Orobanche Caryophyllacea Sm. (OROBANCHACEAE) in POLAND: CURRENT DISTRIBUTION, TAXONOMY, PLANT COMMUNITIES and HOSTS

ACTA AGROBOTANICA Vol. 67 (3), 2014: 97–118 Original Research Paper DOI: 10.5586/aa.2014.035

Orobanche caryophyllacea Sm. (OROBANCHACEAE) IN POLAND: CURRENT DISTRIBUTION, TAXONOMY, PLANT COMMUNITIES AND HOSTS

Renata Piwowarczyk

Department of Botany, Institute of Biology, Jan Kochanowski University ćwiötokrzyska 15, 25-406 Kielce, Poland e-mail: [email protected]

Received: 16.11.2013

Abstract has also been reported on G. schultesii, G. glaucum, The paper presents the current distribution of Orobanche G. album, G. sylvaticum or G. tunetanum from Algie- caryophyllacea Sm. in Poland based on a critical revision of ria [3–5,17,20,23–25]. It prefers xerothermic habitats herbarium and literature data as well as the results of my field of the Festuco-Brometea class and fringe communities studies. The majority of localities are in south and south-east- of the Trifolio-Geranietea sanguinei class, especially ern Poland: Maäopolska Upland, Lublin Upland, Roztocze, of the Geranion sanguinei alliance, chiefly in uplands PrzemyĈl Foothills, Pieniny Mts, rarely in the valleys of the and mountains, e.g. 1700 m in Slovakia (Bielskie Tatra Lower Vistula and Oder rivers or Wolin island. The distribution Mts), up to 2000 m in Switzerland [20,24]. map in Poland is included. The taxonomy, biology and ecology The aim of this study was to identify the present of the species are discussed. distribution of Orobanche caryophyllacea in Poland based on field investigations and verified herbarium Key words: Orobanche caryophyllacea, holoparasite, herbarium and literature data. Preferred habitats, communities, collections, distribution, hosts hosts and threats are also discussed below.

INTRODUCTION TAXONOMIC REMARKS In Poland the holoparasitic genus Orobanche L. Orobanche caryophyllacea J. E. Smith 1798, (Orobanchaceae) contains 19 species, including two Transact. Linn. Soc. London 4: 169. Syn.: O. vulgaris ephemerophytes [1–19]. These taxa are protected and Poiret 1798 (1. Nov.), in Lamarck, Encycl. Meth. 4: their majority grow at rare and threatened localities. 621; O. galii Duby 1828, Bot. Gall. 1: 349. The species chiefly occur within the belt of the Pol- Ind. loc.: “Gathered in shrubby hills near Val- ish Uplands and in the lower Oder and Vistula valleys. cimara [Valcimarra, Caldarola, Macerata Province, Le Orobanche caryophyllacea Sm. [bedstraw (clove- Marche Region, Italy], at the foot of the Apennines scented) broomrape] is a European-western Asiatic [Appennino umbro-marchigiano, Apennine Moun- species [20]. It occurs in Western (from France), Cen- tains], in April 1787. Tour on the Continent, vol. ii. tral and Eastern Europe, from southern England, the 308. Linnaeus received the same from Siberia” [26]. Netherlands, Germany, Poland, Estonia, Mediterra- Lectotype: herb. Smith 1087.23 (LINN). nean Region, to Asia Minor, the Caucasus, and Iran In the traditional division, Orobanche caryo- [21]. The species has also been reported from northern phyllacea is included in grex Galeatae Beck, contain- Africa [22,23]. ing species having the corolla with a curved dorsal Orobanche caryophyllacea is oligophagous and line, a galeate upper lip, large flowers, 17–35 mm, yel- parasitizes species of the family Rubiaceae and the low-brown, purple, lilac, usually purpurescent towards genus Galium. In Central Europe, it mostly occurs on the margin, with high colour variability [22,23]. The G. mollugo, G. verum, G. boreale and G. odoratum; it species is placed in the section Orobanche Teryokhin,

© The Author(s) 2014 Published by Polish Botanical Society 98 Renata Piwowarczyk subsection Galeatae (Beck) Teryokhin [27], in the Södziejowice), corresponding to the described form most recent approach. In Central Europe, Orobanche f. strobiligena (Reichenbach) G. Beck. Intensly-col- lutea and O. teucrii also belong to the subsection oured purple individuals, described as f. orphinia G. Galeatae. In an older approach, these species were Beck, can also be encountered in the same populations included in grex Galeatae together with O. gamose- at scattered localities in Poland. Albinotic specimens pala and O. latisquama [23]. At present, the former is have also been observed in shaded sites, such as the placed in a separate section, Gamosepalae, [27] while specimen in the Grabowiec reserve in the Nida Basin O. latisquama has been placed in the genus Boular- [5]. An albinotic form was described as f. albiflora Ba- dia latisquama [28–30]. O. clausonis Pomel, which is guet and reported so far from Belgium [23]. classified in grex Minores [23], is a very similar species; however, the diagnostic characters indicate it MATERIALS AND METHODS should be included in grex Galeatae. It also parasitizes Rubiaceae (e.g. Asperula hirsuta, Rubia peregrina), Field research was conducted in 1999–2013, occurring in the Mediterranean Region, mainly on the with the investigations being intensified between 2006 Iberian Peninsula [31,32]. A recent phylogenetic study and 2013. All the herbarium materials of Orobanche has shown that O. clausonis (section Orobanche) is caryophyllacea available in Poland were reviewed. The closer to the taxa of the section Trionychon [33]. materials are deposited in the herbaria CHRZ, KRA, In Central Europe, determination errors are KRAM, KTC, KTU, LBL, LOD, OPOL, POZ, TRN, mostly made between Orobanche lutea and O. teucrii. UGDA, WA, WRSL, WSRP, and also in Germany (B, Unlike O. caryophyllacea, O. lutea nearly always has GLM). Herbarium acronyms are given after Mirek a wax-coloured or yellow stigma, the dorsal line of the et al. [37] and Holmgren and Holmgren [38]. corolla is curved from its base, slightly convex, but The nomenclature of vascular plants follows Mirek the corolla is not swollen and with a considerably less et al. [39]. The nomenclature of syntaxa is based on open throat. It parasitizes the Medicago genus and is Matuszkiewicz [40]. The localities are organ- scentless or the scent is very delicate. The dorsal line ized as ATPOL cartogram units, 10 km × 10 km [41, of the corolla in O. teucrii is straight or weakly convex; see http://www.ib.uj.edu.pl/chronpol/], in alphabetical the inflorescence is short and with few flowers. It par- order. Probably incorrect localities or not confirmed by asitizes species of the Teucrium genus. Photographs the field or herbarium investigations are marked with and descriptions of the species are available in several the symbol “?” per square number. The localities are studies [20,21]. O. teucrii was incorrectly reported described as follows: ATPOL grid unit, geographic lo- from the Polish part of the Pieniny Mts [34] and from cation, habitat description, abundance in brackets. The the Lublin region [35]. The species was identified as following information is also given for most localities: O. caryophyllacea on herbarium and field verification. geographic coordinates and altitude (above sea level), Phenotypic variability of shoots is very high in and for revised exsiccata the collector and collection Orobanche caryophyllacea, especially of flowers and date, exsiccatum number and the herbarium acronym. stems, with a wide range of colours from albinotic, Host plants were observed by delicately expos- brigh yellow, light pink and purple to violet. The col- ing the soil with a gardening shovel. A total of 80 soil our of the pistil stigma is also variable. Typically, it is pits were observed in the field and in the analysis of purple-coloured, but forms having orange, yellow and herbarium materials containing an attached host. pink stigmas are known. The intensity of pubescence Symbols and abbreviations: ecol. site – ecologi- and metric characters are also observed. One popula- cal site, leg. – collected by, n. – near, phot. – photog- tion can comprise even a few colour forms. Many va- raphy, res. – reserve, v. – valley, xer. – xerothermic, rieties have been described: var. typica G. Beck, var. gras. – grassland, vid. – seen by, ? – doubtful, uncer- incurvata G. Beck, var. buhsei (Reuter) G. Beck, var. tain, or unconfirmed locality. gymnantha G. Beck, and over 30 lower units depending on the colour, pubescence and calyx structure [22,23]. RESULTS Specimens with large flowers (2.5–3.5 mm), lax and multi-flower inflorescence, were determined in Poland, Distribution in Poland especially in Silesia, as var. laxiflora (= O. laxiflora In Poland Orobanche caryophyllacea occurs Reichenb., f. macrantha (Dietrich) G. Beck), while mainly in the Maäopolska Upland (chiefly the Nida Ba- specimens with a lemon-coloured corolla and a purple sin and the Middle Vistula River valley), Lublin Up- pistil stigma from the area of Krynica were determined land, Roztocze, PrzemyĈl Foothills, Sudeten Foreland as f. citrina (Dietrich) G. Beck [36]. Specimens hav- and Silesian Basin, the valleys of the Lower Vistula and ing a lemon-coloured corolla and a pistil stigma can Oder Rivers, Wolin island, and Pieniny Mts (Fig. 1). also be infrequently observed in the Nida Basin (e.g. Frequent determination errors were made in the past

