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Taxonomic Study of Capparaceae from Egypt: Revisited

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Taxonomic Study of Capparaceae from Egypt: Revisited ® The African Journal of Plant Science and Biotechnology ©2009 Global Science Books Taxonomic Study of Capparaceae from Egypt: Revisited Wafaa M. Kamel1* • Monier M. Abd El-Ghani2 • Mona M. El-Bous1 1 Botany Department, Faculty of Science, Suez Canal University, Ismailia, Egypt 2 The Herbarium, Faculty of Science, Cairo University, Giza 12613, Egypt Corresponding author : * [email protected] ABSTRACT The systematic revision of the Egyptian species belonging to the family Capparaceae based on fresh materials with extensive field observations and herbarium materials is reported. This revision showed the presence of four genera Boscia, Capparis, Cadaba and Maerua. Species delimitation in Capparis was re-evaluated, resulting in the recognition of four species of Capparis: C. decidua (Forssk.) Edgew, C. sinaica Veill., C. spinosa L. (with three varieties viz., spinosa, canescens and deserti) and C. orientalis Veill. The last species is proposed to be elevated from var. inermis to the species level. Our results revealed also that leaves, stipules, flowers and fruit characters are of significant taxonomic value. A key for identification of the genera, species and varieties is provided. Type, synonyms, and selected specimens for each species are also presented. _____________________________________________________________________________________________________________ Keywords: Capparaceae, Boscia, Capparis, Cadaba, Maerua taxonomy, Egypt INTRODUCTION more than one genus from any of the tribes recognized by the authors, monophyly was contradicted (Hall et al. 2002). Capparaceae is a medium-sized family of approximately Hutchinson’s classification is particularly unsatisfactory 40–45 genera and 700–900 species, whose members present because it splits two genera, Maerua and Cadaba, into dif- considerable diversity in habit, fruit, and floral features ferent tribes because they are polymorphic with respect to (Cronquist 1981; Heywood 1993; Mabberley 1997). The the presence/absence of petals. The failure of traditional flowers of Capparaceae are ecologically versatile and aes- tribal classifications suggests that the traditionally impor- thetically exciting (Endress 1992). Floral variation in Cap- tant morphological characters are more homoplasious than paraceae includes both actinomorphy and zygomorphy, a previously considered (Hall et al. 2002). wide range of stamen number (1–250), and pronounced Bremer et al. (1998) proposed to merge the families basal intercalary elongation zones that may produce gyno- Capparaceae and Cruciferae under the common name Bras- phores, androgynophores, and elongated stamens (Endress sicaceae. Undoubtedly, if one takes into account the new 1992). Capparaceae are pantropical in distribution, being results of molecular systematics (Kubitz 2002); it is inap- most conspicuous in tropical seasonally dry habitats. propriate to maintain the two former families as they are The two major subfamilies of Capparaceae, Cleomoi- traditionally defined. However, Hall et al. (2002) recog- deae and Capparoideae, are quite distinct and have even nizing that a family Capparaceae inclusive of Cleomoideae been elevated to familial status by some authors (Airy Shaw would be paraphyletic and recommend the recognition of 1965; Hutchinson 1967). Capparoideae (about 25 genera/ smaller, monophyletic family units instead, and accepted 440 species) are typically woody (shrubs to small trees) and Cruciferae Adans., Capparaceae Adans, and Cleomaceae have dehiscent or indehiscent fruits, which are fleshy. The Horan, as separate families. larger Capparoideae are further divided into four tribes by Fici (2004) investigated leaf surface and pollen features Pax and Hoffmann (1936) and into three tribes by Hut- in the intraspecific taxa of Capparis spinosa by SEM and chinson (1967) equivalent to his (Capparaceae). Although light microscope observations. The micromorphological evi- there is some agreement among these classification systems, dence, coupled with other phenotypic features, supports the most aspects of Capparaceae relationships remain unre- placement of this section at the base of the genus Capparis solved. Cleomoideae (about 8 genera/275 species) are gene- in the palaeotropical area. rally herbaceous and have dehiscent fruits with repla. In Capparaceae are represented in the wild Egyptian flora both subfamilies the type genus is by far the largest and by 4 genera, 10 species of wide ecological and biogeogra- houses the majority of the species: Cleome (200 species) phical range of distribution (Montasir and Hassib 1956; and Capparis (150–200 species). Täckholm 1974) while Boulos (1999) included Capparaceae The traditional tribes in