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39Lichenicolous fungi from the Holarctic. Part V
M. P. Zhurbenko
Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia
zhurb58@gmail.com
Abstract. Endococcus collematis is reported for the first time for Asia and Russia. Xenonectriella physciacearum and X. zimmermanni are reported for the first time for Russia. Sclerococcum zhurbenkoi is recorded for the first time for Asia, North America and the Arctic; the records are based on materials from Norway, Russia, Canada and the USA. A specimen of Xenonectriella cf. physciacearum with unusually long ascospores is briefly described. Distinctive characters of every species with references to previous descriptions and illustrations are provided.
Keywords: Endococcus collematis, Sclerococcum zhurbenkoi, Xenonectriella physciacearum, Xe-nonectriella zimmermanni, biodiversity, biogeography, lichen-dwelling fungi.
Лихенофильные грибы из Голарктики. Часть V М. П. Журбенко
Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия
zhurb58@gmail.com
Резюме. Приведены первые указания Endococcus collematis для Азии и России, Xenonectriella physciacearum и X. zimmermanni — для России, Sclerococcum zhurbenkoi — для Азии, Северной Америки и Арктики, в том числе для Канады, Норвегии, России и США. Образец Xenonectriella cf. physciacearum характеризуется необычно длинными аскоспорами. Для каждого вида приводятся ссылки на описание и иллюстрации, а также отличительные признаки.
Ключевые слова: Endococcus collematis, Sclerococcum zhurbenkoi, Xenonectriella physciacearum, Xenonectriella zimmermanni, биоразнообразие, биогеография, обитающие на лишайниках грибы.
This paper continues the series of publications on noteworthy findings of lichenicolous fungi from the Holarctic (Zhurbenko, 2009a, b, 2020, 2021a). It is the result of a revision of specimens of lichenicolous fungi based on recent taxonomic treatments of some critical groups of these fungi (Berger et al., 2020; Brackel, 2021; Zimmermann, Berger, 2021).
Materials and Methods
Materials from the Mycological herbarium of the Komarov Botanical Institute (LE), mainly collected by the author in different regions of Russia and the world, were studied. The collector of the specimens examined is M. P. Zhurbenko, unless otherwise stated. Voucher specimens are kept in LE or private herbarium of V. Otte (herb. Otte).
Microscopy and imaging were undertaken using a Zeiss Axio Zoom.V16 stereo-microscope fitted with an AxioCam 712 colour digital camera and a Zeiss Axio Imager A1 compound microscope equipped with Nomarski differential interference contrast
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optics and fitted with an AxioCam 506 colour digital camera. Microscopic characters were studied using sections hand-cut with a razor blade and mounted in water, 10% potassium hydroxide (K), Lugol's iodine directly (I) or after a K pretreatment (K/I), and lactic acid. Measurements were taken from water mounts, unless otherwise indicated, and rounded to the nearest 0.5 jm. The length, width and length/width ratio (l/w) of the ascospores are given as (min) (x-SD)-(x+SD) (max), where 'min' and 'max' are the extreme values observed, x is the arithmetic mean and SD is the corresponding standard deviation.
Results
Endococcus collematis Brackel
Distinctive characters, description and illustrations. Detailed description and illustrations of this species are given in the protologue (Brackel, 2021). For more illustrations, see e. g., Zhurbenko (2021b: Fig. 1, as Endococcus pseudocarpus). Its main features are as follows: ascomata perithecioid, 120-200 jm diam.; interascal gel I+ and K/I+ slightly orange; hamathecium composed of periphyses, interascal filaments not observed; asci clavate, 40-55 x 10-13 jm, (4)8-spored; ascospores pale to rarely medium brown, narrowly ellipsoid to fusiform, 1-septate, with ±equal cells, smooth-walled, (8)11-14.5(17) x (3)3.5-4.5(5) jm, l/w = (2)2.6-3.7(5). Reported on species of Collema Weber ex F. H. Wigg., Enchylium (Ach.) Gray, and Lathagrium (Ach.) Gray.
