ANNALES U N IV ERSIT ATIS M ARIAE C U RIE - S K Ł O D O W S K A LUBLIN - POLONIA VOL. LVI SECTIO C 2001

KATARZYNA KOLANKO

Institute of Biology, llniversity of Białystok, Świerkowa 20B, 15-950 Białystok,

Epiphytic lichen-dominated communities in the Knyszyn Forest

Zbiorowiska porostów epifitycznych w Puszczy Knyszyńskiej

SUMMARY

The paper aimed at describing the characteristics of the epiphytic lichen communities in the Knyszyn Forest. The investigations were extended over almost all the trees and shrubs growing in the Forest. The outdoor and phytosociological studies of lichens were carried out according to the principles of the Braun-Blanquet method modified by Klement and Barkman for examining lichen associations. The epiphytic flora was classified among seven associations. Pseudevernietum furfuraceae is the most common in the Knyszyn Forest. Two groups were assigned among epiphytic lichen associations; the first one: Chaenothecetum ferrugineae, Hypocenomycetum scalaris, Parmeliopsidetum ambiguae subass. Parmeliopsidetosum aleuritis and Pseudevernietum furfuraceae growing on , and mixed forests, and the second group: Pertusarietum amarae, Pyrenuletum nitidae and Thelotremetum lepadini growing only in deciduous trees forests on smooth bark. They are characteristic associations of primeval forests.

STRESZCZENIE

Celem badań była charakterystyka fitosocjologiczna epifitycznych zespołów porostów Puszczy Knyszyńskiej. Badania terenowe i fitosocjologiczne wykonano na wszystkich gatunkach drzew występujących w puszczy. Analizę fitosocjologiczną przeprowadzono według zasad szkoły Braun- -Blanęueta, przystosowanych do badań porostów przez K le m e n ta (18) i B a rk m a n a (2). Stwierdzono występowanie 7 zespołów porostów epifitycznych: Chaenothecetum ferrugineae, Hypocenomycetum scalaris, Pertusarietum amarae, Pyrenuletum nitidae, Thelotremetum lepadini, Parmeliopsidetum ambiguae i Pseudevernietum furfuraceae. Key w ords: epiphytic lichens, phytosociology, Knyszyn Forest. 142 KATARZYNA KOLANKO

INTRODUCTION

Togethei\with progress in phytosociology of vascular plants investigations of lichen communi- ties began. M o ty k a (21), G la n c (16), Z ie liń s k a (26), F a b is z e w s k i (14), C ie ś liń s k i and H a lic z (11), B y s tre k (4) and B ie lc z y k (3) were engaged in research into associations of epiphytic lichens in Poland. Phytosociological studies of epiphytic lichens in the Knyszyn Forest were carried out in the following reserves: Budzisk (5), Karczmisko (6) and Jesionowe Góry (9). The aim of investigations presented in this work is phytosociological characteristics of epiphytic lichens of the Knyszyn Forest. The Knyszyn Forest is situated north-east of Białystok and covers a larger part of Wysoczyzna Białostocka formed in a considerable degree from sands of glacial accumulation. Its sculpture is exceptionally varying. It is characterized by numerous morphological levels. There are distinguished three zones of sculpture (1): moraine height surrounding the Forest from the north and east; western and central part of the Forest with characteristic isolated ground eminences surrounded by wide depressions; and the zonę south of the valley of the Supraśl river, where the ground is mildly formed and rises from 150 m to 180 m above sea level. The Forest encompasses mainly the river-basin of the Supraśl. The part which farthest of all advances to the west belongs to the direct drainage area of the Narew. In some regions the Forest area encroaches on the drainage area of the Nietupka and the Nietupa, the tributary of Swisłocz in the river-basin of the Niemen, while from the north the drainage areas of the Brzozówka and the Sidra (tributaries of the Biebrza) appear on the territory of the Forest. The climate is typical of north-eastern region of Poland (19, 23), and its Continental character determines the length of the growing season. In addition, the climatic conditions occurring in the Knyszyn Forest are ditferentiated due to the varying sculpture of the ground, influence of the Białystok agglomeration, and proportionally large regions of forest areas. Over 60% of forest surfaces of the studied region area are occupied by coniłerous forest communities. 21-60 years old stands, whose participation amounts to approximately 47%, pre- dominate in the Forest. Old trees, over 100 years old, are not many; they cover just up to 4% of the area (15). C z e rw iń s k i (12) described 15 forest communities and several supplementary communities from the Knyszyn Forest.

