;. Polish Botanical Studies 22: 53-62, 2006

DISTRIBUTION AND PHYTOCOENOTIC CHARACTERISTICS OF RELICT POPULATIONS OF RHODODENDRON MYRTIFOLIUM (ERICACEAE) IN THE UKRAINIAN CARPATHIANS

ADAM BORATYNSKI, MARCIN PIWCZYNSKI, Y AKIV P. DIDUKH, LIDIA T ASENKEVICH, ANGEL ROMO & HALINA RATYNSKA

Abstract. Rhododendron myrtifolium Sehot & Kotsehy is a high-mountain plant dispersed at several reliet Pleistocene localities in isolated massifs ofthe Carpathians and in the Balkan mountains. The East Carpathians ofUkraine are the northemmost part of its range. This study assessed the current distribution and status of the species by surveying most of its Ukrainian locali­ tieso R. myrtifolium grows in subalpine and alpine belts of the mountains, reaehing the uppermost position at 2020 m a.s.l. on Pietros Mí. and at 2030 m a.s.l. on Hoverla Mí. in the Chomokhora Mts. In the subalpine vegetation belt it fonns the associa­ tion Rhododendretum myrtifolii or enters other closely related eommunities, mostly from the Rhododendro- Vaccinion allianee. Isolated localities outside the Chomokhora range are few and éndangered, especially in the ease of overgrazíng. One of them has disappeared and four others have not been found in the ñeld in reeent decades. The speeies is threatened by global wanning, espeeially at the lowest localities. Key words: Rhododendretum myrtifoííi, plant eonservation, plant geography, ehorology, ecology, phytoeoenology, threat, Erieaceae

Adam Boratyñski. Polish Academy ofSciences, Instítute 01Dendrology, Parkowa 5, PL-62-035 Kórnik, Poland; e-mail: borata@ man.poznan.pl Marcin Píwczyñski, Nicolaus Copernicus Universíty, Institute 01Biology, Gagarina 9, PL-87-100 Iorun, Poland Yakiv P. Didukh, National Academy ofSciences ofUkraine, M G. Kholodny Institute ofBotany, Tereschenkivska 2, 01601 Kyiv, Ukraine: e-mail: [email protected] Lidia Tasenkevich, State Museum ofNatural History, Teatralna 18, 290008 Lviv, Ukraine; e-mail: [email protected] Angel Romo, Consejo Superior de Investigaciones Científicas, Instituto Botánico, Av. deis Muntanyans s/n, Pare de Montjudc, 08030 Barcelona, Spain; e-mall: [email protected] Halina Ratynska, Pedagogical Universíty, Institute ofBiology and Envíronmental Conservation, Chodkiewicza 30, PL-80-064 Bydgoszcz; e-mail: [email protected]

INTRODUCTION

Rhododendron myrtifolium Sehot & Kotsehy, to­ 1888; Komendar 1996). The authors assessed this gether with R. ferrugineum L. and R. hirsutum L., species in severalloeations in 1997-2004. This belong to the series Ferrugineum, section and paper presents the distribution ofR. myrtifolium, subgenus Rhododendron, genus Rhododendron describes the conditions of its occurrence, and (Popova 1972; Davidian 1982; Seithe 1987; analyzes threats. Chamberlain el al. 1996; Chamberlain and Hyam 1998; Kurashige et al. 2001). These three speeies METHODS oeeur in the mountains of Central and Southem Information relating to the distributionof Rhododendron Europe (Fig. 1). In the Ukrainian Carpathians, myrtifolium was collected, following a search ofthe lit­ Rhododendron myrtifolium is known mainly from eratureand herbaria.The generalrange given by Meusel the supraforestal Chornokhora mountain range el al. (IQ78) was supplemented with more recent data (Srodoú 1948). Outside of this range it is arare (Anchev 1982). The distribution of the species in lhe and threatened plant speeies of Ukraine (Weigl Carpathians was analyzed on the basis of literature 54 POLISH BOTANICAL STUDIES 22. 2006.

(Barbarich 1957; Topa 1960; Beldie1967; Coldea 1991; (Meusel el al. 1978). Its range is divided into Komendar 1996) and the authors' field investigations. several isolated populations oceurring in partic­ The occupancy area was estimated foraltitudinal veg­ ular massifs (Fig. 1). The southemmost localities etation layers in particular mountain massifs. of R. myrtifolíum are in the Rila Planina Mts in Detailed data from herbarium materials (preserved Bulgaria (Meusel el al. 1978; Anehev 1982). The at CHER, KOR, KRA, KRAM, KW, KWHA, KWU, LW, LWD and UD; acronyms of herbaria follow Hol­ most elevated are in the Rila Mts, at an altitude mgren el al. 1990 and Vasser 1995)and literature(Re­ of ca 2500 m a.s.\. (Anehev 1982). hman 1873; Weig11888; Woloszczak 1888; Zapalowicz In the Carpathians it was reported from ele­ 1889; Jávorka1924;Kulczyúski el al. 1926; Pawlowski vated mountain ranges with developed subalpine 1937; Barbarich 1957, 1987; Chopik 1976; Kotov and and alpine vegetation belts, mostly on Rumanian Chopik 1976; Malinovsky 1980; Komendar 1996) territory. It reaehes its altitudinal maximum in the were taken as the basis for species distribution in the Carpathians at 2300-2350 m in the Bueegi Mts Ukrainian Carpathians. Mostlocalities were verified in (Beldie 1967). The speeies range is ofreliet (gla­ the fieldduring 1997-2004.Extinctionand threat were eiation) eharaeter. evaluated for particular mountain massifs and for the entire Ukrainian Carpathians area according to IUCN Aeeording to IUCN eriteria (Anonyrnous 2002) criteria (Anonymous 2002). Plant nomenclature follows it falls in the endangered (EN) eategory beeause Mosyakin and Fedoronchuk 1999. its oeeupaney area eovers less than 500 km' and is severely fragmented, and the number ofloeations RESULTS AND DISCUSSION is steadily deelining.

