THE GENUS GVENTHERA ANDR. IN BESS. (BRASSICACEAE, BRASSICEAE)

by CÉSAR GÓMEZ-CAMPO Departamento de Biología Vegetal, ETSIA, Universidad Politécnica de Madrid. E-28040-Madrid (España)

Resumen GÓMEZ-CAMPO. C. (2003). El género Guenthera Andr, in Bess. (Brassicaceae, Brassiceae). Anales Jard. Bot. Madrid 60(2): 301-307 (en inglés). Un grupo de nueve especies actualmente incluidas en Brassica difiere de todas las demás por varios caracteres, sobre todo por la porción estilar de sus pistilos, que siempre carece de pri- mordios seminales. Además, por su tallo subterráneo ramificado, que forma un cáudex con varias rosetas; sus hojas de enteras hasta profundamente pinnatífidas, pero nunca pinnatisec- tas; sus cotiledones solo muy ligeramente escotados, y sus semillas, que tienden a ser elip- soidales o aplanadas. Se propone agruparlas todas bajo la denominación genérica Guenthera Andr, in Bess. Se detallan los nuevos nombres para las especies y Subespecies y se añade una clave para diferenciar las especies. Palabras clave: taxonomía, Guenthera, Brassicaceae, Brassica.

Abstract GÓMEZ-CAMPO, C. (2003). The genus Guenthera Andr, in Bess. (Brassicaceae, Brassiceae). Anales Jard. Bot. Madrid 60(2): 301-307. A group of nine species -now included in Brassica— differ from all the other species in sever- al characters, mainly in the stylar portion of their pistils always without seed primordia. Also in their branched subterranean stem (caudex) with several leaf rosettes, their leaves entire to deeply pinnatifid but never pinnatisect, their shallowly notched cotyledons and their flattened, elliptic or ovoid seed contour. It is suggested to include these species under the generic de- nomination Guenthera Andr, in Bess. New ñames for the species and subspecies are provided, as well as a determination key for the species. Key words: taxonomy, Guenthera. Brassicaceae, Brassica.

INTRODUCTION highly polymorphic and suggests its splitting into no less than five genera: Brassica, Eru- Initially (xvm century), Brassica was an castrum, Brassicaria, Nasturtiops and Mela- ample and polymorphic genus, from which nosinapis, although he does not provide prop- many species were soon withdrawn to form er validations. In line with this view, it can be other new genera {Conringia, Eruca, Hirsch- observed that Brassica and Erucastrum feldia, Moricandia, etc.) or to enlarge other siliques show a seeded beak, while the stylar existing ones (Coincya, Diplotaxis, Erucas- portion of Brassicaria, Nasturtiops and Mela- trum, etc.) occasionally through some now nosinapis is asperm. Although GILLET & abandoned intermediate generic denomina- MAGNE (1879) recognise Brassicaria as a tions. By 1860, Pomel still finds Brassica genus, the last three ñames have been, in gen- 302 ANALES JARDIN BOTANICO DE MADRID, 60(2) 2003

