Orobanche owerinii Beck (Beck), 1922

Piwowarczyk, Renata, Pedraja, Óscar Sánchez, Khutsishvili, Manana & Kharazishvili, Davit, 2023, Holoparasitic Orobanchaceae in Georgia (Caucasus): taxonomic revision, diversity, distribution, habitats and host range, Phytotaxa 604 (1), pp. 1-103 : 63-65

12. Orobanche owerinii Beck (Beck) (1922: 39[471])

. Type:— GEORGIA, Caucasia: prope Tiflis [Tbilisi], Ker Ogli [Kojori], leg. Owerin (LE, lost original material ( Tzvelev 2015: 206)). Russia, Caucasia (neotype designated here):— 1*. “Prope Narzana = Kislovodsk / Mt. Bermamut [Gora Bol’shoy Bermamyt / Гора Большой Бермамыт?]... 4500 фу. [1350 m] ... 17 ... 1889”. – 2**. “Bestau [Mount Beshtau / горы Бештау, near Pyatigorsk / Пятигорск (Stavropol Krai)] / ... 2400 фу. [720 m] 3 ... 1889”. – 3. “I. Akinfiew, Herbarium caucasicum, 1882-1897”. – 4. “ Orob. owerini GB. [m. Beck] / Dr. G. de Beck (* Beck 1930: 141, sub O. owerinii )”. – 5. “ Orob. crenata / f. owerini GB. [m. Beck] / Dr. G. de Beck (** Beck 1930: 141, sub O. owerinii )” (LE s.n. [the neotype is the longest specimen in the centre of the sheet]). Homotypic synonyms:— O. crenata var. owerinii Beck (1890: 227) . Heterotypic synonym:— O. owerinii var. woronowii Beck (1922: 39 [471]). Misapplied names:— O. crenata auct., non O. crenata Forsskal (1775: 113) . Fig. 42 .

General distribution: —The Caucasus, and probably also reaches neighbouring areas of Turkey and Iran.

Distribution: —Most localities are in the northern part, in the Greater Caucasus, in the Mtskheta-Mtianeti, Kakheti, Samegrelo-Zemo Svaneti, Racha-Lechkhumi and Kvemo Svaneti provinces, South Ossetia, Abkhazia, rarely in the central and southern part, in the Imereti, and in the Samtskhe-Javakheti prov. ( Fig. 43 ).

Habitat: —Subalpine meadows (“pratis”), rocky, grassy slopes, pastures, roadsides, rarely forests (e.g., coniferous, birch, mixed), usually (600)1800‒2400(2600) m.

Hosts: — Parasitic on Fabaceae and, more rarely, Asteraceae . Trifolium genus seems to be the main host-plant in the studied area, especially T. canescens Willd. , and T. ambiguum M. Bieb. , and rarely on Leontodon hispidus L. ( Asteraceae ).

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

Conservation status: —Least Concern (LC). The species has a rather wide distribution in Georgia and populations are usually very numerous. EOO is 45,000 km 2 and AOO is ca. 3000 km 2. The main threats are especially overgrazing.

Taxonomic note: —According to ( Beck 1890: 227) the “varietas” was described from Georgia based on material gathered by Owerin which is preserved in LE herbarium (“Transkaukasia: prope Tiflis [Tbilisi, Georgia], Ker Ogli [Kojori fortress] (leg. Owerin), in herb. Petrop. [LE]”, but according to Tzvelev (2015: 206), this material is missing “(укаЗан в LE, но нами не найден) [listed in LE, but not found]”. Beck (1890: 228) mentioned more material from this region (“Transkaukasia”) that could correspond to this same taxon, but this is not explicitly indicated and everything is placed within O. crenata Forssk. Years later, Beck (1922: 39) raised his variety to the rank of species, giving a more elaborate description and indicating more detail about the gatherings that corresponded to the new species. We, after studying the description and the mentioned materials, thought that Beck mixed two species, the one currently discussed ( O. owerinii Beck ) and the one following ( O. laxissima Uhlich & Rätzel ). Since the original material has been lost, and in order to avoid disadvantageous nomenclatural changes entailed by the strict application of the rules ( Turland et al. 2018: Art. 14), we believe it more appropriate to choose a neotype that can maintain the traditional interpretation of the name for the plant of Caucasian subalpine meadows.