© The Author(s) 2014 Published by Polish Botanical Society Orobanche caryophyllacea Sm. (Orobanchaceae) in Poland: current distribution, taxonomy, plant communities and hosts 99 and the list below presents only verified data. As my gras. leg. H. Piotrowska, 25.05.1967 (UGDA 01/01, verification showed, the species was incorrectly report- 01/02), vid. R. Piwowarczyk, 07.09.2007 (<15), ed from the Silesian – Krakow Upland [e.g. 42–46, un- [64,66–75]; Bielinek-Lubichów Dolny [68]; Markocin published data from the ATPOL database]. It was most [68]; ćwierkocin [76]; 31? – Krzymów [60]; 41 – Stare often mistaken with O. lutea and O. kochii, which often Objezierze, grassland NE of the village, between the occur in the area, infrequently with O. bartlingii and margin of a pine forest and a lake, 52o52’43,3”N, O. elatior [47,48]. Many incorrect reports also con- 14o21’37,6”E, 71 m, (<5), meadow with Arrhenather- cerned the Maäopolska Upland and adjacent areas [e.g. um elatius on the margin of a coniferous forest, 49–53, unpublished data from the ATPOL database]. 52o52’47,2”N, 14°21’30,7”E, 65 m, 52°52’53,6”N, Some literature data could not be verified as the her- 14o21’33,7”E, vid. K. Baraþska, P. Pluciþski, R. Piwo- barium material was lacking or despite investigations in warczyk, 25.06.2009; Dolsko, xer. gras. SW of the vil- the field (especially old German data, with not strictly lage towards Górka lake, 52o53’38,6”N, 14o21’32,7”E, defined localities). These uncertain data are marked 66 m, vid. K. Baraþska, P. Pluciþski, R. Piwowarczyk, with a special symbol on the map (Fig. 1). Attempts are 25.06.2009; ēelichów, slope SE of the village (>40), still made to find and verify many of them in the field. [77]; Moryþ (Mohrin) [64]; 66? – ãupowo [60]; 67? – List of localities: AB: 22? – Przytór n. ćwi- Gorzów Wielkopolski (Landsberg): Ruski Stawek noujĈcie [54]; 23 – Western Pomerania, Wolin island, (Schützensee) [64]; Wieprzyce [60]; 68? – Czechów Wicko, sandy fallow land on Trzciñgowska slopes v., (Zechow) n. Gorzów Wielkopolski [64,78]; 83 – Gór- leg. H. Piotrowska, 21.06.1962 (UGDA/Wolin 002); zyca, Kostrzyn on the Oder river district, leg. W. ēu- 32 – Wolin, Lubin, SW slope of an insolated hill, leg. kowski, 27.05.1962 (POZ); Górzyca (Göritz): Góra B. Chmielewski, P. Szkudlarz, 31.05.1985 (POZ); Lu- Pamiöciþska (Frauendorfer Berge), [64,79]; Owczary bin, on the side of Gäówna street, thermophilous slope on the Oder river, on a sunny slope overgrown with xer. of the Szczecin Lagoon, 53o51’49”N, 14o26’08,02”E, vegetation, leg. W. Gugnacka, 29.05.1973 (TRN); Ow- 50 m, (>30), vid. R. Piwowarczyk, 08.07.2012, [54]; czary, in a ravine, on a slope, leg. W. Gugnacka, 33 – Trzciñgowo, NE of Wapnica, vid. B. Kasperkow- 29.05.1973 (TRN); Owczary, xer. gras., 52o28’23,9”N, icz, 2010, [54]; Wapnica, chalk mine [54]; road side in 14o38’43,1”E, 39 m, (<10), vid. R. Piwowarczyk, a mixed coniferous forest, by the blue tourist trail, S of 25.06.2009, [80]; BC: 76? – Szamotuäy (Master) [81]; Turkusowe lake, 53o52’29,4”N, 14o26’25,1”E, 54 m, Piaskowo [82]; BD: 52? – Säawskie lake n. Säawa (2), vid. R. Piwowarczyk, 09.07.2012; Wolin island, ćlñska [83]; 81 – Dalków, Kurów Wielki (Dalkau, cliff on the Szczecin Lagoon, leg. H. Piotrowska, Kauer), leg. Joachim, 26.06.1888 (auch Pinkwart, 15.07.1951 (UGDA/Wolin 001); Wapnica, sunny 07.1899) (WRSL); Kurów Wielki (Gr.-Kauer) n. Gäo- gorge, leg. H. Piotrowska, 18.06.1976 (UGDA, 01/04); gów [84,85]; ćrem (Schrien) n. Gäogów [85]; 82 – Nie- Dargobñdz [54]; Köpa [54]; 96? – Stargard Szczeciþski lubia (unweit des Nilbauer Birken), leg. Klopfer, and Ulikowo [55–57]; AC: 05 – Grödziec on Miedwie 14.08.1928 (WRSL), [86]; 83 – Ogorzelec, Gäogów lake, xer. slope, leg. J. Stasiak, 01.06.1971 (UGDA, district, in a gras., leg. Z. Gäowacki, 11.06.1972 (WSRP, 01/03); Grödziec, Pyrzyce district, steppe “Brodogóry” 00005165); xer. gras. WWN Ogorzelec (>30), vid. res., on a sunny slope, leg. W. Gugnacka, 29.05.1973 E. SzczöĈniak, 2011 (phot.); Lower Silesia, Dalkowskie (TRN), [55,58–60]; roadside escarpment by the Pyr- hills, Szczyglice n. Gäogów, on an insolated slope SE of zyce-Stargard Szczeciþski road [61]; 06 – “Stary Pr- the village in a gras. of the alliance Koelerion glaucae, zylep” res. between Stary Przylep and Janowo villages, leg. E. Kozioä, 21.05.2005 (KTU, 99017); Szczyglice xer. gras., 53o11’27,25”N, 14o59’33,5”E, 80 m, (<5), (Sieglitz) n. Gäogów, 15.07.1871 leg. ? (WRSL), vid. R. Piwowarczyk, 25.06.2009, [58,62]; 12? – Widu- [84,85]; BE: 16 – Stary Woäów, Wrocäaw province, chowa [60]; 15? – Pyrzyce (Pyritz) [56]; 16? – Zabor- sunny slope, leg. E. Kozioä, 17.06.1981 (WRSL); 67 – sko, E. çwikliþski, unpbl, 1979 (after ATPOL); 21 – Górzyce (Gurwitz) n. Wrocäaw, leg. H. Scholtz (WRSL), Krajnik (Kränig), leg. R. Schulz 1903, 1908 (B), [84 (var. laxiflora), 85,87]; 77 – Winna Góra (Wein- [63,64]; Krajnik Dolny, xer. gras. in the SE part of berg) n. Sobótka (Zobten), leg. R. Sadebeck, 06.1840 Górki Krajnickie, n. an excavation pit, 53o01’35”N, (WRSL), Winna Góra n. Sobótka [84,85]; Nasäawice, 14o18’37,9”E, vid. K. Baraþska, P. Pluciþski, R. Pi- xer. gras. and the area around the quarry on a serpent- wowarczyk, 25.06.2009, [60]; Raduþ, leg. R. Schulz inite hill, so-called Kamienny Grzbiet, SzczöĈniak un- 1908, (B), [60,63–65, vid. S. Rätzel, 2010 oral com.); pbl 1997 (after ATPOL), (<300 in 2001), [88], (<80 in 22? – Nawodna [60]; 30 – Bielinek on the Oder river, 2011), 50o53’00,0”N; 16o49’50,8”E, 174 m, vid. R. Pi- Chojna district, leg. J. Schäde 1847, 1850, leg. R. Schulz wowarczyk, 14.07.2011; 87 – Jasinek n. Niemcza 1911, 1914 (B), [63], leg. F. Celiþski, 28.05.1971 (Johannistal), at present Góra Cierniak (Ostseite des (KTU, 1057); “Bielinek” res. on the Oder river, leg. Dörner berges), leg. T. Schube, 14.06.1914 (WRSL), Exc. Inst. Bot. U. P., 06.1952 (POZ); Bielinek, xer. [89]; 88 – Prusy, between Niemcza and Strzelin, on