eral, used to define either sections or subgen- Vella, etc.) is anterior to this evolutionary de- era (GODRON, 1848; O.E. SCHULZ, 1916; velopment while Brassica (as Erucastrum, SALMEEN, 1979; G6MEZ-CAMPO, 1999a). Coincya, Raphanus, etc.) is posterior. We The seedless stylar portion of Brassica ni- avoid to call a sterile stylar portion "beak", gra L. has always been a cause of confusion. since this is the general case with fruits in the In fact, the creation of sect. Melanosinapis Crucifer family. DC. (under Sinapis) was largely endeavored The absence of a beak shows correlations to fit this odd species. However, it is evident with at least five other significant characters. that, apart from the lack of a beak, all other The presence of a caudex is commonplace in characters showed by B. nigra are those Guenthera but never observable in Brassica. of Brassica, particularly those observable A caudex is a vertical buried stem branched in sect. Micropodium DC. which includes under the soil surface, with leaf rosettes at soil for instance B. fruticulosa, B. cossoniana, level, at the tip of each branch. Leaf scars or B. maurorum, B. spinescens, etc. Everything leaf remainings demonstrate its caulinar con- suggests that B. nigra is a true Brassica, sim- dition. A caudex is like a buried bush and it ply renouncing to its fertile beak during the represents an intermediate situation between evolutionary proccess leading to a strong chamaephytes and mono-rossette hemicripto- shortening of its fruit. phytes. The unique exception is the annual When the species corresponding to the habit showed by Guenthera amplexicaulis. Pomel's groups Brassicaria and Nasturtiops However, all other characters of this species are considered together and B. elongata is are those of Gunthera and, particularly, added, the resulting complex of species shows G. amplexicaulis shows strong affinities to a series of important characters in common G. dimorpha. Apart from this case, Gunthera species are hemicriptophytes with several (table 1). In our opinion they deserve the apli- rosettes, although there might be a single one cation of a generic denomination (Guen- in the juvenile stage or sometimes in the adult thera). stage (mostly in G. elongata). The scapiform habit for floriferous stems is very extended though species with scarcely branched leafy DIFFERENTIAL CHARACTERS aerial stems also exist. Rosette-forming habit is already conspicuous in the seedling devel- Among all the other differential characters, opment, a stage where Guenthera is already the absolute absence of seeds -or even seed easy to distinguish from Brassica. primordia- within the stylar portion of the pistil should be emphasised (character 5 in The typical lyrate-pinnatisect leaf silhou- table 1). The absence or presence of a seeded ette -so common in Brassica and most other beak has always been considered a relevant related genera with seeded beak-, can never character to distinguish genera within the be found in Guenthera. Guenthera leaves are tribe Brassiceae, where each option occurs in normally entire, lobed or pinnatifid, often aproximately one half of the 55 existing gen- deeply so. Only within the high variability era. Species with and without a beak had been present in the leaves of the G. repanda sub- maintained together up to the present only in species it is possible to find individual cases Brassica and Diplotaxis. The existence of a approaching a pinnatisect contour. seeded beak (heteroarthrocarpy) is both phy- POMEL (1860) already observed clear dif- logenetically and taxonomically important ferences in cotyledon shape which can be (G6MEZ-CAMPO, 1999b) because it is a singu- "emarginate or not". Such differences are lar morphogenetic achievement which ap- very patent and they were quantified by pears exclusively within the evolutionary ra- G6MEZ-CAMPO & TORTOSA (1974). In other diation of the tribe Brassiceae, while it is words, the cotyledon notch in Guenthera is completely absent in all other cruciferous much shallower than in Brassica. The cotyle- tribes. Guenthera (as Eruca, Sinapidendron, don petiole is also shorter. C. G6MEZ-CAMP0: THE GENUS GUENTHERA 303

TABLE 1 MAIN DIFFERENCES BETWEEN THE GENERA GUENTHERA AND BRASSICA

Guenthera Brassica 1. Mature seeds flattened, ovoid, elipsoidal 1. Mature seeds globose to spherical. or subglobose to globose. 2. Cotyledons only slightly emarginate 2. Cotyledons more deeply emarginate (notch 10 (notch 3-8% of the lamina length). y 25% of the lamina length). 3. Plants perennial, with floriferous stems arising 3. Plants annual, biennial or perennial but never from a caudex with several leaf rosettes. with a caudex. 4. Leaf entire to deeply pinnatifid (but never 4. Leaves lyrate-pinnatisect, sometimes runcinate pinnatisect). or becoming entire by reduction in the number of lateral segments. 5. Stylar portion asperm (seedless). 5. Stylar portion forming a beak with 1 to several seeds (heteroarthrocarpy). 6. Chromosome number n = 10,11. 6. Chromosome number n = 7,8,9 or 10.

With respect to the contour of mature Brassica balearica Koch has a seeded beak seeds, overlappings may exist among the and its polyploid nucleus includes the n = 9 species of both genera (BENGOECHEA & genome shown by B. oleracea and other close G6MEZ-CAMPO, 1975), but a tendency to Mediterranean relatives (SNOGERUP & PERS- sphericity is always much more marked and SON, 1983). However, its dwarfing facies constant within the species of Brassica. and compressed seeds suggest that it might Cromosome number n = 11 is present in be the product of a possible intergeneric cross Guenthera and absent in Brassica. Chromo- Brassica x Guenthera. Further cytogenetic some number n = 10 exists in both genera but studies would be necessary to elucidate this the deep morphological differences among case. The position of Brassica somalensis the species belonging to Brassica and Guen- Hedge & Miller-a suffruticose plant (without thera (as for instance Brassica rapa and a caudex) with asperm stylar portion and el- Guenthera repanda both with n = 10) suggest lipsoidal seeds- is more difficult to interpret. that it is a mere coincidence. HEDGE & MILLER (1977) suggest a possible Not all molecular studies with Brassica in- relationship with the genus Sinapidendron clude Guenthera species. However, dendro- from Madeira Island. In our opinion, this hy- grams obtained by WARWICK & BLACK, pothesis is probable. (1991,1993) with c-DNA. by PRADHAN & al. (1992) with c-DNA and m-DNA, or by INABA Guenthera Andr. in Bess.* Enum. PL Volhyn.: & NISHIO (2002) with sequences of nuclear 83(1822) genes do include a number of species here as- cribed to Guenthera. Constantly, they appear Typus: Guenthera elongata (Ehrh.) Andr. clearly separated from typical Brassica in Bess. species and, significantly, they are often clos- Guenthera [Giinthera] (1922) antecedes er to other taxa with asperm stylar portion be- other possible generic names as Brassicas- longing to genera as Eruca, Diplotaxis, etc. trum (1831) or Brassicaria (1879). On the other hand, differences showed by G. elongata and G. amplexicaulis are still Hemicriptophytes with a caudex (vertical deeper, a fact that is correlated to other mor- subterranean branched stem) with several leaf phological trends and reinforces the distribu- * Besser explicitly attributes the generic name and tion into sections proposed below. description to Andr6anszky. 304 ANALES JARDIN BOTANICO DE MADRID, 60(2) 2003

