The lichen family Parmeliaceae in Poland. I. The genus Parmotrema

the paper presents the results of study on four Parmotrema species in Poland, P. arnoldii, P. crinitum, P. perlatum and P. stuppeum. they are mainly known from southern part of the country, mostly in the Carpathians. P. perlatum is reported also from northern Poland and Central Poland. For P. stuppeum, so far known from single locality only, three new sites are reported. All the species seem to be rare and endangered in Poland. this work is the first part of a larger series which will present data on selected genera of the lichen family Parmeliaceae in Poland.

not always followed and all segregates treated as doubtful (e.g., Purvis et. al. 1992). recently, the taxonomy of Parmeliaceae has been studied with molecular tools, and the importance of chemical, morphological and anatomical diagnostic characters have been re-evaluated (e.g., blanco et al. 2004a, b, 2005; divakar et al. 2006). numerous Hale's genera and those distinguished later by other lichenologists are still accepted, however, some of them appeared to be synonyms (blanco et al. 2004a, b, 2005; divakar et al. 2006).
Parmotrema A. Massal. is one of the genera accepted at present, however in a wider sense then previously, as it also includes Canomaculina elix & Hale, Concamerella w. l. Culb. & C. F. Culb., Parmelaria Awasthi and Rimelia Hale & Fletcher, which were found to be phylogenetically nested within Parmotrema s.str. (blanco et al. 2005). the genus is characterized by usually large foliose thalli with ciliate or eciliate margin, lack of pseudocyphellae, upper cortex consisting of a palisade plectenchyma (or rarely paraplectenchyma), pored or fenestrated epicortex, laminal, perforate or eperforate apothecia and usually simple rhizines (blanco et al. 2005).
As many Parmotrema species are very similar in morphology, the secondary chemistry plays an important role in the identification and taxonomy of these lichens. the substances found in this genus comprises atranorin and rarely usnic acid, which are present in the cortex, and several medullary compounds belonging to orcinol depsides, orcinol depsidones, ß-orcinol depsides, xanthones, aliphatic acids, pulvinic acid and derivatives and antraquinones (cf. Hale 1965;blanco et al. 2005). For the determination of them colour reactions with simple chemical reagents (C, k and Pd) were used in Poland, however this method is not very reliable, as it can not distinguish lichen substances with the same type of reaction. For that reason specimens should be studied by thin-layer chromatography (tlC) .
until now 4 taxa have been reported in Poland (Fałtynowicz 2003), and almost all of them have not been confirmed by tlC. some species were consider to be rather common, e.g., P. perlatum (Huds.) M. Choisy, but some were very rarely reported from few stands only, e.g., P. stuppeum (taylor) Hale. our unpublished results show that frequency for some taxa is different. the aims of this paper, the first dealing with the revision of selected genera of the family Parmeliaceae in Poland, are to present the result of studies on the morphology, chemistry, distribution and habitat requirements of the genus Parmotrema in the country.
MAteriAl And MetHods the present study is based on collection which are deposited in Polish lichen herbaria (krAM, krAP, ktC, lbl, uGdA). the morphology of the specimens were examined under the stereo microscope for thallus colour, shape of lobes, presence and abundance of cilia, and the type of soralia and isidia. the lichen substances were investigated by thin layer chromatography (tlC) following the methods described by orange et al. (2001). the chromatograms were developed in solvent C.
Affinities. the species is distinguished from morphologically similar taxa producing soredia by the production of alectoronic and α-collatolic acids. there is no other species of Parmotrema with these substances in europe (Hale 1965), however this substances are produced in the morphologically somewhat similar Cetrelia chicitae (w. l. Culb.) w. l. Culb. & C. F. Culb. this species differs in the presence of pseudocyphellae and eciliate thallus margin (obermayer, Mayrhofer 2007).
Habitat requirements. According to louwhoff (2009a) P. arnoldii usually grows amongst epiphytic mosses, especially on horizontal branches of trees and old shrubs in ± well-lit, mild and humid, undisturbed woodlands. in Poland this lichen was collected on Acer spp., Alnus spp., Fagus sylvatica, Fraxinus excelsior and Pyrus sp. in beech and black alder forests or in open places.
Distribution. in Poland P. arnoldii is a rare mountain species, known only from the Carpathians (Fig. 1). Most of the analyzed specimens originates from 1950-1960, and perhaps the species is close to extinction.
Affinities. the species is easily recognizable by the presence of isidia, broad bare marginal zone and the production of stictic acid complex. Parmelinopsis horrescens (taylor) elix & Hale also has ciliate-isidia, but it is distinguished by the smaller thallus, the lower surface that is rhizinate to the margins and the lack of stictic acid complex (louwhoff 2009b). so far this species has not been found in Poland.
Marginal and laminal isidia of similar shape to those present in Parmotrema crinitum are present also in Platismatia glauca (l.) w. l. Culb. & C. F. Culb., but that species lacks cilia, does not react with Pd (stictic acid complex absent) and produces caperatic acid (Culberson, Culberson 1968).
Habitat requirements: the species usually grows on bark of deciduous trees in old forests or in open situations (Fałtynowicz 2003; louwhoff 2009a). in Poland it was so far reported only as epiphyte on Abies alba, Acer pseudoplatanus, Alnus spp., Fagus sylvatica (ca 58% of Polish records) and Picea abies, usually in beech forests or along streams.