G. mollugo, 06.1856, leg. R. Uechritz (WRSL); Janow- – Bingsberge hills (at present Góry ãosiowe Mts) be- iczki (Klein Johnsdorf) n. Niemcza (Nimptsch), on tween Zakurzewo (Sackrau) and Weäcz (Wolz), on G. verum, var. laxiflora, leg. Schumann, 05.06.1880 G. molugo [92,94,95]; Grudziñdz, Zakurzewo, Bings- (WSRL), [84,85]; Ksiöginice Wielkie (Gr. Kniegnitz) berge, leg. Peil, 15.06.1880 (TRN), [96]; 81 – Grudziñdz n. Niemcza [84]; 97 – Niemcza, bank of the ćlöĔa riv- (Graudenz) [81], meadows on the high bank of the Vis- er, behind the hospital, abundant (Loheufer hinter dem tula, SW of the fortress [92], on G. mollugo and Genesungsheim in grossen Zahl), 14.06.1935, leg. G. verum], Grudziñdz, fortress, leg. Grütter, 03.07.1887 Frömsdorf; Niemcza (an den Ohleabhangen bei (WA, 33086) [94–96]; DF: 78? – Jurczyce [107]; 88? Nimptsch) [90]; Stasin (Pangel) near Niemcza [85]; – Wola Radziszowska [107]; DG: 00? – Cieszyn (Te- CB: 99? – Cheämno (Kulm) [81,91–93]; Cheämno, schen) [85]; EC: 94? – Modlin [108]; EF: 04 – between area of the Fryba river [94–96]; CC: 08 – between Chomentówek and Södziejowice, xer. gras. and ther- Kieäpand Päutowo, on G. verum [92,95,96]; Päutowo, mophilous scrub on S- and SW-facing hills on the leg. A. Kozäowska, 06.1927 (KRAM, 241975); E and W side of the road (<200 in 2007; <100 in 2013), Päutowo, slope, leg. M. Ceynowa, 07.06.1958 (TRN); 50o34’23”N, 20o39’44”E, 264 m, vid. R. Piwowarczyk, “Päutowo” res., (3), vid. R. Piwowarczyk, 10.09.2004; A. Przemyski, 2007, vid. R. Piwowarczyk 30.05.2013, (6) vid. S. Kusz, 02.06.2013; ćw. Wawrzyniec Mt leg. R. Piwowarczyk, 06.06.2007, 04.06.2009 (KTC), in Starogród n. Cheämo (Althausen am Lorenzberge) [5]; Gartatowice, E part, xer. gras., on gypsum, on both [92,95–97]; “Góra ćw. Wawrzyþca” res., xer. gras. sides of the road (<100), 50o34’34”N, 20o37’33”E, 237 (<5), 53o19’37,5”N, 18o22’57,9”E, 92 m, vid. R. Piwo- m, vid. R. Piwowarczyk, A. Przemyski, 2007, leg. warczyk, 29.05.2009; Szymborno, Cheämno district, R. Piwowarczyk, 06.06.2007 (KTC), [5]; a quarry in leg. M. Ceynowa, 29.07.1961 (TRN), [97]; 18 – slope Borków, leg. ?, 2001 (KTC); Samostrzaäów, SSE part, n. Unisäaw, NWW of Na Skarpö street (4), vid. S. Kusz, xer. gras. (<100), 50o35’18”N, 20o38’41”E, 244 m, leg. 15.06.2013 (phot.); 39? – Gajtowo (Getau), between R. Piwowarczyk, 04.06.2009 (KTC), [5]; 14 – Pasturka, Solec and Toruþ [96,98]; Toruþ, Dybowo (Nieder- in a hole remaining after excavated stone on the forest mühl) [96]; 58? – between ćciborze and Oräowo [96]; margin, leg. J. Mñdalski, 03.06.1966 (KRAM, 494810); 67? – Inowrocäaw [81]; Inowrocäaw, Niedermühle oak-hornbeam forest in the W part of the “Grabowiec” [96]; 68? – Jaronty [96]; 75? – Mogilno [81]; Mogilno/ res. between Gacki and Bogucice I (<100). The species Wiecanowo [82,96,98]; 85? – Trzemeszno n. Poznaþ parasitizes G. odoratum, G. mollugo, G. verum and [99]; CE: 90 – Przeworno n. Strzelin (Im Prieborner Cruciata glabra there (>50), 50o28’21”, 20o34’33”E, Marmorbruch), marble quarry, leg. F. Wimmer, 240 m, vid. A. Przemyski, 2006, leg. R. Piwowarczyk, H. Grabowski, 18?? (WRSL), [84,85,100,101]; 95? – 02.06.2007, 06.06.2007, 28.05.2008 (KTC), vid. R. Pi- Opole (Oppeln, bei dem Salzmagazin), [100], probably wowarczyk, 2012, [5,53]; “Grabowiec” res. n. Krzy- O. lutea; CF: 16? – Kamieþ ćlñski (Grosstein), [100], Ĕanowice, gras., leg. A. Jasiewicz, 07.06.1957 (KRAM, probably O. lutea; 45? – Borzysäawice (Borislawitz) near 414019); Bogucice, Grabowice (probably the “Gra- Pawäowiczki (Gnadenfeld) [84,85]; 59 – Rudy (Rauden), bowiec” res.), leg. K. Kaznowski, 13.06.1931 (POZ); leg. Dziatzko, 02.07.1897 (OPOL, P/3468); 65 – Kietrz res. Bogucice, leg. S. ChroĈcielewska, 15.07.1935 (Katscher), Gipsowa Góra (Gypsgruben), leg. Kelch (WA, 0000028406=018860); 15 – ãagiewniki n. Bus- (GLM-DB, 138814), leg. E. Kozioä, 15.06.1983 (WRSL, ko-Zdrój, xer. gras., S and SW-facing (“Zimne Wody” 96785), [84,85,101]; Kietrz, sunny slope, leg. E. Kozioä, res.) (<50), 1531, 50o28’58,72”N, 20o44’00”E, 254 m, 15.06.1983 (WRSL, 90811); DzierĔysäaw n. Kietrz, xer. leg. R. Piwowarczyk, 21.07.2006 (KTC), [5]; Dobrow- gras. on gypsum areas, leg. Kelch, 1840 (GLM, 138823), oda [109]; Czerwony Chotel [109,110], Busko-Zdrój leg. ?, 14.06.1859, on G. verum and G. mollugo (GLM, [110]; 24 – Skorocice n. Busko Zdrój, gypsum rocks, 106750), leg. Z. Dzwonko, A. Frey, 23.06.1970 (KRA, leg. A. Jasiewicz, 07.06.1957 (KRAM, 414009); 26 – 066868); “Gipsowa Góra” res., DzierĔysäaw between Kików and Suäkowice, xer. gras. on a former n. Kietrz, leg. A. Sendek, 14.05.1972 (KRA, 0138562); vast limestone excavation pit (so-called Kamienna leg. A. Sendek, 14.06.1972 (KTU, 10554), [102]; Gip- Góra), (>200), 50o25’05,51”N, 20o51’24”E, 258 m, sowa Góra n. Kietrz, leg. K. Rostaþski, 01.06.1985 50o25’04”N, 20o51’38”E, 287 m, leg. R. Piwowarczyk, (KTU, 99029); leg. B. Fojcik, 01.06.1990 (KTU, 28.05.2008 (KTC), vid. R. Piwowarczyk 30.05.2013, 99030); leg. M. Buchalik, 05.06.1999 (KTU, 99027); [5]; Suäkowice [111]; 27 – xer. gras. by the Piestrzec – leg. L. Dudek, 14.06.1972 (KTU, 16210), [103,104]; Biechów road junction (>5), leg. A. Przemyski, 08.2008 Kozäówki (Kösling) n. Kietrz, var. laxiflora, leg. (KTC); 34 – WiĈlica, W part, xer. gras. and scrub on M. Wetschky, 06.1876, 18.05.1904 (1907?) (WRSL), a S, SE and SW-facing slope (<50), 50o21’01”N, [85,105]; DA: 80? – Gdaþsk [96]; 87? – Frombork 20o39’50”E, 171 m, leg. R. Piwowarczyk, A. Przemyski, (Frauenburg) [106]; 96? – Kikoäy (Kickelhof) (vicinity 08.2006 (KTC), [5,110]; 35 – Górki n. Szczerbaków, of Janówek at present) between Kadyny (Cadinen) and xer. gras. on a slope on gypsum, S, SW and SE-facing Tolkmicko (Tolkemit), by the road, [92,95,96]; DB: 71 (>100), 50o20’51”, 20o43’54”E, 174 m, 50o20’55”,

