Holoparasitic Orobanchaceae in Georgia (Caucasus): taxonomic revision, diversity, distribution, habitats and host range Author Piwowarczyk, Renata 0000-0003-0507-7835 * Center for Research and Conservation of Biodiversity, Institute of Biology, Jan Kochanowski University, 7 Uniwersytecka St., 25 - 406 Kielce, Poland; renata. piwowarczyk @ ujk. edu. pl; renka 76 @ wp. pl; https: // orcid. org / 0000 - 0003 - 0507 - 7835 renata.piwowarczyk@ujk.edu.pl Author Pedraja, Óscar Sánchez 0000-0002-4383-4588 E- 39722 Liérganes (Cantabria), Spain; osanchez @ farmalierganes. com; https: // orcid. org / 0000 - 0002 - 4383 - 4588 osanchez@farmalierganes.com Author Khutsishvili, Manana 0000-0002-7404-9574 National Herbarium of Georgia, Institute of Botany, Ilia State University, Tbilisi, Georgia; mananakhuts @ yahoo. com; https: // orcid. org / 0000 - 0002 - 7404 - 9574 mananakhuts@yahoo.com Author Kharazishvili, Davit 0000-0003-0507-7835 * Center for Research and Conservation of Biodiversity, Institute of Biology, Jan Kochanowski University, 7 Uniwersytecka St., 25 - 406 Kielce, Poland; renata. piwowarczyk @ ujk. edu. pl; renka 76 @ wp. pl; https: // orcid. org / 0000 - 0003 - 0507 - 7835 & Batumi Botanical Garden, 6411, Batumi, Mtsvane Kontskhi, Georgia; davit. kharazishvili @ gmail. com; https: // orcid. org / 0000 - 0001 - 9254 - 9954 * author for correspondence & * Center for Research and Conservation of Biodiversity, Institute of Biology, Jan Kochanowski University, 7 Uniwersytecka St., 25 - 406 Kielce, Poland; renata. piwowarczyk @ ujk. edu. pl; renka 76 @ wp. pl; https: // orcid. org / 0000 - 0003 - 0507 - 7835 renata.piwowarczyk@ujk.edu.pl text Phytotaxa 2023 2023-07-24 604 1 1 103 http://dx.doi.org/10.11646/phytotaxa.604.1.1 journal article 10.11646/phytotaxa.604.1.1 1179-3163 8181674 6. Phelipanche nana (Rchb. f.) Soják (1972: 130) . Type:— CROATIA ( lectotype designated by Jeanmonod, 2007: 195 ):—Fiume [Rijeka], 1842, Noë no . 1352 (G00001468!; isolectotype: G00074293!). Basionym:— Phelypaea nana Reichenbach f. (1862: 88) . Homotypic synonyms:— Orobanche nana (Rchb. f.) Beck (1890: 91) ; O. ramosa subsp. nana (Rchb. f.) Couthino (1913: 566) ; Phelypaea ramosa var. simplex Visiani (1847: 180) ; P. mutelii var. nana Reut. in Candolle (1847: 9) , nom. superfl. illeg. ( Jeanmonod 2007: 195 ); Phelipanche mutelii var. nana (Reut.) Uhlich & Rätzel in Uhlich (2015: 82) , nom. illeg. ( Turland et al. 2018 : Art. 6.10). Heterotypic synonyms:— O. ramosa var. penicillata O. Kuntze (1887: 225) . Misapplied names:— Phelipanche oxyloba auct. (saltem p.p.), non Phelypaea oxyloba Reut. in Candolle (1847: 9) ; Phelipanche mutelii auct. (saltem p.p.), non Orobanche mutelii F.W. Schultz in Mutel (1835: 353). Fig. 15 . General distribution: —Old World (Mediterranean Basin, Balkans, Asia Minor , the Caucasus). Alien in the Americas. Distribution: —Scattered localities mainly in the western, central and southeastern parts ( Imereti , Samtskhe-Javakheti , Mtskheta-Mtianeti , Kakheti , Kvemo Kartli