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Solanum Insanum L. (Subgenus Leptostemonum Bitter, Solanaceae), the Neglected Wild Progenitor of Eggplant (S

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Solanum Insanum L. (Subgenus Leptostemonum Bitter, Solanaceae), the Neglected Wild Progenitor of Eggplant (S Genet Resour Crop Evol DOI 10.1007/s10722-016-0467-z RESEARCH ARTICLE Solanum insanum L. (subgenus Leptostemonum Bitter, Solanaceae), the neglected wild progenitor of eggplant (S. melongena L.): a review of taxonomy, characteristics and uses aimed at its enhancement for improved eggplant breeding R. H. G. Ranil . J. Prohens . X. Aubriot . H. M. L. Niran . M. Plazas . R. M. Fonseka . S. Vilanova . H. H. Fonseka . P. Gramazio . S. Knapp Received: 27 July 2016 / Accepted: 3 November 2016 Ó The Author(s) 2016. This article is published with open access at Springerlink.com Abstract Solanum insanum L. (Solanaceaeae) is the ethnolinguistic groups. Limited studies carried out on wild progenitor of common eggplant (S. melongena its chemical composition and pharmacognostical L.) and is naturally distributed in south and southeast properties coupled with ethnobotanic studies reveal Asia, Madagascar and Mauritius. Despite interest in its that S. insanum has potential for eggplant improve- potential for eggplant breeding it has not been studied ment with respect to phytochemical properties of extensively. Solanum insanum has been subjected to interest for human health. Further study of morpho- various taxonomic treatments, with several discus- logical and genetic diversity in S. insanum is essential sions and arguments on its delimitation as a species, to promote its utilization in eggplant breeding, in light due to its close similarities with the cultivated S. of its potential to hybridize with cultivated eggplant melongena and the wild species S. incanum L. from successfully. Here we compile and review the taxon- the Middle East and northern Africa. Solanum omy, phytogeography, ecology and habitat character- insanum has long been used in a variety of medicinal istics, cytology, ethnobotany, breeding and and culinary preparations across many different Asian phytochemical analysis of S. insanum. A comprehen- sive botanical description and illustrations for S. insanum are provided. A detailed comparison of S. R. H. G. Ranil Á R. M. Fonseka insanum and its closely allied species, S. melongena Department of Crop Science, Faculty of Agriculture, and S. incanum is also made. The potential of S. University of Peradeniya, Peradeniya, Sri Lanka insanum for crop improvement and future research J. Prohens Á M. Plazas Á S. Vilanova Á P. Gramazio priorities are discussed. Instituto de Conservacio´n y Mejora de la Agrodiversidad ` ` Valenciana, Universitat Politecnica de Valencia, Keywords Solanum insanum Eggplant Valencia, Spain Á Á Taxonomy Á Phytogeography Á Ethnobotany Á X. Aubriot Á S. Knapp (&) Breeding Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK e-mail: [email protected] H. M. L. Niran Introduction Agriculture Research Station, Girandurukotte, Sri Lanka Solanum is the largest and most economically impor- H. H. Fonseka Horticultural Crop Research and Development Institute, tant genus in the family Solanaceae and comprises Gannoruwa, Peradeniya, Sri Lanka around 1400 species worldwide (Bohs 2005). Solanum 123 Genet Resour Crop Evol subgenus Leptostemonum (also referred to as the Vorontsova and Knapp 2016). Samuels (2016) has Leptostemonum clade, sensu Bohs 2005), which is suggested that due to this spelling similarity and to the commonly known as the ‘spiny solanums’, contains previously confused taxonomic usage, the name S. more than 550 currently recognized species, many of insanum should not be used; he, however, did not which are native to the New World (Levin et al. 2006), suggest suppression under the rules of the Interna- where species diversity in the clade has long been tional Code of Nomenclature for algae, fungi and thought to be highest (e.g., Nee 1999; Bohs 2005). plants (McNeill et al. 2012). Others, however, con- Recent discoveries and new taxonomic circumscrip- sider the name not confusable (Knapp et al. 2013; see tions have, however, revealed great spiny solanum below), and recent clarification of both species limits diversity in the Old World. The Old World, including (see below) and relationships means the name is now Africa, Asia and Australia, is home to more than 300 in current use (e.g., Cericola et al. 2013; Wu et al. species, most of which are members of a monophyletic 2013; Meyer et al. 2014; Aubriot