Lichenicolous fungi from the Holarctic. Part IV: New reports and a key to species on Dermatocarpon

Abstract

Endococcus peltigericola is reported for the first time from North America and Epigloea urosperma for the second time. The detection of interascal filaments in Endococcus peltigericola indicates that it may belong to the genus Polycoccum. Adelococcus immersus and Trematosphaeriopsis parmeliana are newly reported for Russia, the former is also first documented from the Arctic. The description of A. immersus, previously known only from a type specimen, is emended. Halecania alpivaga is reported new to the north-central Siberia growing on an unusual host genus (Dermatocarpon). A key to the species of lichenicolous fungi growing on Dermatocarpon is provided.

Full text

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Opuscula Philolichenum, 20: 44–53. 2021.
*pdf effectively published online 9April2021 via (http://sweetgum.nybg.org/philolichenum/)
Lichenicolous fungi from the Holarctic. Part IV: New
reports and a key to species on Dermatocarpon
MIKHAIL P. ZHURBENKO
1
ABSTRACT. – Endococcus peltigericola is reported for the first time from North America and
Epigloea urosperma for the second time. The detection of interascal filaments in Endococcus peltigericola
indicates that it may belong to the genus Polycoccum. Adelococcus immersus and Trematosphaeriopsis
parmeliana are newly reported for Russia, the former is also first documented from the Arctic. The
description of A. immersus, previously known only from a type specimen, is emended. Halecania alpivaga
is reported new to the north-central Siberia growing on an unusual host genus (Dermatocarpon). A key to
the species of lichenicolous fungi growing on Dermatocarpon is provided.
KEYWORDS. – Arctic, biodiversity discovery, biogeography, lichen-dwelling fungi, Russia,
Sweden, United States of America.
INTRODUCTION
This paper continues the author’s publications on noteworthy finds of lichenicolous fungi from the
Holarctic (Zhurbenko 2009a, 2009b, 2020). It is based on collections from Russia, Sweden and the United
States of America, mainly from the Arctic regions. In addition to providing new reports, range extensions,
taxonomic notes, and illustrations of the species studied, the first key to the lichenicolous fungi growing on
species of Dermatocarpon Eschw. is presented.
MATERIALS AND METHODS
Microscopy and imaging were undertaken using a Stemi 2000-CS stereomicroscope and a Zeiss
Axio Imager A1 compound microscope with Nomarski interference contrast, fitted with an AxioCam
MRc5 digital camera. Microscopic characters were studied from hand-made sections produced using a
razor blade and then mounted in water, a 10% aqueous solution of potassium hydroxide (K), Lugol’s Iodine
solution, directly (I), and after KOH pre-treatment (K/I). Measurements were taken from water mounts and
rounded to the nearest 0.5 μm. The length, width, and length/width ratio (l/w) of ascospores are given as
(min‒) (x
̄−SD)‒(x
̄+SD) (‒max), where ‘min’ and ‘max’ are the extreme values observed, ‘x
̄’ the arithmetic
mean, and ‘SD’ the corresponding standard deviation. Colours were determined according to the tables of
Kornerup and Wanscher (1978). Voucher specimens are deposited in LE and UPS.
RESULTS
Adelococcus immersus Etayo & Breuss
FIGURE 1.
NOTES. – The examined material generally fits the protologue (Etayo & Breuss 1998), including
the ascospore size, (9.5–)12.5–16.5(–20) × (5.5–)6.5–8.5(–10) µm, l/w = (1.2–)1.6–2.4(–3.2) (n = 90)
1
MIKHAIL P. ZHURBENKO – Laboratory of the Systematics and Geography of Fungi, Komarov Botanical
Institute, Russian Academy of Sciences, Professor Popov Street, 2, St. Petersburg, 197376, Russia. –
email: zhurb58@gmail.com

