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A Systematic Review of the Genus Abelmoschus (Malvaceae)

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A Systematic Review of the Genus Abelmoschus (Malvaceae) Rheedea Vol. 25(1) 14-30 2015 ISSN: 0971 - 2313 A systematic review of the genus Abelmoschus (Malvaceae) Pravin Patil, Shrikant Sutar1*, Joseph John K.2, Surendra Malik, Satyawada Rao3, Shrirang Yadav1 and K.V. Bhat National Bureau of Plant Genetic Resources, Pusa Campus, New Delhi - 110 012, India. 1Angiosperm Taxonomy Laboratory, Department of Botany, Shivaji University, Kolhapur - 416 004, Maharash- tra, India. 2National Bureau of Plant Genetic Resources, Regional Station, Vellanikkara, Kerala Agriculture University P.O., Thrissur - 680 656, Kerala, India. 3Department of Biotechnology and bioinformatics, North Eastern Hill University, Permanent Campus Mawkynroh, Shillong - 793 022, India. *E-mail: [email protected] Abstract The diverse agro-climatic conditions encompassing variable farming systems, rainfall and soil regimes have promoted diversification of the crops in the Southeast Asia. India is a rich treasure house of various species of Abelmoschus and all 11 species are found either in the wild, semi-wild or under cultivation. The genus Abelmoschus has a tortured systematic history. The ambiguity in nomenclature and lack of strong morphological characters for the delimitation of species has made the circumscription the genus rather controversial and created problems in breeder’s selection efforts. In the present review updated information on its taxonomy, genetic diversity, cytology and interspecific hybridization is discussed. Keywords: Abelmoschus, distribution, Malvaceae, okra, review, systematics Introduction The word Abelmoschus probably originated from origin, distribution, taxonomy, cytology, genetic Arabian abul-l-mosk meaning “father of musk, diversity, interspecific hybridization and economic source of musk” referring to the seeds of the genus importance of genus Abelmoschus to confer the (Don, 1831; Jaeger, 1950; Nayar, 1985; Quattrocchi, needs of those engaged in research on breeding 2000). Genus Abelmoschus is important and well and crop improvement. known because of its cultivated species, viz., A. esculentus and A. caillei. They are grown in many Taxonomical scenario of Abelmoschus parts of the world, especially in tropical and Hibiscus-Abelmoschus complex subtropical countries (Borssum Waalkes, 1966; Velayudhan et al., 1996). Abelmoschus was originally included in the genus Hibiscus under Monadelphia - Polyandria by The genus Abelmoschus has a tortured systematic Linnaeus (1753). On the basis of capsule features, history and it still pose problems in the delimitation Medikus (1787) proposed to raise this section to of species and other infraspecific categories. It is the rank of a distinct genus. This was accepted by mainly due to the polyploid nature of genome, Gaertner (1791), Moench (1794) and Schumann morphological plasticity, intermediate breeding (1890). On the contrary, authors like Masters system, long history of cultivation and dispersion (1874), Prain (1903) and Dunn (1915) did not accept (IBPGR, 1991). Remarkably little attention has been the Medikus view and followed the view of de paid to the wide spread taxa of Abelmoschus in India Candolle (1824). Hence, they treated Abelmoschus (Borssum Waalkes, 1966; Paul & Nair, 1988; IBPGR, as section of Hibiscus. Later, Hochreutiner (1924) 1991; Sivarajan & Pradeep, 1996; Velayudhan et al., distinguished the genus Abelmoschus from Hibiscus 1996; Bhat, 1996). mainly based on its peculiar spathaceous calyx The main objective of this review is to provide splitting to one-side. This was subsequently an overview of the existing information about followed in the taxonomic and contemporary P. Patil et al. 15 literature (Borssum Waalkes, 1966; Terrell & species (Hamon & Charrier, 1983). Sivarajan Winters, 1974; Paul & Nair, 1988; IBPGR, 1991; and Pradeep (1996) have provided detailed Vredebregt, 1991; Sivarajan & Pradeep, 1996; Bhat, morphological features and ecological distribution 1996; Bisht & Bhat, 2006; John et al., 2012; Sutar et for wild species. However, they did not consider al. 2013). infraspecific classification of A. manihot produced by Borssum Waalkes (1966). Paul and Nayar Species circumscription (1988) and Paul (1993) since then used characters The number of species included in the genus for separation, but these seem to be too feeble Abelmoschus by various authors (Hochreutiner, and highly inconsistent. Therefore, Sivarajan 1924; Borsssum Waalkes 1966; Sivarajan & and Pradeep (1996) treated A. manihot as a single Pradeep, 1996; Sutar et al., 2013) are provided in variable species and not attempted for recognising Table 1. Nevertheless, authentic number of species any infraspecific categories. Bates (1968) also included in the genus Abelmoschus