19 - 1 - 25 - Al-Shehbaz New

A SYNOPSIS OF THE GENUS NOCCAEA (COLUTEOCARPEAE, BRASSICACEAE)
Ihsan A. Al-Shehbaz1
Abstract. The controversial limits of Thlaspi are discussed, and all of its segregates proposed or revised during the past four decades
are critically evaluated. As a result, the majority of specific and infraspecific names previously assigned to Thlaspi are currently placed
in Noccaea, a genus the limits of which are expanded to include almost all of Thlaspi segregates. A detailed description of Noccaea, a
synoptic account of its species, along with full bibliography and typification, and key to all 128 species are presented. The new names N.
germanii, N. iranica, and N. meyeri and the following 36 new combinations are proposed: N. abchasica, N. antitaurica, N. atlantica, N.
bornmuelleri; N. bovis, N. bulbosa, N. campylophylla, N. cappadocica, N. capricornuta, N. cilicica, N. cornuticarpa, N. crassiuscula,
N. dolichocarpa, N. eigii, N. elegans, N. huber-morathii, N. jaubertii, N. kurdica, N. lilacina, N. maassoumii, N. natolica, N. orbiculata,
N. oxyceras, N. perfoliata, N. platycarpa, N. pulvinata, N. rechingeri, N. rostrata, N. rosularis, N. stenocarpa, N. szowitsiana, N. trian-
gularis, N. umbellata, N. vesicaria, N. vesicaria subsp. boissieri, and N. zangezurica. The genus Coluteocarpus is reduced to synonymy
of Noccaea. Brossardia papyracea var. kotschyi, Coluteocarpus reticulatus var. boissieri, Hutschinsia trinervia, Iberidella andersonii,
Thlaspi affine, T. japonicum, T. kovatsii, T. szowitsianum, and Vesicaria reticulata are lectotypified.
Keywords: Brassicaceae, Coluteocarpeae, Cruciferae, Noccaea, Thlaspi
The genus Thlaspi L. was once considered among the to the species are not parallel, difficult to follow, rely a great
larger genera of Brassicaceae (Cruciferae) and included deal on vegetative characters and seed anatomy, and some
some 75 species distributed primarily in Eurasia but with of the terminal leads are quite descriptive whereas others are
fewer taxa in North and South America and northern Africa very brief.
(Al-Shehbaz, 1986; Appel & Al-Shehbaz, 2003). Over the Extensive molecular phylogenetic studies (e.g.,
past two and a half centuries, Thlaspi became the dumping Mummenhoff & Zunk, 1991; Koch et al., 1993;
site for any Brassicaceae plant with angustiseptate fruits Mummenhoff & Koch, 1994; Koch, 1995; Zunk et al., 1996;
(flattened at a right angle to the septum), two or more seeds Mummenhoff et al., 1997a, b; Koch et al., 1998a, b; Koch &
per locule, and simple trichomes or no indumentum. Indeed, Mummenhoff, 2001; Koch & Bernhardt, 2004; Koch & Al-
The International Plant Names Index (IPNI) lists under Shehbaz, 2004) have amply demonstrated that the generic
Thlaspi 478 names, of which over 80 are repeated records, circumscription of Thlaspi prior to Meyer’s (1973) work was
though numerous infraspecific names were not listed. clearly unnatural. These studies were the foundation for the
Therefore, the above figure provides only a rough estimate placement of Noccaea Moench in tribe Noccaeeae and the
of the magnitude of excessive naming of taxa in the complex. retention of Meyer’s (1973, 2001a) six species of Thlaspi
A radical departure from the conventional concept of s.str. in tribe Thlaspideae (Al-Shehbaz et al., 2006). Indeed,
Thlaspi was proposed by Meyer (1973, 1979) who relied recent floristic studies (e.g., Al-Shehbaz, 2010, 2012b; Al-
heavily on seed-coat anatomy to divide the genus into 12 Shehbaz & Watson, 2011) have recognized Noccaea as
genera and retained only six species in it. Influenced by distinct from Thlaspi. Subsequent family-wide molecular
tradition, several authors (e.g., Greuter and Raus, 1983; phylogenetic studies (e.g., Khosravi et al., 2009a; Warwick
Greuter et al., 1986; Al-Shehbaz, 1986; Artelari, 2002; Appel et al., 2010) showed that Noccidium F.K.Mey. belongs
& Al-Shehbaz, 2003) did not follow Meyer’s over splitting of in tribe Camelineae, and the remaining ten segregates of
the genus, and Thlaspi remained in a state of mess. Although Meyer (1973) grouped together with Noccaea to form a
seed-coat anatomy offers a few taxonomically useful features rather poorly resolved clade suggesting that they form a
(Meyer, 1991), it is impractical for the practicing taxonomist single genus. Following Al-Shehbaz (2012a), Meyer’s ten
and, therefore, its wide-scale utilization is limited at best. segregates are referred below to the synonymy of Noccaea,
In a series of painstaking and detailed studies, Meyer the earliest-published generic name in the group.
(2001a, b, 2003a–e, 2006a–e, 2010) revised almost all of The tribal name Noccaeeae was reduced by Al-Shehbaz
the Thlaspi segregates he established in 1973. His thorough (2012a) to the synonymy of the earlier-published Coluteo-
synonymy, complete bibliography, full typification of all carpeae (Dorofeyev, 2004), though a formal transfer of the type
taxa, detailed distributions, and excellent illustrations of species of Coluteocarpus Boiss., C. reticulatus Boiss. (=N.
leaves, floral parts, fruits, and seeds are absolutely valuable vesicaria (L.) Al-Shehbaz, to Noccaea was not done until the
for any study on the complex. Unfortunately, he did not deal present study. Coluteocarpus was shown to be nested within
with several New World and Himalayan taxa, and his keys Noccaea (Warwick et al., 2010, 2011; Firat et al., unpubl.).

I am profoundly grateful to Dr. Dmitry A. German for his critical review of the manuscript and valuable comments and help with the literature and typi-
fications. I also thank Dr. Gustavo A. Romero for his editorial advice. I am much indebted to the National Science Foundation for the grant DEB-1252905
that supported the present research. The directors, curators, and collection managers are thanked for loans or allowing studies of specimens from the major-
ity of herbaria cited.
1
Missouri Botanical Garden, 2345 Tower Grove Avenue, St. Louis, Missouri 63110, U.S.A.; ihsan.al-shehbaz@mobot.org
Harvard Papers in Botany, Vol. 19, No. 1, 2014, pp. 25–51.

© President and Fellows of Harvard College, 2014.
ISSN: 1938-2944, DOI: 10.3100/hpib.v19iss1.2014.n3, Published online: 30 June 2014
26 Harvard Papers in Botany Vol. 19, No. 1
Meyer (2001a) placed Thlaspi, all its 11 segregates, and 11 limit their usefulness for identification purposes.
additional genera in tribe Lepidieae subtribe Thlaspidinae as Convergence in the Thlaspi s.l. complex is widespread
delimited by Schulz (1936). However, such circumscription in almost every conceivable character, especially those
is totally artificial, as evidenced from several family-wide of the fruit (Mummenhoff et al., 1997b). The exception,
molecular phylogenetic studies (e.g., German et al., 2009; however, is the sole presence of longitudinally striate and
Khosravi et al., 2009a; Couvreur et al., 2010; Warwick et strongly reticulate seed sculpture in Thlaspi s.str. (Meyer,
al., 2010) and the phylogenetic tribal classification of the 1979, 2001a). Perhaps one of the remarkable examples of
Brassicaceae (Al-Shehbaz et al., 2006; Al-Shehbaz, 2012a). convergence is the evolution of horned fruits in species
Meyer’s (2001a) circumscription of the Thlaspidinae currently assigned to three tribes: T. ceratocarpum (Pall.)
included 23 genera currently assigned to seven tribes. Murray (Thlaspideae), Noccidium hastatulum (DC.)
These are: Aethionema W.T.Aiton in tribe Aethionemeae; F.K.Mey. (Camelineae), and several species of Noccaea
Bivonaea DC. in Bivonaeeae; Brossardia Boiss., Eunomia (Coluteocarpeae) that Meyer (2003b, 2003d) placed in
DC., Noccaea (including Atropatenia F.K.Mey., Callothlaspi Thlaspiceras and Kotschyella, respectively.
F.K.Mey., Kotschyella F.K.Mey., Masmenia F.K.Mey., The limits of Apterigia have recently been expanded by
Microthlaspi F.K.Mey., Neurotropis (DC.) F.K.Mey., Raparia Dorofeyev (2013) to include eight species placed by Meyer
F.K.Mey., Thlaspiceras F.K.Mey., and Vania F.K.Mey) in (1973, 2006e) and Al-Shehbaz (2012) in Noccaea, by Meyer
Coluteocarpeae; Ionopsidium Rchb. and Pastorea Todaro ex (1973, 2006c) in Atropatenia, and by Meyer (1973, 2006d)
Bertol. (=Cochlearia L.) in Cochlearieae; Microcardamum in Vania. The sole character used to unite these species is
O.E.Schulz and Hutchinsiella O.E.Schulz (both synonyms the lack of apical fruit wing. However, as indicated above,
of Hornungia Rchb.) in Descurainieae; Noccidium in fruit morphology is subject to substantial convergence, and
the Camelineae; Teesdalia W.T.Aiton in Iberideae; and molecular data do not support the recognition of more than
Pachyphragma (DC.) Rchb., Physalidium Fenzl. (=Graellsia one entity in the Noccaea complex.
Boiss.), and Thlaspi in Thlaspideae. Meyer (2010) later The present synopsis was undertaken with several goals
described Noccaeopsis F.K.Mey. based on having very short in mind, of which the principal one is providing an updated
siliques instead of silicles, but that division is artificial, and the enumeration of accepted taxa (including major synonymies
genus was referred by Al-Shehbaz (2012a) to the synonymy and typifications) for the World Flora Online currently in
of Noccaea. progress. All names are being made available online on
The infrageneric classification of Meyer’s (1973) BrassiBase (http://brassibase.cos.uni-heidelberg.de), the
segregates requires substantial re-evaluation of major Brassicaceae database housed and continuously
morphological characters in light of comprehensive undated at Heidelberg University, Germany (see Koch
molecular phylogenetic studies. He placed the six species et al., 2012; Kiefer et al., 2013). Furthermore, several
of Thlaspi s.str. in three sections and two series. Such segregates of Thlaspi were reduced to synonymy by Al-
classification can hardly be of any taxonomic value because Shehbaz (2012a), but none of their species has formally
the genus is rather small. By contrast, for the larger Noccaea, been transferred to Noccaea. These necessary nomeclatural
Mayer (2006e) recognized 67 species that he placed in four adjustements are presented below for all taxa except one.
sections and 11 series. Although some of the infrageneric The formal transfer of Thlaspi aghricum P.H.Davis & K.Tan
taxa include closely related species, his heavy reliance on to Noccaea is proposed elsewhere and has been accepted
vegetative features and the seeds epidermal cells to delimit for publication. However, the species is included in the
sections and series, especially in the key, would substantially following key, though it is not listed in the synopsis below.
Taxonomy
Noccaea Moench, Suppl. Methodus 89. 1802, non Kuntze, Brossardia Boiss., Ann. Sci. Nat., Bot. Sér. 2, 16: 380.
Revis. Gen. Pl. 1: 354. 1891; Thlaspi L. sect. Noccaea 1841. Type species: B. papyracea Boiss. (=Noccaea
(Moench) Griseb., Spic. Fl. Rumel. 1: 279. 1843. Type papyracea (Boiss.) Khosravi, Mumm. & Mohsenz.).
species: N. rotundifolia (L.) Moench (=Iberis rotundifolia L.). Callothlaspi F.K.Mey., Feddes Repert. 84: 457. 1973.
Name derivation: for Domenico Nocca, 1758–1841, Type species: C. lilacinum (Boiss. & Huet) F.K.Mey.
Italian clergyman, botanist, and director of the botanic (=Noccaea lilacina (Boiss. & Huet) Al-Shehbaz).
garden at Pavia.
Coluteocarpus Boiss., Ann. Sci. Nat., Bot. sér. 2,
Tribe: Coloteocarpeae V.I.Dorof., Turczaninowia 7(3):
17: 172. 1842. Syn. nov. Type species: C. reticulatus
51. 2004. = Tribe Noccaeeae Al-Shehbaz, Beilstein & E. A.
Boiss. (=Noccaea vesicaria (L.) Al-Shehbaz, based
Kellogg, Pl. Syst. Evol. 259: 112. 2006.
Synonyms: Apterigia Galushko, Novosti Sist. Vyssh. Rast. on Alyssum vesicaria L.).
6: 209. 1970; Thlaspi L. sect. Apterygium Ledeb., Cruciundula Raf., Fl. Tellur. 2: 100. 1837. Type
Fl. Ross. 1: 164. 1841, non Apterygia Baehni, Arch. species: C. minima (Ard.) Raf. (=Noccaea minima
Sci. 17: 79. 1964. Type species: A. pumila (Steven) (Ard.) F.K.Mey.).
Galushko (=Noccaea pumila (Steven) Steud.). Disynoma Raf., Fl. Tell. 3: 81. 1837. Types species: D.
Atropatenia F.K.Mey., Feddes Repert. 84: 467. carnea (Russell) Raf., nom. illeg., based on Thlaspi
1973. Type species: A. rostrata (N.Busch) F.K.Mey. carneum Russell, nom. illeg. (=Noccaea perfoliata
(=Noccaea rostrata (N.Busch) Al-Shehbaz). (L.) Al-Shehbaz).
2014 AL-SHEHBAZ, A SYNOPSIS OF THE GENUS NOCCAEA 27
Iberidella (DC.) Boiss., Ann. Sci. Nat., Bot. Sér. 2, 17: Herbs, perennial with simple or branched caudices or
188. 1842; Hutchinsia W.T.Aiton sect. Iberidella DC., stolons, sometimes biennial, rarely annual, often glaucous.
Syst. Nat. 2: 385. 1821; Thlaspi L. sect. Iberidella Trichomes absent or rarely as short, simple papillae.
(DC.) Janch., Österr. Bot. Z. 58: 207. 1908. Lectotype Multicellular glands absent. Stems erect to ascending or
species designated by Rollins (in Farr et al.: 865. decumbent, simple or branched basally and/or above, rarely
1979, as type): I. trinervia (DC.) Boiss. (=Noccaea woody at base. Basal leaves petiolate, rosulate, simple, entire,
trinervia (DC.) Steud.) denticulate, or dentate; cauline leaves sessile, auriculate,
Kotschyella F.K.Mey., Feddes Repert. 84: 457. sagittate, or amplexicaul or rarely non-auriculate at base,
1973. Type species: K. stenocarpa (Boiss.) F.K.Mey. entire or dentate. Racemes many flowered, ebracteate,
(=Noccaea stenocarpa (Boiss.) Al-Shehbaz). corymbose, elongated considerably or congested in fruit;
Lagowskia Trautv., Bull. Cl. Phys.-Math. Acad. Imp. rachis straight; fruiting pedicels horizontal to divaricate,
Sci. Saint-Pétersbourg ser. 2, 16: 321. 1858. Type rarely ascending or descending, persistent. Sepals oblong,
species: L. pysocarpa Trautv. (=Noccaea vesicaria ovate, or obovate, free, caduceus or very rarely persistent,
(L.) Al-Shehbaz). erect, glabrous, base of lateral pair not saccate; petals white,
Masmenia F.K.Mey., Feddes Repert. 84: 468. 1973. pink, lavender, rose, violet, or purple, usually longer than
Type species: M. rosularis (Boiss. & Balansa) F.K.Mey. sepals; blade obovate, spatulate, oblanceolate, oblong,
(=Noccaea rosularis (Boiss. & Balansa) Al-Shehbaz). or rarely broadly linear, apex obtuse; claw obscurely
Microthlaspi F.K.Mey., Feddes Repert. 84: 452. differentiated from blade, shorter than sepals, glabrous,
1973. Type species: M. perfoliatum (L.) F.K.Mey. unappendaged, entire; stamens 6, erect, tetradynamous or
(=Noccaea perfoliata (L.) Al-Shehbaz, based on subequal; filaments wingless, unappendaged, glabrous,
Thlaspi perfoliatum L.). free; anthers ovate or oblong, obtuse or rarely apiculate at
Neurotropis (DC.) F.K.Mey., Feddes Repert. 84: 467. apex; nectar glands 2, lateral, subtending bases of lateral
1973; Thlaspi L. sect. Neurotropis DC., Syst. Nat. stamens, or 4 and 1 on each side of lateral stamen, median
2: 377. 1821; Thlaspi subgen. Neurotropis (DC.) glands absent; ovules (1 or)2–12 per locule; placentation
Peterm., Deutschl. Fl. 48. 1849. Lectotype species parietal. Fruit dehiscent capsular silicles or rarely siliques,
designated by Meyer (1973: 452): Neurotropis obcordate, obovate, obdeltoid, elliptic, ovate, oblong, or
orbiculata (Steven ex DC.) F.K.Mey. (=Noccaea rarely linear, flattened and usually strongly angustiseptate,
orbiculata (Steven ex DC.) Al-Shehbaz). or nearly subquadrangular, uninflated or very rarely inflated,
Noccaeopsis F.K.Mey., Haussknechtia 12: 15. 2010. unsegmented; valves papery, obscurely to prominently
Type species: N. kamtschatica (Karav.) F.K.Mey. veined, glabrous, strongly keeled, smooth, winged or
(=Noccaea kamtschatica (Karav.) Czerep.). wingless apically and sometimes also around margin,
Raparia F.K.Mey., Feddes Repert. 84: 467. 1973. Type unappendaged; gynophore obsolete; replum rounded,
species: R. bulbosa (Spruner) F.K.Mey. (=Noccaea visible; septum complete, membranous, not veined; style
bulbosa (Spruner) Al-Shehbaz). distinct, included in, subequal, or exserted from and much
Syrenopsis Jaub. & Spach, Ill. Pl. Orient. 1: 6. 1842. exceeding apical notch of fruit; stigma capitate, entire,
Type species: S. stylosa Jaub. & Spach (=Noccaea unappendaged. Seeds uniseriate, wingless, oblong to ovate,
jaubertii (Hedge) Al-Shehbaz). plump or slightly compressed; seed coat minutely reticulate
Thlaspiceras F.K.Mey., Feddes Repert. 84: 467. to nearly smooth, not mucilaginous when wetted; cotyledons
1973. Type species: T. oxyceras (Boiss.) F.K.Mey. accumbent. x = 7.
(=Noccaea oxyceras (Boiss.) Al-Shehbaz). A genus of 128 species distributed primarily in Europe
Vania F.K.Mey., Feddes Repert. 84: 467. 1973. and Asia, with one species each in N. Africa, Patagonia
Type species: V. campylophylla F.K.Mey (=Noccaea (Argentina, Chile), and Mexico and three in the United
campylophylla (F.K.Mey.) Al-Shehbaz). States.
Key to the species of Noccaea
Mature plants are needed for the reliable identification of species, and fruit characters, seed or ovule number, and petals
features are essential. Due to its high variability, vegetative morphology is less reliable and, therefore, used only to a limited
degree in the following key.
1a. Annuals; non-flowering shoots absent by fruiting time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1b. Perennials with caudices and remains of basal rosettes, sometimes biennials; non-flowering shoots usually present by fruiting time. . . . . . . . 10
2a. Fruit 8–17 mm wide; fruit wings prominently reticulate veined, 3–4 mm wide at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2b. Fruit 2–7 mm wide; fruit wings not veined, to 2 mm wide at apex, or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3a. Petals 1.5–2 mm; seeds 1.4–1.8 mm; style obsolete; fruiting raceme dense . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. platycarpa
3b. Petals 2.2–2.8 mm; seeds 2.1–2.3 mm; style 0.1–0.4 mm; fruiting raceme lose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4a. Ovules/seeds 7–9 per locule; style 0.2–0.5 mm; plants stout, branched. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. orbiculata
4b. Ovules/seeds 4–6 per locule; style 0.1–0.2 mm; plants slender, unbranched. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. szowitsiana
5a. Fruits oblong or elliptic, wingless; style 2.5–3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5b. Fruits obcordate, winged distally; style absent or 0.1–1(–1.5) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Key to the species of Noccaea cont.

