Phelipanche pulchella (C.A. Mey.) Soják (1972: 129)

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 : 20-23

3. Phelipanche pulchella (C.A. Mey.) Soják (1972: 129)

. Type:— GEORGIA (lectotype designated by Piwowarczyk et al. 2017d: 122): “Georgia. Imeritia [Imeretiya / Imereti], Eichwald s.n. [herb. Ledebour] (LE!)” [http://re.herbariumle. ru/01015364 – LE1015364 (today numbered)]. Basionym:— Phelypaea pulchella C.A. Mey. in Eichwald (1831: 17). Heterotypic synonyms:— Orobanche bungeana Beck (1890: 119) ; Orobanche caucasica Beck (1922: 34 (466)[– 35(–467)]); Phelipanche bungeana (Beck) Soják (1972: 129) ; Orobanche septemloba (Beck) Tzvelev (1957: 568) ; Phelipanche septemloba (Beck) Soják (1972: 130) ; Orobanche oxyloba var. septemloba (Beck) Beck (1930: 90) ; Orobanche oxyloba f. septemloba Beck (1922: 34 [466]). Misapplied names:— Orobanche coelestis auct. (saltem p.p.), non O. coelestis (Reut.) Beck (1890: 114) ; Phelypaea lanuginosa sensu Boissier (1879: 496) et auct. div. (saltem p.p.), non P. lanuginos a C.A. Mey. in Ledebour (1830: 46). Fig. 10 .

General distribution: —The Caucasus and surrounding areas ( Iran and Turkey).

Distribution: —Scattered localities mainly in the western, central and northern parts of Georgia (Adjara, Imereti, Samtskhe-Javakheti, Mtskheta-Mtianeti, and Samegrelo-Zemo Svaneti provinces (see also: Ledebour 1849: 311; Schneeweiss et al. 2004; Carlón et al. 2005: 67) ( Fig. 11 ).

Habitat: —Rocky slopes, subalpine meadows, rocks, shrublands, forests, old cemeteries, 450–2000(2500) m.

Hosts: —Apparently only parasitic on Lamiaceae , such as Teucrium chamaedrys L. (e.g., in its terra class.); needs confirmation for Thymus .

Phenology: —Flowering (April) May to June (July), fruiting (June) July to August.

Conservation status: —Near Threatened (NT). The species has just over 10 localities, usually not numerous and with high fragmentation. EOO is ca. 22,000 km 2, and AOO is ca. 100 km 2. In some localities, there is a threat of overgrowth, overgrazing or habitat destruction and conversion, as well as landslides.

Taxonomic notes: — Phelipanche pulchella , P. bungeana , P. caucasica and P. septemloba are four poorly known taxa, which are frequently confused with each other and with close species ( Boissier 1879, Novopokrovsky in Grossheim (1949), Novopokrovsky & Tzvelev 1958, Gilli 1966, Teryokhin et al. 1993 [ Phelipanche ser. Cilicicae Teryokhin ], Tzvelev 2015, Piwowarczyk et al. 2017 d, Gagnidze 1999). After seeing the types and studying many fresh (anthesis) and dry materials (post-anthesis) of their classic locations (in natura), we consider them to be synonyms. We now think that the origin of the confusion is that these species were described on the basis of a single specimen and/or poor gathering and their nomenclatural types are not the most typical or representative element of these species ( Turland et al. 2018: Art. 7.2).

Of all these names, the priority name is P. pulchella , this name was typified for us on material of the LE herbarium ( Piwowarczyk et al. 2017d: 122 [at that time not yet numbered]). In the same herbarium, we can observe the isosyntype (perhaps an isolectotype ( Sánchez Pedraja et al. 2016 +)) today with the number LE01015365 (http://re.herbariumle. ru/01015365 [a single specimen divided into two parts (inflorescence and stem)]).

P. bungeana was also typified by us on a single specimen from the LE herbarium ( Beck 1890: 119, “Vidi specimen unicum in herb. Petrop.”, Piwowarczyk et al. 2017d: 122). Now, after seeing more living material of the species, we think that the differences indicated in the key ( Piwowarczyk et al. 2019: 47–48) fall within the variability that this species shows; therefore, both names are synonyms.

