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46Новости систематики высших растений Novitates Systematicae Plantarum Vascularium
2022
53: 89-101
ISSN 0568-5443 (print) ISSN 2687-1564 (online)
Systematic revision of Orobanche pycnostachya and first record of O. filicicola (Orobanchaceae) in Russia
Систематическая ревизия Orobanche pycnostachya и первая находка O. filicicola (Orobanchaceae) в России
Stefan Ratzel1, Holger Uhlich2*
1 Georg-Friedrich-Handel-Str., 13 15234, Frankfurt at Oder, Germany [email protected]
2 Gartenstr., 19, 56357, Welterod, Germany [email protected]
Corresponding author
https://doi.org/10.31111/novitates/2022.53.89
Стефан Рэтцель1, Хольгер Улих2*
1 ул. Георга-Фридриха-Генделя, 13 15234, Франкфурт-на-Одере, Германия [email protected]
2 ул. Гартенштрассе, 19, 56357, Вельтерод, Германия [email protected]
*Автор для переписки
Abstract. Plants whose morphological characteristics correspond to those of Orobanche filicicola Nakai ex Hyun, Lim et Shin as described by Nakai (1952) and validated by Huyn et al. (2003), were in 2012 found for the first time in Russia (Far East, Primorye Territory) growing on the banks of the Ussuri River in the Chuguyevsky District and in a mixed population with O. amurensis (Beck) Kom. on the SE flank of the Sikhote-Alin Mountains (Lazovsky District). The status of the taxon has been revised. As a result, it is reclassified as a new variety of O. pycnostachya Hance, namely as O. pycnostachya var. filicicola (Nakai ex Hyun, Lim et Shin) Rätzel et Uhlich. In this context, O. pycnostachya and O. amurensis are discussed, including the relationship of O. amurensis with O. pycnostachya. In addition, O. coerulescens Stephan, O. coerulescens f. cristata Beck, O. coerules-cens f. lehmannii Beck, O. coerulescens f. pekinensis Beck, O. coerulescens var. arenaria Serg., O. macrolepis Turcz., O. pycnostachya, O. pycnostachya f. amurensis Beck, O. pycnostachya f. albo-lutea Beck, O. quadrivalvis Regel and O. ussuriensis Novopokr. are lectotypified. Three necessary new combinations of infraspecific taxa from O. coerulescens — O. coerulescens var. nipponica (Makino) Uhlich et Rätzel, O. coerulescens var. occidentalis f. multidentata (Beck) Uhlich et Rätzel and O. coerulescens f. arenaria (Serg.) Uhlich et Rätzel — are made. The name O. quadrivalvis is to be used as a synonym of O. kotschyi Reut.
Keywords: Orobanchaceae, Orobanche, O. amurensis, O. coerulescens, O. filicicola, O. kotschyi, O. pycnostachya, Russia, Far East, Primorye Territory, new combinations, lectotypification, synonymization.
Аннотация. Растения, морфологические характеристики которых соответствуют Orobanche filicicola Nakai ex Hyun, Lim et Shin, описанному Nakai (1952) и валидизированному Huyn et al. (2003), были обнаружены впервые в России в Приморском крае на берегах Уссури в Чугуевском районе и в смешанной популяции с O. amurensis (Beck) Kom. на юго-восточном макросклоне Сихотэ-Алиня в Лазовском районе в 2012 году. В результате таксон принят в ранге разновидности — O. pycnostachya Hance var. filicicola (Nakai ex Hyun, Lim et Shin) Rätzel et Uhlich. Даны соответствующие комментарии к O. pycnostachya и O. amurensis, a также к систематической структуре родства O. pycnostachya. Названия O. coerulescens Stephan, O. coerulescens f. cristata Beck, O. coerulescens f. lehmannii Beck, O. coerulescens f. pekinensis Beck, O. coerulescens var. arenaria Serg., O. macrolepis Turcz., O. pycnostachya, O. pycnostachya f. amurensis Beck, O. pycnostachya f. albo-lutea Beck, O. quadrivalvis Regel и O. ussuriensis Novopokr. лектотипифицированы. Обнародованы три новые комбинации внутривидовых таксонов O. coerulescens: O. coerulescens var. nipponica (Makino) Uhlich et Rätzel, O. coerulescens var. occidentalis f. multidentata (Beck) Uhlich et Rätzel и O. coerulescens f. arenaria (Serg.) Uhlich et Rätzel. Название O. quadrivalvis синонимизировано с O. kotschyi Reut.
Ключевые слова: Orobanchaceae, Orobanche, O. amurensis, O. coerulescens, O. filicicola, O. kotschyi, O. pycnostachya, Россия, Дальний Восток, Приморский край, новые комбинации, лектотипификация, синонимизация.
Within the genus Orobanche L., the subsection Coerulescentes (Beck) Novopokr., which has only a few members1, occupies a phylogenetically relatively
1 According to more recent phylogenetic studies (Li et al., 2017; Piwowarczyk et al., 2021), Orobanche boninsimae (Maxim.) Tuyama (= Platypholis boninsimae Maxim.) also belongs to the
Поступила в редакцию | Submitted: 21.12.2021
exceptional position (cf. Schneeweiss et al., 2004; Li et al., 2017; Piwowarczyk et al., 2021). In contrast to
genus Orobanche, in particular to subsect. Coerulescentes. However, that taxon differs very significantly from the other representatives of that subsection, both morphologically and ecologically (e. g. completely different habitats and hosts).
Принята к публикации | Accepted: 20.12.2022
other sections and subsections of the genus Orobanche, it has its main area of distribution in East Asia; only O. coerulescens Stephan emend. Willd., which has an extremely large total distribution area, also reaches Central Europe. From an ecological point of view it should be noted that numerous species of this subsection (but also the sect. Inflatae (Beck) Rouy) parasitize exclusively on members of the genus Artemisia L.; but then in a broad spectrum, that sometimes goes beyond the respective section.
Materials and methods
During a several-week botanical excursion to various regions of the Russian Far East in 2012, the first author of this article collected material from O. coerulescens and taxa from the relatives of O. pycnostachya Hance.
List of herbarium specimens of O. coerulescens, O. pycnostachya var. filicicola (Nakai ex Hyun, Lim et
Shin) Ratzel et Uhlich and var. pycnostachya f. amu-rensis Beck, collected by S. Ratzel in the Russian Far East: Appendix 1.