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Figure 1. Adelococcus immersus growing on Dermatocarpon miniatum var. complicatum (from Zhurbenko
0777). A, habitus of ascomata. B, lateral exciple in cross-section in water. C, asci and ascospores in water.
D, asci and ascospores in I. E, asci and ascospores in K/I. Scale bars: 200 µm in A, 10 µm in B–E.
versus 13–18(–20) × 7–8(–9.5) µm. However, the perithecia are often almost superficial, aggregated to
occasionally concrescent, black, rather glossy, subglobose to occasionally broadly ovoid, with a brown
exciple, 150–320 µm in diameter, and with ostioles 20–30 µm in diameter. In contrast, the protologue
describes the perithecia as completely immersed, ovoid, up to 220 µm in diameter, and with a colourless
exciple. It is likely that the discrepancy in reported excipular pigmentation relates to the degree of
immersion of the perithecia wherein less immersed areas of the perithecia are exposed to sunlight, causing
the exciple to become brown pigmented. Additionally, the hymenial gel is not I− as was described in the
protologue, but instead is I+ pastel red, which is typical for Adelococcus Theiss. & H. Sydow (Hafellner
2004a). The ascospores are also not always ellipsoid as was described in the protologue, but occasionally
can be subglobose, slightly curved and/or attenuated at one end, very rarely aseptate. They are also finely
verruculose (described as “smooth or very weakly ornamented” in the protologue), which is best seen using
the Nomarski interference contrast.
This species was previously known only from the type specimen collected in Turkey on
Dermatocarpon biennense Zsch. (Etayo & Breuss 1998), a species close to D. miniatum (L.) W. Mann

46
Figure 2. Endococcus peltigericola growing on Peltigera aphthosa (from Zhurbenko 0091). A, habitus of
ascomata fringing cephalodia of the host. B, squashed exciple when viewed from above, in water. C,
interascal filaments in water. D, asci in water. E, ascospores in water. F, conidia in water. Scale bars: 200
µm in A, 10 µm in B–F.
(Breuss & John 2004). Here it is reported for the first time from Russia and the Arctic, and D. miniatum
var. complicatum (Lightf.) Th. Fr. is a new host. As an aid in the identification of this and other
lichenicolous fungi on Dermatocarpon, a key to the fungi known from the host is presented here (see
Appendix).
Specimens examined (both on thalli of Dermatocarpon miniatum var. complicatum) – RUSSIA.
KRASNOYARSK TERRITORY: Taimyr Peninsula, Byrranga Mountains, N of Levinson-Lessing Lake,
74°31′N, 98°36′E, elev. 300 m, rocks in tundra, 24.vii.1995, M.P. Zhurbenko 95605 (LE 310199); Taimyr
Peninsula, Byrranga Mountains, W of Levinson-Lessing Lake, 74°30′N, 98°30′E, elev. 120 m, rocks in
tundra, 27.viii.1995, M.P. Zhurbenko 95604 (LE 310200).
Endococcus peltigericola Zhurb. & Stepanchikova
FIGURE 2.
NOTES. – This fungus is characterized by superficial perithecioid, brown to blackish ascomata,
sometimes with indistinct interascal filaments, the presence of conidia, and its confinement to species of
Peltigera Willd. The examined material fits the protologue (Zhurbenko et al. 2012), including its brown to
blackish superficial ascomata (45–)50–80(–90) µm in diameter (n = 21), asci that are 33–45 × 9.5–10 µm
(n = 8), ascospores that are (9.5–)10–12(–12.5) × (3.5–)4–4.5(–5) µm, l/w = (2.0–)2.3–2.9(–3.2) (n = 27),

