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172 Herzogia 31 (1) Teil 1, 2018: 172–189 Noteworthy lichenized and lichenicolous fungi of open-canopy oak stands in east-central Europe Paweł Czarnota * , Helmut Mayrhofer & Andrzej Bobiec Abstract: Czarnota, P., Mayrhofer, H. & Bobiec, A. 2018. Noteworthy lichenized and lichenicolous fungi of open-canopy oak stands in east-central Europe. – Herzogia 31: 172–189. Eighteen species of lichenized and two species of lichenicolous fungi from oak stands of east-central Europe’s rural landscapes are presented. Rinodina isidioides is new to eastern Europe and the Carpathians, Abrothallus microsper- mus is new to the Carpathians, Rinodina excrescens is new to the eastern Carpathians, Biatora pontica is new to the Polish Carpathians, Lecanora substerilis, Ramonia chrysophaea and Verrucaria viridigrana are new to the Polish Carpathians and Poland, and others are new to Hungary, Romania or Ukraine. Distinguishing characters are empha- sised and notes on ecology as well as global and Carpathian distributions are given. Open-canopy oak stands and solitary oaks in wood-pastures are important for the protection of the lichen diversity of the Carpathians and need the special attention of conservation authorities and local stakeholders. Zusammenfassung: Czarnota, P., Mayrhofer, H. & Bobiec, A. 2018. Bemerkenswerte lichenisierte und licheni- cole Pilze von lückigen Eichenbeständen in Zentral-Osteuropa. – Herzogia 31: 172–189. Achtzehn Flechtenarten und zwei lichenicole Pilze werden aus Eichenbeständen der zentralosteuropäischen ländli- chen Landschaft präsentiert. Rinodina isidioides ist neu für Osteuropa und die Karpaten, Abrothallus microspermus ist neu für die Karpaten, Rinodina excrescens ist neu für die östlichen Karpaten, Biatora pontica ist neu für die pol- nischen und östlichen Karpaten, Lecanora substerilis, Ramonia chrysophaea und Verrucaria viridigrana sind neu für die polnischen Karpaten und Polen, während andere Arten für Ungarn, Rumänien oder die Ukraine neu sind. Die Unterscheidungsmerkmale werden hervorgehoben und Notizen zur Ökologie und Verbreitung gemacht. Lückige Eichenbestände und freistehende Eichen der Waldweiden sind wichtig für die Erhaltung der Flechtendiversität der Karpaten und bedürfen besonderer Beachtung durch Naturschutzbehörden und lokaler Interessensgruppen. Key words: Carpathians, epiphytic lichens, lichen diversity, oak wood-pastures, rural landscape. Introduction Lasting for hundreds of years, traditional use of land, involving livestock grazing, in par- ticular cattle and sheep, was a driving factor responsible for the development of semi-natural wooded ecosystems throughout Europe. Although some cultures focused on grain production and used forested areas for grazing and hay-making, other cultures excelled in developing ex- tensive free-range wood-pastures (Rackham 2006, Hartel & Plieninger 2014, Hartel et al. 2015). Common to both systems are open woods or groves, dominated by oaks with wide, low set crowns and substantial parts of boughs and stems exposed to sunlight. In Central and Eastern Europe, legacies of such traditional management have been relatively well preserved in the outskirts of Carpathians, either as marginal oak-dominated former silvopastoral groves * Corresponding author

Transcript of Noteworthy lichenized and lichenicolous fungi of open ... · Noteworthy lichenized and...

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Noteworthy lichenized and lichenicolous fungi of open-canopy oak stands in east-central Europe

Paweł Czarnota*, Helmut Mayrhofer & Andrzej Bobiec

Abstract: Czarnota, P., Mayrhofer, H. & Bobiec, A. 2018. Noteworthy lichenized and lichenicolous fungi of open-canopy oak stands in east-central Europe. – Herzogia 31: 172–189. Eighteen species of lichenized and two species of lichenicolous fungi from oak stands of east-central Europe’s rural landscapes are presented. Rinodina isidioides is new to eastern Europe and the Carpathians, Abrothallus microsper-mus is new to the Carpathians, Rinodina excrescens is new to the eastern Carpathians, Biatora pontica is new to the Polish Carpathians, Lecanora substerilis, Ramonia chrysophaea and Verrucaria viridigrana are new to the Polish Carpathians and Poland, and others are new to Hungary, Romania or Ukraine. Distinguishing characters are empha-sised and notes on ecology as well as global and Carpathian distributions are given. Open-canopy oak stands and solitary oaks in wood-pastures are important for the protection of the lichen diversity of the Carpathians and need the special attention of conservation authorities and local stakeholders.

Zusammenfassung: Czarnota, P., Mayrhofer, H. & Bobiec, A. 2018. Bemerkenswerte lichenisierte und licheni-cole Pilze von lückigen Eichenbeständen in Zentral-Osteuropa. – Herzogia 31: 172–189.Achtzehn Flechtenarten und zwei lichenicole Pilze werden aus Eichenbeständen der zentralosteuropäischen ländli-chen Landschaft präsentiert. Rinodina isidioides ist neu für Osteuropa und die Karpaten, Abrothallus microspermus ist neu für die Karpaten, Rinodina excrescens ist neu für die östlichen Karpaten, Biatora pontica ist neu für die pol-nischen und östlichen Karpaten, Lecanora substerilis, Ramonia chrysophaea und Verrucaria viridigrana sind neu für die polnischen Karpaten und Polen, während andere Arten für Ungarn, Rumänien oder die Ukraine neu sind. Die Unterscheidungsmerkmale werden hervorgehoben und Notizen zur Ökologie und Verbreitung gemacht. Lückige Eichenbestände und freistehende Eichen der Waldweiden sind wichtig für die Erhaltung der Flechtendiversität der Karpaten und bedürfen besonderer Beachtung durch Naturschutzbehörden und lokaler Interessensgruppen.

Key words: Carpathians, epiphytic lichens, lichen diversity, oak wood-pastures, rural landscape.

