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124 Annals of the
Missouri Botanical Garden
C. orientalis and C. sicula and overlap with C. C. parviflora, C. sicula). The Sudano-Zambezian
orientalis. The other two species, C. aegyptia and C. Region (Paleotropical) is inhabited by three species
ovata, are morphologically distinct, characterized by in tropical East Africa, littoral Arabia, western India,
their habit, stipules, and leaves (C. aegyptia: shrubs and southern Pakistan (C. mucronifolia, C. parviflora,
somewhat erect; twigs gray-green or blue-green with C. sicula). The African Karoo-Namib and Indian
waxy cover; stipules curved, retrorse; leaves obovate regions are inhabited by one single species each (C.
to ovate, bases and apices rounded, 2–3 3 1.8–3 cm; hereroensis and C. sicula, respectively). Capparis
C. ovata: shrub pendulous; adult leaves ovate, sicula is the most widespread species, stretching from
texture subcoriaceous; stipules curved, mostly an- the Western Mediterranean to the Himalayan Moun-
trorse). tains and the Rajasthan of India (Indian Region,
The objective of the present revision is to provide Paleotropical). Capparis hereroensis, in western Na-
a general taxonomy, based fundamentally on morpho- mibia, is the only species confined to the African
logical data for Capparis sect. Capparis. Karoo-Namib Region (sensu Takhtajan, 1986). En-
demic taxa are infrequent in this section, although C.
atlantica is endemic to the Atlas range in Morocco, C.
TAXONOMIC CONTEXT OF CAPPARIS SECT. CAPPARIS
mucronifolia Boiss. subsp. rosanoviana (B. Fedtsch.)
Capparis belongs to the subfamily Capparoideae Inocencio, D. Rivera, Obo ´n & Alcaraz is endemic to
(Capparaceae), that also includes Cadaba, Crataeva, central Tajikistan, and C. parviflora Boiss. subsp.
Morisonia, Boscia, and other New World and Old sphaerocarpa Inocencio, D. Rivera, Obo ´n & Alcaraz is
World genera (Hall et al., 2002). The different genera endemic to the western provinces (Nimruz, Farah,
overlap in molecular studies (Hall et al., 2002), Herat) in Afghanistan.
although a marked biogeographical distinction is
found between New World and Old World groups, ECONOMIC IMPORTANCE
recognized at the level of section or subsection
(Candolle, 1824; Bentham & Hooker, 1862; Hall Species of Capparis sect. Capparis are widely used
et al., 2002). A tentative infrageneric division is as food and medicine in the Old World. Capparis
presented in Table 1 in order to contextualize spinosa is almost exclusively known in cultivation.
Capparis sect. Capparis. There are many unresolved The supposed wild individuals of C. spinosa are often
taxonomic and nomenclatural issues for infrastructure remnants of ancient caper fields or escaped from
in Capparis. Therefore taxa in Table 1 are merely cultivation. Local caper cultivars and ethnovarieties
presented as a synthesis of major groups. are recognized throughout the Western and central
Mediterranean region (Spain, France, Italy, continen-
tal and insular). These principally belong to C.
GEOGRAPHIC DISTRIBUTION AND ENDEMISM
spinosa, but also to C. orientalis (Rivera et al., 1999,
Capparis is mostly a Pantropical genus, but section 2003b). Flower buds, consumed as brined product, are
Capparis is almost strictly Holarctic with six exclusive a rich source of the antioxidant phenolic compound
species (C. aegyptia, C. atlantica Inocencio, D. rutin (Inocencio et al., 2000). The use of Capparis
Rivera, Obo ´n & Alcaraz, sp. nov., C. orientalis, C. species by humans has been traced to the Prehistory
ovata, C. spinosa, C. zoharyi Inocencio, D. Rivera, and early historic times of Western Europe (1st–2nd
Obo ´n & Alcaraz, sp. nov.), one paleotropical species cent. A.D., Tongeren, Belgium), the Mediterranean
(C. hereroensis Schinz), and three in holarctic and region (9000–7400 B.C., Franchti Cave, Greece), and
paleotropical areas (C. mucronifolia, C. parviflora West Asia (9th–8th mill. B.C., Tell Mureybit, Syria),
Boiss., C. sicula). Here we name the floristic as evidenced by the presence of these species in
categories (kingdoms and regions) according to archaeological sites (Rivera et al., 2002).
Takhtajan (1986). Capparis sect. Capparis has its
maximum diversity in the Mediterranean Region MATERIAL AND METHODS
(Holarctic) with seven species (C. aegyptia, C.
atlantica, C. orientalis, C. ovata, C. spinosa, C. The herbaria and libraries of the following institu-
zoharyi, C. sicula). It is followed next by the Saharo- tions were consulted: BM, E, JEPS, K, MA, MUB,
Arabian Region (also Holarctic) with six species in OXF, P, RSA, and UMH. Specimens on loan were
North Africa and the Arabian Peninsula (C. aegyptia, received from BISH, BR C, G, HUB, HUJ, MARSSJ,
C. ovata, C. zoharyi, C. mucronifolia, C. parviflora, C. RNG, and US. Special collection trips were conducted
sicula). The Irano-Turanian Region (Holarctic) is in North Africa, western Asia, and Mediterranean
inhabited by four species extending along most of Spain to obtain fresh material for molecular studies,
West and Central Asia (C. aegyptia, C. mucronifolia, for the in vivo study of floral characters, and for