In addition, on several visits to the Komarov Botanical Institute of the Russian Academy of Sciences in Saint Petersburg (LE) since 2015, the second author studied the types and other herbarium material of the members of the subsection Coerulescentes. Besides the relevant herbarium material of the members of the subsection has been researched in various Herbaria (B, DR, FR, GFW, GLM, H, HAL, JE, LD, LZ, Teryokhin's Orobanche herbarium in the Botanical Museum of the Komarov Botanical Institute, a. o.) of the world and other sources (e. g. Chinese Virtual Herbarium, 2021; GBIF, 2022; Herbarium Beroli-nense — Virtual Herbarium, 2022; Naturalis Biodiversity Center, 2022; Vascular Plants Herbarium of the Komarov Botanical Institute RAS — Herbarium LE, 2022; Virtual Herbaria JACQ, 2022) for a number of years.
Results
Until now, O. filicicola Nakai ex Hyun, Lim et Shin was considered endemic to South Korea (Hyun et al., 2003: 64). This is surprising because habitats with clearly ruderal characteristics are named for this taxon (e. g. Hyun et al., 2003: 66; Lee et al., 2011: 273). During the above-mentioned excursion in 2012, material attributable to O. filicicola was found on a bank terrace of the Ussuri River not far from the sett 1 ement of Sokolovka (Fig. 1: A; 2: A, B). The specific dimensions and characteristics agree with the description of O. filici-cola given in the protologue (Hyun et al., 2003) and the information in the table (Hyun et al., 2003: 66). The location on the bank of Ussuri River corresponds to the original locations of the host and parasite (cf. Hyun et al., 2003: 66 — "gravelly place between the river and the forest margin along a roadside". In our view, the plants of O. pycnostachya s. l. are spreading because human activities and increasing urban sprawl are constantly creating suitable habitats for their hosts and the parasites.
Furthermore, O. amurensis (Beck) Kom. was found growing in mixed stands with plants of the O. filicicola-habit on the southeast flank of the Sikhote-Alin Mountains (Fig. 1: B; 2: C, D).
During the excursion it was confirmed that, in contrast to O. pycnostachya s. l., O. coerulescens, which also occurs in the Far East, populates natural or near-natural habitats such as rocks (especially coastal rocks along the Sea of Japan), dunes etc., which, as far as we know,
Fig. 1. Habitats of Orobanche pycnostachya var. filicicola (A, B) and O. pycnostachya var. pycnostachya f. amurensis (B). A — terraces of the Ussuri River near Sokolovka, dominated by Artemisia cf. argyi (host of O. pycnostachya var.filicicola), 15 VI 2012; B — southeast flank of the Sikhote-Alin, Lazovsky District, ruderal vegetation, dominated by Artemisia cf. umbrosa (host of both taxa), 20 VI 2012. Photos by S. Ratzel.
are places that are in fact never ruderally covered or disturbed.
Discussion
The morphological differences between the above three taxa (О. amurensis, O. filicicola and O. pycnostachya) were described and summarized in Hyun et al. (2003).
In our opinion, the inflorescence of O. pycnostachya, which was described as "always dense" in Hyun et al. (2003), cannot be maintained as a characterising feature. In addition, we do not agree, that as described by Hyun et al. (2003: 66, tab. 1), different ratio of the length of the inflorescence to the length of the flowerless part of the stem is a consistent distinguishing feature of O. filicicola from O. amurensis and O. pycnostachya. This ratio is variable in the course of the phenological development of the plant. The specimens of O. pycnostachya available in Chinese Herbaria (Chinese Virtual Herbarium, 2021) show plants with inflorescences which are often shorter, but also sometimes longer than the flower-less part (e. g. PE 01476129, PE 01476083, PE 01476084). Photographs by O. filicicola on the Internet (e. g. Naejangsan National Park Service, 2018) similarly show plants with inflorescences which are often longer and looser, but also sometimes shorter. The first author of this article also found plants which are attributable to O. amurensis but which have a significantly longer inflorescence (Fig. 2: C, D) than that described by Hyun et al. (2003). The flower colour also varies, e. g. in O. filicicola, from a bright dark to pale blue, exceptionally white. In our opinion, this eliminates the differences, which, among other things, were the basis for the differentiation at the rank of species in Hyun et al. (2003) between O. filici-cola on the one hand, and O. pycnostachya and O. amurensis on the other.
The other differences in characteristics between the above three taxa discussed by Hyun et al. (2003) are exclusively of a quantitative nature. They are essentially limited to insignificant differences in size. Other, non-quantitative characteristics that would justify a demarcation between these taxa at species rank are absent. There is also a clear overlap of the spectrum of host plants (Table). Thus all three taxa parasitize Artemisia host species, several of which are the same for all three.
Fig. 2. Habits (A, C) and inflorescences (B, D) of Orobanche pycnostachya subtaxa.
A, B — O. pycnostachya var. filicicola, Ussuri terraces near Soko-lovka, on Artemisia cf. argyi (host), 15 VI 2012; C, D — O. pycnostachya f. amurensis, Sikhote-Alin, Lazovsky District, 20 VI 2012. Photos by S. Rätzel.
In terms of vegetation ecology, too, no significant difference can be found between the taxa under discussion. With their hosts, all three taxa colonize locations with high habitat dynamics (habitat turnover due to
Table. Overview of the hosts used by Orobanche pycnostachya s. l. according to the current state of knowledge
Host (Artemisia L.1) "O. amurensis" "O. filicicola" O. pycnostachya s. str.
Section Series Species
Subgenus Artemisia
Abrotanum (Duhamel) Besser Laciniatae Kitam. A. laciniata Willd. Novopokrovsky, Tzvelev, 1958: 74; Kurbatsky, 1996: 94
Paniculigerae Poljakov, nom. inval. A. keiskeana Miq. Schischkin, 1930: 151
Ponticae [Rydb. unranked] A. sacrorum Ledeb. (= A. gmelinii Web. ex. Stechm.) Novopokrovsky, Tzvelev, 1958: 74 Beck, 1930: 117; Hsu, Wang, 1996: 99; Korsun, 2014
Artemisia Angustilobae Polja-kov, nom. inval. A. argyi H. Lév. et Vaniot PE 01476152, IFP 13202004x0021, IFP13202004x0022 Rätzel, published here (cf.)