6a. Fruiting racemes lax; fruiting pedicels divaricate, 9–14 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. bornmuelleri
6b. Fruiting raceme dense; fruiting pedicels erect-ascending 5–10 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. jaubertii
7a. Style exserted from apical notch of fruit, rarely equal to it; petals unequal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7b. Style well included in apical notch of fruit; petals equal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8a. Apical fruit notch absent or to 0.1 mm deep; ovules/seeds 2–4 per locule; large petals 1–1.5 mm wide . . . . . . . . . . . . . . . . . . . . . . . N. annua
8b. Apical fruit notch 0.2–1 mm deep; ovules/seeds 3–5 per locule; large petals 1.4–2.2 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. natolica
9a. Petals 2–4.7 mm; ovules/seeds 2–4 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. perfoliata
9b. Petals 2–2.5 mm; ovules/seeds 4–6 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. umbellata
10a. Leaves ciliate or coarsely to minutely papillate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10b. Leaves glabrous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
11a. Leaves auriculate; style 1.6–2.5(–3.5) mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. papillosa
11b. Leaves cuneate at base; style ca. 0.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12a. Petals white; fruit obovate-obcordate; style subequaling the ca. 0.5 mm apical fruit notch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. iberidea
12b. Petals pink, rose, or lavender; fruit ovate to elliptic; style exserted from the obsolete apical fruit notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13a. Leaves linear, ciliate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. caespitosa
13b. Leaves orbicular to broadly obovate, minutely papillate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. oppositifolia
14a. Cauline leaves cuneate at base, neither auriculate nor amplexicaul. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14b. At least some cauline leaves distinctly auriculate or amplexicaul at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
15a. Fruits inflated, bladdery, 16–21 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. vesicaria
15b. Fruits flattened, not bladdery, 1–6 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16a. Fruits 1–1.5 mm wide, pyriform-oblong, minutely horned apically; stems woody at base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. iranica
16b. Fruits 2.2–6 mm wide, ovate, obovate, or oblong, not horned apically; stems herbaceous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17a. Fruits 9–20 mm; style 3.2–4.6 mm; ovules/seeds 7–12 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rostrata
17b. Fruits 5–8 mm; style 1–2.1 mm; ovules/seeds 1–6(–8) per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18a. Ovules/seeds 4–8 per locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18b. Ovules/seeds 1 or 2(or 3) per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
19a. Fruits broadly obovate, 5–6 mm wide; style 1.3–2.1 mm; fruiting pedicels 2.3–5 mm; ovules/seeds 4(or 5) per locule. . . . . . . . . N. abchasica
19b. Fruits ovate-oblong, 3.7–3.9 mm wide; style 1–1.2 mm; fruiting pedicels 2–2.5 mm; ovules/seeds 6–8 per locule. . . . . . . . . . . N. zangezurica
20a. Cauline leaves orbicular to obovate-orbicular; fruit apically notched; petals pink; Caucasus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. germanii
20b. Cauline leaves oblong to oblong-linear; fruits not apically notched; petals white; Iraq, Turkey, Greece. . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21a. Fruits 2.2–3.5 mm wide; petals equal, 1.5–2.3 mm wide; Greece . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. epirota
21b. Fruits 3–5 mm wide; petals unequal, 1–1.4 mm wide; Iraq, Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. kurdica
22a. Fruit apex horned. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22b. Fruit apex not horned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
23a. Perennials, usually with woody base and leaf remains of previous seasons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
23b. Biennials, without woody base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
24a. Style exserted from apical notch, 1.8–3 mm; petals 6–7.7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cilicica
24b. Style included or equaling apical notch, 0.8–1.2 mm; petals 2.5–4 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25a. Ovules/seeds 2–4 per locule; petals 2.5–3 mm; Pakistan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cornuticarpa
25b. Ovules/seeds 6–8 per locule; petals 3.5–4 mm; Iran, Turkmenistan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. stenocarpa
26a. Fruits narrowly oblong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
26b. Fruits ovoid- to obovoid-obdeltoid or obdeltoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
27a. Fruits 5–6 mm, horn 0.4–1 mm; petals 4–4.8 × 1.2–17 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cappadocica
27b. Fruits 7.3–10 mm, horn 1.4–2.5 mm; petals 2.5–3.5 × 0.8–1.1 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28a. Seeds 1.7–1.8 mm; fruit horn 1.5–2.5 mm; petals 2.5–2.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. bovis
28b. Seeds 2.7–2.8 mm; fruit horn 1.2–1.5 mm; petals 3–3.7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. meyeri
29a. Style exserted; fruit apex truncate, minutely horned, without apical notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
29b. Style included; fruit apex with distinct horns 1.1–3.7 mm, apical notch 1.1–2 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
30a. Petals 2.8–3.5 × 0.8–1 mm; style 1–1.3 mm; fruits obovoid-obdeltoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rechingeri
30b. Petals 3.5–5 × 1.4–1.8 mm; style 1.3–1.8 mm; fruits obdeltoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. triangularis
31a. Fruits narrowly obdeltoid, horn 3.3–3.7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. capricornuta
31b. Fruits obovate- or ovate-obdeltoid, horn 1.1–2.3 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32a. Seeds tuberculate; style 1.2–2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. huber-morathii
32b. Seeds smooth; style 0.9–1.2 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. oxyceras
33a. Petals purple, violet, mauve, pink, red, or lavender, rarely color-tinged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
33b. Petals white or creamy white (color unknown in N. antitaurica, N. dolichcarpa, N. edinensium, and N. zaffranii). . . . . . . . . . . . . . . . . . . . 58
34a. Fruits (16–)30–35 × (13–)30–40 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. papyracea
34b. Fruits 2.5–12 (–16) × 1.5–9(–12) mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35a. Style included in apical fruit notch or subequaling it. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
35b. Style exserted from apical fruit notch, or fruit not notched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

36a. Perennials with a bulbous base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
36b. Biennials or perennials without bulbous base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
37a. Fruits 10–12 mm wide; Algeria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. atlantica
37b. Fruits 4.5–8.5 mm wide; Greece. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. bulbosa
38a. Petals unequal, 7–9 mm; style 1.5–1.8 mm; ovules/seeds 1 or 2 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. aghrica
38b. Petals equal, 2.3–4.2 mm; style 0.3–1.2 mm; ovules/seeds (2 or)3–6 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39a. Basal leaves dentate; fruits 7.5–13 mm; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. violascens
39b. Basal leaves entire or repand; fruits 5–9(–10) mm; Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40a. Anthers bluish purple; style (0.5–)0.8–1.2 mm; ovules/seeds 4–6 per locule; C, W, N Europe. . . . . . . . . . . . . . . . . . . . . . . . . N. caerulescens
40b. Anthers yellow; style 0.3(–0.5) mm; ovule/seed 2–4 per locule; Cyprus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cypria
41a. Fruits oblong to elliptic, rarely ovate, widest at middle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
41b. Fruits obcordate to obovate, widest above middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
42a. Fruits usually ovate; ovules/seeds 2 per locule; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
42b. Fruits oblong to elliptic; ovules/seeds (3 or)4–6(–9) per locule; W Europe, Himalayas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
43a. Style 0.6–1 mm; fruits 3–3.6 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rubescens
43b. Style 1.2–2 mm; fruits 2.6–3 mm wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. sintenisii
44a. Petals red-violet or purple, 6.5–10 mm; W Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
44b. Petals pale lavender, 3–6(–7) mm; Himalayas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
45a. Fruits (2.5–)3–4(–4.5) mm wide; style (2–)2.5–4.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. corymbosa
45b. Fruits 2–2.7 mm wide; style 0.8–2.5(–3) mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. limosellifolia
46a. Fruits (8–)10–16 mm, apically notched; ovules/seeds 7–9 per locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. yunnanensis
46b. Fruits 5–10 mm, apically not notched; ovules/seeds 3–5 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
47a. Style 0.7–1.5 mm; fruits elliptic, 3–4 mm wide; petals (5–)6–7 × 2.5–3.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. andersonii
47b. Style 3–4 mm; fruits narrowly oblong, 1.5–2 mm wide; petals 4.5–5 × 1.5–2 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. nepalensis
48a. Fruits apically winged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
48b. Fruits apically wingless. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
49a. Petals unequal; ovules/seeds 1 or 2 per locule; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. aghrica
49b. Petals equal; ovules/seeds (1–)3–6 per locule; Europe, U.S.A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
50a. Style 1.2–4.2 mm; petals 4.5–13 mm; U.S.A.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. fendleri
50b. Style 0.5–1.2 mm; petals 2.2–4.8 mm; Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
51a. Raceme (10–)15–90-flowered, distinctly elongated in fruit; stems to 30 cm tall or more. . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. caerulescens
51b. Raceme 4–10-flowered, not or hardly elongated in fruit; stems 1–2.5(–4) cm tall. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. microphylla
52a. Style 0.7–1.3(–1.5) mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
52b. Style (1.5–)2–5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
53a. Fruits oblong-ovate, 2–2.5 mm wide, widest at or below middle; Iran, Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. valerianoides
53b. Fruits oblong-obovate, (2.5–)2.7–4.5 mm wide, widest above middle; Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
54a. Fruits 4.5–6.5 × (2.5–)2.7–3.5 mm; ovules/seeds 2–5 per locule; petals 5–6 × 2.2–2.6 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cepaeifolia
54b. Fruits 6–12 × 3.5–4.5 mm; ovules/seeds 1–3 per locule; petals 6–7.7 × 2.5–3.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rotundifolia
55a. Fruits prominently veined; Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. lilacina
55b. Fruits not or obscurely veined; Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
56a. Seeds 2.3–3 × 1.3–2 mm; petals 6.5–9.5 mm; Albania, Bulgaria, Macedonia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. bellidifolia
56b. Seeds 1.4–2.1(–2.2) × 1–1.4 mm; petals 4–6 mm; Italy, Spain. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
57a. Style 1.5–3(–3.5) mm; ovules/seeds 1 or 2 per locule; seeds 1.4–1.8 mm; Spain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. nevadensis
57b. Style (3–)3.5–5 mm; ovules/seeds 4 or 5 per locule; seeds 2–2.2 mm; Italy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. stilosa
58a. Fruits linear, oblong, elliptic, ovate, or rarely suborbicular, widest at or below middle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
58b. Fruits narrowly to broadly obovate, obcordate, or obdeltoid, widest above middle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
59a. Fruits linear to oblong-linear siliques, 3.5–10 times longer than wide; ovules/seeds (4 or)5–9 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . 60
59b. Fruits oblong, elliptic, ovate, or suborbicular silicles, 1–2.8 times longer than wide; ovules/seeds 1–4(–6) per locule. . . . . . . . . . . . . . . . . 62
60a. Fruits 3–5 mm wide; style (1.2–)1.5–2.4 mm; fruiting pedicels 7–13 mm; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. dolichocarpa
60b. Fruits 1.5–2.5 mm wide; style 0.5–0.6(–1) mm; fruiting pedicels 4–7 mm; Pakistan, Russia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
61a. Petals 2.5–3.5 mm; fruiting pedicels 4–5 mm; fruit 1.8–2.5 mm wide; Russia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. kamtschatica
61b. Petals 4–5 mm; fruiting pedicels 6–7 mm; fruit 1.2–1.5 mm wide; Pakistan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. swatensis
62a. Fruit suborbicular, cochleate, as long as wide; basal leaves coarsely dentate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. crassiuscula
62b. Fruits oblong, elliptic, or ovate, not cochleate, longer than wide; basal leaves if present entire, repand, or rarely dentate. . . . . . . . . . . . . . . 63
63a. Style included or rarely subequaling apical fruit notch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
63b. Style exserted well beyond apical fruit notch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
64a. Fruits contorted; fruiting pedicels spreading to descending; Bhutan, India, Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cochlearioides
64b. Fruits not contorted; fruiting pedicels ascending to divaricate; elsewhere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
65a. Perennials; style 0.5–1.5(–2.5) mm; fruits oblong; China, Mongolia, Russia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. thlaspidioides
65b. Biennials; style absent or to 0.3 mm; fruits elliptic or ovate; Turkey and S Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

66a. Fruits elliptic, 5.5–11 mm; ovules/seeds 3 or 4 per locule; petals 2–3.2 × 0.8–1 mm; fruiting pedicels 2–3.5 mm; S Europe
(Albania, France, Greece, Italy) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. brevistyla
66b. Fruits ovate, 5.5–6 mm; ovules/seeds 2 per locule; petals 6–7 × 2.5–3 mm; fruiting pedicels 7–8 mm; Turkey. . . . . . . . . . . . . . . N. rosularis
67a. Plants with woody lower stems. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
67b. Plants herbaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
68a. Sepals persistent; fruits 3–3.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. apterocarpa
68b. Sepals caducous; fruits 5–9 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
69a. Leaves 1-veined abaxially; petals ca. 5 mm; fruits elliptic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. maassoumii
69b. Leaves strongly 3-veined abaxially; petals 6–10 mm; fruits oblong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. trinervia
70a. Fruits elliptic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
70b. Fruits oblong to ovate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
71a. Fruits (7–)8–11 mm, usually apically winged; style 1.5–2.4 mm; U.S.A. (California) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. fendleri
71b. Fruits 5–8 mm, wingless; style 0.7–1.5 mm; Himalayas, Caucasus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
72a. Ovules/seeds 3–5 per locule; fruits 5–7 mm; Himalayas (Bhutan, China, India, Nepal, Pakistan). . . . . . . . . . . . . . . . . . . . . . . . N. andersonii
72b. Ovules/seeds 2 or 3 per locule; fruits 7–8 mm; Caucasus (Azerbaijan, Georgia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. pumila
73a. Biennials, rarely perennials; fruits 7–11 mm, apex truncate; ovules/seeds 4–6 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. eigii
73b. Perennials; fruits 4.4–7.5(–9) mm, apex cuneate or retuse; ovules/seeds 2 or 3(or 4) per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
74a. Plants pulvinate; anthers apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
74b. Plants not pulvinate; anthers not apiculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
75a. Fruits 6–7 mm, apically not notched; petals ca. 5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. campylophylla
75b. Fruits 4.4–5 mm, apically notched; petals 3–3.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. pulvinata
76a. Petals lingulate, 4.7–5.3 × 1–1.2 mm; Afghanistan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. gardeziana
76b. Petals spatulate, 4.5–7.3 × (1–)1.5–2.5 mm; Greece, Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
77a. Ovules/seeds 1 or 2 per locule; petals 1.5–2.5 mm wide; Greece. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. epirota
77b. Ovules/seeds 4 per locule; petals 1–1.8 mm wide; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. phrygia
78a. Style included in or subequaling apical fruit notch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
78b. Style exserted well beyond apical fruit notch, or apical notch absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
79a. Leaves fleshy, basal broadly obovate, subsessile; cauline leaves suborbicular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. zaffranii
79b. Leaves not fleshy, basal oblanceolate to obovate, long petiolate; cauline leaves ovate to oblong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
80a. Fruits 3–4 × ca. 1.5 mm, wing 0.3–0.6 mm wide apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. wendelboi
80b. Fruits (4–)5–13 × 2.2–7.1 mm, wing (0.5–)0.7–2.5 mm wide apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
81a. Ovules/seeds 7–9 per locule; Caucasus, Iran, NE Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. tatianae
81b. Ovules/seeds 1–6 per locule; S Turkey, Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
82a. Anthers blue, purple, or violet. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
82b. Anthers yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
83a. Petals about as long as sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. brachypetala
83b. Petals distinctly longer than sepals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
84a. Fruits as long as broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. occitanica
84b. Fruits distinctly longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
85a. Fruiting raceme 0.5–1.5 cm, not or hardly elongated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
85b. Fruiting raceme longer, distinctly elongated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
86a. Fruiting pedicels 2–3.5 mm; style 0.5–0.6 mm; fruit apex retuse. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cretica
86b. Fruiting pedicels 3–9 mm; style 0.6–1.2 mm; fruit apex subtruncate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. microphylla
87a. Fruits obovate; style 1–1.8 mm; sepals 2.3–2.6 mm; S Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. amani
87b. Fruits obcordate or rarely narrowly so; style 0.5–1(–1.2) mm; sepals 1–1.9(–2.2) mm; Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
88a. Ovules/seeds 4–6 per locule; petals 2.5–4.5 mm; fruits 6–9 × 3.5–7 mm; widespread in Europe including France. . . . . . . . . N. caerulescens
88b. Ovules/seeds 4 per locule; petals 2–3 mm; fruits 4–6.5 × 2.2–4.8 mm; France. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
89a. Fruits 4–6.5 x 3.7–4.8 mm; seeds 1.6–1.9 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. arenaria
89b. Fruits 4–5 x 2.2–3.2(–3.8) mm; seeds 1.3–1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. firmiensis
90a. Fruits 4.2–5.4 × 2.5–2.8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. eburneosa
90b. Fruits (5–)5.5–10(–13) × (2.5–)3–7.1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
91a. Fruits oblong-obovate; petals 5–7(–8) × (1.5–)2–3 mm; sepals 2.3–3 mm; China, Mongolia, Russia . . . . . . . . . . . . . . . . . . N. thlaspidioides
91b. Fruits broadly obovate to obcordate; petals 1.5–4.5 × 0.5–1.8(–2.1) mm; sepals 1–2.3 mm; elsewhere in Asia and Europe. . . . . . . . . . . . . 92
92a. Ovules/seeds 2–4 per locule; seeds 2.1–2.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
92b. Ovules/seeds (4 or)5 or 6 per locule; seeds 1.3–1.7(–2) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
93a. Fruits 3–6 mm wide; style 0.6–1.3 mm; petals 4.3–5.3 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. libanotica
93b. Fruits 4.5–7.1 mm wide; style 0.3–0.5 mm; petals 2.8–2.9 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. microstyla
94a. Petals 3.5–4.5 mm; ovules/seeds 6 per locule; fruits 2.5–3 mm wide, apical notch 2–3 mm deep; Afghanistan, Pakistan . . . . . . N. griffithiana
94b. Petals 1.5–3.5(–3.8) mm; ovules/seeds (4 or)5(or 6) per locule; fruit 3–6 mm wide, apical notch 0.3–1.3(–1.8) mm deep;
Europe, Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