GENERAL RANGE OCCURRENCE IN THE UKRAIN1AN CARPATHIANS

Rhododendron myrtifolium is restrieted to the high Most of the loealities of Rhododendron myrtifo­ mountains ofthe southem part ofthe Balkan Pe­ lium were reported from the Chomokhora Mts ninsula and to the South and East Carpathians (WeigI1888; Woloszezak 1888; Zapalowiez 1889;

Fig. L Range of Rhododendron myrtifolium Schot& Kotschy (black) andRhododendron ferrugineum L. (hachured) in Europe (after Meusel el al. 1978,supplemented). A. BORATYNSKI ET AL.: RHODODENDRON MYRTIFOLlUMIN THE UKRAINIAN CARPATHIANS 55

24'

":"'. v, '. , ,./23Ó ¡ ~,,_ .. 20 O 20 40 60km (/' ,J E3 - ';) 240

Fig. 2. Distribution of Rhododendron myrtifalium Sehot& Kotschy in the Ukrainian Carpathians.

Barbarich 1957; Chopik 1976). The herbarium ma­ II 00 m on the northem slopes of Smotrych MI. terials and data from the authors' field observa­ aboye Dzhembronya MI. (Piwczyríski 2000, in tions confirm that the most numerous populations sehedis), but the species has been observed only were in mountain ranges where the subalpine belt exceptionally at elevations below 1400 m. is well developed and alpine vegetation is fully formed (Appendix). The main center ofoccurrence ECOLOGICAL CHARACTER OF THE SPECIES is in the Chomokhora Mts (Fig. 2). Other localities have been reported from mountains having a suffi­ The occurrence of Rhododendron myrtifolium is ciently well formed subalpine vegetation belt, such related to north-facing slopes (Fig. 4), but not as Syvula MI., Popadia MI. and Syniak MI. in the so exclusively as in the case of alpine-subalpine Gorgany Mts (Woloszczak 1888) and Blyznytsa plants (e.g., Loiseleuria procumbens and Empetrum MI. in the Swydivec massif. The northernmost hermaphroditum; cf. Boratyríski & Didukh 2000, locality was reported from MI. Borzhava (Juzkiv 2002). It occurs on initial soils developed from & Ivanyckyj 1947, in schedis, LW), but later was siliceous rocks such as sandstones, porphyries believed to have disappeared (Chopik 1976). and crystal shales. This type of soil is shallow, Altitudinally, the greatest number of localities but under dense Rhododendron clumps an evident of Rhododendron myrtifolium have been found humus layer develops, consisting of undecom­ at elevations between 1500 and 1900 m (Fig. 3). posed fragments of shoots, leaves and discarded Altitudinal maxima were reported from 2030 m fruiting capsules of the species. It is acidic, with , on Khoverla MI. (Malynovs'ky & Kricsfalusy pH 3.5-4.9 (Malinovs'ky & Kricsfalusy 2000) . .2000), 2020 m on Pietros MI. (Shyshova 1955, R. myrtifolium individuals grow most frequently in sehedis), 2000 m at Pip Ivan in the Chomokhora in humid places protected against the wind, where Mts (Boratyúski 1997, in sehedis) and 1900 m erosion of humus is relatively limited. The north on Pip Ivan MI. in the Marmaros Mts (Suza ?, in exposures of most localities can be explained by sehedis). The lowest locality was reported from the plant's high demands in regard to soil mois- 56 POLlSH BOTANICAL STUDIES 22. 2006.

Khoverla t------1Il!------t20OO

~e-~-'f-\------+1500 ...... ••.•... • •• ...... ~.... •

..... 23° .....- .....·········24°·· ...... 25° ..

Fíg. 3. Altitudinal distribution ofRhododendron myrtifoliumSehot & Kotschy localities in the Ukrainian Carpathians. ture and humidity. The microrelief of localities degeneration stage of the Vaccinio myrtilli-Pin­ confirms these preferences as well. The species etum mughi. grows most frequently and abundantly in local Rhododendron myrtifolium forrns its own plant microdepressions of terrain, at the foot of cliffs, community composed mostly ofspecies ofthe Eri­ and on steep slopes with water outflows. caceae family, where it dominates or codominates Rhododendron myrtifolium has been reported, in the highest stratnm ofshrubs and shrublets, at­ with various degrees ofconstancy and cover, ftom taining a height of 15-25 cm. The indicator plants eight plant communities, mostly ofthe class Loise­ of Vaccinio-Piceetea (Vaccinium uliginosum, Ju­ /eurio-Vaccinietea Eggler ex Schubert 1960. It oc­ niperus communis subsp. alpina, Homogyne al­ curs in natural plant associations and enters into pina) and of high-mountain acidophilous grassy one anthropogenic one - Vaccinietum myrtylli, the communities of Juncetea trifidi classes (Juncus trifidus, Festuca airoides, Hieracium alpinum N subsp. a/pinum and C/adonia is/andica) are the 50 most common components ofthese plant assocía• tions (Coldea 1991; Malinovs'ky & Kricsfalusy 2000). The most typical phytosociological relevés ofRhododendretum myrtifo/ii consist ofca 20 spe­ cies on average, the number ranging from 14 to 37. In total, 41 species with frequency of at least Wí---+-~o07!(f---+-----)E 20% were observed in Rhododendretum myrtifolii. These data are very similar to those reported by Malinovs'ky and Kricsfalusy (2000). The frequent occurrence of R. myrtifolium in sw SE the Ukrainian Carpathians was also observed in associations of Empetro- Vaccinietum gaulterio­ ides, Pinetum mughi and Cetrario-Loiseleurietum S from the class Loiseleurio-Vaccinietea; Cetrario­ Fig. 4. Bxposition of Rhododendron myrtifoiium Sehot Juncetum trifidi and Primulo-Caricetum curvulae & Kotschy localities in the Chomokhora range, 00 the basis of 107datafrom herbaria, literature and authors' field observa­ from the class Juncetea trifidii; Hyperico grise­ tions (see Appendix); r = 50%. bachii-Ca/amagrostietum villosae from the class A. BORATYNSKI ET AL.: RHODODENDRON MYRTIFOLlUMIN THE UKRAINIAN CARPATHIANS 57

