Lichenopeltella mizerniana sp. nov. from the upper Pliocene of Mizerna (southern Poland)

Abstract

Fungal sporocarps having ostiole with setae were found in the upper Pliocene deposits from Mizerna (borehole Mizerna-Nowa), southern Poland. These remains morphologically correspond to the fossil-genus Trichothyrites Rosend., although the structure of the ostiolar collar with non-septate setae seems unique and is typical for sporocarps (catathecia) of some modern species of the genus Lichenopeltella Höhn. Other contemporary fungal genera with setose sporocarps differ considerably from Lichenopeltella in respect of their morphology. Taking this into consideration, a new fossil-species Lichenopeltella mizerniana G. Worobiec is proposed. Morphologically, Lichenopeltella mizerniana is similar both to some modern lichenicolous [L. peltigericola (D. Hawksw.) R. Sant., L. rangiferinae Brackel, and L. uncialicola Brackel] and non-lichenicolous species [L. ammophilae (J.P. Ellis) P.M. Kirk & Minter, L. palustris (J.P. Ellis) P.M. Kirk & Minter] of this genus. The presence of Lichenopeltella mizerniana suggests that the Pliocene climate of the Mizerna locality was probably at least moderately humid.

Full text

ORIGINAL ARTICLE
Lichenopeltella mizerniana sp. nov. from the upper Pliocene
of Mizerna (southern Poland)
Grzegorz Worobiec
1
&Elżbieta Worobiec
1
Received: 24 January 2020 /Revised: 15 June 2020 / Accepted: 18 June 2020
#The Author(s) 2020
Abstract
Fungal sporocarps having ostiole with setae were found in the upper Pliocene deposits from Mizerna (borehole Mizerna-Nowa),
southern Poland. These remains morphologically correspond to the fossil-genus Trichothyrites Rosend., although the structure of
the ostiolar collar with non-septate setae seems unique and is typical for sporocarps (catathecia) of some modern species of the
genus Lichenopeltella Höhn. Other contemporary fungal genera with setose sporocarps differ considerably from Lichenopeltella
in respect of their morphology. Taking this into consideration, a new fossil-species Lichenopeltella mizerniana G. Worobiec is
proposed. Morphologically, Lichenopeltella mizerniana is similar both to some modern lichenicolous [L. peltigericola (D.
Hawksw.) R. Sant., L. rangiferinae Brackel, and L. uncialicola Brackel] and non-lichenicolous species [L. ammophilae (J.P.
Ellis) P.M. Kirk & Minter, L. palustris (J.P. Ellis) P.M. Kirk & Minter] of this genus. The presence of Lichenopeltella mizerniana
suggests that the Pliocene climate of the Mizerna locality was probably at least moderately humid.
Keywords Lichenopeltella .Trichothyriaceae .Fossil fungi .Palaeoecology .Upper Pliocene .Poland
Introduction
Fungi are an important group of fossils of eukaryotic organ-
isms found as fungal hyphae, sporocarps and spores that ap-
parently appeared in the late Proterozoic (Taylor et al. 2015).
Fossil fungi are useful both as a palaeoecological proxy
(Dilcher 1965; Lange 1976; Elsik 1978; Bera and Mandal
2014; Worobiec and Worobiec 2017;Worobiecetal.2018)
and for calibration of the phylogenetic trees of fungi obtained
using molecular clock methods (Geml et al. 2004;Lopandic
et al. 2005; Beimforde et al. 2014;Morenoetal.2015;Liu
et al. 2017; Halbwachs 2019). Chitinous fungal remains, even
almost unaltered, are frequently found in palynological sam-
ples (Elsik 1996; Worobiec et al. 2009). Microremains of
sporocarps are commonly found both as detached sporocarps
during palynological investigations and on cuticles of
fossilised leaves. These sporocarps, mainly of epiphyllous
fungi, are classified as members of the modern families
Microthyriaceae, Micropeltidaceae, Trichothyriaceae
(Microthyriales) and Asterinaceae (Asterinales) (Cookson
1947; Dilcher 1965; Elsik 1978; Worobiec and Worobiec
2013). However, as most of these sporocarps are preserved
without any spores, and rarely with mycelium attached, they
lack many features important in the taxonomy of these fungi,
based mainly on the morphological features of asci and asco-
spores. Thus, determination of the generic position of sporo-
carps of fossil fungi is rather difficult, and in many cases
resulted in serious mistakes. Contrary to the taxonomy of
present-day fungi that could be based on molecular data ob-
tained from DNA analysis, in the fossil state DNA is almost
never preserved and there is no possibility to use molecular
investigation in the case of fossil fungi. Considering these
limitations, the taxonomy of sporocarps of fossil
(epiphyllous) fungi is usually based on an artificial morpho-
logical taxonomic system (Elsik 1978).
