Taxonomic review of the genus Crambe sect. Crambe (Brassicaceae, Brassiceae)

Prina


Introduction
The genus Crambe L. (Brassicaceae, Brassiceae) has an extensive area of distribution that goes from the Macaronesian archipelagoes to the West of China and North of India and from the Arctic Polar Circle on the Scandinavian Peninsula to 5º Latitude South in the North of Tanzania.It has representation in the Macaronesian, Euro-Siberian, Mediterranean, Sindico-Saharan, Irano-Turkish and Sudan-Zambezian (Ethiopia and Tanzania) regions (Leppik & White, 1975).Candolle (1821), based mainly on the dimensions and shape of the proximal joint of the fruit, made a first subdivision of the genus in three section: sect.Dendrocrambe DC., sect.Leptocrambe DC. and sect.Sarcocrambe DC.This infrageneric division closely corresponds to the geographical areas of distribution of the taxa it includes.Sect.Dendrocrambe is endemic to Macaronesia, Sect.Leptocrambe is distributed throughout the Mediterranean basin as far as the East of Africa and sect.Sarcocrambe has an area of distribution that goes from the East of Europe to the steppes and mountains of Central Asia.The type species of the genus (C.maritima L.) was designated by Green (1925) and belongs to sect.Sarcocrambe (Candolle, 1821) which thus becomes the typical section, Crambe sect.Crambe.

Materials and methods
The classical methodology used in plant taxonomy has been followed, based on the analysis of morphological characters of herbarium material from BM, E, EVIN, K, MA, MPU, VIR and W. In some cases we had living plants obtained from the seeds supplied by the U.S.D.A.-NCRPIS (United States Department of Agriculture-North Central Regional Plant Introduction) and the Bansem-UPM (Banco de Semillas de la Universidad Politécnica de Madrid), and grown in the Campos de Prácticas de la Escuela Técnica Superior de Ingenieros Agrónomos de la Universidad Politécnica de Madrid (ETSIA-UPM).For reasons of space, of the approximately 300 herbarium vouchers studied, only one sample per country is assigned for each taxon.

