Lichen-forming fungi of the genus Montanelia in Poland and their potential distribution in Central Europe

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1 Herzogia 28 (2) Teil 2, 2015: Lichen-forming fungi of the genus Montanelia in Poland and their potential distribution in Central Europe Katarzyna Szczepańska *, Daniel Pruchniewicz, Justyna Sołtysiak & Maria Kossowska Abstract: Szczepańska, K., Pruchniewicz, D., Sołtysiak, J. & Kossowska, M Lichen-forming fungi of the genus Montanelia in Poland and their potential distribution in Central Europe. Herzogia 28: This paper presents the results of studies concerning the distribution and habitat requirements of two species of Montanelia (Parmeliaceae, Ascomycota) in Central Europe. Morphological and chemical analyses of herbarium material confirmed the presence of two species in Poland: M. disjuncta and M. sorediata. These species are relatively rare in the country and occur predominantly in mountainous and upland areas. Due to their morphological similarity, both species are difficult to identify and are also often confused with brown species of Xanthoparmelia. Collection localities and bioclimatic variables were used to develop MaxEnt models of the potential geographical distribution of the studied species in Central Europe. The species distribution model of M. disjuncta indicated mountains as the most suitable habitats, especially the eastern and western Carpathians and the Sudetes, as well as the eastern part of the Alps. The highest potential distribution of M. sorediata is larger than that of M. disjuncta and also includes the Bohemian-Moravian Highlands, and the Podolian and Malopolska Uplands. The least potential distribution of both discussed species was found in the German Lowlands. The potential distribution in both models was strongly influenced by precipitation-related variables. Zusammenfassung: Szczepańska, K., Pruchniewicz, D., Sołtysiak, J. & Kossowska, M Flechtenbildende Pilze der Gattung Montanelia in Polen und ihre potentielle Verbreitung in Mitteleuropa. Herzogia 28: Die Arbeit stellt die Ergebnisse von Untersuchungen zu Verbreitung und Habitatansprüchen von zwei Arten der Gattung Montanelia (Parmeliaceae, Ascomycota) in Mitteleuropa vor. Morphologische und chemische Analysen von Herbarmaterial bestätigen die Präsenz von zwei Arten M. disjuncta und M. sorediata in Polen. Sie sind hier relativ selten und kommen hauptsächlich in Bergregionen und im Hochland vor. Die Arten sind aufgrund ihrer morphologischen Ähnlichkeit schwer auseinanderzuhalten. Sie werden auch häufig mit braunen Arten der Gattung Xanthoparmelia verwechselt. Sammlungsdatensätze und bioklimatische Variablen wurden für die Erstellung von MaxEnt-Modellen der potentiellen geographischen Verbreitung der untersuchten Arten in Mitteleuropa verwendet. In der resultierenden Karte für M. disjuncta treten Gebirge und Hochlandsgebiete im südlichen Teil der modellierten Region als geeignetste Habitate hervor. M. sorediata hat eine viel ausgedehntere potentielle Verbreitung, die sich bis in das nordöstliche und östliche Tiefland erstreckt. Obwohl beide Arten als typische Gebirgsbewohner angesehen werden, zeigen die Modelle deutlich eine Meidung höherer Lagen. Die potentielle Verbreitung ist in beiden Modellen stark abhängig von Variablen, die mit den Niederschlägen in Beziehung stehen. Key words: Parmelioid lichens, Montanelia disjuncta, Montanelia sorediata, taxonomy, ecology, biogeography, potential distribution, MaxEnt. Introduction Although parmelioid lichens produce large thalli and are therefore relatively easy to detect and identify, detailed information concerning the distribution and habitat requirements for * Corresponding author

2 698 Herzogia 28 (2) Teil 2, 2015 many species does not exist. The problem is further complicated by the fact that some species were not previously recognized in the past, whereas others were often incorrectly identified. Numerous studies concerning the taxonomy and phylogeny of this group have recently been conducted to address this knowledge gap. In Poland, the lichens from the genera of Parmotrema (Jabłońska et al. 2009), Cetrelia (Kukwa et al. 2012) and brown species of Xanthoparmelia (Szczepańska & Kossowska 2014) have been reviewed. These studies consisted of verifying the identification of preserved herbarium material and mapping the current distribution of the species. Such revisions allow the potential geographical distribution of individual species to be determined, supporting ecological and biogeographic research, and facilitating risk assessment regarding extinction of the studied species. Another thoroughly studied taxon is the recently described genus Montanelia Divakar, A.Crespo, Wedin & Essl., which accommodates species of the former Melanelia disjuncta group (Divakar et al. 2012). The taxa belonging to this group were initially included within the subgenus Melanoparmelia of the genus Parmelia (Esslinger 1977). Subsequently, Melanoparmelia was raised to genus rank under the name