IntroductionLasting for hundreds of years, traditional use of land, involving livestock grazing, in par-ticular cattle and sheep, was a driving factor responsible for the development of semi-natural wooded ecosystems throughout Europe. Although some cultures focused on grain production and used forested areas for grazing and hay-making, other cultures excelled in developing ex-tensive free-range wood-pastures (Rackham 2006, Hartel & Plieninger 2014, Hartel et al. 2015). Common to both systems are open woods or groves, dominated by oaks with wide, low set crowns and substantial parts of boughs and stems exposed to sunlight. In Central and Eastern Europe, legacies of such traditional management have been relatively well preserved in the outskirts of Carpathians, either as marginal oak-dominated former silvopastoral groves

* Corresponding author

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or still functioning wood-pastures, such as in Transylvania, (Fischer et al. 2012, Hartel et al. 2013, Hanspach et al. 2014). As a conservation measure, this system is being restored on the local scale in Carpathian Basin in Hungary (Varga & Bölöni 2009). Except for their socio-economic importance, wood-pastures are valuable refuges of rich biodiversity due to the com-bination of grasslands and woody plants, including ancient trees (e.g. Bergmeier et al. 2010, Horák & Rébl 2012, Hartel et al. 2014). Traditional rural landscapes with groves and soli-tary veteran trees may be considered as a contemporary analogy of an ancient “savannah” – an important part of “primeval” temperate European landscapes (Vera 2000). In various biogeo-graphical regions of Europe, such trees are very often various species of oaks, Quercus spp., of which longevity, size and shape are conducive to the development of microhabitats used by lichens. Depending on whether a tree layer has a form of dispersed solitary trees, open-canopy woods (grown in the past for pannage, Szabó 2013), or self-established copses filled-in with dense undergrowth of shrubs and younger trees, the corresponding epiphytic communities vary accordingly, particularly with respect to the composition of nitrophilous, heliophilous and forest lichens (e.g. Jönsson et al. 2011, Leppik et al. 2011). Among various interesting species, oaks also harbour national or regional red-listed species, as well as species recognized as relics of ancient forests (e.g. Thor et al. 2010).Whilst the lichenological importance of wood-pastures and open-canopy groves in Western Europe and Scandinavia have long been acknowledged (e.g. Paltto et al. 2011, Leppik et al. 2011), such habitats have been of minor lichenological interest in the central-eastern part of the continent. Our study, being a part of the larger research project on oak woods in rural landscapes of the Carpathians, aimed to fill this gap of knowledge by reference to cultural landscapes of the Carpathian foothills. This paper presents selected results of the inventory, confined to the species, which so far have not been reported from at least one of the four Carpathian countries in which sample plots were investigated.

Material and MethodsCollections were made in oak groves and wood-pastures located at low altitudes of the Eastern or Inner Western Carpathians in four countries: Poland, Ukraine, Romania and Hungary (Fig. 1.). Most of the Polish (PL) studied stands represent abandoned silvopastoral oak groves (me-dian oaks age: 110 years), with dense hazel (Corylus avellana) undergrowth. The Ukrainian (UA) stands are younger (77 years) and are, except one with hazel, more open due to frequent grass burning (Ziobro et al. 2016). Both Hungarian (HU) and Romanian (Transylvanian, RO) study areas represent actual functioning wood-pasture systems, a combination of grasslands with semi-open, park-like groves, solitary trees and abandoned groves with dense undergrowth (median oaks age: 129 years in HU and 107 years in RO – A. Bobiec, unpublished). Certain records are also from neighbouring high forest oak stands, subject to conventional forest man-agement. Only epiphytes of Quercus spp. were collected and examined.The nomenclature of lichen-forming species mostly follows Wirth et al. (2013), with the exception of Agonimia borysthenica (Dymytrova et al. 2011), Biatora pontica (Printzen & Tønsberg 2003), Fuscidea arboricola, Lecidea nylanderi (Tønsberg 1992), Psoroglaena dictyospora (Vondrák et al. 2010), Rinodina isidioides (Giavarini et al. 2009), Rinodina subpariata (Resl et al. 2016, cf. incorrect spelling as R. subparieta) and Verrucaria viridi-grana (Breuss 1998). The nomenclature of lichenicolous fungi, marked with an asterisk (*), follows Czyżewska & Kukwa (2009). Specimens are deposited in the Gorce National Park Herbarium, Poland (GPN) and some duplicates of Rinodina are in the University of Graz

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Herbarium, Austria (GZU). For morphological characters a Zeiss StemiDV4 stereomicroscope was used. Hand-made apothecial or pycnidial sections mounted in water, KOH and HNO3 were analysed using a Zeiss Axiostar plus microscope. Secondary metabolites of sterile lichen species were examined by thin-layer chromatography (TLC) in the solvent system C according to Orange et al. (2001). Aluminum plates were used.

Fig. 1. Geographic location of four study areas: PL – Poland, UA – Ukraine, HU – Hungary, RO – Romania.

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Species list*Abrothallus bertianus De Not.

Hungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96833°N/20.55889°E, alt. c. 300 m, bark of Quercus cerris in oak wood-pasture, 16.05.2015, P. Czarnota 8217 & 8255; ibid., 47.96861°N/20.55694°E, 14.05.2015, P. Czarnota 8224.Host: Melanohalea elegantula.New to Hungary.This is a worldwide distributed species reported mainly from Europe: Great Britain, Ireland Denmark, Sweden, Finland, Lithuania, Estonia, European Russia, the Netherlands, Luxembourg, France, Germany, Poland, Switzerland, Austria, Czech Republic, Slovenia, Spain, Italy, Bosnia and Herzegovina (Motiejūnaitė 1999, Kocourková 2000 and literature cited therein, van den Boom & Clerc 2000, Scholz 2000, Llimona & Hladun 2001, Aptroot et al. 2008, Brackel 2009, 2015, Czyżewska & Kukwa 2009, Seaward 2010, Bilovitz et al. 2010, Zhurbenko & Tugi 2013, Diederich et al. 2016), Ukraine: Podilia (Pirogov & Shovhan 2015), Crimea (Khodosovtsev et al. 2013), as well as from North and South America (Triebel et al. 1991, Esslinger 2016), Asia: Turkey (John & Breuss 2004) and Russian Far East (Zhurbenko & Tugi 2013). A. bertianus was previously reported from the Polish part of Western Carpathians (Alstrup & Olech 1996, Kukwa & Flakus 2009, Bielczyk et al. 2016) and from the Ukrainian Eastern Carpathians (Kondratyuk et al. 2003 without certain locality, Malíček et al. 2018). It has also been reported from Hungary, but according to the latest checklist by Lőkös & Farkas (2009), this record is in fact A. caerulescens Kotte; therefore the records here for Hungary extend the European range of A. bertianus.