O. owerinii is a very polymorphic species that was often confused (or included) with O. crenata and O. laxissima and other Caucasian species ( Piwowarczyk et al. 2019: 77–78).Among the sheets mentioned by Beck as O. owerinii , but which actually correspond to O. laxissima , we can mention, for example C371125 and LE s.n. (Imeretia, leg. Brotherus 1877 ). Another controversial sheet, the LE01013897 (“Окрестности Тифлиса 1871–1874 Оверин [surroundings of Tiflis 1871–1874 Owerin]”), determined by Tzvelev 1988 as O. owerinii and without any note from Beck, also seems to correspond to O. laxissima .

Specimens examined: — GEORGIA. Abkhazia: Sukhumi, s.d., A.V. Pranishnikov (MW0723345) [need confirmation, young specimen]; Abkhazia, auf radices Leontodon , prope Jurjewskoje [Tzebelda area], May 1902, G. Woronow (L2817104); prov. Kutaisi, distr. Suchumi, in silvis ad pag. Jurjevka, 7 May 1902, Alexeenko (TGM53194) [as O. lutea by Melikischvili]; “Transcaucasia: Abchasia prope Smjewoskoje m. Majo (G. Woronow), Tzebelda [Tsebelda], m. Majo (G. Woronow)” [see LE Collections Guide (Collections of Ju.N. Woronow from Caucasus): “Transcaucasia: 1902 (Abkhazia)” https://www.mobot.org/MOBOT/Research/LEguide/collections/119/index.html ( Beck 1922: 39 [471], sub O. owerinii ); bei Smjewoskoje und Tzebelda, May, G. Woronow ( Beck 1930: 141) ; auf dem Berge Mamdzyscha [Khrebet Mamdzyshkhara, 43°19’08.2”N, 40°21’26.3”E], Juli, G. Woronow ( Beck 1930: 141, sub O. owerinii ); Imereti prov.: Kveda Kinchkhi W, 42°29’22.7”N, 42°33’14.7”E, 29 May 2023, M. Ackermann (phot., https://www.inaturalist.org/observations/164750314); Jalaurta SW, 29 May 2023, 42°14’53.6”N, 43°22’44.9”E, M. Ackermann (phot., https://www.inaturalist.org/observations/165287694); Usakhelo NW, 42°14’18.5”N, 43°22’06.8”E, 29 May 2023, M. Ackermann (phot., https://www.inaturalist.org/observations/165291991; between Usakhelo and Tsasri, 42°14’42.0”N, 43°21’29.0”E, 29 May 2023, P. Plänker (phot., https://www.inaturalist.org/ observations/164474767); between Rondishi and Bangveti, 42°27’18.1”N, 42°32’51.6”E, 2 June 2023, S. Czyżewski (phot. https://www.inaturalist.org/observations/165279183); Kakheti prov.: Tusheti, Omalo, pine forest, 1850 m, 4 Aug 1988, D. Mtzkhvetadze (TBI) [as O. caryophyllacea ]; Tusheti Nature Reserve, between Dochu and Omalo, 42°22’23.2”N, 45°33’45.7”E, 2000 m, 27 July 2017, Tapaculo99 [phot., https://www.inaturalist.org/ observations/7458998]; Didikhevi N, 42°26’29.5”N, 45°35’02.7”E, [ca. 1960 m], 28 June 2022, G. Natsvlishvili (phot., https://www.inaturalist.org/observations/124552119, https://www.inaturalist.org/observations/124552110, https://www.inaturalist.org/observations/124547712); W of Tetri Tsklebi, Gombori pass, steppe hills, on L. hispidus , 41°51’54’’N, 45°17’09.3’’E, 1590 m, 23 July 2015, R. Piwowarczyk (KTC); Mtskheta-Mtianeti prov.: Mtiuleti, Kvemo Mletis-mta, 11 August 1936, D. Grigorashvili (TBI); Mtiuleti, in vicin. pag. Kaischauri, ad