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516                GoT mez-Campo et al.—Epicuticular Wax Columns in Brassica

       Chromosome             T     1. Brassica accessions utilised in this research
       number (n)
        8          Accession  Species                                                        Seed source\origin
        9           number

       10          0049       Brassica nigra (L.) Koch                                       Vejer (Spain)
       17          6824       Brassica oleracea L.                                           Granville (France)
       18           —         Brassica oleracea L.                                           Cult. commercial sample
       19          5971       Brassica alboglabra Bailey                                     Gatersleben (Germany)
                   5719       Brassica bourgeaui (Christ) Kunze                              Tijarafe, La Palma, Canary
                   6813       Brassica montana Pourr.                                        Ste. Margueritte (France)
                   6801       Brassica montana Pourr.                                        Palmaiola (Italy)
                   3820       Brassica incana Ten                                            C. Tindari, Sicily (Italy)
                   6558       Brassica incana Ten                                            Capri (Italy)
                   6579       Brassica rupestris Rafin                                        Cinisi (Sicily, Italy)
                   6582       Brassica illosa Biv.                                           Castelmare di Golfo (Italy)
                   3821       Brassica illosa subsp. drepanensis (Caruel) Raim. & Mazz.      Mte. Erice, Sicily (Italy)
                   3819       Brassica macrocarpa Guss.                                      Egadi, Sicily (Italy)
                   3814       Brassica insularis Moris                                       C. Caccia, Sardinia (Italy)
                   6020       Brassica cretica Lam. subsp. cretica                           Diakoftos (Greece)
                   6021       Brassica cretica subsp. aegea (Heldr. & Hal.) Snogerup et al.  Euboea (Greece)
                   6344       Brassica cretica subsp. aegea (ibid.)                          Limnos (Greece)
                   7344       Brassica hilarionis Post                                       Kyrenia range (Cyprus)
                   8431       Brassica rapa L. (canola)                                      Prof. J. P. Tewari
                              Brassica carinata Braun                                        Dr. A. de Haro
                    —         Brassica juncea (L.) Czern                                     Bot. Garden, Upsala
                   0184       Brassica napus L. (canola)                                     Prof. J. P. Tewari
                   8432

(two subspecies). B. macrocarpa (Fig. 1 B), B. montana, B.        Henceforth, at least five lines of partial or total hair loss
cretica and B. hilarionis. Only in B. illosa were they          and parallel development of waxes can be distinguished
accompanied by abundant hairs. B. insularis columns were        (Fig. 2). (a) B. rupestris is geographically confined to the
intermediate between LC and SC.                                 Sicilian area of Palermo, between the distribution areas of
                                                                B. illosa and B. incana. B. rupestris has not completely lost
  Thirdly, there were netted columns (NC) formed by LC-         its hairs and presents no particular difference in the structure
like elements, apparently interlinked and often topped with     of its LC waxes compared with the last two species. (b) B.
transversal columns and wax scraps. They were never             insularis from Corsica, Sardinia and Tunisia has no hairs at
accompanied by hairs. Only B. oleracea, B. bourgaei (Fig.       all, while it exhibits waxes of intermediate size between
1 C) and B. albograbra showed this type of wax structure.       those found in the Sicilian species. (c) B. macrocarpa,
At first sight, the NC pattern appeared as an artefact but       isolated in distribution in the Egadi Archipelago (NW
the same structure was seen whenever observations on            Sicily), has no hairs, while its waxes are SC as in the
B. oleracea were carried out. Transversal columns are           neighbouring B. illosa. (d ) B. cretica and B. hilarionis, from
often shorter than those shown in Fig. 1 C.                     Greece, Turkey and Cyprus, have undergone parallel
                                                                evolution (waxes of SC type and no hairs). The derivation
  No waxes could be found in the leaves of the accessions       of B. cretica from Sicilian material is probably indirect and
of B. nigra, B. rapa and B. juncea studied.                     could be related either to Tunisian B. insularis or to Adriatic
                                                                B. incana. (e) The last (fifth) line apparently derives from B.
                           DISCUSSION                           incana (on geographic and morphological grounds). It
                                                                persists with B. montana, growing on the coasts of Italy,
It has been hypothesized (Go! mez-Campo and Gustafsson,         France and NE Spain. However, it has no hairs and its wax
1991) that the two densely haired Sicilian species, B. illosa   columns are more like the SC type. Further to the west, the
and B. incana, are the most primitive within the group, since   Atlantic kale, B. oleracea, is also glabrous and shows NC
they also show other primitive characters, such as the          waxes.
presence of two (three) seeds in the pod beak (primitive only
for this group). Genetic variation is much higher in B.           NC waxes are also present in B. bourgeaui and B.
 illosa (where four subspecies have been recognized). Other     alboglabra. Although these taxa have been listed as species
members of the group may be considered more evolved             in Table 1, the opinion that both should be included within
since they have less or no hairs and their beak is further      the intraspecific variability of B. oleracea is widespread
reduced to hold no more than one seed. In this respect, we      (Bothmer, Gustafsson and Snogerup, 1995 ; Lanne! r, 1998).
could consider both SC and LC waxes to be associated with       Our results strongly support this opinion. B. bourgeaui is a
the primitive condition represented by B. illosa and B.         rare taxon from the Canary Islands, of uncertain origin and
incana. As shown here, the presence of hairs and waxes is       might merely represent an ancient escaped cultivar of B.
not mutually exclusive in the Sicilian brassicas. On the other  oleracea. The same can be said for B. alboglabra, a cultivated
hand, there is no particular reason why SC and LC waxes         form which is supposed to have reached China in ancient
should be more primitive.                                       times.
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