Notes. Endococcus pseudocarpus Nyl. was described from the Pyrenees growing on squamules of Heppia guepinii (Delise) Nyl. var. nigrolimbata Nyl. (Nylander, 1873) which is probably a species close to Peltula euploca (Ach.) Poelt ex Ozenda et Clauzade (Hawksworth, Diederich, 1988: 309). In the protologue, Endococcus pseudocarpus is characterized by perithecia up to 150 jm diam., 8-spored asci, and slightly blackened, oblong to fusiform, 1-septate ascospores, 9-18 x 5-7 jm (Nylander, 1873). Subsequently, the species was understood in a broad sense and reported also from the lichen genera Callome Otalora et Wedin, Collema, Enchylium, Lathagrium, Placynthium (Ach.) Gray, and Scytinium (Ach.) Gray (Diederich et al., 2018; Brackel, 2021; Zhurbenko, 2021b). However, the heterogeneity of the Endococcus pseudocarpus s. l. complex has already been noted by some authors (Brackel, 2015; Brackel, Puntillo, 2016; Zhurbenko, 2021b). Eventually, Brackel (2021) proposed to narrow the broad concept of E. pseudocarpus and described a new species, E. collematis Brackel, which mainly differs from E. pseudocarpus s. str. in the width of the ascospores (3-5 jm vs 5-7 jm) and the host genera (Collema s. l. vs type genus Peltula Nyl.).
Endococcus collematis was previously known with certainty from Europe — Germany, Italy and Switzerland (Brackel, 2021). The specimens listed below are the first records of Endococcus collematis for Russia and Asia from the Komi Republic and the Irkutsk Region.
Specimens examined [both on thalli of Enchylium polycarpon (Hoffm.) Otalora et al.]: Russia, Komi Republic, 50 km SSE of Vorkuta, near 61 km cabin of the railway Vorkuta-Labytnangi by the right bank of Eletz River, 67°02'N, 64°26'E, 100 m a. s. l., limestone outcrops among shrub tundra, 29 VI 1993, Zhurbenko 93190, LE 260778 [previously published as Endococcuspseudocarpus (Zhur-benko, 2013)]; Irkutsk Region, 4 km SW of Elantsy, 52°46'N, 106°21'E, 590 m a. s. l., calcareous rocks on steppe slope among taiga forest, 8 VI 2005, Zhurbenko 05200, LE 310204 [previously published as E. pseudocarpus (Zhurbenko, 2021b)].
Sclerococcum zhurbenkoi F. Berger et E. Zimm.
Distinctive characters, description and illustrations. Detailed description and illustrations of this species are given in the protologue (Zimmermann, Berger, 2021). It produces galls eventually forming blackish doughnut-like structures, 100-200(-350) jm diam., with concave upper surface, coated with a conidiogenous layer. These structures are associated with a radially sprouting superficial net of thick, dark hyphae. Conidiogenous cells red- to dark-brown, terminal, integrated, of very different shape, hardly discernible. Conidiogenesis mainly monoblastic, rarely oligoblas-tic. Conidia brown, initially 1-septate, (10)12-16(18) x (7)8-11(12) jm, finally with (2-)3(-4) transverse septa, (12)18-23(27) x 9-11 jm, smooth to finely verrucose, normally aggregated in groups and hardly discernible from each other. Reported on Thamnolia vermicularis (Sw.) Schaer.
Notes. In the revision of lichenicolous fungi growing on lichens of the genus Thamnolia Ach. ex Schaer., «a rather common fungus with thick dark superficial hyphal strands and conspicuous black discoid sterile fruit-like structures» was noted (Zhurbenko, 2012). This fungus has recently been described as Sclerococcum zhurbenkoi from the high Alps and Pyrenees in Austria, Italy, France and Switzerland (Zimmermann, Berger, 2021). It is yet known only from the conidial state and is clearly visible macroscopically due to blackish, apothecia-like structures it forms, usually associated with conspicuous dark hyphae (Fig. 1).
Sclerococcum zhurbenkoi was previously known with certainty from Europe (Zimmermann, Berger, 2021). The specimens listed below are its first records for North America from the U. S. A. and Canada, and for Asia from the Krasnoyarsk Territory and the Republic of Sakha (Yakutia) in Russia. Also, it is recorded for the first time for Norway and for the Arctic.
