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Volume 93, Number 1                     Inocencio et al.                    123
            2006                                    Revision of Capparis Sect. Capparis

            Section 6. Quadrella DC. Angular fruit section.  Capparoideae are almost always fleshy in nature in
            Representative species: C. jamaicensis Jacq., C.  contrast to the often dry fruits of Cleomoideae and
            intermedia HBK, C. crotonoides HBK. The species  Brassicaceae. Floral symmetry, stamen number, leaf
            C. decidua (Forssk.) Edgew. with deciduous leaves, up  type, and fruit type all show homoplasy. Clades within
            to 20 3 3 mm in size, is included by Candolle (1824)  Capparoideae show a biogeographical pattern based
            in the genus Sodada Forssk.             on this sampling with a New World clade (all New
              This was followed in part by Bentham & Hooker  World representatives of Capparis, Belencita Karst.,
            (1862), although they created three new sections:  Morisonia L., Boscia Lam., and Cadaba Forssk.) and
            section Sodada (Forssk.) Benth. & Hook. including C.  an Old World clade (Maerua Forssk., Tylachium
            decidua; section Busbeckea (Endl.) Benth. & Hook.  Lour., Ritchiea R. Br., and Cadaba). The Old World
            including Australian species; and section Beautemp-  species Capparis tomentosa Lam. is nested within an
            sia (Gaud.) Benth. & Hook. including one American  otherwise New World clade (based only on trnL-trnF).
            species.                                Old World Capparis spinosa, the type of genus
              Zohary (1960) proposed new systematics based on  Capparis, is nested within an Australasian clade, with
            a partial geographical review of this genus restricted  Apophyllum anomalum F. Muell. (western New South
            to the Mediterranean region and West Asia. He  Wales) and Capparis callophylla Blume (Java) (based
            recognized two biogeographical groups: the tropical,  only on trnL-trnF).
            including Capparis decidua, C. cartilaginea, C.  DNA sequencing of the chloroplast rcbL gene
            mucronifolia Boiss., and the Mediterranean, including  nested New World Capparis hastata Jacq. and Old
            species that have lost their links with the tropical  World C. sandwichiana DC. in the same clade, but
            African stock (C. spinosa, C. sicula Veill., C.  sampling was restricted to these two Capparis species
            leucophylla DC.). All species belong to section  and comparison was made with Crataeva L. species
            Capparis in the sense of Candolle (1824).  (Rodman et al., 1998; Cummings et al., 2003).
              Jacobs (1965) attended to the Capparis species from  Taxonomic confusion of Capparis sect. Capparis is
            India to the Pacific, and organized them into four  reflected in the number of combinations and changes
            sections: 1. section Capparis, monotypic with C.  of rank, with frequent placement under C. spinosa
            spinosa, s.l.; 2. section Sodada, monotypic with C.  (Zohary, 1960; Jacobs, 1965; Higton & Akeroyd,
            decidua; 3. section Monostichocalyx Radlk., in a new  1991). Capparis spinosa has become a blanket
            circumscription containing most of the species  identification used to cover the scarce level of
            formerly included in section Capparis (5 Eucapparis  definition of the taxa in Capparis sect. Capparis
            DC.), with about 65 species in the area under revision;  (Greuter et al., 1984). This has both taxonomic and
            4. section Busbeckea, with 14 species in all.  economic implications, as Capparis flower buds are
              Higton and Akeroyd (1991) reviewed the diversity  the crude material used as commercial capers and
            of Capparis in the Mediterranean region, especially in  variability in this economic product is determined
            connection with C. spinosa, examining six species in  mostly by taxonomic differences (Inocencio, 2001;
            one section, which were finally reduced to a single  Inocencio et al., 2002). Molecular studies of Capparis
            species with two subspecies.            sect. Capparis have been scarce or restricted to small
              Hall et al. (2002) analyzed sequence variation for  sampling (Fici, 2001).
            a large sampling of Brassicaceae and Capparaceae,  A genetic fingerprinting technique (AFLP) was
            using two chloroplast regions, trnL-trnF and ndhF.  used by Inocencio et al. (2005) to examine the
            The results of parsimony and likelihood analyses  relationships among Capparis species. Genetic dis-
            strongly supported the monophyly of Brassicaceae  tances, based on AFLP data, were estimated for 45
            plus Capparaceae and recognized three clades:  accessions of Capparis species from Spain, Morocco,
            Capparaceae subfamily Capparoideae, subfamily  and Syria. The results of this analysis support the
            Cleomoideae, and Brassicaceae. Habit and fruit  differentiation of four (C. orientalis Veill., C. sicula, C.
            characteristics demarcate these three clades. All  aegyptia Lam., and C. ovata Desf.) of the five taxa
            Capparoideae are woody, which is the plesiomorphic  sampled. The fifth, excluded, species was C. spinosa.
            condition for the Brassicaceae and Capparaceae  The group of plants recognized as C. spinosa on the
            clades. The herbaceous habit is generally found in  basis of morphological characters such as shrub
            Cleomoideae and Brassicaceae. Indehiscent, fleshy  procumbent or somewhat erect, stipules usually weak
            fruits are plesiomorphic in Capparaceae and Brassi-  or vestigial, rarely strong, very long and thin (0.3–
            caceae and are the dominant fruit type of Cappa-  0.6 cm long), indument on leaves always very lax, and
            roideae. Brassicaceae and Cleomoideae both have  trichomes thick and long (300–500 mm long), is found
            dehiscent capsules with a replum, a synapomorphy  almost exclusively in cultivation. Several cultivars of
            shared by these two clades. The dehiscent fruits of  C. spinosa appear in an intermediate position between
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