A. selengensis Turcz. ex Besser IFP13202004x0016
Simplicifoliae Poljakov, nom. inval. A. stolonifera (Maxim.) Kom. Schischkin, 1930: 151; IFP13202004x0016
Artemisia A. umbrosa (Turcz. ex Besser) Pamp. Rätzel, published here (cf.) Rätzel, published here (cf.)
A. vulgaris L. Schischkin, 1930: 151
Subgenus Dracunculus (Besser) Peterm.
Commutatae Polja-kov, nom. inval. A. campestris L. Maximowicz, 1859: 211
A. commutata Besser Novopokrovsky, Tzvelev, 1958: 74; Kurbatsky, 1996: 94 Novopokrovsky, Tzvelev, 1958: 73
Japonicae Krasch. ex Poljakov, nom. inval. A. capillaris Thunb. PE 01476155
A. desertorum Spreng. Maximowicz, 1859: 211 Beck, 1930: 117
?Psammophilae Poljakov, nom. inval. A. ordosica Krasch. Hsu, Wang, 1996: 99
natural perturbations or human activity), e. g. river valleys, and especially those with anthropogenic influence (i. e. ± ruderalized locations). Differences in altitude of the populated locations are just as difficult to recognize as their own distribution area. The result of our researches shows that O. filicicola tends to be a small-statured and small-flowered variety of O. pycnostachya. Like O. amurensis, it has ± intensely (purple) blue flowers and usually white stigmas. O. pycnostachya s. str. in contrast, has ± completely yellowish-white flowers and yellow stigmas.
Orobanche filicicola is therefore comparable with taxa such as O. coerulescens var. occidentalis (Beck) O. Fedtsch. et B. Fedtsch. (cf. for the classification of the var. occidentalis also Uhlich, in prep.) from the same subsection, O. owerinii Beck var. woronowii Beck and O. crenata Forssk. var. sylvestris Beck. These taxa are
1 The system of the genus Artemisia follows the treatment in the "Flora URSS" (Poljakov, 1961), including invalidly published names of series.
also small-flowered varieties of what are usually clearly larger-flowered species.
There are no consistent characterising features which justify the present classification of O. filicicola at the rank of species. Instead, it is more appropriate to reclassify it as a variety of O. pycnostachya.
Of the three taxa dealt with herein, the name Orobanche pycnostachya Hance (Hance, 1872) has priority at the rank of species. Since O. filicicola has not been previously named at the rank of variety, we now classify it at that rank.
We consider O. amurensis a colour form of O. pycnostachya. In the subsection Coerulescentes the occurrence of pale-yellowish and ± whitish corollas is also known in other taxa (e. g. O. coerulescens f. albiflora (O. Kuntze) H. Hara = O. korshinskyi Novopokr.). All features of O. amurensis, except for the colours of the corolla and stigma, are identical to, or overlap with those of O. pycnostachya s. str. Characteristics presented as differences in the literature (such as the
representation of style hairiness as "sparsely glandular" for O. pycnostachya versus "subglabrous" for O. amurensis in Hyun et al., 2003; cf. previously Beck, 1890, 1930) fall within the usual variability of these taxa and do not represent consistent or taxonomically usable differences.
We therefore propose an overall breakdown of the three taxa discussed as follows:
Systematic arrangement of Orobanche pycnostachya s. l. and reclassification of O. filicicola as a variety of O. pycnostachya
Orobanche pycnostachya Hance, 1872, J. Linn. Soc., Bot. 13, 66: 84. — Lectotype (designated here): "17048. Orobanche L. pycnostachya Hance. Herb. H. F. Hance. Rec[eive]d. 1887. In m[onti] bus ad occasum Peking sitis, m[ens]. Junio 1870, Dr. E. Bretschneider" (BM: BM000997919 (scan!), the first plant from the right of the specimen sheet; isolec-totypes: BM000997919, all other plants on the specimen sheet). — Fig. 1 in Appendix 2.
O. pycnostachya var. pycnostachya.
O. pycnostachya var. pycnostachya f. pycnos-tachya.
= O. pycnostachya f. typica Beck, 1890, Biblioth. Bot. 19: 141, tab. 2, fig. 32, nom. inval. = O. pycnostachya var. genuina Beck f. typica (Beck) Beck, 1930, in Engl., Pflanzenr. 4 (261): 118, nom. inval.
= O. macrolepis Turcz. 1851, Bull. Soc. Imp. Naturalistes Moscou, 24 (4): 356, nom. illeg., non Coss. 1848. — Lectotype (designated here): "Orobanche macrolepis m[ihi]. Ad radices rupium fl. Schilka cingentium, 1839, Turcz." (LE: LE 01057582 (scan!) — Fig. 2 in Appendix 2); syntype: "Orobanche macrolepis m[ihi]. Ad radices rupium fl. Schilka, cingentium, 1833, Turcz." (LE 01205156!); original specimen: LE 01205154!
= O. pycnostachya var. genuina Beck f. albo-lu-tea Beck, 1890, Biblioth. Bot. 19: 141, tab. 2, fig. 32. — Lectotype (designated here): "560. Herbarium horti Petropolitani. Orobanche. Weiße, wohlriechende Blüthen, mit gelben Pistillen. Gelber Schaft. Braune Blattschuppen. Beinahe 1 Fuß hoch. Tang-shan im Parke. 23. Mai. Flora Pekinensis. Dr. Bretschneider 1877", "O. pycnostachya Hance f. albo-lutea Beck" (LE: LE 01205320! (both plants) — Fig. 3 in Appendix 2); syntype: "561. Blüthen wie die der vorigen, ...Bis 2 Fuß hoch. Yiu shan. 25. Mai. Flora Pekinensis. Dr. Bretschneider 1877" (LE 01205157!).
— O. pycnostachya var. zaisanica Teryokhin, 1993, in Teryokhin et al., Opred. Zaraz. Fl. SSSR: 37, nom. inval., sine typo.
- O. pycnostachya var. pycnostachya auct. non Hance: Park, 2007, Gen. Vasc. Pl. Korea: 895.