Specimens examined: USA, Alaska, Denali National Park, Mt Healy, 63°50'N, 149°00'W, 600 m a. s. l., mountain tundra, on Thamnolia vermicularis var. vermicularis, 29 VIII 2000, Zhurbenko 00332, LE 310247. Canada, Ellesmere Island, Sverdrup Pass, 79°08'N, 81°27'W, 310 m a. s. l., polar desert, on T. vermicularis var. subuliformis (Ehrh.) Schaer., 28 VII 1992, F. Daniels, LE 310233. Norway, Svalbard, Spitsbergen, Dickson Land, W coast of Billefjorden, Mt Garmaksla, 78°37'N, 16°20'E, 200 m a. s. l., arctic tundra, on T. vermicularis var. subuliformis, 23 VII 2003, Zhurbenko 03455, LE 310246. Russia, Arkhangelsk Region, Franz Josef Land Archipelago, Hooker Island, Tikhaya Bay, 80°20'N, 52°47'E, polar desert, on T. vermicularis var. subuliformis, 29 VII 1930, V. P. Savicz, LE 310235; ibid., Hayes Island, 80°36'N, 57°54'E, polar desert, on T. vermicularis var. subuliformis, 11 VIII 2010, D. A. Walker, LE 310234; Krasnoyarsk Territory, Taimyr Peninsula,
Byrranga Mountains, near Levinson-Lessing Lake, 74°33'N, 98°41'E, 120 m a. s. l., limestones, arctic tundra, on T. vermicularis var. subuliformis, 3 VIII 1994, Zhurbenko 94513, LE 310245; ibid., 74°24'N, 98°46'E, 150 m a. s. l., limestones, arctic tundra, on T. vermicularis var. subuliformis, 1 VIII 1995, Zhurbenko 95609, LE 310241, Zhurbenko 95614, LE 310236; ibid, 74°29'N, 98°39'E, 250 m a. s. l., limestones, arctic tundra, on T. vermicularis var. subuliformis, 9 VIII 1995, Zhurbenko 95612, LE 310238; ibid., 74°31'N, 98°36'E, 200-250 m a. s. l., limestones, arctic tundra, on T. vermicularis var. subuliformis, 20 VIII 1995, Zhurbenko 95613, LE 310237; ibid, 26 VIII 1995, Zhurbenko 95611, LE 310239; ibid., 74°30'N, 98°30'E, 120 m a. s. l., arctic tundra, on T. vermicularis var. subuliformis, 27 VIII 1995, Zhurbenko 95608, LE 310242, Zhurbenko 95610, LE 310240; Severnaya Zemlya Archipelago, Bol'shevik Island, peninsula with Cape Baranova, 79°16'N, 101°40'E, 20 m a. s. l., polar desert, on T. vermicularis var. subuliformis, 9 VII 1996, Zhurbenko 961037, LE 310232; Republic of Sakha (Yakutia), Laptev Sea coast, 3 km SW of Tiksi town, near Lake Diring-Kyuel', 71°40'N, 128°40'E, 50 m a. s. l., arctic tundra, on T. vermicularis var. vermicularis, 17 VII 1998, Zhurbenko 98418, LE 310244; ibid., 3 km NW of Tiksi town, 71°39'N, 128°45'E, 70 m a. s. l., arctic tundra, on T. vermicularis var. vermicularis, 24 VIII 1998, Zhurbenko 98419, LE 310243.
Fig. 1. Sclerococcum zhurbenkoi (LE 310238). Habitus of the fruiting body on the thallus of Thamnolia vermicularis var. subuliformis.
Scale bar: 200 |im.
Xenonectriella physciacearum F. Berger et al.
Distinctive characters, description and illustrations. Detailed description and illustrations of this species are given in the protologue (Berger et al., 2020). Its main features are as follows: ascomata perithecioid, 240-320 |im diam., dark red
brown, with dark red to black papilla, 1/2 to 2/3 immersed; exciple K- or K+ violet, lactic acid+ yellow or negative depending on the amount of red pigments; periphy-ses up to 45 jm long; paraphysoids ramifying, highly variable in thickness, sometimes indistinct; asci unitunicate, cylindrical, (80)90-120 x 8-9 jm, 8-spored; asco-spores light brown, broadly ellipsoid, 1-septate, with ±equal cells, (10)11-13(15) x (6)7.5-9(10) jm, l/w = 1.3-1.8, with golden brown, convex tubercules 0.5-1 jm wide, uniseriate in the ascus; conidiomata of Fusarium-type; conidia hyaline, crescent-shaped, 45-65 x 3-4 jm, 4-5-septate. Reported on species of Phaeophyscia Moberg, Physcia (Schreb.) Michx. and Physconia Poelt, always associated with bleached or weakened thallus areas.
