Orobanche ingens (Beck) Tzvelev in Novosti Sist. Vyssh. Rast. 27: 182 (1990)

Piwowarczyk, Renata, Pedraja, Oscar Sanchez, Fateryga, Alexander V. & Svirin, Sergey A., 2022, Orobanche ingens (Orobanchaceae) - a poorly known species of the Greater Caucasus: taxonomic problems, distribution, hosts and habitats, PhytoKeys 193, pp. 55-66 : 55

publication ID	https://dx.doi.org/10.3897/phytokeys.193.79886
persistent identifier	https://treatment.plazi.org/id/B482DFB6-B2D0-51BC-B35B-5ED9D326D53D
treatment provided by	PhytoKeys by Pensoft
scientific name	Orobanche ingens (Beck) Tzvelev in Novosti Sist. Vyssh. Rast. 27: 182 (1990)
status

Treatment

Figs 1 View Figure 1 , 2 View Figure 2

Orobanche alba f. ingens Beck in Repert. Spec. Nov. Regni Veg. 18: 38[470] (1922)

Type.

lectotype ( Tzvelev 1990: 182; Turland et al. 2018: Art. 9.3, 7.11; we do not know if additional elements were used by Beck ): Russia: 1. "Flora Caucasi / 10524 [m. Beck]. / Heracleum parasitica [m. Beck] / Prov. Daghestan, Distr. Kasi-Kumukh [Kazi-Kumykh, Laksky distr.]. / Ad viam inter pagos Chelussun et Kumukh / 13 Jul 1897 6400 ´-7000´ / leg. Th. Alexeenko ". - 2. "Orobanche alba Steph. nov. forma ingens mihi / Luxurians … [description] … / Cum forma bidentata Beck Mon. 211 formis / intermediis conjunta / Beck [m. Beck]". - 3. "Orobanche crenata Forssk.! / var. ingens (Beck) m.c.n. / 15.X.1947 / Determ. I. Novopokrovsky ". - 4. "? O. crenata Forssk. / … / 1955.II.15 Teste: Tzvelev". - 5. " Orobanche ingens (G. Beck) Tzvel / comb. et stat. nov. / Typus! / 1988.III.5 Teste Tzvelev" (LE01015386, Fig. 1 View Figure 1 ) .

Additional non-type material.

Russia: 1. "Flora Caucasi // 10544. Orobanche alba Steph. / f. bidentata Beck [m. Beck] // Prov. Daghestan, Distr. Kasi-Kumukh [Kazi-Kumykh, Laksky distr.]. / Ad viam inter pagos Chelussun et Kumukh / 13 Jul 1897 6400 ´-7000´ / leg. Th. Alexeenko ". - 2. "Af. O. owerini G. Beck [m. Beck, handwritten on the sheet]". - 3. "I. Novopokrovsky. Notae criticae / Orobanche crenata / Forssk.! / 1948.8.XI". - 4. " Orobanche ingens (G. Beck) Tzvel / Isotypus! / 1988.III Teste Tzvelev" (LE01015385, Fig. 2 View Figure 2 ).

1. "Flora Caucasi / 10544. / Orobanche alba Steph. / f. bidentata Beck / Prov. Daghestan, Distr. Kasi-Kumukh [Kazi-Kumykh, Laksky distr.]. / Ad viam inter pagos Chelussum [sic] et Kumukh / 6400 ´-7000´ / 13 Jul 1897 / leg. Th. Alexeenko [m. Beck]". - 2. "an. O. picridis - hieraciodes / Holandre [handwritten on the sheet]". - 3. " Orobanche ingens (G. Beck) Tzvel / Isotypus! / O. crenata Forsk. / 1955.II.2 [date of the first revision] Teste Tzvelev III.1988" (LE01015387, Fig. 2 View Figure 2 ).

Synonyms.

Orobanche alsatica var. heraclei Tzvelev, Fl. Azerb. 7: 591 (1957), nom. inval. ( Turland et al. 2018: Art. 39.1); Orobanche alsatica var. heraclei Tzvelev in Schischk., Fl. SSSR 23: 111 (1958), nom. inval. ( Tzvelev 2015: 211, nom. nud. [sic]).