Betulo-Adenostyletea; Polytrichetum sexangularis, reduces the area available to high-mountain plant Festucetum picturatae and Salicetum retuso-re­ taxa, via the development of Picea abies and aboye ticulatae from the ciass Salicetea herbaceae; and the forest-line also Pinus mugo communities (Risch Senecio carpatici-Seslerietum bielzii from the et al. 2003; Dullinger et al. 2003, 2004). class Elyno-Seslerietea (Malinovs'ky & Kricsfa­ lysy 2000). ACKNOWLEDGEMENTS. The authors gratefully acknowl­ edge the help and valuable cornments provided by the curators andstaffofthe reviewed herbaria during col­ CONSERVATION lection ofthe material. Thestudy wasa cooperative sci­ entific and technological project ofthe Polish Academy The species Rhododenron myrtifolium, plant asso­ of Sciences and the National Academy of Sciencesof ciation Rhododendretum myrtifolii, and sorne other Ukraine, and was funded in part by the Institute of plant communities where it is also found (e.g., Ce­ Dendrology of the Polish Academy of Sciences. The trario-Loiseleurietum, Empetro-Vaccinietum) are material was prepared primarily by M. Piwczyáski as ciassified as rare and vulnerable in the Ukrainian his MSc dissertation at the Nicholas Copemicus Uni­ versity in Toruú. Carpathians (Kulczyúski et al. 1926; Wodziczko 1932; Shelyag-Sosonko et al. 1987; Khamor et al. 1994; Komendar 1996; Stoyko et al. 1998). REFERENCES The populations ofR. myrtifolium in the Ukrainian Carpathians are not numerous, except those from ANCHEV M. 1982. Ericaceae. In: D. JORDANOV (ed.), Flora the Chornokhora mountain range. All other locali­ Reipublicae PopularisBulgaricae 8: 288-301.Izdatelstwo na Bylgarskata Akademija naNaukite, Sofija. ties should be considered endangered, one oflhem in the Borzhava massif was extinct, and at least ANONYMOUS 2002. 2002 IVeN Red List of Threatened Species (Categories & Criteria, version 3.1). http://www. four others in the Gorgany Mts (Syvula, Popadija, redlist.org. Strymba, Steryshora) have not been confirmed in BARBARTCHA. 1. 1957. Ericaceae. In: M. 1. KOTOW &A. 1.BAR­ the field in the last three decades (Chopik 1976). BARICH (eds), Flora URSR 8: 29-59. Vidawnictwo Aka­ Following IUCN criteria (Anonymous 2002), the demij Nauk Ukrajns'koj RSR, Kijew (in Ukrainian). species should be considered critically endangered BARBARTCH A. 1. 1987. Ericaceae. In: Y. N. PROKUDlN & in Ukraine. For the reasons given, Rhododendron D. N. DOBROCHAEVA (eds), OpredeliteJ vysszikh rastenij myrtifolium was included in the 'Ukrainian red Ukrainy, pp. 134-136. Naukowa Dumka, Kijew.

data book' (Komendar 1996), and its plant associa­ BELDlE A. 1967. Flora ~i vegetatia muntilor Bucegi. Editura tions in the 'Green data book' (Shelyag-Sosonko Academiei Republicii Socialiste Romana, Bucuresti. et al. 1987). The rnain area of its occurrence in BORATYÑSKI A. & DIDUKH Y. P. 2000. The genusEmpetrum Ukraine is conserved within the Carpathians Na­ (Empetraceae) in Ukraine. Fragm. Florist. Geobot. 45: ture Reserve, but marginal ones are exposed to 423-430.

pasturing in grazed areas and could disappear. BORATYÑSKI A. & DIDUKH Y. P. 2002.Loiseleuría procumbens The plant associations ofRhododendron myr­ (Ericaceae) in the Ukrainian Carpathians. Dendrobiology tifolium can be easily degraded by grazing. Then 47:3~. they evolve into communities from alliances of CHAMBERLAIN D. F. & HYAM R. 1998. The genusRhododen­ Nardion in the subalpine and of Juncion trifidi dron: acasestudy to test thevalue ofvarious molecular tech­ in the alpine belts. Also, the absence of moderate niques asmeasures ofbiodiversity. In: A. KARP, P. G.ISAAC & D. S. INGRAM (eds),Molecular toolsfor screening biodi­ traditional pasture activity in the protected areas of versity, pp. 441-448. Chapman & Hall, London. , the Ukrainian Carpathians can accelerate invasions . oftrees and suppress R. myrtifolium, especially at CHAMBERLAIN D. F.,HYAM R.,ARGENT G., FAIRWEATHER G. & WALTER K. S. 1996. ThegenusRhododendron, itsclas­ its lower locations. sification andsynonymy. Royal Botanic Garden Edinburgh, Global warming should be seen as a factor in Edinburgh. the shrinking ofthe range ofRhododendron myr­ CHOPIK V. 1. 1976. Vysokogirna flora Ukrajns'kikh Karpat. tifolium. Climate warming shifts the treeline and Naukowa Dumka, Kiev. 58 POLISH BOTANICALSTUDIES 22. 2006.