95a. Biennials; fruits obcordate; fruiting raceme distinctly elongated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
95b. Perennials; fruits obovate; fruiting raceme not or hardly elongated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
96a. Petals 3–4 mm; sepals 2–2.3 mm; seeds 1.5–1.6 mm; Romania. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. banatica
96b. Petals 1.5–3 mm; sepals 1–1.8 mm; seeds 1.7–2 mm; Austria, Italy, Switzerland. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. salisii
97a. Basal leaves entire; fruits 5–7 mm; Montenegro, Romania . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. dacica
97b. Basal leaves denticulate; fruits 6.5–8.5 mm; Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. densiflora
98a. Fruits apex obtuse to acute or truncate to subtruncate, notch absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
98b. Fruit apex retuse to emarginate, notch distinct. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
99a. Fruit valves prominently veined; seeds/ovules 8–10 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. antitaurica
99b. Fruit valves not or obscurely veined; seeds/ovules 1–6 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
100a. Petals 2–2.8(–3) mm; sepals 0.8–1.8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
100b. Petals (3–)4–7.6(–13) mm; sepals (1.6–)1.8–4(–5.3) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
101a. Style 1–1.3 mm; fruit 7–8 mm; fruiting pedicels 6–8 mm; Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. japonica
101b. Style 0.2–0.5(–0.6) mm; fruit 3.5–7 mm; fruiting pedicels 2–5.5 mm; U.S.A.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. parviflora
102a. Fruits apically winged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
102b. Fruits apically wingless. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
103a. Ovules/seeds 1 or 2 per locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. epirota
103b. Ovules/seeds 4–6 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
104a. Fruits 2.5–3 mm wide, wing 0.2–0.7 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. phrygia
104b. Fruits 3.8–5.8 mm wide, wing 1–1.6 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
105a. Fruits obovate, 5.8–8 × 3.8–4 mm; petals 3–4.4 mm; sepals 1.7–2.4 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. sarmatica
105b. Fruits obdeltoid, 7.5–9.2 × 3.8–5.8 mm; petals 4.7–7.6 mm; sepals 2–4.4 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. versicolor
106a. Fruits narrowly obovate, about 3 times longer than broad; petals 3–5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
106b. Fruits broadly obovate to obcordate, up to 2 times longer than broad; petals 4–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
107a. Style 0.3–1 mm; petals 2–2.5 mm wide; sepals 1.5–2.3 mm; Alaska and Canada. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. arctica
107b. Style 1–2 mm; petals 1–2 mm wide; sepals 2–3 mm; Iran. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. tenuis
108a. Fruiting pedicels 9–10 mm; fruits about as long as wide; seeds 2–2.3 mm; Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. elegans
108b. Fruiting pedicels 2.5–7 mm; fruits about twice longer than wide; seeds 1.1–1.8 mm; Spain, U.S.A.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
109a. Fruits 3–4.5 mm wide; ovules/seeds 3–5 per locule; style 0.8–2 mm; U.S.A.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. fendleri
109b. Fruits 2.3–3.3 mm wide; ovules/seeds 2 per locule; style 1.5–3 mm; Spain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. nevadensis
110a. Anthers blue, purple, or violet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
110b. Anthers yellow (not known in N. edinensium). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
111a. Petals 0.8–2.5 mm, about as long as sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. brachypetala
111b. Petals (2–)2.5–8 mm, distinctly longer than sepals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
112a. Style (1.2–)1.5–2.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
112b. Style 0.5–1(–1.2) mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
113a. Petals 5–8 mm; sepals 2–3.8 mm; fruiting pedicels 5–10 mm; Greece. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. greaeca
113b. Petals 3–4 mm; sepals 1.5–2.3 mm; fruiting pedicels 2.5–7 mm; France, Italy, Switzerland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. virens
114a. Fruiting racemes 0.5–1.5 cm, not or hardly elongated; fruits obovate; Greece. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
114b. Fruiting racemes longer, distinctly elongated; fruits obcordate; C, W, and N Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
115a. Fruiting pedicels 2–3.5 mm; style 0.5–0.6 mm; fruit apex retuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cretica
115b. Fruiting pedicels 3–9 mm; style 0.6–1.2 mm; fruit apex subtruncate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. microphylla
116a. Ovules/seeds 4–6 per locule; petals 2.5–4.5 mm; fruits 6–9 × 3.5–7 mm; widespread in Europe including France. . . . . . . . . N. caerulescens
116b. Ovules/seeds 4 per locule; petals 2–3 mm; fruits 4–5 × 2.2–3.8 mm; France . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. firmiensis
117a. Biennials. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
117b. Perennials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
118a. Petals 1.5–3.1 mm; sepals 1.2–1.8 mm; fruiting pedicels 2–6 mm; Austria, Italy, Switzerland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. salisii
118b. Petals 5–7 mm; sepals 2–3 mm; fruiting pedicels 6.5–12 mm; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
119a. Fruits 6–7(–8) mm; ovules/seeds (1 or)2(or 3) per locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. camlikensis
119b. Fruits 11–12 mm; ovules/seeds 3–5 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cariensis
120a. Fruits wingless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
120b. Fruits winged. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
121a. Style 0.2–0.5(–0.6) mm; petals 2–2.8(–3) mm; ovules/seeds 2–6 per locule; U.S.A.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. parviflora
121b. Style 1–3 mm; petals 3–8 mm; ovules/seeds 2 or 3 per locule; Europe, Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
122a. Cauline leaves broadly obovate-orbicular; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. haussknecktii
122b. Cauline leaves lanceolate to oblong-ovate; Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
123a. Seeds 2.2–2.8 mm; fruits 3–4.5(–6) mm wide; sepals 1.7–2.9 mm; basal leaves repand or entire; Austria, Italy. . . . . . . . . . . . . . N. minima
123b. Seeds 1.7–1.8 mm; fruits 2.3–3.3 mm wide; sepals 2.6–3.7 mm; basal leaves often dentate; Spain. . . . . . . . . . . . . . . . . . . . . . N. nevadensis
124a. Fruits oblanceolate siliques, (3–)4–5 times longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. ferganensis
124b. Fruits broadly to narrowly obovate or obcordate silicles, 0.9–2.4 times longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