MATERIAŁ AND METHODS

Outdoor and phytosociological investigations were carried out on 262 stands distributed uniformly enough over the whole studied area. Over 400 phytosociological records were performed, out of which 123 were used for elaboration of epiphytic lichens communities. Phytosociological analysis was madę following the method of Braun-Blanquet, which was modified and adapted for the reaearch into lichens by K le m e n t (18) and B a rk m a n (2). Phytosociological investigations were carried out on bark of pine, spruce, oak, birch, hornbeam, alder, linden and mapie. A list of of phytosociological records stands: 1. Budzisk, deciduous forest; 10. Remuczewo; deciduous forest; 11. Trzy Słupki; deciduous forest; 17. Sokole, mixed forest; 19. Majdan, 1 km to S; 20. Żednia, spruce forest; 21. Road Zednia-Michałowo; 24-25. Wzgórza Świętojańskie, mixed forest, 27. Pieszczaniki, 0.5 km from the village; 29. Pieszczana Góra, mixed forest; 30. Lipowy Most, pine-spruce forest; 33. Kopna Góra, 1 km to SE of arboretum; 38. Las Cieliczański, deciduous forest; 39. Cieliczanka, spruce forest; 41. EPIPHYTIC L1CHEN-DOMINATED COMMUNITIES IN THE KNYSZYN FOREST 143

Supraśl, mixed forest; 50. Supraśl, mixed forest; 52. Pólko, Krasne, mixed forest; 61. Wierzchlesie, 2 km to W of the village; 63. Międzyrzecze, deciduous forest; 64. Woronicze, deciduous forest; 69. Jesionowe Góry, deciduous forest; 70. Machnacz, station; 73. Góry, mixed forest; 78. Starodrzew Szyndzielski, deciduous forest; 82. Przewalanka, spruce forest; 84. Podkrzemianka, mixed forest; 85. Karczmisko, mixed forest; 86. Katrynka, spruce forest; 88. Kopisk, near the village; 91. Kopisk, mixed forest; 103. Obrubniki, birches; 111. Zalesie, mixed forest; 112. Bobrowa, mixed forest; 116. Stara Dębina, deciduous forest; 125. Zielona, mixed forest; 129. Czołnowo, deciduous forest; 131. Sosnowik, mixed forest; 145. Kol. Mostek, mixed forest; 152. Krasny Las, mixed forest; 160. Talkowszczyzna, spruce-pine forest; 175. Stara Grzybowszczyzna, mixed forest; 179. Górany, birches; 187. Michalowo-Kolonia, spruce forest; 188. Kopce, mixed forest; 197. Lebiedzin, spruce forest; 198. Lebiedzin, old pines; 201. Kumiałka, 2 km to SE of the village; 202. Kumiałka, spruce-pine forest; 208. Cimoszka, mixed forest; 220. Kwasówka, 1.5 km to E; 235. Leśniczówka Mostek, mixed forest; 242. Szaciły, pine-spruce forest; 245. Surażkowo, spruce forest; 249. Ośrodek, mixed forest; 255. Sochonie, pine forest; 258. Woroszyły, mixed forest; 262. Czarny Blok, spruce forest.

RESULTS AND DISCUSSION

In the Knyszyn Forest there were ascertained 7 associations of epiphytic lichens and some communities, which cannot rank among associations. Classification of lichens communities in the Knyszyn Forest is as follows: I. Order; Leprarietalia Barkman 1958 emend. W irth 1972 Alliance: Calicion viridis (H ad a 1944) emend. B a rk m a n 1958 Association: Chaenothecetum ferrugineae Barkman 1958 II. Order: Lecanoretalia variae Barkman 1958 Alliance: Lecanorion variae Barkman 1958 Association: Hypocenomycetum scalaris H ilitzer 1925 III. Order: Arthoniethalia radiatae Barkman 1958 Alliance: Graphidion scriptae Ochsner 1928 emend. Barkman 1958 Association: Pertusarietum amarae H i 1 i t z e r 1925 emend. B a rk ­ m an 1958 Association: Pyremdetum nitidae H ilitzer 1925 Association: Thelotremetum lepadini H ilitzer 1925 IV. Order: Parmelietalia physodo-tubulosae Barkman 1958 Alliance: Parmelion saxatilis Barkman 1958 Association: Pseudevernietum furfuraceae ( H ilitz e r 1925) Ochsner 1928 Association: Parmeliopsidetum ambiguae H ilitzer 1925 Subass.: Parmeliopsidetum ambiguae-Parmeliopsidetosum aleu- ritis Barkman 1958 144 KATARZYNA KOLANKO