CaLDEA G. H. 1991.Prodromedes associaciones végétalesdes MOSYAKIN S. L. & FEDORONCHUK M. M. 1999.Vascularplants Carpates de Sud-Bst(Carpates roumaineses). Documentes of Ukraine.A nomenclaturalchecklist. Nationa1 Academy Phytosociologiques, N.S. (Camerino) 13: 317-540. ofSciences ofUkraine, Kiev. DAVIDIAN H.H. 1982. TheRhododendron species. 1. Lepidotes. PAWLOWSKI B. 1937.Einfiihrung indie Pftanzenwelt derCzamo­ Batsford,London. hora in den Ostkarpaten.InstituteofBotany of Jagiellonian University,Krak6w. DULLINGER S., DIRNBOCK T. & GRABHERR G. 2003. Pattems oí shrub invasion into high mountain grasslands of the POPQVA T.N. 1972.Rhododendron L. In:T.G.1'uTJN, V.H. HEY­ Northern calcareous Alps, Austria. Arctic, Antarctic and WOOD, N. A. BURGES, D. M. MOORE, D. H. VALENTINE, Alpine Research35: 434-441. S. M. WALTERS & D. A. WEBB (eds), Flora Europaea. 3: 8-9. Cambridge University Press, Cambridge. DULLINGER S., DIRNBOCK T. & GRABHERR G. 2004. Model­ ling climate change-driven treeline shifts: relative effects REHMAN A. 1873. Materyaly do Flory wschodnich Karpat ze­ of temperature increase,dispersal and invasibility. J. Ecol. brane w r. 1871 i 1872.Sprmv. Komis.Fizjogr. 7: 1-39. 92: 241-252. RISCHA. C., NAGEL L. M., SCHÜTZ M., KRÜSI B. O., KIENAST HOLMGREN P. K., HOLMGREN N. H. & BARNEIT L. C. 1990. F. & BUGMANN H. 2003. Structureand long-termdeve1op-.­ Index Herbariorum. Ed. 8. New York Botanical Garden, ment ofsubalpinePinusmontanaMiller and Pinuseembra New York. L. forests in the Central European Alps. Forstwiss. Zen­ tralbl. 122: 219-230. JÁVORKA S. 1924. Magyar Flóra 2. Studium Kiadas, Buda­ pesto SEITHE A. 1987. Systematik und Nomenklatur bei Rhadoden­ dron.Rhododendron undimmergrune LaubgehO/ze (1987): KHAMOR F.D., VAINAGIY 1.V.&ANTOSYAK V.M. 1994.Protec­ 37-<55. tion situation with plants species from the red book of the Ukraine in the preserved territories of the Ukrainian Car­ SHELYAG-SOSONKO y. R., STOYKO S. M.,DUDUKH Y.P.,DUBYNA pathians. Ukrainsky BotanieheskyZhurnaI51(6): 122-129 D. V., ANDRIENKO T. L., TKACHENKO V. S. & BEZUS'KO (in Ukrainian with English summary). L. F. 1987.Ze1ena knigaUkrainskoySSR.NaukovaDumka, Kiev. KOMENDAR V. 1. 1996. Rhododendron kotsehyi Simonk. In: y. R. SHELYAG~SOSONKO (ed.), Chervona kntga Ukrajni, STOYKO S., MILKINA L. & YASHCHENKO P. 1998. Raritetni fi­ Roslinnij Swit, p. 112. Vidavnitstwo 'Ukrajns'ka enciklo­ tocenozi zakhidnikh regioniv Ukraini. Polli, L'viv.

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Karpat. Naukova Dumka, Kyiv. Skarby przyrody í ieh ochrona, pp. 81-107. Paústwowa Rada Ochrony Przyrody & Kasa im. Mianowskiego, War~ MALINOVS'KY K. A. & ICRlCSFALUSY V. V.2000. Visokogirna szawa. roslinnist' . In: K, A. MALINOVS'KY & Y.P. DIDUKH (eds), Roslinnist' Ukraini 1: 5-231. Phytosociocentre, Kyiv. WOLOSZCZAK E. 1888. O rododendronie na Syniaku. Kosmos 13: 374. MEUSEL H., JAGER B., RAUSCHERT S. & WEINERT E. 1978. Verg1eichende Chorologie der Zentraleuropaischen Flora. ZAPALOWICZ H. 1889. Roélinna szata g6r Pokucko-Marmaro­ Pischer, Jena. skich. Spraw. Komis. Fizjogr. 24: 1-389.