rosettes, more rarely biennial (G. elongata, Syn.: Brassica elongata subsp. pinnatifida partly) or annual (G. amplexicaulis). Florifer- (Schmalh.) Greuter & Burdet, Willde- ous stems leafy or leafless growing annually nowial5:64(1985) from the rosettes. Leaves entire, sinuate, or deeply pinnatifid, never pinnatisect. Silique Guenthera elongata subsp. subscaposa with a prominent mid-nerve in each valve and (Maire & Weiller) G6mez-Campo, comb, a stylar portion always sterile. Mature seeds nov. somehow flattened, ovoid, ellipsoidal or sub- Bas.: Brassica subscaposa Maire & Weiller, globose, rarely globose. Cotyledons only very Bull. Soc. Hist. Nat. Afrique N. 31:9 (1940) slightly notched. Chromosome number In = Syn.: Brassica elongata subsp. subscaposa 20 or 22, with some cases of polyploidy. (Maire & Weiller) Maire, Fl. Afrique N. 12: 168(1965) Three sections are distinguished and are defined by the characters which are expressed Sect. Nasturtiops G6mez-Campo, sect nov. in the proper place. Caules floriferi parce tantum ramosi, fo- liosi, foliis ovatis, amplexicaulibus. Gyno- Sections, species and subspecies phorum breve (infra 1.5 mm longum). Typus: Guenthera dimorpha (Coss. & Sect. Guenthera Dur.) Gomez-Campo. Aerial stems branched and leafy with Aerial stems only slightly branched, leafy leaves not amplexicaul. Gynophore 1,5- with leaves ovate in contour, amplexicaul. 6 mm. Gynophore < 1.5 mm. Guenthera elongata (Ehrh.) Andr. in Bess. Guenthera amplexicaulis (Desf.) Gomez- Enum. PI. Volhyn.: 83 (1822) Campo, comb. nov. Bas.: Brassica elongata Ehrh., Beitr. Naturk. Bas.: Sisymbrium amplexicaule Desf., Fl. 7: 159(1790) Atlant.2:81(1798) Syn.: Brasskastrum elongatum (Ehrh.) Link, Syn.: Brassica amplexicaulis (Desf.) Pomel Handb.2:318(1831) Mat. Fl. Atlant. 15 (1860), non A. Richard, Ten. Fl. Abyss. 1: 23 (1847), nom. illeg. Guenthera elongata (Ehrh.) Andr. subsp. Syn.: Brassica souliei subsp. amplexicaulis elongata (Desf.) Greuter & Burdet, Willdenowia 13: 86(1983) Guenthera elongata subsp. imdrahsiana (Quezel) Gomez-Campo, comb. nov. Guenthera amplexicaulis (Desf.) G6mez- Bas.: Brassica elongata subsp. imdrahsiana Campo subsp. amplexicaulis Quezel, Bull. Soc. Sci. Nat. Maroc 34: 304 Guenthera amplexicaulis subsp. souliei (1955) (Batt.) Gomez-Campo, comb. nov. Bas.: Camelina souliei Batt., Bull. Soc. Bot. Guenthera elongata subsp. integrifolia France 36: 218 (1889) (Boiss.) Gomez-Campo, comb. nov. Syn.: B. souliei (Batt.) Batt., Bull. Soc. Bot. Bas.: Brassica elongata var. integrifolia France 40: 262 (1893); B. amplexicaulis Boiss., Fl. Orient. 1: 394 (1867) subsp. souliei (Batt.) Maire & Weiller in Syn.: Brassica elongata subsp. integrifolia Maire, Fl. Afrique N. 12:191 (1965) (nom. (Boiss.) Breistr., Not. Syst. Bot. 13 (1942) illeg.) Guenthera elongata subsp. pinnatifida Guenthera dimorpha (Coss. & Dur.) (Schmalh.) Gomez-Campo, comb. nov. Gomez-Campo. comb. nov. Bas.: Brassica elongata var. pinnatifida Bas.: Brassica dimorpha Coss. & Dur., Bull. Schmalh., Fl. Ssredn. Jushn. Rosii 1:80 (1895) Soc. Bot. France 2: 306 (1855) C. G6MEZ-CAMP0: THE GENUS GUENTHERA 305