Misapplied names.

Orobanche alba var. bidentata sensu Beck in Engl., Pflanzenr. 96: 155 (1930) [saltem p.p.], non O. alba f. bidentata Beck in Biblioth. Bot. 19: 211 (1890).

Distribution, ecology and taxonomic problems

General distribution. Caucasus, mainly Greater Caucasus range, Russia (Krasnodar Krai, Karachay-Cherkessia, North Ossetia, Dagestan) and Georgia. Needs confirmation in Azerbaijan. Endemic to the Caucasus (Fig. 3 View Figure 3 ).

Specimens examined. Georgia. Racha-Lechkhumi and Kvemo Svaneti prov., tall herb communities, forest edges and glade, near the trail to Udziro lake, 1.5-2 km S of Shovi Village, 42°41'07"N, 43°39'56"E, 2050-2100 m a.s.l., on Heracleum leskovii , but probably also on Ligusticum alatum , 18 July 2018, R. Piwowarczyk (KTC); Russia: Dagestan. Distr. Kasi-Kumukh [Laksky distr.], ad viam inter pagos Chelussun et Kumukh, 6400 ´-7000´, 13 July 1897, Th. Alexeenko (LE) [as O. ingens by Tzvelev in 1988, Typus]; prov. Dagestan, distr. Samur, ad riparian fl. Dulty-chaj, 7300', 12 July 1897, Th. Alexeenko (LE) [as “10526” O. alba f. rubiginosa by Beck ( Beck 1930: 148, sub O. alba ), O. alsatica by Novopokrovsky in 1949, O. flava by Tzvelev in 1955, O. ingens by Tzvelev in 1988]; Dagestan, Dokuzparinskiy distr., Kiler, meadow near road, parasitising on Heracleum sosnowskyi [root attachment verified], 41°22'50"N, 47°53'55"E, [ca. 1130 m a.s.l.], 26 June 2021, A.V. Fateryga & S.A. Svirin (KTC, YALT); Karachay-Cherkessia. North Caucasus, upper reaches of the Bolshaya Laba river, upper part of Zagedanka river, tall grassy vegetation in subalpine meadow, on the roots of Heracleum , 2300 m a.s.l., 19 July 1946, Blyumental & Karpova (LE) [as O. owerinii by Novopokrovsky in 1946, O. alsatica Kirschl. var. heraclei Tzvel. by Tzvelev in 1955, O. ingens (G. Beck) Tzvel. by Tzvelev in 1988]; North Caucasus, basin of the Urup river [Kuban distr.], 9 Aug 1945, V.I. Grubov & L.I. Ivanina (LE) [as O. ingens by Tzvelev in 1988]; near Dombay, Alibek river valley, 29 July 2014, M. Skotnikova [phot., https://www.plantarium.ru/page/image/id/268327.html, https://www.plantarium.ru/page/image/id/256514.html, https://www.plantarium.ru/page/image/id/256512.html, as O. alsatica ]; Dombay, Alibek river valley, 26 August 2013, S. Banketov [phot., https://www.plantarium.ru/page/image/id/205049.html, https://www.plantarium.ru/page/image/id/205050.html, https://www.plantarium.ru/page/image/id/205051.html, https://www.plantarium.ru/page/image/id/205052.html, as O. alsatica ]; Karachay-Cherkessia, Dombay, Dombay-Ulgen river valley, above the Russian glade, about 2000 m a.s.l., 21 July 2010, E. Komarov [phot., https://www.plantarium.ru/page/image/id/66557.html, https://www.plantarium.ru/page/image/id/66558.html]; Karachayevsky distr., near Dombay, 11 Aug 2010, E. Suslova [phot., https://www.inaturalist.org/observations/70214391, as O. alsatica ]; Karachay-Cherkessia, Malokarachaevsky distr., slope of Bolshoi Bermamyt peak, ca. 2000 m a.s.l., subalpine meadow, 5 Aug 2017, T. Gaidash [phot., https://www.plantarium.ru/page/image/id/527245.html, https://www.plantarium.ru/page/image/id/527246.html, as O. grossheimii ]; Karachay-Cherkessia, Malokarachaevsky distr., Eshkakon river valley, 15 June 2013, I. Tabunova [phot., https://www.plantarium.ru/page/image/id/193231.html]; Krasnodar Krai. North Caucasus, Krasnodar territory, Mostovskoi distr., 6-7 km N from Shedok, vicinity of Dyatlovo, small piece of fallow land at the field margin, not far from dried-up stream, on Heracleum mantegazzianum , 4 June 2002, D.V. Geltman (LE) [as O. alsatica ]; North Ossetia. Vladikavkaz, July 1888, Akinfiew (LE) [as O. flava ? by Tzvelev in 1955, as O. ingens by Tzvelev in 1988]; North Ossetia, Irafsky distr., upper part of Urukh river, ca. 1700 m a.s.l., subalpine meadow, 9 July 2016, M. Skotnikova [phot., https://www.plantarium.ru/page/image/id/479013.html, as O. alsatica ].