125a. Fruits wider than or about as broad as long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
125b. Fruits distinctly longer than broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
126a. Fruits 4–4.2 × 4.2–4.5 mm; petals 6–8.2 mm; style 1.5–2 mm; Albania. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. cikaea
126b. Fruits 3.8–7.8 × 4–8 mm; petals 4.5–6.5 mm; style 0.9–1.8 mm; C Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. montana
127a. Petals at least 3 times longer than sepals; cauline leaves strongly 3-veined abaxially. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. trinervia
127b. Petals less than 3 times longer than sepals; cauline leaves with only midvein distinct. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
128a. Wing more than 1 mm wide at fruit apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
128b. Wing to 1 mm wide at fruit apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
129a. Fruits 2.2–2.5 times longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. goesingensis
129b. Fruits 1.2–1.7(–1.9) times longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
130a. Fruits obdeltoid-obovate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
130b. Fruits obcordate, rarely obovate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
131a. Seeds 1.2–1.6 mm; sepals 1.8–2.5 mm; style 0.7–2 mm; fruits 5–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. janakae
131b. Seeds 1.7–2.6 mm; sepals 2–4 mm; style 1.5–4 mm; fruit 5–12 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
132a. Basal and lowermost cauline leaves dentate; petals 1.5–2.2 mm wide; anthers 0.7–0.9 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. praecox
132b. Basal and lowermost cauline leaves entire; petals 2–3 mm wide; anthers 1–1.3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. tymphaea
133a. Ovules/seeds 2–4 per locule; Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
133b. Ovules/seeds (4 or)5–8 per locule; Balkan Peninsula. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
134a. Plants glaucous; style 2–2.4 mm; fruits 3.5–4.5 mm wide; petals remaining white when dry. . . . . . . . . . . . . . . . . . . . . . . . . . N. edinensium
134b. Plants not glaucous; style (1.8–)2.4–3.6 mm; fruits (3.5–)4–6 mm wide; petals turning yellow when dry. . . . . . . . . . . . . . . . . N. ochroleuca
135a. Fruits (4.5–)5–7.3 mm wide; style 2.3–4.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
135b. Fruits 3.5–4.5(–5) mm wide; style 0.9–3 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
136a. Fruits oblong-obcordate, wing 1–2 mm wide; ovules/seeds 4–8 per locule; style 1.6–3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. lutescens
136b. Fruit obovate, wing 0.7–1.2 mm wide; ovules/seeds 5 or 6 per locule; style 0.9–2.4 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rhodopensis
137a. Seeds 2–2.5 mm; fruits 6–8 mm; petals 2–2.5 times longer than sepals; ovules/seeds 4–8 per locule Albania. . . . . . . . . . . . . . . N. albanica
137b. Seeds 1.5–2 mm; fruits 7–12 mm; petals less than twice longer than sepals; ovules/seeds 4 or 5 per locule Greece. . . . . . . . . . . N. boeotica
138a. Seeds/ovules (7–)9–14 per locule; China. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. flagellifera
138b. Seeds/ovules 1–6(–8) per locule; SW Asia, Europe, North and South America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
139a. Style 0.2–0.6 mm; Argentina, Chile, Mexico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
139b. Style (0.6–)1–4.2 mm; Europe, SW Asia, North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
140a. Fruiting pedicels 4–10 mm; fruits usually 2 times longer than broad; petals more than twice longer than sepals;
Argentina, Chile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. magellanica
140b. Fruiting pedicels 2–3 mm; fruits less than twice longer than broad; petals less than twice longer than sepals; Mexico. . . . . . . . . N. mexicana
141a. Fruits obdeltoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
141b. Fruits obovate to obcordate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
142a. Ovules/seeds 4–8 per locule; fruit 5–7.3 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. janakae
142b. Ovules/seeds 2–4 per locule; fruits 3–5(–6) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. kovatsii
143a. Ovules/seeds 1 or 2(or 3) per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
143b. Ovules/seeds (3 or)4–6 per locule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
144a. Fruit more than twice longer than broad; style 1–2 mm; Greece. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. epirota
144b. Fruits less than twice longer than broad; style 1.5–3.4 mm; Austria, Spain. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
145a. Seeds broadly ovate, 1.3–1.6 mm wide, brown; ovules/seeds 2 or 3 per locule; Austria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. alpestris
145b. Seeds ovate, 1.2–1.3 mm wide, orange; ovules/seeds 2 per locule; Spain. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. stenoptera
146a. Sepals 3.5–5.3 mm; petals 7–13 × (2.5–)3–5 mm; U.S.A.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. fendleri
146b. Sepals 1.6–3.2(–4) mm; petals 3–7(–9) × 0.7–2.7(–3) mm; Europe, Russia, Turkey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
147a. Fruits oblong-obovate, 2–2.5 × longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
147b. Fruits obovate or obcordate to broadly so, usually less than twice longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
148a. Petals equal, 2–3.2 mm wide; ovules/seeds 4–6 per locule; Austria, Balkan Peninsula. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. goesingensis
148b. Petals unequal, 1–1.8 mm wide; ovules/seeds 4 per locule; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. phrygia
149a. Seeds oblong, about twice longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. borealis
149b. Seeds ovate to broadly so, to 1.7 time longer than broad. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
150a. Stamens tetradynamous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
150b. Stamens equal or rarely subequal in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
151a. Petals 5–9 mm; seeds orange; fruits 6–8 mm; Russia, Ukraine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. macrantha
151b. Petals 4–5.5(–6) mm; seeds red-brown; fruits 5.5–6.2 mm; France, Italy, Switzerland. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. sylvia
152a. Seeds yellow- to orange-brown; ovules/seeds 4 per locule; anthers 0.5–0.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. aptera
152b. Seeds dark or red- brown; ovules/seeds 4–6 per locule; anthers (0.7–)0.8–9 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
153a. Sepals 1.7–2.2 mm; petals 3–4.5 mm; fruit 3.2–4 mm wide; style 1.2–1.9 mm; Turkey. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. angustifolia
153b. Sepals 2–3 mm; petals 4.2–6.4 mm; fruits 2.5–3.5 mm wide; style 1.5–3 mm; Balkan Peninsula, Bulgaria . . . . . . . . . . . . . . N. viridisepala
Synopsis of Noccaea
Unless otherwise indicated, Meyer accounts (1973, Hedge (1965) reduced the species to synonymy of Noccaea
2001b, 2003a–e, 2006a–e, 2010) were used as the primary microstyla (both as Thlaspi). However, N. amani differs from
source for typification of the majority of taxa. However, the latter by having reddish violet (vs. yellow) anthers, longer
the citation of type collections in various herbaria was (1–1.8 vs. 0.3–0.5 mm) styles, 4 or 5 (vs. 2 or 3) ovules/seeds
also enriched by checking JSTOR (http://plants.jstor.org/) per locule, and larger (3.8–4 × 1–1.5 vs. 2.8–2.9 × 0.7–0.8
and all major herbaria that have the images of their types mm) petals. In my opinion, the differences are substantial to
online. The presence in a given herbarium of more than one justify their recognition as independent species.
isotype or isolectotype is shown by a number (two and up)
preceding the herbarium acronym. Noccaea andersonii (Hook.f. & Thomson) Al-Shehbaz,
Adansonia sér. 3, 24(1): 91. 2002.
Noccaea abchasica (F.K.Mey.) Al-Shehbaz, comb. nov. Basionym: Iberidella andersonii Hook.f. & Thomson,
Basionym: Callothlaspi abchasicum F.K.Mey., Feddes J. Linn. Soc., Bot. 5: 177. 1861. TYPE: INDIA.
Repert. 84: 458. 1973. TYPE: GEORGIA. Abchazia, Garhwal [Uttarakhand], Chor hoti pass [Uttaranchal],
Gora Achaguvash, ca. 6,700 m, 29 Jul 1905, J. N. 16,000 ft [4,877 m], R. Strachey & J. E. Winterbottom
Woronow 249 (Holotype: LE). 12 (Lectotype here designated: K-000484215;
Homotypic synonym: Thlaspi abchasicum (F.K.Mey.) Isolectotype: GH).
V.I.Dorof., Konspekt. Fl. Kavkaza 3(2): 457. 2012. Homotypic synonm: Thlaspi andersonii (Hook.f. &
Distribution: Georgia. Thomson) O.E.Schulz, Anz. Akad. Wiss. Wien,
Math.-Naturwiss. Kl. 43: 98. 1926.
Noccaea albanica F.K.Mey., Feddes Repert. 84: 461. 1973. Heterotypic synonym: Iberidella tibetica C. Marquand &
TYPE: ALBANIA. Mali I Brasit, ca. 800 m, 7 May 1956, X. Airy Shaw, J. Linn. Soc., Bot. 48: 163. 1929. TYPE:
Qosia & M. Demiri s.n. (Holotype: JE). CHINA. Tibet, Nymia La, 14,000–15,000 ft [4,267–
Homotypic synonym: Thlaspi albanicum (F.K.Mey.) 4,572 m], 21 Jun 1924, F. K. Ward 5827 (Holotype:
Greuter & Burdet, Willdenowia 13: 95. 1983. K-000484217).
Distribution: Albania. Distribution: Bhutan, China (Xizang), India (Sikkim),
Nepal, Pakistan.
Noccaea alpestris (Jacq.) Kerguélen, Coll. Patrim. Nat. 8: Although Jafri (1973: 90), followed by Al-Shehbaz
14. 1993. (2002), designated Strachey & Winterbottom 11 (K) as the
Basionym: Thlaspi alpestre Jacq., Enum. Stirp. Vindob. lectotype of N. andersonii, this collection number was not
116, 260. 1762, non L., Sp. Pl., ed. 2. 2: 903. 1763. cited among the four syntypes listed by Hooker & Thomson
TYPE: not located. (1861). Therefore, a new lectotypification, proposed above,
Heterotypic synonym: Thlaspi alpinum Crantz, Stirp. was needed.
Austr., 1: 23. 1762; Thlaspi montanum L. var. alpinum
(Crantz) DC., Syst. Nat. 2: 380. 1821; Noccaea Noccaea angustifolia F.K.Mey., Feddes Repert. 84: 465.
crantzii F.K.Mey., Feddes Repert. 84: 461. 1973, non 1973. TYPE: TURKEY. Karagoelldagh, above Artabir, 6 Aug
N. alpina (L.) Rchb., Fl. Germ. Excurs. 663. 1832; 1894, P. Sintenis 7319 (Holotype: JE; Isotypes: E, G, W, Z).
TYPE: Illustration (T. 3, fig. 1) in Crantz (1762) Homotypic synonym: Thlaspi angustifolium (F.K.Mey.)
above. Greuter & Burdet, Willdenowia 13(1): 95. 1983.
Distribution: Austria. Distribution: Turkey.
Both Thlaspi alpestre and T. alpinum are generally
accepted as conspecific (Jalas et al., 1994). Meyer (1973, Noccaea annua (K.Koch) F.K.Mey., Feddes Repert. 84:
2006e) overlooked the fact that Jacquin’s Thlaspi alpestre 465. 1973.
was published prior to Crantz’s T. alpinum (Stafleu & Basionym: Thlaspi annuum K.Koch, Linnaea 15: 258.
Cowan, 1979). Therefore, the older epithet alpestre should 1841. TYPE: ARMENIA. Dschelaloglu, 24 Apr 1837,
have been taken instead of proposing a new name in K. Koch s.n. (Lectotype designated by Meyer (2006e:
Noccaea. 164): B-10077011; Isolectotype: B).
Heterotypic synonyms: Thlaspi stenopterum Conrath &
Noccaea amani (Post) F.K.Mey., Feddes Repert. 84: 465. Freyn, Bull. Herb. Boissier 3: 38. 1895, non Boiss.
1973. & Reut., Diagn. Pl. Orient. ser. 1, 8: 40. 1849; T.
Basionym: Carpoceras amani Post, Bull. Herb. Boissier freynii N.Busch in Kuznecov et al., Fl. Caucas. Crit.
1: 16. 1893. TYPE: TURKEY. Amanus, Jul 1891, 3(4): 171. 1908; Noccaea freynii (N.Busch) Czerep.,
G. Post s.n. (Lectotype designated by Meyer (2006e: Sosud. Rast. SSR 140. 1981. TYPE: ARMENIA.
182): JE). Ljalwar and Lok, 2,200–2,600 m, 1888 and 1891, P.
Homotypic synonym: Thlaspi amani (Post) Bornm., Mitt. Conrath s.n. (Lectotype designated by Meyer (2006e:
Thüring. Bot. Vereins 38: 58. 1929; T. amani (Post) 166): BRNM; Isolectotype: GZU).
Greuter & Burdet, Willdenowia 13: 95. 1983, comb. Distribution: Armenia, Georgia.
superfl. Several additional synonyms given by Meyer (2006e)
Distribution: Turkey. are not repeated here.
Noccaea antitaurica (F.K.Mey.) Al-Shehbaz, comb. nov. Homotypic synonym: Thlaspi arenarium (Duby) Jord. ex
Basionym: Callothlaspi antitauricum F.K.Mey., Feddes F.W.Schultz, Arch. Fl. France Allemagne 1: 162.
Repert. 84: 457. 1973. TYPE: TURKEY. [Kayseri], 1850.
Región montagneuse superieure du Dédé-Dagh Distribution: France.
(l’undes pics de l’anti-Tauris), à 12 lieues à l’ESE du
Mont-Argée, 8 Aug 1856, B. Balansa 439 (Holotype: Noccaea atlantica (Batt.) Al-Shehbaz, comb. nov.
G-BOIS). Basionym: Thlaspi atlanticum Batt. in Batt. & Trab., Fl. de
Homotypic synonym: Thlaspi antitauricum (F.K.Mey.) l’Alger. 1(Append. 2): 2. 1890. TYPE: ALGERIA.
Greuter & Burdet, Willdenowia 13: 95. 1983. Djebel-Tamesguida, 13 Jun 1890, J. A. Battandier s.n.
Distribution: Turkey. (Holotype: MPU-007905; Isotype: MPU).
Homotypic synonyn: Raparia atlantica (Batt.) F.K.Mey.,
Noccaea aptera (Velen.) F.K.Mey., Feddes Repert. 84: 462. Feddes Repert. 84: 458. 1973.
1973. Distribution: Algeria.
Basionym: Thlaspi apterum Velen., Allg. Bot. Z. Syst. 33.
1904. TYPE: BULGARIA. Sliven, 1887, Skorph s.n. Noccaea banatica (R.Uechtr.) F.K.Mey., Feddes Repert.
(Holotype: PRC). 84: 464. 1973.
Distribution: Albania, Bosnia and Herzegovina, Basionym: Thlaspi banaticum R.Uechtr., Oesterr. Bot. Z.
Bulgaria, Macedonia, Montenegro, Serbia, Turkey. 25: 186. 1875. TYPE: ROMANIA. Banat, Dumugled,
Simkovicz s.n. (Lectotype designated by Meyer
Noccaea apterocarpa (Rech.f. & Aellen) Al-Shehbaz & (2006e: 159): JE; Isolectotypes: LI, W).
Menke, Taxon 61: 949. 2012. Distribution: Romania.
Basionym: Aethionema apterocarpum Rech.f. & Aellen,
Phyton (Horn) 3: 49. 1951. TYPE: IRAN. Khorasan, Noccaea bellidifolia (Griseb.) F.K.Mey., Feddes Repert.
Kuh-e Hazar Masjed, 2,000–2,500 m, 8–9 Jun 1948, 84: 460. 1973.
K. H. Rechinger 5063 (Holotype: W; Isotypes: C, E, Basionym: Thlaspi bellidifolium Griseb., Spic. Fl. Rumel.
KD, M, MO, UC, US, WU, ZT). 2: 505. 1846. TYPE: MACEDONIA. Ljubatrin, A.
Homotypic synonyms: Aethionema trinervium (DC.) Boiss. Grisebach s.n. (Holotype: GOET).
var. apterocarpum (Rech.f. & Aellen) Hedge in Distribution: Albania, Bulgaria, Macedonia.
Rech.f., Fl. Iran. 57: 107. 1968; Thlaspi apterocarpum
(Rech.f. & Aellen) Mozaff., Iranian J. Bot. 7: 139. Noccaea boeotica F.K.Mey., Feddes Repert. 84: 462.
1996. 1973. TYPE: GREECE. Mt. Helicone Boeotica, 19 May
Distribution: Iran. 187?, T. G. Orphanides 1053 (Holotype: JE-00000206:
Noccaea apterocarpa was treated by Hedge (1968) as a Isotypes: G, S, W).
variety of Aethionema trinervium (=N. trinervia). However, Distribution: Greece.
it differs from the latter by having smaller (3–3.5 × 2–2.5
vs. 5–9 × 3–4 mm) fruits without (vs. with) apical notch Noccaea borealis F.K.Mey., Haussknechtia 12: 11. 2010.
and persistent (vs. caducous) sepals. Persistent sepals are TYPE: RUSSIA. Poljarnyj Ural, Bass. R. Syni, Verchovja r.
not known elsewhere in Noccaea. Lopta, 19 Jul 1926, B. N. Gorodkov s.n. (Holotype: LE).
Distribution: Russia.
Noccaea arctica (A.E.Porsild) Holub, Preslia 70: 107. 1998;
Thlaspi arcticum A.E.Porsild, Sargentia 4: 40. 1943. TYPE: Noccaea bornmuelleri (Rech.f.) Al-Shehbaz, comb. nov.
CANADA. Yukon Territories, Arctic coast of Mackenzie Basionym: Syrenopsis bornmuelleri Rech.f., Ann. Naturhist.
River delta, Kay Point, 69˚12'N, 138˚30'W, 23–25 Jul 1934, Mus. Wien 4: 49. 264. 1939. TYPE: TURKEY.
A. E. Porsild 7144 (Holotype: CAN; Isotypes: GH, S). Kurdistan, Siirt, Parklansschaft auf dem Daharaköl
Distribution: Canada (Yukon), U.S.A. (Alaska). Dağ, 1,700 m, 15 Jun 1936, J. Frödin 264 (Holotype:
I have not examined the material on which Noccaea W; Isotype: JE).
arctica is reported for Russia in the Panarctic Flora Homotypic synonym: Thlaspi bornmuelleri (Rech.f.)
(http://nhm2.uio.no/paf/results?biogeographic=&biocli Hedge, Notes Roy. Bot. Gard. Edinburgh 26: 184.
matic=&region=&name=noccaea+arctica). However, if the 1965.
fruit apex in that material is obtuse, then the Panarctic re- Distribution: Turkey.
cord is correctly based on N. arctica. However, if the apex is
distinctly notched, then that record is based on N. borealis. Noccaea bovis (F.K.Mey.) Al-Shehbaz, comb. nov.
Basionym: Thlaspiceras bovis F.K.Mey., Feddes Repert. 84:
Noccaea arenaria (Duby) F.K.Mey., Feddes Repert. 84: 454. 1973. TYPE: TURKEY. Amanus Mts., between
463. 1973. Achagi Zarkoun and Bagajak, 1,800 m, 1 Jul 1932, A.
Basionym: Thlaspi alpestre L. var. arenarium Duby, Bot. Eig & M. Zoharay s.n. (Holotype: HUJ).
Gall. 1: 38. 1828. TYPE: FRANCE. Landes, Mont- Homotypic synonym: Thlaspi bovis (F.K.Mey.) Greuter &
de-Marsan, Perris (Neotype designated by Meyer Burdet, Willdenowia 13(1): 95. 1983.
(2006e: 127): LY; Isoneotypes: JE, S, W). Distribution: Turkey.
Noccaea brachypetala (Jord.) F.K.Mey., Feddes Repert. treated as a polymorphic entity as done here.
84: 464. 1973.
Basionym: Thlaspi brachypetalum Jord., Observ. Pl. Nouv. Noccaea bulbosa (Spruner) Al-Shehbaz, comb. nov.
3: 5. 1846. TYPE: FRANCE. Grenoble et de Gap, Basionym: Thlaspi bulbosum Spruner in Boiss., Diagn.
A. Jordan (Lectotype designated by Meyer (2006e: Pl. Orient. ser. 1, 1: 74. 1843. TYPE: GREECE.
14): LY). “Hab. in umbrosis regionis montanae, mons Parnes