and Shida Kartli provinces), rarely in Adjara and Abkhazia , as well as in the north in Samegrelo-Zemo Svaneti prov. ( Fig. 16 ). Probably more frequent, but because of its early flowering and small habit, it might be overlooked. Habitat: —Rocky slopes, steppe, meadows, pastures, ruderal habitats, roadsides, railway areas, lawns, city greens, edges of forests and shrublands, from a few meters above sea level (in Sokhumi on the Black Sea in Abkhazia ) to usually 500‒1200(2000) m. Hosts: —Parasitic usually on native plants, here mainly on Caucalis platycarpos L., Orlaya grandiflora (L.) Hoffm. (syn. Caucalis daucoides L.) ( Apiaceae ), Erodium cicutarium (L.) L’Hér., Geranium robertianum L. ( Geraniaceae ), Sherardia arvensis L. ( Rubiaceae ), Plantago lanceolata L. ( Plantaginaceae ), Trifolium aureum Pollich , Medicago brachycarpa M. Bieb. ( Fabaceae ), Alyssum hirsutum M. Bieb. , A. murale Waldst. & Kit. , Diplotaxis muralis (L.) DC. ( Brassicaceae ), Potentilla recta L. ( Rosaceae ). Phenology: —Flowering (April) May–June, fruiting June to July. Conservation status: —Least Concern (LC). The species has a rather wide distribution in Georgia (extent of occurrence [EOO] is ca. 43,000 km 2 , and area of occupancy [AOO] is ca. 5000 km 2 ). Populations consist usually of a few to over 100 shoots. The species may be endangered in some localities, especially those in ruderal habitats, especially urban, industrial, which may undergo dynamic transformations. Notes :—Within the entire range of the species, various host families are shown, in Georgia it has been found on such hosts as Apiaceae , Geraniaceae , Rubiaceae , Plantaginaceae , Fabaceae , Brassicaceae , and Rosaceae . Sometimes, within one population, it parasitises on two or more hosts from different families. In the area of the Caucasus, P. nana finds the limit of its natural range. The P. nana collected in Georgia are very variable and combine typical forms with less typical forms (erect-patent flowers versus patents and corollas with less curved dorsal line). All of the Georgia host-plants mentioned here are common to this species throughout the rest of its range. The species described from Croatia shows very typical forms in the central and western Mediterranean but these forms are less typical towards the east (Eastern Mediterranean - Middle East). On the other hand, P. mutelii was described in NW Africa ( Algeria ) and is a completely different species from a morphological and genetic point of view ( Carlón et al. 2005: 60 ; Carlón et al. 2008: 11–12 ). Its area of distribution is smaller and most likely will not reach Asia. Supporting our previous comment, Kuntze (1887) recorded this species from Astara ( Azerbaijan ), sub “ O. ramosa L. var. nana Reut. in DC.” and described from Tbilis [ Tbilisi , Georgia ] a new var. that he called “ penicillata ”, which is subordinate to O. ramosa . According to this author, this variety would be characterised, among other morphological characteristics, due to its hairy