Noccaea venusta (Schischk.) German (2017: 297)

Özüdoğru, Bariş & German, Dmitry A., 2018, Taxonomic remarks on Pseudosempervivum (Brassicaceae), Phytotaxa 383 (1), pp. 103-110 : 104-107

Noccaea venusta (Schischk.) German (2017: 297) ≡ Cochlearia venusta Schischkin (1929: 459) ≡ Pseudosempervivum venustum (Schischk.) Pobedimova (1971: 195). Lectotype ( German 2017: 297):—[ TURKEY, B9 Ağrı]: Turkish

Armenia. Bayazet sandzhak. Dutakh – Burnubulak, steppe, [fl., fr. immat.]. 13 [26] May 1916. B. Schischkin s.n. (LE

01017936!; isolectotype ERE 0004298 !).

= Syrenopsis bornmuelleri Rechinger (1939: 264) ≡ Thlaspi bornmuelleri (Rech. f.) Hedge ( Davis et al. 1965: 184) ≡ Noccaea bornmuelleri (Rech. f.) Al-Shehbaz (2014: 34) . Lectotype ( Hedge 1965: 340, as “type”):—[ TURKEY, C9 Siirt]: Kurdistan: Parklandschaft auf dem Daharakol Dag, östl. vom Shat, 1700 m, [fl., fr. immat.]. 15 Juni [19]36. J. Frödin 264 (W 1938-0005888!; isolectotypes JE 00001515! [http://herbarium.univie.ac.at/database/detail.php?ID=114760], UPS V-089164 [http://herbarium.emg.umu.se/record. php?Genus= Syrenopsis &Page=1&AaccNr=V-089164&Ainst=UPS&Acoll=&Aid=56045620&nrRecords=1], photo in JE!).

= Cochlearia gurulkanii Yıldırımlı (2006: 3) ≡ Pseudosempervivum gurulkanii (Yıld.) Mutlu, Al-Shehbaz & Dönmez ( Al-Shehbaz et al. 2007: 329), syn. nov. Type:— TURKEY, C9 Şırnak: Foot of Namaz dağı, from yaylabaşı via limekiln to centrum, steppe, rocky quercetum, (Q. brantii Lindley) places, 1700–1900 m, 21 June 2005, [fr.]. Ş. Yıldırımlı 30169 (Hb. Yıldırımlı; isotype HUB!).

It was established just recently that the plant long known as Thlaspi bornmuelleri has first been described as Cochlearia venusta and correct name for relevant species under the current taxonomic concept is Noccaea venusta ( German 2017) . Further studies demonstrated that one more name, Pseudosempervivum gurulkanii , refers to the same species and should be added to its synonymy. Just like C. venusta ( Schischkin 1929) , C. gurulkanii was initially ( Yıldırımlı 2006) assigned to Cochlearia sect. Pseudosempervivum , and based on well-developed (to 1.5 mm long) styles – unique in species ever assigned to Pseudosempervivum – and similar distribution, their conspecificity or at least close affinity could be suggested. However, other features reported for C. gurulkanii , such as perennial habit, mandatory development of several stems, and strictly 1-seeded locules of silicles were discordant with the morphology of N. venusta . As evidenced by the studied material, report of such characters is a result of incomplete description and misinterpretation of observed phenomena. For example, the claimed perenniality is based on the presence of “fibrous petiolar collar” and ignorance of the fact that the plant which is clearly monocarpic has simple and rather short taproot typical for annuals and that remnants of petioles forming the “collar” belong to rosette leaves of current year which had completely withered by fruiting time (type gathering of C. gurulkanii is in the stage of predominantly mature fruits). Similarly, “many-stemmed” habit is clearly a result of abundant compensatory low branching as a consequence of damage of the main stem (like in isotype of N. venusta or, even more pronounced, in Ekim 7794 [GAZI], see Fig. 1 , B) that is otherwise well-developed and usually anyway much-branched, either from lower half or above, as in case of the isotype of C. gurulkanii ( Fig. 1 , A). Finally, the number of developed seeds (1–2) in 2-ovulate locules of the latter taxon ( Fig. 1 , C) fits well the range of (0)1 to usually 2 seeds in 2–4-ovulated locules observed in N. venusta . Hence, there is no way to distinguish P. gurulkanii from N. venusta based on the above-mentioned characters and otherwise the two are equally the same. In general, the species possesses quite distinctive morphology and can be hardily confused with any of its numerous congeners. Its distribution area is confined to E and SE Turkey (provinces Tunceli, Ağrı, Bingöl (near the border with Elazığ), Siirt, Şırnak, and Muş including border with Erzurum) within the grid squares B7, B8, B9, and C9 ( Schischkin 1929, Rechinger 1939, Hedge 1965, Adıgüzel & Ekim 1991, Yıldırımlı 2002, 2006, German 2017).