Taxonomic treatment
The species corresponding to sect.Crambe s.l., appear in the system proposed by Khalilov (1991a), distributed in four different sections: sect.Crambe s.str., sect.Orientecrambe I.I.Khalilov, sect.Astrocrambe I.I.Khalilov and sect.Flavocrambe I.I.Khalilov.The distinctive characters considered for sect.Orientecrambe are the fruits with a conspicuous, cylindershaped proximal joint, and an ellipsoidal, four-ribbed distal joint; but none of these traits are exclusive to the species included by Khalilov in this section.In a later study based on the characteristics of the pollen grain Khalilov & Archangelsky (1991) recognizes that neither can these characters distinguish this section of sect.Crambe s. str.Khalilov (1991b) subdivides sect.Orientecrambe into two subsections, subsect.Orientecrambe and the subsect.Junceae I.I.Khalilov differentiating between them by the presence of strengthened edges on the distal joint of the fruits (in subsection Orientecrambe) or their absence (in subsect.Junceae), in direct contradiction with the main trait with taxonomic value on which the creation of the new section was previously based.
The monotypic section Astrocrambe is characterised, according to the author, by its particular fasciculate hair indument, of C. shugnana Korsh.The presence of this type of indumenta has also been found in C. alutacea Hand.-Mazz., a species which Khalilov (1991a) does not mention in his work, which led to the preliminary solution of including this last species in the new section (Prina, 1998).However, neither do later taxonomic analysis of the entire genus (Prina, 2000) sustain sect.Astrocrambe; in fact, C. alutacea, apart from the traits of indument, is morphologically closer to C. orientalis L., C. koktebelica (A.Junge) N. Busch and C. hedgei I.I.Khalilov.Sect.Crambe ϵ sect.Sarcocrambe DC., Syst. Nat. 2: 650. 1821, nom.inval.
Hemicriptophytes or chamaephytes, from glabrous to densely pubescent, with either simple or fasciculate hairs.Fruit with proximal joint from obconical to cylin-der-shaped, poorly developed and sterile, except for C. shugnana and C. koktebelica, and the distal joint well developed, spheric or ellipsoidal, from four strengthened edges to ± rugose, bearing a single seed.Spherical or bifacial seed, smooth testa or slightly reticulate, with highly developed surrounding funiculus.
KEY FOR THE SPECIES OF THE SECT.CRAMBE C. cordata Willd., Enum Pl. Suppl.: 43. 1813, nom. inval.Hemicriptophyte, profusely branched from the base, 1-1.5 m, lignified caudex up to 15-20 cm in diameter.Stem glabrous.Basal leaves bearing long petioles, with cordate, individed and irregularly serrated blade, of 10-15 × 10-12 cm, leaves on stem ovate or obovate, interrupted, the terminal segment with cordate base and 1-2 lateral segments.Glabrous sepals, of 2.5 × 1.5 mm, the middle ones oblong and the lateral oblong-ovate.White petals, of 4.5-5.5 × 2.5-3 mm, with oblong blade and ruffed margin, narrowed in a nail of 0.8-1 mm.Stamen filaments of 2.8-3 mm with the concave ventral face applied to the ovary with one subapical tooth, the sides shorter and without tooth.Fruit with very reduced and sterile proximal joint; distal joint globose, 4.5-5 mm in diameter, slightly strengthened at the edges.
Habitat and distribution.Native to the steppes and plains to the north of the Caucasus.Can be found spontaneous in the S of Great Britain (Jalas & al., 1996).Hemicriptophyte 1.5-2 m, with a monopodial stem, branched in its terminal part with single stem and with intensely lignified basal region, with scattered hairs.Basal leaves 30-50 × 50-55 cm, bearing long petioles, cordate to sub-kidney shaped blade and lobed or serrate margin, cauline leaves subsessile, glabrous or with sparse hair.Sepals 4-4.5 mm, all very similar, with stiff antrorse hairs on the back.Petals 6-7 × 4-4.5 mm, white, with oblong blade, narrowed to The subordination of this entity to C. cordifolia is based solely on the difference of greater thickness and less hairiness of the leaves and that its fruit is somewhat smaller.Jafri (1973) states that some specimens from the region of Baluchistan show intermediary characters, which has been confirmed in the study of the specimen J. H. Lace 3751.Boissier himself, in the protologue of C. kotschyana, points to its affinity with C. cordifolia and only points to differences of little significance and great variance such as the size of the flowers and the degree of pubescence.Hedge (1968), while assigning it a specific rank, recognizes this affinity and the little relevance of the characters that differentiate them.Furthermore, both taxa have a different geographic distribution.The specimens with less hair, with flowers and fruit of greater size and leaves less profoundly divided, which are here designated to the subsp.kotschyana, are distributed around the eastern region of Iran and S of Afghanistan to Pakistan and to India, whereas those of smaller size, in general with more profoundly divided leaves and more densely-haired, corresponding to the subsp.cordifolia, are limited to the region N of the Caucasus.
Habitat and distribution.Disjunct distribution, on the coasts of the Northern Sea and the Baltic Sea, from France to Finland, spreading towards Eastern Europe, probably through cultivation, and on the coasts of the Black Sea, in the region of Crimea.Grown as a vegetable in the Caucasus region (Czerniakowskaya, 1939).The fruit is adapted to hydrocory (Scott & Randall, 1958).The materials from the Black Sea have been considered by Schulz (1919), among others, as a different taxon, be this as a species under the denomination A. Prina Crambe pontica Stev.ex Rupr.or with this species name in a rank subordinate to C. maritima.No differentiating morphological traits have been found between samples from one region or the other, for which reason we consider it convenient to keep all this material under the same species without acknowledging intraspecific taxa.

Representative specimens
There has been one citation for the coast of the Dead Sea; however, according to Dinsmore (1932) this must be an erroneous citation, possibly confusing it with Cakile maritima L. Schulz (1919) points to the same mistake in older citations of the species on the coast of the Mediterranean.The possible occurrence of this species on the coast of Galicia (Spain), cited by Planellas (1852), although without mentioning the herbarium voucher, is more that questionable.with one subapical tooth of 0.2-0.3mm.Fruit with proximal joint obconic and sterile, and distal joint spherical, of 5.5-6 mm in diameter, smooth or slightly rugose, single-seeded.