*Abrothallus microspermus Tul.Romania: Eastern Carpathians, Transylvania, Oklánd county, 3 km S of Homoródújfalu village, 46.13146°N/25.41587°E, alt. c. 545 m, on solitary old Quercus robur in existing oak wood-pasture, 18.05.2015, P. Czarnota 7863; ibid., 46.13125°N/25.41579°E, P. Czarnota 7866; ibid., 46.12800°N/25.41460°E, alt. 560 m, P. Czarnota 8095.Host: Flavoparmelia caperata.New to Romania and the Carpathians.It is known from Belgium, France and Luxembourg (Diederich et al. 2016), Italy (Brackel 2015) Austria (Wittmann & Türk 1989), Germany (Scholz 2000), Poland (Czyżewska & Kukwa 2009), Belarus (Tsurykau 2017), Hungary, Ireland (Seaward 2010), Spain (Longán & Gómez-Bolea 1999), Switzerland, Great Britain, Southern Ural Mts (Urbanavichene et al. 2013), North America (e.g. Cole & Hawksworth 2001, Diederich 2003, Kocourková et al. 2012), Asia: North-East Caucasus (Urbanavichus & Ismailov 2013), South Korea (Kondratyuk et al. 2013) and from New Zealand (Longán & Gómez-Bolea 1999).

Agonimia borysthenica Dymytrova, Breuss & S.Y.Kondr.Ukraine: Ivano-Frankivsk region, 2 km S of Rakiv village, 48.97720°N/24.12312°E, alt. 430 m, on bark of Quercus petraea in forested wood-pasture close to the edge of forest, 11.06.2015, P. Czarnota 8209 & 8211.A description, illustrated habitus and comparison to other most similar species of the genus Agonimia are presented in details by Dymytrova et al. (2011) and Dymytrova (2013) based on Ukrainian collections. It is also morphologically similar to Verrucaria viridigrana Breuss but differs in muriform large ascospores while in V. viridigrana they are simple.This recently described species was previously collected several times in European, wet, old-growth deciduous forests in Ukraine (Kyiv Region: Dymytrova et al. 2011, Dymytrova 2013, Ukrainian Carpathians: Malíček et al. 2018), Switzerland (Kanton of Zug, Kanton of Ticino: Dymytrova & Scheidegger 2012) and the Czech Republic (Cahnov-Soutok National Nature Reserve: Vondrák et al. 2016). Our locality at the foot of Ukrainian Carpathians is situated at the edge of an oak forest which was previously a wood-pasture, and except for sparsely distributed old oaks, this area is partly kept open through spring grass burning, partly overgrown by hazel undergrowth. This shows that this species may have a greater ecological plasticity.

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Anisomeridium polypori (Ellis & Everh.) M.E.BarrHungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96806°N/20.56139°E, alt. 300 m, on bark of Quercus cerris in oak wood-pasture, 16.05.2015, P. Czarnota 8245. Romania: Eastern Carpathians, Transylvania, Oklánd county, c. 3 km S of Homoródújfalu village, 46.12800°N/25.41460°E, alt. 560 m, on bark of Quercus sp. in semi-forested former wood-pasture close to the edge of forest, 19.05.2015, P. Czarnota 7840, 7844 & 7845.

New to Hungary and Romania.This worldwide distributed species, found in Europe, North & Central America, Africa, Asia, Australia and Oceania, has been described several times under different names from different regions of the world both as a lichen-forming species as well as a lichenicolous fungus (Kashiwadani & Thor 1995, Aptroot 1999, Etayo & van den Boom 2006, Yazici & Aptroot 2008). Here it is reported as new to Hungary and Romania (not recorded in the checklists of Lőkös & Farkas 2009 and Ciurcea 2007 and later contributions to both countries). The genus Anisomeridium in both countries is repre-sented by A. biforme. Perhaps some collections of this species belong to A. polypori which is currently widespread in Europe and common in Western Carpathians (e.g., Bielczyk et al. 2016, Adamčík et al. 2016) and also reported in Polish, Slovak and Ukrainian Eastern Carpathians (e.g., Kondratyuk & Coppins 2000, Pišút et al. 2007, Kościelniak 2013, Vondrák et al. 2015, Adamčík et al. 2016, Malíček et al. 2018).

Bacidina sulphurella (Samp.) Hauck & V.WirthUkraine: Eastern Carpathians, Ivano-Frankivsk region, Rozhniativ county, c. 1.5 km W of Ivanivka village, 48.88606°N/24.09161°E, alt. 490 m, on bark of Quercus sp. in young oak-hornbeam forest, a former wood-pasture, 7.06.2015, P. Czarnota 8151.

The species is common in Europe mainly in woodlands; it has also been frequently recorded in old-growth forests in the East Carpathians Biosphere Reserve, including its Ukrainian part, as Bacidina arnoldiana (Körb.) V.Wirth & Vězda (Kondratyuk & Coppins 2000, Kościelniak 2013; see also Vondrák et al. 2015). Only recently Malíček et al. (2018) have reported B. sulphurella from this Carpathian region based on a species concept adopted by Brand et al. (2009) and Czarnota & Guzow-Krzemińska (2012). The latest discovery is new to the Ukraine and extends the distribution of B. sulphurella in the Eastern Carpathians.