Melanelia (Esslinger 1978), with Melanelia stygia as a type species. In subsequent years, molecular analyses have indicated that the genus Melanelia was polyphyletic (Blanco et al. 2004, 2006, Crespo et al. 2010, Thell et al. 2012), and consequently, some new genera were recognized, such as Melanelixia O.Blanco, A.Crespo, Divakar, Essl., D.Hawksw. & Lumbsch and Melanohalea O.Blanco, A.Crespo, Divakar, Essl., D.Hawksw. & Lumbsch. Furthermore, the Melanelia disjuncta group proved to be unrelated to Melanelia stygia, which was clustered in a cetrarioid clade as sister to Melanelia hepatizon (Crespo et al. 2010, Thell et al. 2009, Nelsen et al. 2011). Finally, Melanelia s. str. was restricted to a small clade of saxicolous cetrarioid lichens, when the taxa of Melanelia disjuncta group, representing the parmelioid clade, were reclassified into a new genus, Montanelia (Divakar et al. 2012). At present, the genus Montanelia includes five taxa: M. disjuncta, M. panniformis, M. predisjuncta, M. sorediata, and M. tominii. However, recent molecular studies imply that the nominal taxa M. panniformis and M. tominii include previously unrecognized species-level diversity. Additionally, Leavitt et al. (2015) suggested that M. predisjuncta is potentially a synonym of M. disjuncta. Therefore, in the authors opinion, a further, formal, taxonomic revision in Montanelia is required. As currently defined, the characteristic features of the Montanelia genus are short and narrow lobes, with plane to convex lobe margins, a non-pored epicortex, cylindrical to fusiform conidia, a medulla containing orcinol depsides (perlatolic, stenosporic or gyrophoric acids), and flat, effigurate pseudocyphellae (absent only in M. sorediata; Divakar et al. 2012). According to Otte et al. (2005), all the species of this genus are arctic alpine and circumpolar and occur mainly on silicate rocks in the mountain areas of the northern hemisphere, including Arctic regions (Divakar et al. 2012). The only exception is M. panniformis, which was also reported by Esslinger (1977) from the southern hemisphere. To date, three species of Montanelia have been reported from Poland: M. disjuncta, M. panniformis, and M. sorediata (Fałtynowicz 2003). Two of these are considered to be rare and endangered in the country, and are listed on the Polish red list of lichens (Cieśliński et al. 2006), as Critically Endangered (M. panniformis) and Vulnerable (M. disjuncta). Montanelia sorediata was considered to be the most common, at numerous localities throughout the country (Fałtynowicz 2003). However, Montanelia has not been the subject of a critical revision

3 Szczepańska et al.: Lichen-forming fungi of the genus Montanelia in Poland 699 in Poland, and data concerning the distribution, ecology and conservation status for individual taxa are incomplete and require revision. Therefore, we undertook a detailed review of herbarium material from Poland to verify the data on lichens in this group. Further study objectives were to estimate the potential geographical distribution of the species in Central Europe and to identify the most suitable ecological conditions for these taxa and the factors that determine their occurrence. Material and methods Chorological and taxonomical analyses The studied herbarium specimens originated from the following Polish herbaria and private collections: GPN, KRAM-L, KRAP, KTC, LOD-L, POZ-L, TRN, UGDA, WA, WRSL, hb. Kossowska, hb. Lipnicki and hb. Szczepańska. A total of 67 specimens were examined. Each specimen was analyzed for the presence of secondary metabolites in the thallus using thin layer chromatography (in solvents A and C), according to methods described by Orange et al. (2001). Additionally, a thorough examination of morphological features under a stereoscopic microscope was conducted, with particular focus on the shape and size of lobes and soralia and the presence of pseudocyphellae and pycnidia. Brief descriptions of the species provided in this paper are based on personal observations. The names of physiogeographic mesoregions are given according to Kondracki (2002). The distribution of the species in Poland is shown on the maps based on the ATPOL grid square system (Zając 1978), modified by Cieśliński & Fałtynowicz (1993). Modeling of a potential range of species using MaxEnt MaxEnt is a maximum-likelihood modeling method based on the maximum entropy principle (Elith et al. 2011, Philips et al. 2006, Parolo et al. 2008, Syfert et al. 2013). This method is used for niche modeling, also known as species or habitat potential distribution modeling (Guisan & Thuiller 2005), and it enables the identification of areas where ecological factors, including climatic conditions, are the most favorable for the species outside their known localities. It is especially suitable for lichens, when the available data on their distribution are incomplete and do not fully reflect the entire potential range of the taxon. Environmental variables and data preparation. Twenty topographic and bioclimatic variables