One of the most frequently found fossil sporocarps are
those classified as an artificial fossil-genus Trichothyrites
Rosend., related to the modern family Trichothyriaceae.
Fossil-species Trichothyrites hordlensis P.H. Sm., based on
the morphological similarity of their ostiolar collar to the con-
temporary Trichothyrina ammophillae Ellis [=Lichenopeltella
ammophilae (J.P. Ellis) P.M. Kirk & Minter], suggested by
Section Editor: Gerhard Rambold
*Grzegorz Worobiec
g.worobiec@botany.pl
1
W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz
46, PL-31-512 Kraków, Poland
https://doi.org/10.1007/s11557-020-01598-0
Mycological Progress (2020) 19:799–804

Smith (1980), has been proposed by Samarakoon et al. (2019)
as a representative of genus Lichenopeltella Höhn.
Samarakoon et al. (2019) did not propose the use of the
fossil-genus Trichothyrites for divergence calibration,
avoiding the discussion about the inclusion of Trichothyrites
in modern genus Lichenopeltella. In addition, morphological-
ly identical to Trichothyrites hordlensis sporocarps can be
found not only in the genus Lichenopeltella, but also in other
genera of the Trichothyriaceae and in the Microthyriaceae
family (see Wu et al. 2011). Thus, the taxonomical relation-
ships of Trichothyrites hordlensis and genus Lichenopeltella
based only on ostiolar collar morphology seem rather weak.
During palynological investigations of pollen samples tak-
en from upper Pliocene deposits from Mizerna, Poland, we
found remains of fungal sporocarps corresponding to the
fossil-genus Trichothyrites,althoughhavinganostiolarcollar
with preserved setae. The morphology of ostiole with setae of
these trichothyrites-like fossil sporocarps is rather character-
istic and seems unique for sporocarps of some modern species
of genus Lichenopeltella (Spooner and Kirk 1990). After con-
sidering this, we decided to describe a new fossil-species of
genus Lichenopeltella. We discuss its recent counterparts and
the importance of presence of the remains of genus
Lichenopeltella for the palaeoecology of fossil assemblages.
Material and methods
The Mizerna-Nowa borehole was drilled in 1979 in the east-
ern part of the Nowy Targ Intramontane Depression, Polish
Western Carpathians (49° 28′N, 20° 18′E), southern Poland.
The borehole was 39 m deep and the core contained deposits
of a shallow late Pliocene palaeolake (Birkenmajer and
Worobiec 2013; Worobiec and Birkenmajer 2014). In the
fine-grained sediments of the Mizerna palaeolake, well-
preserved spores, pollen grains, and non-pollen
palynomorphs, including algae and fungal remains, were re-
corded (Worobiec et al. 2017). The results of the palynologi-
cal analysis were used to reconstruct vegetation in and around
the palaeolake (Birkenmajer and Worobiec 2013). The water
body was surrounded by herbaceous vegetation, with sedges
and grasses, as well as by mixed and coniferous forests.
The samples from Mizerna-Nowa were processed using
successively hydrochloric acid, potassium hydroxide and
hydrofluoric acid to remove silicates (Moore et al. 1991). A
total of 160 samples have been studied. Remains of the stud-
ied fungal sporocarps were recorded in one sample from 26.4-
m depth. From this sample, 22 microscope slides were made,
using glycerine jelly as a mounting medium. Fossil specimens
are housed in the W. Szafer Institute of Botany, Polish
Academy of Sciences (Kraków), under catalogue number
Mizerna-Nowa 1979/244.