marg. silvae, 18 June 1966, Melikischvili (TGM15816) [as O. lutea by Melikischvili]; near Gveleti, on Trifolium , 17 July 2015, R. Piwowarczyk (KTC); Khevsureti,Ardoti, in region subalp. prop. pag. Čimgha, 12 July 1970, D. Otschiauri (TGM56629) [as O. picridis by D. Otschiauri]; distr. Tianeti, in decliv. Chaschari, 10 July 1962, Melikischvili (TGM13579) [as O. gamosepala by Melikischvili]; Gudauri, roadsides, slopes, meadows, on T. canescens , 42°28’27’’N, 44°28’45’’E, 2160‒2200 m, 16 July 2015, R. Piwowarczyk (KTC); Kobi, meadows, pastures, on Trifolium , 42°33’17’’N, 44°29’51’’E, ca. 2000 m, 17 July 2015, R. Piwowarczyk (KTC); Garbani, roadside in the village, meadows, 42°36’38’’N, 44°35’37’’E, 1850 m, 17 July 2015, R. Piwowarczyk (KTC); Sioni, meadow in the village, roadside, 42°35’52’’N, 44°35’07’’E, 1940 m, 17 July 2015, R. Piwowarczyk (KTC); Pkhelshe, meadows, pastures, 42°35’45’’N, 44°33’56’’E, 1890 m, 18 July 2015, R. Piwowarczyk (KTC); Mtskheta-Mtianeti, Great Caucasus, village of Kazbegi, 1840 m, 44°39’56’’E, 42°39’17’’N, 17 July 2002, G. Schneeweiss et al. (WU0027648, WU0027649); prov. Chwei (Khevi, Chevi), Greater Caucasus (Kavkasioni, Ķavķasioni), eastern surroundings of the village Kazbegi (Qazbegi) from the building of the Botanical Research Station (1850 m) to the foot of the steep slopes, 44°40’E, 42°39’N, 1850‒1950 m, 20 July 1997, G. Schneeweiss et al. (WU0027674, WU0027675); Stemantsminda (Kazbegi) NE of Hotel Kazbegi, subalpine meadow on the slope, 42°39’39’’N, 44°39’10’’E, 1870 m, 18 July 2015, R. Piwowarczyk (KTC); Kazbegi, slope of the hill in the town, 18 June 2019, J. Marciniuk (herb. J. & P. Marciniuk); near Kazbegi (Stepancminda), July 2011, M. Podsiedlik [phot.]; Kazbegi, Truso Valley, pasture by the river Terek, on Leontodon ?, 42°35’5.2’’N, 44°27’36.2’’E, 16 June 2019, G. Łazarski (KTC); Terek, Truso Valley, 19 June 2019, J. Marciniuk (herb. J. & P. Marciniuk); Tusheti distr., near Shatili, Mutsoje-Chala [Mutso, Mutsos-Chala], 20 July 1985, Ju. Menickij (LE); Kazbegi distr., between Kabakhi and Dariali, 42.722431N, 44.624444E, 17 June 2019, S. Thomas [phot., https://www.inaturalist.org/people/sjthomas]; Georgia, province of Mtskheta-Mtianeti, Great Caucasus, road from Terek valley to the village Juta, on Trifolium sp. (Fabaceae) , 18 July 2002, G. Schneeweiss et al. (WU0027608, WU0027609, WU0027610, WU0027611, WU0032459); prov. Chewi (Khevi, Chevi), Greater Caucasus (Kavkasioni, Ķavķasioni), ascent from the valley of the Snos-tskali (Sonsçqali) [= Gugushauris-tskali (Gudušaurisçqali)] towards the village of Jut’a (Džuta), approx. 