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Figure 3. Epigloea urosperma growing on Placynthiella icmalea (from Magnusson 12934). A, habitus of
ascomata. B, asci and ascospores in K/I. Scale bars: 200 µm in A, 10 µm in B.
and conidia that are aseptate, pale brown, ellipsoid, oblong or broadly clavate, often somewhat truncated at
the base, 5–6.5 × 2.5–3.5 µm in size. The presence of branched interascal filaments reported in this species
by Zhurbenko and Vershinina (2014) is also confirmed here, which suggests it belongs to the genus
Polycoccum Körb. Compared with the other species of Polycoccum that grow on Peltigera, E. peltigericola
is quite similar to P. superficiale D. Hawksw. & Miądl. also characterized by superficial ascomata, which
are unusual for the genus, and is known from Finland, Germany, Poland, Asian and European parts of
Russia, and Sweden (Czyźewska 2003, Hawksworth & Miądlikowska 1997, Puolasmaa et al. 2012,
Zhurbenko & Vershinina 2014). However, according to the protologue (Hawksworth & Miądlikowska
1997) and the description in Zhurbenko and Vershinina (2014), the latter species differs in having larger
asci, (43–)50–64(–73) × (12–)14–17(–20) µm, and ascospores (10.5–)13.5–17(–19) × (5–)5.5–7(–8.5) µm.
Further studies are needed to determine whether these taxa are conspecific. Until this issue is resolved, I
prefer not to change the generic position of E. peltigericola.
This species was previously known only from the type specimen collected in the Kamchatka
Territory of Russia growing on Peltigera membranacea (Ach.) Nyl. (Zhurbenko et al. 2012) and is newly
reported here for North America. Peltigera aphthosa (L.) Willd. is a new host species.
Specimen examined. – U.S.A. ALASKA. INTERIOR: ca. 50 km ENE of Fairbanks, Caribou-
Poker Creeks Watershed, Alnus crispa-Betula papyrifera-Populus tremuloides forest, 65°10.133′N,
147°30.241′W, elev. 305 m, on Peltigera aphthosa (cephalodia, mainly fringing their margins; thallus),
26.viii.2000, M.P. Zhurbenko 0091 (LE 310184).
Epigloea urosperma Döbbeler
FIGURE 3.
NOTES. – According to Döbbeler (1994), this fungus is characterized by immersed to superficial
perithecia that are minute, inconspicuous, blackish, glabrous, glossy, subglobose, usually with a well-
developed depression around the ostiole, 60–150 µm in diameter, and with a thin, light brown, gelatinous
exciple consisting of periclinally-oriented hyphae. The paraphyses are filiform, simple or rarely branched,
not capitate at the apex, and surrounded by gelatinous sheaths. The asci are elongate-clavate, short-stalked,
rounded at the apex, without any distinct apical apparatus, discharging by longitudinal splits, mostly 32-
spored, with I+ and K/I+ evenly blue wall. The ascospores are hyaline, narrowly ellipsoid, 5–9 × 2–2.5 µm,
1-septate, smooth-walled, non-halonate, irregularly arranged in the ascus, and with thread-like appendages
at both ends that are up to 6 µm long. The specimens cited here match these characters.
The species was previously known from Africa (Republic of Seychelles; Diederich et al., 2017),
Asia (Russia; Urbanavichus et al. 2020), Europe (Austria, Czech Republic, England, Estonia, Finland,
Germany, Luxembourg, Poland, Slovakia, Spain, Sweden, Switzerland; Brackel 2014, Diederich et al.
2012, Döbbeler 1994, Kukwa & Flakus 2009, Malíček et al. 2018, Myllys & Launis 2018, Pérez-Ortega &
Barreno 2006, Suija et al. 2007, Vondrák et al. 2015), North America (U.S.A.; Spribille et al. 2020) and
South America (Bolivia; Flakus & Kukwa 2012) where it grows on Placynthiella dasaea (Stirt.) Tønsberg

48
Figure 4. Halecania alpivaga growing on Dermatocarpon miniatum var. complicatum (from Zhurbenko
961034). A, habitus of apothecia. B, apothecium in cross-section in water. C, paraphyses and ascus with
spores, in water. D, hymenium with ascus in K/I. E, ascospores with perispore in K. Scale bars: 200 µm in
A, 20 µm in B, 10 µm in C–E.
and P. uliginosa (Schrad.) Coppins & P. James (the latter is the host of the type specimen). Here it is
reported for the second time from North America based on a second collection from Alaska. As the species
is only known from a small number of occurrences in Sweden, an additional specimen located while
studying material at UPS is included here.
Specimens examined. – SWEDEN. VÄSTERGÖTLAND: Ale-Skövde par., Prässebo, on thallus
of Placynthiella icmalea (Ach.) Coppins & P. James growing on a stump, 27.viii.1931, A.H. Magnusson
12934 (UPS F-857610). U.S.A. ALASKA. INTERIOR: ca. 50 km ENE of Fairbanks, Caribou-Poker
Creeks Watershed, Picea mariana forest, 65°09.391′N, 147°29.388′W, elev. 295 m, on sterile lichen thallus
growing on top of Peltigera malacea (Ach.) Funck, 13.ix.2000, M.P. Zhurbenko 00410 (LE 310185).
Halecania alpivaga (Th. Fr.) M. Mayrhofer
FIGURE 4.
NOTES. – Information on the biology of this species is somewhat contradictory. According to
Fletcher and Coppins (2009) it is a lichen, that grows on calcareous rocks in arctic-alpine environments of