O. pycnostachya var. pycnostachya f. amurensis
Beck, 1890, Biblioth. Bot. 19: 141.
= O. pycnostachya var. amurensis (Beck) Beck, 1930, in Engl., Pflanzenr. 4 (261): 118. = O. amurensis (Beck) Kom. 1907, Trudy Imp. S.-Peterburgsk. Bot. Sada, 25: 469; id. 1907, Fl. Manshur. 3, 2: 469. — Lectotype (designated here): "Orobanche ammophila C. A. Mey. 2 VII 1860. Bai Possjet, auf Wiesenland nicht selten, fl. azurei. Bai Victoria, Mandshuria, Maximowicz [hand of Maximowicz]"; "Залив Посьета, на лугах, не редко, 2 VII 1860, Максимович [hand of Komarov]" (LE: LE 01015418!, the complete specimen). — Fig. 4 in Appendix 2.
N o t e . Beck (1890) cites the collections by Maxi-mowicz: "Mandshuria et territ. fl. Amur: ad Bureiae ostium, prope Possjet, Poriph, Poddale, Dsikiri prope Blagowe-stschensk (f. 3 [i. e. f. amurensis] leg. Maximowicz)", as well as refers to the specimens that Maximowicz (1859) mentioned in his "Primitiae florae amurensis" for the most part under the name O. ammophila. Thus, the collection LE 01015418 labelled by Tzvelev as "authentic specimen" is therefore, from our point of view, entirely suitable as a type material.
We select the entire specimen as a unit, as individual fragments cannot be reliably assigned.
= O. ussuriensis Novopokr. 1950, Bot. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk SSSR, 12: 274. -Lectotype (designated here): "Переселенческое Управление; экспедиция В. Л. Комарова в ЮжноУссурийский край. V. Komarov, Iter Austro-Ussu-riense. № 431. Orobanche amurensis G. Beck. Бассейн р. Сучана. Селение Владимирская падь во Влад.-Александровске. Формация Кустарников[ая] заросль с высокой трав[янистой] рас[тительност]ью. Почва Перегной. 25 VI 1913, А. Булавкина"; "Orobanche ussuriensis Novopokr. Determ. I. Novopokrovsky 12 XII 1948"; "= O. amurensis (G. Beck) Kom. (Typus O. ussuriensis Novopokr.). Teste: Цвелёв 19 II 1955" (LE: LE 01015423!, the complete specimen). — Fig. 5 in Appendix 2.
Orobanche pycnostachya Hance var. filicicola (Na-
kai ex Hyun, Lim et Shin) Rätzel et Uhlich, comb. et stat. nov.
= O. filicicola Nakai ex Hyun, Lim et Shin, 2003, Novon, 13, 1: 64; Nakai, 1952, Bull. Natl. Sci. Mus. 31: 104, nom. nud. — Holotype: "Korea. Chunnam: ca. 1 km N of the Office of National Parks Authority at Mt. Bailyang, along roadside, running parallel with the Hwangryong River, Changsung-Gun. 1 VI 1999, Jin O. Hyun 1999" (SNU; isotypes: KB (from AJOU), MO5694578, scan!, SNU; paratype: TI).
Fig. 3. Lectotype (right plant) and isolectotype of Orobanche coerulescens (HAL0090414; https://hal.jacq.org/ HAL0090414). Courtesy of the Herbarium of the University Halle/Saale (CC-BY 4.0).
HAL0090414
Further reclassifications and lectotypifications
Orobanche coerulescens Stephan emend. Willd. 1800, in Willd., Sp. Pl. 3: 349. — Lectotype (designated here): "Orobanche coerulescens e Siberia versus mare Caspium (Stephan), W[illdenow] det. [hand of D. F. L. v. Schlechtendal, pat.]" (HAL: HAL0090414!, right plant; isolectotype: HAL0090414, left plant). — Fig. 3.
Note 1 . In view of the author citation "Stephan emend. Willd.", it follows from the protologue (Willdenow, 1800: 349) that Stephan provided both the name O. coerulescens and brief diagnosis ("Steph. in litt."; see the second paragraph of the protologue), while Willdenow completed this diagnosis in the first and last paragraphs of the protologue (each with his abbreviation "W.").
In Berlin (B-WILLD: B -W 11594 -01 0!, B -W 11594 -02 0!), there are probably isolectotypes.
Note 2 . According to Novopokrovsky and Tzvelev (1958), and Tzvelev (2006), the locus classicus is assumed to be in western Kazakhstan in the vicinity of Lake Inder ("Описан из района Индерского озера"). This assumption is based on a mixed collection, the left hand plant of which was collected by Stephan: "Orobanche coerulescens mihi, selten, [на] Индерской горе [Inderskoi gore. Hand of Stephan]", "Herb. Stephan. Orobanche caerul[escens]. Clas. XIV, Montes Inderienses [hand?]", "Orig. Stephan. Orobanche (Trionychon) coerulescens Stephan, typ-ica, det. Dr. G. Beck [hand of Beck?]", "Typus! 14 V 1949, I. Novopokrovsky. Not. crit. [hand of Novopokrovsky]" (LE 00050163! — Fig. 4). However, the specimen does not reveal that it was available to Willdenow for the protologue (compare to: Turland et al., 2018: Art. 9.4); the location is also not mentioned in the protologue. Beck's assignment of the species to the "Trionychon" section is apparently an oversight. The reference to a type located in St. Petersburg (LE) in Novopokrovsky and Tzvelev (1958) obviously goes back to this specimen.
Compared to the herbarium specimen LE 00050163, we consider the herbarium specimen HAL0090414 to be more suitable as a lectotype: the specimen contains the phrase from the protologue almost verbatim. There is no evidence that Willdenow saw the St. Petersburg specimen. Had he seen it, he would probably have chosen the relatively precise locality "Lake Inder" or "Inderskoi gore" as the classic locality and not the broad formulation "in Siberia versus Mare Caspium". The writer of the label of the herbarium from Halle, D. F. L. von Schlechtendal (1767-1842), was a contemporary of Willdenow and was in close contact with him. The herbarium specimen probably reached Halle from Willdenow. The abbreviation "W." on the label implied that Will-denow, who abbreviates himself that way, saw the receipt himself.