20°44’01”E, 179 m, vid. A. Przemyski, 2007, leg. czyk, 01.08.2009, 49o25’04,6”N, 20o27’35,1”E, 505 m; R. Piwowarczyk, 2007 (KTC), vid. R. Piwowarczyk Góra Zamkowa, gras. near pines by the summit, on 30.05.2013, [5]; 86? – Janowice, on a dry meadow on G. mollugo, leg. K. Zarzycki, 13.07.1967 (KRAM, a steep slope on the Dunajec river [112]; EG: 14? – 363774), [34]; small mid-forest meadow on the slope of Góra Cheäm, Kñclowa [113]; 17? – Cheämiecka Góra Góra Zamkowa by the Potok Pieniþski stream, leg. [36, vid. 06.1916); 32 – Pieniny Mts, between Macelak K. Kostrakiewicz, 24.06.1952 (KRAM, 241650); in and a unnamed peak towards the Szopka pass, by the scrub on Góra Zamkowa, leg. J. Nowak, 19.07.1954 trail near a group of trees (Galium, Laserpitium), leg. (CHRZ); gras. and scrub n. castle ruins, (2), W. Gugnacka, L. Rutkowski, 14.06.1983 (TRN); Mac- 49°25’12,5”N, 20o25’12,9”E, 724 m, vid. R. Piwowar- elak, N slopes, (5), 49o25’14”N, 20o22’32”E, 840 m, czyk, 30.07.2009; Pieniny Mts, „Górka” n. Däugi Gro- vid. I. Dembicz, ã. Kozub, 03.07.2013 (phot.); Kozia nik, meadow with Brachypodium, leg. E. Kotejowa, Góra 790 m, Origano-Brachypodietum [114]; Kozia 15.06.1967 (KRAM, 363776, 363777); warm meadow Góra, warm meadow, (<15), vid. R. Piwowarczyk, n. Däugi Gronik, (<5), 49o25’08,2”N, 20o26’56,6”E, 23.07.2010, 49o25’15,6”N, 20o22’09,4”E, 791 m; Ko- 515 m, leg. R. Piwowarczyk, 20.07.2010, [34]; ãaĒne zia Góra, SW-facing roadsides, (<20), 49o25’13,1”N, Skaäy, meadow, vid. W. and L. Frey (phot.), 2009 20o22’21,4”E, 822 m, vid. R. Piwowarczyk, 23.07.2010; (<30), on G. mollugo; between KroĈcienko and Tyl- 33 – Pieniny Mts [115], Pieniny Mts, woody scrub, leg. manowa [119]; Stajkowa Góra over KroĈcienko, on the Woäoszyn, 07.1928 (KRA, 012171); Pieniny Mts, leg. Dunajec on the Dunajec SW, dry meadow, 670 m, leg. R. Kobendza, 1926 (WA, 0000028444=018927); Pien- K. Stawowczyk, 09.06.2009; thermophilous gras. S of iny Mts, leg. A. Jasiewicz, 24.06.1952 (KRAM, an unnamed rock, 872 m, N of Cyrylowa Skaäa, leg. 414023); cfr. Pieniny Mts, rocks on the Dunajec river, K. Zarzycki, 13.06.1966 (KRAM, 363780); ãñczana, leg. M. Wayda, 23.06.1994 (KRA, 0128530); Pieniny n. Przeäöcz Trzy Kopce, small hill N of the trail, (<10), Mts, Biaäe Skaäy, beech forest on S slopes, leg. A. Ja- 49o25’08,4”N, 20o23’12,7”E, 780 m, vid. R. Piwowar- siewicz, 25.06.1964 (KRAM, 414020, 095244); forest czyk, 23.07.2010; Nowa Góra 850 m, Origano-Brachy- below Biaäe Skaäy, leg. E. Koteja, 196? (KRAM, podietum [114]; Kras 460 m, Origano-Brachypo- 363773), [34]; Ociemny Potok, on the upper side of the dietum, [114, as O. lutea]; Polana Pieniny Mts, 940 m, green trail, Fagetum carpaticum, leg. K. Zarzycki, 1965 Veratrum lobelianum-Laserpitium latifolium, [114, as (KRAM, 363778, 363779); Ociemny Potok (Pieniþski), O. lutea]; Polana Zaosice, 630 m [120, as O. alba]; in the Carpathian beech grove on G. odoratum, leg. Wñwóz Gorczyþski gorge (Macelowy) [121]; Facimiech K. Zarzycki, 26.06.1966 (KRAM, 363771), [34]; Pien- [122]; Slovakia: Maäe Pieniny Mts, immediately past iny Mts, thermophilous beech groves below Czerwone the Slovak border, by the yellow tourist trail descending Skaäy, towards the Pieniþski Potok stream, 500 m a.s.l., to LeĈnica and diverging from the blue ridge trail be- leg. K. Zarzycki, 08.06.1966 (KRAM, 363772); Potok tween Szafranówka and Bystrzyk, vid. W. and L. Frey, Pieniþski [116]; Potok Pieniþski, below the Limerczyki 02.07.2006 and 2008; Wylizana Mt, by the blue trail, glade, Phyllitido-Aceretum, leg. E. Koteja, 07.1963 S-facing, 49o24’49,8”N, 20o27’12,3”E, 447 m, vid. (KRAM, 363775), [34]; Potok Pieniþski, below Czer- R. Piwowarczyk, 01.08.2009; 34 – Maäe Pieniny Mts, tezik, on gruss in scrub, leg. A. Jasiewicz, 24.06.1952 “Wñwóz Homole” res., gras. on a rock slope, leg. (KRAM, 414010); by the bank of the Pieniþski Potok B. Wójcikiewicz, 27.06.1973, 12.07.1973 (KRA, 71416, stream, opposite Ociemny Wierch, meadows, leg. 71417); “Wñwóz Homole” res., Gras. at the foot of rocks A. Jasiewicz, 07.07.1953 (KRAM, 414028); scrub be- over less than 100 meters to the left of the entrance, low the Pieniþski Potok stream, leg. Szajna, 10.05.1930 W-facing, (<10), 49o24’20,1”N, 20o32’52,4”E, 568 m, (KRAM, 180355), [117]; scrub on rocks below the vid. R. Piwowarczyk, 31.07.2009; Wñwóz Homole on Pieniþski Potok stream, leg. K. Piech, 09.06.1930 the west slope of the gorge, rock grassland, E-facing, (KRAM, 2200105); Dolina Czarnego Potoku valley, 49o24’16,6”N, 20o32’51,9”E, 560 m, vid. R. Piwowar- 650 m, cleared patch with Calamagrostis varia, leg. czyk, 31.07.2009; 37? – Krynica [36,123], var. citrina; K. Zarzycki (KRAM, 363769), as O. teucrii; Sokolica Kopciowa [36,123], var. citrina; 44 – Wysokie Skaäki, [116]; beech forest below Sokolica, leg. K. Piech, summit [124]; Wysoka Mt, summit, rock grassland, 09.06.1930 (KRAM, 220106); Trzy Korony, leg. Szaj- (>5), 49o22’49,4” N,20o33’20”E, 1050 m, vid. R. Piwo- na, 09.05.1930 (KRAM, 180356); Trzy Korony, gras. warczyk, 31.07.2009; 46? – Mikowa n. Muszyna [36]; near the summit, leg. K. Zarzycki, 02.07.1967 (KRAM, FC: 97? – Drohiczyn [125]; FE: 13 – Vicinity of Puäawy, 363776); scree below Okrñglica [118]; Okrñglica, on hillsides, leg. A. K. Simionow, 06.1887 (WA, 49o24’56.9”N, 20o25’20”E, 947 m, vid. R. Piwowar- 000002845= 018925), Wäostowice near Puäawy [126]; czyk, 30.07.2009, 22.07.2010; 07.2012, [4]; Orlica, on 23 – Miöèmierz n. Kazimierz Dolny, grassy chalky gruss on the Dunajec, leg. A. Jasiewicz, 24.06.1952 slope, leg. H. Przychodzeþ, 19.06.1980 (LBL); xer. (KRAM, 414024), as O. teucrii; behind the „Orlica” gras., Möèmierz, Albrechtówka (>5), vid. R. Piwo- hostel n. Szczawnica, meadow (<15), leg. R. Piwowar- warczyk, 04.06.2011; 27 – Rury, Lublin, gorges, leg.