C. putaria
Habitat and distribution.It grows on steppes and hills rich in clay and limestone from Eastern Europe to the Caucasus.The adult plant usually breaks up at the base and thus becomes an organ of dispersal (tumbleweed).It has been considered endemism in the region of the Lower Volga and the desertic area between the Caucasus and the Caspian Sea (Czerniakowskaya, 1939); however samples have been studied that were gathered in Turkey and Hungary whose characters correspond to those of this variety, thus considerably extending its area of distribution.

Representative specimens
The differentiating morphological characters do not justify species status, nor present a differential area of distribution, for which reason we confirm its status as a variety, following Boissier.

Habitat and distribution.
Steppes and clay hills in the region of Crimea and S of the Caucasus.
Rarely collected, only type specimen studied.Czerniakowskaya (1939) recognises f. oxycarpa, and brings it up to the rank of variety on the basis of the acuminate fruits at both ends.Only one of the isotypes examined contains a sheath with several mature fruit which possess this trait, which however is not present in the fruits still attached to the specimen, for which reason the identity of this sample remains doubtful.As other samples with these traits have not been found and the remaining isotypes were in a very bad state of conservation, we felt it convenient to follow Schulz (1919), who does not recognize this variety.Curiously the fruit of this species is extremely similar to several Macaronesian species of sect.Dendrocrambe, especially in their acuminate and rugose distal joint, although the size of the fruit of C. steveniana is considerably larger.Branched hemicriptophyte of 60-100 cm, stems with strengthened edges, glabrous or with sparse retrorse and patent hairs.Glabrous basal leaves or with very sparse stiff cilia along the veins and margin, blade ovate or ovate-lanceolate, highly pinnatipartite, with 4-6 lateral segments oblong to lanceolate, markedly sinuate-serrate to highly pinnatipartite, the caulinar leaves oblong, serrate, the distal leaves sublinear, generally whole.Sepals 3.4-4 mm, oblong.Petals (5)-5.5-6× 3-4 mm, white, with oblong blade sharply narrowing to a nail.Staminal filaments of 3.5-4 mm, with one tooth in the apical area.Fruit with proximal joint obconical 3.5-4 mm, the distal joint tetragonous spherical or barely ovoid, of 7-8 mm in diameter, clearly veined and rugose.

Crambe grandiflora
Habitat and distribution.Known on the Taman peninsula and the estuary of the Kuban river in the Russian region of Krasnodar and in the region of Dagestan, its area now spreads considerably towards the S, including Turkey and Iran.In accordance with Schulz (1919), C. grandiflora is close to C. tataria.While the fruit with four strengthened and somewhat rugose edges remind us of the fruits of the latter, the rest of the plant is morphologically more similar to C. koktebelica and C. orientalis, especially in certain glabrous forms of this last species.The greater size of the flowers and in general of the fruit, however, indicate the convenience of keeping this taxon.

Representative specimens
We have been able to study the 8 vouchers collected by N.A. Busch and determined by Czerniakowskaya to be C. lipskii Czerniak.On one of these, in the handwriting of that author, the word typus is accompanied by a brief description and a drawing in pencil of a petal, preliminary elements of a diagnosis which however would seem not to have been published.From the study of this material and their comparison with the type specimen of C. grandiflora DC. we deduce that we are dealing with the same taxonomical entity.Three varieties are distinguished and are differentiated by the following characters.7a.C. edentula var.edentula (Fig. 4) Chamaephyte of 45-70 cm, with thin root, lightly lignified, and stems with multiple strengthened edges, with leaves at proximal end.Basal and proximal cauline leaves with stiff hairs, especially along the veins and in the petiole, blade 4-12 × 4-10 cm, lyratepinnatifid, with l terminal segment ovate-cordate, distal caulines linear-subulate, subsessile or with short Taxonomic review of Crambe petiole, of 2-3.5 cm.Sepals 2.8-3 mm, oblong, subequal, glabrous.Petals 4.5-6 × 2.5-3 mm, white, with blade oblong-obovate.Staminal filaments 2.3 mm, with no apical tooth and with small wings.Fruit with proximal joint 0.5-0.8mm, distal joint widely ovate to sub-spherical 3.5-4 mm in diameter, glaucous, with smooth surface, with a viable seed and, sometimes, also vestiges of an accompanying seed.