Distribution. According to Fabiszewski (1968) and Fałtynowicz (2003 and literature cited therein) P. crinitum was reported from the Carpathians and sudety Mts., however this study confirm its occurrence in the Carpathians (material from sudety Mts. not traced). known distribution of the species is presented on figure 2. this lichen is known only from old collections, and perhaps it is already extinct in the territory of Poland. P. crinitum is a cosmopolitan species, widespread throughout tropical and temperate areas and even subboreal forests (Hale 1965;louwhoff 2009a). it was reported in europe from, e.g., Austria (Hafellner, türk   Characteristic of the species. the diagnostic characters of P. perlatum are the conspicuous submarginal soralia, revolute lobes, narrow, shiny and often rugose naked zone of lower side along the margin. the species contains atranorin, stictic acid complex, including stictic, constictic and menegazziaic acids with other related substances. Cortex reacts k+ yellow, whereas medulla is C−, k+ yellow and Pd+ orange(louwhoff 2009a). the chemistry of Polish agrees with the previously reported results.
Affinities. Parmotrema perlatum can be separated from other sorediate and marginally ciliate species, P. arnoldii (du rietz) Hale and P. robustum (degel.) Hale, by the presence of stictic acid complex (Hale 1965;louwhoff 2009a). in the spot test Fig. 3. distribution of Parmotrema perlatum in Poland in the AtPol grid square system. reactions, P. perlatum is similar to P. reticulatum (taylor) M. Choisy, but the latter is readily distinguished by the network of hair-line cracks on the upper surface, the soredia at tips of incised lobe margins and the presence of salazanic acid (louwhoff 2009a); so far it has not been reported from Poland.
Habitat requirements. Parmotrema perlatum is sensitive to air pollution and occurs on well-lit, neutral to somewhat acid-barked, broad-leaved trees, also frequently on siliceous rocks and walls, coastal rocks, usually where illumination is moderate to good (lowhoff 2009a). in Poland this lichen was found mostly on bark of Alnus spp. and Fagus sylvatica, but single specimens were also collected on bark of Fraxinus excelsior and Picea abies.
Characteristic of the species. Parmotrema stuppeum is characterized by thallus loosely attached to the bark, black and sparsely rhizinate lower surface, continuous (not finely reticulately cracked) upper cortex, soralia formed on the top of laciniae and the presence of atranorin in cortex and salazinic acid in medulla (Hale 1965;sipman 2005). Cortex reacts k+ yellow, whereas medulla is C and kC-, k+ yellow turning red, and Pd+ orange-red. the chemistry of Polish specimens agrees with previously reported information.
Affinities. the species can be separated from other sorediate Parmotrema species with continuous cortex by the presence of salazinic acid. this substance is produced by P. reticulatum (tayl.) M. Choisy [syn. Rimelia reticulata (tayl.) Hale & Fletcher] and P. subreticulatum (tav.) Hale, but these taxa has finely, but distinctly reticulately cracked and maculate upper cortex (Hale 1965;sipman 2005;louwhoff 2009a); so far they have not been reported from Poland.
in Poland most specimens of P. stuppeum were determined as P. chinense, a synonym of P. perlatum (see above). both taxa have Pd+ orange to orange-red medulla, but it is due to salazinic acid in P. stuppeum, whereas in P. perlatum the reaction is caused by stictic acid and related substances. they can also be separated without tlC; medulla of P. perlatum is k+ yellow, but in P. stuppeum the yellow colour turns red (Hale 1965;sipman 2005;louwhoff 2009a).
Habitat requirements. in Poland P. stuppeum is a corticolous lichen. it was collected almost exclusively on Fagus sylvatica, but once it was found on bark of Pyrus communis.
Distribution. in Poland the species has been so far reported only once by sulma and Fałtynowicz (1988). Here we report three new localities, two from bieszczady Mts. and one from roztocze. the known distribution of the species is presented on figure 4. it appears as a rare and an endangered lichen in Poland, as only one contemporary record is known.
According to Hale (1965) P. stuppeum is common in the mountain of north America, central America and europe; in Africa and Asia it is more rare. in europe the species has been reported from Austria (Hafellner, türk 2001), the Czech republic (liška et al. 2008), Germany (Hale 1965;scholtz 2000), Portugal (Hale 1965), spain (llimona, Hladun 2001(llimona, Hladun ), switzerland (scheidegger et al. 2002(llimona, Hladun ) and ukraine (kondratyuk et al. 1998. it was also reported from the netherlands, but it has been recently considered as misidentification; the records belong to P. subreticulatum (Aptroot et al. 2008).

ConClusions
Four species of Parmotrema are known to occur in Poland, P. arnoldii, P. crinitum, P. perlatum and P. stuppeum. they have been mainly found in southern part of the country, mostly in the Carpathians. only P. perlatum is known from two localities in northern and central Poland.
the study has shown, that the secondary lichen metabolites are very useful character in determination of species; the analyzes of lichen secondary metabolites led to discovery of 3 new localities of P. stuppeum. so far the species has been reported only from one locality. At present, only P. perlatum and P. stuppeum are known to be represented by recent collections. it appeared, that P. arnoldii and P. crinitum have not been collected since 1962. thus, all Parmotrema species should be considered as in great danger of extinction in Poland. Possibly two of them, P. arnoldii and P. crinitum, are already extinct in Poland.