Capparoideae described by Pax and Cleomaceae in one family (Capparaceae) containing 7 and Hoffmann (1936) divided Capparoideae into Cappari- genera and 21 species and 4 varieties. The Egyptian taxa of deae, with rounded buds, stellate or simple hairs, scales, sel- Capparaceae belong to the xerophytic communities (Zahran dom with glandular hairs, and no sap (16 genera); Maerueae, and Willis 1992; Abd El-Ghani and Marei 2006). Earlier with elongated cylindrical buds (3 genera); and Stixeae, studies on Capparaceae in Egypt focused mainly on leaf with flowers that typically have six sepals (5 genera). Simi- anatomy (Al-Gohary 1982), seed morphology (Al-Gohary larly, Hutchinson (1967) divided Capparoideae (equivalent 1997) and pollen morphology (Khafagi and Al-Gohary to his Capparaceae) into Capparideae, with bisexual flowers 1998). The systematic treatment and phylogenetic affinities and more than two ovules (25 genera); Cadabeae, with bi- of the Egyptian species of Capparis is still obscured, and sexual apetalous flowers and more than two ovules (8 unsolved problems concerning their biology and taxonomy genera); and Apophylleae, with unisexual flowers with one need further verification and confirmation. or two ovules (2 genera). In all cases where they sampled Capparis is a large and polymorphic genus with a Received: 3 December, 2008. Accepted: 30 October, 2009. Original Research Paper The African Journal of Plant Science and Biotechnology 3 (1), 27-35 ©2009 Global Science Books world-wide distribution in the tropics and subtropical zones. Microscope. Phytogeographical territories for the revised speci- Infrageneric treatments have been proposed for it by vari- mens are those proposed by El Hadidi (2000, Fig. 1). ous authors ( De Candolle 1824; Endlicher 1836; Grisebach 1859; Eichler 1865; Pax and Hoffmann 1936), whereas SYSTEMATIC TREATMENT Jacobs (1965) published a revision of the south Asian and Pacific taxa, no modern global classification exists other Division: Magnoliophyta than Hutchinson’s 1967, who atomized Capparis into seve- Class: Magnoliopsida ral small genera that few other than himself have accepted. Order Type: Eudicots-Rosids II In so far as they are natural units, we consider them to be Order: Brassicales sections of Capparis. According to Iltis (2001) there are Family: Capparaceae (APG 2003) about 160 Capparis taxa world-wide, of which more than two third are present in the old World. Täckholm (1974) 1. Fruit dry, globose, indehiscent, warty or pitted, with recognized 6 species of Egyptian Capparis, whereas, Bou- crustaceous exocarp …………………………….. Boscia los (1999) classified the genus as 3 species and 4 varieties. – Fruit succulent, baccate, with seeds surrounded by a little Ahmed et al. (2007) investigated two Egyptian Cap- pulp …………………………………………………..... 2 paris species namely Capparis cartilaginea and C. deserti 2. Leaves with hooked spiny stipules .....................Capparis for their glucosiolates and rutin content. From C. cartila- – Leaves exstipulate ………………..…………................. 3 ginea four isothiocynates were isolated and identified using 3. Stamens many 20 – 40 inserted on torus …….… Maerua GC and EI/MS techniques. In addition, two other compounds – Stamens 5, inserted on petaloid androphore… Cadaba were isolated and identified from Capparis deserti. Abd El-Ghani et al. (2007) investigated the leaf archi- CAPPARACEAE Juss. tecture of 19 species belonging to 7 genera of Egyptian Capparaceae. The venation pattern showed mostly pinnate Herbs, shrubs or trees, sometimes woody climbers; leaves brochidodromous or craspidodoromous. Leaf architecture alternate, or rarely opposite (sometimes deciduous), glab- helped to distinguish the investigated species rous or furnished with glandular or eglandular hairs (Figs. 2, The objectives of the systematic treatment are to give an 3); hairs one to many celled, simple or branched, sometimes updated survey of the occurrence of members of the Cap- peltate, thorn like or glandula. Leaves, if present, simple or paraceae in Egypt including a key to all genera and species palmately compound, petiolate, stipulate or exstipulate; and descriptions, data on distribution and additional infor- stipules, if present, inconspicuous or thorn-like. Inflores- mation on the ecology in order to permit easier identifica- cence of terminal or axillary, racemes, corymbs, sometimes tion of material for the area and form a base for further of solitary or clustered flowers. Flowers actinomorphic or investigations. zygomorphic, hypogenous, with open or closed aestivation, bisexual or very rarely unisexual. Perianth usually 4-merous MATERIALS AND METHODS (sometimes tubular or ± perigynous basally, or very rarely undifferentiated); sepals 4 (-8), free or connate; petals 4-16 The materials used in this study were fresh materials collected or absent, free and
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