anthers [“Antherae apice tantum pilis paucis longis penicillatae”] in front of the glabrous or almost of the type [“α. normalis ”]. We believe this taxon must be treated as a morphological variant included in P. nana , which would be characterised, besides, by its flowers, erect-patent with corollas with longer floral tube and less curved dorsal line than what is normal in this species and calyces, bracts and scale-leaves longer than the type of this species. This variety, which we tipify below, is relatively frequent in the region and its surroundings. Orobanche ramosa var. penicillata O. Kuntze in Trudy Imp. S.-Peterburgsk. Bot. Sada 10: 225 (1887) [“β”]. Ind. loc.: “ Tiflis [ Tbilisi , Georgia ]” ( Kuntze 1887: 225 ) . Lectotype (here designated):—1. “( Trionychon ) Phelipaea auct. / Orobanche ramosa L. / v. penicillata m. / Antherae apice tantum pilis paucis longis / penicillatae; var. intermedia / α. normalis et γ. indica antheris lanatis / Tiflis IV 86 / Herbarium of Otto Kuntze ”. – 2. “ Holotype of: Orobanche ramosa β penicillata O. Kuntze / 1887 Acta horti Petroppol. 10: 225 / Tiflis / Apr 1886 / verif. T. Zanoni ”. – 3. “ Herbarium of Otto Kuntze / Presented by Mr. Andrew Carnegie / 1908”. – 4. “ Tiflis / = Tbilisi , Georgia ” (NY312476 - ID: c2bc3417-3ffe-488d-9757-106d5ac0b6a0, the sheet contains two specimens , we consider the well-developed specimen to the left of the sheet as the lectotype of the name, the other poorly developed specimen would be the isolectotype, also in the upper right corner we can see part of the base of the first). FIGURE 15. Phelipanche nana . A–D. General habit. E–J. Flower, side view. K–O. Flower, front view. P–U, Y. Representative hosts (P. Medicago brachycarpa . R. Trifolium aureum . S. Caucalis platycarpos . T. Erodium cicutarium . U. Plantago lanceolata . Y. Sherardia arvensis ). W–Z. Representative habitats (W. Tbilisi, Abanotubani distr. Y. Sighnaghi. Z. Tbilisi–Turtle Lake). Photos by R. Piwowarczyk. FIGURE 16. Distribution of Phelipanche nana in Georgia. Specimens examined :— GEORGIA . Abkhazia : surroundings Sokhumi, 15April 1888 , A.P.Ivanov (MW0717069) [as P. ramosa ]; prov. Kutaisi, Suchum., in fruticetis, 7 May 1902 , Alexeenko (TGM53205) [as O. ramosa by Melikischvili]; Abchasien ( Albow !), und bei Jurjewskoje, ( G. Woronow !) ( Beck 1930: 72 , sub O. nana Noë ); Adjara : Batumi, railway station, J. Zalewska-Gałosz , May 2015 (phot.); Imereti prov. : Kutaisi, near monastery, 13 July 2011 , M. Podsiedlik (phot.); Kakheti prov. : S of Udabno, Davit Gareja monastery, rocky slopes, on Galium ?, Alyssum sp. , 41°26’56’’N , 45°22’30’’E , 700 m , 26 May 2014 , R. Piwowarczyk (KTC); Gare Kakheti , Davit Gareja, Fischer et al. (2018) [phot.] [as O. mutelii ]; S and SE of Kvemo Bodbe, steppe hills, 41°32.21’N , 45°54.081’E , 765 m , 41°32.351’N , 45°54.326’E , 750 m , on Alyssum , Caucalis , etc., 9 May 2019 , 12 May 2019 , R. Piwowarczyk (KTC); between Zemo Kedi and Kalvata, rocky slopes, roadsides, on Medicago , Erodium , 