Pseudosempervivum sempervivum (Boiss. & Balansa) Pobedimova (1971: 195) ≡ Cochlearia sempervivum Boiss. & Balansa ( Boissier 1856: 28). Original locality designation: “Hab. in regione montanâ superiori montis Masmeneudagh cl. Balansa. Augusto jam fructiferum”. Lectotype (designated here by Al-Shehbaz):—[ TURKEY, C5 Niğde] Région montagneuse supérieure du Masmeneu-Dagh, à 25 lieues au SSO de Césarée. 8 août 1855, B. Balansa 439 (G-BOISS: G 00332260; isolectotypes: (BM 000582868 [http://plants.jstor.org/stable/viewer/10.5555/ al.ap.specimen.bm000582868], C!, G 00389743 [http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen. g00389743], GOET 002739! [http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.goet002739], JE 00001803! [https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.je00001803], JE 00001804! [https://plants.jstor.org/ stable/viewer/10.5555/al.ap.specimen.je00001804], K 000484388 [http://plants.jstor.org/stable/viewer/10.5555/al.ap. specimen.k000484388], K 000484389 [http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.k000484389], KW 000128007!, P 02272525 [http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.p02272525], P 02272526 [http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.p02272526], P 02272527 [http://plants.jstor.org/stable/ viewer/10.5555/al.ap.specimen.p02272527], P 02272528[http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen. p02272528], W 1889-11478!, W1889-76132!, WAG 0004283 [http://plants.jstor.org/stable/viewer/10.5555/al.ap. specimen.wag0004283]).

= Masmenia crassiuscula Meyer (1973: 468) ≡ Thlaspi crassiusculum (F.K. Mey.) Greuter & Burdet ( Greuter & Raus 1983: 95) ≡ Noccaea crassiuscula (F.K. Mey.) Al-Shehbaz (2014: 37) , syn. nov. Type:—[ TURKEY, C6 Adana] Osmaniye, Koyun meleten dag. Amanos dağları, ca. 1960 m, 9 VI 1968. Y. Akman 203 (G 00371946 [http://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen. g00371946]).

This species has been described based on the single specimen represented by a plant with torn apart upper part of the stem compensated by several branchlets with buds and only few first flowers ( Fig. 2 , A). Notwithstanding nearly complete absence of well-developed generative organs, the plant is peculiar for its robust habit, fleshy, numerous, and dense leaves, lower broadly spatulate and attenuate into a wide petiole, middle and upper sessile, broadly cordatereniform and semiamplexicaul, and is immediately recognizable as a member of Pseudosempervivum and, more specifically, as P. sempervivum , the species not only best fitting morphologically ( Fig. 2 , B–D) but also geographically. Furthermore, since long time this relationship is confirmed phylogenetically ( Koch & Mummenhoff 2001). It is very likely that the author of Masmenia crassiuscula came to the same conclusion regarding the identity of his species which can be assumed based on indirect reducing Masmenia Meyer (1973: 468) to synonymy of Pseudosempervivum proposed by him in subsequent publications ( Meyer 1991: 15, 2001: 8) but he never published it. As a result, with a single doubt by Davis et al. (1988) who mentioned M. crassiuscula [as Thlaspi crassiusculum ], along with a number of other Meyer’s species, as incertae sedis taxon, the species is hitherto treated as accepted in all respective publications and databases ( Greuter et al. 1986, Marhold 2011, Mutlu 2012, Al-Shehbaz 2014, Plants of the World online 2018, Tropicos.org, etc.). As evidenced herein, this concept cannot be followed anymore.