with a highest resolution of 30 arc-seconds (~1 km) were downloaded from World- Clim ( and were used for MaxEnt modeling of the current distribution of Montanelia species in Central Europe. The bioclimatic variables were interpolations of recorded monthly data from the years (Hijmans et al. 2005). Before using the environmental variables in MaxEnt modeling, a correlation matrix was prepared to determine multicollinearity between the study variables. Statistical calculations were conducted using ENMTools software (Warren et al. 2010). For the model building, we used altitude, seasonal temperature (bio 4) and seasonal precipitation (bio 15), which in our analysis were not cross-correlated and play an important role, especially for the distribution of montane species (Percequillo 2003, Anderson & Raza 2010). Model building. The model was constructed using MaxEnt v software (Phillips et al. 2006, Phillips & Dudik 2008). To avoid spatial bias, the localities that were less than 10 km

4 700 Herzogia 28 (2) Teil 2, 2015 apart were randomly removed (Kramer-Schadt et al. 2013). The final data set, which was used in the modeling contains 14 locations for M. disjuncta and 28 for M. sorediata from Poland, the Czech Republic and the Ukraine. The analyses were carried out from the logistic output format with default settings as follows: random test percentage 25, maximum number of background points 10,000, regularization multiplier 1 (Kramer-Schadt et al. 2013). To optimize the settings in MaxEnt modeling using the distribution of species from few localities, the cross-validated method for both models was used (Muscarella et al. 2014). The area under the receiving operator curve for the training (AUCr) and for testing (AUCt) data was used to evaluate the resulting models. If the difference between the AUCr and AUCt was low, the model was not overfitted (Shcheglovitova & Anderson 2013). The effects of the environmental variables on the distribution of the studied species were estimated with the jackknife test. The final maps of the potential distribution of Montanelia species were characterized by values ranging from 0 to 1, where the values >0.6 represented a high potential, those between a good potential, those between a moderate potential, and the values <0.2 represented the category with the least potential (Yang et al. 2013). Results Montanelia disjuncta (Erichsen) Divakar, A.Crespo, Wedin & Essl. Am. J. Bot. 99: Parmelia disjuncta Erichsen, Ann. Mycol. 37: Melanelia disjuncta (Erichsen) Essl., Mycotaxon 7: Diagnostic characters. Thallus foliose, loosely adnate to the substrate. Lobes mm broad, flat to slightly convex, usually distinctly shiny, broadened and rounded at the ends. Upper surface olivebrown, dark-brown to reddish-brown, smooth and usually shiny. Pseudocyphellae small and rather indistinct, submarginal. Soralia mm in diameter, arising on the surface of the lobes and at the margins, punctiform, irregular to capitate or eroded and then flat or crateriform, older becoming confluent. Soredia granular to isidioid, dark but appearing white when abraded (Fig. 1A D). Pycnidia rare, conidia µm. Apothecia not seen in the examined material. Secondary metabolites: perlatolic and stenosporic acids. Habitat. Montanelia disjuncta is a saxicolous lichen that grows on non-calcareous substrates. In Poland, the species has been recorded on natural siliceous rock outcrops of various chemical and mineral composition (basalt, gneiss, granite, quartzite, and sandstone) in well-lit, warm and dry places. According to the jackknife test, performed in the MaxEnt software, its distribution within the modeled area is mostly determined by seasonal precipitation. General distribution. The geographical range of Melanelia disjuncta consists of both continental and oceanic areas of Europe and North America (Esslinger 1977, Otte et al. 2005, Hansen 2013). In central and eastern Europe, this species has been recorded in Austria, the Czech Republic, Germany, Hungary, Poland, Russia, Slovakia, Switzerland and the Ukraine (Hawksworth et al. 2008). Distribution in Poland. Montanelia disjuncta occurs in the southern, mountainous part of the country, with a distinct concentration of localities in the Sudetes and their foreland (SW Poland; Fig. 2). In this region, the species was recorded only at lower altitudes (up to about 800 m above sea level), especially on mountain slopes and in the valleys. It was not reported from the highest summits of the Karkonosze (Giant) Mountains. Single and dispersed localities of M. disjuncta are known from the Carpathians and Góry Świętokrzyskie (Holy Cross Mountains). In the Carpathians, this species was also recorded in higher parts of the mountains. It reaches the highest altitude in the Bieszczady Mountains (Eastern Carpathians), occurring at about 1150 m above sea level. Our study did not confirm the occurrence of the species outside of mountainous areas (see Fałtynowicz 2003). All the examined specimens collected in lowland and upland areas were M. sorediata.