Biatora pontica Printzen & TønsbergUkraine: Eastern Carpathians, Ivano-Frankivsk region, 2 km SW of Rozhniativ town, 48.92103°N/24.12278°E, alt. 430 m, on bark of Quercus sp. in forested former wood-pasture at the edge of forest, 9.06.2015, P. Czarnota 8143 & 8147; ibid., Rozhniativ county, 1.5 km W of Ivanivka village, 48.88672°N/24.09631°E, alt. 490 m, on bark of Quercus sp. in forested former wood-pasture, 8.06.2015, P. Czarnota 8166; ibid., Ivano-Frankivsk region, 2 km S of Rakiv village, 48.97683°N/24.12208°E, alt. c. 430 m, on bark of Quercus sp. in semi-forested wood-pasture at the edge of broadleaved forest, P. Czarnota 8193 & 8206; ibid., 48.97720°N/24.12312°E, alt. c. 430 m, on bark of Quercus sp. in hardly forested former wood-pasture at the edge of broadleaved forest, 11.06.2015, P. Czarnota 8208. Additional records. Poland: Atpol Ge-09, Eastern Carpathians, Góry Sanocko-Turczańskie Mts, c. 2 km NE of Huwniki village, 49.65819°N/22.73157°E, on bark of Quercus sp. inside young oak-beech forest, alt. c. 360 m, 19.07.2015, P. Czarnota 7930 & 7931; ibid., c. 1.3 km S of Pacław village, 49.61163°N/22.70721°E, alt. c. 470 m, on bark of Quercus sp. at the edge of deciduous forest, 24.04.2015, P. Czarnota 7823.

Chemistry: thiophanic acid, asemone and ‘pontica unknown’ (identified by M. Kukwa).New to Polish Carpathians.Biatora pontica in its sterile stage resembles Lecidella subviridis which occupies similar substrates and woodland habitats and also produces a C+ orange xanthone-containing sorediate thallus. The tin-ge of B. pontica thallus is usually a little bit more greenish since it grows in shady forests. However, it is just possible, based on their chemistry, to distinguish the two species since B. pontica produces asemone and ‘pontica unknown’ substances, while L. subviridis contains atranorin, arthothelin and ‘expallens unknown’ metabolites (both species contain thiophanic acid). For detailed descriptions and similarities to such species as Biatora chrysantha, B. efflorescens, B. epixanthoides and Phyrrospora

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quernea, when they are fertile or only sterile, see Printzen & Tønsberg (2003), Kukwa et al. (2012) and Malíček et al. (2014). Additionally, small patches of B. pontica can also be confused with Ropalospora viridis, Fuscidea arboricola or F. pusilla which in central Europe often grow together on the same trunks of broadleaved trees; however, these associate, usually sterile species have a C– reaction.Despite the limited data on the ecology and habitat preferences of B. pontica, this species is inclu-ded in the list of indicators of the coniferous-broadleaved forest zone in Central European Russia (Muchnik 2015). Undoubtedly, B. pontica represents the group of forest lichens and perhaps even prefers old-growth forests (as many its records were confined to those conditions), but in the Eastern Carpathians it is also found in the margins of broadleaved forests, including those which were for-merly used as wooded pastures. Today such areas form loose stands composed of sparsely distributed old trees (mainly Quercus robur) and often dense saplings of many deciduous tree and shrub species. However, B. pontica seems to avoid overly shaded microhabitats.Biatora pontica is a widespread though rarely reported forest species, known from Europe: Norway, Austria (Printzen & Tønsberg 2003, Hafellner et al. 2008), Italy (Tretiach 2004), Slovenia (Mayrhofer & Coppins 2013), Russia: Caucasus (Urbanavichus 2010, Urbanavichene & Urbanavichus 2014) and S Ural Mts (Urbanavichene et al. 2013), Asia: Turkey and North America: USA, North Carolina (Printzen & Tønsberg 2003). It has recently been reported several times from lowlands in northern Poland (e.g. Kukwa et al. 2012, Kowalewska & Kukwa 2013, Kubiak et al. 2014), and from the Czech Republic (Vondrák et al. 2016), the Slovak Western Carpathians (Malíček et al. 2014) and the Ukrainian Carpathians (Malíček et al. 2018).

Fuscidea arboricola Coppins & TønsbergHungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96806°N/20.56139°E, alt. 300 m, on bark of Quercus cerris in oak wood-pasture, 16.05.2017, P. Czarnota 8247 & 8258; ibid., 47.96730° N, 20.55011° E, alt. 290 m, on bark of Quercus sp. in a young oak forest, 21.05.2015, P. Czarnota 7895 & 7913 (det. M. Kukwa by TLC ).

Chemistry: fumarprotocetraric acid, protocetraric acid (trace). New to Hungary. Second report from the Western Carpathians.A sorediate green lichen with a slight brown tinge, found usually in its sterile, soraliate stage and thus recognizable only by a detection of fumarprotocetraric acid (P+ ginger), is similar in morphology and the P+ reaction to Violella fucata which produces, however, atranorin as an additional secondary sub-stance and forms a thallus with a greyish tinge.This forest lichen known from many countries in northern and central Europe (e.g. Fałtynowicz 2003, Coppins et al. 2005, Motiejūnaitė 2011, Malíček & Palice 2013) has recently been reported from Poland as new to the Western Carpathians (Bielczyk et al. 2016) and moreover collected from trunks of various deciduous trees from several localities in the Eastern Carpathians (Kondratyuk & Coppins 2000, Coppins et al. 2005, Ardelean et al. 2013, Malíček et al. 2018). According to Fryday (2008) it is frequent in North America.