Sect. Brassicaria (Godr.) G6mez-Campo, Guenthera repanda (Willd.) G6mez-Campo comb. nov. subsp. repanda Bas.: Brassica sect. Brassicaria, Godr. in Gren & Godr., Fl. Fr. 1: 78 (1848) Guenthera repanda subsp. africana (Maire) Gomez-Campo, comb. nov. Plants scapose with aerial stems either Bas.: Brassica saxatilis subsp. africana aphyll or with 1-2(3) poorly developed Maire, Bull. Soc. Hist. Nat. Afrique N. 20: leaves. Gynophore < 1.5 mm. 13(1929) Syn.: Brassica repanda subsp. africana Guenthera desnottesii (Emb. & Maire) (Maire) Greuter & Burdet, Wildenowia 13: Gomez-Campo, comb. nov. 86 (1983); Brassica repanda subsp. nudi- Bas.: Brassica desnottesii Emb. & Maire, PI. caulis (Lag.) Heywood, Feddes Repert. Marocc. Nov. 2: 1 (1929) Spec. Nov. Regni Veg. 69: 121 (1964) Guenthera gravinae (Ten.) Gomez-Campo, comb. nov. Guenthera repanda subsp. almeriensis Bas.: Brassica gravinae Ten., Fl. Napol. 1: 39 (Gomez-Campo) Gomez-Campo, comb, nov. (1811) Bas.: Brassica repanda subsp. almeriensis Guenthera ioncholoma (Pomel) Gomez- G6mez-Campo, Anales Inst. Bot. Ca- Campo, comb. nov. vanilles 33:154(1976) Bas.: Brassica Ioncholoma Pomel, Nouv. Guenthera repanda subsp. blancoana Mat. Fl.Atlant: 360 (1875) (Boiss.) Gomez-Campo, comb. nov. Syn.: Eruca Ioncholoma (Pomel) O.E. Schulz, Bas.: Brassica blancoana Boiss., Diagn. PI. Bot. Jahrb. Syst. 54(119): 56 (1916) Orient, ser. 2,1:29(1854) Syn.: Brassica repanda subsp. blancoana Guenthera nivalis (Boiss. & Heldr.) G6mez- (Boiss.) Heywood, Feddes Repert. Spec. Campo, comb. nov. Nov. Regni Veg. 66: 153 (1962) Bas.: Brassica nivalis Boiss. & Heldr., Diagn. PI. Orient., Ser. 2,1:32(1854) Guenthera repanda subsp. cadevallii (Font Syn.: Brassicella nivalis (Boiss. & Heldr.) Quer) Gomez-Campo, comb. nov. O.E. Schulz, Bot. Jahrb. Syst. 54(110): 53 Bas.: Brassica saxatilis var. cadevallii Font (1916) Quer, Cavanillesia 7: 72 (1935) Syn.: Brassica repanda subsp. cadevallii Guenthera nivalis (Boiss. & Heldr.) Gomez- (Font Quer) Heywood, Feddes Repert. Campo subsp. nivalis Spec. Nov. Regni Veg. 66:154 (1962) Guenthera nivalis subsp. jordanoffii (O.E. Guenthera repanda subsp. cantabrica Schulz) G6mez-Campo, comb. nov. (Font Quer) Gomez-Campo, comb. nov. Bas.: Brassica jordanoffii O.E. Schulz, Bas.: Brassica saxatilis var. cantabrica Font Notizbl. Bot. Gait. Berlin-Dahlem 10: 111 Quer, Cavanillesia 7: 73 (1935) (1927) Syn.: Brassica repanda subsp. cantabrica Syn.: Brassica nivalis subsp. jordanoffii (Font Quer) Heywood, Feddes Repert. (O.E. Schulz) Akeroyd & Leadlay, Bot. J. Spec. Nov. Regni Veg. 69: 151 (1964) LinneanSoc. 106: 102(1991) Guenthera repanda subsp. confusa (Emb. Guenthera repanda (Willd.) G6mez-Cam- & Maire) Gomez-Campo, comb. nov. po, comb. nov. Bas.: Brassica saxatilis subsp. confusa Emb. Bas.: Sisymbrium repandum Willd., Sp. PI. 3: & Maire, Cat. PI. Maroc: 1007 (1941) 497(1800) Syn.: Brassica repanda subsp. confusa (Emb. Syn.: Brassica repanda (Willd.) DC, Syst. & Maire) Heywood, Feddes Repert. Spec. Nat. 2: 598 (1821) Nov. Regni Veg. 66: 153 (1962) 306 ANALES JARDIN BOTANICO DE MADRID, 60(2) 2003