Received 8 April 2004 A. BORAIYNSK1 ETAL.: RHODODENDRON MYRTIFOLIUM1N THE UKRA1N1AN CARPAIHlANS 59

ApPENDIX ibidem, lego Zerova, 1956(KW); ibidem, leg. Lazebna, 1955 (LWD); ibídem, leg. Fírczuk, 1956 (LWD); ibídem, leg. Zhdanova, 1959 (LW); ibidem, leg. Rozenko, 1959 Localities of Rhododendrom mytrifolium Schot (LW); ibidem, leg. Karpa, 1961 (LW); ibidem, leg. Naj­ & Kotschy in the Ukrainian Carpathians. czuk, 1966 (LW); ibidem, leg. Andryjezyn, 1968 (LW); POLONINA BORZHAVA: [Wolovets, on the Borzhava] ibidem,leg. Zeman & Kritska, 1981 (KW); ibídem, SW Wolovec, na poloninach Borzawskich, leg. Juz 'kiv slope,leg. Lazebna, 1954 (LWD); ibídem, 1750 m, leg. & Ivanye'kyj, 1947 (LW); extinet (Chopik 1976). Koliszczuk, 1956 (LWD); ibídem, S slope, leg. Vaynagiy, 1956 (LWD); ibídem, 2020 m, leg. ?, 1955 (KRAM); GORGAN MIS: [Syniak Mt.], on Siniak near Tar­ ibídem, on top, 2025 m, leg. Szyszowa, 1955 (LW); tarow, leg. Wolaszezak, 1888 (LW); ibídem, (Wo­ íbidem, subalpine meadows, lego Melnyk, 1988 loszezak 1888); ibídem, 1660 m, leg. Srodon, 1937 (KWHA); ibidem, rare onN slope,2020 m Cetrarietum (KRAM); íbidem, rare (Barbarieh 1957); [Syvnla Mt.] sempervirentis (Malinovsky & Kricsfalusy 2000); Syvula (Barbarieh 1957); [Popadiya Mt.] Popadija ibidem, S s1ope, 1760 m, Rhododendretum myrtifolii (Chopik 1976); [StrymbaMt.] Strymba(Chopik 1976); (Malinovsky & Kriesfalusy 2000); ibídem, SW slope, [Steryshora Mt.] Steryszora (in the Polonina Czarna 1840-1850 m, Rhododendretum myrtifolii (Malinovsky massil), leg. Kuts, 1950, (UD); ibidem, leg. Hryhora, & Kricsfalusy 2000); ibídem, NW slope, 1700 m, Em­ 1952, (UU), ibídem (Barbarieh 1957). petro-Vaccinietum (Ma1inovsky & Kricsfa1usy 2000); ibídem, W slope (Barbarich 1957); [Gropa Mt.], on SVIDOVEC: [Svidova Mt.], Swidowiec, Swidowa, Gropa,leg. Rehman, ?, (LW); ibídem, leg. Choptk, 1963 1735-1770 m alt., (Zapalowiez 1889); [Bliznitsa Mt.] (KWHA); [Khoverla Mt.], Howerla, leg. Turczyñski, Blizniea,leg. Zapalowiez, 1904 (KRAM); ibídem, Kar 1864(LWD); ibidem, in regione mughi, leg. Woloszczak, Trofanieeki,leg. Fedorowicz & Wilczyñski, 1911 (LW, 1888 (LWD); ibídem, top of mountain, 2058 m alt LWD); ibídem, subalpine zone, lego Barbarích, 1947 (Zapalowiez 1889), ibídem, N glacial cirque, 1505 m (KW); ibídem, leg. Gryn', 1947 (KW); ibídem, leg. (Zapalowiez 1889), ibídem, stony plaees in alpine belt, Rudenko, 1948 (UU); ibidem, leg. Fodor, 1957 (UU); in places abundantly, leg. Paczoski, 1895 (LW); ibídem, ibidem, leg. Kuts, 1950, (UU); ibídem, leg. Kosee, 1950 leg. Janowsko, 1901 (LWD); ibídem, leg. Kulczyñski, (KW); ibídem, leg. Stojko, 1956 (LWD); ibídem, leg. 1925, (LW); ibídem, leg. Kuzma, 1928 (LWD); ibídem, Melnyk, 1977 (KW); ibídem, leg. Kharkevieh, 1977 leg. Prygrodski, 1932, (LWD); ibídem, leg. Blaszczyk, (KWHA), ibidem (Barbarieh 1957); [Kraehuniska Mt.], 1932 (KRA); ibídem, leg. Kotov, 1947 (KW); ibídem, Kraezunivska PoIonina, 1460 m alto (Zapalowiez 1889), subalpine lego ibídem, (Barbariez 1957); [Gropa Mt.], Polonina Gropa, meadows, 1800-2000 m, Dobroczaeva, 1947 (KW); ibídem, at mountain base, leg. Dobro­ leg. Malinowski, 1948 (LWD); czaeva, 1947 (KW); ibídem, leg. Rudenko, 1947 (UU); CHORNOHORA: [Chornokhora range], Czamohora, ibídem, S slope, 1800 m, leg. Bilik, 1948 (KW); ibídem, in thesubalpine andalpinebelts, leg. Kozikowski, 1909 leg. Kryvonos, 1950 (UU); ibídem, leg. Pavliv, 1950 (LW); ibídem, leg. Kulczynski, 1925 LW; ibídem, leg. (LW); ibídem, leg. Fodor 1952, 1958, 1980 (UU); Mqdalski, 1927 (KW); ibídem, leg. Liedl, 1927 (LW); ibídem, leg. Komendar, 1952 (KW); ibídem, leg. Ar­ ibidem,leg. Komendar, 1953, (KW), íbtdem, leg. Dorn­ temezuk, 1957 (CHER); ibídem, leg. Kondratyuk, 1952 stein, 1961 (UU); ibídem, leg. Palezyk, 1961 (UU); (UU); ibídem, leg. Vtalanova, 1952 (UD); íbidem, 2000 ibídem, leg. Fedoronczuk, 1986 (KW), Bela Tysa Fo­ m,leg. Chopik, 1954 (KWHA); ibídem, leg. Szyszova, rest Resort, leg. ?, ? (KW), Kvasy, NE of Kvasy, on 1955 (LW); ibídem, leg. Kondratyuk, 1957 (KW); back wall of the glacier car, sandstone, lego Kardasz, ibídem, leg. Pawliv & Rozhyeka, 1958 (LW); ibídem, 1985 (LW).; ibídem, N slope, 1850 m, Rhododendretum leg. Gazude, 1961 (UU); ibídem, leg. Mosyakin, 1985 myrtifolii (Malinovsky & Kriesfalusy 2000); ibídem, (KW); ibídem, at the Prut souree, leg. Artyuszenko, 1957 SW slope, 1850 ro, Empetro-Vaccinietum (Malinovsky (KW); ibídem, at top, leg. Vaynagiy, 1958 (LWD); & Kriesfalusy 2000); [Sheshul Mt.], Szeszul, leg. La- ibídem, N slope, leg. Vaynagiy, 1958, (LWD); ibidem, " zebna, 1955 (LWD); ibidem, leg. Kardasz, 1985 (LW); 1800 m, leg. Szyszowa, 1955 (LW); ibidem, leg. ?, 1955 [Petros Mt.], Pietros, S slope, 1600 m (Zapalowicz (KRAM); ibídem, 1800 m, leg. Koliszczuk, 1957 (LWD); 1889), ibídem, leg. Paczoski, 1895, (LW); ibídem, ibídem, 2020 m, frequent1y, Primulo-Caricetum cu­ 1700 m, leg. Malinowski, 1948 (KW, LWD), 1952 rvulae (Malinovsky & Kricsfalusy 2000); ibidem, SSE (LWD); ibidem, leg. Fodor: 1949 (UU); ibídem, leg. slope, 1950 m, Calamagrostis villosa (Malinovsky Tiaeej, 1953 (UU), ibídem, leg. Ulyczna, 1953 (LW); & Kriesfalusy 2000); ibídem, W slope, 1980 m, Rho- 60 POLISH BOTANICAL STUDIES 22. 2006. ._------