H. Koporska, 06.1916 (LBL); 28 – Sobianowice n. Lu- czyþski, 20.07.1870 (WA, 0000028399=018863); 61/62 blin, loess slope, leg. D. Fijaäkowski, 09.06.1949 (LBL); – Tarnogóra, gras. belonging to Thalictro-Salvietum prat- sunny loess slopes, leg. D. Fijaäkowski, 30.05.1959 ensis association in the escarpment complex in the (LBL); Däugie n. Lublin, loess slope, leg. D. Fijaäkowski, Wieprz v. (>15), 50o53’47”N, 23o07’10”E, leg. A. Cwen- 03.06.1949 (LBL); Lublin, a gras. belonging to Thalic- er, 24.06.2008 (LBL), [17]; 62 – Dworzyska, E-facing tro-Salvietum pratensis association on a high baulk on steep slopes of the Wieprz v., an overgrowing Thalictro- the slope of the Bystrzyca valley (>10), 51o16’23”N, Salvietum pratensis association (<20), 50°55’47”N, 22o37’57”E, leg. A. Rysiak, 2006 (LBL); 52 – Ciszyca 23°08’38”E, leg. A. Cwener, 09.06.2008 (LBL), [17]; Kolonia, xer. gras. by the edge and on the SE-facing 64 – Skomorochy Maäe n. Grabowiec, chalky slopes, leg. slope of the Vistula v. (>100), 51o01’08”N, 21o46’38”E, D. Fijaäkowski, 14.06.1958 (LBL); Hajowniki (Refor- 153 m, leg. R. Piwowarczyk, 25.07.2001, 01.06.2002, ma), xer. gras. and scrub on the chalky slope of the Woli- 21.06.2005 (KTC), [1,3,5,17,127]; Dorotka n. Taräów, ca v., on the northern side of the Skierbieszów – Grabo- xer. gras. and scrub N, S and SE of Dorotka, S, SE and wiec road (>50), 50o50’26”N, 23o26’35”E, leg. A. Cwe- SW-facing (<30), 51o00’02”N, 21o47’05”E, 148 m, leg. ner, 22.06.2007 (LBL), [17]; 67 – Szpikoäosy n. Hrbie- R. Piwowarczyk, 01.06.2002, 18.06.2006 (KTC), szów, scrub of a loess slope, leg. D. Fijaäkowski, [1,3,5,17,127]; 59 – Kolonia ēuków n. Krzczonów, 13.06.1958 (LBL); 72 – Zäojec, a xer. gras. belonging to a partly forested xer. gras. n. a closed limestone quarry Thalictro-Salvietum pratensis association on a midforest (>5), 51o02’15”N, 22o47’40”E, vid. P. Chmielewski, hill in the ãabuþka v., ca. 300 m N of the village church 17.08.2009, [17]; 62 – Wesoäówka, NNE part, S-facing (>30), 50o46’55”N, 23o07’12”E, vid. W. Michalczuk wasteland on chalky rendzina (>15), 50o58’37”N, 17.06.2006, [17]; 77 – Czumów n. Hrubieszów, loess 21o47’49”E, vid. R. Piwowarczyk, 1999–2006, slope, leg. D. Fijaäkowski, 20.06.1955 (LBL); 81? – [1,3,5,17,127,128], it did not occur in 2008–2013; Szczebrzeszyn [108]; Dziewcza Góra S of Niedzieliska Säupia NadbrzeĔna [50,129], SE-facing xer. gras., [133]; 82 – Kñty n. ZamoĈè, clearing in an oak forest on 250 m NW of the church (>10), vid. R. Piwowarczyk, rendzina, leg. D. Fijaäkowski, 05.06.1955 (LBL); oak for- 2001, 2005; [O. caryophyllacea: Biedrzychów reported est on chalk, leg. D. Fijaäkowski, 29.06.1951 (LBL); in the literature [129]; literature data verified in the field, sunny chalky hill, leg. D. Fijaäkowski, 04.06.1959 (LBL); a locality of O. lutea recorded, leg. R. Piwowarczyk, Kñty II, xer. gras. adjacent to a dirt road from the 2007 (KTC); Baätów [51,126], a locality of O. alba re- Wymysäówka village to Wychody, on a midfield hill corded, leg. R. Piwowarczyk, 24.06.2007; “Ulów” res. known as Wieprzecka Góra (>50), 50o40’22”N, [130], a locality of O. lutea recorded, vid. R. Piwowarc- 23o07’22”E, leg. R. Piwowarczyk, W. Michalczuk, zyk, 2007]; Popów n. Józefów, xer. gras. on a chalky P. Chmielewski, 15.07.2006 (KTC), [17]; Hubale, xer. slope, leg. M. Kucharczyk, 30.06.1980 (LBL); 68? – gras. on a calcareous hill, ca. 0.5 km N of the village Wysokie n. Turobin [126]; 73 – Opoka n. Annopol on (>15). The locality was partly damaged by ploughing in the Vistula, xer. gras. on a chalky slope, leg. M. Kucha- 2009. 50o41’07”N, 23o09’47”E, vid. W. Michalczuk, rczyk, 26.06.1980 (LBL); 82 – Dwikozy ecol. site, 2007, [17]; 94 – ãabunie n. ZamoĈè, forest glade on Dwikozy n. Sandomierz, N of the stadium, xer. gras. on chalk, leg. D. Fijaäkowski, 15.07.1957 (LBL); 96 – Do- the S-facing slope of the Opatówka river v. (>20), buĔek, “Skarpa DobuĔaþska” res., ca. 1 km NW of the 50o44’20”N, 21o47’03,9”E, 174 m, leg. R. Piwowarc- DobuĔek village, on the steep slope of the Huczwa v., zyk, 27.07.2006 (KTC), [5,17,50,131]; Dwikozy n. San- xer. gras. belonging to Thalictro-Salvietum pratensis as- domierz, (on G. mollugo) leg. R. Kobendza, 05.1930 sociation, on chalky rendzina and chernozems (>50), (WA, 0000028402=018858), [5]; GD: 23? – Bohukaäy 50o34’50”N, 23o42’55”E, leg. P. Chmielewski, R. Piwo- n. Terespol, xer. gras. [132]; GE: 23 – Stawska Góra n. warczyk, 12.07.2006 (KTC), [17]; GF: 07 – Posadów, Cheäm, summit of a chalky hill, leg. K. Izdebski, xer. gras. on embankments of an early medieval earth- 10.07.1958 (LBL); ãöczyca-Wäodawa lakeland, “Staw- work, on the edge of the Kamionka v., ca. 0.5 km N of the ska Góra” res., leg. B. Chwastowski, 08.06.1978 village (>30), 50o30’27”N, 23o48’57”E, vid. E. Gorgol (KRAM, 455835); Cheäm district, leg. M. Hempel, 1880 and P. Chmielewski, 24.05.2009, [17]; 14? – Biaäa Góra (WA, 0000028404 (10898); 55/56 – Kolonia Teresin, n. Tomaszów [126]; 15 – Przeorsk, xer. gras. on the ãysa xer. gras. and roadsides on chalky rendzinas on a hill ca Góra hillside and on the bottom of a dry small v. joining 0.5 km S of the village (>50), 50o59’21”N, 23o41’56”E; the Soäokija river (>50), 50o25’27”N, 23o31’06”E, leg. 50o59’14”N, 23o42’20”E; 50o59’20”N, 23o41’30”E; P. Chmielewski, R. Piwowarczyk, 09.06.2007 (KTC), 50o59’00”N, 23o40’46”E, vid. P. Chmielewski, A. Cwe- [17]; 25 – Machnów Stary, overgrowing xer. gras. in the ner, R. Krawczyk, 2005, leg. P. Chmielewski, R. Piwo- „Machnowska Góra” res. (>10), 50o22’06”N, 23o35’18”E, warczyk, 15.07.2006 (KTC), [17]; Teresin, along the vid. P. Chmielewski, R. Piwowarczyk, 13.07.2006, [17]; margins of a small forest n. Teresin farm, leg. F. Karo, 80? – Winna Góra n. PrzemyĈl, S slope; ravine behind the 29.06.1883 (WA, 0000028413=018891); 56 – Strzelce DruĔbacki farm [134]; 90 – PrzemyĈl, Grochowska street, (near Biaäopole) in Hrubieszowskie district, leg. K. ãap- dry roadside, leg. M. Wolanin, 05.06.2009 (KRA).

Fig. 1. Distribution of Orobanche caryophyllacea in Poland: Ɣ – localities confirmed after 2000; ż – localities documented with herbarium collections, not confirmed after 2000; ? – localities not confirmed in the field, unsertain or probably doubtful, lack of herbarium materials.

Host preferences G. mollugo in 80% of cases, seldom G. verum and very Prior to my study, Orobanche caryophyllacea was rarely G. boreale, e.g. in Södziejowice or Kików in the reported from Poland as a parasite of the Rubiaceae fam- Nida Basin. Parasitism on G. odoratum was noted only ily, genera Galium, Asperula and Sherardia [135]. A total in the “Grabowiec” reserve in the Nida Basin [5] and in of 80 soil pits were investigated in the field between 2006 Carpathian beech forests in the Pieniny Mts (leg. K. Zar- and 2013. Herbarium materials were analyzed if a speci- zycki, 26.06.1966, KRAM; see list of localities). The spe- men was collected together with the host. My analysis cies parasitized Cruciata glabra only at one locality on has shown that in Poland O. caryophyllacea parasitizes the margin of the “Grabowiec” reserve [5] (Fig. 2). Symp- exclusively species of the family Rubiaceae. It infects toms of host weakening by the parasite were not observed.

7700

50 s e liti 4400 oca l of r 30 be m u number of localitie of number n 2200

1100

0 GaliumGalium mollumollugogo G.G. verum G.G. boreale G.G. odoratum CruciataCruciata glabraglabralabra Fig. 2. A comparison of preferred host species of Orobanche caryophyllacea in Poland (on the basis of field observations and herbarium research).

Preferred habitats and plant communities the Querco-Fagetea class where it parasitizes Ga- In Poland Orobanche caryophyllacea mainly lium odoratum. The species has been rarely recorded occurs in thermophilous habitats, from the southern in grassland communities of the Sisymbrio-Stipetum sector of the horizon, seldom on west- or north-facing capillatae association. These preferred communities slopes, i.e. xerothermic grasslands of the Festuco- are often in mosaic segetal communities of the Stel- Brometea class, thermophilous meadows Molinio-Ar- larietea mediae, shrub communities Rhamno-Prunetea, rhenatheretea class. These communities vary consider- fringe communities Trifolio-Geranietea sanguinei and ably depending on the substrate, altitudinal aspect and ruderal communities Artemisietea vulgaris classes [5] regional location (Table 1). A description of phytocoe- (Table 1). noses preferred by O. caryophyllacea in the regions Lublin Upland and Roztocze. The species is re- with the highest density of its localities in Poland is corded on sun-exposed hillsides (e.g. Machnowska Mt, given below. Wieprzecka Mt) and in river valleys (Vistula, Wieprz, Pieniny Mts. The properties of soils in the Pien- Huczwa, Kamionka, ãabuþka, Soäokija, Bystrzyca) or iny Mts are different from the soils in the Carpathian on embankments of early medieval earthwork, on cal- Mts which formed from Carpathian flysch [136]. This careous or loess substrate (especially chalky rendzina had a fundamental influence on the distrubtion of local- and rarely chernozems). It prefers open sites where it ities of the species belonging to the genus Orobanche. occurs in xerothermic grasslands of Cirsio-Brachypo- dion pinnati alliance, especially communities Thalic- O. caryophyllacea occurs here on lime soils, shallow tro-Salvietum pratensis, Inuletum ensifoliae, Adonido- or medium deep and highly skeletal rendzinas, less of- Brachypodietum pinnati associations, with a high con- ten on brown soils [34]. The species has been reported tribution of thermophilous fringe species and shrubs of from thermophilous meadows, especially herbaceous the Trifolio-Geranietea sanguinei or Rhamno-Prunetea meadows in the Pieniny Mts (Arrhenatheretum ela- classes [17]. tioris, Gladiolo-Agrostietum capillaris, Anthyllidi- Lower Silesia and Opole region. At present Trifolietum montani associations), at higher altitudes Orobanche caryophyllacea occurs in post-excava- Veratrum lobelianum-Laserpitium latifolium association serpentinite pits in the Kamienny Grzbiet hill tion, xerothermic grassland of the Festuco-Brometea (Nasäawice region) in the ćlöĔa Massif and in the class and alpine grasslands developing on calcareous Ogorzelec (Gäogów Glacial Valley). A unique com- substrate Seslerietea variae class (ecotones of the ther- munity determined as Viscario-Avenuletum pratensis mophilous rock grassland Festucetum pallentis, alpine association has formed there. The community com- rock grassland, endemic community Dendranthemo- prises species of the Festuco-Brometea, Molinio- Seslerietum variae associations, Seslerietea variae Arrhenatheretea, Koelerio glaucae-Corynephoretea class) – community Calamagrostis varia: on slopes canescentis and Trifolio-Geranietea sanguinei classes and screes of S-facing rock gorges, relict pine groves, [88], (Table 1). thermophilous beech groves (Carici albae-Fagetum as- In the Opole province, it occurs at one locality sociation), mountain sycamore maple forest Phylliti- on Góra Gipsowa Mt near Kietrz. The site has persist- do-Aceretum association (Tilio platyphyllis-Acerion ed since the 19th century. The species grows on former pseudoplatani alliance) [4,34,114,118,121]. The cur- gypsum extraction workings in xerothermic grassland rent field research done by the authors could suggest of the Festuco-Brometea class with a high contribution that at present O. caryphyllacea occurs mostly in the of fringe species of the Geranion sanguinei alliance Anthyllidi-Trifolietum montani association, thermo- [104], (Table 1). philous meadows with xerothermic plants, rarely in Pomerania. On the Lower Oder and Vistula riv- communities from Trifolio-Geranietea sanguinei and ers, Orobanche caryophyllacea is mostly found on rich Rhamno-Prunetea classes, usually on mild slopes or at moraine calcium carbonate in xerothermic grasslands flat sites (Table 1). of the Festuco-Stipion alliance, mainly in Potentillo- Nida Basin. Orobanche caryophyllacea occurs Stipetum capillatae associations, often occurring as a in xerothermic grasslands, on hills, in pits often re- mosaic with meadow communities of the Molinio-Ar- maining after excavation or in quarries still operating in rhenatheretea, fringe communities Trifolio-Geranietea the area, of the Cirsio-Brachypodion pinnati alliance, sanguinei or shrub communities of the Rhamno-Pru- especially in the Adonido-Brachypodietum pinnati, netea classes (Table 1). Thalictro-Salvietum pratensis, rarely Inuletum ensifo- On Wolin island, the species occurs on strongly liae, and also on gypsum hills in locally differentiated disturbed slopes of the Szczecin Lagoon (Lubin area), communities Seslerio-Scorzoneretum purpureae and in thermophilous grassland communities with a high Carex glauca-Tetragonolobus maritimus ssp. siliquo- contribution of meadow and ruderal species. It is also sus associations. The only locality in the “Grabowiec” encountered on the roadside in a cleared mixed forest reserve is on gypsum in the oak-hornbeam forest of towards Wapnica (see list of localities, Table 1).