5 Szczepańska et al.: Lichen-forming fungi of the genus Montanelia in Poland 701 Fig. 1: Montanelia disjuncta. A thallus; B lobes broadened at the ends and shiny; C lobes with laminal or submarginal soralia; D punctiform to irregular soralia. Scales: A 0.5 cm, B C 2 mm, D 1 mm. Potential distribution in Central Europe. In MaxEnt modeling, the AUC value for the training and testing data was 0.885, thus confirming the high accuracy and that the potential distribution model for M. disjuncta was not overfitted. A high potential distribution of the studied species (>0.6) was found in the southern part of Poland, especially in the Western and Eastern Carpathians (the Beskids Mountains) and throughout the Central Sudetes (Fig. 3). Outside Poland, optimal conditions for M. disjuncta were found in the highest parts of the Eastern and Western Carpathians (Slovakia, the Ukraine, Romania). A high potential distribution was also found in the Alps, especially in the eastern part of the range (Austria and Germany). Good conditions ( ) were found in most of the modeled area, whereas unsuitable conditions (the least potential distribution, <0.2) of M. disjuncta occurred within the German Lowland. Number of specimens examined: 20. Specimens examined: [Ea 78] Karkonosze Mts: Szklarska Poręba town, Marianki rocks, on granite, alt. ca 700 m, 7 Oct. 2007, M. Kossowska 52 (hb. M. Kossowska). [Ea 79] Karkonosze Mts: Grabowiec Mt., Patelnia rock, on granite, alt. ca 720 m, 7 Jul. 2013, M. Kossowska 1371, 1372 (hb. M. Kossowska, two specimens). [Eb 60] Kotlina Jeleniogórska Basin: Jelenia Góra city, Zamkowa Góra Mt., siliceous boulder on the meadow, alt. ca 347 m, 3 May 2005, K. Szczepańska 517 (hb. K. Szczepańska). [Eb 64] Wzgórza Strzegomskie Hills: Góra Krzyżowa Mt., alt. 350 m, on basalt outcrop, 4 Oct. 2013, K. Szczepańska 969 (hb. K. Szczepańska). [Eb 84] Góry Sowie Mts: Choina Mt., gneiss rock in the beech forest, alt. ca 400 m, 17 Feb. 2014, K. Szczepańska 989 (hb. K. Szczepańska). [Ee 77] Góry Świętokrzyskie Mts: Łysa Góra Mt. forest section No. 196, on quartzite, 8 Jul. 1982, S. Cieśliński (KTC). [Fb 36] Masyw Śnieżnika Massif: Pasterskie Skały rocks near Idzików village, alt. ca 600 m, on siliceous conglomerate rocks in the forest, 25 Aug. 2003, K. Szczepańska 299 (hb. K. Szczepańska). [Fb 38] Góry Bialskie Mts: Trzy Siostry rocks, on siliceous boulder field, 7 Aug. 2003, K. Szczepańska (WRSL- 2887). [Fb 46] Masyw Śnieżnika Massif: valley of Szklarzynka stream, siliceous rocks above the road, 28 Aug.

6 702 Herzogia 28 (2) Teil 2, 2015 Fig. 2: Topographic map of distribution of Montanelia disjuncta in Poland. 2003, K. Szczepańska (WRSL-2008). [Fb 47] Masyw Śnieżnika Massif: Młyńsko Mt, alt. 978 m, on gneiss, 26 May 2004, K. Szczepańska 482 (hb. K. Szczepańska). [Fe 97] Pogórze Ciężkowickie Foothills: vicinity of Grybów village, on rocks, 12 Sept.1926, J. Motyka (KRAM-L); Ciężkowice town, nature reserve, alt. ca 280 m, on sandstone rock, 16 May 1973, R. Kozik (KRAP). [Gd 15] Beskid Żywiecki Mts: Pilsko range, Sopotnia Mała village, on rocks, 23 Sept. 1964, J. Nowak (KRAM-L-16711). [Gd 58] Western Tatra Mts: Dolina Chochołowska valley, Polana Chochołowska glade, alt m, 16 Jul. 2004, L. Śliwa 3069 (KRAM-L-54415); the same locality, alt. ca 1115 m, on siliceous rock, Aug. 1982, K. Toborowicz (KTC). [Gg 60] Western Bieszczady Mts: Bukowe Berdo Mt., insolated sandstone rocks on the western slope, alt m, 11 Aug. 1958, K. Glanc (KRAM-L-36981). Additional material examined: Ukraine: Kamianets-Podilskyi, on stones, 1909, V. Kutak (WRSL); Gorgany Mts, Strymba Mt, on rocks, alt m, 26 Jul. 1934, T. Sulma (UGDA-L-2427); Gorgany Mts, Preluky, on boulders, 20 Jul. 1934, T. Sulma (UGDA-L-2435). Taxonomical comments. A thorough examination of the specimens preserved in Polish herbaria showed that identification of M. disjuncta could be relatively difficult. Among 42 analyzed specimens previously labeled as M. disjuncta, only 14 actually represented this species. Montanelia disjuncta is most commonly confused with another taxon of this group, namely M. sorediata. Both species exhibit very similar morphology; they have an olive-brown upper surface, produce soralia and grow on silice-

7 Szczepańska et al.: Lichen-forming fungi of the genus Montanelia in Poland 703 Fig. 3: Potential distribution modelling of Montanelia disjuncta in Central Europe. ous rocky substrates. According to Esslinger (1977), the most important feature distinguishing these two species is the presence of pseudocyphellae on the thalli of M. disjuncta. However, the pseudocyphellae in Polish specimens were usually poorly developed and difficult to observe. In contrast to M. sorediata, M. disjuncta also has the ability to produce pycnidia, but these are not always present. The most useful diagnostic features appear to be the location and shape of soralia. Montanelia disjuncta produces laminal or submarginal soralia, which can be punctiform, capitate or even flat and irregular. The soralia of M. sorediata are terminal, located on the tips of small, erect, lateral branches, and are always distinctly capitate