49
the Holarctic. However, Poelt and Mayrhofer (1988) cited it as an example of a cyanotrophic lichen
growing on saxicolous cyanolichens of the genus Placynthium (Ach.) Gray or on free-living cyanobacteria
of the genera Stigonema Bornet & Flahault and Gloeocapsa Kützing. It was also reported growing on
species of Physcia (Schreb.) Michx. (Mayrhofer 1987, Poelt 1958, Santesson et al. 2004), Collema Weber
ex F.H. Wigg. agg., Dermatocarpon and Toninia A. Massal. (Rambold & Triebel 1992). As the specimen
reported here was found on an unusual host, namely on D. miniatum var. complicatum, which is lichenized
with green algae, a brief description is given below. This is the first report of the species from north-central
Siberia (Kotlov 2003, Zhurbenko 2009a).
DESCRIPTION. – Lichenized thallus indistinct. Apothecia sessile, constricted at the base, 370–620
µm in diameter, disc dark brown, ±flat, thalline margin medium brownish orange to pale orange, mostly
prominent, 50–100 µm wide, 100–180 µm thick, in cross-section with brown outer edge 20–30 µm wide;
aggregated, occasionally contiguous. Hypothecium colourless, 100–150 µm tall, composed of isodiametric
or slightly elongated, thick-walled cells. Hymenium 60–100 µm tall, including epihymenium 15–20 µm
tall, not inspersed, colourless except for the brown, K− epihymenium, I+ and K/I+ blue. Paraphyses
filiform, 1.5–2 µm in diameter, often with swollen brown apices, 3.5–6.5 µm in diameter, occasionally
branched. Asci of Catillaria-type, with amyloid tholus without axial structures, with amyloid outer coat,
elongate clavate, (49–)53–60(–61) × (9.5–)10.5–12.5(–14) µm (n = 11), 8-spored. Ascospores colourless,
very narrowly obovate (wider above), 1-septate, not constricted at the septum, (13–)13.5–16.5(–18) × (4–
)4.5–6(–6.5) µm, l/w = (2.2–)2.5–3.5(–4.6) (n = 24), guttulate, with perispore clearly visible in K, ca. 2 µm
thick (in K), irregularly biseriate in the ascus.
Specimen examined. – RUSSIA. KRASNOYARSK TERRITORY: Severnaya Zemlya
Archipelago, Bol’shevik Island, near western coast of Akhmatov Bay, 10 km S of the Bazovaya River
mouth, 79°03′N, 102°42′E, elev. 60 m, polar desert, semi-dried bed of a stream, on thallus of
Dermatocarpon miniatum var. complicatum growing on rock, 16.vii.1996, M.P. Zhurbenko 961034 (LE
310198).
Trematosphaeriopsis parmeliana (Jacz.) Elenkin
FIGURE 5.
NOTES. – This fungus is characterized by its occurrence on Xanthoparmelia (Vain.) Hale,
pseudothecia that are completely immersed in bullate galls, a hyaline to pale brown upper exciple, simple
to occasionally septate, apically not swollen interascal filaments, I− and K/I− hymenial gel, fissitunicate,
subcylindrical, 8-spored, I− and K/I− asci, and ascospores that are very narrowly ellipsoid to fusiform,
initially smooth and hyaline, finally becoming verruculose and occasionally pale yellow, sometimes
halonate, (0–)3-septate, sometimes slightly constricted at the septa (Hafellner 2004b; specimen cited
below). It is known from scattered localities in Africa (Canary Islands; Hafellner 2001), Asia (Mongolia;
Elenkin 1901, Hafellner 2001), Europe (Austria, Germany, Italy; Brackel 2010; Hafellner 2001, 2004b),
North America (Mexico, U.S.A.; Hafellner 2001, 2004b) and South America (Venezuela; Hafellner 2001)
and is here newly reported for the Asian part of Russia from the Mongolian border region.
Specimen examined. – RUSSIA. REPUBLIC OF BURYATIA: Dzhida District, Borgoi steppe, 4
km W of Beloozersk, 50º39′N, 105º40′E, elev. 650 m, steppe, on Xanthoparmelia camtschadalis (Ach.)
Hale (thallus) growing on soil, 21.vi.2005, M.P. Zhurbenko 0538a (LE 310189a).
ACKNOWLEDGEMENTS
The Rolf Santesson Foundation, Roland Moberg and Martin Westberg are thanked for giving the author the
opportunity to revise Rolf Santesson's collection of lichenicolous fungi in the herbarium of Uppsala University. The
author’s work was carried out within the framework of the research project of the V. L. Komarov Botanical Institute of
the Russian Academy of Sciences “Biodiversity, ecology, structural and functional features of fungi and fungus-like
protists” (АААА-А19-119020890079-6) using the equipment of its Core Facility Center “Cell and Molecular
Technologies in Plant Science”. Javier Etayo provided valuable comments on the identification of Adelococcus
immersus.