Grabowiec Mt Zamkowa Kozia Mt ugi ä D Gronik glica ñ Homole Okr ki ä 7.2009 31.07.2009 22.07.2012 29.07.2009 23.07.2010 30.07.2009 28.05.2011 Ska Wysokie 9°22’49,4” 49°24’20,1” 49°24’56,9” 49°25’08,2” 49°25’15,6” 49°25’12,5” 50°28’21” C ” 20°33’20” 20°32’52,4” 20°25’20” 20°26’56,6” 20°22’09,4” 20°25’12,9” 20°34’33” o awice ä in Poland...... II ...... II Lubin Nas Dolny Krajnik Stare Table 1 Objezierze Orobanche caryophyllacea Owczary Stary Przylep Mt Plant communities with Wawrzyniec ty II Dwikozy ñ K ów ä Samo- strza dziejowice ö .++.+++.++3+...... III .1+..1++++.+...... III . .1.14211+ .+..+++.4+...... +.II .++...++.+.1...... II -1------252085n - ---10--- 515555555 55551520151051520 55551520151051520 515555555 1++22+1.+3...... III S SW SE SE S S NW SW SE SE S SE S W SE NE SW SW SW t + ++++++ + +++++++++++V +1+.+++++.++1..1+..IV ...1.1..++.III +2+.+++. +++...... +.+.+.++..III + ++++ . . . ++ 00 1015201010101515156040401520205a 25 5010105 5015 5085 15 30 85 - 2557 50 15 77 100 15 25 57 100 25 - 34 90 25 - 42 95 25 27 - 95 25 31 - 80 15 28 10 95 40 31 100 - 20 35 95 - 25 25 60 50 - 35 95 25 20 22 75 50 15 60 100 s 15 23 100 20 80 45 35 90 56 60 y 34 c 33 258 271 244 272 174 92 39 80 65 26 50 174 1052 568 947 515 791 724 240 n kowice S ä 20°51’24” 20°39’45” 20°38’41,6” 23°07’22”18°22’57,9” 21°47’03,9” 14°38’43,1” 14°59’33,5” 14°21’33,7” 14°18’37,9” 14°26’08,2” 16°49’50,8 Su 28.05.2008 19.06.2012 04.06.2009 15.07.2006 27.07.2006 29.05.2009 25.06.2009 25.06.2009 25.06.2009 25.06.2009 08.07.2012 14.07.2011 31.0 50°25’05,51” 50°34’2,8” 50°35’18,2” 50°40’22” 50°44’20” 53°19’37,5” 52°28’23,9” 53°11’27,25” 52°52’53,6” 53°01’35” 53°51’49” 50°53’00” 4 ) 2 ] o rissias Location Date Latitude [N] Longitude [E] Altitude a.s.l. [m] Relevé area (m Exposure Inclination [ Cover of tree layer (%) A Cover of shrub layer B (%) Cover of herbaceous layer C (%) Cover of moss layer D (%) Number of species Orobanche caryophyllacea Ch. Festuco-Brometea, Festucetalia valesiacae Centaurea scabiosa Euphorbia cypa Brachypodium pinnatum Centaurea stoebe Potentilla arenaria Plantago media Stipa capillata Artemisia campestris Phleum phleoides Veronica spicata

© The Author(s) 2014 Published by Polish Botanical Society 106 Renata Piwowarczyk Grabowiec Mt Zamkowa Kozia Mt ugi ä D Gronik glica ñ Homole Okr ki ä ...... II Ska Wysokie .1.1....I awice ä Lubin Nas Dolny Krajnik Stare Objezierze Owczary Stary Przylep Mt Wawrzyniec ty II Dwikozy ñ K ów ä Samo- strza dziejowice ö +1...... +..+..II . .++1+.. .+...+++...+...... II .+...++.+..+...... II .+....+.+..+...... II . +.+...... +. .+. . . .II . .+.+. . . .+.+ ...... 2+2....I ...... ++...... I ...... I .+....+...... 4.3...... I ...... ++..I ...... I .2..+.. .+...... +...... I ...... I .1+..... 1 .+3...... +.+. . .II ...... II 11+.1.. ++++....+...... II .....+.....II +++...... I +1..1...... I ++..+...... I +++...... I +.++...... I +21..... +3...... I +...... +...... I kowice S ä Su a Location Salvia verticillata Achillea pannonica Dianthus carthusianorum Carex flacca Melampyrum arvense Koeleria macrantha Scabiosa ochroleuca Festuca rupicola Filipendula vulgaris Acinos arvensis Adonis vernalis Asperula tinctoria Campanula sibirica Festuca pallens Onobrychis viciifolia Sesleria uliginosa Allium montanum Allium oleraceum Asperula cynan- chica Avenula pratensis Campanula glomerat Carex humilis Carlina vulgaris Scorzonera purpurea Stachys recta Stipa joannis

© The Author(s) 2014 Published by Polish Botanical Society Orobanche caryophyllacea Sm. (Orobanchaceae) in Poland: current distribution, taxonomy, plant communities and hosts 107 Grabowiec Mt Zamkowa Kozia Mt ugi ä D Gronik glica ñ Homole Okr ki ä ....++.III Ska Wysokie awice ä ...... II Lubin Nas Dolny Krajnik Stare Objezierze Owczary Stary Przylep Mt Wawrzyniec ty II Dwikozy ñ K ów ä Samo- strza dziejowice ö .+...... +...... I . ++++ . + . + . . + ...+...++..II . .++..+...... 11..II . .+1+... . 1. .+. . . .+. . .+. .+. .II ...... +++1..II . ++.+.+...... I .2..+.1...... ++....I ...... I .+.2...... +...+.I .+...... +...... I .+...... +.....I 2..2...... I 1 1+12 . 2 + +1 . . .+. .+. .III +.1+1...... I ...... I +1...... I ++...... I +1+.12211132...+...IV +.+..+.3.+...... II +..+.... .+...... I ++...... +.....I ++...... I kowice S ä Su . subsp Location Tetragonolobus maritimus Thesium linophyllon Thymus pannonicus Inula ensifolia Sisymbrium polymorphum Seseli annuum Ch. Trifolio-Geranietea sanguinei Galium verum Medicago falcata Coronilla varia Agrimonia eupatoria Origanum vulgare Trifolium montanum Thalictrum minus Clinopodium vulgare Fragaria viridis Peucedanum cervaria Peucedanum oreoselinum Viola hirta Bupleurum falcatum Anemone sylvestris Campanula rapunculoides Veronica teucrium Vicia tenuifolia Valeriana angustifolia siliquosus

Grabowiec Mt Zamkowa Kozia Mt ugi ä D Gronik glica ñ Homole Okr ki ä Ska Wysokie ....+1+.IV awice ä ...... I Lubin Nas Dolny Krajnik Stare Objezierze Owczary Stary Przylep Mt Wawrzyniec ty II Dwikozy ñ K ów ä Samo- strza dziejowice ö