dodendretum myrtifolii (Malinovsky & Kricsfalusy Rhododendretum myrtifolii (Malinovsky & Kricsfalusy 2000); ibídem, NNE slope, 1550 m, Empetro-Vaccí• 2000);ibídem, SE slope, 1740 m, Rhododendretum myr­ nietum (Ma1inovsky & Kricsfalusy 2000), tbídem, tífolti (Malinovsky & Kricsfalusy 2000); ibídem, W slope (Barbarich 1957); [Kozmeska Mt.], Kozmeska, E slope, 1500 ro, Vaccinietum myrtilli (Malinovsky 1485 m, (Zapa1owicz 1889), ibídem, NW slope, 1500 m, & Kricsfa1usy 2000); ibídem, SE slope, 1610 m, PI­ Empetro-Vaccinietum (Malinovsky & K.ricsfalusy 2000), netum mughi (Malinovsky & Kricsfalusy 2000); [Ma­ NNW slope, 1700ro, rare in Cetrario-Juncetum (Ma­ ryshevska Velka Mt.], Maryszewska Wielka, Jaremcze, linovsky & Kricsfalusy 2000); [Kostrich Mt.], Kostrycz, in sparse forest on N slope below top, lego Koliszczuk, 1547 ro, NE slope, rare on forest edge, lego Srodoñ, 1960 (KW); [Homul Mt.], Homu1, 1470 m, (Zapalowicz 1936 (KRAM); [Brescu1 Mt.], Bresku1, lego Kozij, 1889), íbidem, leg. Artemczuk, 1957 (CHER); [Between (LWD); ibídem, on the slopes, leg. Miczynski, 1933 Dancer MI. and Turku1], Dancerz - Turku1, SE slope, (KRAM); ibídem, lego Grin', 1948 (KW); ibídem, SE 1910 m, frequent in Cetrario-Juncetum (Malinovsky slope, lego Szewczuk, 1961 (LWD); ibídem, lego Lo­ & Kricsfa1usy 2000); íbtdem, NE slope, 1800 m, Rho­ zebna, 1962 (LWD); ibídem, S lope, frequently in sub­ dodendretum myrtífolii (Malinovsky & Kricsfalusy a1pine belt, 1800 m, lego Boratynski, 1998 (KOR); 2000); ibídem (Barbarich 1957); [Nesamovite lake], Je­ ibidem, NW slope, 1700ro, Calamagrostis villosa (Ma­ zioro Níesamowite, in Pinus mugo thicket aboye lake, linovsky & Kricsfalusy 2000); [Zaros1ak], Zaroslak, leg. Boratynski & Dldukh, 1998 (KOR); [Turkul Mt.], 1425 m, (Zapalowicz 1889), íbidem, lower glacial Turkul (Drabiny below Turkul), 1580 m a1t. (Zapalo­ cirque above Zaros1ak, lego Powlowskl 1925 (KRA), wicz 1889), Turku1ska Polonina, 1570 m alt. (Zapato­ ibídem, Plecze, 1420 m (Zapalowicz 1889); [Dancer wicz 1889); ibídem. lego Szofer, 1908 (KRA); íbidem, MI.] Dancerz, 1700 m, lego Kozij, 1930 (LWD); ibídem, fonnsdensethicket belowTurkut, lego Wycieczka LB.B., S slope,lg. Lozebna, 1967 (LWD); ibídem, lego Du­ 1910, KRA, KRAM, LW; ibídem, 1680-1780 m, dense bovik, 1972 (KW); ibídem, 1650 m, NE slope, aboye thicket orundergrowrh ín Penc mugo thicket on sand­ glacial cirque, lego Boratyñski & Dídukh, 1998 (KOR); stone, lego Pawlowskl, Sulma & Walas, 1929 (KRA, ibídem, NE slope, 1850ro, ftequent, Cetrario-Juncetum LW); íbidem, lego Morozyuk, 1970 (KW); ibídem, N (Malinovsky & Kricsfa1usy 2000); ibídem, S slope, slopes, leg. Stopkan', 1970 (KWHA); ibídem, lego Do­ 1750 ro and SW slope, 1800 ro, Calamagrostis villosa broczaeva, 1977 (KW); ibídem, lego Vynogradova, 1977 (Malinovsky & Kricsfalusy 2000), ibídem, SE slope, (herb. CBR); ibídem, lego Loveljus, 1979 (KW); íbidem, 1550 ro, Calamagrostis villosa (Malinovsky & Krics­ S slope, 1950 m, Primulo-Caricetum curvulae (Mali­ fa1usy 2000); ibídem, NW slope, 1870 m, Rhododen­ novsky & Kricsfalusy 2000); ibídem, S slope, 1700 m, dretum myrtifolii (Ma1inovsky & Kricsfalusy 2000); rare with Festuca amethystina (Malinovsky & Krics­ [Breskulets Mt.], Breskulec, subalpine belt, Empetro­ falusy 2000); ibídem, NE slope, 1740 m, Soldanello­ Vaccínietum, NW, 1550 ro, leg. Boratyfzski & Didukh, Nardetum (Malinovsky & Kricsfalusy 2000); ibidem, 1998 (KOR); [Between Breskul and Pozhyzhevska] NNE slope, 1920 m, with Calamagrostís vil/osa (Ma­ Breskul - Pozyzevska, NE slope, 1800 m, rare in Ce­ linovsky & Kricsfalusy 2000); ibídem, N slope, 1800 m, trario-Juncetum (Malinovsky & Kricsfalusy 2000); Rhododendretum myrtifolii(Malinovsky & Kricsfalusy ibídem, NE slope, 1840 m, Rhododendretum myrtifolii 2000); ibídem, NE slope, 1910 m, Rhododendretum (Malinovsky & Kricsfa1usy 2000); ibídem, W slope myrtifolii (Malinovsky & Kricsfa1usy 2000); ibidem, N (Barbarich 1957); [Pozhyzhewska], Pczyzewska, in slope, 1980m, Rhododendretum myrtifolii(Malinovsky upper part of polonina with dwarfmountain pine, leg. & Kricsfalusy 2000); ibídem, NW slope, 1740 m, PI­ Janowskl, 1901 (LWD); ibídem, leg. Motyka, 1933 netum mughl (Malinovsky & Kricsfalusy 2000); [Be­ (LW); íbídem, lego Kharkevích, 1961 (KWHA); ibídem, tween Shpytsi and Turkul Mt.], Szpyci - Turkul, N lego Andriuszczenko, 1962 (LWD); ibídem, lego Bedej, slope, 1910 ro, frequentIy in Cetrarto-Juncetum (Ma­ 1962 (LW); ibídem,lego Choplk, 1966 (KWHA); ibídem, linovsky & Kricsfalusy 2000); [Shpytsi Mt.], Szpyci, lego Zawada, 1969 (LWD); ibidem, lego Holubec, 1970 on stony top, lego Lobarzewskí. 1854 (LWD); íbidem, (LWD); íbidem, lego Vaynagly, 1971 (herb. CBR); below top near one spring of Gadzhyna stream, with ibídem, lego Vynogrodova, 1977 (herb. CBR); íbidem, white flower, leg. Turczynski, 1883 (LWD); ibídem, lego Vynogradzkl, 1973 (LWD); ibídem, leg. Kric'ko, covers aH the mountains, lego Jonowski, 1901 (LWD); 1979 (KW); ibídem, leg. Loveijus, 1979 (KW); ibídem, íbidem.Leg. Zapalowlcz, 1906 (KRAM); ibídem, on the subalpine meadows, lego Zelenczuk, 1981 (LW); ibídem, slopes exposed toWielkie Kozly, lego Fedorowicz & Wil­ 1400 m, lego Jermaczenko, 1958 (LWD); ibídem, 1650­ czynski, 1929 (LWD); ibídem, 1800 m, among Pinus 1750 m, leg. Bojarczuk, 1997 (KOR); ibídem, 1800 m, mugo, lego Tasienkiewícz; 1976 (LWD); ibídem; 2000 m, A. BORATYNSKI ETAL.: RHODODENDRON MYRTIFOLlUM IN THE UKRAINIAN CARPATHIANS 61