12 km SE from the village of Kazbegi (Qazbegi), 44°45’E, 42°35’N, 1900‒2100 m, 24 July 1997, G. Schneeweiss et al. (WU0027677); Mtskheta-Mtianeti, Great Caucasus: gorge of river Dariali close to the Russian border, 44°37’53’’E, 42°44’03’’N, 1300–1450 m, 20 July 2002, G. Schneeweiss et al. (WU0027618, WU0027631, WU0027632, WU0027633, WU0027634, WU0027635); Mtskheta-Mtianeti, Great Caucasus, valley of small river Tibais-tskali near Gveleti i.e. gorge of river Dariali, 44°37’28’’E, 42°42’29’’N, 1480–1600 m, 20 July 2002, G. Schneeweiss et al. (WU0027639); Mughure NW, 42°28’26.8”N, 44°31’38.8”E, [ca. 1880 m], 17 July 2022, M. Svoboda (phot., https://www.inaturalist. org/observations/128589171); Racha-Lechkhumi and Kvemo Svaneti prov.: ad fontes fl. Tskhenistskali, Zemo Lapuri [42.822256°, 42.710874°], exp. austr.-orient., 2300 m, 15 July 1978, K. Kimeridze & G. Arabuli (TGM20123, TGM60839) [as O. raddeana by K. Kimeridze & G. Arabuli]; Svanethi, in vallis Zeskho, in decl. septentr., 24 July 1978, K. Kimeridze & G. Arabuli (TGM60938, TGM60939, TGM20121, TGM201222) [as O. raddeana by K. Kimeridze & G. Arabuli]; Great Caucasus: from Shovi along the road to Mamisoni pass [Mamison Pass], 43°43’40’’E, 42°40’45’’N, 1860 m, 13 July 2002, G. Schneeweiss et al. (WU0027620, WU0027619, WU0027620, WU0027621, WU0027622); Racha-Lechkhumi, Great Caucasus, from Shovi along the road to Mamisoni pass, above treeline, 43°43’E, 42°39’N, 2280–2500 m, 13 July 2002, G. Schneeweiss et al. (WU0027616, WU0027617, WU0027623); Samegrelo-Zemo Svaneti prov.: slopes of the Natakhtash pass [ca. 42.830023°, 42.287050°], 8000’, border of the Svaneti and Mingreli [Samegrelo, Samargalo, Megrelia, Mingrelia], Kutaisskaja prov., 12 July 1911, A.B. Schelkovnikov (TGM53180) [as O. caesia by Melikischvili]; N of Cheliri (near Iskari), N of the tower, slopes, roadsides, pastures, on Trifolium, 1600 –1700 m, 23 July 2018, R. Piwowarczyk (KTC); subalpine grassland, rocky places near Shdugra (Mazeri) waterfall, 4–5 km N of Mazeri village, 43°07’46”N, 42°35’42”E, 2200 m, on Trifolium , 25 July 2018, R. Piwowarczyk (KTC); Ushguli NE to Mt Shkhara, subalpine meadows, slopes of Enguri river valley, on Trifolium , 42°56’53”N, 43°03’59”E, 2200‒2600 m, 24 July 2018, R. Piwowarczyk (KTC); Davberi NW, 42°56’11.5”N, 42°55’13.4”E, [ca. 1860 m], 16 June 2022, Marcelpolling (phot., https://www.inaturalist.org/observations/122028517); Mestia NE, 43°03’42.5”N, 42°45’58.8”E, [ca. 1740 m], 11 June 2022, J. Pietruszewski (phot., https://www.inaturalist. org/observations/121284978); Mestia, 43°03’34.4”N, 42°43’17.8”E, [ca 2170 m], 43°03’17.9”N, 42°43’19.4”E, [ca. 1900], 6 July 2019, Chew_on_my_cholla (phot., https://www.inaturalist.org/observations/120221328, https://www. inaturalist.org/observations/120221281); Adishi, 42°59’21.2”N, 42°56’23.5”E, [ca. 2200 m], 2 July 2019, Chew_on_ my_cholla (phot., https://www.inaturalist.org/observations/120156693); Samtskhe-Javakheti prov.: between Skhvilisi and Didi Pamaji, in direction to Vale, meadows, edges of the field, on Trifolium , 41°37’37.6”N, 42°54’24.1”E, 1090 m, 13 July 2015, R. Piwowarczyk (KTC); between Naokhrebi and Turkish border, meadow, on Trifolium , 41°35’41.6”N, 42°49’43.1”E, 1230 m, 13 July 2015, R. Piwowarczyk (KTC); Lesser Caucasus, Trialeti range, Tetrobi