+...... +.++..II .+....+. ..2+...1...II .++...... +...... 1.+...I ...... +.++..I ...... +.++..I . . . .+...... +. . . .+.I .+...... +.+...I ...... +..+..+..I ...... +...I .+..+.. ..+...... I .++...... +.+...... +...... +...I ...... ++.I . .+...... +...I ...... ++..I . .+....3+...+...... II . ....+.+..+...... I .+...... +...... I 4 23122. . 1+11+2333+.V ..3....22..II 11+..... + ++++++ + ++3 +++.. .++ . . .1+1. .+. .III +++2.+11...... 23..III ...... ++..II +.+2...... I +2+..... +.+...... +.....I kowice S ä Su . agg Galium mollugo Location Ch. Molinio-Arrhenatheretea Dactylis glomerata Achillea millefolium Arrhenatherum elatius Festuca rubra Knautia arvensis Leucanthemum vulgare Plantago lanceolata Anthyllis vulneraria Campanula patula Centaurea jacea Galium boreale Leontodon hispidus Taraxacum officinale Trifolium pratense Trifolium repens Vicia cracca Rumex thyrsiflora Festuca pratensis Lathyrus pratensis Phleum pratense Prunella vulgaris Tragopogon pratensis Ch. Artemisietea vulgaris Echium vulgare Anchusa officinalis Artemisia vulgaris

Grabowiec Mt Zamkowa Kozia Mt ugi ä D Gronik glica ñ Homole Okr ki ä Ska Wysokie .+.+.+. .I awice ä Lubin Nas Dolny Krajnik Stare Objezierze Owczary Stary Przylep Mt Wawrzyniec ty II Dwikozy ñ K ów ä Samo- strza dziejowice ö . ....+.+...... I ...... +....+I ...... +....1I . .+.....++1....++.+II ...... +++..1.II ...... 1.....+..+I ...... +...... +I ...... 12..I ...... +.++..I ...... +..++I ...... 23I ...... ++.....I ...... ++.I ...... 2...1I ...... ++I . .+...... +2+.++.II ...... +..1+...... 1.....II ...... +.....I .+..+.. . .+...... ++...... II . ..+.... .+...... I +...... +2..I kowice S ä Su Location Descurainia sophia Geranium robertianum Geum urbanum Ch. Rhamno-Prunetea Prunus spinosa C Cotoneaster integerrimus B Crataegus monogyna B Prunus spinosa B Crataegus monogyna C Rosa dumalis B Ch. Querco-Fagetea Acer pseudoplatanus C Corylus avellana C Corylus avellana B Epipactis helleborine Lonicera xylosteum C Melica nutans Pulmonaria obscura Ch. Epilobietea angustifolii Fragaria vesca Calamagrostis epigejos Rubus idaeus Verbascum nigrum Ch. Agropyretea intermedio-repentis Convolvulus arvensis Falcaria vulgaris

Grabowiec Mt Zamkowa Kozia Mt ugi ä D Gronik glica ñ Homole Okr ki ä Ska Wysokie awice ä ...... II ...... II .....++..II Lubin Nas Dolny Krajnik Stare Objezierze Owczary Stary Przylep Mt Wawrzyniec ty II Dwikozy ñ K ów ä Samo- strza dziejowice ö ...... +34....I ...... ++.....I .+...... +...I ...... ++..I ...... +.+...+...... I ...... +..2...... I . ++...... +.++2++.III ...... ++...+++.II . . .1+...... +.++..II ...... +....+1..I ...... +.2.+..I ...... +.22.I ...... +...++.....I .+...... +...+.I ++.+. . . . .+ +++.. .+ . ++.1++.++..III .+..+++...+III +++...+. ++11..+.++...... II + ++++ . + . . . + .+.+. . . .+ ...... II +1.++.. +.+....+...... +..II ....+...... II +++..... kowice S ä Su Location Ch. Seslerietea variae Sesleria varia Saxifraga paniculata Ch. Nardo-Callunetea Polygala vulgaris Cuscuta epithymum Potentilla erecta Ch. Koelerio glaucae-Corynephoretea canescentis Sedum acre Trifolium arvense Others Thymus pulegioides Vincetoxicum hirundinaria Pimpinella saxifraga Salvia pratensis Orobanche lutea Briza media Hypericum perforatum Ranunculus acer Festuca trachyphylla Primula veris Sanguisorba minor Agrostis capillaris Digitalis grandiflora Laserpitium latifolium Medicago lupulina Sedum maximum

2, 1, 19, Ge- P. py- 12 (1). A 5(1), A Fraxinus Cynoglos- 10(1), Melandrium 8, Frangula alnus Papaver rhoeas Grabowiec A 5, B 1, A 19, 2, Elymus hispidus Gladiolus imbricatus Ch. Nardo-Callune- A. platanoides Sanicula europaea Eryngium campestre A 17(2), B 14(1), C 14; A A 18(2), C 18, A Viscaria vulgaris Mt 16, 15; s.l. 12(2), 6, 18, Zamkowa 19, B 16, Anthericum ramosum Pyrus communis Conyza canadensis Picea abies 2, 17, Lathyrus tuberosus Kozia Mt Festuca ovina Fagus sylvatica Scabiosa lucida Carex praecox Melampyrum nemorosum Salvia glutinosa Viola mirabilis 14,

5, 14, ugi B 19, sp. ä 11, 6(1), 19, D Acer pseudoplatanus Gronik Gentiana pneumonanthe A 4, B 4(1), C 9, A 17, Pulsatilla pratensis glica Euphrasia ñ 12, Ch. Agropyretea intermedio-repentis: 18, Poa nemoralis Ch. Stellarietea mediae: Chaerophyllum aromaticum 18, Phyteuma orbiculare Verbascum thapsus 12; Euonymus verrucosa Asparagus officinalis Pinus sylvestris Lithospermum officinale Ch. Trifolio-Geranietea sanguinei: 15, 17, 8(2), Ch. Querco-Fagetea: 14, 4; Homole Okr 14, 19(1), B 18, B 19(3), C 19; 2; Carex spicata Deschampsia caespitosa ki ä C. nutans Potentilla heptaphylla 14, 15(2), 6, Ska 4, Wysokie Abies alba Phyteuma spicatum Listera ovata Arabis hirsuta Carduus glaucus Trifolium campestre Stellaria graminea 19, 9, 6, Cruciata glabra Sambucus nigra

9(1), awice ä Teucrium chamaedrys Verbascum lychnitis B 19, Bellis perennis 8, Carduus crispus 16, Silene otites 17(2), 9, Calamagrostis varia 6, ...... I Millium effusum Lubin Nas Peucedanum alsaticum 18(1), B 18, C 14, Ch. Vaccinio-Piceetea: 19, 15; Ch. Seslerietea variae: S. columbaria Cerasus avium Anthericum liliago Lavathera thuringiaca T. medium sp. A 11; 2, 18, 2, 8, 18, 5, B C 12, A Dolny Berteroa incana Krajnik Angelica sylvestris Sorbus Senecio vernalis Ch. Epilobietea angustifolii: O. mayeri 9, 2, 16, 11(2), C 14; Urtica dioica Stare Clematis alpina 12, Objezierze Betula pendula O. kochii Melittis melisophyllum Scabiosa canescens Alyssum montanum Lappula squarrosa Trifolium alpestre Senecio nemorensis 4, 10, 12, B 15, sp. 17, A 19(2), A 19(1), Senecio jacobaea 18(1), major Viburnum opulus 14, Helichrysum arenarium Artemisia absinthium B 14(1), C14, subsp. Tanacetum vulgare Alchemilla Ch. Molinio-Arrhenatheretea: B 2(1), 5, Oxytropis pilosa Owczary Stary Przylep Ajuga genevensis B 1, 4, Carpinus betulus 5, Lonicera xylosteum B 5(1); 18, Calamagrostis villosa Mt Rhinanthus glaber 19, Polygonatum odoratum Orobanche alba 2, 17, Wawrzyniec 1, Juniperus communis 15, A. setacea Linum flavum Solidago gigantea Rhamnus cathartica 9, 14, Tilia cordata 4, Ch. Artemisietea vulgaris: 11, 19, Astrantia major 12; Aesculus hippocastanum

Lilium martagon 14, B 1, C 14, Orchis ustulata Reseda phyteuma 17, ty II Dwikozy 7, nemorum ñ 1, . Achillea collina

K S Others: Bupleurum longifolium Libanotis pyrenaica Melilotus albus 8; Jovibarba sobolifera 16, 19, Hieracium bauhinii 8, ów Rhinanthus minor 1, ä O. spinosa 18, Ch. Koelerio glaucae-Corynephoretea canescentis: Samo- 12, strza Cornus sanguinea 17, Asarum europaeum 13; Geranium phaeum 19, Viola arvensis B 5, Ranunculus bulbosus 7, 19(4), Stellaria holostea dziejowice Lathyrus sylvestris Melandrium album ö 19, 5, 1, Ch. Betulo-Adenstyletea: Ononis arvensis A 19(4), A Hieracium murorum H. umbellatum Lotus corniculatus 4, +...... +..I .++...... +.+...... I ...... ++..I ...... +....+...... I . ....+.+...... I . .+.+...... +..+..I . ....3..1...... I ...... ++..I . . .+...... +. . . . .I . . ...+.. ..+...... I ...... +....+I ...... I . .+..+...... +.+...... I ...... +...+.I ...... +...2.I ...... I . .+..+.. 11; +..+...... I +...... +.....I 18, Vicia hirsuta kowice S 17, ä Cerasus fruticosa