Chaenothecetum ferrugineae B a r k m a n (2) (Tab. 1) occurs in the investi- gated area only on spruce bark. It is most often located on lower parts of trunks and avoids habitats with immediate effect of sunshine. It occupies smali surfaces in places which are not exposed to an immediate flow of rain water. It locates on shell segments of bark as well as on crevices between them. In plots of this association dominates Chaenotheca ferruginea, the only characteristic species of this group, and together with other leprose lichens determines its physiognomy. In the Knyszyn Forest it occurs in fragments of natural, decimated woods. There are some differences in comparison with afore-published descriptions of the Forest (5, 6). They refer to the number of species and coverage degree. Chaenothecetum ferrugineae is a widespread association, common on the lowland of Europę in the subalpine zonę. The community is ombrophobus, aerohygrophilous, acidophilous and photophilous, but avoids habitats with the immediate effect of sunshine (2, 26). Chaenothecetum ferrugineae in Poland was recorded in similar habitats and described, among others, in the works of G 1 a n c (16), Z i e li ń s k a (26), C i e ś li ń s k i and H a lic z (11), B y s tr e k et al. (4, 5, 6, 7, 8) and B ie lc z y k (3). Hypocenomycetum scalaris H i 1 i t z e r (17) (Tab. 2) develops on bark of trees growing on borders of forests and clearings. It occupies the base and lower parts of trunks of birch and pine. The forming species locate mostly in bark crevices. Floristically it is a poor association. Hypocenomyce scalaris grows in masses within this association and achieves high degrees of coverage and the highest constancy. Among the accompanying species worth noting is the participation of the species of the order Parmelietalia physodo-tubulosae. The occurrence of one- -species plots of Hypocenomyce scalaris is a generał phenomen. Such populations are not included in the association. Hypocenomycetum scalaris is remarkably photophilous community. K 1 e m e n t. (18) considered Hypocenomycetum scalaris as photophilous, xero- and acidiphilous. The association from the Knyszyn Forest described in the paper does not essentially differ from the communities in other regions of Poland. It occurs on similar habitats and has the same appearance. Most of all it approximatesHypocenomycetum scalaris from the Kampinos Forest, described by Z i e li ń s k a (26). Hypocenomycetum scalaris from the Świętokrzyskie Mts., described by C ieśliński and H alicz (II), is richer in species. Special attention should be focused on the occurrence of the following several species of this association, growing in patches. These are: Bryoria crispa, Platismatia glauca, Usnea hirta from the order Parmelietalia physodo-tubulosae. In this association described from the Beskidy Mts, by B ielczyk (3) the following species do not grow: Lecanora expallens, L. varia and Pseudevernia furfuracea. Information about the occurrence of this association in Poland can be found also in ANN. UNIV. MARIAE CURIE-SKŁODOWSKA, sectio C, vol. LVI T a b l . I

Fig. 1. Pertusarietum amarae

Fig. 2. Pyrenoletum nitidae

Katarzyna Kolanko ANN. UNIV. MARIAE CURIE-SKŁODOWSKA, sectio C, vol. LVI Tabl. II

Fig. 3. Thelotremetum lepadini

Fig. 4. Pseudevernietum f urfuraceae Katarzyna Kolanko EPIPHYT1C LICHEN-DOMINATED COMMUNITIES IN THE KNYSZYN FOREST 145