Heterotypic synonyms: Thlaspi alpestre L., Sp. Pl., ed. 2. 2: Atticae,” 4,000 ft [1,219 m], 1840, W. von Spruner
903. 1763, nom. illeg., non Jacq, Enum. Stirp. Vindob. s.n. (Lectotype designated by Meyer (2006b: 197):
116, 260. 1762. TYPE: “Habitat in Austria. Stamina G-BOIS; Isolectotypes: JE, W).
flore longiora,” (Lectotye designated by Marhold & Homotypic synonym: Raparia bulbosa (Spruner) F.K.Mey.,
Mátonfi in Cafferty & Jarvis (2002: 536): Herb Linn. Feddes Repert. 84: 458. 1973.
No. 825.14 (LINN)). Heterotypic synonym: Raparia bulbosa (Spruner) F.K.Mey.
Thlaspi tatrense Zapał., Bull. Acad. Cracovie, Cl. Sci. subsp. aegaea F.K.Mey., Haussknechtia 11: 199. 2006.
Math 1913: 431. 1913; Noccaea brachypetala subsp. TYPE: GREECE. Ikaria (Nikaria), Hagios Kirykos,
tatrensis (Zapał.) F.K.Mey., Feddes Repert. 84: 464. ca. 900 m, 18 Apr 1934, K. H. & F. Rechinger 4443
1973. TYPE: SLOVAKIA. Tatry, 1,585 m, 30 Jul (Holotype: W).
1879, E. Janota s.n. (Lectotype designated by Meyer Distribution: Greece.
(2006e: 148): KRAM). Meyer (2006b) distinguished subsp. aegaea from subsp.
Thlaspi huteri Kerner, Sched. Fl. Austro-Hung. 7: 39. bulbosa solely on the slight differences in floral size and
1896; Noccaea brachycarpa subsp. huteri (Kerner) style length. In my opinion, these are insignificant to support
F.K.Mey., Feddes Repert. 84: 464. 1973. TYPE: the recognition of infraspecific taxa.
AUSTRIA. Tirol, 1,300–1,500 m, R. Huter 2523
(Holotype: WU; Isotypes: GOET, 2GZU, K, W, Z). Noccaea caerulescens (J.Presl & C.Presl) F.K.Mey., Feddes
Distribution: Austria, Czech Republic, Finland, France, Repert. 84: 463. 1973.
Italy, Slovakia, Spain, Sweden, Switzerland. Basionym: Thlaspi caerulescens J.Presl & C.Presl, Fl. Čech.
This highly variable species was divided by Meyer 133. 1819. TYPE: CZECH REPUBLIC. Prague, W.
(1973; 2006e) into three subspecies under which he listed Mann (Neotype designated by Meyer (2006e: 134):
over two dozen synonyms. Nonetheless, the distinguishing PR).
characters used, especially the size measurements of leaves, Heterotypic synonyms: Thlaspi sylvestre Jord., Observ. Pl.
are part of a continuum over a large geographical range. Nouv. 3: 9. 1846; T. caerulescens J.Presl & C.Presl
subsp. sylvestre (Jord.) M.Laínz, Bol. Soc. Brot.
Noccaea brevistyla (DC.) Steud., Nomencl. Bot., ed. 2. 2: sér. 2, 53: 38. 1979; Noccaea caerulescens subsp.
196. 1841. sylvestris (Jord.) F.K.Mey., Feddes Repert. 84: 463.
Basionym: Hutchinsia brevistyla DC., Syst. Nat. 2: 387. 1973. TYPE: FRANCE. Lyon, Soucieux, A. Jordan
1821. TYPE: FRANCE. Corsica, Labillardiére s.n. s.n. (Lectotype designated by Meyer (2006e: 138):
(Holotype: G-DC). LY; Isolectotype: JE).
Homotypic synonym: Thlaspi brevistylum (DC.) Jord., Thlaspi sylvestre var. oligospermum Merino, Fl.
Observ. Pl. Nouv. 3: 27. 1846. Galicia 3: 510. 1909; T. oligospermum (Merino)
Heterotypic synonyms: Thlaspi rivale C.Presl, Delic. Greuter & Burdet, Willdenowia 13(1): 96. 1983;
Prag. 1: 12. 1822; Noccaea brevistyla subsp. rivalis Noccaea oligosperma (Merino) Holub, Preslia 70:
(C.Presl) F.K.Mey., Haussknechtia 12: 195. 2006. 108. 1998. TYPE: FRANCE. Coruna, R. P. B. Merino
TYPE: ITALY. Sicily, Curma grandi Nebrodum, s.n. (Holotype: not seen).
1817, C. B. Presl s.n. (Holotype: PRC). Distribution: Austria, Belgium, Czech Republic,
Thlaspi pseudorivulare Hausskn. ex Bornm., Mitth. England, Finland, Germany, Netherlands, Norway, Poland,
Thüring. Bot. Vereins 38: 56. 1929; Noccaea Russia, Scotland, Sweden, Switzerland, Wales.
pseudorivularis (Hausskn. ex Bornm.) F.K.Mey., An extremely variable species that was divided into
Feddes Repert. 84: 466. 1973; N. brevistyla subsp. numerous entities recognized at the specific, subspecific,
pseudorivularis (C.Presl) F.K.Mey., Haussknechtia and varietal ranks. The complex is in much need of thorough
12: 196. 2006. TYPE: GREECE: Pinus Tymphaeus, systematic and molecular studies to resolve its taxonomy. It
Chaliki, Mt. Plaka, 4 Jul 1896, P. Sintenis 802 is beyond this study to deal with such complexity, and most
(Lectotype designated by Meyer (2006e: 196): JE- of the three dozen synonyms listed by Meyer (2006e) are
00010170; Isolectotypes: B, S, W). not repeated herein.
Distribution: Albania, France, Greece, Italy.
This variable species complex is in need of further Noccaea caespitosa (Boiss.) Al-Shehbaz & Menke, Taxon
studies, and it seems that the variation is somewhat 61: 949. 2012.
continuous in every character except for distribution. It Basionym: Iberidella caespitosa Boiss., Ann. Sci. Nat.,
remains to be seen whether or not the species merits the Bot., ser. 2 17: 190. 1842. TYPE: TURKEY. Alpibus
division into subspecies, as done by Meyer (2006e), or Armeniae, P. M. R. Aucher-Eloy 155 (Holotype:
G-BOIS; Isotypes: BM, K).
Homotypic synonyms: Aethionema caespitosum (Boiss.) Noccaea cariensis (Carlström) Parolly, Nordt & Aytaç, Bot.
Boiss., Fl. Orient. 1: 343. 1867; Eunomia caespitosa J. Linn. Soc. 150: 413. 2006.
(Boiss.) O.E.Schulz, Bot. Jahrb. Syst. 66: 95. 1933; Basionym: Thlaspi cariensis Carlström, Willdenowia 16: 73.
Apterigia caespitosa (Boiss.) V.I.Dorof., Konspekt. 1986. TYPE: TURKEY. Mugla, 1 km E of Marmaris,
Fl. Kavkaza 3(2): 459. 2012. 19 Apr 1983, A. Carlström 9293 (Holotype: LD).
Distribution: Turkey. Homotypic synonym: Callothlaspi cariense (Carlström)
F.K.Mey., Haussknechtia 11: 183. 2006.
Noccaea camlikensis Aytaç, Nordt & Parolly, Bot. J. Linn. Distribution: Turkey.
Soc. 150: 410. 2006. TYPE: TURKEY. Konya, Derebucak,
Çamlik village, Kizildag, 1,400–1,600 m, 4 Jun 2000, H. Noccaea cepaeifolia (Wulfen) Rchb., Fl. Germ. Excurs.
Duman 7306 (Holotype: GAZI; Isotype: ANK). 663. 1832.
Homotypic synonym: Callothlaspi camlikense (Aytaç, Basionym: Iberis caepaifolia Wulfen in Jacqin, Misc.
Nordt & Parolly) F.K.Mey., Haussknechtia 11: 185. Austriac. 2: 28. 1781. TYPE: ITALY. “In valle Rabl,”
2006. F. X. Wulfen s.n. (Lectotype designated by Meyer
Distribution: Turkey. (2006e: 40): W).
Homotypic synonym: Thlaspi cepaeifolium (Wulfen)
Noccaea campylophylla (F.K.Mey.) Al-Shehbaz, comb. W.D.J.Koch, Deutschl. Fl., ed. 3. 4: 534. 1833.
nov. Distribution: Austria, Italy.
Basionym: Vania campylophylla F.K.Mey., Feddes Repert.
84: 467. 1973. TYPE: TURKEY. Prov. Van, Distr. Noccaea cikaea F.K.Mey., Feddes Repert. 84: 461. 1973.
Satak, Kavuṣṣahap Dag, 3,100 m, 23 Jul 1954, P. H. TYPE: ALBANIA. Çika, Mali e Qorres, ca. 1,220 m, F. K.
Davis & O. Polunin 23132 (Holotype: JE; Isotype: E). Meyer 5206 (Holotype: JE).
Distribution: Iran, Turkey. Homotypic synonym: Thlaspi cikaeum (F.K.Mey.) Greuter
& Burdet, Willdenowia 13: 95. 1983.
Noccaea cappadocica (Boiss. & Balansa) Al-Shehbaz, Distribution: Albania.
comb. nov.
Basionym: Carpoceras cappadocicum Boiss., Diagn. Pl. Noccaea cilicica (Schott & Kotschy ex Boiss.) Al-Shehbaz,
Orient. ser. 2, 6: 19. 1859. TYPE: TURKEY. Region comb. nov.
montagneuse sperieure du Dédé-Dagh (lʼun des pics Basionym: Carpoceras cilicicum Schott & Kotschy ex
de lʼanti-Tauris), à 12 lieues à lʼESE du Mont Argée, 6 Boiss., Fl. Orient. 1: 332. 1867. TYPE: TURKEY.
Aug 1856, B. Balansa 438 (Holotype: G-BOIS). Taurus, Bulghar Dagg, above Gülek, 7,000 ft [2,130
Homotypic synoynms: Thlaspiceras cappadocicum (Boiss. m], 8 Jul 1853, C. G. T. Kotschy 70 (Holotype:
& Balansa) F.K.Mey., Feddes Repert. 84: 455. 1973; G-BOIS; Isotypes: B, K, 2P, S, W, WAG).
Thlaspi cappadocicum (Boiss. & Balansa) Bornm., Homotypic synonym: Kotschyella cilicica (Schott &
Repert. Spec. Nov. Regni Veg. Beih. 89: 70. 1936. Kotschy ex Boiss.) F.K.Mey., Feddes Repert. 84: 457.
Heterotypic synonyms: Thlaspiceras cappadocicum (Boiss. 1973; Thlaspi cilicicum (Boiss.) Hayek, Ann. Nat.
& Balansa) F.K.Mey. subsp. microspermum F.K.Mey., Hofm. Wien 28: 155. 1914.
Feddes Repert. 84: 455. 1973; Thlaspi cappadocicum Distribution: Turkey.
(Boiss. & Balansa) Bornm. subsp. microspermum
(F.K.Mey.) Greuter & Burdet, Willdenowia 13(1): Noccaea cochlearioides (Hook.f. & Thomson) Al-Shehbaz,
95. 1983. TYPE: TURKEY. Amanus Mts., Achagi Adansonia sér. 3, 24: 91. 2002.
Zarkoun, ca. 1,800 m, 1 Aug 1932, A. Eig & M. Basionym: Thlaspi cochlearioides Hook.f. & Thomson,
Zohary s.n. (Holotype: HUJ). J. Linn. Soc., Bot. 5: 177. 1861. TYPE: [INDIA],
Distribution: Turkey. Sikkim, 14,000–16,000 ft, J. D. Hooker s.n.
Five specimens of Noccaea cappadocica (at GOET, JE, (Holotype: K-00077284).
3P) differ from the holotype only in the collection number Distribution: Bhutan, India, Nepal, Pakistan.
(Balansa 1004 instead of Balansa 438), and they carry the
exact collection data. Noccaea cornuticarpa (Naqshi) Al-Shehbaz, comb. nov.
Basionym: Thlaspi cornuticarpum Naqshi, J. Econ. Tax.
Noccaea capricornuta (F.K.Mey.) Al-Shehbaz, comb. nov. Bot. 5: 717. 1984. TYPE: PAKISTAN. Apharwat
Basionym: Thlaspiceras capricornutum F.K.Mey., Feddes top, A. R. Naqshi 3462 (Holotype: KASH; Isotype:
Repert. 84: 454. 1973. TYPE: TURKEY. Cassius, KASH).
Ain el Aramie, 700 m, 17 May 1933, E. Wall s.n. Distribution: Pakistan.
(Holotype: S).
Homotypic synonym: Thlaspi capricornutum (F.K.Mey.) Noccaea corymbosa (J.Gay) F.K.Mey., Feddes Repert. 84:
Greuter & Burdet, Willdenowia 13(1): 95. 1983. 460. 1973.
Distribution: Syria, Turkey. Basionym: Hutschinsia corymbosa J.Gay, Syll. Pl. Nov.
1: 169. 1824. TYPE: SWITZERLAND. Wallis, near
Zermatten, J. Gay s.n. (Holotype: K; Isotypes: JE, W).
Homotypic synonym: Thlaspi corymbosum (J.Gay) Rchb., Kassan Oghlu, Gorumse, 5,600 ft [1,707 m], 13 May
Ic. Fl. Germ. Helv. 2: 3. 1837, non Molina, Sag. Stor. 1859, C. G. T. Kotschy 62 (Holotype: W; Isotypes:
Nat. Chili, ed. 2. 292. 1810. 2B, 2BM, G, JE, 3P, S, W, ZT).
Distribution: France, Italy, Switzerland. Distribution: Turkey.
Noccaea crassiuscula (F.K.Mey.) Al-Shehbaz, comb. nov. Noccaea dolichocarpa (Zohary) Al-Shehbaz, comb. nov.
Basionym: Masmenia crassiuscula F.K.Mey., Feddes Basionym: Carpoceras dolichocarpum Zohary, Palestine J.
Repert. 84: 468. 1973. TYPE: TURKEY. Osmaniye, Bot., Jerusalem Ser. 2: 159. 1941. TYPE: TURKEY.
Koyun meleten dag Amanos daglari, ca. 1960 m, 9 Hatay, Amanus, E slopes between Karagouz and
Jun 1968, Y. Akman 203 (Holotype: BASB). Bagajak, 800–1,600 m, A. Eig & M. Zohary s.n.
Homotypic synonym: Thlaspi crassiusculum (F.K.Mey.) (Holotype: HUJ).
Greuter & Burdet, Willdenowia 13: 95. 1983. Homotypic synonyms: Thlaspiceras dolichocarpum
Distribution: Turkey. (Zohary) F.K.Mey., Feddes Repert. 84: 454. 1973;
Thlaspi dolichocarpum (Zohary) Greuter & Burdet,
Noccaea cretica (Degen & Jáv.) F.K.Mey., Feddes Repert. Willdenowia 13(1): 95. 1983.
84: 466. 1973. Distribution: Turkey.
Basionym: Thlaspi microphyllum Boiss. & Orph. subsp.
creticum Degen & Jáv., Magyar Bot. Lapok 21: 25. Noccaea eburneosa F.K.Mey., Feddes Repert. 84: 464.
1923. TYPE: GREECE. Crete, Dist. Mylopotamos, 1973. TYPE: SWITZERLAND. Avers, below Juppa, 2,010
above Jovis, Baldacci 53 (Lectotype designated by m, 17 Jul 1941, W. Koch s.n. (Holotype: ZT; Isotype: JE).
Meyer (2006e: 191): BP; Isolectotypes: BPU, G, W, Distribution: Switzerland.
WU).
Homotypic synonmys: Thlaspi graecum Jord. subsp. Noccaea edinensium F.K.Mey., Feddes Repert. 84: 460.
creticum (Degen & Javorka) Greuter, Flor. Rep. 1973. TYPE: TURKEY. Prov. Kütahya, Murat Dag, above
Cretan Area 44. 1972; Thlaspi creticum (Degen & Gediz, above Kesik Sögüt, 1,900 m, 5 Jul 1962, P. H. Davis
Jáv.) Greuter & Burdet, Willdenowia 13(1): 95. 1983. & M. J. E. Coode 36781 (Holotype: E).
Distribution: Greece. Homotypic synonym: Thlaspi edinensium (F.K.Mey.)
Greuter & Burdet, Willdenowia 13: 95. 1983.
Noccaea cypria (Bornm.) F.K.Mey., Feddes Repert. 84: Distribution: Turkey.
466. 1973.
Basionym: Thlaspi cyprium Bornm., Mitth. Thüring. Bot. Noccaea eigii (Zohary) Al-Shehbaz, comb. nov.
Vereins 38: 54. 1929. TYPE: CYPRUS. Summit of Basionym: Carpoceras eigii Zohary, Palestine J. Bot., Jeru-
Mt. Troodos and at Riv. Maschinari, 6,000 ft [1,829 salem Ser. 2: 158. 1941. TYPE: SYRIA. Amanus Mts.,
m], 20 May 1862, C. G. T. Kotschy s.n. (Holotype: JE; between Achagi Zarkoun and Bagajak, ca. 1,800 m, 1
Isotypes: G-BOIS, W). Jul 1932, A. Eig & M. Zohary s.n. (Holotype: HUJ).
Distribution: Cyprus. Homotypic synonyms: Thlaspiceras eigii (Zohary) F.K.Mey.,
Feddes Repert. 84: 454. 1973; Thlaspi eigii (Zohary)
Noccaea dacica (Heuff.) F.K.Mey., Feddes Repert. 84: 464. Greuter & Burdet, Willdenowia 13(1): 95. 1983.
1973. Heterotypic synonyms: Thlaspiceras eigii subsp.
Basionym: Thlaspi dacicum Heuff., Oesterr. Bot. Z. 8: 26. samuelssonii F.K.Mey., Feddes Repert. 84: 454.
1858. TYPE: ROMANIA. Retjeszath, 6 Aug 1836, 1973; Thlaspi eigii subsp. samuelssonii (F.K.Mey.)
Rochel & Heuffel s.n. (Neotype designated by Meyer Greuter & Burdet, Willdenowia 13: 95. 1983.
(2996e: 157): DR). TYPE: TURKEY. Cassius, prope transitum ad Ain el
Heterotypic synonyms: Noccaea dacica subsp. Aramie, ca. 750 m, 26 Apr 1933, G. Samuelsson 4087
montenegrina F.K.Mey., Feddes Repert. 84: 464. (Holotype: S S-G-8723).
1973; Thlaspi dacicum Heuff. subsp. montenegrinum Distribution: Syria, Turkey.
(F.K.Mey.) Greuter & Burdet, Willdenowia 13: 95. The slight differences in flower size between subspp.
1983. TYPE: MONTENEGRO. Biele Carini, 7 Jul samuelssonii and eigii are quantitative in nature and represent
1872, J. Pantocsek s.n. (Holotype: W). extremes of an otherwise morphological continuum.
Distribution: Montenegro, Romania.
The morphologically overlapping quantitative characters Noccaea elegans (Boiss.) Al-Shehbaz, comb. nov.
used by Meyer (2006e) do not justify the division of the Basionym: Thlaspi elegans Boiss., Diagn. Pl. Orient. ser. 1,
species into infraspecific taxa. 5: 82. 1844. TYPE: TURKEY. Caria, 1842, C. Pinard
s.n. (Holotype: G-BOIS; Isotypes: B, G, GOET, JE,
Noccaea densiflora (Boiss. & Kotschy) F.K.Mey., Feddes 2K, W).
Repert. 84: 465. 1973. Homotypic synonym: Thlaspiceras elegans (Boiss.)
Basionym: Thlaspi densiflorum Boiss. & Kotschy in Unger F.K.Mey., Feddes Repert. 84: 456. 1973.
& Kotschy, Ins. Cyp. 328. 1865. TYPE: TURKEY. Distribution: Turkey.
Noccaea epirota (Halácsy) F.K.Mey., Feddes Repert. 84: Noccaea fendleri (A.Gray) Holub subsp. fendleri
459. 1973. Heterotypic synonyms: Thlaspi prolixum A.Nelson, Amer.
Basionym: Thlaspi epirotum Halácsy, Consp. Fl. Graec. 1: J. Bot. 32: 287. 1945. TYPE: U.S.A. Arizona, San
109. 1900. TYPE: GREECE. [N Pindos], “in lapidosis Carlos Indian Reservation, about 20 miles W of Rice,
mobil. Alp. M. Smolika dist. Konitsa,” 18 Jul 1896, 9 May 1935, A. & R. Nelson 1832 (Holotype: RM;
A. Baldacci 208 (Lectotype designated by Franzén in Isotype: MO).
Strid (1986: 324) and, apparently unaware of that, by Thlaspi stipitatum A.Nelson, Amer. J. Bot. 32: 288. 1945.
Meyer (2006e: 42): WU; Isolectotypes: WU, Z). TYPE: U.S.A. Arizona, Arizona Mts., between Ruby
Distribution: Greece. and Tucson-Nogales Hwy, Pena Banca Mts, S of
Tucson, not far from Ruby, 15 Mar 1935, A. & R.
Noccaea fendleri (A.Gray) Holub, Preslia 70: 108. 1998. Nelson 1196 (Holotype: RM; Isotypes: GH, MO, NY,
Basionym: Thlaspi fendleri A.Gray, Smithsonian Contr. S, UC, US, UTC, WTU).
Knowl. 5(6): 14. 1853. TYPE: U.S.A. New Mexico, Distribution: U.S.A. (Arizona, New Mexico, Texas).
[on the Organ Mts., NE of El Paso, 1852, C. Wright
1322 (Lectotype partially designated by Wooton & Noccaea fendleri (A.Gray) Holub subsp. glauca (A.Nelson)
Standley (1915: 273) and completed by Holmgren Al-Shehbaz & M.Koch, Syst. Bot. 29: 382. 2004.
(1971: 88): GH; Isolectotypes: GH, K, MO, PH, UC). Basionym: Thlaspi alpestre L. var. glaucum A.Nelson,
Homotypic synonym: T. montanum L. var. fendleri (A.Gray) First Rep. Fl. Wyoming, 84. 1896. TYPE: U.S.A.
P.K.Holmgren, Mem. New York Bot. Gard. 21(2): 87. Wyoming, Albany Co., La Plata Mines, 21 Aug 1895,
1971. A. Nelson 1777 (Holotype: RM; Isotypes: GH, ILL,
Distribution: U.S.A. ISC, NY, UC, US).
Meyer (2006e) did not treat the New World Noccaea Homotypic synonyms: Noccaea glauca (A.Nelson) Holub,
in his account and, therefore, full synonymies and Preslia 70: 108. 1998; Thlaspi fendleri var. glaucum
typifications are provided herein. North American authors (A.Nelson) C.L.Hitchc., Vasc. Pl. Pacif. N. W. 2: 554.
(e.g., Holmgren, 1971; Rollins, 1993) treated this species 1964; T. glaucum (A.Nelson) A.Nelson, Bull. Torrey
as conspecific with the European Thlaspi montanum, Bot. Club 25: 275. 1898.
whereas Payson (1926) correctly recognized the North Heterotypic synonyms: Thlaspi coloradense Rydb., Bull.
American taxa (five subspecies are cited below) as native Torrey Bot. Club 28: 280. 1901; T. fendleri A.Gray var.
to the continent. Molecular phylogenetic studies by Koch & coloradense (Rydb.) Maguire, Amer. Midl. Naturalist
Al-Shehbaz (2004) amply supported Payson’s view of the 27: 469. 1942; Noccaea coloradensis (Rydb.) Holub,
independent status of the North American representatives. Preslia 70: 107. 1998. TYPE: U.S.A. Colorado, El
In addition to the substantial disjunction between the Paso Co., Bald Mt., near Pikes Peak, 11,500 ft [3,505
central European Noccaea montana and the western North m], 1 Aug 1896, E. A. Bessy s.n. (Holotype: NY).
American N. fendleri, the former differs by having fruits Thlaspi purpurascens Rydb., Bull. Torrey Bot.
shorter than wide to rarely as long as wide, whereas the Club 28: 281. 1901; T. alpestre var. purpurascens
latter has fruits 1.5–2 times longer than wide. Both Payson (Rydb.) Ostenf., Skr. Vidensk.-Selsk. Christiana,
