E Stuttgart, November 8, 2011

Plant Div. Evol. Vol. 129/3 4, 317 327 E Stuttgart, November 8, 2011 Article Distribution changes of endangered peatland vascular plants in Western Pomerania (Poland) By Zofia Sotek With 2 figures Abstract Sotek, Z.: Distribution changes of endangered peatland vascular plants in Western Pomerania (Poland). Plant Div. Evol. 129: 317 327. 2011. ISSN 1869-6155. Peatland degradation destroys the habitats of rare and endangered plants, which is reflected in the decrease of their local range. This paper discusses how the distribution of the vascular plant species Carex pulicaris and Pedicularis sceptrum-carolinum has changed in Western Pomerania, Poland, since the 19 th century. These species are especially affected by peatland melioration and the resulting encroachment by tall perennials and shrubs. Keywords: Carex pulicaris, Pedicularis sceptrum-carolinum, local range, locality. Introduction An enormous number of Polish peatlands has been been drained since the 19th century, and in particular in the second half of the 20th century, to harvest peat or to convert them into agricultural land (Herbichowa 1976, Jasnowska & Jasnowski 1977). The vegetation type that was most decimated by this development is the natural fen vegetation. Fens are the most fertile type of peatland and, for this reason, were the first to be converted into grasslands (Jasnowski 1972). However, the most endangered plant species are those of the raised and transitional bogs, which are very susceptible to disturbances in hydrological conditions. The disappearance of peatland plant species has become a widespread phenomenon (Herbichowa 1976, Jasnowska & Jasnowski 1977), and the rarest species are most threatened by extinction (Kornaś 1976). This paper discusses how the distribution of the vascular plant species Carex pulicaris and Pedicularis sceptrum-carolinum, of the phytosociological class Scheuchzerio-Caricetea nigrae, has changed in Western Pomerania since the 19th century. They are included in the list of endangered and threatened vascular plant species of Western Pomerania (Żukowski & Jackowiaka 1995), under category E (extinct). In addition, C. pulicaris is included in the Polish Red Book of Plants (Kaźmierczakowa & Zarzycki 2001). Received January 24, 2011, in revised form May 7, 2011, accepted June 22, 2011 2011 E. Schweizerbart sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de DOI: 10.1127/1869-6155/2011/0129-0056 1869-6155/2011/0129-0056 $ 02.75

318 Z. Sotek, Distribution changes of endangered peatland vascular plants Methods Distribution maps of the selected species are based on the author s field research, on historical and current data, both published and unpublished, and on records from Polish herbaria. All localities were mapped according to the ATPOL 5 5 km grid square system (Zając 1978). In the list of localities, current Polish names are used throughout instead of former German names. Three time intervals were selected for the comparison of distribution ranges: the 19th century, the first half of the 20th century (until 1950), and the period since 1950. Abbreviations refer to the following herbaria: POZ Herbarium of the Adam Mickiewicz University in Poznań; TRN Herbarium of the Nicolaus Copernicus University in Toruń. Results Carex pulicaris L. Carex pulicaris belongs to European-temperate (western) sub-element, extending to the Atlantic region of Europe (Zając M. & Zając A. 2009). It range reaches northern Spain and Ireland in the west, southern Scandinavia in the north and Poland as well Estonia in the east. Its southern