Notes. The concept of Xenonectriella leptaleae (J. Steiner) Rossman et Lowen (Rossman et al, 1999) has long been a source of incertainty in identifying specimens of Hypocreales growing on Physciaceae. These difficulties were successfully solved by Berger et al. (2020), who proposed to return to the original concept of Pronectria leptaleae (J. Steiner) Lowen (Steiner, 1900; Lowen, 1990) and also described several new species of Xenonectriella, including X. physciacearum F. Berger et al. and X. zimmer-manni F. Berger et Brackel, which fit well with many specimens previously assigned to X. leptaleae. According to Berger et al. (2020) Pronectria leptaleae, growing on species of Physcia, mainly differs from Xenonectriella physciacearum in the smaller asco-mata, 140-200 jm diam., almost exclusively confined to apothecial discs of the host lichens, consistently K- exciple, shorter asci up to 65 jm long, and hyaline, smooth or very finely verruculose, smaller ascospores, 6-11 x 4.5-7 jm. The specimens examined match well the Xenonectriella physciacearum concept, including the size of the ascospores, (8)10-14(18) x (4.5)6-8(9.5) jm, l/w = (1.2)1.3-2.1(2.9) (n = 141); they show a negative reaction of the exciple with K and lactic acid.
This species was previously known with certainty from Europe — Austria, Germany, Great Britain, Italy and Switzerland (Berger et al., 2020; Hitch, 2022) and Asia — China (Zhurbenko, 2022). The specimens listed below are the first records of Xenonectriella physciacearum for Russia from the Caucasus.
Specimens examined: Russia, Karachayevo-Circassian Republic, Teberda town, left bank of Teberda River, 43°26'41"N, 41°44'06"E, 1350 m a. s. l., freestanding deciduous tree in a pasture, on Physcia aipolia (Ehrh. ex Humb.) Furnr. (more or less damaged thallus), 27 VIII 2012, Zhurbenko 12199, LE 261334 [previously published as Xenonectriella cf. leptaleae (Zhurbenko, Kobzeva, 2014)]; Republic of Adygeya, vicinities of Mt Bol'shoy Tkhach, abandoned village Brilevo, 44°07'N, 40°20'E, 750 m a. s. l., on Physconia distorta (With.) J. R. Laundon (thallus), 26 VIII 1999, V. Otte L 452, LE 260846, herb. Otte [previously published as Xenonectriella cf. leptaleae (Zhurbenko, Otte, 2012)]; ibid, Abago pastureland, Dom Kotova cabin, 43°56'55"N, 40°12'31"E, 1770 m a. s. l., mixed forest, on P. distorta (decaying apothecia, thallus), 10 VIII 2014, Zhurbenko 14262c, LE 264415c [previously published as Xenonectriella cf. leptaleae (Zhurbenko, Kobzeva, 2016)]; ibid, on neighbouring Physcia aipolia and Physconia distorta (bleached thalli), 10 VIII 2014, Zhurbenko 14244, LE 308472 [previously published as Xenonectriella cf. leptaleae (Zhurbenko, Kobzeva, 2016)]; ibid., Lagonaki Upland, Armyanskii pass, 43°58'15"N, 39°56'30"E, 1750 m a. s. l., Fagus orientalis forest, on Physconia distorta (decaying apothecia, thallus), 12 VIII 2014, Zhurbenko 14245b, LE 264345b [previously published as Xenonectriella cf. leptaleae (Zhurbenko, Kobzeva, 2016)].
Specimen of Pronectria leptaleae examined for comparison: Russia, Pskov Region, Lok-nyanskii Region, near Bashovo village, 56°38'35"N, 30°09'02"E, on Physcia stellaris (L.) Nyl. (apo-thecia) growing on Populus tremula, 25 I 2000, E. S. Popov, LE 232160.
