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23Новости систематики высших растений 2017
Novitates Systematicae Píantarum Vascuíarium 48: 78-83 Î I ISSN 0568-5443
A taxonomic note on Sterigmostemum and related genera (Anchonieae, Cruciferae)
Заметка о Sterigmostemum и близких родах (Anchonieae, Cruciferae)
D. A. German1, I. A. Al-Shehbaz2
1 Heidelberg University, Centre for Organismal Studies, Department of Biodiversity and Plant Systematics
Im Neuenheimer Feld, 345, D-69120, Heidelberg, Germany; South-Siberian Botanical Garden, Altai State University Lenina Ave., 61, Barnaul, 656049, Russia [email protected]
2 Missouri Botanical Garden
P. O. Box 299, St. Louis, Missouri, 63166-0299, USA [email protected]
Д. А. Герман1, И. А. Аль-Шебаз2
1 Гейдельбергский университет, Центр исследований организмов, кафедра биоразнообразия и систематики растений
Им Нойенхаймер Фельд, 345, D-69120, Гейдельберг, Германия; Южно-Сибирский ботанический сад, Алтайский государственный университет пр. Ленина, 61, Барнаул, 656049, Россия [email protected]
2 Миссурийский ботанический сад
а/я 299, Сент-Луис, Миссури, 63166-0299, США [email protected]
Abstract. Our critical re-evaluation of diagnostic characters of Anchonium DC., Oreoloma Botsch., Sterigmostemum M. Bieb., and Zerdana Boiss. on the basis of phylogenetic evidence resulted in merging these genera. Anchonium and Zerdana are synony-mized with Sterigmostemum, and relevant combinations are validated. A checklist and a key for the expanded Sterigmostemum are presented. Synstemon Botsch. is found unrelated to this genus and should be transferred from the tribe Anchonieae DC. to Euclidieae DC.
Keywords: Anchonium, Brassicaceae, Middle East, Oreoloma, SW and Central Asia, taxonomy, Zerdana.
Аннотация. На основании критической ревизии диагностических признаков родов Anchonium DC., Oreoloma Botsch., Sterigmostemum M. Bieb. и Zerdana Boiss. в свете имеющихся молекулярно-филогенетических данных сделан вывод о целесообразности признания единственного таксона родового ранга вместо четырех. Anchonium и Zerdana впервые отнесены к синонимам Sterigmostemum, обнародованы соответствующие комбинации. Приведен конспект и ключ для определения видов Sterigmostemum s. l. Род Synstemon Botsch. предложено исключить из трибы Anchonieae DC. и перевести в Euclidieae DC.
Ключевые слова: Ближний Восток, систематика, Юго-Западная и Центральная Азия, Anchonium, Brassicaceae, Oreoloma, Zerdana.
Sterigmostemum M. Bieb. was established to accommodate two species originally described under Chei-ranthus L. (= Erysimum L.) that have connate pairs of median (long) stamens and presumably indehiscent (in fact partly dehiscent in S. caspicum (Lam.) Rupr.) fruits (Marschall von Bieberstein, 1819). Unlike the majority of genera of the mustard family (Brassicaceae Burnett or Cruciferae Juss.), both morphological limits and affinities of Sterigmostemum did not undergone drastic changes during the history of its systematics. The main update of the generic characteristics was adding the species with completely dehiscent fruit (Jacquemoud, 1988; Appel, Al-Shehbaz, 2003). It is also a relatively rare case within Cruciferae that the very first tribal affiliation proposed for the genus (Anchonieae DC.; Candolle, 1821a, b) is being accepted nowadays (Al-Shehbaz, 2012). Three genera, Anchonium DC. (Candolle, 1821a, b),
Поступила в редакцию | Submitted: 11.12.2017
Zerdana Boiss. (Boissier, 1842), and Oreoloma Botsch. (Botschantzev, 1980) were described and traditionally considered as closely related to Sterigmostemum, and this morphology-based affinity was subsequently confirmed by molecular phylogenetic studies (Warwick et al., 2007; German et al., 2009; Couvreur et al., 2010) proving the monophyly of the group in question.