Fuscidea pusilla TønsbergHungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96806°N/20.56139°E, alt. 300 m, on bark of Quercus cerris in oak wood-pasture, 16.05.2017, P. Czarnota 8260; ibid., 47.96833°N/20.55889°E, P. Czarnota 8242; ibid., 47.96730°N/20.55011°E, alt. 290 m, on bark of Quercus sp. in young oak forest, 23.05.2015, P. Czarnota 7914 & 8243 (det. M. Kukwa by TLC).

Chemistry: divaricatic acid (Czarnota 7914 & 8243) or divaricatic and nordivaricatic (trace) acids (Czarnota 8243 & 8260).New to Hungary.This sorediate, whitish-green, crustose species resembles in morphology several common lichens with similar ecology in Europe including Fuscidea arboricola, F. lightfootii and Ropalospora viridis. F. arboricola can be easily separated by its P+ ginger spot test reaction (due to fumarprotocetraric acid), but the distinction of F. pusilla and Ropalospora viridis, both with negative spot tests, needs to be

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supported by TLC data to differentiate divaricatic and perlatolic acids. For further details including the note on Fuscidea lightfootii see, for example, Czarnota & Węgrzyn (2012).Fuscidea pusilla is widely distributed in continental Europe (e.g., Coppins et al. 2005) including whole Polish and Slovak Carpathians (e.g., Czarnota & Węgrzyn 2012, Guttová et al. 2012, Kościelniak 2013, Vondrák et al. 2015, Bielczyk et al. 2016), Eastern Carpathians in Ukraine (Coppins et al. 2005) and Romania: Rodnei Mts (Ardelean et al. 2013). Moreover the species has been reported from the Pacific and Atlantic sides of North America (e.g. Fryday 2008, Lendemer 2011).

Lecanora substerilis Malíček & Vondrák Hungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96730°N/20.55011°E, alt. c. 290 m, on bark of Quercus sp. within a young dense oak monoculture close to oak wood-pasture, 23.05.2015, P. Czarnota 7909. Additional records. Poland: Western Carpathians, Western Beskidy Mts, Gorce Mts, Gorce National Park, valley of Kamienica river below Mostownica Mt, Zapadłe area, 49.55231°N/20.13639°E, alt. 1110 m, on bark of Fagus sylvatica in the Carpathian beech forest, 31.07.2015, P. Czarnota 7978, L. Widak & K. Wąsik; ibid., valley of Roztoka stream below former Szyja Turbacza glade, 49.55610°N/20.11561°E, alt. 1180 m, on bark of Fagus sylva-tica in the Carpathian beech forest, 30.07.2015, P. Czarnota 7980, L. Widak & K. Wąsik; ibid., valley of Kamienica river below Bieniowe glade, 49.55461°N/20.21640°E, alt. 1110 m, on bark of Fagus sylvatica in the Carpathian beech forest, 29.07.2015, P. Czarnota 8048 & 8054, L. Widak & K. Wąsik; ibid., valley of Kamienica river below Spaleniec glade, 49.57827°N/20.23027°E, alt. 970 m, on bark of Fagus sylvatica in the Carpathian beech forest, 27.07.2015, P. Czarnota 8086, L. Widak & K. Wąsik.

Chemistry: atranorin, roccellic acid.New to Hungary and Poland.Specimens mentioned above have previously been recognized as a sterile stage of Lecanora farinaria Borrer, but following a recent description of L. substerilis (Malíček et al. 2017), supported by mole-cular evidence, there are no doubt that they represent this new species. The main differences between both species are in their thallus morphologies and distribution ranges: thallus endosubstratal to thin in the oceanic L. farinaria vs. areolate to pustulate or thin, but at least locally pustulate in the continental L. substerilis (Malíček et al. 2017). The species differ also in chemical characters. Lecanora subste-rilis contains atranorin, roccellic acid and some unidentified fatty acids (sometimes present in traces), while Tønsberg (1992) also detected chloratranorin in L. farinacea. All examined Hungarian and Polish specimens form small patches of ± thick thalli which are often continuous or cracked in non-sorediate parts and usually develop convex, whitish to greyish-white, separate soralia. In such forms the species can be mistaken in the field with, for example, Pertusaria pupillaris, Loxospora elatina or Ochrolechia arborea; each of these species differ distinctly from L. substerilis in their chemistry and simple spot test reactions, since they all have no atranorin and P. pupillaris reacts P+ ginger (presence of fumarprotocetraric acid, L. elatina P+ orange (presence of thamnolic acid), and O. arborea C+ red (presence of gyrophoric and lecanoric acids).The distribution of L. substerilis is poorly known since the species was only described very recently. Malíček et al. (2017) reported it as an old-growth beech forest lichen from several Carpathian coun-tries: the Czech Republic, Slovakia, Ukraine and Romania. The Polish records presented here confirm its ecological preferences as growing in natural, well-preserved forests in the lower mountain belt, but the Hungarian discovery shows that it is perhaps more ecologically tolerant.

Lecidea nylanderi (Anzi) Th.Fr.Hungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96730°N/20.55011°E, alt. 290 m, on bark of Quercus sp. in young oak forest, 21.05.2015, P. Czarnota 7898.

Chemistry: divaricatic acid.New to Hungary.This usually sterile, sorediate species resembles thin, leprose crusts of some representatives of the ge-nus Lepraria, mostly L. incana and L. elobata because of the bluish tinge of their thalli. Both Lecidea

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nylanderi and Lepraria incana produce divaricatic acid, but zeorin is also found as the main com-pound in the latter (Tønsberg 1992). L. elobata has different chemistry, including stictic acid complex (Tønsberg 1992), which reacts P+ orange.Lecidea nylanderi is a common but not so frequently reported species in the Western Carpathians, mostly confined to acid bark or dead wood of Picea abies in upper mountain spruce forests (Czarnota 2012); also recently found in Romanian Eastern and Southern Carpathians (Vondrák & Liška 2013, Ardelean et al. 2013, Malíček et al. 2015). Moreover in Europe it is widespread from Fennoscandia to the Mediterranean region (e.g., Dietrich & Scheidegger 1996, Llimona & Hladun 2001, Printzen et al. 2002, Marmor et al. 2011, Motiejūnaitė 2011), and recorded also in North America (Lendemer 2006) and Asia: Turkey (Breuss & John 2004), Mongolia (Hauck & Javkhlan 2006).