Crambe edentula
Habitat and distribution.Rocky slopes, holes and fissures in the N of the Caucasus and gypsum and limestone soils on the coast of the Caspian Sea.Although this presents morphological similarities to C. kralikii Coss.and C. hispanica L., both of which belong to sect.Leptocrambe, the scarce development in length of the proximal joint leaves little doubt that it should be included in the section Crambe.from the typical variety in the greater size of its flower.The sample studied did not have fruit.

Representative specimens
Habitat and distribution.Shares area with typical variety.

Representative specimens
Only type specimen was studied.Czerniakowskaya (1925) describes C. edentula var.cretacea and later (Czerniakowskaya, 1929) raises it to the rank of species.Among the several samples studied therein, the first to be cited is the sample Sintenis 1581, which had been used to describe C. juncea b glabrata (Freyn, 1903).Later, Czerniakowskaya (1939) omits any allusion of the species name cretacea but undertakes a critical analysis of the sample Sintenis 1581 and on this basis proposes C. edentula var.freynii Czerniak., which is an illegitimate name, since in the varietal range, the epithet used by Freyn & Sintenis holds priority.These authors seem to have been unconvinced of the systematic position of their variety, noting that the absence of teeth on the stamen distanced it from C. juncea, but the indicated no affinity with C. edentula.Distinct from the typical variety in its crass leaves at the base, with petioles of 5-8 cm, slightly widening at base and blade 10-12 × 9-10 cm, widely ovate and reduced at terminal segment, with sinuate-serrate margin with adpressed and rough hairs.
Habitat and distribution.Grows in the region of the Balkhan range (Turkmenistan) and in the surroundings of the eastern coast of the Caspian Sea.
Habitat and distribution.Grows in salty soils in the region of the Caucasus and to W of Turkmenistan, form which it was cited for the first time.
Habitat and distribution.Grows in salty or marlstone soils in the region of the Pamir Alai in Kyrgyzstan and Taxonomic review of Crambe on the steppe of Kyzyryk-Dara, in Uzbekistan; it is here documented for the first time for Afghanistan.
Habitat and distribution.Widely distributed from the Eastern Mediterranean to Central Asia, in steppes and mountains with clay soils, between 300 and 2500 m.Usually behaves as a weed (Zohary, 1966).Is differentiated from the subsp.orientalis by the greater size of its petals of 3.8-4.2mm, the sepals are also tainted yellow, and the general glabrescence of the plant.Given that plants of these characteristics have a defined area of distribution, it was decided to propose the subspecific status.Bornmüller (1905) describes C. persica var.glaberrima based on the specimen Bornmüller 6305, indicating in the protologue "Tota planta glaberrima".Later, Schulz (1919) transfers the variety to C. orientalis and Mouterde (1970) gives it the rank of species.The indumentum of the species is extremely variable, from totally glabrous to totally hairy plants.The stems are in general glabrous and smooth, although there are specimens which have striated stems with retrorse hairs.The presence or absence of indumentum on the stems is independent of the hairiness of the leaves, just as it is independent of the variety to which they belong.There are also totally glabrous specimens which may be confused with C. pinnatifida.The following material studied presents hairs on the stems: 13.Crambe grossheimii I.I.Khalilov, Bot. Zhurn. 75(11): 1572. 1990 Taxonomic review of Crambe Ind. loc.: "Caucasus, Transcaucasia australis, RSSA Nakhitschevan, prope salifodinam".