and Rollins (1993) recognized four species in this complex, Math.-Naturvidensk. Kl. 1909(8): 47. 1910. TYPE:
but I accept their treatment by Holmgren as infraspecific U.S.A. Arizona, near Prescott, 1876, E. Palmer 571
taxa at the subspecific instead of varietal rank. For keys (Holotype: NY-00185801; Isotypes: MO, 2NY, YU).
distinguishing the five subspecies of Noccaea fendleri, see
Thlaspi australe A.Nelson, Amer. J. Bot. 32: 287.
Al-Shehbaz (2010) and (as Thlaspi) Holmgren (1971).
1945. TYPE: U.S.A. Arizona, W side of Baboquivari
Peak, 16 Apr 1935, A. & R. Nelson 1545 (Holotype:
Noccaea fendleri (A.Gray) Holub subsp. californica
RM; Isotypes: GH, MO, NY, S, UC, US, WIS).
(S.Watson) Al-Shehbaz & M.Koch, Syst. Bot. 29: 382.
2004. Thlaspi glaucum var. hesperium Payson, Univ.
Basionym: Thlaspi californicum S.Watson, Proc. Amer. Wyoming Publ. Sci., Bot. 1: 154. 1926; T. hesperium
Acad. Arts 17: 365. 1882. TYPE: U.S.A. California, (Payson) G.N.Jones, Univ. Wash. Publ. Biol. 5: 161.
Humboldt Co., Kneeland Prairie, 2,500 ft [762 m], 1936; T. fendleri var. hesperium (Payson) C.L.Hitchc.,
Jun 1882, B. Rattan s.n. (Holotype: GH; Isotypes: DS, Univ. Wash. Publ. Biol. 17(2): 554. 1964. TYPE:
NDG, NY, P, 2US). U.S.A. California, Trinity Co., Doreleska, Salmon
Homotypic synonyms: Thlaspi alpestre L. var. californicum Mts., 2,000 m, 18 Jul 1909, H. M. Hall 8570
(S.Watson) Jepson, Man. Fl. Pl. Calif. [Jepson] (Holotype: RM; Isotypes: 2C, DS, GH, MO, 2NY, S,
436. 1925; T. glaucum (A.Nelson) A.Nelson subsp. UC, US).
californicum (S.Watson) Munz, Aliso 4: 91. 1958; Thlaspi fendleri var. tenuipes Maguire, Amer. Midl.
T. montanum L. var. californicum (S.Watson) Naturalist 27: 469. 1942. TYPE: U.S.A. Utah, Sanpete
P.K.Holmgren, Mem. New York Bot. Gard. 21(2): 80. Co., Mayfield Canyon, 0.5 mile above Ranger Station,
1971. 10,928 ft [3,331 m], 8 Aug 1940, B. Maguire 19998
Distribution: U.S.A. (California). (Holotype: UTC).
Thlaspi glaucum A.Nelson var. pedunculatum Noccaea firmiensis F.K.Mey., Feddes Repert. 84: 463. 1973.
Payson, Univ. Wyoming Publ. Sci., Bot. 1: 152. 1926. TYPE: FRANCE. Aveyron, Firmy, 12 Apr–20 May 1882, L.
TYPE: U.S.A., Idaho, [Nez Perce Co. or Latah Co.], Giraudias s.n. (Holotype: JE-00010185; Isotype: JE).
near Kendrick, 21 Apr 1894, L. F. Henderson 2893 Homotypic synonym: Thlaspi firmiense (F.K.Mey.) Greuter
(Holotype: RM). & Burdet, Willdenowia 13: 96. 1983.
Distribution: Mexico, U.S.A. (Arizona, California, Distribution: France.
Colorado, Idaho, Montana, Nevada, New Mexico, Oregon,
Texas, Utah, Washington, Wyoming). Noccaea flagellifera (O.E.Schulz) Al-Shehbaz, Adansonia
sér. 3, 24: 91. 2002.
Noccaea fendleri (A.Gray) Holub subsp. idahoense Basionym: Thlaspi flagelliferum O.E.Schulz, Anz. Akad.
(Payson) Al-Shehbaz & M.Koch, Syst. Bot. 29: 383. 2004. Wiss. Wien, Math.-Naturwiss. Kl. 43: 98. 1926.
Basionym: Thlaspi idahoense Payson, Univ. Wyoming TYPE: China. Sichuan. Wa-schan, Yadschou, May
Publ. Sci., Bot. 1: 159. 1926. TYPE: U.S.A. Idaho, 1915, H. Weigold s.n. (Holotype: W; Isotype: B).
Custer Co., Bonanza, alt. 7,000 ft [2,134 m], 25 Jul Distribution: China (Sichuan).
1916, J. F. Macbride & E. B. Payson 3436 (Holotype:
RM; Isotypes: CAS, CM, GH, MO, NY, P, UC, US). Noccae gardeziana F.K.Mey., Haussknechtia 12: 172.
Homotypic synonyms: Thlaspi fendleri var. idahoense 2006. TYPE: AFGHANISTAN. Prov. Gardez, Paktia, in
(Payson) C.L.Hitchc., Vasc. Pl. Pacif. N. W. 2: 554. jugo Sata Kandao, between Gardez and Khost, 2,900 m, 8
1964; T. montanum L. var. idahoense (Payson) P.K. Jul 1965, K. H. Rechinger 32117 (Holotype: W).
Holmgren, Mem. New York Bot. Gard. 21(2): 81. 1971. Distribution: Afghanistan.
Heterotypic synonyms: Thlaspi aileeniae Rollins, Contr.
Gray Herb. 214: 16. 1984; Thlaspi idahoense Payson Noccaea germanii Al-Shehbaz, nom. nov.
var. aileeniae (Rollins) Rollins, Crucifer. Continental Replaced name: Eunomia rotundifolia C.A.Mey., Verz. Pfl.
N. Amer. 904. 1993. TYPE: U.S.A. Idaho, Custer Co., Casp. Meer. 194. 1831, non Noccaea rotundifolia (L.)
off road to Cape Horn, between Knopp Creek and Moench, Suppl. Meth. 89. 1802. TYPE: “In alpibus
Valley Creek, ca. 16 miles NW of Stanley, 15 Jul 1983, Elborus Caucasi occidentalis, inter fragmina lapidum
R. C. & K. W. Rollins & A. G. Roads 83287 (Holotype: (alt. 1500 hexap.).” Additional data on syntypes: “locis
GH; Isotypes: CAS, E, GH, ID, NY, RSA, US). lapidosis (alt. 9,000 p.), 10 July 1829, Enum. Cauc.
Distribution: U.S.A. (Idaho). Casp. N. 1678, D. Meyer” (Lectotype designated by
Dorofeyev (2012: 460): LE; Isolectotypes: 2LE).
Noccaea fendleri (A.Gray) Holub subsp. siskiyouense Distribution: Armenia, Azerbaijan, Georgia.
(P.K.Holmgren) Al-Shehbaz & M.Koch, Syst. Bot. 29: 383. The species is named in honor of Dr. Dmitry A. German
2004. (12 August 1976–), an outstanding student of the Brassicaceae.
Basionym: Thlaspi montanum L. var. siskiyoense
P.K.Holmgren, Mem. New York Bot. Gard. 21(2): 84. Noccaea goesingensis (Halácsy) F.K.Mey., Feddes Repert.
1971. TYPE: U.S.A. Oregon, Kerby Flat at base of 84: 462. 1973.
Eight Dollar Mt., above Illinois River, 2 miles W of Basionym: Thlaspi goesingense Halácsy, Oesterr. Bot. Z. 30:
Selma, 18 May 1942, L. Constance & R. C. Rollins 173. 1880. TYPE: AUSTRIA. Mt. Goesing, ad pagum
2972 (Holotype: WTU; Isotypes: GH, MO, NY, RM, Ternits, 4 May 1880, E. von Halácsy s.n. (Holotype:
UC, US, UTC). WU; Isotypes: 3B, F, 2FR, GOET, GT, 2GZU, 2HBG,
Distribution: U.S.A. (Oregon). JE, 2K, MA, US, Z).
Heterotypic synonym: Thlaspi umbrosum Waisbecker,
Noccaea ferganensis (N.Busch) Czerep., Sosud. Rast. Oesterr. Bot. Z. 43: 318; 44: 74. 1893. TYPE:
SSSR 140. 1981. AUSTRIA. Bernstein, 700 m, 8 May 1892, A.
Basionym: Thlaspi ferganense N.Busch, Bot. Zhurn. Waisbecker s.n. (Holotype: BP; Isotypes: B, JE, K).
S.S.S.R. 21: 551. 1936. TYPE: “Fergana, jugum Distribution: Austria, Bosnia and Herzegovina,
Alaicum, prope Ak-basoga. In decliv. Angustiarum Bulgaria, Serbia.
inter Juniperus, 9,500 ft [2,896 m]. 30 Jun 1901. Leg.
Alexeenko (Flora Turkestanica Ex. 1983” (Lectotype Noccaea graeca (Jord.) F.K.Mey., Feddes Repert. 84: 463.
designated by German (2008: 9): LE; Isolectotype: 1973.
LE). Basionym: Thlaspi graecum Jord., Observ. Pl. Nouv. 3:
Distribution: W China, Kazakhstan, Kyrgyzstan, 30. 1846. TYPE: GREECE. [Peloponnes], Taygeto,
Mongolia, Tajikistan, Uzbekistan. Jul 1844, T. H. H. von Heldreich s.n. (Holotype: LY;
Meyer (2006e) listed as a holotype another specimen Isotypes: G, JE, W).
collected on 31 June 1901, but due to the presence at Heterotypic synonym: Thlaspi taygeteum Boiss., Diagn.
LE of more than one collection by Alexeenko, the above Pl. Orient. ser. 1, 8: 40. 1849. TYPE: GREECE.
lectotypification by German is adopted herein. The species [Peloponnes], Laconia, Taygetos, above Mistra, Apr
has recently been first recorded for Mongolia (German, 1842, E. Boissier s.n. (Holotype: G; Isotypes: JE, W).
2009) and Uzbekistan (Tojibaev, 2011). Distribution: Greece.
Noccaea griffithiana (Boiss.) F.K.Mey., Haussknechtia 12: Noccaea iranica, described and illustrated by Mozaffarian
170. 2006. (1996) as the later homonym Thlaspi pulvinatum, is
Basionym: Carpoceras griffithianum Boiss., Diagn. Pl. somewhat anomalous in Noccaea for being woody at base
Orient., ser. 2, 1: 40. 1854. TYPE: AFGHANISTAN. and for having non-auriculate cauline leaves and pyrifom-
Bharowul, W. Griffith 1473 (Holotype: G; Isotype: K). oblong fruits with truncate, minutely horned apex.
Homotypic synonym: Thlaspi griffithianum (Boiss.) Boiss.,
Fl. Orient. 1: 329. 1867. Noccaea jankae (Kerner) F.K.Mey., Feddes Repert. 84:
Distribution: Afghanistan, Pakistan. 462. 1973.
Basionym: Thlaspi jankae Kern., Oesterr. Bot. Z. 17: 35.
Noccaea haussknechtii (Boiss.) F.K.Mey., Feddes Repert. 1867. TYPE: SLOVAKIA. Nitra, zobor, 587 m,
84: 460. 1973. 28 May 1975, F. K. & J. Meyer 11461 (Neotype
Basionym: Thlaspi haussknechtii Boiss., Fl. Orient. 1: 330. designated by Meyer (2006e: 110): JE).
1867. TYPE: TURKEY. Taurus Cataonicus, Beryt Heterotypic synonyms: Thlaspi finitimum Dvořáková, Folia
Dagh, 9,000–10,000 ft [2,740–3,050 m], 10 Aug. Geobot. Phytotax. 8: 150. 1973. TYPE: SLOVAKIA.
1865, C. Haussknecht 1102 (Holotype: JE; Isotype: Nitra, Mt. Žibrica, ca. 500 m, 1 May 1936, V. Krist
G-BOIS). s.n. (Holotype: BRNU-276631).
Distribution: Turkey. Thlaspi hungaricum M.Dvorakova, Preslia 50: 15. 1978;
Noccaea hungarica (Dvořáková) Holub, Folia
Noccaea huber-morathii (F.K.Mey.) Al-Shehbaz, comb. Geobot. Phytotax. 18: 204. 1983. TYPE: HUNGARY.
nov. Budapest, 6 May 1911, S. Jávroka s.n. (Holotype: BP-
Basionym: Thlaspiceras huber-morathii F.K.Mey., 68164).
Feddes Repert. 84: 455. 1973. TYPE: TURKEY. Distribution: Hungary, Slovakia.
Prov. Erzincan, Dist. Refahiye, 1 km N Refahiye,
1,540–1,560 m, 2 Jul 1953, A. Huber-Morath 12974 Noccaea japonica (H.Boissieu) F.K.Mey., Haussknechtia
(Holotype: G). 12: 14. 2010.
Homotypic synonym: Thlaspi huber-morathii (F.K.Mey.) Basionym: Thlaspi japonicum H.Boissieu, Bull. Herb.
Greuter & Burdet, Willdenowia 13(1): 96. 1983. Boissier 7: 797. 1899. TYPE: JAPAN. Shakotan, 9 Jun
Heterotypic synonyms: Thlaspiceras huber-morathii 1893, U. J. Faurie 9908 (Lectotype here designated:
F.K.Mey. subsp. divergens F.K.Mey., Feddes Repert. P-01817547).
84: 455. 1973; Thlaspi huber-morathii subsp. divergens Distribution: Japan.
(F.K.Mey.) Greuter & Burdet, Willdenowia 13(1): 96. Although cited the two syntypes of Thlaspi japonicum,
1983. TYPE: TURKEY. Erzincan, Kara Dag, Rfahiye- Meyer (2010) did not designate a lectotype. I have examined
Erzincan, 27 km E Refahiye, 1,940–1,960 m, 2 Jul the lectotype above, and it is also available on the JSTOR
1953, A. Huber-Morath 12973 (Holotype: G). website (http://plants.jstor.org/).
Distribution: Turkey.
The differences between subsp. huber-morathii and Noccaea jaubertii (Hedge) Al-Shehbaz, comb. nov.
subsp. divergens are only quantitative in nature, and the latter Basionym: Thlaspi jaubertii Hedge, Notes Roy. Bot. Gard.
is somewhat bigger than the former. Both were collected Edinburgh 26: 185. 1965; based on Syrenopsis stylosa
from nearby localities, which make their independent status Jaub. & Spach, Ill. Pl. Orient. 1: 6. t. 3. 1842; non T.
rather doubtful. stylosum Rchb., Ic. Fl. Germ. 2, 3: t. 6. 1837–1838;
nec T. stilosum (Ten.) Mutel, Fl. Franç.1: 99. 1834
Noccaea iberidea (Boiss.) Al-Shehbaz & Menke, Taxon (as stylosum), nec Noccaea stilosa (Ten.) Rchb.,
61: 949. 2012. Fl. Germ. Excurs. 663. 1832 (as stylosa). TYPE:
Basionym: Eunomia iberidea Boiss., Ann. Sci. Nat., Bot., TURKEY: Bursa, in Olympo bithyno, Jul 1839,
ser. 2, 17: 191. 1842; Aethionema iberideum (Boiss.) Jaubert s.n. (Holotype: P?).
Boiss., Fl. Orient. 1: 351. 1867. TYPE: TURKEY. Distribution: Turkey.
Olympus Bithynus, P. M. R. Aucher-Eloy 345
(Holotype: G-BOIS; Isotypes: E, K, P). Noccaea kamtschatica (Karav.) Czerep., Sosud. Rast.
Distribution: Turkey. SSSR 140. 1981.
Basionym: Thlaspi kamtschaticum Karav., Sist.
Noccaea iranica Al-Shehbaz, nom. nov. Zametki Mater. Gerb. Krylova Tomsk. Gosud.
Replaced name: Thlaspi pulvinatum Mozaff., Iranian J. Univ. Kuybysheva 83: 4. 1965. TYPE: RUSSIA.
Bot. 7: 138. 1996, non T. pulvinatum (F.K.Mey.) Kamtschatka, ad limitem superiorem fluvium Pankara,
Greuter & Burdet, Willdenowia 13: 96. 1983. TYPE: 17 Aug 1930, P. Novograblenov 2533 (Lectotype
IRAN. Azarbaijan, between Shhindege and Takab, designated by Meyer (2010: 16): MW: Isolectotypes:
kuh-e Gharadash (Aghdash), from Ghezghapan and LE, MW).
Auchdare, 2,500–2,800 m, 4 Jul 1991, V. Mozaffarian Homotypic synonym: Noccaeopsis kamtschatica (Karav.)
69874 (Holotype: TARI, n.v.) F.K.Mey., Haussknechtia 12: 16. 2010.
Distribution: Iran. Distribution: Russia.
Meyer (2010) placed the species in his monospecific Noccaea limosellifolia (Reut. ex Burnat) F.K.Mey., Feddes
Noccaeopsis because it has silique fruits with length: width Repert. 84: 460. 1973.
ratio more than 3:1 instead of silicles with a ratio < 3:1, Basionym: Thlaspi rotundifolium (L.) Gaudin var.
a feature characteristic of the vast majority of Noccaea limosellifolium Reut. ex Burnat, Fl. Alp. Marit. 1:
species. Similar fruit types occur in numerous genera of 142. 1892. TYPE: France. Col de la Madone delle
the Brassicaceae and sometimes within a given species. Fenestre, 13 Aug 1852, G. Reuter (Holotype: G).
Therefore, such a difference in the fruit type is usually Distribution: France, Italy.
unreliable when used alone in the recognition of genera in
the family. Noccaea lutescens (Velen.) F.K.Mey., Feddes Repert. 84:
462. 1973.
Noccaea kovatsii (Heuff.) F.K.Mey., Feddes Repert. 84: Basionym: Thlaspi lutescens Velen., Sitzungsber. Königl.
461. 1973. Böhm. Ges. Wiss. 1903(28): 2. 1903. TYPE: GREECE.
Basionym: Thlaspi kovatsii Heuff., Flora 36: 624. 1853. Thessaloniki, Panórama, 350 m, F. K. & J. Meyer
TYPE: ROMANIA. Mt. Bagys ad Lunkány, May 21887 (Neotype designated by Meyer (2006e: 118): JE).
1832, Heuffel s.n. (Holotype: BP). Heterotypic synonym: Thlaspi lutescens subsp. drenorskii
Heterotypic synonyms: Thlaspi affine Schott & Kotschy ex F.K.Mey., Feddes Repert. 84: 462. 1973. TYPE:
Boiss., Fl. Orient. 1: 327. 1867; T. alpinum Crantz var. BULGARIA. Alibotush, 1,600 m, 6 Jun 1933, A. K.
affine (Schott & Kotschy ex Boiss.) Stoj. & Stef., Fl. Drenovski s.n. (Holotype: SOM).
Balg. 1: 491. 1924; T. kovatsii var. affine (Schott & Distribution: Bulgaria, Greece, Macedonia.
Kotschy ex Boiss.) Nyár. In Savulescu, Fl. Rep. Pop.
Romine 3: 403. 1955. TYPE: ROMANIA. Krajuluj, Noccaea maassoumii (Mozaff.) Al-Shehbaz, comb. nov.
6,000 ft [1,828 m], 24 Jun 1850, C. G. T. Kotschy 375 Basionym: Thlaspi maassoumii Mozaff., Iranian J. Bot. 7:
(Lectotype here designated: B). 135. 1996. TYPE: IRAN. Mazandaran, S. of Ramsar,
Thlaspi longiracemosum Schur, Oesterr. Bot. Z. 8: 22. Siemam Mt., 3,600 m, Runemark & Maassoumi
1858. TYPE: ROMANIA. Near Corona, Schur s.n. 21825 (Holotype: TARI, n.v.).
(Lectotype designated by Meyer (2006e: 85): WU). Distribution: Iran.
Although the type collection of the species was not
Thlaspi pawlowskii Dvořáková, Preslia 45: 315. 1973; examined, the illustration and description given by
Noccaea pawlowskii (Dvořáková) Holub, Preslia 70: Mozaffarian (1996) clearly place it in Noccaea, though in
108. 1998. TYPE: ROMANIA. Suliguł, 2 Jun 1936, somewhat isolated position due to having woody lower stems.
B. Pawlowski s.n. (Holotype: KRA).
Distribution: Hungary, Romania. Noccaea macrantha (Lipsky) F.K.Mey., Feddes Repert.
84: 464. 1973.
Noccaea kurdica (Hedge) Al-Shehbaz, comb. nov. Basionym: Thlaspi praecox Wulfen var. macranthum
Basionym: Thlaspi kurdicum Hedge, Notes Roy. Bot. Gard. Lipsky, Zap. Kievsk. Obsc. Estestvoip. 13: 271. 1894.
Edinburgh 23: 547. 1961. TYPE: TURKEY. Prov. TYPE: RUSSIA. Tshernomor okr., Novorossijsk, 20
Van, Distr. Gevaṣ, Artas Dag, 11,500 ft [3,500 m], 16 Apr 1889, V. Lipsky s.n. (Lectotype designated by
Jul 1954, P. H. Davis & O. Polunin 22806 (Holotype: Dorofeyev (2003: 121): LE).
E; Isotypes: BM, JE, K). Homotypic synonym: Thlaspi macranthum (Lipsky)
Homotypic synonym: Vania kurdica (Hedge) F.K.Mey., N.Busch, Trudy Bot. Sada Imp. Yur’evsk. Univ.
Feddes Repert. 84: 467. 1973; Apterigia kurdica 7: 142. 1906.
(Hedge) V.I.Dorof., Novosti Sist. Vyssh. Rast. 44: 94. Distribution: Russia, Ukraine.
2013.
Distribution: Iraq, Turkey. Noccaea magellanica (Comm. ex Poir.) Holub, Preslia 70:
108. 1998.
Noccaea libanotica F.K.Mey., Feddes Repert. 84: 466. Basionym: Thlaspi magellanicum Comm. ex Poir., Encycl.
1973. TYPE: LEBANON. Sanin, 1,800 m, 17 Jun 1897, J. 7: 541. 1806. TYPE: [Chile or Argentina] Magallan,
Bornmüller 92 (Holotype: JE-00004194). 1767, P. Commerson s.n. (Holotype: P-JUSS; Isotype:
Homotypic synonym: Thlaspi libanoticum (F.K.Mey.) BAA!).
Greuter & Burdet, Willdenowia 13: 96. 1983. Distribution: Patagonian Argentina and Chile.
Distribution: Lebanon, Syria. For a complete synonymy and typification, see Al-
Shehbaz (2012b)
Noccaea lilacina (Boiss. & A. Huet) Al-Shehbaz, comb. nov.
Basionym: Thlaspi lilacinum Boiss. & A.Huet, Diagn. Noccaea mexicana (Standl.) Holub, Preslia 70: 108. 1998.
Pl. Orient. ser. 2, 5: 41. 1856. TYPE: TURKEY [as Basionym: Thlaspi mexicanum Standl., Publ. Field Mus.
Armenia]. Inter Gumuchchané et Zazalarchane, May Nat. Hist., Bot. Ser. 22: 76. 1940. TYPE: MEXICO.
1843, A. Huet du Pavillion s.n. (Holotype: G-BOIS). Peak of Cerro Potosi, Municipio de Galeana, 21 Jul
Homotypic synonym: Callothlaspi lilacinum (Boiss. & 1935, C. H. Müller 2244 (Holotype: F; Isotypes: GH,
A.Huet) F.K.Mey., Feddes Repert. 84: 457. 1973. MICH, MO).
Distribution: Turkey. Distribution: Mexico.
Noccaea meyeri Al-Shehbaz, nom. nov. Contrary to the North American literature (e.g., Holmgren,
Replaced name: Thlaspiceras crassifolium Hub.-Mor. 1971; Rollins, 1993; Holmgren, 2005), Koch and Al-Shehbaz
& F.K.Mey., Feddes Repert. 84: 454. 1973, non (2004) and Meyer (2006e, 2010) amply demonstrated (as
Noccaea crassifolia Rouy & Foucaud, Fl. France 2: Thlaspi or Noccaea) that the native American N. fendleri is
91 et 92. 1895. TYPE: TURKEY. Prov. Seyhan, Distr. unrelated molecularly, morphologically, and geographically
Osmaniye, Yaglipinar S of Yarpus, 1,350–1,400 m, 2 to the central European N. montana. The taxonomy of the
Jul 1959, A. Huber-Morath 16241 (Holotype: G). North American plants has recently been corrected byAl-