Orobanche coerulescens var. coerulescens f. arenaria (Serg.) Uhlich et Rätzel, stat. nov.
= O. coerulescens var. arenaria Serg. 1939, in Krylov et al., Fl. Zapadnoi Sibiri, 10: 2540. — Lectotype
(designated here): "Гербарий Томского университета. Orobanche coerulescens C. A. Mey. var. arenaria m[ihi] (?). Семипалатинская] губ., Зайсанск[ий] у. Между д. д. Николаевкой и Александровкой -48^° с. ш. и 55%° в. д., на песках Бланды-кум. 22 VII 1929, П. Крылов и Л. Сергиевская" (TK: TK-002057 (the second plant from the right; scan!) — Fig. 5; isolec-totypes: LE 00053850! ("I. Novopokrovsky. Notae criti-cae. Orobanche coerulescens Steph. f. antheris pilosiuscu-lis. 15 VII 1948"), TK-002057 (all other plants)).
This plant differs from typical O. coerulescens only in its relatively tall habit (35-50 cm high), a longer, above rounded spike (16-20 cm) and hairy anthers (cf. Kry-lov et al., 1939). The height of the plant and the length of the inflorescence are within the variability of the species. Only the hairy anthers are a systematically and ta-xonomically relevant feature. The taxon is only known from the type locality. These circumstances justify the classification in the rank of a form.
O. coerulescens var. coerulescens f. cristata Beck, 1890, Biblioth. Bot. 19: 138.
Lectotype (designated here): "Orobanche: radicibus Artemisiae campestris innascens in insul. arenos. Wolgae. Bunge. Herb. Fischer", "a) O. coerulescens Steph. b) f. cristata Beck [hand of Beck]", "Lectoty-pus Orobanche coerulescens f. cristata Beck, Mon. Orob.: 138. 1890. Holger Uhlich, Welterod, Germany (2020)" (LE: LE 01060143! — Fig. 6 in Appendix 2).
O. coerulescens var. coerulescens f. lehmannii Beck, 1890, Biblioth. Bot. 19: 139 ("lehmanni").
Lectotype (designated here): "Alexandri Lehmann Reliquiae botanicae. Orobanche coerulescens. Al. Bunge", "O. coerulescens Steph. f. O. Lehmanni Beck [hand of Beck]", "Holotypus O. coerulescens f. lehmannii Beck. Holger Uhlich, Welterod, Germany (2019)" (LE: LE 00053849 — Fig. 7 in Appendix 2).
O. coerulescens var. coerulescens f. pekinensis
Beck, 1890, Biblioth. Bot. 19: 138.
Lectotype (designated here): "556. Herbarium horti Petropolitani. Orobanche. Blau, geruchlos. — Pan shan. 29. Mai. Flora Pekinensis. Dr. Bretsch-neider 1877", "O. coerulescens Steph. f. O. pekinensis mihi. Beck [hand of Beck]", "Orobanche coerulescens Steph. (non O. ombrochares Hance!). Teste N. Tzvelev (LE) XII 1995", "№ 556. Orobanche coerulescens f. pekinensis Beck, Monogr. Orob.: 138. 1890. Lectotypus! (the middle larger plant) Hic designatus: H. Uhlich 2 IV 2021" (LE: LE 01072663!, the middle larger plant; isolectotypes: LE 01072663, the other two plants from the collection). — Fig. 8 in Appendix 2.
Fig. 4. Original specimen of Orobanche coerulescens (LE 00050163, left plant; http://rr.herbariumle.ru/00050163). Courtesy of the Komarov Botanical Institute of the Russian Academy of Sciences.
LE 00050163
Fig. 5. Lectotype (second plant from the right) and isolectotypes of Orobanche coerulescens var. arenaria (TK-002057; https://tk.botdb.ru/r/002057). Courtesy of the P. N. Krylov Herbarium, Biological Institute, Tomsk State University.
TK-002057
Note. There are two gatherings on the sheet, the second one is: "558. Herbarium horti Petropolitani. Orobanche. Blau, geruchlos. — Miao feng shan. 15. Juni. Flora Pekinensis. Dr. Bretschneider 1877", "O. coerulescens Steph. f. O. ombro-chares Hance. Beck [hand of Beck]" (LE 01072664!).
Orobanche coerulescens var. nipponica (Makino) Uhlich et Rätzel, stat. nov.
= O. nipponica Makino, 1928, J. Jap. Bot. 5, 10: 40. = O. coerulescens f. nipponica (Makino) Kitam. 1957, Acta Phytotax. Geobot. 17, 1: 8.
Type specimen unknown, possible in Makino Herbarium (Tokyo Metropolitan University), not seen.
The classification as a variety is justified because of its own East Asian distribution area and because of significant differences between the taxon (the entire plant is glabrous) and typical O. coerulescens. In this context, reference must be made to O. coerulescens var. glaberrima Sakata (1935, Bull. Agric. Forest. Coll. Suigen (Suwon), 91: 32). The latter name might have priority as the oldest combination in the rank of variety. A clarification was not possible, however, since the literature was not accessible to the authors.
Orobanche coerulescens var. occidentalis (Beck) O. Fedtsch. et B. Fedtsch. f. multidentata (Beck) Uhlich et Rätzel, stat. nov.
= O. coerulescens f. occidentalis subf. multidentata Beck, 1930, in Engl., Pflanzenr. 4 (261): 121.
Protologue citation: "Rußland: Permer Gubernement (Kryloff!)". Type specimen possibly in LE, not seen.
The recombination was necessary because Fed-tschenko and Fedtschenko (1913) upgraded the f. occidentalis to a variety — rightly in our opinion.
Note on Orobanche coerulescens var. coerulescens f. ombrochares (Hance) Beck (1930, in Engl., Pflanzenr. 4 (261): 121. = O. ombrochares Hance, 1872, J. Linn. Soc., Bot. 13: 84).
Beck (1930: 121) has the plants, collected under № 15872 by S. W. Williams and Bretschneider (BM000997917 (scan!), BM000997918 (scan!), K000061378 (scan!), K000061379 (scan!); Hance refers only to S. W. Williams!) seen and considers them to be typical specimens of O. coerulescens. In the original description, the plant has all the characteristics of the typical O. coerulescens, except for the purple-violet colour of the flowers. Although Hance (1873: 84) states glabrous anthers, it is coded with very hairy anthers as a distinguishing feature in more modern "Floras" (e. g. Zhang, Tzvelev, 1998), which often place them in the rank of the species. Further investigations are required for a critical assessment of the taxon.