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ULIOJ qiMOJX) C f i <2 3 f i f i o* 03 o CU CU 03 EPIPHYTIC LICHEN-DOMINATED COMMUNITIES IN THE KNYSZYN FOREST 147 the works of M attick (20), Glanc (16) and Piaczyńska (24). According to Barkman (2) it is a mesophilous and acidddophilous community. Pertusarietum amarae H i l i t z e r 1925 (17) emend. B a rk m a n 1958 (2) (Tab. 3, Phot. 1) occurs on bark of hornbeam, linden and mapie. Floristically it is a rich community. Characteristic species are at the same time dominating species. First of all, abundantly sorediose lichens of the genus Pertusaria (13) occur there. They determine the physiognomy of the association. Pertusarietum amarae is a moderately photophilous association which prefers the Southern side of trunks. Floristic composition of the association on the particular phorophytes is similar, but the best developed and most differentiated plots can be found on bark of hornbeam. In the Knyszyn Forest Pertusarietum amarae occurs exclusively in highly natural deciduous communities. Pertusarietum amarae was described by H i l i t z e r in 1925 (17). O c h s n e r (22), B a rk m a n (20), W ilm a n n s (25), G la n c (16) described it under different names, completing the list of typical and distinguishing species at the same time. The comparative analysis of Pertusarietum amarae over the territory of Poland is also difficult, considering the diversity of phorophytes on which it was found. In the natural beech forests in Roztocze (4), the Świętokrzyskie Mountains (11) and the Beskidy Mountains (3) there occur characteristic species on bark of beech, which are not present on other phorophytes. C i e ś li ń s k i and H a lic z (11) suggest that we have to do with the association which is regionally differentiated to smaller units. As regards the studied area the information given by C ieśliń sk i and H alicz (11) about its sensitivity to dust and air pollutions is confirmed. Plots of this association in the Forest differ from those described in the literaturę, first of all in the rangę of characteristic species. Within the investigated area these are: Pertusaria amara, Phlyctis argena and Pertusaria coccodes, just as in the Świętokrzyskie Mountains (11). Among the characteristic species of this community in Roztocze there is also Pertusaria albescens (4) and in Wigry National Park also Pertusaria coccodes var. phymatodes and P. albescens var. globulifera (8). F a b is z e w s k i (14) records Menegazzia terebrata as a local characteristic species, which in the Knyszyn Forest is a rare species. Besides Pertusaria amara and Phlyctis argena B i e 1 c z y k (3) mentions Pertusaria hemisphaerica among characteristic species. Pertusarietum amarae from the Knyszyn Forest was described earlier, from the reserves B u d z is k (5) and Jesionowe Góry (9). Pyrenuletum nitidae H i l it z e r 1925 (17) (Tab. 4, Phot. 2) develops on the studied area in a typical form on hornbeam and on dying, often protruding bark of hazel, forming multicoloured mosaic. It is formed from crustaceous, nonsore- diate lichens with numerous apothecia, including the species with perithecia. Specific appearance of this association is characteristic of the species of the 148 KATARZYNA KOLANKO families Graphidaceae, Pyrenulaceae and Pertusariaceae. The community occu- pies smali surfaces, mainly in the eastern and north-eastern side of trunks. The highest coefficients of coverage and constancy are achieved by the characteristic species Graphis scripta, Pyrenula nitida and Pertusaria leioplaca. On hornbeams Pyrenuletum nitidae develops among the plots of Pertusarietum amarae. This as- sociation grows far inside the primeval broad-leaved forest. Pyrenuletum nitidae was earlier described from Budzisk (5) on bark of mapie and ash as well as from Jesionowe Góry (9). New localities found in the Knyszyn Forest differ in floristic composition, first of all among the accompanying species. B a rk m a n (2) as well as C i e ś li ń s k i and H a lic z (11) consider Pertusaria leioplaca a typical species of the alliance Graphidion scriptae. In Jesionowe Góry it was placed among the accompanying species due to a lower degree of coverage and also poorly formed thalli, difficult for identification. M o ty k a (21) described this community from the Western Beskidy Mountains, G lanc (16) — from the experimental forest inspectorate Zielonka near Poznań. F ab iszew sk i (14) characterized that association as one of poorer, overgrowing broad-leaved trees. According to C ie ś 1 iń s k i and H a lic z (11) Pyrenuletum nitidae is a frequent association in the communities Abieteto-Fagetum and Abietetum polonicum with addition of beech. B y s tre k et al. (5, 9) recorded this species from bark of hazel and hornbeam in oak-linden-hornbeam forests, B i e 1 c z y k (3) — from bark of beech and mapie. Pyrenuletum nitidae is cited from various regions of Europę (2, 17, 18, 22, 25). Differences in floristic composition stem from different geographical, habitat and microclimate factors. Thelotremetum lepadini H i 1 i t z e r (17) (Tab. 5, Phot. 3) is a typical epiphytic association. It develops wide patches on bark of hoary hornbeams and lindens, located mainly in the north and north-eastern side of the trunk. Its occurrence is restricted to primeval forests. The coverage of the plot by thalli achieves 100%. The physiognomy of Thelotremetum lepadini is determined by crustaceous lichens. It is a moderate photophilous and hygrophilous association. Thelotrema lepadinum is the decided dominant of this community. At the same time it is a typical species. Graphis scripta and Opegrapha viridis achieve high degrees of constancy. Coefficients of other species coverage are Iow. Thelotremetum lepadini in the Knyszyn Forest occurs only in the reserve Budzisk. In comparison with the patches earlier described from this locality (5) there is a distincly marked difference in the number of forming them species. Now the plots are poorer even by about 50%. Equally distinct is the second difference — an increasing surface occupied by Thelotrema lepadinum. This may suggest the expansion of this species within the plot. The association from the Knyszyn Forest also proves differences in relation to the plots described by B ielczyk (3) from the Western Beskidy Mountains. As a typical species she records, among Tab. 3. Pertusarietum amarae Hil. 1925 emend. Barkman 1958