and never confluent. The appearance of the lobes of these two taxa is also somewhat different. The lobes of M. disjuncta are slightly shorter, wider, broadened at the ends and shiny, whereas those of M. sorediata are narrower, often elongated and generally dull. Identification of these species always requires considering all of these traits simultaneously, because single features might not be visible on individual specimens. A species similar to M. disjuncta is M. tominii (Oxner) Essl., which has not been recorded in Poland to date, but has been recorded in many European countries, e.g., Austria, Switzerland, Spain, Italy, Latvia, Norway, Russia and Sweden (Hawksworth et al. 2008). Therefore, this species is probably also present in Poland. Both of these taxa are saxicolous, form laminal and marginal, punctiform to capitate soralia and have pseudocyphellae, which are more numerous and distinctly visible in M. tominii (Esslinger 1977). Moreover, unlike M. disjuncta, M. tominii usually produces apothecia and its medulla reacts C+ rose to red, due to the presence of gyrophoric acid. Other taxa that are morphologically similar to M. disjuncta and are therefore sometimes confused with this species, are brown representatives of the genus Xanthoparmelia (Vain.) Hale, especially X. loxodes (Nyl.) O.Blanco, A.Crespo, Elix, D.Hawksw. & Lumbsch and X. verruculifera (Nyl.) O.Blanco, A.Crespo, Elix, D.Hawksw. & Lumbsch. Both of these species produce pustular isidia, which form branched-coralloid structures that might slightly resemble confluent soralia on the older parts of M. disjuncta thalli. The main features that allow to distinguish Montanelia species from Xanthoparmelia

8 704 Herzogia 28 (2) Teil 2, 2015 include a generally smaller thallus, an upper cortex that does not react with HNO 3, and a different chemical composition of the thallus. Apart from stenosporic and perlatolic acids, which are present in all the discussed taxa, X. verruculifera also contains divaricatic acid, whereas X. loxodes contains glomelliferic and glomellic acids. In doubtful cases the specimens should be analyzed using thin-layer chromatography (TLC). In the examined herbarium material, M. disjuncta was also confused with Melanelixia fuliginosa (Duby) O.Blanco, A.Crespo, Divakar, Essl., D.Hawksw. & Lumbsch, which, however, develops fine, cylindrical isidia on the thallus surface and its medulla reacts C+ red. It was also confused with Melanelia stygia (L.) Essl., which does not produce soralia, forms distinct, numerous pseudocyphellae on the upper surface and has a different chemical composition of the thallus (fumarprotocetraric and protocetraric acids). Montanelia sorediata (Ach.) Divakar, A.Crespo, Wedin & Essl. Am. J. Bot. 99: Parmelia stygia var. sorediata Ach., Lichenogr. Universalis.: Melanelia sorediosa (Almb.) Essl., Mycotaxon 7: Melanelia sorediata (Ach.) Goward & Ahti, Mycotaxon 28: Diagnostic characters. Thallus foliose, loosely adnate to the substrate. Lobes mm broad, flat to slightly convex, sometimes elongate and linear, distinctly rugged and pitted at the ends. Upper surface olive-brown to dark-brown, dull but sometimes shiny at the lobe-ends, generally smooth. Soralia mm, arising on the ends of the main lobes or on the smaller, erect side lobes, usually distinctly convex and capitate, older crowded (Fig. 4A D). Soredia granular to isidioid, dark but appearing white Fig. 4: Montanelia sorediata. A thallus; B lobes narrow, elongated and dull; C lobes with terminal soralia, located on the ends of the main lobes or on the tips of small, erect, lateral branches; D convex and capitate soralia. Scales: A 0.5 cm, B C 2 mm, D 1 mm.

9 Szczepańska et al.: Lichen-forming fungi of the genus Montanelia in Poland 705 when abraded. Pseudocyphellae and pycnidia absent. Apothecia not found in the examined material. Secondary metabolites: perlatolic and stenosporic acids. Habitat. Montanelia sorediata is a saxicolous lichen that prefers siliceous substrates. In the lowland part of Poland, it has been recorded on siliceous erratic boulders, usually in open and well-lit places. In mountain areas, it occurs on natural rock outcrops composed of various minerals (granite, sandstone, gneiss etc.). The jackknife test calculated in the MaxEnt software, indicated that seasonal precipitation was the most important factor affecting M. sorediata distribution in Central Europe. General distribution. The species occurs in North America and Europe (Esslinger 1977, Otte et al. 2005). In central and eastern Europe, it has been reported from Austria, the Czech Republic, Germany, Lithuania, Poland, Romania, Russia, Slovakia, Switzerland and the Ukraine (Hawksworth et al. 2008). Distribution in Poland. Montanelia sorediata is more common and widely distributed than the previous species. It occurs in dispersed localities mainly in the east of the country, in both mountainous and lowland areas (Fig. 5). However, it appears to be rare in the western part of the Polish lowlands, and has been reported from only two sites. Fig. 5: Topographic map of distribution of Montanelia sorediata in Poland.