leg. Kyjak, 1976 (LWD); Ibldem, E slope, 1900 m (Ma­ abícs forest, 1100 m (Piwczynski 2000, obs.); [Mun­ linovsky & Kricsfalusy 2000); ibidem, SW slope, 1950 che1 Mt.], Munczel, N slope, 1930 m, rare, Cetrarto­ ID, rare, Cetrarietum sempervirentis (Malinovsky Juneetum (Malinovsky & Kricsfalusy 2000); ibídem,N & Kriesfalusy 2000); íbidem (Barbarich 1957); [Be­ slope, 1900 m, rare with Sesleria bielzii (Malinovsky tween Shpytsi and Rebra Mt.] Szpyci - Rebra, N slope, & Kricsfalusy 2000); ibídem, N slope, 1995 m, Rho­ 1900 ID, Rhododendretum myrtifolii (Malinovsky dodendretum myrtifolii (Malinovsky & Kricsfalusy & Kricsfalusy 2000); [Rebra], Rebra, NE slopes, 2000); ibídem, NE slope, 1750m, Cetrario-Vaccinietum 1750 m, sparse in Festucetum picturatae community (Malinovsky & Kricsfalusy 2000); [Pip lvan Mt.], Pop (Malinovsky & Kricsfalusy 2000); ibídem, N slope, Ivan Mt., Czama Góra, inalpae PopIwan, leg. Herbich, 1920 ro Primulo-Carícetum curvulae (Malinovsky 1840 (LW); ibídem, near top, leg. Turczynski, 1883 & Kriesfalusy 2000); ibídem, SWW slope, 1950 m, fre­ (LWD); ibídem, ieg. Dzieduszycki, ?, (LWD); ibídem, quent with Sesleria bíelzii (Malinovsky & Kricsfalusy 2026 m (Zapalowicz 1889), ibídem, leg. Kotula, 1935 2000); ibídem, S slope, 2000 m, with Festuca amety­ (LWD); ibídem, leg. Klokov & Zozulin, 1940 (KW); stina (Malinovsky & Kriesfalusy 2000); ibídem, NW ibidem, 1956, lego Popov & Khrzanowsky, 1946 (KW); slope, 1870ID, Rhododendretum myrtifolii(Malinovsky ibidem, SE slope, leg. Bradis 1947 (KW); ibídem, leg. & Kricsfalusy 2000); ibídem, N slope, 1950 m, Rho­ Kotov, 1947 (KW); ibídem, 1600 m, leg. Grin', 1948 dodendretum myrtifolii (Malinovsky & Kricsfalusy (KW); ibídem, leg. Kosec, 1950 (KW); leg. Barbarice, 2000); ibídem, NE slope, 2007 m, Rhododendretum 1954 (KW); ibídem, leg. Artemczuk; 1962 (CHER); myrtifolii (Malinovsky & Kriesfalusy 2000); ibídem, ibidem, SE slopebelow the top, 2000 m, leg. Boratynski, NW slope, 1750 ID, Cetrario-Loiseleurietum (Malino­ 1997 (KüR); ibídem, NW slopes, 1980 m (Malinovsky vsky & Kricsfalusy 2000); [Velyki Kozly Mt.], leg. Za­ & Kricsfalusy 2000); ibídem, 1700 and 1750 m, N wada, 1972 (LWD); NE and E slopes, 1700 m (Ma­ slopes (Malinovsky & Kricsfalusy 2000), ibidem, SW linovsky & Kricsfalusy 2000); [Gutyn-Tomnatek Mt., slope, 1970-2000 m, frequently in Salicetum retuso­ Tomnatek Mt.], Tomnatek Maly, 2018 m alt. (Zapato­ reticuíatae (Malinovsky & Kricsfalusy 2000); ibídem, wicz 1889); Gutyn-Tomnatek, N slopes, 1900 ro, rare N slope, 1800 m, Rhododendretum myrtifolii (Ma­ in Festucetum pícturatae (Malinovsky & Kricsfalusy linovsky & Kricsfalusy 2000); ibídem, SE slope, 2000); ibídem, SWW slopes, 1910 m, rare in Primulo­ 1800 m, Pinetum mughi (Malinovsky & Kricsfalusy Caricetumcurvulae(Malinovsky & Kricsfalusy 2000), 2000); ibídem, W slope (Barbarich 1957); [Pokhorylets ibídem, NW slopes, 2000 m, Primulo-Carícetum cu­ Mt.], Pohorylec, 1460 m (Zapalowicz 1889); [Smotrich rvulae (Malinovsky & Kricsfalusy 2000); ibidem, NNW Mt.], Smotrycz, Dzembronia near Zabie, in dwarf slope, 1930 m, rare, Cetrarío-Juncetum (Malinovsky mountain pine thicket, leg. Dzieduszyeki, 1870 (LWD); & Kricsfalusy 2000); [Gadzhyna Mt.],Gadzyna, 1355 m ibídem, leg. Andrijenko & Bradis (KW), Smotrycz (Zapalowicz 1889), ibídem, leg. Szewezuk, 1960 (LWD); (Smotreé) Mt., 1400 m