Note on Orobanche quadrivalvis Regel (1880, Trudy Imp. S.-Peterburgsk. Bot. Sada, 6: 354).
Beck (1890: 138, 1930: 120) and Sánchez Pedraja et al. (2022) place this name in the synonymy of O. coerulescens. As part of the research at the Herbarium LE, the type specimen of O. quadrivalvis was found and is selected here as the lectotype of this name.
Lectotype (designated here): "A. Regel. Iter Turkestanicum 1878 VIII", "Orobanche quadrivalvis Rgl., Obere Borotala, 8500', 18 VIII [1878], A. Regel", "Orobanche kotschyi Reuter? 13 VIII 1947. De-term. I. Novopokrovsky", "Orobanche solmsii Clarke. 1995. Teste N. Tzvelev (LE)", "Orobanche quadrivalvis A. Regel, ... Lectotypus! Hic designatus: H. Uhlich 04. April 2021" (LE: LE 01072662! — Fig. 9 in Appendix 2). Based on the type, it can be determined that the collected plant clearly belongs to O. kotschyi Reut. The name O. quadrivalvis is therefore to be used as a synonym of O. kotschyi.
Identification key for the differentiation of the taxa of the Orobanche subsection Coerulescentes in the Russian Far East (excl. O. boninsimae (Maxim.) Tuyama)
1. Inflorescences (incl. bracts, calyces and corollas) glandular pubescent, rarely sparsely villous. Lobes of the corolla at margin ± long glandular ciliate. Stigma yellow or white, often projecting. Anthers at suture dense woolly to (almost) glabrous (check several flowers or plants)..................
..................................................................... O. pycnostachya ... 2.
+ Inflorescences (incl. bracts, calyces and corollas) whitish arachnoid-villous, rarely glabrous. Lobes of the corolla glabrous at margin. Stigma white, usually enclosed in the corolla. Anthers glabrous or slightly hairy at suture, rarely hirsute or densely white tomentose-villous along suture (check several flowers or plants)......... O. coerulescens... 4.
2. Corolla yellow or white. Stigma yellow. Inflorescence
tends to ± dense ..............................................................................
............. O. pycnostachya var. pycnostachya f. pycnostachya.
+ Corolla ± intensive bluish or azure (rarely also plants with a white corollas in otherwise typically coloured populations). Stigma white. Inflorescence tends to ± lax (especially in the lower part) .......................................................... 3.
3. Plant 10-45 cm tall. Bracts 15-20 mm long, broadly lanceolate. Calyx 8-18 mm long. Corolla 18-25 mm long.
Style almost glabrous ...................................................................
.................... O. pycnostachya var. pycnostachya f. amurensis.
+ Plant 13-30 cm tall. Bracts 10-16 mm long, linear-lanceolate. Calyx 8-11 mm long. Corolla 13-22 mm long. Style sparse glandular hairy................O. pycnostachya var. filicicola.
4. Inflorescences (like the whole plant) glabrous......................
.................................................. O. coerulescens var. nipponica}
+ Inflorescences (incl. bracts, calyces and corollas) whitish arachnoid-villous ...................................................................... 5.
5. Anthers almost glabrous, very rarely hairy ........................ 6.
1 See above "Further reclassifications".
+ Anthers hirsute or densely white tomentose-villous along suture......................................................................................... 12.
6. Corolla large, 20-23 mm long............................................... 7.
+ Corolla small, 10-18(21) mm long...................................... 8.
7. Corolla 20-22 mm long, conspicuous prominent. Lobes of the lower lip mostly distinctly denticulate. Calyx segments scarcely divided to the middle. Bracts a little shorter than or as long as the corolla ................................................
.....................O. coerulescens var. coerulescens f. coerulescens.
+ Corolla large 22-23 mm long. Lobes of the lower lip broadly elliptical. Calyx segments deep (often down below
the middle) 2-toothed. Bracts as long as the corolla ............
......................... O. coerulescens var. coerulescens f. lehmannii.
8. Corolla very small, 10-15 mm long, dark blue......................
........................ O. coerulescens var. coerulescens f. pekinensis.
+ Corolla 16-21 mm long, intense blue-violet or blue to almost whitish .............................................................................. 9.
9. Calyx and bracts dense white lanate. Corolla often intensive coloured ...................................................................................
.................... O. coerulescens var. coerulescens f. ombrochares.
+ Calyx and bracts not dense white lanate.......................... 10.
10. Bracts longer than the corolla and therefore inflorescences
at apex comose. Calyx teeth as long as the corolla tube ......
............................. O. coerulescens var. coerulescens f. cristata.
+ Bracts as long as the corolla or shorter. Flowers small, barely 20 mm long, scarcely exceeding the bracts, often
hid by bracts and calyx segments ..............................................
.......................................O. coerulescens var. occidentalis... 11.
11. Calyx segments 2-toothed ..........................................................
....................... O. coerulescens var. occidentalis f. occidentalis.
+ Each calyx segment 3-toothed, dorsally slightly connate.
Plant robust. Inflorescences multi-flowered...........................
.....................O. coerulescens var. occidentalis f. multidentata.
12. Anthers hirsute. Stem 35-50 cm tall. Inflorescences 1620 cm long, obtuse at apex ..........................................................
........................... O. coerulescens var. coerulescens f. arenaria.
+ Anthers densely white tomentose-villous along suture. Stem 15-30 cm tall, densely white arachnoid-tomentose throughout. Inflorescences 5.08-6.35 cm (sec. Hance, 1873: "2-2.5 pollicaris") or 8-15 cm (sec. Zhang et Tzve-
lev, 1998) long, rounded at apex ................................................
.................... O. coerulescens var. coerulescens f. ombrochares.
The other species of the subsection Coerules-centes — O. megalantha Harry Sm., O. mupinen-sis Hu, O. sinensis Harry Sm., incl. var. cyanescens (Harry Sm.) Zhi Y. Zhang — are not to be found in the Russian Far East, but in southwestern China (SE Tibet, S Qinghai, N, NW and W Sichuan). O. ombrochares is included in O. coerulescens (see above). O. pycnostachya var. yuennanensis Beck ("yunnanen-sis") according to Harry Smith (1933) is also attributable to O. coerulescens s. str.