10 706 Herzogia 28 (2) Teil 2, 2015 To date, M. sorediata has not been found in the Sudetes. In the Carpathians, the species appears to avoid higher altitudes. All known localities are concentrated in the mountain valleys and lower parts of the slopes, up to 800 m (localities in the Żywiec Beskids and the Gorce Mountains). Potential distribution in Central Europe. In the MaxEnt modeling, the AUC value for the training data was and was for testing data, which confirms the high accuracy of the potential distribution model of M. sorediata. The highest potential distribution of the species (>0.6) was found in the Western Carpathians (Poland and Slovakia), Eastern Carpathians (Ukraine), as well as in the eastern part of the Alps (Austria). Besides the mountain regions, the highest potential distribution of the species was found in the Bohemian-Moravian Highlands (Czech Republic), the Podolian Upland (Ukraine) and Malopolska Upland (Poland). Good conditions ( ) for M. sorediata include the central and eastern parts of the modeled area, whereas unsuitable conditions (the least potential distribution, <0.2) occur in the German Lowland (Fig. 6). Fig. 6: Potential distribution modelling of Montanelia sorediata in Central Europe. Number of specimens examined: 34. Specimens examined: [Af-97] Pojezierze Wschodniosuwalskie Lakeland: Głazowisko Bachanowo nature reserve, on a granite boulder, 17 May 1985, S. Cieśliński & Z. Tobolewski (KTC); Turtul settlement, on stones, 18 Jul. 1987, W. Fałtynowicz (KRAM-L & UGDA-L-3131, two specimens). [Bc-36] Bory Tucholskie Woodland: Kręgi Kamienne nature reserve, on a siliceous boulder, Jul. 1985, L. Lipnicki 1163 (hb. L. Lipnicki); the same locality, 18 Aug. 1984, L. Lipnicki (TRN). [Bd-79] Garb Lubawski Hummock: 2 km NNW from Klonowo village, on a granite erratic boulder, Sept. 2002, Sz. Karandys, J. Kiszka, M. Kukwa, & A. Łubek (UGDA-L 9075). [Be-71] Pojezierze Olsztyńskie Lakeland: Nadrowo settlement near Waplewo village, on a stone, 23 Jun. 1996, S. Cieśliński (KTC). [Bf-73] Wysoczyzna Kolneńska Plateau: ca 1 km NWW of Kurki village, on an erratic boulder, 24 Aug. 1990, S. Cieśliński (KTC); ca 2 km E of Glinki village, on a boulder, 24 Aug. 1990, S. Cieśliński (KTC). [Bf-95] Wysoczyzna Kolneńska Plateau: Wagi Gnaty settlement, on a boulder, 23 Aug. 1990, S. Cieśliński (KTC). [Cg-14] Wysoczyzna Białostocka Plateau: Puszcza Knyszyńska Forest, ca 1,5 km E of Grzybowce village, on a boulder in the forest, 14 Aug. 1992, S. Cieśliński (KTC). [Cg-55] Równia Bielska Plain: Puszcza Białowieska Primeval Forest, Teremiski village, on a granite boulder, 1982, S. Cieśliński & Z. Tobolewski (KTC).

11 Szczepańska et al.: Lichen-forming fungi of the genus Montanelia in Poland 707 [Da-07] Pojezierze Lubuskie Lakeland: Łagówek village, on an erratic boulder by the country road, 6 Sept. 1950, Z. Tobolewski (POZ-L). [Df-25] Wysoczyzna Siedlecka Plateau: Leszczyny settlement, on an erratic boulder by the country road, 16. Sept. 1966, J. Zielińska (WA). [Df-40] Radachówka village, on an erratic boulder, 23. Sept. 1967, J. Zielińska (WA). [Ed-41] Wyżyna Wieluńska Upland: Lisowice village near Działoszyn town, on stones, 1 Jun. 1963, J. Nowak (KRAM-L-12172). [Ed-51] Wyżyna Wieluńska Upland: Patoki settlement near Działoszyn town, on quartzite erratic stones, 3 Jun. 1963, J. Nowak (KRAM-L-12187). [Ee-45] Płaskowyż Suchedniowski Plateau: Bliżyn settlement, glade in a forest S of village, on a sandstone boulder, 4 May 1959, J. Nowak (KRAM-L-2303). [Ee-53] Płaskowyż Suchedniowski Plateau: Wzgórza Kołomańskie Hills, Serbinów village, on siliceous stones, Jul. 1985, M. Bidzińska & K. Toborowicz (KTC). [Ee-54] Płaskowyż Suchedniowski Plateau: Świnia Góra nature reserve, on a sandstone boulder, 4 May 1959, J. Nowak (KRAM-L-2331). [Ee76] Góry Świętokrzyskie Mts: Łysica Mt, summit and top part of Agata boulder field, on quartzites, 27 Nov. 1982, S. Cieśliński (KTC). [Fd -94] Beskid Mały Mts: Maleckie Mt., alt. ca 800 m, on sandstones, 7 Aug. 1960, J. Nowak (KRAM-L-7692); Czernichów village, Pod Magurką locality on the slope above Dolina Roztoki valley, alt. ca 500 m, on stones, 24 Aug. 1960, J. Nowak (KRAM-L-7698). [Fd-96] Beskid Mały Mts: Gancarz Mt., alt. ca 750 m, on sandstones, 13 May 1960, J. Nowak (KRAM-L-7693, 7694, 7695, three specimens); Targoszów village, alt. ca 600 m, on stones in a cultivated field, 11 May 1960, J. Nowak (KRAM-L-7696); the same locality, alt. ca 550 m, on sandstones, 11 May 1960, J. Nowak (KRAM-L-7699). Ponikiew village, ravine by Królowa Wyżnia Mt., alt. ca 450 m, on sandstones, 12 May 1960, J. Nowak (KRAM-L-7697 & LOD-L , two specimens). [Fd- 99] Beskid Makowski Mts: valley of Kaczanka stream, on a stone, 23 Apr. 1966, J. Nowak (KRAM-L-17443). [Gd-07] Beskid Makowski Mts: Jałowiec range, east Skupniówka Mt. on a stone wall, 3 Sept. 1965, J. Nowak (KRAM-L-15508). [Ge-22] Gorce Mts: Barbarówka settlement in Ochotnica Dolna village, by the Ochotnica river, outcrop of calcareous sandstone, alt. 430 m, 5 Oct. 2000, P. Czarnota (GPN 2347); Ochotnica Dolna village, S slope of