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Figure 5. Trematosphaeriopsis parmeliana growing on Xanthoparmelia camtschadalis (from Zhurbenko
0538a). A, galls with immersed pseudothecia. B, gall with immersed pseudothecia in cross-section in water.
C, interascal filaments in K. D, ascospores in water. Scale bars: 1 mm in A, 50 µm in B, 10 µm in C & D.
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APPENDIX – KEY TO THE LICHENICOLOUS FUNGI GROWING ON DERMATOCARPON
This key was compiled from the literature cited under each species below under ‘Lit.’. References
to taxonomic descriptions of the species are given in bold font, those that exclusively related to the
occurrence of the species on Dermatocarpon are given in regular font. The names of species that are
exclusively or mainly confined to species of Dermatocarpon are given in bold. Lichenized species are
denoted by ‘L’. Homostegia dermatocarponis Alstrup & M. Cole and Phaeospora dermatocarponis (R.G.
Werner) S.Y. Kondr., reported from species of Dermatocarpon by Alstrup and Cole (1998) and Werner
(1963) respectively, are considered to be synonyms of Opegrapha pulvinata Rehm var. pulvinata
(Diederich et al. 2018). Non-lichenicolous lichens occasionally found on species of Dermatocarpon (e.g.,
Agonimia tristicula (Nyl.) Zahlbr. fide Hafellner et al. 2005; Myriolecis semipallida (H. Magn.) Sliwa,
Zhao Xin & Lumbsch fide Poelt et al. 1995, Valadbeigi & Sipman 2010) are not included.
1. Ascomata apothecia ..................................................................................................................................... 2
2. Apothecia lirellate, shortly elongate, aggregated in dense clusters; hymenium opening with a slit;
ascospores soon becoming light brown, 3-septate, (18–)20–24(–26) × 6–7(–8) µm. Lit.: Ertz & Egea
(2007) ................................................................................... Opegrapha pulvinata Rehm var. pulvinata
2. Apothecia discoid, hymenium wide-open, ascospores colourless ........................................................ 3
3. Ascospores mostly or exclusively 3-septate .................................................................................. 4
4. Apothecial disc with a bluish-grey pruina, up to 0.5 mm in diameter; hypothecium
colourless; ascospores 3-septate, 18.5–23 × 4–4.5 µm. Note. This is a poorly known species
described from a find on thallus of Dermatocarpon sp. growing on a calcareous rock in
the Kyrgyz Republic. It was characterized as a parasite that does not develop its own thallus,
except for the apothecial thalline margin. Lit.: Makarevich (1971)
LLecania ferganae Oksner
4. Apothecial disc without a bluish-grey pruina, up to 1.1 mm in diameter; hypothecium
reddish-brown; ascospores (0–)3-septate, 10.5–23.5 × 3–7 µm. Lit.: Ladd & Morse (2018) ....
............................................................................................ Toninia tecta C.A. Morse & Ladd
3. Ascospores 0 or 1-septate .............................................................................................................. 5
5. Ascospores 1-septate, halonate (best seen in K), 13–18 × 4–6.5 µm. Lit.: Mayrhofer
(1987), present paper ..........................................
LHalecania alpivaga (Th. Fr.) M. Mayrhofer
5. Ascospores 0-septate, non-halonate ....................................................................................... 6
6. Apothecia up to 1 mm in diameter, disc brownish or olive-brown, ascospores 9–14 ×
5.5–6 µm. Lit.: Poelt (1974), Zhurbenko (2009a) ................................................................
.................................... LLecanora dispersa (Pers.) Sommerf. f. parasitans (Wedd.) Harm.
6. Apothecia up to 2.7 mm in diameter, disc shades of yellow, ascospores 9–12 × 4.5–6
µm. Lit.: Ryan et al. (2004) ............................................
LLecanora thallophila H. Magn.
1. Ascomata perithecia .................................................................................................................................... 7

53
7. Ascospores hyaline, 1-septate, 16–18 × 4.5–5.5 µm. Lit.: Keissler (1930), Santesson et al. (2004) .....
............................................................................................. Stigmidium stygnospilum (Minks) R. Sant.
7. Ascospores brown, (0–)1 or 3-septate .................................................................................................. 8
8. Ascospores (0–)1-septate, 9.5–20 × 5.5–10 µm. Lit.: Etayo & Breuss (1998), present paper .....
............................................................................................ Adelococcus immersus Etayo & Breuss
8. Ascospores 3-septate, 24–26 × 6.5–9 µm. Lit.: Hafellner (1999), Hawksworth (1986) ................
.................................................................................... Pyrenidium hetairizans (Leight.) D. Hawksw.