The terminology for the morphology of fungal remains of
Lichenopeltella follows Korf (1958), Kirk et al. (2008)and
Wu et al. (2011). The method of measuring the size of fungal
structures depended on their shape. Diameter measurements
were used for regular, round or broadly elliptical structures,
and length and width for quadrangular structures.
Microphotographs were taken with a Nikon Eclipse E400 mi-
croscope fitted with a Canon A640 digital camera.
The classification of fossil and recent fungi follows
Kalgutkar and Jansonius (2000), Wu et al. (2011), Hyde
et al. (2013) and Wijayawardene et al. (2018).
Results
Taxonomy
Dothideomycetes, families incertae sedis
Trichothyriaceae Theiss. (1914)
Lichenopeltella Höhn. (1919)
Lichenopeltella mizerniana G. Worobiec sp. nov. Fig. 1–6
MycoBank: MB835847
Type: Slide Mizerna-Nowa 244(2), slide location 6/24.
Stored in W. Szafer Institute of Botany, Polish Academy of
Sciences, Kraków, collection Mizerna-Nowa 1979/No
244(2). Illustrated on Fig. 1–3
Etymology: The species name refers to the type locality
Type locality: Mizerna-Nowa, southern Poland
Stratigraphic horizon: upper Pliocene
Distribution: Mizerna-Nowa, southern Poland
Other specimens examined: Collection Mizerna-Nowa
1979/No 244(10), W. Szafer Institute of Botany, Polish
Academy of Sciences, Kraków; Poland
Repository: W. Szafer Institute of Botany, Polish Academy
of Sciences, Kraków, Poland, collection Mizerna-Nowa 1979
Diagnosis. Sporocarps orbicular, scutate, ostiolate, margin
slightly sinuate. Upper wall of scutellum composed of hardly
visible cells, arranged in radiating rows extending outward
(textura prismatica). Ostiole central, surrounded by a clearly
delimited collar composed of dark cells. Setae attached to the
ostiole, divergent, smooth, non-septate, thick-walled, with
flask-shaped base and acute apex.
Description. Sporocarps (catathecioid ascoma) orbicular,
scutate, ostiolate, about 80 μm in diameter, margin slightly
sinuate. Upper wall of scutellum composed of hardly visible
cells, arranged in radiating rows extending outward (textura
prismatica), about 5 μm in size. Lower wall (basal plate) not
visible. Ostiole central, presumably 10 μm in diameter,
surrounded by clearly delimited collar composed of dark,
more or less isodiametric cells, 2–4μminsize.Setaepresent,
attached to ostiole, divergent, smooth, non-septate, with rather
thick walls (2.0–2.5 μm), 15–50 μm long and about 5 μm
800 Mycol Progress (2020) 19:799–804

wide. Base of setae flask-shaped, about 7.5 μm in size, apex
acute. Vegetative hyphae and ascospores absent.
Discussion
The above described fungal sporocarps morphologically cor-
respond to the fossil-genus Trichothyrites.Sporocarpsofthis
genus are roundish, thyriothecium-like with a centrally locat-
ed ostiole usually surrounded by a distinct collar composed of
some rows of thick-walled and melanized cells. Fossils of this
type were reported starting from the beginning of the twenti-
eth century and were usually classified as representatives of
the Microthyriaceae family (e.g. fig. 7,Kräusel1920).
Rosendahl (1943), on the basis of fossil sporocarps from early
Pleistocene deposits at Springfield, Minnesota, created a new
fossil-genus Trichothyrites to accommodate fossil sporocarps
with a central ostiole. For the same type of fossils, Cookson
(1947) created a new fossil-genus Notothyrites Cookson and
assigned it to the Microthyriaceae family. Later, Venkatachala
and Kar (1969), for trichothyriaceous sporocarps, proposed a
new fossil-genus Sphaerialites Venkatach. & R.K. Kar.