Number of locality 1 1 10 10 11 11 63 63 64 69 78 78 78 116 116 116 116 129 Tree species Cb Tc Tc Cb Tc Cb Cb Tc Cb Cb Tc Cb Cb Cb Cb Tc AP AP - J§ 1 Tree diameter m 0.5 0.2 0.2 0.4 0.3 0.4 0.4 0.3 0.4 0.5 0.2 0.5 0.4 0.5 0.4 0.2 0.3 0.4 Heigit on trunk m 1.5 1.4 1.2 1.2 1.5 1.6 1.4 1.6 1.5 1.7 1.3 1.1 1.6 1.2 1.3 1.6 1.5 1.3 Surface of record dm2 12 6 6 9 8 12 8 8 12 15 6 16 12 12 9 6 9 8 Exposure of plot N E E E S E E S SE SE E SE SW S S S‘ SW W Number of species 12 10 9 8 6 11 12 10 10 12 9 7 11 9 10 8 10 6 Characteristic species ofthe association Pertusaria amara 2 1 2 1 2 2 1 2 2 2 2 2 1 2 2 2 2 2 V LP Phlyctis argena + 4- 1 + 4- 4- 1 4- 4- 4- III I ł Pertusaria coccodes + 1 + 4- 4- 4- II Characteristic species ofthe alliance Graphidion scriptae Cr Graphis scripta 1 2 1 1 1 4- 1 1 1 1 1 1 4- 1 .4- 1 1 V Cr Opegrapha rufescens + + + + + + 4- 4- 4- III Cr Lecanora argentata + + + + 4- 4- 4- 4- III Cr Pyrenula nitidella + + + -i- + + 4- 4- 4- III Cr Pertusaria albescens v. globulifera + + + 4- II Cr Opegrapha atra 1 4- + 1 + 4- + 4- 4- III III Cr Lecanora intumescens + 1 + + 1 4- 1 1 Characteristic species of the order Arthonietalia radiatae Cr Pertusaria pertusa + 1 1 4- 1 II Cr Lecanora carpinea + + + 1 1 4- 1 4- 4- 4- III Cr Lecidella elaeochroma 1 1 + 1 4- 1 1 1 4- III Cr Arthonia radiata + 4- 4- 4- II Characteristic soecies of the order Parmelietalia physodo-tubulosae Pa Hypogymnia physodes + + 4- 4- 4- II Pa Platismatia glauca + + + 4- 4- 4- 4- II Aocompanyine species Pa Melanelia fiiliginosa + + 4- 4- 4- + 4- II Pa M. exasperatula + + 4- 4- 4- II Ra Evernia prunastri + 4- 4- I Cr Lecanora pulicaris + 4- 4- 4- II Lepraria incana + + 4- I Pa Parmelia sulcata + 4- . I I Ra Ramahna farinacea + 4- IT Cr L. allophana + 4- iT Lp L. chloropolia 4- TI Cr Arthothelium ruanideum + 1T Lp Ochrolechia microstictoides + Tab. 4. Pyrenuletum nitidae Hil. 1925

Number of locality 1 1 1 10 11 11 11 63 63 64 69 78 78 78 116 116 116 116 (1 Tree species Cb Ca Cb Cb Cb Ca Ca Ca Cb Cb Ca Ca Ca Cb Cb Ca Ca Ca Tree diameter m 0.4 0.3 0.4 0.5 - 0.3 0.4 - - - 0.3 0.4 - - - Heigfrt on trunk m 1.3 1.5 1.7 1.1 - - 1.5 1.3 - - - 1.4 1.6 - - - Surfece of record dm2 4 2 4 6 4 2 2 3 2 4 2 4 2 4 4 3 3 2 Exposure of plot NE SE E NE - - SW NE - - - SE NE - - - Number of species 10 8 6 9 6 5 6 8 7 6 6 6 5 6 7 7 5 5 Characteristic species of the associatian Cr Pyrenula nitida 2 + 1 1 1 1 1 1 2 1 1 1 2 1 1 1 1 1 V Cr Graphis scripta 2 2 3 2 3 2 1 2 2 3 1 1 1 3 2 2 1 2 V Cr Pertusaria leioplaca + + 1 + + + 1 + 1 1 III Characteristic species ofthe allianoe Graphidion scriptae Cr Opegrapha viridis 1 1 + 1 + + + 1 1 + 1 IV Cr Pertusaria pertusa 4- + 1 + + 1 + + III Cr Opegrapha atra + I Cr O. rufescens + -i- + + + + 1 + + III Cr Thelotrema lepadinum + + I Characteristic species of the order Arthonietalia radiatae Cr Arthonia radiota + 1 + + + 1 + + III Cr Lecidella elaeochroma + 1 1 1 + + + + + 1 III Acoompanying species Lp Phlyctis argena + + + I Cr Arthothelium ruanideum + + + + II Cr Lecanora carpinea + + + + + + + II Łp Pertusaria coccodes + + + I Cr Porina aenea + I Pa Hypogymnia physodes + + I Lp Lepraria incana + + + I EP1PHYTIC LICHEN-DOMINATED COMMUNITIES IN THE KNYSZYN FOREST 149