41°24.729’N , 46°26.733’E , 560 m , 12 May 2019 , R. Piwowarczyk (KTC); rocky slopes in Singhaghi in direction to Tsnori, near serpentine road, on Sherardia , Plantago , 41°37.4’N , 45°55.7’E , 13 May 2019 , R. Piwowarczyk (KTC); Sighnaghi NE, side path from main road to Tsnori, rocky slopes, shrublands, 41°37.2’N , 45°55.8’E , 640 m , 13 May 2019 , R. Piwowarczyk (KTC); road between Singhaghi and Anaga, rocky slopes near serpentine road, on Plantago , 41°38.115’N , 45°54.605’E , 630 m , 14 May 2019 , R. Piwowarczyk (KTC); Sighnaghi N, near old defense wall, roadside, near Lolashvili St., on Sherardia , 41°37.3’N , 45°55.2’E , 788 m , 14 May 2019 , R. Piwowarczyk (KTC); Kachreti, [in the village, near Veronica , and probably also Petroselinum crispum ], 41°38’42.5”N , 45°40’51.4”E , [ca. 550 m ], 5 May 2021 , Naturalist55708 (phot., https://www. inaturalist.org/observations/77434378); Dedoplistskaro NW, [Eagle Canyon], 41°29’24.1”N , 46°05’48.4”E , 26 April 2023 , Hughkent (phot., https://www.inaturalist.org/observations/156774443); Kvemo Kartli prov. : Rustavi NW, lawn at the petrol station Socar, Davit Kldiashvili St., on Trifolium aureum , 41°33.849’N , 44°58.58’E , 387 m , 8 May 2019 , R. Piwowarczyk (KTC); near the village of Malkhazovka [until 1940 - Krasnogorsk, from 2012 - the new name is Sataple], Tifl. distr., 1 June 1924 , M. Timofeev (TBI); Kvemo Kartli , Umgebung von Tbilisi , Kumisi tba (kleiner See bei Kumisi), S Tbilisi , 17 May 2001 , G. Schneeweiss & A. Tribsch (WU0022589) [as P. aegyptiaca ]; Didgori [between Kojori and Kiketi, W of Azeula Fortress], 41°38’29.5”N , 44°40’24.4”E , A. Kvelashvili (phot., https://www.inaturalist. org/observations/163693149); Mtskheta-Mtianeti prov. : prov. Tiflis , distr. Duschet., inter Mtschet et p. Karsani, in fruticetis, 3 May 1922 , B. Schischkin (TGM53193); Mtskheta-Mtianeti , Talgebiet des Flusses Mtkvari (Kura), Shio Mqvime bei Mtsxeta, Umgebung der Kirche, 17 May 2001 , G. Schneeweiss & A. Tribsch (WU0022590, WU0022601, WU0022602, WU0022603, WU0022604, WU0022605) [as P. aegyptiaca ]; between Zahesi and Mtskheta, below Jvari monastery, rocky slopes, steppe slopes of Mtkvari river, on Erodium , 41°50’17”N , 44°43’56”E , 550 m , 5 May 2019 , R. Piwowarczyk (KTC); rocky slopes, roadsides, near the road from Mtskheta to Karsani, on Erodium , 41°50.037’N , 44°42.878’E , 585 m , 5 May 2019 , R. Piwowarczyk (KTC); Kartli, Shiomgvime, 2018, Fischer et al. (2018) [phot., as O. mutelii ]; Mestia, Samegrelo-Zemo Svaneti prov. : 43°02’53.1”N , 42°43’13.7”E , [ca. 1500 m ), 22 June 2022 , Gymnasticus (phot., https://www.inaturalist.org/observations/123382181); Gorovani, [in the village, home garden?], 41°52’59.8”N , 44°39’52.6”E , [ca. 640], 26 August 2021 , G. Natsvlishvili (phot., https://www.inaturalist.org/ observations/93239040); Samegrelo-Zemo Svaneti prov. : the route from Mestia towards the Koruldi lakes, on the side of the trail, 43°03’45.5”N , 42°43’58.2”E , 1950 m , 30 June 2015 , M. Zając (KTC); between Mestia and