14(1), Su 11, Heliantchemum nummularium Inula hirta Galium odoratum 2, Aegopodium podagraria Stachys sylvatica 14, Galium aparine Quercus robur 19, 8, Rumex crispus Mycelis muralis C 18, sp. B C 17, B 1, C 14, 18, sp. sp. Gymnadenia conopsea B 2, C 11, Hieracium pilosella Potentilla argentea Leontodon autumnalis Location Veronica chamaedrys Alyssum alyssoides Anthoxanthum odoratum Arenaria serpyllifolia Camelina microcarpa Carex hirta Carlina acaulis Festuca Heracleum sphondyllium Linum catharticum Orchis militaris Quercus robur C Rosa Rosa Rubus caesius Senecio vulgaris Solidago virgaurea Polypodium vulgare Turritis glabra Actaea spicata excelsior raster rubrum E. repens Ch. Rhamno-Prunetea: ranium sanguineum 10, sum officinale 16, Gentiana cruciata tea: Sorbus aria Sporadic: Ch. Festuco-Brometea, Festucetalia valesiacae: B 19,

DISCUSSION though the genus Myrmica (Formicidae) and other genera were also noted (Piwowarczyk unpublished). In Poland Orobanche caryophyllacea mainly The abundance of Orobanche caryophyllacea occurs in the Maäopolska Upland (chiefly the Nida Ba- populations varies regionally in Poland and fluctuates sin and Middle Vistula River Valley), Lublin Upland, considerably annually. Its populations are usually not Roztocze, PrzemyĈl Foothills, Sudeten Foreland, Sile- very abundant and mostly comprise 5 to 20 specimens. sian Basin, Pieniny Mts and Pomerania: Lower Vistula The most abundant populations are recorded in Low- and Oder rivers, Wolin island. A total of 160 localities er Silesia (Kamienny Grzbiet) – over 300 specimens of O. caryophyllacea are known in Poland, of which [88], in the Nida Basin, e.g. Kików and Chomentówek ca. 120 have been confirmed and documented by her- – 200 specimens at each, Gartatowice, Samostrzaäów, barium specimens or photographs. Approximately Grabowiec, Górki – 100 specimens each [5]. The pop- 60 sites have been confirmed after 2000. Many sites ulations in the Lublin Upland are also quite abundant were noted in the 19th century and are now historical. and 50 specimens were recorded in Kñty, DobuĔek or O. caryophyllacea is one of the most sensitive species Przeorsk [17]. to changes in the phytoconoesis structure among spe- Orobanche caryophyllacea is strictly protect- cies of the genus Orobanche found in Poland. The ver- ed in Poland (Regulation 2012). It is also included in ification showed that the species had been incorrectly the regional red lists and books: in Gdaþsk Pomera- reported, for instance, from the ćlñsko-Krakowska nia, CR [139]; Western Pomerania and Wielkopol- Upland, Maäopolska Upland and adjacent areas. It was ska, E [140]; Sudeten Mts, EN [141]; Opole prov- usually confused with O. lutea and O. kochii, which ince, CR [104], Lower Silesia province, endangered, are frequent there. A verified and updated distribution EN [142], Kujawy-Pomerania region, R (rare) [143], map (Fig. 1) of the ATPOL map [137,138] shows con- Poäudniowopodlaska Lowland, DD [144], Proszowicki siderable differences and it is a result of errors and dif- Plateau, EN [145], Maäopolska Upland, VU [146]. The ficulties in determining the genus Orobanche. species is also vulnerable in neighbouring countries, The herbarium revision and field investigations i.e. Germany [147,148] and the Czech Republic [149]. also showed that Orobanche teucrii reported from Po- The progressing process of secondary succes- land in the Polish part of the Pieniny Mts [34, after sion of tree and shrub vegetation, the density of the Jasiewicz] or in the Lublin region [35] was reported herb layer and the cessation of former management incorrectly and the species was in fact O. caryophyl- methods are the main threats to Orobanche caryophyllacea. lacea. The localities are also often situated near arable Orobanche caryophyllacea prefers xerothermic fields where they are exposed to chemical agents. The grasslands of the Festuco-Brometea class, fringe com- influence of invasive species also poses a consider- munities of the Trifolio-Geranietea sanguinei class, able threat. Heracleum sosnowskyi has been moving thermophilous meadows (Arrhenatherion elatioris alli- worryingly close to the O. caryophyllacea popula- ance) of the Molinio-Arrhenatheretea class, rarely al- tion (Gipsowa Góra). Mass self-sowing of Solidago pine grasslands Seslerietea variae class developing on gigantea and Calamagrostis epigejos are observed at calcareous substrate, cleared oak-hornbeam forests of many sites. Many localities are at disturbed sites or at the Querco-Fagetea class, such as sycamore maple in operating excavation sites (limestone, serpentinite). the Pieniny Mts, “Grabowiec” reserve in the Nida Ba- Only a minority of localities are protected: within sin. It colonizes soils such as shallow rendzinas, loess national parks – Wolin and Pieniny National Parks; and chernozems. The species is recorded in lowland reserves – e.g. Brodogóry, Stary Przylep, Bielinek, areas, from 29 m in Pomerania and Wielkopolska, 140– Zbocza Päutowskie, Góra ćw. Wawrzyþca, Gipso- 270 m in the belt of the Polish Uplands to 1050 m a.s.l. wa Góra, Stawska Góra, Machnowska Góra, Skarpa in the Pieniny Mts (Wysokie Skaäki). DobuĔaþska, Homole, Grabowiec, Zimne Wody; as Orobanche caryophyllacea is oligophagous and ecological sites, e.g. Dwikozy, Owczary, or within parasitizes species of the family Rubiaceae, genus Ga- Natura 2000 sites. This does not ensure the preserva- lium: G. mollugo, G. verum, rarely G. boreale, spo- tion of the species as active protection, which is not al- radically G. odoratum and Cruciata glabra. O. lutea ways introduced, is needed. At least some of the locali- and O. caryophyllacea are some of the earlierst flow- ties should be protected as ecological sites and active ering species of the genus Orobanche in Poland. The protection measures should be implemented: mowing flowering peak is observed in late May and early June. or grazing of excessively developing field and herba- O. caryophyllacea is characterized by the fact that it ceous vegetation, while preserving the unique mosaic emits a strong scent resembling the smell of cloves. systems of grasslands and fringe communities, and na- Representatives of the genus Bombus (Apidae) were ture monitoring. The abundance of the host species of the most frequently observed pollinating insects, al- the genus Galium should also be monitored.

Species biodiversity at the localities occupied by 9. P iwowarczyk R. Orobanche alba subsp. alba and sub- Orobanche caryophyllacea is also high. Many species sp. major (Orobanchaceae) in Poland: current distribution, extremely rare in Poland were observed when phyto- taxonomy, plant communities, hosts, and seed micromor- sociological relevés were performed in the communi- phology. Biodiv Res Conserv. 2012; 26: 23–38. http://dx. ties with O. caryophyllacea. These include: Achillea doi.org/10.2478/v10119-012-0005-6 setacea, Adonis vernalis, Anthericum liliago, Cerasus 10. P i w o w a r c z y k R . Orobanche purpurea (Orobanchace- fruticosa, Linum flavum, Orchis ustulata, Orobanche ae) in Poland: current distribution, taxonomy, plant com- alba subsp. major, O. elatior, O. mayeri, Oxytropis munities, and preferred hosts. Biodiv Res Conserv. 2012; pilosa, Phyteuma orbiculare, Peucedanum alsaticum, 26: 73–81. http://dx.doi.org/10.2478/v10119-012-0006-5 Reseda phyteuma, Scorzonera purpurea, Sesleria var- 11. P i w o w a r c z y k R . A revision of distribution and his- ia, Stipa joannis, and other (Tab. 1). torical analysis of preferred hosts of Orobanche ramosa (Orobanchaceae) in Poland. Acta Agrobot. 2012; 65(1): 53–62. http://dx.doi.org/10.5586/aa.2012.043 Acknowledgements 12. P i w o w a r c z y k R . A revision of distribution and the eco- The author thanks the herbarium curators and logical description of Orobanche picridis (Orobanchaceae) the contributors who made the data on the species avail- at the NE limit of its geographical range from Poland and able and helped during the field investigations. This Ukraine. Acta Agrobot. 2012; 65(1): 91–106. http://dx.doi. work was supported by the Polish State Committee org/10.5586/aa.2012.047 for Scientific Research (KBN grant no. NN303357733 13. 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© The Author(s) 2014 Published by Polish Botanical Society 118 Renata Piwowarczyk rozwijajñce siö na podäoĔu wapiennym Seslerietea nach (Wysokie Skaäki). NaleĔy do gatunków ĈciĈle variae, przeĈwietlone zbiorowiska grñdowe (np. jawo- chronionych w Polsce. Znajduje siö równieĔ na regio- rzyny w Pieninach, grñd na gipsie w rezerwacie Gra- nalnych czerwonych listach i w ksiögach. ZagroĔe- bowiec w Niecce Nidziaþskiej) z klasy Querco-Fage- niem dla gatunku jest przede wszystkim postöpujñcy tea. Gatunek notowany jest na terenach nizinnych, od proces sukcesji wtórnej roĈlinnoĈci drzewiastej i krze- 29 m na Pomorzu i Wielkopolsce, poprzez 140–270 m wiastej, a takĔe zwarcie warstwy zielnej, oraz zanie- w pasie WyĔyn Polskich, do 1050 m n.p.m. w Pieni- chanie dawnych metod uĔytkowania.

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