Orobanche gamosepala Reut.

4. Orobanche gamosepala Reut. View in CoL View at ENA in Candolle (1847: 15)

. Type:— RUSSIA ( lectotype designated by Piwowarczyk et al. 2019: 54): North Caucasus, Stavropol Krai, near Pyatigorsk, “pr. Karass [colony two miles away from Mount Beshtau] & in m. Maschuka [Mashuk / Maschuka mountain], majo, Fl. Caucasica, [leg. Hohenacker]” (P02968131! [two righthand specimens (herb. Dr. a Ķhlewein)]); isolectotypes: P02968131! [only one specimen on left side of the sheet (herb. Al de Bunge)], G, LE [s.n., today LE01013898, LE01013900, LE01013899, LE01015360], COI00056887 - herb. Ķhlewein! Homotypic synonyms:— Ceratocalyx reuteri Lange (1864: 52) . Heterotypic synonyms:— Orobanche phoenicea Fisch. ex Beck (1890: 154) , pro syn. ( Turland et al. 2018: Art. 36.1b); Necranthus orobanchoides Gilli (1968: 297) . Fig. 26 View FIGURE 26 .

General distribution: —the Caucasus, mainly in Ciscaucasia (European Russia), less abundant in Transcaucasia ( Georgia, Armenia, Azerbaijan and NE Turkey). Endemic to the Caucasus.

Distribution: —High mountain ranges of the Greater Caucasus mainly in the northern parts (Samegrelo-Zemo Svaneti and Racha-Lechkhumi and Kvemo Svaneti provinces, as well as Abkhazia and South Ossetia), and rarely in the central parts of the Lesser Caucasus (Samtskhe-Javakheti prov.). Needs confirmation in Adjara ( Fig. 27 View FIGURE 27 ).

Habitat: —Here, in subalpine meadows, the upper mountain belt, rocky, grassy slopes, screes, shrublands, forests, usually (1700)2000–2600(2900) m.

Hosts: —In Georgia, it parasitises Geranium sp. (Geraniaceae) , such as G. gracile Ledeb. (probably a new host for it ( Sánchez Pedraja et al. 2016 +)), G. gymnocaulon DC. , G. ibericum Cav. , G. psilostemon Ledeb. (probably a new host for it ( Sánchez Pedraja et al. 2016 +)), and G. sylvaticum L.

Phenology: —Flowering (end May) June–July (August), fruiting end July–August.

Conservation status: —Near Threatened (NT). Species is known to occur in fewer than 20 localities and subpopulations are fragmented and do not consist of numerous individuals. EOO is ca. 25,000 km 2 and AOO is less than 200 km 2. Limiting factors are restricted loss/degradation of habitats as a result of overgrazing, felling of trees, landslides.

Taxonomic notes: — O. gamosepala ( Orobanche sect. Gamosepalae Teryokhin (1993: 40)) is genetically very distinct, forming a clade with O. anatolica / O. colorata ( Orobanche subsect. Arcuatae Teryokhin (1993: 40) [ Orobanche sect. Orobanche ]), simultaneously this clade is one of the oldest lineages of Orobancheae ( Piwowarczyk et al. 2021a).

Beck placed the two species in his Orobanche trib. Galeatae Beck in Biblioth. Bot. 19: 134 [152] (1890), nom. invalid. ( Turland et al. 2018: Art. 37.6). Syn: ≡ Orobanche [unranked] Galeatae Beck in Halácsy & Braun, Nachtr. Fl. Nieder-Österr.: 125 (1882) ( Beck 1890: 152); ≡ Orobanche Grex Galeatae (Beck) Beck in Engl., Pflanzenr. 96: 222 (1930), nom. invalid. Based on genetic studies ( Piwowarczyk et al. 2021a), we think that the two species should be considered as a different subgenus ( Gamosepalae) within the genus Orobanche . This new subgenus would contain the two sections based on the section ( Gamosepalae) and subsection ( Arcuatae) published by Teryokhin and placed by him in two different sections ( Gamosepalae and Orobanche ). The first comprises a single species ( O. gamosepala ), while the second comprises two ( O. colorata and O. anatolica ) or one ( O. colorata ) if we consider them synonyms.

Orobanche subg. Gamosepalae (Teryokhin) Ó. Sánchez & Piwow. , comb. & stat. nov. ( Turland et al. 2018: Art. 5A.1 and 6.10).

Basionym: Orobanche sect. Gamosepalae Teryokhin in Opred. Zarazikhovykh Fl. SSSR (ed. L.Yu. Budantsev): 40 (1993).

Type Name: Orobanche gamosepala Reut. in DC., Prodr. 11: 15 (1847).

— Orobanche sect. Gamosepalae Teryokhin in Opred. Zarazikhovykh Fl. SSSR (ed. L.Yu. Budantsev): 40 (1993).

— Orobanche sect. Arcuatae (Teryokhin) Ó. Sánchez & Piwow. , comb. & stat. nov.

Basionym: Orobanche subsect. Arcuatae Teryokhin in Opred. Zarazikhovykh Fl. SSSR (ed. L.Yu. Budantsev): 40 (1993).

Type Name: Orobanche colorata K. Koch. View in CoL in Linnaea 17: 291 (1843).