Orobanche caucasica was described by Beck (1922) from the Caucasus and surrounding areas. This species has frequently been confused or synonymised with P. pulchella and P. bungeana as we have already indicated previously. Thus, in Novopokrovsky & Tzvelev (1958:48) it is considered synonymous with O. bungeana . However, it is considered synonymous with O. pulchella by Novopokrovsky in Grossheim (1949 [see revision label]) ( Novopokrovsky & Tzvelev 1958: 49) or in Tzvelev (2015: 200). We also believe that it was never combined correctly in the genus Phelipanche ( P. caucasica (Beck) Uhlich ( Hassler 2022 +) and so is not effectively published ( Turland et al. 2018: Art. 29 and 30).

The single specimen that we choose here as a lectotype was determined by Beck himself and presents flowers with calyx-teeth longer than the tube and corolla ± tubular with acute lobes; these morphological characters of the specimen indicated here would incite future confusion ( Gilli 1982: 12). However, the specimen we choose here as the lectotype represents a poor and not very typical form of the P. pulchella to which we think O. caucasica is synonym.

Lectotype (here designated): 1. “ Orobanche caucasica Beck … / vide Beck 1930 … / БорЖоми, на горе на юг 5000’ / 28 июня 1888 / КавкаЗский гербарий, И. R. Акинфиев [Borjomi, on the mountain to the South 5000’, 28 June 1888, Caucasian herbarium, I. Ya. Akinfiev]”. – 2. “I. Akinfiew Herbarium Caucasicum / 1882 — 1897”. – 3. “ Orob. (Trionychon) caucasica n. sp. / Dr. G. de Beck [m. Beck]”. – 4. “I. Novopokrovsky. Notae criticae. / Orobanche pulchella (C.A. Mey.) Novopokr. / (syn.: O. caucasica G. Beck ) / 1947.30.XII”. – 5. “ Orobanche caucasica G. Beck / (= O. Bungeana G. Beck ) / Typus! / 1956.IX.28 Teste: Tzvelev”. – 5. “Typus!”. – 6. “10.XI.82 Seed examples…” (LE01014672 [http://re.herbariumle.ru/01014672]). Note see P. pulchella for review labels ( Novopokrovsky & Tzvelev 1958: 49).

Beck (1922) describes another new taxon from Georgia from the same place or proximity ( Beck 1930: 101). This new taxon is the O. oxyloba f. septemloba .

We have been able to study the types in the LE herbarium and, in addition, one of us (R. Piwowarczyk) visited the classical location and collected material similar to the types. After studying all the materials at our disposal and performing the typification of both taxa, we believe that they certainly correspond to the same species.

When Beck (1922) describes this taxon as a form (“f.”) of O. oxyloba , we believe that he does so on the basis of the five specimens (two of them joined by the base) contained in a sheet of LE—I. Akinfiew Herbarium Caucasicum 1882–1897 (s.n., Piwowarczyk!), This single sheet, in addition, contains other specimens up to a total of nine, which correspond to different species (eight to the genus Phelipanche and one to Orobanche ) and gatherings (four); apparently, Akinfiew completed all of them in the vicinity of Borjomi during the years 1887 and 1888. The sheet has recently been numbered with four numbers, one for each gathering (LE01015367 – LE01015368 – LE01015369 – LE01015370). This sheet, in addition to the original Akinfiew labels, contains another manuscript by Beck. In this label, he gives a diagnosis of its new form, and this diagnosis would be repeated with a lot of fidelity by the same Beck (1922) when he published the new form and by Beck (1930) when he raised it to the rank of “var.”. The Beck label is placed near to one of “ 26 June 1887 [26 июня 1887 г.]”; this last contains four specimens, the first one on the left corresponds to an Orobanche sp. and three others (on the right side and two of them joined at the base [“scapus subimplex”]) correspond, undoubtedly to this species, with simple stem ( Novopokrovsky & Tzvelev 1958: 58, “up to 25 cm tall”), scale leaves ± broadly oval-lanceolate and acuminate at the apex, inflorescence, ± long, dense and polyanthous and flowers with corolla tubular to tubular-infundibuliform, ± widened at the mouth ( Novopokrovsky & Tzvelev 1958: 51, t. 4 f. 4), dorsal line straight, but ± curved after anthesis, upper lip slightly retracted with respect to the lower lip, lobes ± acute and calyx teeth lanceolate-subulate longer than tube. These three specimens belong to the same single gathering (LE01015370) and are designated here as the lectotype of the name. The other two specimens of the gathering of “ 4 June 1888 ” are considered as isosyntypes (LE01015368). The rest of the gatherings, “ 3 July 1887 ” (LE01015367) and “ 9 June 1888 ” (LE01015369), apparently, do not contain specimens corresponding to this species. Tzvelev, on his review label of January 10, 1955, marked with an “α” the label of the specimens that considered to be the “α-typus!” (LE01015368) of the species, but we believe that both the Beck label and the description of it are better suited to the indicated specimens (LE01015370).