Homotypic synonym: Thlaspi crassifolium (Hub.-Mor. & Shehbaz (2010) and above (see under N. fendleri).
F.K.Mey.) Greuter & Burdet, Willdenowia 13: 95.
1983. Noccaea natolica (Boiss.) Al-Shehbaz, comb. nov.
Distribution: Turkey. Basionym: Thlaspi natolicum Boiss., Ann. Sci. Nat., Bot.
The transfer of Thlaspiceras crassifolium to Noccaea sér. 2, 17: 180. 1842. TYPE: Turkey, P. M. R.
would have created a later homonym. Therefore, the new Aucher-Eloy 4151 (Lectotype designated by Meyer
name N. meyeri is proposed in honor of Dr. Friedrich Karl (2003a: 7): G-BOIS; Isolectotypes: G, P, W).
Meyer (1926–2013) who worked extensively on Thlaspi s.l. Homotypic synonyms: Thlaspi perfoliatum L. var. natolicum
and its dozen segregates, including Noccaea. (Boiss.) Post, Fl. Syr. Pal. Sin. 94. 1896; Microthlaspi
natolicum (Boiss.) F.K.Mey., Feddes Repert. 84: 452.
Noccaea microphylla (Boiss. & Orph.) F.K.Mey., Feddes 1973.
Repert. 84: 460. 1973. Heterotypic synonyms: Microthlaspi natolicum (Boiss.)
Basionym: Thlaspi microphyllum Boiss. & Orph., Diagn. F.K.Mey. var. drabiflorum (Fenzl) F.K.Mey., Feddes
Pl. Orient. ser. 2, 6: 19. 1859. TYPE: GREECE. Mt. Repert. 84: 453. 1973; Thlaspi drabiflorum Fenzl,
Parnass, Carcaria, T. G. Orphanides s.n. (Lectotype Pug. Pl. Nov. Syr. 14. 1842. TYPE: SYRIA. Prope
designated by Meyer (2006e: 65): G-BOIS; Suedia, C. G. T. Kotschy s.n. (Holotype: W).
Isolectotype: JE). Microthlaspi natolicum (Boiss.) F.K.Mey. subsp.
Distribution: Albania, Greece, Macedonia. gaillardotii F.K.Mey., Feddes Repert. 84: 452. 1973;
Thlaspi perfoliatum L. subsp. gaillardotii (F.K.Mey.)
Noccaea microstyla (Boiss.) F.K.Mey., Feddes Repert. 84: Greuter & Burdet, Willdenowia 13(1): 96. 1983.
466. 1973. TYPE: LEBANON, “Champs calcaires au pied du
Basionym: Thlaspi microstylum Boiss., Diagn. Pl. Orient. Liban près Saida,” C. Gaillardot s.n. (Holotype: JE).
ser. 1, 8: 38. 1849. TYPE: TURKEY. Cassius, May– Thlaspi perfoliatum L. var. longistylum Post, Fl. Syr.
Jul 1846, E. Boissier s.n. (Holotype: G-BOIS). Pal. Sin. 94. 1896; Microthlaspi natolicum (Boiss.)
Distribution: Syria, Turkey. F.K.Mey. subsp. longistylum (Post) F.K.Mey., Feddes
Repert. 84: 453. 1973; T. annuum K.Koch var.
Noccaea minima (Ard.) F.K.Mey., Feddes Repert. 84: 459. longistylum (Post) Dinsmore in Post, Fl. Syr. Pal. Sin.,
1973. ed. 2, 1: 99. 1932. TYPE: SYRIA. Shuway in Jab.
Basionym: Thlaspi minimum Ard., Animadv. Bot. Spec. Alt. Nusayriyyah, G. E. Post s.n. (Lectotype designated by
33. 1764. TYPE: ITALY. [Belluno], Mt. del Vette, Meyer (2003a: 12): BEI).
Arduino s.n. (Lectotype designated by Meyer (2006e:
Microthlaspi natolicum (Boiss.) F.K.Mey. var.
44): LINN 827.5).
pentadactylonis F.K.Mey., Feddes Repert. 84: 453.
Heterotypic synonym: Thlaspi kerneri Huter, Oesterr. Bot.
1973. TYPE: CYPRUS, Petacactylo, C. G. T. Kotschy
Z. 24: 82. 1874. TYPE: ITALY. Mt. Raut, Huter 587
367 (Holotype: W).
(Lectotype designated by Meyer (2006e: 46): WU;
Isolectotypes: HBG, JE, LI, W). Microthlaspi natolicum (Boiss.) F.K.Mey. subsp.
Distribution: Austria, Italy. sporadium F.K.Mey., Feddes Repert. 84: 453. 1973.
TYPE: GREECE, “in umbrosis Rhodi,” Mar 1845,
Noccaea montana (L.) F.K.Mey., Feddes Repert. 84: 461. T.T.H. Von Heldreich s.n. (Holotype: JE; Isotypes: E, W).
1973. Distribution: Cyprus, Greece, Lebanon, Syria, Turkey.
Basionym: Thlaspi montanum L., Sp. Pl. 2: 747. 1753. Several authors (e.g., Greuter & Burdet in Greuter &
TYPE: “Habitat in Helvetiae, Austriae, Italiae, Raus, 1983; Greuter et al., 1986; Artelari, 2002) have
Monspelii petrosis.” (Lectotype desinganted by confused the limits of this species and Noccaea perfoliata
Marhold & Mártonfi (2001: 189), Herb. Linn. No. (both as Thlaspi) and recognized some of the infraspecific
825: 10, LINN). taxa above as part of the latter species. The flowers of N.
Distribution: Austria, Belgium, Czech Republic, France, natolica have larger and unequal petals, whereas those of
Germany, Hungary, Slovakia, Switzerland, N. perfoliata have smaller and equal petals. Meyer (1973,
Mayer (2006e) listed some two dozen synonyms under 2003a) over divided N. natolica (as Microthlaspi) into
the species and gave full citation and typification. These are four subspecies and two additional varieties based almost
not repeated here, and the interested reader should consult solely on microscopic differences in seed size, style length,
that reference. and petal size. This excessive division of the species into
infraspecific taxa is impractical and not useful. Therefore, Noccaea oppositifolia (Pers.) Al-Shehbaz & Menke, Taxon
the five infraspecific taxa above are considered herein as 61: 949. 2012.
only minor variants of the species. Basionym: Iberis oppositifolia Pers., Syn. Pl. 2: 186.
1806. TYPE: LEBANON. J.J. H. Labillardière s.n.
Noccaea nepalensis Al-Shehbaz, Adansonia sér. 3, 24: 89. (Holotype: G).
2002. TYPE: NEPAL. Dozam Khola, near Simikot, 10,500 Homotypic synonyms: Lepidium oppositifolium
ft. [3,200 m], 29 May 1952, O. V. Polunin, W. R. Sykes & L. (Pers.) Labill., Icon. Pl. Syr. 5: 14. 1812; Thlaspi
H. J. Williams 4199 (Holotype: BM; Isotypes: A, E). oppositifolium (Pers.) Poir. in Lamarck, Encycl. Suppl.
Distribution: Nepal. 5: 277. 1817; Eunomia oppositifolia (Pers.) DC., Syst.
Nat. 2: 556. 1821; Aethionema oppositifolium (Pers.)
Noccaea nevadensis (Boiss. & Reut.) F.K.Mey., Feddes Boiss., Fl. Orient. 1: 344. 1867.
Repert. 84: 460. 1973. Distribution: Lebanon, Turkey.
Basionym: Thlaspi nevadense Boiss. & Reut., Pugill. Pl.
Afr. Bor. Hispan. 11. 1852. TYPE: SPAIN. Sierra Noccaea orbiculata (Steven ex DC.) Al-Shehbaz, comb. nov.
Nevada, below Cueva de Panderon, Jul 1849, Reuter Basionym: Thlaspi orbiculatum Steven ex DC., Syst. Nat.
s.n. (Holotype: G). 2: 377. 1821. TYPE: GEORGIA. Iberia, 1820, C.
Distribution: Spain. Steven s.n. (Holotype: G-DC; Isotype: LE).
Homotypic synonym: Neurotropis orbiculata (Steven ex
Noccaea occitanica (Jord.) F.K.Mey., Feddes Repert. 84: DC.) F.K.Mey., Feddes Repert. 84: 452. 1973.
463. 1973. Distribution: Georgia, Turkey.
Basionym: Thlaspi occitanicum Jord., Observ. Pl. Nouv. 3:
12. 1846. TYPE: FRANCE. La Séranne près Ganges, Noccaea oxyceras (Boiss.) Al-Shehbaz, comb. nov.
A. Jordan s.n. (Holotype: LY). Basionym: Carpoceras oxyceras Boiss., Diagn. Pl. Orient.
Homotypic synonyms: Thlaspi sylvestre Jord. subsp. ser. 1, 8: 37. 1849. TYPE: TURKEY. Cassius, Jun
occitanicum (Jord.) Rouy & Foucaud, Fl. France 2: 1846, E. Boissier 51 (Holotype: G-BOIS).
151. 1895; T. alpestre L. subsp. occitanicum (Jord.) Homotypic synonyms: Thlaspi oxyceras (Boiss.) Hedge, Fl.
O.Bolòs & Vigo, Butl. Inst. Catalana Hist. Nat., Turkey 1: 339. 1965; Thlaspiceras oxyceras (Boiss.)
Secc. Bot. 38(1): 77. 1974; T. caerulescens J.Presl F.K.Mey., Feddes Repert. 84: 454. 1973.
& C.Presl subsp. occitanicum (Jord.) M.Laínz, Bol. Distribution: Turkey.
Soc. Brot. sér. 2, 53: 38. 1979; Noccaea caerulescens
(J.Presl & C.Presl) F.K.Mey. subsp. occitanicum Noccaea papillosa (Boiss.) F.K.Mey., Feddes Repert. 84:
(Jord.) Kerguélen, Collect. Patrim. Nat. 8: 14. 1993. 460. 1973.
Heterotypic synonym: Thlaspi tallonis Sennen, Plantes Basionym: Thlaspi papillosum Boiss., Fl. Orient. 1: 330.
d’Espagne nr. 6039. 1927; Thlaspi alpestre L. var. 1867. TYPE: TURKEY. [Prov. Denizli], Cadmus,
tallonis (Senn.) O.Bolòs & Vigo, Butl. Inst. Catalana Dewileh, Jun 1942, E. Boissier s.n. (Holotype:
Hist. Nat., Secc. Bot. 38: 77. 1974; Noccaea occitanica G-BOIS).
subsp. tallonis (Sennen) F.K.Mey., Feddes Repert. 84: Homotypic synonym: Apterigia papillosa (Boiss.)
463. 1973. TYPE: SPAIN. Gerona, Llivia, Néguila, V.I.Dorof., Novosti Sist. Vyssh. Rast. 44: 94. 2013.
1,270 m, 24 Jun 1927, F. Sennen 6039 (Holotype: BC; Distribution: Turkey.
Isotype: W).
Distribution: France, Spain. Noccaea papyracea (Boiss.) Khosravi, Mumm. &
Mohsenz., Feddes Repert. 119: 21. 2008.
Noccaea ochroleuca (Boiss. & Heldr.) F.K.Mey., Feddes Basionym: Brossardia papyracea Boiss., Ann. Sci. Nat.,
Repert. 84: 461. 1973. Bot. sér. 1, 16: 380. 1841. TYPE: IRAN. Ispahan,
Basionym: Thlaspi ochroleucum Boiss. & Heldr., Diagn. P. M. R. Aucher-Eloy 4127 (Holotype: G-BOIS;
Pl. Orient. ser. 1, 8: 39. 1849. TYPE: TURKEY. Prov. Isotypes: K, 2P).
Isparta, Mons Davros Dag Pisidiae, May 1845, T. H. Heterotypic synonyms: Brossardia papyracea Boiss.
H. von Heldreich s.n. (Lectotype designated initially var. kotschyi Boiss., Diagn. Pl. sér. 1, 8: 41. 1849;
as type by Hedge (1965: 335) and finalized by Meyer Brossardia retusa Boiss., Fl. Orient. 1: 336. 1867.
(2006e: 95): G-BOIS; Isolectotypes: BM, G, GOET, TYPE: IRAN. Kuhdelu, 3 May 1842, C. G. T.
JE, K, P, W). Kotschy 327 (Lectotype here designated: G-BOIS;
Homotypic synonyms: Thlaspi alpinum Crantz var. Isolectotypes: B, E, K, US).
ochroleucum (Boiss. & Heldr.) Stoj. & Stef., Fl. Balg. Distribution: Iran, Iraq, Turkey.
1: 491. 1924; T. praecox Wulfen var. ochroleucum Both Brossardia retusa and B. papyracea var. kotschyi
(Boiss. & Heldr.) Stoj. & Stef., Fl. Balg., ed. 2, 1: 448. are based on the same mixed collection. The species has only
1933. recently been recorded for Turkey (Adigüzel & Koyuncu,
Distribution: Turkey. 1999) and placed in Noccaea by Khosravi et al. (2007).
Noccaea parviflora (A.Nelson) Holub, Preslia 70: 108. Heterotypic synonyms: Thlaspi kotschyanum Boiss.
1998. & Hohen., Diagn. Pl. Orient. ser. 1, 8: 39. 1849;
Basionym: Thlaspi parviflorum A.Nelson, Bull. Torrey Neurotropis kotschyana (Boiss. & Hohen.) Czerep.,
Bot. Club 27: 265. 1900. TYPE: U.S.A. Wyoming, Sosud. Rast. SSSR 140. 1981. TYPE: IRAN.
Yellowstone National Park, Golden Gate, 28 Jun Elburs, Passgala, 6 May 1843, C. G. T. Kotschy 102
1899, A. & E. Nelson 5554 (Holotype: RM; Isotypes: (Holotype: G-BOIS; Isotypes: B, BM, GOET, H, JE,
CM, DS, GH, 2ISC, MO, 2NY, P, US). K, 3P, W, WAG).
Distribution: U.S.A. (Idaho, Montana, Wyoming). Thlaspi brevicaule Boiss. ex Kotschy, Diagn. Pl.
Orient. ser. 2, 5: 40. 1856. TYPE: SYRIA. “Hab. in
Noccaea perfoliata (L.) Al-Shehbaz, comb. nov. subalpinis Bludan suprá Zebdani Antilibani,” near
Basionym: Thlaspi perfoliatum L., Sp. Pl. 2: 646. 1753. Damascus, 6,000 ft [1,829 m], 6 Jun 1855, C. G. T.
TYPE: “Habitat in Germaniae, Helvetiae, Galliae Kotschy 47 (Holotype: G-BOIS; Isotypes: B, GZU,
apricis, agris” (Lectotype designated by Jafri (1973: JE, K, 3P, S, W).
85) as type: Herb. Linn. No. 825.9 (LINN); same Thlaspi cardiocarpum Hook.f. & Thomson, J. Proc.
lectotypification was proposed thirty years later by Linn. Soc., Bot. 5: 176. 1861. TYPE: AFGHANISTAN.
Meyer (2003a: 17) who overlooked the earlier one). 1838–1840, Griffith 1402 (Lectotype designated by
Homotypic synonym: Microthlaspi perfoliatum (L.) Meyer (2001b: 49): K; isolectotype: W).
F.K.Mey., Feddes Repert. 84: 453. 1973.
Heterotypic synonyms: Thlaspi granatense Boiss. & Reut., Thlaspi armenum N.Busch, Trudy Bot. Sada Imp.
Diagn. Pl. Orient. ser. 2, 1: 40. 1854; Microthlaspi Yur’evsk. Univ. 7: 142. 1906; Neurotropis armena
granatense (Boiss. & Reut.) F.K.Mey., Feddes Repert. (N.Busch) Czerep., Sosud. Rast. SSSR 140. 1981.
84: 453. 1973. TYPE: SPAIN. In regione alpina TYPE: AZERBAIJAN. Prop Nachiczevan, Szovits
superiori montis Sierra de Baza, 21 May 1851, E. s.n. (Holotype: LE).
Bourgeau 1025 (Holotype: G-BOIS; Isotype: WAG). Distribution: Afghanistan, Azerbaijan, China, Greece,
Distribution: Native to S and C Europe, NW Africa, and Iran, Iraq, Kazakhstan Lebanon, Pakistan, Syria, Tajikistann,
SW Asia eastward into W China; naturalized in NC Europe Turkey, Turkmenistan, Uzbekistan.
and North America. The species has recently been recorded for China by
This is an extremely widespread and morphologically German and Chen (2009).
variable species that includes several chromosomal races
(Koch, 1995; Koch et al., 1998a, b; Koch & Bernhardt, Noccaea praecox (Wulfen) F.K.Mey., Feddes Repert. 84:
2004). Several dozens of species and infraspecific taxa were 462. 1973.
described in this complex, but I follow Meyer (2003a), who Basionym: Thlaspi praecox Wulfen in Jacqin, Coll. 2:
listed and typified all of them, in treating them as minor 124. 1789. TYPE: “Thlaspi montanum praecox,”
variants of the species. The variation does not seem to fall (Lectotype designated by Meyer (2006e: W)
into geographically and morphologically defined groups, Heterotypic synonyms: Thlaspi cuneifolium Griseb.,
and the division of the species into infraspecific or specific Oesterr. Bot. Z. 23: 268. 1873; Noccaea cuneifolia
segregates would not be practical at this point. Therefore, a (Griseb.) Holub, Preslia 70: 108. 1998. TYPE:
broad species concept is adopted herein. The synonymy at MONTENEGRO. Bnjela Gora, Jastrebica, 27 Apr
the specific and infraspecific levels are extensive and need 1872, J. Pantocsek s.n. (Lectotype designated by
not be repeated here, and only those at the specific rank are Meyer (2006e: 102): GOET-002665; Isolectotypes:
listed. B, DR, G, JE, W).
Distribution: Albania, Austria, Bosnia and Herzegovina,
Noccaea phrygia (Bornm.) F.K.Mey., Feddes Repert. 84: Croatia, Italy, Montenegro, Serbia, Slovenia.
465. 1973. Meyer (2006e) listed a dozen additional synonyms (all in
Basionym: Thlaspi phrygium Bornm., Mitth. Thüring. Bot. Thlaspi) that are not repeated herein.
Vereins 38: 57. 1929. TYPE: TURKEY. Phrygia, Mt.
Sultandagh, 1,850 m, 16 Jun 1899, J. Bornmüller Noccaea pulvinata (F.K.Mey.) Al-Shehbaz, comb. nov.
4093 (Holotype: B). Basionym: Vania pulvinata F.K.Mey., Feddes Repert. 84:
Distribution: Turkey. 467. 1973. TYPE: TUREKY. Prov. Hakkari, Cilo Dag,
10 km W of Cilo Tepe, 12,000 ft [3,660 m], 9 Aug.
Noccaea platycarpa (Fisch. & C.A.Mey.) Al-Shehbaz, 1954, P. H. Davis & O. V. Polunin 24207 (Holotype:
comb. nov. E; Isotype: JE).
Basionym: Thlaspi platycarpum Fisch. & C.A.Mey., Index Homotypic synonym: Thlaspi pulvinatum (F.K.Mey.)
Seminum Hort. Petrop. 7. 57. 1840. TYPE: TURKEY. Greuter & Burdet, Willdenowia 13: 96. 1983, non T.
Grown at LE from seeds collected by Wiedemann s.n. pulvinatum Mozaff., Iranian J. Bot. 7: 138. 1996.
(Lectotype designated by Dorofeyev (2012: 456): LE; Distribution: Turkey.
Isolectotype: K).
Homotypic synonym: Neurotropis platycarpa (Fisch. & Noccaea pumila (Steven) Steud., Nomencl. Bot., ed. 2. 2:
C.A.Mey.) F.K.Mey., Feddes Repert. 84: 452. 1973. 197. 1841.
Basionym: Iberis pumila Steven, Mém. Soc. Imp. 1983; T. cepaeifolium (Wulfen) W.D.J.Koch subsp.
Naturalistes Moscou 3: 269. 1812. TYPE: grignense (F.K.Mey.) Greuter & Burdet, Willdenowia
AZERBAIJAN. Schahdagh, Jun 1810, Steven s.n. 15: 70. 1985; T. grignense (F.K.Mey.) Landolt, Fl.
(Lectotype designated by Dorofeyev (2012: 459): H). Indicativa 270. 2010. TYPE: ITALY. Grigna-Gruppe
Homotypic synonyms: Thlaspi pumilum (Steven) Ledeb., am Comer See, ca. 2,000 m, 12 Jul 1904, G. Geilinger
Fl. Ross. 1(1): 164. 1841; Apterigia pumila (Steven) s.n. (Holotype: Z).
Galushko, Novosti Sist. Vyss. Rast. 6: 209. 1970. Noccaea rotundifolia subsp. intermedia F.K.Mey.,
Distribution: Azerbaijan, Georgia. Feddes Repert. 84: 459. 1973. TYPE: FRANCE.
Beuil au Mont Monnier, 20 Jun and 4 Aug 1894, G. P.
Noccaea rechingeri (F.K.Mey) Al-Shehbaz, comb. nov. Vidal s.n. (Holotype: JE).
Basionym: Thlaspiceras rechingeri F.K.Mey., Feddes Distribution: Austria, France, Germany, Italy, Slovenia,
Repert. 84: 456. 1973. TYPE: TURKEY. Prov. Hatay, Switzerland
Akma Dag, 3,600 ft [1,097m], Jun 1862, (Pl. Syriae Mayer (2006e) gave an extensive list of 16 synonyms
bor. Ex Aano occidental supra Arsus), C. G. T. Kotschy that are not repeated herein, and only the above additional
181 (Holotype: W; Isotypes: G, JE). synonyms are given. The two subspecies of Noccaea
Homotypic synonym: Thlaspi rechingeri (F.K.Mey.) rotundifolia described by Meyer (1973) differ only slightly
Greuter & Burdet, Willdenowia 13: 96. 1983. in floral measurements and style length.
Distribution: Turkey.
Noccaea rubescens (Schott & Kotschy ex Boiss.) F.K.Mey.,
Noccaea rhodopensis F.K.Mey., Feddes Repert. 84: 462. Feddes Repert. 84: 459. 1973.
1973. TYPE: BULGARIA. Between Bačkovo and Kuru- Basionym: Aethionema rubescens Schott & Kotschy ex
dere, 2 May 1967, F. K. & J. Meyer 8718 (Holotype: JE; Boiss., Fl. Orient. 1: 343. 1867. TYPE: TURKEY.
Isotype: JE). Iter cilicicum in Tauri alpes, Bulgar Dagh, summit of
Distribution: Bulgaria. Metdesis, 11,800 ft [3,597 m], 3 Jul 1853, C. G. T.
Kotschy 160 (Holotype: G-BOIS).
Noccaea rostrata (N.Busch) Al-Shehbaz, comb. nov. Homotypic synonym: Thlaspi rubescens (Boiss.) Greuter &
Basionym: Thlaspi rostratum N.Busch, Monit. Jard. Bot. Burdet, Willdenowia 13: 96. 1983.
Tiflis, n.s. 1926–27(3–4): 10. 1927. TYPE: AZERBAIJAN. Heterotypic synonyms: Thlaspi crassum P.H.Davis, Fl.
Nachitshevan, Ordubad, 26 May 1923, A. A. Grossheim s.n. Turkey 10: 41. 1988; T. sintenisii Hausskn. ex Bornm.
(Holotype: LE). subsp. crassum (P.H.Davis) Parolly, Willdenowia 25:
Homotypic synonym: Apterigia rostrata (N.Busch) 248. 1995. TYPE: TURKEY. Niğde, Ala Dağlari, 29
Galushko, Novosti Sist. Vyssh. Rast. 6: 209. 1970; Aug 1965, G. W. Findlay 163 (Holotype: E).
Atropatenia rostrata (N.Busch) F.K.Mey., Feddes Distribution: Turkey.
Repert. 84: 467. 1973.
Distribution: Azerbaijan. Noccaea salisii (Brügger) F.K.Mey., Feddes Repert. 84:
464. 1973.
Noccaea rosularis (Boiss. & Balansa) Al-Shehbaz, comb. Basionym: Thlaspi salisii Brügger, Z. Ferdinandeums