Note. So far this is the only confirmed specimen of Pronectria leptaleae from Russia. It was published in Popov et al. (2008: 145).
Xenonectriella cf. physciacearum F. Berger et al.
Notes. The species identification is somewhat uncertain as the material examined differs from the species protologue (Berger et al., 2020) in having longer ascospores, (12.5)13-18.5(23.5) x (6.5)8-9(9.5) ^m, l/w = 1.4-2.4(3.4) (n = 39), which are not only broadly ellipsoid, with ± equal cells, but also sometimes narrowly ovate, with a wider upper cell (Fig. 2). It should be noted that the range of ascospore size variation given in the protologue for this species is based on material from Central Europe, while this specimen was collected in the Caucasus.
Specimen examined: Russia, Caucasus, Karachayevo-Circassian Republic, Teberda town, left bank of Teberda River at 1 km upstream of Teberda Reserve office, 43°26'41"N, 41°44'06"E, 1350 m a. s. l., on freestanding deciduous tree in the pasture along the river, on Physconia distorta (bleached parts of thallus and discs of apothecia), 29 VIII 2012, Zhurbenko 12194, LE 261424 [previously published as Xenonectriella cf. leptaleae (Zhurbenko, Kobzeva, 2014)].
Fig. 2. Xenonectriella cf.physciacearum (LE 261424). Ascospores, in water. Scale bar: 10 |im.
Xenonectriella zimmermanni F. Berger et Brackel
Distinctive characters, description and illustrations. Detailed description and illustrations of this species are given in the protologue (Berger et al., 2020). Its main features are as follows: ascomata perithecioid, 140-240 |m diam., vividly orange to deep red, papilla inconspicuous, 1/2 immersed; exciple K+ violet, lactic acid+
yellow; periphyses up to 35 jm long; paraphysoids ramifying, highly variable in thickness, sometimes indistinct; asci unitunicate, cylindrical, (45)65-85 x 6-7(7.5) jm, (6)8-spored; ascospores uniseriate in the ascus, with flat tubercules ca 0.5 jm wide, 1-septate, broadly ellipsoid, with ±equal cells, hyaline to pale brown, (7)8-10(12) x 5-6(7) jm, l/w = (1.4)1.5-1.7(2) or aseptate, broadly ellipsoid to spherical, pale brown, (7)8-9.5(10) x (5.5)8-9(10) jm, l/w = (1)1.2-1.3(1.5); conidiomata of Fu-sarium-type; conidia hyaline, crescent-shaped, ca 25-30 x 2-4 jm, 3-septate. Reported on Physcia spp.
Notes. Xenonectriellaphysciacearum also occurs on Physcia spp. and can be distinguished from X. zimmermanni by its darker and larger ascomata with distinct papilla, sometimes negative reaction of the exciple with K, larger asci, and larger, always septate ascospores with smaller tubercules. Only hyaline, smooth (probably immature) ascospores were observed, but otherwise the specimen examined corresponds well to the species protologue, including the reaction of the exciple with lactic acid and K, and the size of the 1-septate ascospores, (8)8.5-10.5(11.5) x (5)5.5-6 jm, l/w = (1.4)1.5-1.9(2.2) (n = 12, in lactic acid).
This species was previously known with certainty from Europe (Austria, Germany, Switzerland) and Bermuda (Berger et al., 2020). It is reported here for the first time for Russia from the Caucasus.
Specimen examined: Russia, Krasnodar Territory, Lagonaki Upland, foot of Mt Fisht at the headwaters of Belaya River, 43°57'34"N, 39°55'48"E, 1580 m a. s. l., mixed forest, on Physcia stellaris (bleached thallus), 18 VIII 2014, P. M. Zhurbenko, LE 308473 [previously published as Xenonectriella cf. leptaleae (Zhurbenko, Kobzeva, 2016)].
Acknowledgments
This study 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" (122011900033-4) using the equipment of its Core Facility Center 'Cell and Molecular Technologies in Plant Science'. Franz Berger and Wolfgang von Brackel are thanked for their comments on the Xenonectriella species.
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