Page 2 - CAPPARIS_2006

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