alt. (Zapalowicz 1889), ibídem, ibidem,N slope,2007 m, Cetrario-Loiseleurietum (Ma­ aboye W ridge of Skoruszny, 1310 m (Zapalowicz linovsky & Kricsfalusy 2000); [Kizie Ulokhy], Kizie 1889), ibidem, 1350m, on slopesinPínus mugothicket, Ulohy, 1470 m (Zapalowicz 1889), ibídem, in regíoni leg. Koliszczuk, 1956 (LWD); ibídem, 1500 m, NW Píneü mughi, leg. Mqdalski, 1927 (LW); ibídem, leg. slopes, (Boratyóski 1997, obs.); íbidem, between Smo­ Zerov, 1962 (KW); [Lemska Mt.], Lemska, 1495 m (Za­ trich and Stepanec, leg. Zahulskij & Gojda, 1986 (LW); palowicz 1889); [Brebenska Mt.], Brebenieska, 1540 m [Lopata Mt.] Lopata, SW slope, 1600 m, Junlperetum ¡ and 2039 m alto (Zapalowicz 1889), Brebeneskul, leg. sibiriceae(Malinovsky & Kricsfalusy 2000); [Negrovets i;¡ Dobroczaeva,?, (KW); ibídem, withwhiteflowers (Bar­ Mt.], Negrovec', NE síope, 1720 m, rare in Cetrario­ barich 1957); bidem, N slope, 1995 m, rare in Solda­ Festueetum (Malinovsky & Kricsfalusy 2000); [Rog­ 1 nello hungaricae-Salicetum kitaibelianae (Malinovsky neska Mt.], Rogneska, leg. Susip, 1960 (LW); ibídem, & Kricsfalusy 2000); ibídem, NW and W slopes, N slope, 1900 m, Rhododendretum myrtifolii (Ma­ 1980m, Cetrario-Loiseleurietum (Malinovsky & Krics­ linovsky & Kricsfalusy 2000); íbidem, NW slope, falusy 2000); [Rozshybenyk Mt.], Rozszybenyk, 2000 m 1780 m, Empetro-Vaccinietum (Malinovsky & Kricsfa­ (Zapalowiez 1889); [Dzhembronya Mt.] Dzembronia, lusy 2000); [Skopeska Mt.], Skopeska, lego Vaynagiy, 1470 m (Zapalowicz 1889), ibídem, at Dzembronia 1993 (herb. CBR). stream, 1360 m and 1225 m alt.,(Zapalowicz 1889), ibídem, at Mincholek stream, 1335 m (Zapalowicz MARMAROS ALPS: Marmaros Mts,lego Zapalowicz, 1889); [Dzhembronya] rare on tbe granite rocks, N 1907 (KRAM); [Gropa Mt.], Gropa, 1360 m, to Pop slopes aboye Dzhembronya village, in sparse Picea Ivan (Zapalowicz 1889), ibídem, Gropszora Polonyna, 62 POLISHBOTANICAL STUDIES 22. 2006. leg. Tovt, 1964 (UD); LatundurMt., leg Vaynogiy, 1977 barich 1957); [Petros MI. of Marmaros Mts], Petros (CBR); [Pip 1van of Marmaros Mts], Pop 1van Mar­ (Barbarich 1957). maroski: aufder Fejérpataker AlpePop Ivan Marmaros, leg. Vagner, 1873 (LW); RossiaSubcarpatica: inrupibus CHYVCHIN MTS: [Lozdun Mt.], Polonyna Lozdun schistosis monttsPop Ivon, ca. 1800 m, leg. Suza, 1927 in Chyvchin Mts, lego Kharkevich, 1965 (KWHA); (KRA, LW); ibídem, leg. Hruby, 1932 (KRAM); ibidem, [Preluki MI.] Polonyna Preluki, leg. Chopik, 1964 leg. Popov, 1946 (LWD); ibidem, leg. Czernecki, 1947 (KWHA); [Hnitesa Mt.J, Hnitesa (Gnetesa), subalpine (UU); ibídem, alpine meadows, 1940 m, leg. Soko­ meadows,leg. Chopik, 1963 (KWHA); [Koman Mt.], Iovsky, 1947 (KWHA); ibídem, leg. Komendar: 1956 Koman, 1705-1730 m (Zapa1owicz 1889), ibídem, (UU); ibidem, leg. Chopik, 1964 (KWHA); ibidem, E slope, on Baltagu1 Polonina, 1615 m, leg. Mqdolski, leg. Voynagiy, 1992, 1993, 1994 (herb CBR); ibídem 1934 (LWD); ibidem, 1600 m, N slope, at foothill of (Barbarich 1957); [Zerban Mt.], Zerban, 1450 m alt., Koman, leg. Pawlowski, 1934 (KRA); ibidem, 1500 m, N slope (Zapa1owicz 1889), ibidem, (Barbarich 1957); leg. Powlowski, 1934 (KRA); [Between Stewiora and [Neneska Mt.], Nienieska, S slope, 1785 m (Zapa1owicz Koman], on slopes in devastated dwarf-pine scrub; lego 1889), ibídem, leg. Juz'kiv, 1947 (LW); tbtdem, (Bar- Kluskówno & Walos, 1934 (KRA).