Considering the priority of the name Trichothyrites,both
Notothyrites and Sphaerialites should be considered younger
synonyms of the fossil-genus Trichothyrites (Kalgutkar and
Jansonius 2000). Smith (1980) described a new fossil-species
T.hordlensis, along with some other morphotypes of sporo-
carps of trichothyriaceous fungi, and emended diagnosis of
fossil-genus Trichothyrites. He wrote that the “Uppermost tier
(ostiolar margin) of cells may have short prolongations (setae)
in some cases”. Most of the fossil-species of Trichothyrites,
however, have no setae around the ostiole (Kalgutkar and
Jansonius 2000;Samarakoonetal.2019). The fossil-species
T. setifer (Cookson) R.K. Saxena & N.K. Misra has up to
eight setae, up to 13 μm long, non-septate, arising from ostiole
cells (Cookson 1947). Another two fossil-taxa, T. denticulatus
(Ramanujam & K.P. Rao) Kalgutkar & Janson. and
T. echinatus (K.P. Rao & Ramanujam) Kalgutkar & Janson.
have short setae-like processes oriented towards the ostiole
cavity (Kalgutkar and Jansonius 2000). All fossil-taxa listed
above were described from pre-Quaternary deposits.
Thyriothecioid sporocarps with prominent setae were found
21 3
456
Fig. 1–6Lichenopeltella
mizerniana sp. nov. 1–2
sporocarp. 3Detail of ostiole and
setae. 4–6Detached collar of
ostiole with attached setae, note
structure of setae. Bars 1–2, 4–
5=20μm, 3, 6 = 10 μm
9
78
Fig. 7–9Diverse
trichothyriaceous sporocarps
from the upper Pliocene of
Mizerna-Nowa. Bars = 20 μm
801Mycol Progress (2020) 19:799–804

in Quaternary deposits as well, e.g. in Pleistocene deposits in
Great Britain (Godwin and Andrew 1951), Finland (Eriksson
1978) and in the Holocene of the Pyrenees (López-Vila et al.
2014). Fungal sporocarps in form of thyriothecium (or
catathecium) with setae around the ostiole have only the mod-
ern fungal genera Chaetothyriothecium Hongsanan & K.D.
Hyde and Lichenopeltella. Among them, catathecia with setae
around the ostiole with its morphology directly corresponding
to the setose representatives of fossil-genus Trichothyrites
have only some species of the modern genus
Lichenopeltella.Chaetothyriothecium elegans Hongsanan &
K.D. Hyde, a representative of Microthyriales, has
thyriothecia similar to Lichenopeltella, but differs consider-
ably, having septate and more numerous setae (Hongsanan
et al. 2014). Many other fungal genera have setae around or
close to the ostiole [e.g. Capronia Sacc., Coniochaeta (Sacc.)
Cooke, Phoma Sacc., Pleospora Rabenh. ex Ces. & De Not,
Podospora Ces., Pyrenochaeta De Not., Setosphaeria K.J.
Leonard & Suggs, Trichomerium Speg., and Wentiomyces
Koord.] but differ essentially in having perithecium-like asco-
carps. Thus, fossil fungal thyriothecia (or catathecia) with a
ostiole possessing non-septate setae could be directly related
to the Lichenopeltella genus. Taking into account the above
deductions we decided to describe a new fossil-species,
Lichenopeltella mizerniana. Earlier mentioned setae-bearing
fossil-species Trichothyrites setifer, most probably also relat-
ed to Lichenopeltella, differs in having shorter setae (Cookson
1947). Relationships of fossil Trichothyrites denticulatus and
T. echinatus that have short setae-like outgrowths to the genus
Lichenopeltella seem unclear. Fossil Lichenopeltella
mizerniana could be compared with a modern species of
Lichenopeltella, bearing setae around the ostiole. Aptroot
et al. (1997) listed 7 lichenicolous species of Lichenopeltella
with ostiolar setae known at the time [L. bunodophoronis
Diederich, L. lobariae Etayo & Diederich,
L. pannariacearum Diederich, L. peltigericola (D.
Hawksw.) R. Sant., L. santessonii (P.M. Kirk & Spooner) R.