Morphology and taxonomy of Sterigmostemum and related genera were the focus of a number of studies during past four decades (Dvorak, 1972; Botschantzev, 1980; Léonard, 1980, 1988; Kamelin, German, 2001; Sonboli et al., 2001, 2006; Appel, Al-Shehbaz, 2003) among which the most extensive and detailed are the monographic treatments of Jacquemoud (1984b, 1985, 1988). As evidenced by thorough analyses of the last author, accompanied by the data of Dvorak (1972), Zhou et al. (2001), Appel, Al-Shehbaz (2003) and own obser-
Принята к публикации | Accepted: 18.12.2017
vations, Sterigmostemum (with exclusion of more distant Petiniotia J. Leonard) shares with Anchonium, Oreoloma, and Zerdana nearly all features, both common to the tribe (uniform dense indumentum of many-branched dendritic trichomes usually throughout the plant, often presence of multicellular glands, non-auriculate cau-line leaves poorly differentiated into petiole and blade, distinctly lobed stigmas, base chromosome number of n = 7, etc.) and specific of all the four genera (filaments of median stamens united into pairs, terete to rarely sub-quadrangular, somewhat torulose fruits often gradually tapering towards apex, diverged stigma lobes, oblong and wingless seeds, and incumbent cotyledons). The latter character combination is not found elsewhere in the tribe Anchonieae, and the connation of median sta-minal filaments does not occur in any other genus of the tribe (except for problematic Synstemon Botsch.; see below). Most of the characters used in differentiating these genera are mosaically distributed among them, and very few unique characters are minor and lacking a strong taxonomic value. In particular, slightly apiculate anthers, leafless stems, zygomorphic flowers, and (sub) scapose habit are diagnostic of Zerdana, though specimens of the highly polymorphic Anchonium elichrysifo-lium (DC.) Boiss. may share such a habit as well. Furthermore, the reduction of the ovule number to 4-6 per ovary in Anchonium is found in Sterigmostemum ra-mosissimum (O. E. Schulz) Rech. f., a species typically with 4-8 ovules per ovary. Finally, the sepals are erect and saccate in Oreoloma, erect and subsaccate in Ancho-nium, erect and non-saccate in Zerdana, and ascending to erect-ascending and non-saccate (but rarely erect and subsaccate) in Sterigmostemum. The differentiation of petals into a claw and a limb is strong in Oreoloma, weak in Zerdana, weak to strong in Anchonium, and weak (but rarely strong) in Sterigmostemum. In a similar way, fruits are completely dehiscent in monospecific Zerdana, tardily dehiscent distally in three species of Oreoloma, and completely indehiscent in both members of Anchonium, whereas all these fruit types can be found in Sterigmoste-mum s. str. (seven species). Many other characters, e. g., leaf dissection (entire to pinnatisect), style form (cylin-dric or conical) and length (obsolete or to 5 mm long), stigma lobes (elongate up to linear to shortened), etc., reveal the same picture.
In the absence of strong morphological differences among these closely related genera, it would be far more practical and scientifically sound that species of Sterigmostemum, Anchonium, Oreoloma, and Zerdana are treated as members of one instead of four genera. However, by now a merge with Sterigmostemum has only been proposed for Oreoloma (Yang, 1987; Kamelin, German, 2001), a viewpoint followed by some (Al-
Shehbaz, 2012; present study) but not others (Zhou et al., 2001; Appel, Al-Shehbaz, 2003; Warwick et al., 2006, 2007; Yang, 2009). Surprisingly, the status of Anchonium and Zerdana has never been questioned.