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others, Menegazzia terebrata, which on the examined area occurs at individual localities. Thelotremetum lepadini is a community not by all distinguished. It has many species that are common with Pyrenuletum nitidae. High degrees of constancy and distinctly prevailing Thelotrema lepadinum are the reason why this community is treated as an independent association. B a rk m a n (2) described it as strongly aerohygrophytic species, which needs clean, moist air to develop. In this connection its occurrence is restricted to the best preserved primeval species. Parmeliopsidetum ambiguae H i l it z e r 1925 (17) (Tab. 6) in the Knyszyn Forest is represented by only one of several subassociations distinguished in Europę — subass. Parmeliopsidetosum aleuritis. In the studied area it grows on bark of spruce and pine in light places, on borders of forests, near glades and forest roads. Its patches develop at the base of trunks, often on the northern side. Imshaugia aleurites is a typical species of the lowland variety of the association Parmeliopsidetum ambiguae. The highest degree of constancy and coverage achieve typical species of Imshaugia aleurites and Parmeliopsis ambigua. The participation of crustose and leprose lichens is big. The plots of Parmeliopsidetum ambiguae in the Knyszyn Forest lack char- acteristic species such as Tuckermanopsis sepincola and Parmeliopsis hyperop- ta recorded from mountainous regions (3, 14). However, Imshaugia aleurites, a species which characterizes well this association was recorded from lowlands (16, 26) and the Świętokrzyskie Mountains (11). Distinctly marked is the pres- ence of Hypogymnia physodes, and some plots of Tuckermanopsis chlorophylla. In comparison with the plots described earlier from the examined area (5, 6) no significant differences were found. B a rk m a n (2) and K 1 e m e n t (18) stated that this association, developing above all in mountainous and piedmont regions was on lowlands replaced by Hypocenomycetum scalaris. B a rk m a n (2) and F a b is z e w s k i (14) also characterized Parmeliopsidetum ambiguae as chinophylic species, i. e. locating in basal parts of trees. According to B a rk m a n (2), this fact may be explained by avoidance of the coldest habitats. F a b is z e w s k i (14), however, considers morę probable the fact that humidity of bark in basal parts of trunks is the highest. The association prefers bark of the conifers (3, 11, 14, 26), although K le m e n t (18) described it also on rough-barked deciduous trees. Pseudevernietum furfuraceae H ilitzer (17), Ochsner (22) in the Kny­ szyn Forest (Tab. 7, 8, Phot. 4) is the most common, but also the only association predominated by macrolichens. It occurs in pine-woods and mixed woods on bark of pine, birch, morę rarely on spruce bark as well as on their dead, non- decorticated branches. In oak-linden-hornbeam forests it develops abundantly on iao. o. Parmeliopsidetum ambiguae Hil. 1925 subass. Parmeliopsidetosum aleuritis3axkxnx& 1958