Murshkeli, W of Murshkeli, 43°03’22.6”N , 42°46’49.9”E , [ca. 1850 m ], 11 June 2022 , J. Pietruszewski (phot., https://www. inaturalist.org/observations/121288023); Samtskhe-Javakheti prov. : Prov. Tiflis , distr.Gori, surroundings of Borjomi, between villages Tsaghveri and Tsemi, 3 June 1917 , W. Kozlovsky (TBI); Borjomi, park, 13 June 1943 , A. Dmytrieva (BATU); NE of Borjomi, Kortaneti, a rocky slope above an asphalt road, on Caucalis , 41°53’45”N , 43°26’55”E , 780–790 m , 31 May 2014 , R. Piwowarczyk (KTC); 10 km ESE of Borjomi, roadsides, rocky slopes, S exposure, on the road between Tsaghveri and Timotesubani, on Caucalis , 41°48’15”N , 43°29’52”E , 1075 m , 2 June 2014 , R. Piwowarczyk (KTC); on direction to Vale, slope near the road, on probably Hypochaeris , 13 July 2015 , R. Piwowarczyk (KTC); between Daba and Tsaghveri, NW of Tsaghveri, rocky slope with tragacanth on Gujaretistskali river valley, on Medicago , 41°48’21.9”N , 43°28’24.8”E , 1030 m , 15 June 2022 , R. Piwowarczyk (KTC); Daba E, overgrown slopes with a predominant hornbeam, on Caucalis , 41°48’44.6”N , 43°27’34.1”E , 1005 m , 15 June 2022 , R. Piwowarczyk (phot.); SE of Rustavi, rocky slopes of Mtkvari river, on Alyssum hirsutum , 41°36’20”N , 43°08’29”E , 1030–1100 m , 3 June 2014 , R. Piwowarczyk (KTC); between Skhvilisi and Didi Pamaji, in direction to Vale, steppe, rocky, hills, on Plantago , 41°37’37.6”N , 42°54’24.1”E , 1090 m , 13 July 2015 , R. Piwowarczyk (KTC); Shida Kartli prov. : Khashuri distr., Vakha, 42°02’26.7”N , 43°42’04.4”E , 31 May 2014 , C. Hough [phot., https://www.inaturalist.org/ observations/1375978]; Kartli, Gori, Fischer et al. (2018) [phot., as O. mutelii ]; between Mrgvali Chala and Kaspi, N of Kaspi, E parts of Kvernaki range, semi-arid eroded hills, probably on Alyssum , 41°57’14.3”N , 44°25’08.6”E , 645 m , 8 June 2022 , R. Piwowarczyk (KTC); Tblisi prov.: Hortus Botanicus Tiflisiensis, 3 July 1927 , Koenig (TBI); Caucasus, Tiflis district , 5 May 1883 , Anonymous (MW0723346) [ P. nana var. penicillata ]; Tiflis , Dabachanka [part of r. Dabakhanka gorge in Botanical Garden], 30 May 1919 , Grossheim (ERE7317) [ P. nana var. penicillata ; as O. ramosa ]; Transcaucasia, Tiflis in familis fl. Dabachanca, 4 May 1919 , 13 May 1919 , 30 May 1919 , 6 June 1920 , Grossheim (ERE7330, ERE7331, ERE7333, TGM42222) [as O. ramosa , O. purpurea , O. aegyptiaca ]; Transcaucasia, Tiflis , in rute st-Davidi [Mama Daviti Church], 1 June 1919 , Grossheim (ERE7318) [as O. aegyptiaca ]; Transcaucasia, Tiflis , in fruticetus, 25 May 1919 , A. Grossheim (ERE7319) [as O. aegyptiaca ]; Tbilisi , Botanical Garden, 41°41’04.5”N , 44°47’55.8”E , [ca. 555 m ], 19 May 2021 , Mariam2222 (phot., https://www.inaturalist.org/observations/79371706); Tbilisi , Botanical Garden N part, by the path near rocky slope in gorge, behind the Mediterranean Collection, 41°41’13.9”N , 44°47’58.5”E , 475 m , 3 June 2022 , R. Piwowarczyk (KTC); Tbilisi , Botanical Garden, rocky hill west of the stone bridge and