The above-mentioned molecular phylogenetic studies, which included collectively 11 out of the 13 species currently accepted in Sterigmostemum and its relatives, clearly highlight the concerns of recognizing four, poorly defined genera. The overall similarities among these genera in intermingling habit, floral and fruit morphology are perfectly matched by the evident paraphyly of Sterigmostemum in which Anchonium, Oreoloma, and Zerdana are partly or completely embedded (Warwick et al., 2007; German et al., 2009). Possible taxonomic solutions were briefly mentioned in these works and are further discussed and summarized here. One option could be the further splitting and recognition of all terminal monophyletic clades as separate genera. This would, in particular, mean the transfer of S. longisty-lum (Boiss.) Kuntze to Zerdana and establishing a new genus for S. ramosissimum, a species morphologically intermediate between Sterigmostemum and Anchonium (Jacquemoud, 1988). On the other hand, A. elichry-sifolium would be either placed back in Sterigmostemum (German et al., 2009) in agreement with initial concept of Candolle (1821b, as Sterigma DC., nom. illeg.), or also recognized as a genus by its own. Such a reshuffling would result in some morphological grouping (e. g., assembling all species combining completely dehiscent fruits, erect sepals, narrow and elongated stigma lobes, and long, narrowly cylindrical styles under the generic name Zerdana). However, such changes would barely outweigh the shortcomings of this decision because in general delimitation between the newly defined genera will be even more blurred than before. For instance, among the above features characterizing the potentially dispecific Zerdana, only relatively readily dehiscent fruit is unique for both species while other characters occur, individually or in various combinations, in some representatives of Sterigmostemum s. l. However, even fruit dehiscence is not fully diagnostic in this case because very tardily dehiscent fruits may be observed in S. acanthocarpum (Fisch. et C. A. Mey.) Kuntze (Jacquemoud, 1988; Sonboli et al., 2006), another species with long cylindrical styles and elongated stigma lobes which revealed closest phylogenetic affinity with S. sul-phureum (Banks et Soland.) Bornm., a species with indehiscent fruit and short style. Meanwhile, states of a number of characters (such as base of sepals, petal differentiation into a claw and a blade, flower symmetry, etc.), separate Z. anchonioides Boiss. from S. longistylum and either make each of them morphologically closer to other species of Sterigmostemum s. l. or (e. g., flower
zygomorphy) emphasize exclusive nature of Z. anchoni-oides. As a result, expanded Zerdana will be more hete-rogenic rather than separated from the related genera. With the recognition of S. ramosissimum and A. elichry-sifolium as two new monospecific genera to maintain Sterigmostemum s. str. (including Oreoloma) as mono-phyletic, the situation will get even worse. In particular, completely indehiscent fruits will be found in Anchoni-um, Sterigmostemum, and in both newly established genera, and the rest of characters will be distributed similarly. In our opinion, splitting is not a good taxonomic decision in Sterigmostemum s. l.
Hence, the alternative approach of lumping the four genera under the earliest-published Sterigmostemum seems far more justified because it requires only minor changes in the morphological characteristics of the expanded genus, and the support of the common clade is rather high (Warwick et al., 2007; German et al., 2009). The only concern regarding this taxonomic decision is the phylogenetic position of Synstemon as a sister taxon to the type species of Anchonium, A. billardierei, which was found by Warwick et al. (2007). This finding was extremely unexpected because Synstemon differs from the Anchonieae by the lack (vs. usually presence) of multicellular glands and by having strongly crisped simple and forked trichomes (vs. exclusively or predominantly dendritic with usually straight branches) and pilose (vs. glabrous) petal claws and sometimes filaments (Al-Shehbaz et al., 2000). The only feature which could be considered as a hint of the possible relationship of Synstemon with the rest of the respective clade is its basally united median filaments pairs (vs. connate above the middle in Anchonium, Oreoloma, Sterigmostemum, and Zerdana). Possible explanations of the strange position of Synstemon are either a laboratory mistake or, in view of several cases of successful ancient inter-tribal hybridization within the family (e. g., Joly et al., 2009; Mandakova et al., 2010, 2017; German, Friesen, 2014), a hybrid origin of Synstemon. Additional sequencing of nuclear (ITS) and chloroplast (trnL-F) fragments of both species of Synstemon (German & Koch, unpubl. data) revealed its unambiguous relationship with Euclidieae DC. and no sign of inter-tribal hybridization. This agrees well with the data of Chen et al. (2016) who, based on the analysis of several chloroplast markers, found Synstemon petrovii Botsch. within the clade formed also by Pycnoplinthus uniflora (Hook. f. et Thoms.) O. E. Schulz and Cryptospora falcata Kar. et Kir., both of which are members of the Euclidieae. Apparently, Synstemon is much at home in Euclidieae rather than in Anchonieae, and the sequence DQ357599 of S. petrovii, the generic type, in Warwick et al. (2007), which was the basis for assigning the genus to the An-
chonieae, is nearly identical to DQ357512 of A. billar-dierei and to which it should be attributed.