= Cochlearia amana Contandriopoulos & Quézel (1976: 418) ≡ Pseudosempervivum amanum (Contandr. & Quézel) Al-Shehbaz, Mutlu & Dönmez (2007: 329) , syn. nov. Type:—[ TURKEY, C6 Hatay] Amanus, 2000 m, sommets ophiolitiques à 15 km au S.-E. d’Osmanyie. P. Quézel et J. Contandriopoulos 73-434. 30 VI 1973 (Herbier de la Faculté des Sciences de Marseille Saint-Jérôme)” (MARS? AIX?).

Note. The type of C. amana is believed to be stored, according to the original publication, in MARS ( Davis et al. 1988, Al-Shehbaz et al. 2007, Al-Shehbaz 2014). However, as kindly reported by the curator (Bruno Vila, pers. comm. to D. A. G. by e-mail, March 30, 2017), it is absent from there and the only other option could be AIX where, unfortunately, it was also not found (Isabelle Chanaron, pers. comm. by e-mail to D. A. G., April 18, 2017). Hence, any conclusions on the identity of the species could be based on the analysis of protologue [with slight correction regarding sepal and petal size by Davis et al. (1988)], because no other gatherings are known [one specimen was reported by Yıldırımlı (2006) as C. sempervivum with the note “might be C. amana ”] and all efforts to collect P. amanum in the classical locality failed ( Reeves & Adıgüzel 2008).

By the present time, P. amanum is treated as accepted species in any relevant works ( Davis et al. 1988, Yıldırımlı 2006, Al-Shehbaz et al. 2007, Mutlu 2012, The Plant List 2013, GBIF 2017, Plants of the World online 2018, Tropicos. org, etc.). However, Yıldırımlı (l. c.: 8) admitted that “ C. amana might be a synonym of C. sempervivum ”, but no evidences proving this suggestion have been presented and formal synonymy was not established. According to Contandriopoulos and Quézel (1976) and Davis et al. (1998), C. amana is closest to C. sempervivum (which indeed follows from the description) but differs from it in a life form of perennial herb with branched rootstock [vs. biennial with simple root], narrower basal leaves (ca. 4 [vs. 2–2.5] times as long as wide) and pink-tipped sepals. Among three closely related species of Pseudosempervivum , P. aucheri ( Boissier 1842: 168) Pobedimova (1971: 194) , P. sempervivum , and P. sintenisii (Hausskn. ex Bornmüller 1936: 40) Pobedimova (1971: 195) , perennial habit is only peculiar for the latter species, otherwise more distant morphologically and geographically from P. amanum than the other two, both known as robust biennials. However, the specimens of P. aucheri ( Fig. 3 ), a species very close to (if not conspecific with; see below) P. sempervivum , were studied, e. g. those collected by F. Sorger in 14 km north of Sivas on 12 VII 1969, No. 69-49-70 (W 1992-8258!) and in 17 km south of Kangal on 2 VIII 1976, No. 76-17-15 (W 1992-8260!) clearly showing the underground structures ( Fig 3. C, D ) described by Contandriopoulos and Quézel, viz., woody rootstocks branched above and bearing numerous leaf rosettes. Stems (stout, much branched, ca. 40–50 cm high), which in both specimens are mounted separately, are typical for P. aucheri and cannot belong to P. sintenisii characterized by relatively slender and moderately branched stems to 20 cm tall and more north-eastern distribution. Based on this observation it can be admitted that normally biennial P. aucheri may under certain conditions (or it is not so uncommon?) grow as at least short-leaved perennial, and apparently the same phenomenon was observed by the authors of C. amana . Pseudosempervivum aucheri is also very helpful for demonstrating the polymorphism of shape of basal leaves which vary from broadly spatulate to linear (“linear-spatulate”) with the ratio length to width ranging from 1: 2 to 1:>10; a narrow-leaved form was described as P. diversifolium ( Schulz 1936: 462) Pobedimova (1971: 194) but it is usually ( Hedge 1965, Warwick et al. 2006, Mutlu 2012) considered a synonym of P. aucheri . Having in mind these facts it should not be surprising at all if the closest species would reveal more or less similar pattern of variation. Finally, pinkish sepals are observed sometimes on specimens of both P. aucheri and P. sempervivum . Therefore, in agreement with the above-mentioned assumption of Yıldırımlı (2006), we see no reason to keep treating P. amanum as an enigmatic endemic of Amanus and rather add it to synonymy of sympatric P. sempervivum .