is still uncertain, suggested that all subspecies and varieties of A. as they have not been studied in detail with respect manihot should be merged under the species. Later, to their taxonomic delimitation. Over fifty species Hamon et al. (1987) and Vredebregt (1991) pointed were described from the world (Charrier, 1984) that out that A. manihot subsp. manihot, A. manihot comprised many synonyms and misidentifications subsp. tetraphyllus var. tetraphyllus and A. manihot (Vredebregt, 1991). subsp. tetraphyllus var. pungens (Roxb.) Hochr. complex lack discrete species boundaries among Hochreutiner (1924) described 14 species, of them, which further contradicts Hochreutiner A. moschatus A. manihot which Medik. and (L.) (1900), Borssum Waalkes (1966), Paul and Nayar Medik. comprises several varieties. However, (1988) and Paul (1993). Intraspecific taxonomy of A. esculentus , only six species viz., (L.) Moench A. moschatus is also a matter of debate as many A. moschatus , A. ficulneus Medik. (Linn.) Wight subspecies and varieties have been recognized by , A. manihot , A. crinitus & Arn. (L.) Medik. Wall. Masters (1874), Hochreutiner (1900) and Borssum A. angulosus and Wall. ex Wight & Arn. were Waalkes (1966). maintained by Borssum Waalkes (1966). Further, A. manihot has been divided into two subspecies, Initially, A. caillei has been considered as a species namely subsp. manihot and subsp. tetraphyllus resembling to A. esculentus (Chevalier, 1940) and (Roxb. ex Hornem.) Borss. whereas, A. moschatus therefore, with restrictive approach Borssum with three subspecies, namely subsp. moschatus, Waalkes (1966) ignored this species in his revision subsp. tuberosus (Span.) Borss. and subsp. biakensis of Malesian Malvaceae. Later, A. caillei was (Hochr.) Borss. Later, this classification was elevated to a distinct species by Stevels (1988) adopted by International Okra workshop held at and this was further supported by Martin and NBPGR, Delhi (IBPGR, 1991) with some minor Rhodes (1983) and Sunday et al. (2008). However, changes. In this modified treatment A. manihot A. esculentus (Asian genotype) and A. caillei (West subsp. tetraphyllus was raised to the specific rank African genotype) generally pose problems as to and two more species viz., A. tuberculatus Pal & their determination. Singh and A. caillei (A. Chev.) Stevels were included. The recognition of A. tuberculatus as the closest Three varieties of A. angulosus Wall. ex Wight & progenitor of A. esculentus has been widely Arn., viz., var. angulosus, var. grandiflorusThwaites accepted, but Bates (1968) in his description and var. purpureus Thwaites are reported from of Abelmoschus, suggested that A. tuberculatus India (Sivarajan et al., 1994; Sivarajan & Pradeep, should be included under A. esculentus. However, 1996). Recently, John et al. (2012) described a new his argument for this is questionable since A. species, A. enbeepeegearense from the Western tuberculatus has already been recognized as a Ghats and Sutar et al. (2013) described A. palianus distinct species by Pal et al. (1952). The sequence from Chhattisgarh, India. Thus presently there are information based on internal transcribed spacer 11 species, 3 subspecies and 4 varieties in India as (ITS) gene also confirms the species status of A. well as for the world (Sutar et al., 2013). tuberculatus (authors unpublished data). Sterility However, the taxonomic treatment for some of direct and reciprocal F1 hybrids of A. esculentus species of Abelmoschus is not consistent. A. manihot and A. tuberculatus give additional evidence to their and A. moschatus are the most polymorphic distinctiveness as two different biological species. Table 1. Systematic treatment of the genus Abelmoschus by various authors. Hochreutiner (1924) Borssum Waalkes (1966) Paul & Nayar (1988) IBPGR (1991) Sivarajan & Pradeep (1996) Sutar et al. (2013) A. crinitus A. crinitus A. crinitus A. crinitus A. crinitus A. crinitus 16 A. ficulneus A. ficulneus A. ficulneus A. ficulneus A. ficulneus A. ficulneus R eview ofthegenus A. angulosus A. angulosus A. angulosus A. angulosus A. angulosus A. angulosus var. grandiflorus var. grandiflorus var. angulosus var. angulosus var. purpureus var. purpureus A. haenkeanus Abelmoschus A. moschatus A. moschatus A. moschatus A. moschatus A. moschatus A. moschatus var. genuinus subsp. moschatus subsp. moschatus var. moschatus var. moschatus var. betulifolius var. betulifolius var. betulifolius var. multiformis var. rugosus A. todayensis subsp. tuberosus subsp. tuberosus A. rhodopetalus A. brevicapsulatus A. sharpei A. biankensis subsp. biakensis subsp. biakensis A. manihot A. manihot A. manihot A. manihot A. manihot A. manihot var. genuinus subsp. manihot subsp. manihot var. timorensis subsp. tetraphyllus subsp. tetraphyllus var. tetraphyllus var. tetraphyllus var. tetraphyllus A. tetraphyllus
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