The removal of this Synstemon obstacle proves the congruence of morphological and phylogenetic data and supports a broader concept of Sterigmostemum. The last item to be mentioned is the sister position of A. bil-lardierei to the rest of the discussed group (Warwick et al., 2007). One might argue for keeping Anchonium as a monospecific genus separate from Sterigmostemum because it has bracteate (vs. ebracteate) racemes, but this character is usually trivial taxonomically at the generic level because both raceme types are found within numerous genera of the family. Therefore, an expanded concept of Sterigmostemum is proposed here with a formal synonymization of Anchonium and Zerdana and necessary combinations for their three species and three subspecies when transferred to Sterigmostemum. The united genus encompasses 13 species of predominantly SW and Central Asian distributions, of which a complete checklist is provided below.
Sterigmostemum M. Bieb. 1819, Fl. Taur.-Cauc. 3:
444.
= Sterigma DC. IV 1821, Mem. Mus. Hist. Nat. 7: 242; id. V 1821, Reg. Veg. Syst. Nat. 2: 579, nom. illeg. superfl.
Lectotypus (Jacquemoud, 1984a: 303): S. incanum M. Bieb.
= Anchonium DC. IV 1821, Mem. Mus. Hist. Nat. 7: 242; id. V 1821, Reg. Veg. Syst. Nat. 2: 578, syn. nov. — Ty p u s: A. billardierei DC.
= Zerdana Boiss. 1842, Ann. Sci. Nat. Bot., ser. 2, 17: 84, syn. nov. — Ty p u s: Z. anchonioides Boiss.
= Oreoloma Botsch. 1980, Bot. Zhurn. 65, 3: 425. — Ty p u s: O. matthioloides (Franch.) Botsch.
1. Sterigmostemum acanthocarpum (Fisch. et
C. A. Mey.) Kuntze, 1891, Revis. Gen. Pl. 1: 36. = Sterigma acanthocarpum Fisch. et C. A. Mey. 1835, Index Sem. Hort. Petropol. 1: 38.
2. Sterigmostemum anchonioides (Boiss.)
D. A. German et Al-Shehbaz, comb. nova. = Zerdana anchonioides Boiss. 1842, Ann. Sci. Nat. Bot., ser. 2, 17: 84.
2a. Sterigmostemum anchonioides subsp. steno-carpum (Jacquemoud) D. A. German et Al-Shehbaz, comb. nova. = Zerdana anchonioides subsp. stenocarpa Jacquemoud, 1985, Candollea, 40, 2: 369.
2b. Sterigmostemum anchonioides subsp. ste-nophyllum (Boiss. et Hausskn.) D. A. German et Al-Shehbaz, comb. nova. = Zerdana anchonioides [var.] p. stenophylla Boiss. et Hausskn. 1888, Fl. Orient. Suppl.: 46. = Z. anchonioides subsp. stenophylla
(Boiss. et Hausskn.) Jacquemoud, 1985, Candollea, 40, 2: 368.