Guenthera repanda subsp. dertosensis (Boiss. & Reut.) Heywood, Feddes Repert. (Molero & Rovira) Gomez-Campo, comb, Spec. Nov. Regni Veg. 66: 153 (1962) nov. Bas.: Brassica repanda subsp. dertosensis Guenthera repanda subsp. maritima Molero & Rovira, Collect. Bot. (Barce- (Willk.) G6mez-Campo, comb. nov. lona) 17(1): 99-101 (1987) Bas.: Diplotaxis brassicoides var. maritima Willk., 111. FLHisp. 1:141(1885) Guenthera repanda subsp. diplotaxiformis Syn.: Brassica repanda subsp. maritima (Maire) G6mez-Campo, comb. & stat. (Willk.) Heywood, Feddes Repert. Spec. nov. Nov. Regni Veg. 66: 153 (1962) Bas.: Brassica saxatilis subsp. repanda var. diplotaxiformis Maire, Bull Soc. Hist. Nat. Guenthera repanda subsp. saxatilis (DC.) AfriqueN. 19:82(1928) Gomez-Campo, comb. nov. Bas.: Diplotaxis saxatilis DC., Syst. Nat. 2: Guenthera repanda subsp. galissieri (Gi- 636(1821) raud) G6mez-Campo, comb. nov. Syn.: Brassica repanda subsp. saxatilis (DC.) Bas.: Diplotaxis galissieri Giraud, Bull. As- Heywood, Feddes Repert. Spec. Nov. soc. Pyren. Echange PI. 1-2: 21 (1892) Regni Veg. 69: 151 (1964) Syn.: Brassica repanda subsp. galissieri (Gi- raud) Heywood, Feddes Repert. Spec. Guenthera repanda subsp. silenifolia Nov. Regni Veg. 69: 150 (1964) (Emb.) G6mez-Campo, comb. nov. Bas.: Brassica saxatilis subsp. silenifolia Guenthera repanda subsp. glabrescens Emb., Bull. Soc. Sci. Nat. Maroc 15: 200 (Poldini) G6mez-Campo, comb. nov. (1936) Bas.: Brassica glabrescens Poldini, Giorn. Bot. Ital. 10: 181 (1973) Syn.: Brassica repanda subsp. silenifolia Syn.: Brassica repanda subsp. glabrescens (Emb.) Greuter & Burdet, Willdenowia 13: (Poldini) G6mez-Campo, Anales Jard. 86(1983) Bot. Madrid 56: 379 (1998) Guenthera repanda subsp. turbonis Guenthera repanda subsp. gypsicola (P. Monts.) G6mez-Campo, comb. nov. (G6mez-Campo) G6mez-Campo, comb, Bas.: Brassica saxatilis subsp. turbonis nov. P. Monts., Pirineos 28-30: 181 (1953) Bas.: Brassica repanda subsp. gypsicola Syn.: Brassica repanda subsp. turbonis Gomez-Campo, Anales Jard. Bot. Madrid (P. Monts.) J.M. Monts. & Romo, Willde- 50: 145 (1992) nowia 15: 64(1985)