Twarogi Mt., on sandstone, 5 Oct. 2000, P. Czarnota (GPN 4799). Additional material examined: Czech Republic: Bohemian Forest, Zahvozdi settlement, near forester s lodge, on granite, 14 Apr. 2000, P. Czarnota (GPN 4795). Taxonomical comments. Montanelia sorediata proved to be the most common species of the discussed group in Poland. However, it is rarely reported, probably due to the difficulties in its identification. Among the 67 analyzed specimens, only 18 had been previously labeled as M. sorediata. The specimens of this species were usually referred to as M. disjuncta, which has a very similar morphology. According to Esslinger (1977), M. disjuncta and M. sorediata can be primarily distinguished based on the position of soralia, the appearance of their lobes and the presence of pseudocyphellae and pycnidia on M. disjuncta thalli (see comments on M. disjuncta). Similar to the previous species, M. sorediata is sometimes confused with brown representatives of Xanthoparmelia genus, especially X. loxodes and X. verruculifera. However, the species of the genus Xanthoparmelia have a different chemical composition, possess a generally larger thallus with wider lobes, and instead of soralia, produce pustular isidia, which sometimes resemble soralia. Discussion This study confirmed the occurrence of two species of the genus Montanelia previously reported from Poland, M. disjuncta and M. sorediata. The third of the reported taxa, M. panniformis, was not found among the analyzed specimens, and thus, a revision of its current distribution in the country is impossible. Montanelia panniformis should accordingly be treated at least as a critically endangered species in Poland and its high category of threat on the red list of Polish lichens should be maintained. Furthermore, due to a lack of contemporary records since 1992 (Alstrup & Olech 1992), its current status cannot be confirmed and there is a possibility that this taxon has become extinct in Poland. The most frequent and widely distributed species in the studied herbarium material was M. sorediata. However, the number of confirmed localities is still small and the species should not be considered as common in the country. The second species, M. disjuncta, appeared to be a less frequent taxon than previously reported, with a small number of confirmed localities and very limited range in the country. Most of the analyzed herbarium specimens labeled as M. disjuncta belonged to other taxa, e.g., M. sorediata and Xanthoparmelia verruculifera. Therefore,

12 708 Herzogia 28 (2) Teil 2, 2015 a change in the status of this species on the red list of Polish lichens from VU (Vulnerable) to EN (Endangered) should to be considered. The taxonomic analysis of specimens of the genus Montanelia showed that proper identification of brown parmelioid species based on their morphological traits was sometimes relatively difficult. Chemotaxonomic methods (e.g., TLC) should also be applied to distinguish taxa. The most important diagnostic features that enabled the identification of morphologically similar and therefore often confused taxa occurring in Poland are presented in Table 1. Table 1: Overview of the main distinguishing features of the vegetatively reproducing Montanelia and brown Xanthoparmelia species occurring in Poland. Species Isidia/soralia Pseudocyphellae Chemistry M. disjuncta soralia laminal, flat to capitate present perlatolic and stenosporic acid M. sorediata soralia convex, on erect side lobes absent perlatolic and stenosporic acid X. loxodes X. verruculifera isidia strongly pustular isidia slightly pustular absent absent glomelliferic, glomellic, perlatolic, stenosporic and occasionally gyrophoric acid stenosporic, divaricatic, perlatolic and occasionally gyrophoric acid Similar to the other representatives of the genus Montanelia, M. disjuncta and M. sorediata have a montane type of geographical range. This feature is even reflected in the name of the newly described genus, segregated from Melanelia s. str. (Divakar et al. 2012). Otte et al. (2005) considered both discussed taxa to be essentially arctic alpine, although they are also recorded in lower and middle montane belts and (rarely) in the lowlands. However, in the light of this study, these taxa appear to avoid the highest mountain sites. Their known localities from Polish territory are concentrated on mountain slopes and in valleys, only rarely exceeding an altitude of 1,000 m. Of the two studied species, the range of M. disjuncta shows a more montane character, because it is reported from higher altitudes and does not occur in the Polish lowlands. In contrast, most Polish records of M. sorediata came from erratic boulders in the lowlands and in the mountains; this species was found only in single and dispersed localities. These observations overlap to some extent with the results of the models constructed using MaxEnt for both species in Central Europe. The models of the geographical distribution of M. disjuncta include mainly mountains in the central and southern part of the modeled area as