Short Communication |
Corresponding author: Renata Piwowarczyk ( renata.piwowarczyk@ujk.edu.pl ) Academic editor: Daniel Nickrent
© 2022 Renata Piwowarczyk, Óscar Sánchez Pedraja, Alexander V. Fateryga, Sergey A. Svirin.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Piwowarczyk R, Sánchez Pedraja Ó, Fateryga AV, Svirin SA (2022) Orobanche ingens (Orobanchaceae) – a poorly known species of the Greater Caucasus: taxonomic problems, distribution, hosts and habitats. PhytoKeys 193: 55-66. https://doi.org/10.3897/phytokeys.193.79886
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Orobanche ingens is an endemic species from the Caucasus, especially the Greater Caucasus, parasitising on large Apiaceae (usually Heracleum). This species was misclassified over the years and little was known about its range and habitats. Here, we clarify the typification, as well as provide notes about the taxonomy of this species. Additionally, we presented distribution, habit and host range of O. ingens and morphological features that distinguish it from similar species.
Caucasus, distribution range, Heracleum, holoparasitic Orobanchaceae, host, Orobanche, typification clarification
This taxon was described by
Years later,
Moreover, the new variety (var. heraclei), described by
Orobanche alba f. ingens Beck in Repert. Spec. Nov. Regni Veg. 18: 38[470] (1922)
lectotype (
Lectotype of Orobanche ingens (LE01015386) (http://re.herbariumle.ru/01015386).
Russia: 1. “Flora Caucasi // 10544. Orobanche alba 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.
Additional non-type material of Orobanche ingens (LE01015385, LE01015387) (http://re.herbariumle.ru/01015385, http://re.herbariumle.ru/01015387).
1. “Flora Caucasi / 10544. / Orobanche alba 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.
Orobanche alsatica var. heraclei Tzvelev, Fl. Azerb. 7: 591 (1957), nom. inval. (
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).
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.
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 (
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 (
Phenology. Flowering (June) July–August, fruiting (July) August (September).
Note. At first, the species was mistakenly described as a form of O. alba (subsect. Glandulosae); next
Distinctive morphological characters and hosts of the studied species and its Caucasian relatives.
O. bartlingii | O. ingens | O. owerinii | O. laxissima | O. minor | |
---|---|---|---|---|---|
Inflorescence | cylindrical to ovate, ± dense, usually shorter than the rest of the stem | usually long, rarely short (small specimens), cylindrical, somewhat lax, usually many-flowered | usually short-cylindrical to ovate, ± lax, usually few-flowered | usually long, or less frequently short (small specimens), cylindrical, very lax, usually many-flowered | usually long, rarely short (small specimens), cylindrical, dense at the first of anthesis, becoming lax later, usually many-flowered (except small specimens) |
Floral bract | about two thirds of the length of the corolla | as long as the corolla tube, slightly shorter than the corolla | as long as the corolla or slightly shorter or longer than the corolla | as long as the corolla tube, slightly shorter than the corolla | as long as the corolla |
Calyx segments | bidentate, teeth short, usually subulate at tip | deeply bidentate, teeth frequently filiform | segments entire or less frequently bidentate, narrowly subulate | usually entire or less frequently bidentate, rarely with 4 teeth, teeth narrowly subulate to filiform | bidentate or less frequently entire, teeth subulate to narrowly subulate but not filiform |
Corolla, length | 12‒18 mm | (20–) 23–24 (–30) mm | 15–20 (‒30) mm | (20) 22–24 (–25) mm | 10–19 mm |
Corolla, form | tubular | campanulate | campanulate-infundibuliform | tubular-infudibuliform | tubular |
Corolla, dorsal line | evenly curved at the proximal and distal part, almost straight at the middle | almost straight, except evenly curved at the proximal part | almost straight, except evenly curved at the proximal part and, sometimes, slightly bent forward at the distal part | almost straight, except evenly curved at the proximal part and, sometimes, slightly bent forward at the distal part | evenly curved at the proximal part, almost straight at the middle and slightly bent forward at the distal part |
Corolla, colour | yellow, brownish-yellow, pinkish ± tinged with purple | yellow, rarely violet, dark pink | pale violet to purple, rarely cream | purple, dark or light pink, rarely dirty yellow, light brown, sometimes with ± dark purplish veins | whitish or yellowish-white, more or less coloured purple, with dark purplish veins, rarely entirely yellow |
Filaments | inserted 1–3 mm above the corolla base; with long hairs at the basal half and sparsely glandular pubescent under the anthers | inserted 1 (–2) mm above the corolla base; with long hairs at the basal half and sparsely glandular pubescent (almost glabrous) under the anther | inserted 3–4 mm above the corolla base; hairy at the basal half (or third) and sparsely glandular pubescent under the anthers | inserted (2–)3–5 mm above the corolla base; hairy at the base and sparsely glandular pubescent under the anthers | inserted 2–3 (–5) mm above the corolla base, glabrous or sparsely hairy at the base and sparsely glandular pubescent under the anthers |
Stigma | yellow | mainly yellow or orange, rarely reddish, violet, dark pink | reddish to deep purple | dark pink, purple, violet, rarely orange, pale to dark yellow | pinkish, ± deep purple, rarely yellow or whitish |
Host | Apiaceae (Seseli) | Apiaceae (Heracleum) | Fabaceae (Trifolium and Vicia) and rarely Asteraceae (Leontodon and Lactuca) | trees and shrubs, Betulaceae, Oleaceae, Fagaceae, Sapindaceae, Cornaceae, Lythraceae, Fabaceae, Anacardiaceae (e.g., Fraxinus, Fagus, Carpinus, Rhus) | Fabaceae, Asteraceae, Apiaceae, etc. (e.g., Trifolium, Medicago, Hypochaeris, Leontodon, Chondrilla, Daucus) |
Our thanks are due to the LE herbarium and other herbaria consulted. Images of the herbarium specimens of Orobanche ingens were obtained from the Virtual herbarium of the Komarov Botanical Institute RAS (http://re.herbariumle.ru). Our thanks are due to Irina V. Sokolova for help in clarifying some taxonomic uncertainties. Helpful suggestions to improve the first version of this paper by Gerald M. Schneeweiss, Gianniantonio Domina and an anonymous reviewer are highly appreciated. This work was supported by the Research Projects of the Jan Kochanowski University in Kielce no. SUPB. RN.21.244 for R. Piwowarczyk. The field research in the Caucasus was partially financed by the National Geographic Society grant no. GEFNE 192–16 for R. Piwowarczyk. The work of A.V. Fateryga was a part of the State research project No. 121032300023-7.