Lecidella flavosorediata (Vězda) Hertel & LeuckertRomania: Eastern Carpathians, Transylvania, Oklánd county, 3 km S of Homoródújfalu village, 46.13125°N/25.41579°E, alt. 545 m, on bark of Quercus robur in oak wood-pasture, 18.05.2015, P. Czarnota 7867; ibid., 1 km W of Homoródújfalu village, 46.17667°N/25.41306°E, on bark of Quercus sp. in oak wood-pasture, 21.05.2015, P. Czarnota 7879 & 7881 (det. M. Kukwa); both in a sterile stage.

Chemistry: arthothelin, granulosin.New to Romania.This usually sterile, sorediate species is widely distributed in Europe, but in the Carpathians it is ra-rely recorded, perhaps due to its morphological similarity to the more common, yellowish-coloured Lecidella subviridis Tønsberg or Lecanora expallens Ach. Sterile examples need to be checked by TLC to find above mentioned composition of secondary substances (for further details see for ex-ample Zduńczyk & Kukwa 2014). It was collected in Polish Western Carpathians (e.g. Bielczyk 2003, Czarnota 2010, Śliwa & Kukwa 2012) and recently also in Eastern Carpathians in Slovakia, Ukraine (e.g. Vondrák et al. 2015, Malíček et al. 2018) and Poland (Kościelniak 2013).

Lepraria rigidula (de Lesd.) TønsbergHungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96806°N/20.56139°E, alt. 300 m, on bark of Quercus cerris in oak wood-pasture, 16.05.2015, P. Czarnota 8250; ibid., 47.96730°N/20.55011°E, alt. 290 m, on bark of Quercus sp. in young oak forest, 21.05.2015, P. Czarnota 7893.

Chemistry: atranorin, nephrosteranic acid (rigidula unknown).New to Hungary.This European Lepraria species is morphologically recognizable by the presence of long, projecting hyphae and usually lack of visible standard spot test reactions. The K+ yellow reaction mentioned in some literature (e.g. Wirth et al. 2013) due to atranorin is usually not observed.Lepraria rigidula is common and widespread throughout the world, reported from almost all conti-nents (Orange & Laundon 2009).

Protoparmelia hypotremella Herk, Spier & V.WirthHungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96806°N/20.56139°E, alt. 300 m, on bark of Quercus cerris in oak wood-pasture, 16.05.2015, P. Czarnota 8259, 8261 & 8263.

New to Hungary.According to Aptroot et al. (1997), Kukwa (2000) and Palice et al. (2006), P. hypotremella is simi-lar to several species forming subsquamulose to isidia-like thalli, namely Hypocenomyce caradocen-sis, Bacidia rubella and especially Protoparmelia oleagina. Rinodina excrescens is also considered to be very similar to P. hypotremella. Both species can grow in the same habitats in warmer regions of Europe, but R. excrescens contains P+ red pannarin. The known worldwide distribution of P. hypotremella includes Europe: the Netherlands, Belgium, France, Switzerland, Germany, Austria, Sweden (Kubiak et al. 2010 and literature cited therein), the Czech Republic (Malíček 2008), Poland (Kukwa 2000, Kubiak et al. 2010), European Russia:

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Leningrad region (Himelbrant et al. 2014) and North America (Brodo & Aptroot 2005). In the Carpathians, P. hypotremella was recorded in Slovakia: Muranska planina (Palice et al. 2006) and Poland (Śliwa 2006, Czarnota & Wojnarowicz 2008, Kościelniak 2013).

Psoroglaena dictyospora (Orange) H.HaradaRomania: Eastern Carpathians, Transylvania, Oklánd county, 3 km S of Homoródújfalu village, 46.131°N/25.415°E, alt. 580 m, on bark of Quercus sp. in oak shady forest close to the wood-pasture, 20.05.2015, P. Czarnota 7872.

New to Romania.Psoroglaena dictyospora is an epiphytic lichen on various organic substrata, e.g. bark of deciduous trees or plant debris (e.g. Dietrich 1991, Orange 1991, Longán & Gómez-Bolea 1998, Printzen et al. 2002). Printzen et al. (2002) suggested that although ecologicaly variable, P. dictyospora pre-fers undisturbed habitats, but it was also recorded from highly polluted and metal-reach area in Polish Upper Silesia (Kiszka & Kościelniak 2006 as Leucocarpia biatorella, Czarnota unpubl.) and ur-banized area of Slovak Bratislava (Palice 1999). Here it is reported from the edge of a former grazed oak woodland; after abandonment dense hornbeam (Carpinus betulus) has developed, over-shadowing the oak trunks.It is widespread in Europe, known from Spain (Longan & Gomez-Bolea 1998), Austria (Hafellner & Türk 2016), Switzerland (Dietrich 1991), Germany, Finland (Printzen et al. 2002), Sweden (Orange 1991, Santesson 1993), Czech Republic (Svoboda et al. 2014), Poland (Kubiak 2013), Slovakia and Ukraine (Palice 1999, Vondrak et al. 2010).

Ramonia chrysophaea (Pers.) VězdaPoland: Atpol Gf– 09, Eastern Carpathians, Góry Sanocko-Turczańskie Mts, c. 0.5 km W of Kalwaria Pacławska village, 49.62980°N/22.69901°E, on bark of Quercus robur in a deciduous forest, 18.07.2015, P. Czarnota 7948.