Acknowledgements
We would like to thank the scientists and the staff of the V. L. Komarov Botanical Institute of the Russian
Academy of Sciences (LE), in particular Irina Illa-rionova, Marina Legchenko, Denis Melnikov, Larisa Raenko, Lidia Ryazanova, Valeria Shvanova, Ivan Ta-tanov, Roman Ufimov and Peter Efimov. We would also like to thank the ladies and gentlemen of the Herbaria of Berlin (B) and Halle (HAL) for supporting our research. We would like to thank Kristin Meyer (Frankfurt at Oder) and Volker Otte (Görlitz) for the joint planning and implementation of the excursion to the Russian Far East in 2012. We would like to thank the team of the Botanical Garden-Institute of the Far Eastern Branch of the Russian Academy of Sciences (Vladivostok), namely Pavel Krestov, Vadim Bakalin, Natalia Labetskaya, and Natalya Fishchenko (Benevskoe) for the diverse support and warm welcome during the excursion. Thanks to Leslie Lewis (Chepstow, UK) for linguistic revision and Andreas Fleischmann for helpful hints.
Supplementary materials
Appendix 1. Herbarium specimens of Orobanche coerulescens, O. pycnostachya var. filicicola and O. pycnostachya var. pycnostachya f. amurensis, collected by S. Rätzel in the Russian Far East. https://www.binran. ru/files/journals/Novitates/2022_53/NSPV-53_11-Ratzel_Uhlich-Appendix_1.pdf
Appendix 2. Images of lectotypes of the names of some Orobanche taxa, designated in the article. https:// www.binran.ru/files/journals/Novitates/2022_53/ NSPV-53_11-Ratzel_Uhlich-Appendix_2.pdf
References
[Beck] Beck von Mannagetta G. 1890. Monographie der Gattung Orobanche // Biblioth. Bot. Bd. 4, H. 19. S. 1-275. [Beck] Beck-Mannagetta G. 1930. Orobanchaceae // Das Pflanzenreich. Regni vegetabilis conspectus / A. Engler (Hrsg.). Bd. IV.261 [H. 96]. Leipzig: W. Engelmann. S. 1-348.
Chang C. S., Kim H., Chang K. S. 2014. Provisional checklist of vascular plants of the Korea peninsula flora. Seoul, Korea: Designpost. 660 p. Chinese Virtual Herbarium. 2021.
https://www.cvh.ac.cn (Accessed 15.11.2021). Fedtschenko O. A., Fedtschenko B. A. 1913. Conspectus Florae Turkestanicae. Pt. 5. Yuryev: Typogr. K. Mattisena. 393 p. [In Russian] (Федченко О. А., Федченко Б. А. 1913. Перечень растений дико растущих в русском Туркестане. Ч. 5. Юрьев: Типогр. К. Маттисена. 393 с.). GBIF. 2022. Global Biodiversity Information Facility.
https://www.gbif.org (Accessed 24.10.2022). Hance H. F. 1873 [publ. 1872]. Notes on some plants from Northern China // J. Linn. Soc., Bot. Vol. 13. P. 74-94. https: //doi.org/10.1111/j. 1095-8339.1872.tb00089.x Herbarium Berolinense — Virtual Herbarium. 2022.
https://ww2.bgbm.org/herbarium/ (Accessed 24.10.2022).
Hsu Y.-P., Wang K.-Z. 1996. Orobanchaceae // Flora Sinensis in Area Tan-Yang. T. 4. Typis Ningxiae Popularis. P. 92100. [In Chinese].
Hyun J.-O, Lim Y., Shin H. 2003. Validation of Orobanche fi-licicola (Orobanchaceae) from Korea // Novon. Vol. 13, № 1. P. 64-67.
https://doi.org/10.2307/3393566
Korsun O. 2014. Vot takoi tsvetok [Here is such a flower]. [In Russian] (Корсун О. 2014. Вот такой цветок). https://olegkorsun.livejournal.com/183351.html (Accessed 22.10.2019).
Krylov P. N., Schischkin B. K., Sergievskaya L. P., Steinberg E. I., Sumnevich G. P. 1939. Flora Sibiriae occiden-talis. Iss. 10. Solanaceae — Dipsacaceae. Tomsk: Krasnoye Znamya. P. 2401-2627. [In Russian] (Крылов П. Н., Шишкин Б. К., Сергиевская Л. П., Штейнберг Е. И., Сумневич Г. П. 1939. Флора западной Сибири. Вып. 10. Solanaceae — Dipsacaceae. Томск: Красное Знамя. С. 2401-2627).
Kurbatsky V. I. 1996. Semeistvo Orobanchaceae — Zarazi-khovyye [Orobanchaceae — Broomrape family] // Flora Sibiriae. T. 12. Solanaceae — Lobeliaceae / A. V. Polozhij, G. A. Peschkova (eds.). Novosibirsk: Nauka. P. 93-99. [In Russian] (Курбатский В. И. 1996. Семейство Orobanchaceae — Заразиховые // Флора Сибири. Т. 12 / А. В. Положий, Г. А. Пешкова (ред.). Новосибирск: Наука. C. 93-99).
Lee H.-С., Chekar E.-K., Lim D.-O. 2011. The specific plant species and naturalized plants in the area of Naejangsan National Park, Korea // Korean J. Environm. Ecol. Vol. 25, iss. 3. P. 267-283.
Li X., Jang T.-S., Temsch E. M., Kato H., Takayama K., Schnee-weiss G. M. 2017. Molecular and karyological data confirm that the enigmatic genus Platypholis from Bonin-Islands (SE Japan) is phylogenetically nested within Orobanche (Orobanchaceae) // J. Pl. Res. Vol. 130, iss. 2. P. 273-280. https://doi.org/10.1007/s10265-016-0888-y
Maximowicz C. J. 1859. Primitiae Florae Amurensis. Versuch einer Flora des Amurlandes // Mém. Acad. Imp. Sci. St.-Pétersbourg Divers Savans. T. 9. P. [1]-504; 10 Taf.