range boundary runs across Transsylvania, Hungary, northern Croatia, the Italian Alps and southern France, up to the Pyrenees (Schultze- Motel 1980; Hultén, Fries 1986, Tutin et al. 1989). The eastern border of the distribution area crosses Poland. It has been reported in Western Pomerania, Lower Silesia and the Małopolska Uplands, as well as in the Western Carpathians (Zając A., Zając M. (Eds.) 2001). In Pomerania, the range of Carex pulicaris covers mostly northern areas, not reaching far to the south (Fig.1). This species has been recorded so far in 74 localities, with most of them being observed in the north-eastern part of the region. From among 27 localities reported prior to 1900, seven were not confirmed later, while two of them persist until today. Unconfirmed localities were reported from the north-western and the north-eastern part of Pomerania. In 1900 1950, C. pulicaris was known from 54 localities in this region, but only three of them, in the north-eastern part of the region, were confirmed later. At present, there are 19 known occurrences in Pomerania, scattered across the entire northern part of the region, and almost all of them were first reported after 1950. The Sowno Moor ( Wrzosowisko Sowno ) occurrence near Płoty was discovered in 1973 and confirmed in 1977 (Jasnowska & Jasnowski 1973, 1977 unpublished), but could not be found again in 1998 and 2003, despite great efforts. C. pulicaris occurs in fen communities in the phytosociological order Caricetalia davallianae. It is also found in wet heaths, peat-covered meadows, montane Nardus grasslands of the alliance Nardion, and in sallow thickets (Myrico-Salicetum auritae) (Matuszkiewicz 2001). Localities [AB-1411] Międzywodzie (Piotrowska 1966); [AB-1510] Dziwnów (Piotrowska 1966); [AB-2311] near Lake Linówek (Lucas 1860 (1861)); [AB- 2400] Boryń (Lucas 1860 (1861)); [AB-3500] Rozwarowo Marshes, SE part (Jurzyk & Wróbel 2003); [AB-4800] Wrzosowisko Sowno (Sowno Moor) (Jasnowska

Z. Sotek, Distribution changes of endangered peatland vascular plants 319 Fig. 1. Distribution of Carex pulicaris L. in Polish Pomerania. locality until 1900, locality in 1901 1950, locality after 1950, + extinct locality. & Jasnowski 1973, 1977 unpublished), not found recently (Sotek & Stasińska 1998 unpublished, 2003 unpublished); [AB-6510] Goleniów (Müller 1911); [AB-7201] Pilichowo (Müller 1911); [AB-7300] Goślice near Przęsocin (Müller 1911); [AB- 7301] Szczecin-Glinki (Müller 1911); [AC-0300] Szczawno (Müller 1911); [AC- 1710] Gardziec (Müller 1911); [BA-6910] Pęplino (Misiewicz 1977); [BA-7811] c. 2 km SE of Nosalin (Sotek, Stasińska 2005 unpublished); [BA-8601] Słowino (Müller 1911); [BA-9801] Łętowo (Załuski 1975 unpublished); [BB-2711] near Drzewiany (Osadowski 1999); [BB-2810] near Zarzewie (Osadowski 1999); [BB- 3201] Głodzino (Winkelmann 1889; Müller 1911); [BB-8600] in the vicinity of Lake Busino Małe near Nadarzyce (Abromeit et al. 1940); [CA-3511] Osieckie Marsh (Abromeit et al. 1940); [CA-3601] reserve Piaśnickie Łąki (Preuss 1911; Herbichowa, Herbich 1993); [CA-3611] Wierzchucino (Wnorowski 1986 unpublished); [CA-3710] Żarnowiec (Abromeit et al. 1940); Łąki Żarnowieckie (Abromeit et al. 1940); [CA-3810] Czarny Młyn (Graebner 1895, 1895 TRN; Abromeit et al. 1940);