Lectotype (here designated): 1. “ Имеретия [Imereti], Даба [Daba, village near Borjomi, in Samtskhe-Javakheti prov.], на высоте 2000 футов [at 2000 feet] / 26 June 1887 [26 июня 1887 г.]”. – 2. “ O. oxyloba / f. O. septemloba scapus subsimplex / flores 16–21 mm longi … det. Dr. G. Beck”. – 3. “+ O. barbata Poir. / minor Sutt. 1955 .I.10 Teste Tzvelev” (LE01015370 [http://re.herbariumle.ru/01015370], lectotype are the three specimens on the right side, two of them joined at the base [“scapus subimplex”], the remaining specimen, the first one on the left, corresponds to an Orobanche sp. , cf. O. lutea ).

Syntypes: 1. “α [m. Tzvelev] / БорЖоми, Тбилиси провинция, воЗле кладбища / 4 июня 1888 [Borjomi, Tbilisi province, near the cemetery 4 June 1888]”. – 2. “ Orobanche septemloba (G. Beck) Tzvel. / α-Typus! / 1955.I.10 Teste : Tzvelev ”. – 3. “α.- Typus !” (LE01015368, the syntypes are two specimens close their barcode); 1. “БордЖоми [ Borjomi , Likani] 3000’ [28 June 1888] I. Akinfiew Herbarium ”. 2. “ Orobanche oxyloba / (Reut.) G. Beck var. O. septem - / loba G. Beck / 18.IX,1947 / Det. I. Novopokrovsky ”. – 3. “ Orobanche septemloba (Beck) Tzvel. / 1955.I.10 Teste: Tzvelev” (LE, s.n. [sheet with a single specimen]) .

Other gatherings of this LE sheet: 1. “ЗакавкаЗья. / БорЖоми на высоте 3000 футов, 3 июля 1887 [Transcaucasia. Borjomi at an elevation of 3000 ft, 3 July 1887]” (LE01015367 [http://re.herbariumle.ru/01015367], this unique, small and non-representative specimen could correspond to the same species). 1. “ЗакавкаЗья. Borjom 2500 f. [locality and elevation in black ink with Latin alphabet, m. ign.] / БорЖоми Тбилиси провинция 2500 футов, 19 июня 1888 [Transcaucasia. Borjomi Tbilisi province 2500 ft, 19 June 1888]” (LE01015369, two thin and small specimens that could correspond to P. nana . But, Beck (1922), could have used this label [the black ink text], for understanding it, to indicate the locality and elevation of his new form ( Beck 1930: 90)).

Beck considered his new “f.” included in O. oxyloba and close to O. coelestis f. persica Beck (1890) [ Phelipanche persica (Beck) Piwow., Ó. Sánchez & Moreno Mor. (2017) ] and to O. orientalis Beck (1890) [ Phelipanche libanotica (Schweinf. ex Boiss.) Soják (1972) ], probably by the morphological characters indicated above (e.g., calyx-teeth longer than tube), ± common to the three taxa. However, as indicated before, the specimens we choose here as the lectotype represents a ± dry (post-anthesis) and not very typical form of the species to which we think it is synonym.