3. Sterigmostemum billardierei (DC.) D. A. German, comb. nova. = Anchonium billardierei DC. IV 1821, Mém. Mus. Hist. Nat. 7: 242; id. V 1821, Reg. Veg. Syst. Nat. 2: 578.
4. Sterigmostemum caspicum (Lam.) Rupr. 1869, Mém. Acad. Sci. Pétersb., sér. 7, 15: 95. = Cheiranthus caspicus Lam. 1794, in Pall., Voy. Plus. Prov. Russ., App. 8: 348.
= Cheiranthus nitrarius Pall. 1799, Reise Südl. Russ. Reichs, 1: 105, 124, 141.
= Cheiranthus tomentosus Willd. 1800, Sp. Pl. 3: 523. = Sterigmostemum tomentosum (Willd.) M. Bieb. 1819, Fl. Taur.-Cauc. 3: 444. = Sterigma tomentosum (Willd.) DC. 1821, Reg. Veg. Syst. Nat. 2: 579.
5. Sterigmostemum eglandulosum (Botsch.) H. L. Yang, 1987, Fl. Desert. Reipubl. Popul. Sin. 2: 66. = Oreoloma eglandulosum Botsch. 1980, Bot. Zhurn. 65, 3: 427.
= Sterigmostemum grandiflorum K. C. Kuan, 1980, Bull. Bot. Lab. N. E. Forest. Inst., Harbin, 8: 43. = Oreoloma grandiflorum (K. C. Kuan) H. L. Yang, 2009, J. Desert Res. 29, 3: 433.
6. Sterigmostemum elichrysifolium (DC.)
D. A. German et Al-Shehbaz, comb. nova. = Sterigma elichrysifolium DC. 1821, Reg. Veg. Syst. Nat. 2: 581. = Anchonium elichrysifolium (DC.) Boiss. 1867, Fl. Orient. 1: 240. = A. tournefortii Boiss. 1842, Ann. Sci. Nat. Bot., sér. 2, 17: 386, nom. illeg. superfl.
= Matthiola persica DC. 1821, Reg. Veg. Syst. Nat. 2: 168. = Anchonium persicum (DC.) Bornm. 1904, Bull. Herb. Boissier, sér. 2, 4: 1265. = A. elichrysifolium subsp. persicum (DC.) Coode et Cullen, 1965, Notes Roy. Bot. Gard. Edinburgh, 26, 2: 193.
= Anchonium elichrysifolium var. brachycarpum Trautv. 1873, Acta Horti Petrop. 2: 503. = A. brachycar-pum (Trautv.) Vass. 1939, Fl. URSS, 8: 321.
= Anchonium tournefortii var. canescens Hausskn. ex Bornm. 1904, Bull. Herb. Boissier, sér. 2, 4: 1265. = A. elichrysifolium subsp. canescens (Hausskn. ex Bornm.) Coode et Cullen, 1965, Notes Roy. Bot. Gard. Edinburgh, 26, 2: 194.
= Anchonium elichrysifolium [var.] ß. tournefortii (Boiss.) Bornm. subvar. cilicicum Hausskn. ex Bornm. 1936, Feddes Repert., Beih. 89, 1: 39. = A. elichrysifo-lium subsp. cilicicum (Hausskn. ex Bornm.) Coode et Cullen, 1965, Notes Roy. Bot. Gard. Edinburgh, 26, 2: 193.
= Anchonium elichrysifolium subsp. glandulosum Coode et Cullen, 1965, Notes Roy. Bot. Gard. Edinburgh, 26, 2: 193.
6a. Sterigmostemum elichrysifolium subsp. villo-sum (Coode et Cullen) D. A. German et Al-Shehbaz, comb. nova. = Anchonium elichrysifolium subsp. villosum Coode et Cullen, 1965, Notes Roy. Bot. Gard. Edinburgh, 26, 2: 194.
