be taken into consideration in all available
national biodiversity monitoring and protection ac-
tions. Regarding the status of S. fagicola, we consid-
er it should be included in the Preliminary national
red list of fungi and for the future Red Book of fungi
deserves a status of a critically endangered (CR)
species B2b(iii) according to IUCN categorisation,
as well as its only known habitat to be protected.
Similar status for S. fagicola has been already fore-
seen in Bosnia and Herzegovina (Jukić & Omerović,
The rare Neolecta vitellina and Sowerbyella fagicola
have been registered during the period 2016-2018 in
two selected regions in southern (Suva Planina Mt.)
and central (Kopaonik NP) Serbia. Although pro-
tected, both areas where these rare species persist
are inuenced by harmful anthropogenic activities
which may be a possible threat to these ascomycetes.
Data on species with such status will increase the
insights into better understanding of further fungal
conservation approaches. Therefore, morphological
description based on found specimens, along with
macro- and microscopic images and biogeographi-
cal data from Serbia have been fully presented in
this paper. Since it is already within a protected area,
as it is situated in a nature reserve in the National
Park, it is an area of high recreational use. Accord-
ingly, we anticipate that all recreational activities on
this site could be an important threat to the survival
of this fungal species.
The authors express sincere gratitude to Dirk Wi-
eschollek for his useful information and generous
help with the identication of S. fagicola. Many
thanks, we express to Aleksandar Trailović, Dani-
jela Sokolović and Marko Blagojević, members of
the Mycological Society ,,Vilino kolo, Petrovac na
Mlavi”, for providing us with fresh material and in
situ photographs of N. vitellina.
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Full-text available
Neolecta vitellina is a rarely reported ascomycete in Central Europe. A collection from the Untersulzbachtal in Salzburg is presumably the first record of this species in Austria. A description and photographs of this collection are provided based on living material. The distribution, ecology, and phylogeny of Neolecta vitellina are briefly discussed. Neolecta vitellina ist ein seltener Schlauchpilz, der in Mitteleuropa nur von we-nigen Funden bekannt ist. Eine Kollektion aus dem Untersulzbachtal in Salzburg dürfte der erste Nachweis dieser Art in Österreich sein. Der Fund wird anhand von lebendem Material beschrieben und illustriert. Die bekannte Verbreitung, Ökologie und Phylogenie dieser Art werden kurz diskutiert.
Full-text available
Étude macroscopique et microscopique, avec photographies (in situ et au laboratoire), dessins, commentaires écologiques et taxinomiques, de Sowerbyella fagicola Moravec (Discomycètes), espèce signalée pour la première fois au Monténégro.
Were the first ascomycetes yeast-like (unicellular) or filamentous with complex multicellular growth? Molecular studies have indicated that the earliest lineages of ascomycetes were mostly yeast-like and without complex, multicellular fruit bodies. The genus Neolecta stands out as an exception. Neolecta spp. have filamentous growth and discomycete-type of fruit bodies, but constitute one of the basal lineages both in rRNA and RPB2 gene trees. In this paper, we have used 10 previously unpublished and 30 GenBank β-tubulin genes to test the phylogenetic position of Neolecta spp. We compared the phylogenetic information from the amino acid sequences (485 characters) with the phylogenetic information content of 33 characters from intron gains and losses. Due to gene duplications, two paralogous versions of β-tubulin genes occurred in four species included in our analysis. Although phylogenetic interpretation of β-tubulin gene trees was complicated by a history of gene duplications, intron gains and losses, and by unequal rates of amino acid substitution, Neolecta species never formed a monophyletic group with any of the sequences from the filamentous ascomycetes in the Pezizomycotina (euascomycetes) in parsimony or distance analyses. Like the RPB2 and rRNA genes, the β-tubulin genes support the hypotheses of an early divergence of Neolecta from superficially similar filamentous ascomycetes. Neolecta could become a key taxon particularly in comparative studies between the fungal model organisms in the mainly unicellular (=yeast-like) taxa Saccharomyces and Schizosaccharomyces and the filamentous fruit-body forming taxa Neurospora and Aspergillus.