Nakai T. 1952. A synoptical sketch of Korean flora // Bull. Natl. Sci. Mus. Tokyo. Vol. 31. P. 1-152.
Naturalis Biodiversity Center. 2022.
https://www.naturalis.nl (Accessed 24.10.2022).
Naejangsan National Park Service. 2018. Orobanche filici-cola Nakai (In Korean). https://www.knps.or.kr/front/ portal/photo/photoDtl.do?photoId=SSPM015220&page (Accessed 23.10.2022).
Novopokrovsky I. V., Tzvelev N. N. 1958. Sem. Zarazikhovyye — Orobanchaceae Lindl. [Fam. Broomrapes — Orobanchaceae Lindl.] // Flora URSS. T. 23 / B. K. Schischkin (ed.). Mos-qua; Leningrad: Ed. Acad. Sci. URSS. P. 19-117, 685-687. [In Russian] (Новопокровский И. В., Цвелёв Н. Н. 1958. Сем. Заразиховые — Orobanchaceae Lindl. // Флора СССР. Т. 23 / Б. К. Шишкин (ред.). М.; Л.: Изд-во Акад. наук СССР. С. 19-117, 685-687).
Piwowarczyk R., Schneider A. C., Góralski G., Kwolek D., Denysenko-Bennett M., Burda A., Ruraz K., Joachim-iak A. J., Sánchez Pedraja Ó. 2021. Phylogeny and his-
torical biogeography analysis support Caucasian and Mediterranean centres of origin of key holoparasitic Orobancheae (Orobanchaceae) lineages // PhytoKeys. Vol. 174. P. 165-194.
https: //doi.org/10.3897/phytokeys. 174.62524
Poljakov P. P. 1961. Rod Polyn — Artemisia L. [Genus Sagebrush — Artemisia L.] // Flora URSS. T. 26 / B. K. Schischkin, E. G. Bobrov (eds.). Mosqua; Leningrad: Ed. Acad. Sci. URSS. P. 425-631. [In Russian] (Поляков П. П. 1961. Род Полынь — Artemisia L. // Флора СССР. Т. 26 / Б. К. Шишкин, Е. Г. Бобров (ред.). М.; Л.: Изд-во Акад. наук СССР. С. 425-631).
Sánchez Pedraja Ó., Moreno Moral G., Carlón L., Piwowarczyk R., Laínz M., Schneeweiss G. M. 2022. Index of Orobanchaceae. https://www.farmalierganes.com/ 0trospdf/publica/0robanchaceae%20Index.htm (Accessed 20.10.2022).
Schischkin I. K. 1930. Materialy k flore basseina reki Imana (Dalnevostochnyi Krai, Khabaroskii Okrug) [Materials for the flora of the Imana River (Far Eastern Territory, Khabarovsk District)] // Zapiski Vladivostokskogo otdela Gosudarstvennogo Russkogo geograficheskogo obshchestva [Notes of the Vladivostok Department of the State Russian Geographical Society]. Vol. 5, iss. 2. P. 5-173. [In Russian] (Шишкин И. К. 1930. Материалы к флоре р. Имана (ДВК, Хабаровский округ) // Зап. Владивосток. отд. Госуд. Русск. геогр. общ. Т. 5, вып. 2. С. 5-173).
Schneeweiss G. M., Palomeque T., Colwell A. E., Weiss-Schnee-weiss H. 2004. Chromosome numbers and karyotype evolution in holoparasitic Orobanche (Orobanchaceae) and related genera // Amer. J. Bot. Vol. 91, iss. 3. P. 439-448. https://doi.org/10.3732/ajb.91.3.439
Smith H. 1933. Plantae Sinenses: 26. Orobanchaceae // Acta Horti Gothob. Vol. 8. P. 127-135.
Turland N. J., Wiersema J. H., Barrie F. R., Greuter W., Hawksworth D. L., Herendeen P. S., Knapp S., Kus-ber W.-H., Li D.-Z., Marhold K., May T. W., McNeill J., Monro A. M., Prado J., Price M. J., Smith G. F. (eds.). 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress, Shenzhen, China, July 2017. Glashütten: Koeltz Botanical Books. xxxviii + 254 p. (Regnum Vegetabile. Vol. 159). https://doi.org/10.12705/Code.2018
Tzvelev N. N. 1996. Semeistvo Zarazikhovyye — Orobanchaceae Vent. [Broomrape family — Orobanchaceae Vent.] // Plantae vasculares Orientis Extremi sovie-tici. T. 8 / S. S. Charkevicz (ed.). Saint Petersburg: Nau-ka. P. 249-251. [In Russian] (Цвелёв Н. Н. 1996. Сем. Заразиховые — Orobanchaceae Vent. // Сосудистые растения советского Дальнего Востока. Т. 8 / отв. ред. С. С. Харкевич. СПб.: Наука. С. 249-251).
Tzvelev N. N. 2006. Sem. [Fam.] Orobanchaceae Vent. // Plantae Asiae Centralis. Fasc. 15 / V. I. Grubov (ed.-in-chief). Moscow: KMK Sci. Press. P. 11-23. [In Russian] (Цвелёв Н. Н. 2006. Сем. Orobanchaceae Vent. // Растения Центральной Азии. Вып. 15 / В. И. Грубов (отв. ред.). М.: Т-во науч. изд. КМК. С. 11-23).
Uhlich H. (in prep.). Orobanchaceae (Orobanche, Pheli-panche) // Exkursionsflora für Österreich und die gesamten Ostalpen. 4. Ed. / M. A. Fischer, K. Oswald, W. Adler (eds.). Linz: Biologiezentrum der Oberösterreichischen Landesmuseen.
Vascular Plants Herbarium of the Komarov Botanical Institute RAS — Herbarium LE. 2022. https://en.herbariumle.ru/ (Accessed 23.10.2022).
Virtual Herbaria JACQ. 2022. https://www.jacq.org/ (Accessed 24.10.2022). Willdenow C. L. 1800. Species plantarum. T. 3, ps. 1. Berolini: G. C. Nauk. [ii] + 847 p. https: //doi.org/10.5962/bhl.title.37657 Zhang Z.-Y., Tzvelev N. N. 1998. Orobanchaceae // Flora of China. Vol. 18 / Z.-Y. Whu, P. H. Raven (eds). Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. P. 229-243.