Specimens examined: — GEORGIA. Abkhazia: Abchasien: in Alpenwiesen an den Quellen des Flusses Seken [Sakeni River (Kodori Range)] ( Alboff!)” ( Beck 1930: 226, sub O. gamosepala ); Azikhba Mt, 17 July 1981, Sh. Kvaratskhelia (BATU); Bzipi [Bzyp, Gagra district] range, Kosiro Mt., S slopes, 2000 m, 9 August 1930, R.I. Tetlev [Potlev?] (LE); Abkhazia, Ritsa Strict Nature Reserve, mountain Agura, Kamennaya Polyana tract, Ritsa relic national park, ca. 2400 m, subalpine meadow, 43°29′48.4”N, 40°44′38.0”E, 14 July 2017, M. Skotnikova [phot. https://www. plantarium.ru/page/image/id/553690.html]; Adjara: cited by Gagnidze (1999), need confirmation; according to Dmitryeva (1990) O. gamosepala is rare in Adjara [no herbarium sheets kept from Adjara at TBI, BATU, etc.]; Racha-Lechkhumi and Kvemo Svaneti prov.: ad font fl. Tskhenistskali, Laphuri, ac dextea vallis, in prat. subalp., 2150 m, 19 July 1978, K. Kimeridze & G. Arabuli (TGM60937); Shovi, alpine grasslands and screes near the tourist path Glola-Lake Udziro, 4 km S from the village, 42.6669000°N, 43.6569778°E, ca. 2200 m, 6 July 2017, K. Chytrý & V. Kalníková (BRNU); near the forest path, the trail to Udziro lake, 1.5–2 km S of Shovi village, 42°41’31”N, 43°40’04”E, 1950 m, 18 July 2018, R. Piwowarczyk (KTC); Samegrelo-Zemo Svaneti prov.: bei Dadiasch [Svaneti Range] und Latpar [Latpari Pass], 2100 m, ( Akinfiew!) ( Beck 1930: 226, sub O. gamosepala ); Latpari [Latpari Pass], Svanetie, 13 July 2011, M. Kesl [phot. biolib.cz/en/taxonimage/id201833/?taxonid=784321]; Svaneti, in pratis alp. ad font. fl. Dewlura, loc. Schawlura, in vallis Čuberi [Chubersky Gorge], 2500 m, 10 August 1977, K. Kimeridze (TGM5951); Adishi N, 43°01’15.9”N, 42°54’58.8”E, [ca. 2690 m], 2 July 2019, Chew_on_my_cholla (phot., https:// www.inaturalist.org/observations/120156045); Mestia distr., SE of Tsaldashi, 43°1’29.513”N, 42°53’32.678”E, 43°1’27.880”N, 42°53’26.950”E, 2535 m, 18 Augst 2022, P. Novák, V. Sedláček, D. Szokala & G. Štětková (phot.); Localities difficult to determine: Samegrelo-Zemo Svaneti and Racha-Lechkhumi and Kvemo Svaneti provinces: in Swanetia [Svaneti] ( Radde) ( Beck 1890: 155, sub O. gamosepala ); Mingrelien [Samegrelo/Mingrelia]: an den Quellen des Flusses Ingur[Enguri River], 2900 m, ( Radde!); Swanetien[Svaneti]( Radde!) ( Beck 1930:226, sub O. gamosepala ); Samtskhe-Javakheti prov.: Borjomi-Bakuriani-Akhalkalaki Highway, 2 km south from Bakuriani, 41°44’08.0”N, 43°30’33.0”E, [ 1750 m], 6 June 2005, C. Davidson [phot., floraoftheworld.org/taxon/flora/17592186258585]; by the road from Bakuriani to the Tskhratskaro pass [ 2454 m], edge of a deciduous forest, an overgrown subalpine meadow, 41°43’52.2”N, 43°30’10.7”E, 1780–1790 m, 13 June 2022, R. Piwowarczyk (KTC); South Ossetia: subalpine meadow, on the eastern slope of the Bibility range [the Malaya Liakhva River basin], 2300 m, 11 July 1930, E. & N. Busch (LE); Ermani, in birch forest, near Dzoar-Shad Lake, on Geranium sylvaticum , 2 July 1937, I. Abramov (LE); birch forest, left slope of upper Ermani gorge, 2500 m, 4 July 1940, det. N. Tzvelev, (LE); left bank of the Upper Ermani, 17 July 1940,?, det. Tzvelev (LE); Бриттатское ущелье [Britat gorge], left slope, Haranasar урочище [Haranasar tract], 2 August 1929, E. & N. Busch (LE); Ermani, in a birch forest, left bank of the Middle Ermani gorge, 2200 m, 28 July 1939, E.E. Rut (LE); Шуацхури [Shuatskhuri; Bass. Maloi Liakhvy], hay mowing subalpine meadow, Nekh Mt., 2200 m, 9 July 1930, E. & N. Busch (LE); Ermani area, right bank of the Sredne-Erman gorge, subalpine meadow, on roots of Geranium ibericum , 12 July 1939, V.D. Mozgova (W1941-0003773; Ex Herb. Inst. Bot. Ac. Sc. URSS).