Habitat. In tall herbaceous habitats, mainly in subalpine meadows, pastures, edges of forests and shrubs, forest glades, near river valleys, as well as fallow lands, usually (1000) 2000-2300 (2500) m a.s.l.

In Georgia, in the locality on a subalpine glade, numerous pollinators of this species - workers of Dolichovespula sylvestris (Scop.) ( Hymenoptera , Vespidae ) - have been observed.

Host. Parasitises Heracleum species ( Apiaceae ), such as H. leskovii Grossh., H. sosnowskyi Manden. and H. mantegazzianum Sommier & Levier. Probably also on other large Apiaceae species, such as Ligusticum alatum (M. Bieb.) Spreng. (= Cnidiocarpa alata (M. Bieb.) Pimenov & Kljuykov), but this needs confirmation (observed by R. Piwowarczyk in Georgia, but root attachment was not verified).

Apart from O. ingens , only one Heracleum parasite is known - Phelipanche sevanensis Piwow., Ó. Sánchez & Moreno Mor., described from the slopes of Sevan lake in Armenia, where it parasitises H. trachyloma Fischer & Meyer ( Piwowarczyk et al. 2017, 2019).

Phenology. Flowering (June) July-August, fruiting (July) August (September).

Note. At first, the species was mistakenly described as a form of O. alba (subsect. Orobanche Glandulosae ); next Tzvelev (1990, 2015) continued to indicate the relationship of O. ingens with O. alsatica aggr., especially with O. bartlingii Griseb. (subsect. Orobanche Curvatae ). However, according to morphological features (Figs 1 View Figure 1 , 2 View Figure 2 , 4 View Figure 4 , Table 1 View Table 1 ), it is far from the O. alsatica group and subsect. Orobanche Curvatae , but belongs to the Minores / Speciosae subsection. In the Minores / Speciosae subsection, the corolla is usually tubular or campanulate, almost straight in the middle part, the pubescence of the corolla is usually a mixture of longer and stiff whitish hairs, the calyx segments are long, entire or bidentate, usually subulate or filiform at the tip. In contrast, in the subsect. Orobanche Curvatae , the corolla is usually tubular-infundibuliform or ± broadly tubular, the dorsal line of the corolla is evenly curved over its entire length, the pubescence is rarely a mixture of whitish hairs. Molecular studies do not indicate the validity of dividing the species into the subsect. Orobanche Speciosae because the species included here are both morphologically and genetically very similar to those of the subsect. Orobanche Minores ( Piwowarczyk et al. 2021). It is worth emphasising that individuals of O. ingens are usually very tall, over 60 to 100 cm, with corolla (20-) 23-24 (-30) cm long, usually with a mixture of longer and stiff whitish hairs (Fig. 4F View Figure 4 ; without dark glandular hairs which are characteristic for the subsect. Orobanche Glandulosae ) and both the corolla and stem can be very variable in colour, even within one population, from yellow to pink and purple; additionally, the colour of the stigma can be variable, from yellow through to orange, pink to purple (Fig. 4 View Figure 4 and photos from Plantarium.ru and iNaturalist.org, links above).