Sant., L. swaminathaniana Hariharan, Mibey & D. Hawksw.,
and L. setifera Matzer]. Among these, only L. peltigericola,
similarly to Lichenopeltella mizerniana, has divergent setae
around the ostiole (Aptroot et al. 1997). Later, von Brackel
(2010) described two other setae-bearing lichenicolous spe-
cies, L.uncialicola Brackel and L.rangiferinae Brackel, both
with divergent setae around the ostiole. In terms of the collar
structure (see fig. 1,vonBrackel2010, and fig. 2,vonBrackel
2011), fossil L. mizerniana (Fig. 1–3) seems more similar to
L. uncialicola than to L. peltigericola. Considering the non-
lichenicolous taxa of Lichenopeltella, there are four modern
species which have an ostiolar collar surrounded by setae:
L. alpestris (Sacc.) P.M. Kirk & Minter, L. ammophilae (J.P.
Ellis) P.M. Kirk & Minter, L. palustris (J.P. Ellis) P.M. Kirk &
Minter, and Micropeltopsis (= Lichenopeltella,
Wijayawardene et al. 2017)cannabis McPartl. (Ellis 1977;
Kirk and Spooner 1989; McPartland and Cubeta 1997).
Lichenopeltella mizerniana, having divergent setae (Fig. 1–
3), seems closest to L. ammophilae,L. palustris, and less so to
Micropeltopsis cannabis,contrarytoL. alpestris with its con-
vergent setae (Ellis 1977). The overall shape and orientation
of the setae of L. palustris (see fig. 10 a, b, Ellis 1977)are
more similar to Lichenopeltella mizerniana (Fig. 1–6)thanto
L. ammophilae (see fig. 2c, Kirk and Spooner 1989).
L. ammophilae and L. palustris are found on the dead stems
of mono- and dicotyledons (Ammophila arenaria,Filipendula
ulmaria,Phalaris arundinacea,andThalictrum flavum, Ellis
1977)andL. palustris on peatmoss Sphagnum (Laukka 2005).
Following Smith (1980) and Samarakoon et al. (2019), we
can agree that some of the remaining fossil-species of
Trichothyrites that have an ostiolar collar without setae could
probably also represent sporocarps of Lichenopeltella. Smith
(1980), on the basis of the ostiolar collar structure, compared
the fossil Trichothyrites hordlensis with the contemporary
species Lichenopeltella ammophilae (syn. Trichothyrina
ammophilae) (see Samarakoon et al. 2019). In Mizerna, L.
mizerniana was found accompanied by other
trichothyriaceous sporocarps (Fig. 7–9), resembling modern
Lichenopeltella ammophilae,Lichenopeltella nigroannulata
(J.Webster)P.M.Kirk&Minter,andLichenopeltella
pinophylla (Höhn.) P.M. Kirk & Minter as well. These sporo-
carps were found without any traces of setae around the osti-
ole. Similarly to the case of modern L. palustris (Ellis 1977)
and L. ammophilae (Kirk and Spooner 1989), which have
catathecia both with and without setae, it should not be
overruled that at least some of these sporocarps could also
represent the genus Lichenopeltella. On the other hand, the
thyriothecia of some other than Lichenopeltella modern fungi
without collar setae, e.g. from genus Arnaudiella Petr., espe-
cially species A.caronnae (Pass.) Petr. (family
Microthyriaceae, see Wu et al. 2011) have an ostiolar collar
rather similar to those fossil-species of Trichothyrites.
Genus Lichenopeltella Höhn.[syn. Didymopyrostroma
Bat. & Cavalc., Micropeltopsis Vain., Microthyris Clem.,
Trichothyrina (Petr.) Petr., Wu et al. 2011] belongs to the
family Trichothyriaceae Theiss. (Wu et al. 2011;
Wijayawardene et al. 2017) and comprises 48 modern species.
Among them, 39 are parasitic on lichens (Wu et al. 2011;
Diederich et al. 2018). The remaining taxa are partly
bryophilous and are partly found as saprobes on dying or dead
vascular plants (Marsh et al. 2010).