320 Z. Sotek, Distribution changes of endangered peatland vascular plants Bielawskie Marsh (between Sławoszyn and Mieroszyn) (Graebner 1895, 1895 TRN, Enderlein 1908, Wangerin 1916, Abromeit et al. 1940); between Karwia and Ostrowo and Sławoszyno (Preuss 1911); [CA-4400] Szczenurze (Müller 1911); [CA-4501] Osieki (Graebner 1895, 1895 TRN; Müller 1911; Abromeit et al. 1940); [CA-4700] Kartoszyno (Wnorowski 1986 unpublished); [CA-4801] Werblińskie Marsh (Bail 18?? TRN, Graebner 1895, 1895 TRN, Enderlein 1908, Preuss 1911, Abromeit et al. 1940); [CA-4811] between Darżlubie and Miechowo (Graebner 1895, Abromeit et al. 1940); [CA-5000] Dębina (Müller 1911); [CA-5510] near Brzeźno Lęborskie (Klinggraeff 1883 TRN); [CA-5700] Rybno (Klinggraeff 1881, Abromeit et al. 1940); Czajcze Marsh (Kniewo) (Herweg 1914); [CA-5711] Wejherowo (Künzer 1878, Abromeit et al. 1940); [CA-5810] near Nananice (Ruhmer 1879, Klinggraeff 1881, Abromeit et al. 1940); [CA-5900] Mrzezino (Klinggraeff 1883 TRN, Graebner 1895, 1895 TRN, Preuss 1911, Abromeit et al. 1940, Markowski & Szlachetko 1993); [CA-5901] S of Osłonino (Graebner 1895, Abromeit et al. 1940); by the Reda river, near Beka (Graebner 1895, Abromeit et al. 1940, Lenartowicz et al. 1996); Rewa (Abromeit et al. 1940); [CA-5910] Moście Marsh, W of Rewa (Bail 1869 TRN, Preuss 1911; Abromeit et al. 1940); between Rewa and Kazimierz (Abromeit et al. 1940); between Reda and Dębogórze (Abromeit et al. 1940); near Połchowo (Graebner 1895, 1895 TRN, Abromeit et al. 1940); [CA-6701] Dąbrowski Młyn (Konopelska-Feliks 1987 unpublished); [CA-6900] between Dębogórze and Cisowa (Abromeit et al. 1940); between Kazimierz and Suchy Dwór (Abromeit et al. 1940); [CA-6911] near Gdynia (Lange 1887 TRN); [CA-7000] Słupsk (Müller 1911); [CA-7500] Okalice (Lemcke 1887, Herweg 1914, Abromeit et al. 1940); [CA-7510] between Okalice and Łebunia (Lemcke 1887, Herweg 1914, Abromeit et al. 1940); [CA-7611] Strzepcz (Abromeit et al. 1940); [CA-8101] Starnice (Czubiński 1950); [CA-8211] Budowo (Müller 1911); [CA-8601] Wysoczyzna Staniszewska (Herbich 1976 unpublished); [CA-8901] Las Oliwski (Abromeit et al. 1940); between Klukowo and Oliwa (Bethke 1884); [CA-8911] Kiełpino (Preuss 1911, Abromeit et al. 1940); Kiełpinek (Klinggraeff 1881, Abromeit et al. 1940); [CA-9200] Niepoględzie (Czubiński 1950); [CA-9810] Borcz (Abromeit et al. 1940). [CB-0300] Bytowo, 3 km NE (Herbich 1994 unpublished); [CB-0601] Gołubie near Lake Patulskie (Herbich 1994 unpublished; Herbich 2000); [CB-2210] in the vicinity of Lake Trzebielsk near Lipnica (Lisowski et al. 1969, 1968 POZ); [DA-6010] Gdynia (Preuss 1910; Abromeit et al. 1940); [DA-7010] Sopot (Klinggraeff 1881, Herweg 1914, Preuss 1928); Świemirowo (Bail 1881 TRN, Bethke 1884, Lützow 1891, 1883 TRN, Preuss 1911, Herweg 1914, Wangerin 1928 TRN); [DA-8000] Oliwa (Lützow 1884 TRN); [DA-8001] Nawitzerthal (Herwig 1878 TRN); [DA-8010] Gdańsk Brętowo (Klinsmann 1852 TRN, Klinggraeff 1881, Abromeit et al. 1940); near Matęblewo (Preuss 1911, Abromeit et al. 1940). Pedicularis sceptrum-carolinum L. P. sceptrum-carolinum is a Euro-Siberian species (Zając & Zając 2009) that occurs in northern and central Europe as well as in Siberia. In Europe, it has been reported from Scandinavia, Estonia, Lithuania, Poland, Denmark, Germany, Austria, Hungary, east-

Z. Sotek, Distribution changes of endangered peatland vascular plants 321 Fig. 2. Distribution of Pedicularis sceptrum-carolinum L. in Polish Pomerania. locality until 1900, locality in 1901 1950, locality after 1950. ern Romania, northern Ukraine, and Belarus, and from the European part of Russia (Tutin et al. 1990, Hartl 1975, Meusel et al. 1978, Hultén & Fries 1986). Its western range boundary cuts accross Poland. The localities are concentrated in two areas, i.e. north-eastern Poland and the Lublin Region (Zając & Zając 2001). In Pomerania, sparse occurences of P. sceptrum-carolinum have been recorded at 26 localities, mostly in the in the Brda River valley in the south-eastern part of the region (Fig. 2). The species does not reach far west, with the westernmost locality being situated in Barwice of Central Pomerania (Czubiński 1950). Until 1900, P. sceptrumcarolinum was known from 15 localities, nine of which have not been confirmed since. In 1900 1950, eleven of the 17 localities known in this period were reported for the first time, including three solitary, isolated localities. No new localities have been re-