in front of a modern glass building, N of Mirza Shafi Str., on Potentilla , 41°41’12.5”N , 44°47’50.7”E , 510 m , 3 June 2022 , R. Piwowarczyk (KTC); Tbilisi , Botanical Garden, rocky places on the edge of the thicket below the stone bridge, 41°41’13.4”N 44°47’52.4”E , 500 m , 3 June 2022 , R. Piwowarczyk (KTC); Tbilisi , near Abanotubani district, Taboris-Mta, SW of the Tabor Monastery of the Transformation, rocky slopes, on Caucalis , 41°41’07.3”N , 44°48’45.2”E , 502 m , 3 June 2022 , R. Piwowarczyk (KTC); Tbilisi , trial to the Turtle (Kutsba) lake, 41°42’01.3”N , 44°45’39.0”E , [ca. 700 m ], 22 May 2022 , M. Kolesnik (phot., https://www.inaturalist.org/ observations/118283596); Tbilisi , trial from the Marabda Str. to the Turtle (Kutsba) lake, slopes, rocky places, pine groves, along the paths, on Medicago , Caucalis , 41°42’06.4”N , 44°45’50.1”E , 41°42’00.8”N , 44°45’37.0”E , 640–715 m , 4 June 2022 , R. Piwowarczyk (KTC); Tbilisi , Tbilisi Sea, slopes, 41°45’19.2”N , 44°51’24.7”E , [ca. 650 m ], 18 May 2021 , N. Botchorishvili (phot., https://www.inaturalist.org/observations/79260693); Tbilisi , Tbilisi Sea, NE part, steppe slopes of the Kvirikobis Mt., on Medicago , not exluded also Geranium and Alyssum , 41°45’08.3”N , 44°51’24.4”E , 575 m , 7 June 2022 , R. Piwowarczyk (KTC); Tbilisi , Tbilisi Sea, N part, steppe slopes above road, probably on Alyssum murale , 41°45’46.4”N , 44°50’31.9”E , 570 m , 7 June 2022 , R. Piwowarczyk (KTC); Tiflis , Teleti, 5 May 1925 , Kikodze & Kara-Murza (TGM672) [as O. purpurea ]; Tbilisi , Krtsanisi distr., Teleti range, road to the St. George of Khandzta monastery of Krtsanisi, a rocky slope of the paved road with a sparse growing Rhus , probably on Alyssum , 41°40’09.3”N , 44°48’49.7”E , 500 m , 9 June 2022 , R. Piwowarczyk (KTC); Tbilisi , Vake distr., Tbilisi Zoo, 41°42’45.7”N , 44°46’46.5”E , [ca. 415 m ], 27 May 2022 , N. Tiunov (phot., https://www.inaturalist.org/ observations/119021392); Tbilisi , Rike Park, 41°41’38.2”N , 44°48’36.9”E , [ca. 400 m ], 15 June 2021 , M. Svoboda (phot., https://www.inaturalist.org/observations/107137505); Tbilisi , Avlabari distr., left bank of the river Mtkvari, Rike Park, Rike Park—Music Theatre and Exhibition Hall, mowed lawns around modern buildings and paths, on Caucalis , but also on other hosts, such as Diplotaxis , Plantago , Veronica ?, Sherardia , the precise determination of this hosts was not possible due to the very dense vegetation of the lawn, 41°41’39”N , 44°48’36”E , 41°41’38.2”N , 44°48’36.9”E , 398 m , 3 June 2022 , R. Piwowarczyk (KTC); Tbilisi , Mtatsminda Mt., Mtatsminda Park, 41°41’38.1”N , 44°47’00.2”E , [ca. 745 m ], 20 May 2021 , N. Botchorishvili (phot., https://www.inaturalist.org/observations/79512607); Tiflis , near Arsenalis Mt., 15 May 1923 , Kara Murza (TGM6003) [as O. lanuginosa ]; Tbilisi , Queen Tamar Avenue [near King David Residences], 41°42’59.1”N , 44°47’09.1”E , 28 April 2023 , B. Vardeh (phot., https://www.inaturalist. org/observations/157523997).