Ascocarps of Lichenopeltella which have both the upper
and lower wall (basal plate) are classified as both catathecia
(Pérez-Ortega and Spribille 2009) and as thryriothecia (Wu
et al. 2011; Hyde et al. 2013). The catathecia of
Lichenopeltella are circular, in section lenticular, with a most-
ly entire margin [very rarely fimbriate, as in case of
L. fimbriata (J.P. Ellis) P.M. Kirk & Minter], superficial,
black, and with a central ostiole. The upper wall (scutellum)
802 Mycol Progress (2020) 19:799–804

of these catathecia is composed of rectangular cells, arranged
in parallel lines radiating from the central ostiole (textura
prismatica). The lower wall (basal plate) cells have a similar
arrangement. Asci 8-spored, ascospores 2–3-seriate, hyaline,
ellipsoid, and 2-celled (comp. Wu et al. 2011). The presence
and orientation of setae is considered an important diagnostic
feature in modern Lichenopeltella (Ellis 1977; Spooner and
Kirk 1990; Aptroot et al. 1997; Earland-Bennett and
Hawksworth 1999;vonBrackel2010).
Genus Lichenopeltella is worldwide distributed from
tropical to polar areas (Aptroot et al. 1997; von Brackel
2010; Wijayawardene et al. 2017), and thus, annual
temperatures seem to not limit their range. On the other
hand, as there are no records of this genus from arid
areas, its distribution probably depends on significant
moisture of the environment resulting from an adequate
amount of precipitation. When considering this in rela-
tion to Lichenopeltella mizerniana,wecanonlyinfer
that the Pliocene climate of the Mizerna locality was
probably at least moderately humid. The results of ear-
lier palynological investigations of deposits from
Mizerna confirm a humid climate and point to mild,
temperate climatic conditions. In the sub-mountainous
area, the annual rainfall was probably higher than in
the lowlands (Birkenmajer and Worobiec 2013).
A significant problem in the palaeoecology of
Lichenopeltella mizerniana is the presumed host on which
the fossil-taxon existed at that time. As was noted before, most
of the modern species of Lichenopeltella are parasitic on li-
chens, but there are some bryophilous taxa and also sapro-
phytic on dying or dead vascular plants. Taking into account
that the morphology of Lichenopeltella mizerniana is most
similar both to some modern lichenicolous (L. peltigericola,
L. rangiferinae, and L. uncialicola) and non-lichenicolous
species (L. ammophilae,L. palustris), it is extremely difficult
to make a decision about the presumed host of Lichenopeltella
from Mizerna. No features other than the morphology of spo-
rocarp are preserved in fossil Lichenopeltella mizerniana (lack
of asci and ascospores), thus precluding any close comparison
with the listed modern Lichenopeltella species. Nonetheless,
we can make some assumptions on the host of Lichenopeltella
mizerniana after taking into account the results of palynolog-
ical investigations of the Mizerna deposits. The palynological
profile from Mizerna-Nowa (Birkenmajer and Worobiec
2013) shows the considerable presence of pollen from grasses
and sedges. Among the above suggested modern counterparts
of fossil Lichenopeltella mizerniana, only non-lichenicolous
L. ammophilae and L. palustris were found on the body of
grasses. Similarly in the case of the closest modern counter-
parts, it is probable that the presumed host/s of Lichenopeltella
mizerniana could be some grass/es.
Results of the investigation on fossil Lichenopeltella
from Mizerna are of great importance for mycologists.
We proved the presence of genus Lichenopeltella in the
past, dating it back at least to the late Pliocene (earlier
than 2.6 Ma BP). This dating is of importance for the
calibration of the divergence time estimations in the
phylogenetic trees of genus Lichenopeltella,aswellas
the Trichothyriaceae family.
Acknowledgements Dr. hab. Marcin Piątek (Department of Mycology,
W. Szafer Institute of Botany, Polish Academy of Sciences) is acknowl-
edged for valuable suggestions to the manuscript.We would like to thank
the anonymous reviewer for critically reading the manuscript and for his
suggestions.
Funding information This work was supported by the W. Szafer Institute
of Botany, Polish Academy of Sciences, Kraków, Poland through the
statutory funds.
Open Access This article is licensed under a Creative Commons
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