322 Z. Sotek, Distribution changes of endangered peatland vascular plants corded after 1950, and none of the earlier localities has been confirmed, although floristic studies were carried out in the respective areas. P. sceptrum-carolinum is characteristic for the phytosociological class Scheuchzerio-Caricetea nigrae (Matuszkiewicz 2001). In north-eastern Poland, it occurs in fen communities of the order Caricetalia davallianae. In Pomerania, it occurs in fens, wet forests, and on river banks, but its phytosociological affiliation is unknown. Localities [BB-5501] Barwice (Czubiński 1950); [CA-3800] Ostrowo (Preuss 1907, Preuss 1911, Herweg 1914); between Karwia and Ostrowo (Preuss 1911); [CA-4301] Łeba (Müller 1911); [CB-2711] Zamek Kiszewski (Treichel 1878, 1879 TRN, Abromeit 1896, Abromeit et al. 1903, Kalkreuth 1917); Stara Kiszewa (Preuss 1899 TRN); Wilcze Błoto (Abromeit 1896); Bartoszylas (Abromeit et al. 1903, Abromeit et al. 1940); [CB-7400] Obrowo (Schriften 1880, 1883; Klinggraeff 1881, Brick 1883 TRN, Grütter 1891, Abromeit 1897, Preuss 1906, Wangerin 1919, Abromeit et al. 1940), not found recently (Ceynowa-Giełdon 1993); between Dąbrówka and Obrowo (Preuss 1906 TRN); between Żalno and Tucholka (Preuss 1906); [CB- 7410] near Kęsowo (Abromeit 1897); [CB-7610] near Huta (Grütter 1889, Schriften 1890, Abromeit et al. 1903); [CB-8410] near Komierowo (Abromeit 1915); [CB- 8700] between Ostrowite and Cisiny (Peil & Grütter 1885 TRN, Abromeit et al. 1903); Ostrowite (Hohenfeltt 1885 TRN; Grütter 1891); [CB-9300] near Zboże (Abromeit et al. 1903; Preuss 1905 TRN); Messy, N of Zboże (Abromeit 1882); [CB- 9400] near Przepałkowo (Koppe 1928); [CC-0401] Słupowo (Kuhling 1866); [CC- 0411] Krąpiewo (Kuhling 1866, Wodziczko et al. 1938); [CC-1411] Strzelewo (Kuhling 1866, Klinggraeff 1881); Minikowo (Kuhling 1866); [CC-2201] Śmielin (Kuhling 1866); [CC-2400] Ślesin (Kuhling 1866, Wodziczko et al. 1938); Gorzeń (Wodziczko et al. 1938). Discussion Many historical localities of Carex pulicaris and Pedicularis sceptrum-carolinum have not been confirmed later on. This does not necessarily reflect their disappearance, but might also be due to a lack of research. For similar reasons, it is difficult to determine when newly discovered localities were first colonized. However, if a locality has not been confirmed for nearly 60 years in spite of dedicated research, the respective population has probably disappeared. Therefore, it can be assumed that P. sceptrum-carolinum, which has not been found after 1950 (Fig. 2), is now totally extinct in Pomerania. As a perennial which can grow as high as 80 100 cm, it would be difficult to overlook. P. sceptrum-carolinum is a hemi-parasite that grows on grass and sedge roots. Its basal leaves are clustered in a rosette, while the stalk leaves are much smaller and sparser. Its physiognomy could explain why it is so prone to being overgrown by shrubs and tall perennials (e.g. reeds), and this may have happened at the localities in the Noteć River valley and at least some of those in the Brda River valley. Moreover, the Noteć River valley was meliorated in the post-war period, which lead to gradual soil drainage and increasingly disfavourable conditions for P. sceptrum-carolinum. The area was also