New to Poland and Polish Carpathians; the third record for the Eastern Carpathians.The species was found mainly in wet, deciduous forests and for that reason is sometimes regarded as an old-growth forest indicator (Larsen & Søchting 2006, Vondrák et al. 2015) or species of major importance for biodiversity conservation (Woods & Coppins 2012). Most of its known, but realtively rare, records come from western part of Europe, i.e. Great Britain, Ireland (Seaward 2010), France, Spain (Coppins 1987, Sanderson & Purvis 2009, Coste 2014), Portugal (Jones 2002), Luxembourg and Germany (Hessen; Eichler et al. 2010), Denmark (Larsen & Søchting 2006), Italy (Gambera & Tretiach 2003, Tretiach 2004) and from Fennoscandia (Coppins et al. 1994, Thor & Hultengren 2001, Pykälä 2006) suggesting that R. chrysophaea prefers an Atlantic cli-mate; a Mediterranean-Atlantic species according to Nimis & Martellos (2017) and Sanderson & Purvis (2009). Recently it has been found in Central Europe, in the Czech, Slovak and Ukrainian parts of the Carpathians (Kondratyuk & Coppins 2000, Coppins et al. 2005, Svoboda et al. 2007, Vondrák et al. 2015) and also in Southern Ural Mts (Urbanavichene et al. 2013).

Rinodina excrescens Vain. Ukraine: Eastern Carpathians, Ivano-Frankivsk region, Rozhniativ county, 1.5 km W of Ivanivka village, 48.88672°N/24.09631°E, alt. c. 490 m, 08.06.2015, P. Czarnota 8124; ibid., 2.5 km SW of Rozhniativ town, 48.91917°N/24.11389°E, alt. 430 m, 6.06.2015, P. Czarnota 8183 (fertile).

New to the Eastern Carpathians and Ukraine.The above collections were found in a foothill of the Ukrainian Eastern Carpathians growing on seve-ral well-lit, as well as more shaded, 80 –100-years old oak trees forming open woods in outer parts of broadleaved forests, formerly used as wood-pastures.Galanina et al. (2011) discussing the worldwide occurrence of R. excrescens concluded that this species seems to be the rarest Rinodina member in Europe due to only few reports having been made at that time. This is a disjunctive species recorded in three continents with main distribution centres in the Russian Far East and Japan (Galanina et al. 2011, Sheard et al. 2017) and the North American region of Great Lakes in Canada and USA (Sheard 2010). In Europe R. excrescens has been reported from Austria (Giralt et al. 1994), Mediterranean region: Croatian island of Mljet (Giralt et al. 1995),

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Crete (Spribille et al. 2006), Spain (Aragón et al. 2004) and more recently several times from the Czech Republic (Malíček 2013, Malíček & Palice 2013).

Rinodina isidioides (Borrer) H.OlivierRomania: Eastern Carpathians, Transylvania, Oklánd county, 3 km S of Homoródújfalu village, 46.12800°N/25.41460°E, alt. 560 m, on bark of Quercus sp. in outer part of deciduous forest close to the oak wood-pasture, 19.05.2015, P. Czarnota 7846 (GPN; dupl. in GZU).

New to Eastern Europe, Carpathians and Romania.According to Nimis & Martellos (2017) R. isidioides is a mild-temperate, Mediterranean-Atlantic species confined to ancient, undisturbed woodlands (also e.g. Giavarini et al. 2009), and thus regi-onally regarded as a species indicative of ecological continuity (e.g. Coppins & Coppins 2002). The Romanian Eastern Carpathians record above is from an outer part of a broadleaved forest complex formerly used by a livestock for grazing, and currently closely bordering a typical wood-pasture with large, solitary oak trees. The species was accompanied by other forest species such as Chrysothix candelaris, Lecanora albella, Normandina pulchella and Parmotrema perlatum, as well as lichens of rather open and drier habitats, namely Amandinea punctata, Candelariella xanthostigma, Parmelia sulcata, Flavoparmelia caperata and Ramalina pollinaria. These components of the present lichen biota show that this forest habitat, formerly extensively grazed and naturally fertilized, still has an ancient woodland character.Rinodina isidioides is a rare but widespread species in the World, known mainly from Western Europe: Norway, Great Britain, Ireland, France, Spain, Portugal, Switzerland and Italy (Tønsberg 1994, Martinez & Aragon 1998, Nowak 1998, Jones 2002, Scheidegger & Clerc 2002, Giavarini et al. 2009, Seaward 2010, Nimis & Martellos 2017), but also reported from North America: Mexico, Greater Sonoran Desert Region (Sheard et al. 2004) and Reunion Island in Indian Ocean (van den Boom et al. 2011). It has also been recorded from Kenia and Russia: South Caucasus (Giavarini et al. 2009, GIF Data Base, on-line 2015).

Rinodina subpariata (Nyl.) Zahlbr. [= R. degeliana Coppins; see Resl et al. 2016] Hungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu villa-ge, 47.96833°N/20.55889°E, alt. 300 m, on bark of Quercus cerris in oak wood-pasture, 16.05.2015, P. Czarnota 8239. Romania: Eastern Carpathians, Transylvania, Oklánd county, 3 km S of Homoródújfalu village, 46.13172°N/25.41517°E, alt. c. 530 m, on bark of Quercus sp. in oak wood-pasture, 18.05.2015, P. Czarnota 7861; ibid., 46.12800°N/25.41460°E, alt. c. 560 m, on bark of Quercus sp. in semi-forested former wood-pasture close to the edge of forest, 19.05.2015, P. Czarnota 7844. Ukraine: Eastern Carpathians, Ivano-Frankivsk region, Rozhniativ county, 1.5 km W of Ivanivka village, 48.88606°N/24.09161°E, alt. 490 m, on bark of Quercus sp. in young oak-hornbeam forest, a former wood-pasture, 07.06.2015, P. Czarnota 8120; ibid., 48.88672°N/24.09631°E, on bark of Quercus sp. in forested former wood-pasture, alt. c. 490 m, 08.06.2015, P. Czarnota 8131; ibid., 2.5 km SW of Rozhniativ town, 48.91917°N/24.11389°E, alt. 430 m, on bark of Quercus sp. in semi-forested former wood-pasture at the edge of forest, 06.06.2015, P. Czarnota 8181; ibid., 2 km S of Rakiv village, 48.97683°N/24.12208°E, alt. 430 m, on bark of Quercus sp. in semi-forested wood-pasture at the edge of forest, 10.06.2015, P. Czarnota 8195.