7. Sterigmostemum fuhaiense H. L. Yang, 1987, Fl. Desert. Reipubl. Popul. Sin. 2: 445. = Oreoloma fuhai-ense (H. L. Yang) H. L. Yang, 2009, J. Desert Res. 29, 3: 433.
= Sterigmostemum schmakovii Kamelin et D. A. German, 2001, Turczaninowia, 4, 3: 5.
8. Sterigmostemum incanum M. Bieb. 1819, Fl. Taur.-Cauc. 3: 444. = Cheiranthus torulosus M. Bieb. 1808, Fl. Taur.-Cauc. 2: 121, non Thunb. 1800. = Sterigma torulosum (M. Bieb.) DC. 1821, Reg. Veg. Syst. Nat. 2: 580, comb. illeg. = Sterigmostemum torulosum (M. Bieb.) Stapf, 1886, Denkschr. Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl. 51: 301, comb. illeg.
= Sterigma contortuplicatum Boiss., 1842, Ann. Sci. Nat. Bot., sér. 2, 17: 387. = Sterigmostemum contortuplicatum (Boiss.) Kuntze, 1891, Revis. Gen. Pl. 1: 36.
9. Sterigmostemum longistylum (Boiss.) Kuntze, 1891, Revis. Gen. Pl. 1: 36. = Sterigma longistylum Boiss. 1842, Ann. Sci. Nat. Bot., sér. 2, 17: 387.
10. Sterigmostemum matthioloides (Franch.) Botsch. 1959, Bot. Zhurn. 44, 10: 1487. = Dontostemon matthioloides Franch, 1883, Pl. David. 1: 35. = Oreoloma matthioloides (Franch.) Botsch. 1980, Bot. Zhurn. 65, 3: 426.
11. Sterigmostemum ramosissimum (O. E. Schulz) Rech. f. 1968, Fl. Iranica, 57: 280. = Anchonium ramosis-simum O. E. Schulz, 1933, Bot. Jahrb. Syst. 66, 1: 97.
- Anchonium sterigmoides Lipsky ex Vass. 1939, Fl. URSS, 8: 322, nom. inval.
12. Sterigmostemum sulphureum (Banks et Soland.) Bornm. 1911, Beih. Bot. Centralbl. 28, 2: 110. = Cheiranthus sulphureus Banks et Soland. 1794, in Russell, Nat. Hist. Aleppo, ed. 2, 2: 257. = Sterigma sulphureum (Banks et Soland.) DC. 1821, Reg. Veg. Syst. Nat. 2: 580.
= Sterigma brachypetalum Boiss. 1842, Ann. Sci. Nat. Bot., sér. 2, 17: 386.
13. Sterigmostemum violaceum (Botsch.) H. L. Yang, 1987, Fl. Desert. Reipubl. Popul. Sin. 2: 65.
= Oreoloma violaceum Botsch. 1980, Bot. Zhurn. 65, 3: 426.
= Oreoloma sulphureum Botsch. 1980, Bot. Zhurn.
65, 3: 427. = Sterigmostemum regeliorum Kamelin et
D. A. German, 2001, Turczaninowia, 4, 3: 8, non S. sulphureum (Banks et Soland.) Bornm. 1911.
Key for species of Sterigmostemum
1. Ovules 4-8 per ovary; fruit indehiscent, 3-4 times wider than pedicels.............................................................................. 2.
+ Ovules (8)10-50 per ovary; fruit partially to completely dehiscent, or indehiscent, as wide as or up to 2(2.5) times wider than pedicels .................................................................. 4.
2. Fruit oblong to linear, terete-quadrangular in cross section, usually pendulous or patent-pendulous on reflexed or recurved pedicels.....................................6. S. elichrysifolium.
+ Fruit oblong-conical, suberect, divaricate-ascending or spreading, sometimes recurved, terete, on erect-ascending to spreading fruiting pedicels ................................................ 3.