Guenthera repanda subsp. humilis (DC.) Guenthera setulosa (Boiss. & Reuter) G6mez-Campo, comb. nov. G6mez-Campo, comb. nov. Bas.: Brassica humilis DC, Syst. Nat. 2: 598 Bas.: Eruca setulosa Boiss. & Reuter in (1821) Boiss., Diagn. PI. Orient., ser. 2, 5: 26 Syn.: Brassica saxatilis subsp. humilis (DC.) (1856) Maire, Fl. Afrique N. 12:202 (1965) (nom. Syn.: Brassica setulosa (Boiss. & Reuter) inval.) Cosson, 111. Fl. Atlant. 1: 32 (1882) Guenthera repanda subsp. latisiliqua (Boiss. & Reut.) G6mez-Campo, comb. KEY TO THE SPECIES OF GUENTHERA nov. Bas.: Brassica latisiliqua Boiss. & Reut. in 1. Aerial stems branched, leafy 2 Boiss., Diagn. PI. Orient, ser. 2,1:30 (1854) - Aerial stems unbranched, aphyll or with some Syn.: Brassica repanda subsp. latisiliqua small leaves 4 C. G6MEZ-CAMP0: THE GENUS GUENTHERA 307

2. Cauline leaves sessile, not amplexicaul, nomic and evolutionary significance of some juvenile gynophore 1.5-6 mm G. elongate characters in the Brassiceae. Bot. Journal Linnean - Cauline leaves amplexicaul; gynophore Society 69: 105-124. < 1.5 mm 3 GREN, J.C.M. & D.A. GODRON (1848). Flore de la France. 3. Caudex well developed, with branches up to HEDGE, I.C. & A.G. MILLER (1977). New and interesting 5-10(15) cm G. dimorpha taxa from NE Tropical Africa. Notes from the Royal - Caudex absent; plant annual G. souliei Botanic Garden. Edinburg. 35(2): 179-193. 4. Valves linear, 25-70(90) mm in length 5 INABA, R. & T. NISHIO (2002). Phylogenetic analysis of - Valves ellipsoidal, up to 20(25) mm in length... 7 Brassiceae based on the nucleotide sequences of the 5. Leaves all basal G. repanda S-locus related gene, SLR1. Theor. Appl. Genet. 105: - Some small caulinar leaves present 6 1159-1165. 6. Plant glabrous or subglabrous. Silique up to POMEL, A.N. (1860). Materiaux pour la flore atlantique. Caen. 2 mm wide G. nivalis PRADHAN, A.K., S. PRAKASH, A. MUKHOPADHYAY & - Plant hairy. Silique more than 2 mm wide D. PENTAL (1992). Phylogeny of Brassica and allied G.gravinae genera based on variation in chloroplast and mito- 7. Flowers purple or violaceous G. setulosa chondrial DNA patterns. Molecular and taxonomic - Flowers yellow 8 classifications are incongruous. Theor. Appl. Genet. 8. Plant 4-20 cm. Siliques c. 15 mm on pedicels of 85:331-340. 4-7 mm G. loncholoma SALMEEN, O. (1979). A systematic revision of the genus - Plant up to 40 cm. Siliques < 10 mm on pedicels Brassica L in the Mediterranean region. PhD. The- sis. Univ. of Reading (unpublished). 10-17 mm G. desnottesii SCHULZ, O.E. (1919). Cruciferae-Brassiceae. I. In: A. En- gler (ed.), Das Pflanzenreich. 70:19. SNOGERUP, S. & D. PERSSON (1983). Hybridization be- REFERENCES tween Brassica insularis Moris and B. balearica Pers. Hereditas 99: 187-190. BENGOECHEA, G. & C. GOMEZ-CAMPO (1975). Algunos WARWICK, S.I. & L.D. BLACK (1991). Molecular system- caracteres de la semilla en la tribu Brassiceae. Anales atics of Brassica and allied genera (Subtribe Brassici- last. Bot. Cavanilles 32: 793-841. nae, Brassiceae) - chloroplast genome and cytodeme GILLET, C.C. & J.H. MAONE (1879). Nouvelle flore congruence. Theor. Appl. Genet. 82: 81-92. frangaise, ed. 4. WARWICK, S.I. & L.D. BLACK (1993). Molecular rela- G6MEZ-CAMPO, C. (1999a). Taxonomy. In: C. G6mez- tionships in subtribe Brassicinae (Cruciferae, tribe Campo (ed.), Biology ofBrassica coenospecies: 3-32. Brassiceae). Can. J. Bot. 71:906-918. Elsevier. GOMEZ-CAMPO, C. (1999b). Seedless and seeded beaks in the tribe Brassiceae. Cruciferae Newsletter 21:11-12. Editado por Santiago Castroviejo GOMEZ-CAMPO, C. & M.E. TORTOSA (1974). The taxo- Aceptado para publicacidn: 28-VII-2003