nov. Tirol, ser. 3, 9: 45. 1860. TYPE: SWITZERLAND.
Basionym: Thlaspi rosulare Boiss. & Balansa, Diagn. Silvaplana, Brügger s.n. (Lectotype designated by
Pl. Orient. ser. 2, 5: 39. 1856. TYPE: TURKEY. Meyer (2006e: 155): CHUR).
[Niğde], “in regione montana superiore jugi montis Distribution: Austria, Italy, Switzerland.
Masmeneudagh,” 8 Aug 1855, B. Balansa s.n.
(Holotype: G-BOIS; Isotypes: GH, K, 2P, US). Noccaea sarmatica F.K.Mey., Feddes Repert. 84: 464.
Homotypic synonyms: Masmenia rosularis (Boiss. & 1973. TYPE: MOLDOVA. Dshamano, Bulbokskogo
Balansa) F.K.Mey., Feddes Repert. 84: 468. 1973. rajonna, 25 Apr 1950, V. N. Andrejev & L. I. Posharinkaja
Distribution: Turkey. s.n. (Holotype: LE).
Homotypic synonym: Thlaspi sarmaticum (F.K.Mey.)
Noccaea rotundifolia (L.) Moench, Suppl. Meth. 89. 1802. V.I.Dorof., Turczaninowia 5(3): 113. 2002.
Basionym: Iberis rotundifolia L., Sp. Pl. 2: 649. 1753. Distribution: Moldova, Ukraine.
TYPE: “Habitat in Helvetica,” not yet designated
(Jarvis, 2007). It is likely that Herb. Linn. No. 827.7 Noccaea sintenisii (Hausskn. ex Bornm.) F.K.Mey., Feddes
(LINN) can serve as the lectotype, but without Repert. 84: 458. 1973.
examining that collection, a formal lectotypification Basionym: Thlaspi sintenisii Hausskn. ex Bornm., Mitth.
will have to wait. Thüring. Bot. Vereins 38: 52. 1929. TYPE: TURKEY.
Heterotypic synonyms: Noccaea rotundifolia subsp. Prov. Pontus, Karagoelldagh, Mt. Manator, 3 Aug
grignensis F.K.Mey., Feddes Repert. 84: 459. 1973; 1894, P. E. E. Sintenis 7316 (Holotype: JE-00004211;
Thlaspi rotundifolium (L.) Gaudin subsp. grignense Isotypes: B, BM, DS, FR, JE, 2K, 3LD, S, 2TBI,
(F.K.Mey) Greuter & Burdet, Willdenowia 13: 96. W, Z).
Homotypic synonym: Apterigia sintenisii (Hausskn. ex 553. 1979; T. alpestre Jacq. subsp. sylvium (Gaudin)
Bornm.) V.I.Dorof., Novosti Sist. Vyssh. Rast. 44: 94- Kerguélen, Lejeunia 120: 175. 1987; Noccaea
95. 2013. alpestris (Jacq.) Kerguélen subsp. sylvium (Gaudin)
Distribution: Turkey. Kerguélen, Collect. Patrim. Nat. 8: 14. 1993.
Distribution: France, Italy, Switzerland.
Noccaea stenocarpa (Boiss.) Al-Shehbaz, comb. nov.
Basionym: Carpoceras stenocarpum Boiss., Diagn. Pl. Noccaea szowitsiana (Boiss.) Al-Shehbaz, comb. nov.
Orient. ser. 1, 8: 37. 1849. TYPE: IRAN, near Tehran, Basionym: Thlaspi szowitsianum Boiss., Fl. Orient. 1: 324.
Mt. Totschal, 17 May 1843, C. G. T. Kotschy 165 1867. TYPE: ARMENIA. “Hab. in humidis montis
(Holotype: G-BOIS; Isotypes: B, GOET, JE, K, 4P, Karagiol prov. Karabagh,” 1 Jun 1829, Szovits 295
W). (Lectotype here designated: G-BOIS; Isotypes: 2LE).
Homotypic synonyms: Kotschyella stenocarpa (Boiss.) Homotypic synonym: Neurotropis szowitsiana (Boiss.)
F.K.Mey., Feddes Repert. 84: 457. 1973; Thlaspi F.K.Mey., Feddes Repert. 84: 452. 1973.
stenocarpum (Boiss.) Hedge, Fl. Iranica 57: 116. Distribution: Armenia, Azerbaijan.
1968. Meyer (2001b) indicated that the type is at G-BOIS,
Distribution: Iran, Turkmenistan. whereas Dorofeyev (2003: 123) designated one of the two
sheets at LE as the lectotype. Both authors did not meet the
Noccaea stenoptera (Boiss. & Reut.) F.K.Mey., Feddes requirements of lectotypification in Article 7.10 of ICN in
Repert. 84: 461. 1973. McNeill et al. (2012) and, therefore, a lectotypification of
Basionym: Thlaspi stenopterum Boiss. & Reut., Diagn. Pl. the species is needed.
Orient. ser. 1, 8: 40. 1849, non Conrath & Freyn, Bull.
Herb. Boissier 3(1): 38. 1895. TYPE: SPAIN. Sierra Noccaea tatianae (Bordz.) F.K.Mey., Feddes Repert. 84:
de Guadarrama Castellae, above El Paular, 1841, G. F. 465. 1973.
Reuter s.n. (Holotype: G). Basionym: Thlaspi tatianae Bordz., Izv. Kievsk. Bot. Sada
Heterotypic synonyms: Thlaspi suffruticosum Asso ex 12-13: 114. 1931. ARMENIA. Two syntypes were
Loscos & J.Pardo, Ser. Inconf. Pl. Aragon. ed. 2, cited by Meyer (2006e), none of which was examined
38. 1867; Noccaea suffruticosa (Asso ex Loscos & by him or the present author. Lectotypification of the
J.Pardo) Holub, Preslia 70: 108. 1998. TYPE: Not species is needed.
located. Distribution: Armenia, Azerbaijan, Iran, Turkey.
Distribution: Spain.
Noccaea tenuis (Boiss. & Buhse) F.K.Mey., Feddes Repert.
Noccaea stilosa (Ten.) Rchb., Fl. Germ. Excurs. 663. 1832. 84: 459. 1973.
Basionym: Iberis stilosa Ten., Fl. Napol. 1: 37. 1812. Basionym: Iberidella tenuis Boiss. & Buhse, Nouv. Mém.
TYPE: ITALY. La Majella, Mt. Amaro, Tenore s.n. So. Imp. Natualistes Moscou 12: 25. 1860. TYPE:
(Holotype: NAP). IRAN. Kuh-e Sahand, Damirdagh, 10,000 ft [ca.
Homotypic synonym: Thlaspi stilosum (Ten.) Mutel, Fl. 3050 m], F. A. Bushse 589/2 (lectotype designated by
Franç. 1: 99. 1834 (as stylosum). Hedge 1968: 117 as type: G-BOIS; Isolectotype: LE).
Distribution: Italy. Homotypic synonyms: Aethionema tenue (Boiss. & Buhse)
Boiss., Fl. Orient. 1: 343. 1867; Thlaspi tenue (Boiss.
Noccaea swatensis F.K.Mey., Haussknechtia 12: 178. 2006. & Buhse) Hedge, Fl. Iranica 57: 117. 1968.
TYPE: PAKISTAN. Swat, E Kalam, 3,000 m, 6 Jun 1965, J. Distribution: Iran.
Lamond & K. H. Rechinger 30855 (Holotype: W).
Distribution: Pakistan. Noccaea thlaspidioides (Pall.) F.K.Mey., Haussknechtia
Meyer (2006e) described Noccaea swatensis solely on 12: 5. 2010.
flowering material. More complete material of the species Basionym: Lepidium thlaspidioides Pall., Reise Russ.
was erroneously interpreted by Jafri (1955, 1973) as Thlaspi Reich. 3: 161. 1776. LECTOTYPE designated by
septigerum (Bunge) Jafri. The latter name is based on Meyer (2010: 5): Plate 56, Fig. 1 in Gmelin, Fl. Sibir.
disjunct Altai plants known as Eutrema septigera Bunge 3. 1768.
which was reduced by Busch (1939) to synonymy of E. Homotypic synonym: Thlaspi thlaspidioides (Pall.) Kitag.,
edwardsii R.Br. Rep. Inst. Sci. Res. Manchoukuo 2: 293. 1938.
Heterotypic synonyms: Thlaspi cochleariforme DC., Syst.
Noccaea sylvia (Gaudin) F.K.Mey., Feddes Repert. 84: 461. Nat. 2: 381. 1821; Noccaea cochleariformis (DC.)
1973. Á. & D. Löve, Bot. Not. 128(4): 513. 1976. TYPE:
Basionym: Thlaspi sylvium Gaudin, Fl. Helv. 4: 221. 1829. RUSSIA. Sibiria et Dahuria ad jugum Yabloni-
TYPE: SWITZERLAND. Zermatt, Murith s.n. Chrebet, Patrin s.n. (Holotype: G-DC).
(Lectotype designated by Meyer (2006e: 91): LAU). Thlaspi exauriculatum Kom., Repert. Spec. Nov.
Homotypic synonyms: Thlaspi minimum Ard. subsp. Regni Veg. 9: 392. 1911; Noccaea exauriculata
sylvium (Gaudin) H.P.Fuchs, Feddes Repert. 90: (Kom.) Czerep., Sosud. Rast. SSSR 140. 1981.
TYPE: RUSSIA. Insula Sachalin, 18 Jun 1909, M. E. Noccaea umbellata (Steven ex DC.) Al-Shehbaz, comb. nov.
Semjagin s.n. (Holotype: LE). Basionym: Thlaspi umbellatum Steven ex DC., Syst. Nat.
Distribution: N China, Kazakhstan, Mongolia, Russia. 2: 377. 1821. TYPE: IRAN. Gilan, S. G. Gmelin s.n.?
Holmgren (1971) listed Thlaspi cochleariforme as a (Holotype: H; isotypes, LE, MW).
synonym of T. montanum, but as shown by Meyer (2010) Homotypic synonym: Microthlaspi umbellatum (Steven ex
the former is a synonym of Noccaea thlaspidioides, a DC.) F.K.Mey., Feddes Repert. 84: 453. 1973.
species clearly unrelated to N. montana. Records of N. Distribution: Armenia, Azerbaijan, Iran.
thlaspidioides (as T. cochleariforme) from Tibet (Xizang) The present author has not seen the type collection of the
in Zhou et al. (2001) are likely based on plants of N. species to justify if the lectotypification of the species, as
yunnanensis. done by Dorofeyev (2012: 457), is needed or not.
Noccaea triangularis (F.K.Mey.) Al-Shehbaz, comb. nov. Noccaea valerianoides (Rech.f.) F.K.Mey., Feddes Repert.
Basionym: Thlaspiceras triangulare F.K.Mey., Feddes 84: 460. 1973.
Repert. 84: 455. 1973. TYPE: TURKEY. Prov. Basionym: Thlaspi valerianoides Rech.f., Phyton (Horn) 3:
Seyhan, Osmaniye, near Zortum, Amanus, 1,500 m, 50. 1951. TYPE: TURKEY. Kurdistan, Passet Aghzi
17 Jun 1967, Y. Akman 220 (Holotype: G). Gedik, 13 km N of Mukus, 2,900 m, 24 Jun 1939, J.
Homotypic synonym: Thlaspi triangulare (F.K.Mey.) Frödin (Holotype: W; Isotypes: JE, UPS).
Greuter & Burdet, Willdenowia 13: 96. 1983. Homotypic synonym: Apterigia valerianoides (Rech.f.)
Distribution: Turkey. V.I.Dorof., Novosti Sist. Vyssh. Rast. 44: 94. 2013.
Heterotypic synonym: Noccaea valerianoides subsp.
Noccaea trinervia (DC.) Steud., Nomencl. Bot., ed. 2. 2: rechingeri F.K.Mey., Haussknechtia 12: 69. 2006.
TYPE: IRAN, Prov. Mazanderan, Distr. Kudjur,
197. 1841.
Mt. Ulodj, 3,200–3,400 m, 9 Aug 1948, K. H. & F.
Basionym: Hutchinsia trinervia DC., Syst. Nat. 2: 387.
Rechinger 6522 (Holotype: W).
1821. TYPE: IRAN. Mt. Elwind, A. Michaux s.n.
Distribution: Iran, Turkey.
(Lectotype here designated: G-DC-00131230;
The species is poorly collected in both Iran and Turkey,
Isolectotype: P). and the disjunct population in Iran is hardly distinct
Homotypic synonyms: Aethionema trinervium (DC.) Boiss., morphologically to merit its recognition as an independent
Fl. Orient. 1: 342. 1867; Iberidella trinervia (DC.) subspecies.
Boiss., Ann. Sci. Nat., Bot. Sér. 2, 17: 188. 1842;
Vania trinervia (DC.) Khosravi et al., Ann. Missouri Noccaea versicolor (Stoj. & Kitan.) F.K.Mey.,
Bot. Gard. 96: 570. 2009; Thlaspi trinervium (DC.) Haussknechtia 12: 116. 2006.
Mozaff., Iranian J. Bot. 7: 139. 1996. Basionym: Thlaspi lutescens Velen. f. versicolor Stoj. &
Distribution: Afghanistan, Armenia, Azerbaijan, Iran, Kitan., Izv. Bulg. Bot. Druzh. 9: 97. 1943. TYPE:
Iraq, Kazakhstan, Turkey, Turkmenistan, Uzbekistan. GREECE. Tasos, Ilija, 1,100 m, 8 Apr 1942, N.
Khosravi et al. (2009b) placed the species in genus Vania, Stojanov & B. Kitanov s.n. (Holotype: SOM).
which is reduced herein to synonymy of Noccaea. They Distribution: Greece, Turkey.
designated Oliver’s collection at G-DC as the lectotype, but
that collection was made in 1822, a year after the original Noccaea vesicaria (L.) Al-Shehbaz, comb. nov.
publication of the species. Therefore, a new lectotypification Basionym: Alyssum vesicaria L., Sp. Pl. 2: 910. 1753.
is proposed above. TYPE: “Vesicaria Orientalis foliis dentatis Coroll.”
In Tournefort, Rel. Voy. Levant 2:[“t. 14”]. 1717
Noccaea tymphaea (Hausskn.) F.K.Mey., Feddes Repert. (Lectotype designated by Al-Shehbaz & Turland in
84: 462. 1973. Cafferty & Jarvis (2002: 531)).
Basionym: Thlaspi tymphaeum Hausskn., Mitth. Thüring. Homotypic synonyms: Coluteocarpus vesicaria (L.)
Bot. Vereins N. Folge 3-4: 115. 1893. TYPE: Holmboe, Bergens Mus. Ǻrbok 13: 6. 1907.
GREECE. Pindus Tymphaeus, Mt Zygos, above Heterotypic synonym: Coluteocarpus reticulatus (Lam.)
Metzovo, 4,500–5,000 ft [1,371–1,524 m], Jul Boiss., Ann. Sci. Nat., Bot. sér. 2, 17: 162. 1842;
1885, C. Haussknecht s.n. (Holotype: JE-00004213; Vesicaria reticulata Lam. ex Poir., Encycl. (Lamarck)
Isotypes: B, JE, W). 8: 572. 1808. TYPE: TURKEY. J. P. de Tournefort
Heterotypic synonyms: Thlaspi pindicum Hausskn., Mitth. s.n. (Lectotype here designated: MPU).
Thur. Bot. Ver. N. Folge 3-4: 115. 1893; Noccaea Distribution: Armenia, Azerbaijan, Georgia, Turkey.
pindica (Hausskn.) Holub, Preslia 70: 108. 1998.
TYPE: GREECE. Pindus Tympaeus, Zygos, 4,500– Noccaea vesicaria subsp. boissieri (Bornm.) Al-Shehbaz,
5,000 ft [1,371–1,524 m], July 1885, C. Haussknecht comb. nov.
s.n. (Holotype: JE-00004235; Isotypes: JE, K). Basionym: Coluteocarpus reticulatus var. boissieri Bornm.,
Distribution: Albania, Bosnia and Herzegovina, Greece, Beih. Bot. Centralbl. 19(2): 207. 1906. TYPE:
Macedonia. IRAN. Mt. Schuturun Kuh, May 1897, T. Strauss s.n.
(Lectotype here designated: JE-00006910).
Homotypic synonyms: Coluteocarpus vesicaria subsp. Noccaea wendelboi (Rech.f.) F.K.Mey., Haussknechtia 12:
boissieri (Bornm.) Hedge, Notes Roy. Bot. Gard. 179. 2006
Edinburgh 26: 181. 1965; C. boissieri (Bornm.) Basionym: Thlaspi wendelboi Rech.f., Biol. Skr. 10(3): 26.
O.E.Schulz in Engler & Prantl, Nat. Pflanzenfam., ed. 1959. TYPE: PAKISTAN. Chitral, Barum Gol, Soth
2, 17B: 428. 1936. Barum Glacier, ca. 4,500 m, 27 Jul 1950, P. Wendelbo
Distribution: Iran, Iraq, Lebanon, Turkey. s.n. (Holotype: W).
Subspecies boissieri differs from subsp. vesicaria Distribution: Afghanistan, Pakistan.
by having (1 or)2-seeded fruits with a complete septum Both Hedge (1968) and Jafri (1973) reduced this species
and ovate to spatulate or rhomboid rosette leaves (vs. to synonymy of Noccaea griffithiana (as Thlaspi). However,
4–10(–12)-seeded fruits with the septum reduced to a narrow N. wendelboi is a diminutive plant that grows at much
rim and linear to narrowly oblanceolate rosette leaves). higher altitudes, and it has smaller flowers and smaller and
narrower fruits than those of N. griffithiana. Furthermore,
Noccaea violascens (Schott & Kotschy) F.K.Mey., Feddes the style in N. wendelboi subequals (vs. much shorter)
Repert. 84: 465. 1973. than the apical notch of the very narrowly (vs. rather
Basionym: Thlaspi violascens Schott & Kotschy in Unger broadly) winged fruits. Without thorough studies on these
& Kotschy, Ins. Cyp. 328. 1862. TYPE: TURKEY. narrow endemics, I prefer to maintain both of them as
Taurus, Bulgar Dagh, 7,000 ft [2,134 m], 8 Jul distinct species.
1853, C. G. T. Kotschy 70a (Holotype: W; Isotypes: B,
G, WAG). Noccaea yunnanensis (Franch.) Al-Shehbaz, Adansonia
Distribution: Turkey. sér. 3, 24(1): 91. 2002.
Basionym: Thlaspi yunnanense Franch., Bull. Soc. Bot.
Noccaea virens (Jord.) F.K.Mey., Feddes Repert. 84: 463. France 33: 407. 1886. TYPE: China. Yunnan. Yen-
1973. tze-hay, Lankong, 3,200 m, 1 June 1886, J. M. Delavay
Basionym: Thlaspi virens Jord., Observ. Pl. Nouv. 3: 17. 2077 [as 4077 in the original descritption] (Holotype:
1846. TYPE: FRANCE. A loctotype should be P-02141454; Isotypes: MO, NY, P, US, W).
selected from the four syntypes collected by A. Jordan Distribution: China.
and presumably housed at LY: Mont Lozère près
Villefort, Mont Mèzin, Mont Pilat près Lyon, and Noccaea zaffranii F.K.Mey., Willdenowia 15: 389. 1986.
Pierre-sur Haute. None of these were examined by the TYPE: GREECE. Crete, Lefa Ori, Volkias, 1,700 m, 3 Aug.
present author. 1966, J. Zaffran 1684 (Holotype: B).
Homotypic synonyms: Pterotropis virens (Jord.) Four., Homotypic synonym: Thlaspi zaffranii (F.K.Mey.) Greuter
Ann. Soc. Linn. Lyon, sér. 2, 16: 337. 1868; Noccaea & Burdet, Willdenowia 15: 420. 1986.
caerulescens (J.Presl & C.Presl) F.K.Mey. subsp. Distribution: Greece.
virens (Jord.) Kerguélen, Collect. Patrim. Nat. 8: 15.
1993. Noccaea zangezurica (Tzvelev) Al-Shehbaz, comb. nov.
Distribution: France, Italy, Switzerland. Basionym: Thlaspi zangezuricum Tzvelev, Bot. Mater.
Gerb. Bot. Inst. Komarova Akad. Nauk S.S.S.R. 19:
Noccaea viridisepala (Podp.) F.K.Mey., Feddes Repert. 84: 9. 1959. TYPE: ARMENIA. Dist. Megri, “in rupibus
461. 1973. regionis montanae superioris prope trajectum trans
Basionym: Thlaspi praecox Wulfen var. viridisepalum jugum Zangezuricum supra pag. Vardanadzor,” 5
Podp., Verh. K. K. Zool.-Bot. Ges. Wien 52: 632. July 1956, T. V. Egorova, N. N. Tzvelev, and S. K.
1902. TYPE: BULGARIA. Vitosha, ca. 1500 m, 7 Czerepanov 1941 (Holotype: LE).
May 1967, F. K. & J. Meyer 8863 (Neotype designated Homotypic synonym: Atropatenia zangezurica (Tzvelev)
by Meyer (2006e: 88): JE). F.K.Mey., Feddes Repert. 84: 467. 1973; Apterigia
Homotypic synonyms: Thlaspi viridisepalum (Podp.) zangezurica (Tzvelev) V.I.Dorof., Konspekt. Fl.
Greuter & Burdet, Willdenowia 13: 96. 1983. Kavkaza 3(2): 459. 2012.
Distribution: Albania, Bosnia and Herzegovina, Distribution: Armenia.
Bulgaria, Greece, Macedonia, Montenegro.
Excluded Names
Names in boldface are the currently accepted placements of excluded Noccaea taxa, and all are currently assigned to tribe
Descurainieae.
Noccaea affinis (F.W.Schultz) Rouy & Foucaud, Fl. France Noccae maritima (Jord.) Rouy & Foucaud, Fl. France 2: 91,
2: 90. 1895. = Hornungia alpina (L.) O.Appel. 92. 1895.. = Hornungia procumbens (L.) Hayek.
Noccaea alpina (L.) Rchb., Fl. Germ. Excurs. 663. 1832. = Noccaea pauciflora (Koch) Rouy & Foucaud, Fl. France 2:
Hornungia alpina (L.) O.Appel. 92. 1895. = Hornungia procumbens (L.) Hayek.
Noccaea brevicaulis (Spreng.) Rchb., Fl. Germ. Excurs. Noccaea procumbens (L.) Rchb., Fl. Germ. Excurs. 663. =
663. 1832. = Hornungia alpina (L.) O.Appel. Hornungia procumbens (L.) Hayek.
Noccaea auerswaldii Willk., Willk. & Lange, Prod. Fl. Noccaea prostii (Jord.) Rouy & Foucaud, Fl. France 2: 92,
Hisp. 3: 780. 1880. = Hornungia alpina (L.) O.Appel. 93. 1895. = Hornungia procumbens (L.) Hayek.
Noccaea crassifolia (Corbiière) Rouy & Foucaud, Fl. Noccaea revelieri (Jord.) Rouy & Foucaud, Fl. France 2: 91,
France 2: 91, 92. 1895. = Hornungia procumbens (L.) 93. 1895. = Hornungia procumbens (L.) Hayek.
Hayek. Noccaea speluncarum (Jord.) Rouy & Foucaud, Fl. France
Noccaea diffusa (Jord.) Rouy & Foucaud, Fl. France 2: 91, 2: 91, 93. 1895. = Hornungia procumbens (L.) Hayek.
92. 1895. = Hornungia procumbens (L.) Hayek. Noccaea tournefortii (DC.) Rouy & Foucaud, Fl. France 2:
91, 92. 1895. = Hornungia procumbens (L.) Hayek.
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A SYNOPSIS OF THE GENUS NOCCAEA (COLUTEOCARPEAE, BRASSICACEAE)

Harvard Papers in Botany, Vol. 19, No. 1, 2014, pp. 25–51.

Key to the species of Noccaea cont.

Key to the species of Noccaea cont.

Key to the species of Noccaea cont.

Key to the species of Noccaea cont.

Key to the species of Noccaea cont.

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