Number of locality 11 21 29 33 39 41 53 63 64 84 91 112 125 145 175 202 208 Tree species Pe Pe Pe Ps Pe Pe Ps Pe Ps Pe Ps Pe Pe Ps Pe Pe Ps & Tree diameter m 0.5 0.4 0.4 0.4 0.4 0.5 0.4 0.4 0.5 0.4 0.4 0.5 0.5 0.5 0.4 0.4 0.4 Heigfrt on trunk m 0.5 0.3 0.6 0.6 0.6 0.5 0.5 0.8 0.5 0.4 0.6 0.5 0.6 0.8 0.3 0.6 0.6 1 1 Surfaoe of reoord dm2 3 4 8 4 6 6 3 4 4 8 4 8 6 6 6 6 4 <5 Exposure of plot NE SW W N S S S NE E NE N N NE N SW N N Number of species 5 4 5 6 6 5 6 4 6 4 6 4 5 4 4 5 4 Characteristic species ofthe association P Parmeliopsis ambigua 1 2 2 2 2 1 1 2 2 1 2 1 2 1 1 2 1 V P Imshaugia aleurites + 1 1 2 2 + 1 + 1 2 1 1 IV P Vulpicida pinastri + I Characteristic species of the alliance Parmelion saxatilis Pa | Tuckermanopsis chlorophylla L L L± L± L □ • r r r RR n n ri nL L L L!!! Characteristic species ofthe order Parmelietalia physodo-tubulosae Pa 1 Hypogymnia physodes 1 2 . 2 L 2 1 2 J l 2 J RR r r H 1 2 | 2 |I? | 2 2 2 L? 1 i - 1 V Accompanying species Lp Lepraria incana + + + -i- + + + '+ + + III Lp Lecanora conizaeoides + + + + II Sq Cladonia coniocraea + + + + + + + III Sq Hypocenomyce scalaris + + + + + + + III Lp Chaenotheca ferruginea + + + I Lp Lecanora expallens + + I Sq Cladonia chlorophaea + + I

Tab. 7. Pseudevemietum furfuraceae (Hil. 1925) Ochsn. 1928

Number of locality 10 20 24 29 53 82 85 116 116 129 131 152 188 198 202 242 249 255 258 Tree species Pe Pe Pe Pe Pe Pe Ps Ps Ps Ps Ps Ps Pe Pe Ps Pe Ps Ps Ps (1 i Tree diameter m 0.2 0.2 0.2 0.4 0.4 Ó.4 0.4 0.3 0.5 0.4 0.3 0.4 0.4 0.4 0.5 0.4 0.3 0.4 0.4 Heigfrt of trunk m 1.7 1.5 1.5 1.6 1.6 1.7 1.5 1.7 1.7 0.9 1.2 1.5 1.4 1.6 1.9 1.6 1.8 1.6 1.6 Surfaoe of reoord dm2 4 4 3 8 8 9 12 9 16 12 12 9 8 6 16 8 8 12 9 Exposure of plot S SW S SE S W S S W SE SW N SW SW SW S S S SE Number of species 9 4 4 9 4 5 7 7 5 4 7 5 5 6 5 4 3 5 5 Characteristic species of the association An Pseudevernia furfuracea 2 1 1 2 1 1 2 1 2 2 2 2 1 1 1 2 2 2 1 V Pa Hypogymnia tubulosa + + + + + II Characteristic species ofthe alliance Parmelion saxatilis Us 1 Usneahirta RR RR r RR r r r RR RR RR RR RR RR RR RR RR RR RR RR 1 n Characteristic species ofthe order Parmelietalia physodo-tubulosae Pa Hypogymnia physodes 3 3 2 3 3 3 2 3 2 3 3 3 2 3 2 3 2 1 3 V Pa Platismatia glauca + 1 + 1 1 + 1 + + + + + + 1 IV Pa Tuckermanopsis chlorophylla 1 + + + + + + + III Us Bryoria implexa + + . I Us II Usnea tuberculata + 1 + + + Pa II Parmelia sulcata -i- + + + Accompanying species Lp Lecanora conizaeoides + + + + + + . + III Cr L. varia + + + + + + + II Lp Lepraria incana + + + I J p Scoliciosporum chlorococcum + + I Tab. 8. Pseudevemietum furfuraceae (Hil. 1925) Ochsn. 1928