Z. Sotek, Distribution changes of endangered peatland vascular plants 323 undersown with various grass species in the 1970s, before land use was abandoned in the 1980s, which caused a rapid succession to tall perennials and shrubs. On the other hand, many localities in the Brda River valley that were mown in the past are now overgrown with reeds. P. sceptrum-carolinum, which in Poland reaches its south-western distribution limit, is sporadically found to the west of Vistula River (Zając & Zając 2001), but its range has been shrinking. It has rarely been recorded in eastern Germany, and before 1949 was known only from Mecklenburg-Vorpommern (Benkert et al. 1998). In Poland, it is considered an endangered species, both in the region of Pomerania (Żukowski & Jackowiak 1995, Markowski & Buliński 2004) and in the country as a whole. The Polish Red List of Plants (Zarzycki & Szeląg 2006) includes it among the critically endangered species, and so does the List of Threatened Bog Flora Species (Jasnowska & Jasnowski 1977). C. pulicaris grows 5 30 cm tall and has setaceous leaf blades. It is a rather inconspicuous plant which can easily be overlooked, even when flowering or fruiting. Accordingly, in some localities where it has not been confirmed, it may just have slipped the researchers attention. The ears of this sedge contain few female flowers, so that seed production is low and the potential for generative reproduction weak. In some localities, for example near Nosalin, C. pulicaris forms small local populations. Small populations (small in terms of area and / or number of individuals) are particularly endangered in case when living conditions become worse (Szlachetko 1987, Tyszkowski 1995). It is possible that some of the unconfirmed populations died out because of drainage or encroachment by tall perennials. Some of the localities were certainly destroyed as a result of urban development, for example at Gdynia (Preuss 1910) or Sopot (Preuss 1928) near Puck Bay, or at Pilichowo and Glinki (Müller 1911), which today are city quarters of Szczecin. The eastern range boundary of C. pulicaris cuts across Poland, and it has been observed that there is a disjunction between the Atlantic and the East Baltic populations east of Puck Bay (Kaźmierczakowa & Zarzycki 2001). In Pomerania, this species shows recessive trends. Most localities in the north-eastern part of the region are lost, and further extinction would increase the East West disjunction. Moreover, there is also a marked gap between the north-western localities on the one hand and the central and north-eastern localities on the other. Scattered occurrences of C. pulicaris have been reported from all over eastern Germany, but are relatively frequent only in mountainous areas. In the period after 1949, the localities in the mountains and in Mecklenburg-Vorpommern proved to be the most stable ones (Benkert et al. 1998). In Poland, C. pulicaris has endangered status both regionally in Pomerania (Żukowski & Jackowiak 1995, Markowski & Buliński 2004) and nationally. The Polish Red Book of Plants includes it among endangered species, while the Red List of Plants in Poland gives it critically endangered status (Zarzycki & Szeląg 2006), and the List of Threatened Bog Flora Species cites it as a very endangered species (Jasnowska & Jasnowski 1977). The changes in the distribution of C. pulicaris and P. sceptrum-carolinum seem to confirm the view that isolated populations, or those situated at range boundaries, are most prone to extinction (Kornaś 1976, Wołejko 1983). C. pulicaris had more numer-