It can be noted that relationship of P. sempervivum with P. aucheri needs to be clarified in view of quite unclear distinction among them. Most of the characters previously used as diagnostic, such as degree of development of auricles of stem leaves, morphology of fruit base and septum ( Boissier 1856, 1867) were not utilized subsequently ( Hedge 1965) as unreliable and currently the species are separated by the size of basal leaves (bigger and less toothed in P. sempervivum ) and style length (smaller in the latter species). However, these characters considerably interfere as well. Although generally more north-eastern P. aucheri indeed often develops relatively small, to 2.5 cm long, basal leaves, some specimens produce those 5.5–6 cm long (e.g., PH 00006403 [http://plants.jstor.org/stable/viewer/10.5555/al.ap. specimen.ph00006403]) which even exceeds the value (by 5 cm) usually reported for and observed in P. sempervivum . Style length in typical plants of the latter species is 0.2–0.3 mm (vs. 0.5–0.8, rarely to 1.1 in P. aucheri ), but in other specimens (e.g., Haussknecht’s gathering from Beritdagh [JE!, LE!] cited by Boissier (1867: 246)) some styles reach 0.6 mm long while even in the type gathering of C. aucheri the style length varies from 0.2 to 0.8 mm. Noteworthy, similar range of variation is observed in P. sintenisii . Thus, one could be tempted to merge P. sempervivum with P. aucheri but this does not agree with available, though quite scarce, ITS-based phylogenetic data ( Koch et al. 1999, Koch & Mummenhoff 2001, Özüdoğru 2018a) showing closer relationship of P. aucheri with morphologically more distant P. sintenisii rather than with habitually more similar P. sempervivum . The problem therefore stays open till a comprehensive morphological/phylogenetic study of the group based on sufficient sampling is performed.

Figures

FIGURE 1. Representative specimens of Noccaea venusta. A. Isotype of Pseudosempervivum gurulkanii with undamaged main stem (Yıldırımlı 30169, HUB!); B. “Many-stemmed form” of Noccaea venusta (Ekim 7794, GAZI!); C. Fruits of Noccaea venusta (Pseudosempervivum gurulkanii, isotype).

FIGURE 2. Representative specimens of Pseudosempervivum sempervivum with variously damaged stems. A. Type of Masmenia crassiuscula (G 00371946, http://www.ville-ge.ch/musinfo/bd/cjb/chg). B. Isolectotype of P. sempervivum (K 000484389, http:// specimens.kew.org/herbarium/K000484389). C. Özüdoğru 5006 & Özgişi (HUB). D. Özüdoğru 3610, Özgişi & Açıcı (HUB).

FIGURE 3. Representative specimens of Pseudosempervivum aucheri with brached rootstock. A, C. Sorger 69-49-70 (W 1992-8258!) B, D. Sorger 76-17-15 (W 1992-8260!)