Number of locality 10 19 29 125 38 52 63 86 63 78 125 39 78 129 131 160 179 254 Js Tree species qr BP BP Bp Ag Ag Ag Ag Bp Bp BP Bp Bp Tree diameter m 0.3 0.4 0.4 0.3 0.5 0.2 0.5 0.2 0.3 0.3 0.2 0.4 0.4 0.3 0.4 0.3 0.3 0.4 Heigjit on tnmk m 1.6 1.5 1.6 1.7 1.5 0.8 1.8 1.1 1.6 1.6 1.7 1.4 1.7 1.5 1.8 1.5 1.6 1.5 Surface of record dm2 9 12 12 12 16 6 16 6 9 12 8 9 15 12 12 8 12 15 Exposure of plot Constancy SE S SW SE SE E S NE N SW E S SE S E SE S SW Number of species 10 10 10 6 8 10 9 8 5 8 9 5 9 11 6 6 7 5 Charactcristic species of the associations An Pseudevemia furfiiracea 1 2 1 + 2 1 1 1 2 1 1 2 1 1 1 1 1 1 V Pa Hypogymnia tubulosa + + + + + + + + + + III Charactcristic species of the variant Ra Ramabna farinacea + + + 2 2 2 2 Ra Evemia prunastri + 2 2 2 2 1 + II II Charactcristic species ofthe alliance Parmelion saxatilis P Parmeliopsis ambigua + + I Us Usnea hirta + + 4- I P Yulpicida pinastri + I Charactcristic species of the order Parmelietalia physodo-tubulosae Pa Hypogymnia physodes 3 3 3 2 2 3 3 3 2 1 2 3 3 2 3 2 2 3 V Pa Platismatia glauca 1 + + 1 1 + + 1 1 + + 1 1 IV Us Usnea tuberculata 2 2 2 1 + + + + III Pa Tuckermanopsis chlorophylla + + + + + + 1 1 1 + III Us Bryoria crispa + I Us Br. implexa + + + I Pa II Parmelia saxatilis + + + + + Pa II P. sulcata + + + + + Us I Bryoria positiva + + Aocompanying species Lp Lecanora expallens + + I Lp Scoliciosporum chlorococcum + + + I Cr Lecanora varia + + + + + II Lp Lepraria incana + + I Lp Lecanora conizaeoides + + + + II Pa Melanelia exasperatula + + I Pa I M. fubginosa + + + Cr II Lecanora pulicaris + + + Us II Usnea subfloridana + Cr Lecidella elaeochroma I + + Lp Phlyctis argena I + I I EPIPHYTIC LICHEN-DOMINATED COMMUNITIES IN THE KNYSZYN FOREST 151

bark of oak. It also grows on bark of mapie and alder. Plots and species of this association also occur in other habitats (fences, roofs, balustrades), and the distinguishing species belong to pioneer ones in the communities and on each phorophyte. The initial stages of Pseudevernietum furfuraceae form as early as in young pine forests, but on very smali plots. In older forests the structure of association and the occupied area depend on microhabitat conditions, species and age of the tree as well as on the height at which it grows on the trunk. On pine and spruce this association covers smali surfaces and is poor in species. Hypogymnia physodes distinctly dominates and achieves high degrees of coverage. On bark of deciduous trees the association is floristically richer and covers larger surfaces. There are distinguished varieties with large participation of Evernia prunastri (stands: 38, 52, 63, 86) and Ramalina farinacea (stands: 63, 78, 125). On birch the presence of rare in Poland species Bryoria and Usnea (stands: 10, 19, 29, 125). Usnea tuberculata forms large populations there. Pseudevernietum furfuraceae is a very common community, often described in the literaturę (2, 17, 18, 22, 25). It is generally considered nitro-, conio- and toxophobous, because it is formed far from towns and communication routes. It is photophilous and needs high air humidity. According to most of the researchers it lacks typical species (2, 18, 22). Instead, a combination of the species occurs: Pseudevernia furfuracea, Hypogymnia physodes, H. tubulosa, Tuckermanopsis chlorophylla, Platismatia glauca, Usnea and others. hirta First information about Pseudevernietum furfuraceae in Poland was published by M o ty k a (21). Among common associations growing on various phorophytes this association is men- tioned by G1 anc (16), Zielińska (26), Fabiszew ski (14), C ieśliński and H a lic z (II), B y s tre k et al. (4, 5, 6, 7, 8), B ie lc z y k (3). In comparison with the earlier described patches of this association on the studied area (5, 6) the present data indicate its impoverishment. Extinct or endangered are the following characteristic species: Bryoria crispa, Br. implexa, Br. subcana, Evernia mesomorpha, Hypogymnia and tubulosa some Usnea.

CONCLUSIONS

The associations Pertusarietum amarae, Pyrenuletum nitidae and Thelotreme- tum lepadini are the communities characteristic of natural deciduous forests and should be taken into consideration in the estimation of the degree of naturalness of forest communities. The most common association in the Knyszyn Forest is Pseudevernietum furfuraceae, with wide ecological amplitudę. Differentiation of this species, both in the examined area and in other regions of Poland depends on microclimatic 152 KATARZYNA KOLANKO factors, species and age of the tree, naturalness degree of forest communities and anthropogenic factors. Acknowledgements. I wish to thank Professor Jan Bystrek for his revision of the species Bryoria and Usnea as well as for his valuable advice in the course of my work on the study. This study was financially supported by the Committee for Scientific Research (grant 6PO4C 03 612) realized in Institute of Biology, University in Białystok.

REFERENCES

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PHOTOGRAMS

Phot. 1. Pertusarietuin amarae. Phot. 2. Pyrenuletum nitidae. Phot. 3. Tholotremetum lepadini. Phot. 4. Pseudevernietum furfuraceae.