Chemistry: atranorin, zeorin, unidentified terpenoid (Rf-1 in solvent C) (examined specimen: Czarnota 8131).New to Hungary and Romania.Rinodina subpariata is usually found in open old-growth deciduous forests, but probably often over-looked due to its inconspicuous, areolate to sublobate thallus which often dissolve into the thin leprose crust resembling at first glance other sterile sorediate species, such as young thalli of some representa-tives of the genus Lepraria Ach. (Kubiak 2010). For excellent illustrations of the characteristic labri-form soraliate thallus and a description of re-revised characters of R. subpariata see Resl et al. (2016) and, for example, Tønsberg (1992) and Kubiak (2010), in both cases as R. degeliana.Despite rarely reported (always as R. degeliana), it is a widely distributed species in the Northern Hemisphere: Europe, North America and Asia (Hauck & Javkhlan 2006, Kubiak 2010 and litera-ture cited therein, Urbanavichus & Ismailov 2013, Resl et al. 2016, Sheard et al. 2017). European

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range of R. subpariata includes Great Britain, Norway, Sweden, Finland, Lithuania, Estonia, Austria, Italy, Poland and Russia (Kukwa & Kubiak 2007, Otte 2007, Kubiak 2010 and literature cited therein, Nascimbene 2014) and also the Czech Republic (Malíček & Palice 2013) and Germany (Wirth et al. 2013). This species has recently been collected also in primeval beech forests in the Slovak (Vondrák et al. 2015) and Ukrainian parts of East Carpathians Biosphere Reserve (Malíček et al. 2018).

Ropalospora viridis (Tønsberg) TønsbergHungary: Inner Western Carpathians, Bükk Mts, Bükk National Park, c. 3 km NE of Cserepfalu village, 47.96730°N/20.55011°E, on bark of Quercus sp. in young oak plantation close to oak wood-pasture, P. Czarnota 7896, 7912, 7919, 7920b (det. M. Kukwa).

Chemistry: perlatolic acid.New to Hungary.Recently this sorediate species has increasingly been reported from the Carpathians, including Polish (e.g. Fałtynowicz 2003 and literature cited therein), Slovak (Pišút et al. 2007, Vondrák et al. 2015, Adamčík et al. 2016), Ukrainian (e.g. Coppins et al. 2005, Dymytrova et al. 2013, Malíček et al. 2018) and Romanian parts (Ardelean et al. 2013, Vondrák & Liška 2013, Malíček et al. 2015), thus the Hungarian locality above fills the gap in the Carpathian distribution of R. viridis. It is locally frequent on trunks of deciduous trees in woodlands being sometimes overlooked due to its morpho-logical similarity to several other sorediate species including, for example, Biatora pontica, Buellia griseovirens, Fuscidea arboricola, F. pusilla, Lecidella subviridis or Violella fucata. The thallus of Ropalospora viridis does not react with C, K and P, while in above-mentioned species at least one reagent shows a conspicuous positive colour reaction, except for F. pusilla which produces divaricatic acid, while R. viridis has perlatolic acid (see also notes under Biatora pontica and Fuscidea pusilla, and Tønsberg 1992).

Verrucaria viridigrana BreussPoland: Atpol Gf-09, Eastern Carpathians, Góry Sanocko-Turczańskie Mts, c. 0.5 km W of Kalwaria Pacławska village, 49.62980°N/22.69901°E, on bark of Quercus robur, 18.07.2015, Czarnota 7953.

New to Poland and Polish Carpathians.Considering its ecological preferences, V. viridigrana could be used as an indicator of old-growth forests or/and indicator of forest ecosystem continuity (see references below). The new Polish locality is from an open-canopy old oak grove with dense hazel Corylus avellana undergrowth. There is no evidence that the locality is a remnant of a former ancient forest, but many vascular plants characteris-tic of a fertile deciduous submountain forest are present, indicating the spontaneous process of forest habitat recovery.This pyrenocarpous microlichen has been rarely recorded to date, always within old, wet, deciduous forests, in several central European localities, including Ukrainian Eastern Carpathians (Breuss 1998, Dymytrova et al. 2013, Malíček et al. 2018), Muránská planina in Slovak part of Western Carpathians (Breuss 1998, Guttova & Palice 1999) and old-growth lowland forest Cahnov-Soutok National Nature Reserve in Southern Moravia (Vondrak et al. 2016). The only other European re-cords are from Upper Austria (Breuss 1998) and Ukraine, Kyiv Region (Dymytrova 2013). It was once reported from Canada (Lay 2004).

AcknowledgementsAuthors thank Prof. Martin Kukwa (University of Gdańsk, Poland) for making available TLC analyses in his labora-tory and his help with determination of sterile species. We are also grateful to Prof. Mark Seaward (Bradford, UK) for the English improvements and J. Vondrák (AV ČR, Průhonice, Czech Republic) and the second reviewer (anonymous) for very constructive criticism. Financial support for this study was received through the project ‘Oak woods in rural landscapes of the Carpathian region: origin, dynamics and conservation values’, financed by the National Science Centre, Poland, following the decision DEC-2013/11/B/NZ9/00793.

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Manuscript accepted: 24 January 2018.Communicated by: Christian Printzen

Addresses of the authorsPaweł Czarnota, Department of Agroecology, Faculty of Biology and Agriculture, University of Rzeszów, Ćwiklińskiej 1A, 35-601 Rzeszów, Poland. E-mail: [email protected] Helmut Mayrhofer, Institute of Plant Sciences, NAWI Graz, University of Graz, Holteigasse 6, 8010 Graz, Austria. E-mail: [email protected] Bobiec, Department of Agroecology, Faculty of Biology and Agriculture, University of Rzeszów, Ćwiklińskiej 1A, 35-601 Rzeszów, Poland. E-mail: [email protected]