Representative specimens
Habitat and distribution.Grows on rocky and arid hills of N Afghanistan, between 2000 and 3000 m.Hemicriptophyte of 1.5-2.5 m; base with very dense and retrorse pubescence in young plants, glabrous or with very short hairs and sparse in adult plants.Basal leaves petiolate, blade 20-30(60) × 9-8(25) cm, discolour, lighter on the lower side, ovate-elliptical, pinnatisect, with 4-6 pairs of lateral segments, with dense pubescence of hairs from stiff to lying on both surfaces, the caulines of 2-5 cm, sparse, very reduced, from linear to oblanceolate.Upright pedicels, never adpressed.Sepals 2-2.5 mm, glabrous, colour green glabrous.Petals 4.5-5.5 mm, with obovate blade, sharply narrowing in a nail of c. 0.3 mm, white.Staminal filaments with one subapical tooth of c. 0.3 mm.Fruit with proximal joint short obconical, of 1-1.4 mm, with aborted seed primordia, the distal joint globose, with dark and prominent veins which give it a slightly rugose quality, of 3.5-4 mm in diameter, single-seeded.

Representative specimens
Habitat and distribution.Grows on the Taman peninsula and the coasts of Ukraine on the Azov Sea and the Black Sea, and in Russia in the region of the Volga delta on limestone-clay soils.Tarchankut, ca. Olenevka, 26-V-1984, Tsevelev & al. 1251 (LE).This is a species related to C. juncea and to C. orientalis.It was described as a variety of the first taxon and later elevated by Busch (1909) to the rank of species, a criterion followed by Czerniakowskaya (1939); it was also considered as a variety of C. orientalis (Schulz, 1919).Its specific status is supported by the larger size of its flowers compared to C. orientalis, its rougher and shorter hairiness than that of C. juncea; furthermore it grows in coastal areas of marine influence where neither of the other species are to be found.Hemicriptophyte up to 1.2 m., with roots with capacity to resprout, and hirsute basal rosette.Basal leaves with hirsute and pungent petiole, with blade 15-40 × 8-20 cm, ovate-elliptical, individed, irregularly sinuate-serrate in proximal leaves, later lyrate-pinnafitid to pinnatipartite, with 7-9 lateral segments, hirsute specially in the beam, the caulines similar although smaller and tending towards glabrous.Pedicel adpressed.Sepals 1.5-2 mm, glabrous, yellowish-green, lightly edged white.Petals white oblong, with blade 4.5-5 mm, sharply narrowing in a nail of 0.5 mm.Staminal filaments 2.5 mm, with one subapical tooth facing that of the opposite filament.Fruit with proximal joint cylindrical 1-1.5 mm, the distal joint subspherical, smooth, 3-4 mm in diameter, single-seeded.-NCRPIS 325.274, 25-IV-1998, A. Prina 1003 (MA).Busch (1909) described the var.aculeolata based on aculeate petioles and stems, later it was elevated to the rank of species by Czerniakowskaya (1939) and maintained at that rank by Khalilov (1993).The study of the type specimens of C. juncea and C. aculeolata has demonstrated that the taxonomical value of the latter is also to be found in the type specimen of C. juncea, for which reason they have been considered synonyms.Huber-Morath & Reese (1940) described C. parviflora on the basis of a sample collected in the region of Phrygia, in W Turkey.They pointed to similarities with C. aspera and differentiated it from C. tataria by the smaller size of its organs, especially the flowers.Later, this species was reduced to a variety of C. tataria by Hedge & Huber-Morath (1965).A few specimens of this region have been detected with small flowers, but do not coincide with C. parviflora in its other traits, above all because some of these individuals have mature fruit of the same size as that frequently found in C. tataria.Khalilov (1993) syonimizes it with the typical variety of this species.Given the great morphological variability of the group "tataria", we think it convenient to consider this taxon as dubious for the moment, until such time as the study of a greater number of specimens may confirm its status.
Taxonomic review of Crambe Sea Cabbage with leaves deeply cut, and an upright bran king stalk".This is probably a synonym of C. maritima L., but as the description offers so little information and as the author in the same work consigns C. maritima to a different taxon, we have decided to keep this name as dubious.