324 Z. Sotek, Distribution changes of endangered peatland vascular plants ous localities and was found in a larger area of Pomerania, covering with its range limits the northern part of the region and the belt of Lakelands. This could significantly affect preservation of its presence, despite the lack of confirmation with respect to its many localities. In case of the retreat of P. sceptrum-carolinum from the area of Pomerania however, the fact that it was found in few localities only, mostly concentrated within a small area at the edge of Pomerania, played probably a major role. More generally, this study underlines the vulnerability of peatland species to anthropogenic habitat changes. Streszczenie Zmiany w rozmieszczeniu zagrożonych roślin naczyniowych torfowisk na Pomorzu. Procesy degradacji torfowisk zaznaczające się już w XIX wieku, nasiliły się zwłaszcza w drugiej połowie XX wieku, na skutek antropopresji (m.in. melioracji torfowisk, eksploatacji złóż torfu, rolniczego użytkowania). Wszelkie zaburzenia stosunków hydrologicznych w nieodwracalny sposób wpływają na skład gatunkowy specyficznej flory torfowisk. Prowadzi to do trwałej utraty wielu stanowisk roślin rzadkich i zagrożonych, co niejednokrotnie znajduje odzwierciedlenie w zmniejszaniu się ich lokalnych zasięgów. Dostępne dane florystyczne z Pomorza, historyczne i współczesne, pozwoliły na uchwycenie zmian w zasięgu lokalnym i w rozmieszczeniu stanowisk gatunków zagrożonych na Pomorzu: Carex pulicaris i Pedicularis sceptrum-carolinum. Stwierdzono, że prawdopodobnie całkowicie z tych terenów ustąpił Pedicularis sceptrum-carolinum, gdyż po 1949 roku nie podano nowego, ani nie potwierdzono żadnego z dotychczasowych jego stanowisk. Z kolei w lokalnym zasięgu Carex pulicaris obecnie wyraźnie zaznacza się przerwa między stanowiskami północno-zachodnimi, a pozostałymi, znajdującymi się w części środkowej i północno-wschodniej regionu. Zamieszczone mapy rozmieszczenia gatunków i wykazy ich stanowisk pokazują rangę problemu, jakim jest zanikanie rzadkich roślin ekosystemów torfowiskowych. References Abromeit, J. 1882: Bericht über die Untersuchung des Kreises Flatow. Schriften der Physikalisch- Ökonomischen Gesellschaft zu Königsberg 22: 20 32. Abromeit, J. 1896: Bericht über die 34. Jahresversammlung des Preußischen Botanischen Vereins am 8. Oktober 1895 zu Rastenburg. Schriften der Physikalisch-Ökonomischen Gesellschaft zu Königsberg 37: 139 145. Abromeit, J. 1897: Bericht über die 35. Jahresversammlung des Preußischen Botanischen Vereins am 6. Oktober 1896 zu Konitz. Schriften der Physikalisch-Ökonomischen Gesellschaft zu Königsberg 38: 37 47. Abromeit, J. 1915: Bericht über die wissenschaftlichen Verhandlungen auf der 52. Jahresversammlung in Marienwerder, Westpreußen, am 11. Oktober 1913 sowie über Tätigkeit des Preußischen Botanischen Vereins. Schriften der Physikalisch-Ökonomischen Gesellschaft Königsberg 55: 143 188. Abromeit, J., Jentzsch, A. & Vogel, G. 1903: Flora von Ost- und Westpreußen. I. Samenpflanzen oder Phanerogamen. Preußischer Botanischer Verein zu Königsberg, Berlin. Abromeit, J., Neuhoff, W. & Steffen H. 1940: Flora von Ost- und Westpreußen 2(5). Kommissionsverlag Gräfe und Unzer, Königsberg. Benkert, D., Fukarek, F. & Korsch, H. (eds.) 1996: Verbreitungsatlas der Farn- und Blütenpflanzen Ostdeutschlands. G. Fischer, Jena etc.

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