the most suitable habitats. It is notable that the areas with the most favorable habitat conditions (>0.6) for M. disjuncta are located at much higher altitudes than those for M. sorediata. The geographical distribution of M. sorediata appears to be much wider than that of M. disjuncta and it also includes the northeastern and eastern parts of the modeled area. The most suitable habitat conditions for this taxon (>0.6) are found in the submountain and mountain areas, but unlike M. disjuncta, this species definitely avoids higher altitudes, which is especially apparent in the Alps. The occurrence of M. sorediata in the lowlands might be the result of favorable local microclimate conditions not included in the models, or due to the availability of suitable silicate substrates. The factor that most strongly affects the potential geographic ranges of the two modeled taxa is precipitation. We can conclude that the variables related to precipitation are most often rep-

13 Szczepańska et al.: Lichen-forming fungi of the genus Montanelia in Poland 709 resented in the models concerning lichens and most often hold the highest predictive power in the presented MaxEnt analyses, compared, for example, to variables related to temperature (Szczepańska et al. in press). This can probably be explained by the fact that all physiological processes that occur in lichen thalli (gas exchange, nitrogen fixation and photosynthesis) depend on the availability of water, with its deficit preventing the proper functioning of the organism (Green et al. 2008, Nash 2008, Palmqvist et al. 2008). Therefore, climatic factors related to precipitation have the greatest influence on the potential distribution of lichens. According to the MaxEnt model, the most suitable ecological niches (>0.6) for M. sorediata in Poland are concentrated in the southeastern part of the country, especially in the mountain areas. The results of modeling confirm those of previous field studies. It should be mentioned that this species has not yet been recorded in the Sudetes, but due to suitable habitat conditions, it can probably be found there in the future. Beyond the Sudetes, new locations of M. sorediata might also be expected in the eastern Polish Carpathians and in the Małopolska Upland. According to the MaxEnt model, suitable habitat conditions for M. sorediata exist in most countries in Central Europe. These results confirm the known records of this species (Hawksworth et al. 2008). Montanelia disjuncta occurs especially in the southern, mountainous and upland regions of Poland. According to the MaxEnt model, these areas represent the most suitable climatic conditions for this taxa (>0.6). Montanelia disjuncta has been noted in most Central European countries (Hawksworth et al. 2008), which is also confirmed by the MaxEnt model. However, this species is also likely to occur in the Romanian Carpathians, where it has not yet been recorded. Potential ecological niche modeling is not yet widely used for lichens, although it is a tool that can be successfully applied to support research studies in the fields of biogeography, ecology and conservation biology of this group of organisms (Braidwood & Ellis 2012, Carlsen et al. 2012, Ellis et al. 2014). Recognition of the species distribution and the parameters that affect it are also necessary for the estimation and protection of local and global biodiversity (Newbold 2010). However, the interpretation of models of potential distribution always requires particular caution, because these models can be influenced by many different factors (e.g., agriculture, urbanization, geographical barriers that reduce dispersion and edaphic factors), most of which might not be included in the analyses (Newbold 2010). To prevent potential errors in the generated models, it is also necessary to consider several important principles, such as the correct identification of taxa, collecting specimens proportionally in different types of environments, using sufficiently large datasets and excluding an excessive number of environmental variables, to avoid overfitting the models (Chatfield 1995, Loiselle et al. 2008, Newbold 2010). This study attempted to take into account all of the mentioned principles, and the obtained models probably genuinely reflect the actual distribution of species of the genus Montanelia in Central Europe. The next step towards a validation of the obtained models could include field studies that should confirm the presence of the investigated taxa in new areas that represent the most suitable habitat conditions. Acknowledgments We are grateful to the all Polish lichenologists and curators of Polish herbaria for the loan of specimens for this study and to anonymous reviewers for their valuable remarks and corrections. The authors would like to thank also Dr. Volker Otte from the Senckenberg Museum für Naturkunde Görlitz and Dr. Magda Podlaska from Wroclaw University of Environmental and Life Sciences, for their help with the abstract translation.

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