3. Fruit straight, smoothly torulose; raceme bracteate; petals purplish to pale yellow with dark veins, slightly exceeding erect purple sepals..........................................3. S. billardierei.
+ Fruit curved, ribbed-moniliform; raceme ebracteate; petals uniformly bright yellow, distinctly exceeding ascending pale green sepals ..................................... 11. S. ramosissimum.
4. Styles in fruit (2.5)3-8 mm long, stigma lobes oblong to linear ............................................................................................ 5.
+ Styles in fruit to 2.5(3) mm long, stigma lobes semicircular to oblong............................................................................. 10.
5. Plants scapose; leaves exclusively basal; flowers zygomor-phic.................................................................2. S. anchonioides.
+ Plants with well-developed, branched, and usually leafy stems; flowers actinomorphic ................................................ 6.
6. Annuals; flowers yellow; styles narrowly cylindrical; fruit not or rarely slightly widened at base, completely dehiscent or irregularly transversely breaking into fragments ....
........................................................................................................7.
+ Perennials, occasionally biennials; flowers yellow, violet, or while; styles narrowly conical; fruit widest at base, dehiscent only in the upper part ............................................... 8.
7. Sepals erect-ascending; fruiting pedicels (2)3-8(10) mm long; fruit usually covered throughout with long glandular setae, very late dehiscent or transversely breaking into fragments................................................. 1. S. acanthocarpum.
+ Sepals erect; fruiting pedicels to 3 mm long; fruit not setose, dehiscent ................................................ 9. S. longistylum.
8. Plants sparsely glandular on sepals and pedicels, rarely throughout; petals 15-22 mm long; stigma lobes 2.54.5 mm long; ovules 40-50 per ovary ... 5. S. eglandulosum.
+ Plants densely glandular throughout; petals 8-18 mm long; stigma lobes 1-3 mm long; ovules 20-30 per ovary ...
........................................................................................................ 9.
9. Petals 14-18 x 5-7 mm; sepals 8-10 mm long, lateral pair distinctly saccate at base; stigma lobes 2-3 mm long;
leaves mostly deeply sinuate-dentate to pinnatisect ............
............................................................................. 13. S. violaceum.
+ Petals 8-12 x 3-4 mm; sepals 5-7 mm long, lateral pair not or slightly saccate; stigma lobes to 1.5 mm long; leaves
mostly shallowly sinuate-dentate to entire ............................
................................................................................7. S. fuhaiense.
10. Stigma not wider than style; lateral sepals saccate at base;
petals pink, creamy white or yellow, 14-18 mm long ..........
....................................................................... 10. S. matthioloides.
+ Stigma wider than style; lateral sepals not saccate; petals bright to dull yellow, (5)7-11(12.5) mm long................ 11.
11. Plants perennial, completely eglandular; fruit not widened at base, terete, straight or slightly curved, late dehiscent at least in the upper part ......................................4. S. caspicum.
+ Plants annual or biennial, occasionally perennial, glandular throughout or in the lower part, rarely eglandular; fruits not or widened at base, usually slightly tetragonal, straight or often contorted or circinnate, normally indehiscent ....................................................................................... 12.
12. Petals 6-8(9) mm long; fruiting pedicels erect-ascending to subappressed; fruit glandular .....................8. S. incanum.
+ Petals 9-11(12.5) mm long; fruiting pedicels divaricate-
ascending to divaricate; fruit eglandular .................................
.......................................................................... 12. S. sulphureum.
Acknowledgements
Curators and collection managers of the herbaria consulted (ALTB, G, LE, M, MW, OSBU, P, W, WU) are acknowledged for providing the opportunity of studying the collections. Maria S. Ivanova and Lisa Kretz are much thanked for laboratory work. Careful reviewing by Alexander N. Sennikov is highly appreciated. The study was partly supported by the German Research Foundation (Deutsche Forschungsgemeinschaft, DFG) (grant K02302-13 to Marcus A. Koch).
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