Specimens examined: — GEORGIA. Adjara prov.: Transkaukasien: in Lasistan, Provinz Batum, distr. Artwin im Tale des Flusses Ardanuczsu [Turkey], im Mai 1907 (Woronow!) auf Teucrium chamaedrys ( Beck 1930: 98, as O. bungeana ); distr. Shuakhevi, Quercus and Pine forest with Cistus and Teucrium , on Teucrium , 450 m, 22 April 1960, A. Dmytrieva (BATU) [as O. pulchella ); distr. Shuakhevi, Ajara, vil. Chanchkhalo, Quercus forest with Cistus and Teucrium , 29 April 2015, N. Memiadze (BATU) [as O. pulchella ]; Ajara-Shavsheti range, Ajara, mount. Tbeti, subalpine rocks, 2400 m, 2 Aug 2006, J. Jakeli & N. Memiadze (BATU) [as O. pulchella ]; Imereti prov.: Georgia. Imeritia [Imeretiya / Imereti], Eichwald s.n. (LE1015364 - hb. Ledebour ( Eichwald 1831: 17, as P. pulchella ; Reuter 1847: 5, as P. pulchella ; Boissier 1879: 497, sub, as P. pulchella ; Piwowarczyk et al. 2017: 122, lectotype )); In Imeritia [Imeretiya / Imereti], legit cel: Dr.: Eichwald, Acc. 1827 (LE01015365, isosyntype); Imeritia [Imeretiya / Imereti], Szovits [1827–1830, m. C.A. Meyer] (P02968079 [as Phelipaea coelestis Reut. ] ( Beck 1890: 115, O. aegyptiaca var. typica f. pulchella )); Mtskheta-Mtianeti prov.: Khevsureti, a sinistr. angust. Arghuni, in decl. siccis circa p. Schatili (loc. Vake), 18 July 1976, D. Otschiauri (TGM60001); Gudamakari, circa pag. Magartha (Makarta], in pratis, 4 July 1973, D. Otschiauri (TGM68198, TGM68199); Georgia, Great Caucasus, valley of river Terek (Tergi), between Achkhoti, Panseti [P’anshet’i / Pancheti] and Kasbegi [Kazbegi district / Qazbegi district / Stepantsminda district], 42°39’07’’N, 44°38’04’’E, 1750–1780 m, 20 July 2002, G. Schneeweiss et al. (WU: S&T 8581) ( Schneeweiss et al. 2004: Supplementary Data 1 Page 10); AY209326: Georgia, province of Chewi (Khevi, Chevi), Great Caucasus, Terek valley, 1–2 km S of Gerget‘i, that is towards the village P’anshet’i [Kazbegi district / Qazbegi district / Stepantsminda district], 1750–1850 m, G. Schneeweiss (WU: s.n.); AY960741: Georgia, province of Chewi (Khevi, Chevi), Great Caucasus, Terek valley, 1–2 km S of Gerget‘i, that is towards the village P’anshet’i [Kazbegi district / Qazbegi district / Stepantsminda district], 1750–1850 m, G. Schneeweiss (WU: S&T 5610) ( Carlón et al. 2005: 67, as P. pulchella ); 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); Iskari N, [in the Thymus clump], 43°03’19.2”N, 42°31’44.8”E, [ca. 1700 m], 9 June 2022, J. Pietruszewski (phot., https://www.inaturalist.org/ observations/121042627); Samtskhe –Javakheti prov.: “α” Borjomi, Tbilisi province, near the cemetery, 4 June 1888, Akinfiew (LE01015368). This herbarium sheet also includes the following specimens: Transcaucasia, Borjomi, at an elevation of 3000 ft, 3 July 1887, Akinfiew; Borjomi, Tbilisi province, 2500 ft, 9 June 1888, Akinfiew (LE01015369); Borjomi, Likani, 3000’, 28 June 1888, Akinfiew (LE); prov. Tiflis, distr. Gori, Borjomi, in silva prope Likani, 27 June 1920, B. Schischkin (TGM8425, left specimen) [as O. arenaria by Melikischvili]; Borjomi, slopes of the Mtkvari (Kura) river, the overgrown rocky slopes above the cemetery as well as on the graves and near the paths directly in the cemetery, on Teucrium , 41°50’15”N, 43°22’23”E, 830–840 m, 14 June 2022, R. Piwowarczyk (KTC); between Borjomi and Kvibisi, NE end of Borjomi, rocky slopes of the Mtkvari (Kura) river, on Teucrium , 41°51’23.7”N, 43°24’50.1”E, 810 m, 15 June 2022, R. Piwowarczyk (KTC). TURKEY. Rize prov.: Lazistan, près Djimil-région [Cimil, Pontic Mountains], sous alpine alt. v. 2000 m, Juillet 1906 [see B. Balansa, Plantes d’Orient, 1866] (L2817723, as Phelypaea araneosa Reuter and as Orobanche caesia var. borealis ]); Ph. bungeana (Beck) Soják - AY209330: Turkey, NE Anatolia, between Ardanuç and Bulanik, 1 km N of Bulanik, 1120 m, on Teucrium sp. (Lamiaceae) , G. Schneeweiss & C.J. (WU: S&T 7768) ( Carlón et al. 2005: 66, as P. bungeana ).

Figures

FIGURE 10. Phelipanche pulchella. A–D. General habit. E–G. Flower, side view. H–J. Flower, front view. K, M. Representative habitats in Borjomi (Samtskhe-Javakheti prov.). L. Host, Teucrium chamaedrys. Photos by R. Piwowarczyk.

FIGURE 11. Distribution of Phelipanche pulchella in Georgia.