massif [Tetrobi-Chobareti], Akhalkalaki district, on the river bank, in a meadow, 15 July 1961, E.W. Wołkowa & L.I. Ivanina (LE); Bogdanovskij [Ninotsminda] distr., between Bogdanovka [Ninotsminda] and Jigrasheni, left slope 2‒3 km NW from Bogdanovka, 1800 m, 12 July 1960, N. Tzvelev & S. Cherepanov (LE); Samtskhe-Javakheti, western part of the moutain range of Trialeti: surroundings of the village Azavreti, 43°23’12’’E, 41°36’23’’N, 2170 m, 15 July 2002, G. Schneeweiss et al. (WU0027606); Samtskhe-Javakheti, western part of the mountain range Trialeti, surroundings of the village Azavreti, 43°25’07’’E, 41°35’12’’N, 1940 m, 15 July 2002, G. Schneeweiss et al. (WU0027612, WU0027613); Borjomi W, roadside and the edge of the thickets, in a grove, in the area of former post- Soviet buildings, perhaps an old sanatorium, on Trifolium , 41°49’58.3”N, 43°21’38.3”E, 880 m, 14 June 2022, R. Piwowarczyk (KTC); South Ossetia: Dzau distr., Ermani, left bank of Ermani-don, opposite Srednie [Middle] Ermani vil., 21 July 1937, 25 July 1937, I. Abramov (LE) [attached T. canescens ]; Ermani, left bank of Britatskoye gorge, birch forest above Upper Britaty vil., 18 August 1946, I. Abramov (LE); Dzau distr., Ermani, birch forest near vil. Middle Ermani, 4 September, 1937, I. Abramov (LE); near Zgubiri, 1700 m, 26 June 1957, Bovodilin (MW0723349); Ermani, on Trifolium canescens , right bank of Ermani-don [Ermanistskali River], S exposure, 195?, T. Moiseeva (MW0723350); coniferous forest above Zgubiri, 26 June 1955, Gatsuk et al. (MW0723352); Ermani, 1954, Vitollo (MW0723353); right bank of the river Kalasani-don, subalpine meadow, 1900 m, 5 July 1948, E. Gognina (MW0723354); Ermani, right bank of Kalasani-don, subalpine meadow, 1900 m, 5 July 1948, V. Surova (MW0723355, MW0723358, MW0723359); right bank of Ermani-don, 2000 m, 30 June 1948, V. Surova (MW0723356); Ermani, right bank Middle Ermani Don, east slope, subalpine meadow, 2300 m, 4 June 1947, E.E. Gognina (MW0723357); right bank of river Kalasani-don, 1900 m, 5 July 1958 (MW0723360); right bank of Ermani-don, birch forest, 2000 m, 20 July 1948, E. Gognina (MW0723361); right bank of river Kalisani-don, subalpine meadow, on clover, 1800 m, E. Gognina (MW0717003); Гну́хское ущелье [Gnukh gorge], Khankoyty, meadow above vil. Disigin, 1900 m, 1 Aug 1930, E. & N. Busch (TBI) [as O. alba ]; Kamennik between Lower Ermani and Khodzem, 2300 m, 22 August 1929, E. & N. Busch (TBI) [as O. alba ]; Beloti rayon, lower pass Vanuri-vtsyag, subalpine meadow, 2100 m, 16 July 1930, E. & N. Busch (TBI).

Figures

FIGURE 42. Orobanche owerinii. A–D. General habit. E–G. Flower, side view. H–J. Flower, front view. K, M, O. Representative habitats (Mtskheta-Mtianeti prov., slopes of the Greater Caucasus along Georgian Military Road, Gudauri–Kobi). L–N. Representative hosts (L, M. Trifolium canescens, N. T. ambiguum). Photos by R. Piwowarczyk.

FIGURE 43. Distribution of Orobanche owerinii in Georgia.