et al. 2016). Until clade (Levin et al. 2006; Weese and Bohs 2007; Knapp et al. (2013) re-visited species limits and Vorontsova and Knapp 2016; but see Aubriot et al. identities in the eggplant clade, S. insanum was treated 2016 for exceptions). The spiny solanums from Africa in two of the morphotypes of S. melongena,‘‘S. have been extensively studied taxonomically (Bitter melongena group E’’ and ‘‘S. melongena group F’’ 1923; Jaeger 1985; Jaeger and Hepper 1986; Lester (e.g., Lester and Hasan 1991; Daunay and Hazra and Hasan 1990, 1991; Samuels 1996, 2010, 2012). For a review of the difficulties with the previous 2012a, 2013b; Levin et al. 2006; Vorontsova et al. informal group classification see Knapp et al. (2013) 2013), but the only comprehensive modern mono- and Syfert et al. (2016). graph is that of Vorontsova and Knapp (2016). Studies Many early phylogenetic analyses using molecular on the spiny solanums in Asia have been more limited data did not explicitly include S. insanum (Levin et al. (e.g., Deb 1989; Karihaloo and Rai 1995; Davidar 2006; Weese and Bohs 2007, 2010), but have included et al. 2015; Mutegi et al. 2015), or limited to broad material that we would now identify as that species. phylogenetic analyses with less discussion of species Meyer et al. (2012) examined samples of S. insanum boundaries (Aubriot et al. 2016). (as S. undatum Lam. or S. incanum) from India, Sri The identity of the wild progenitor of the eggplant Lanka, China, Indonesia and Thailand for their has been the subject of considerable taxonomic analysis of phylogenetic relationships among Asian confusion. Solanum insanum was described in 1767 eggplants, and Vorontsova et al. (2013) also included (Linneaus 1767). Since its first description, S. insanum samples from China, India and Indonesia for their has been treated in a wide variety of ways, from being phylogenetic analysis of African spiny solanums. ignored entirely, to being given infraspecific status, to More recent phylogenetic analyses using additional being recognized at the species level under a variety of molecular markers and a broad sampling of Asian and different names (Roxburgh 1832; Prain 1903; Duthie African Solanum species (Aubriot et al. 2016) confirm 1911; Gamble 1921; Bhaduri 1951; Deb 1979; Hepper previous hypotheses (Knapp et al. 2013; Meyer et al. and Jaeger 1985; Lester and Hasan 1990, 1991; 2015) that S. insanum is the wild progenitor of Karihaloo and Rai 1995; Weese and Bohs 2010; eggplant (S. melongena). These studies and unpub- Meyer et al. 2012; Knapp et al. 2013; Samuels lished work (X. Aubriot pers. comm.) do not support 2010, 2012a, 2013a; Vorontsova et al. 2013; Aubriot the proposed linear progression of relationships from et al. 2016; Vorontsova and Knapp 2016; see our Africa to Asia that has long been the paradigm for current view of synonymy in taxonomic treatment eggplant evolution (Lester and Hasan 1991; Weese below). Such differences in status and taxonomic and Bohs 2010; Samuels 2010, 2012b, 2013b). Sup- treatment were largely due to its morphological port for the relationships amongst species of the similarity to and close affinity with cultivated eggplant eggplant clade (sensu Vorontsova et al. 2013) is weak, (S. melongena) and other quite similar wild relatives but new data from whole plastid sequences of (Knapp et al. 2013), and to the unfortunate similarity additional species and involving denser sampling of the spelling of the specific epithet to that of S. indicates the African species (including S. incanum) incanum L. (Hepper and Jaeger 1985), another mem- are, as a group, sister to S. insanum ? S. melongena ber of the eggplant clade (Aubriot et al. 2016; (X. Aubriot unpublished). 123 Genet Resour Crop Evol Although data from molecular phylogeny together that such information will be a useful information with evidence from chemical composition (Meyer resource for taxonomists, plant breeders and conser- et al. 2015) support S. insanum as closely related but vation biologists who are working with S. insanum and distinct from S. melongena, future molecular phylo- will spark interest in this previously confused and genetic analysis including an ample representation of largely neglected species. S. insanum accessions is needed to provide in-depth information about relationships of populations across Asia, and on the geographical origins of S. melongena. Materials and methods Eggplant is thought to have several centres of domes- tication across Asia (India, China and Malesia), based Our circumscription of S. insanum is based on on genetic differences (Meyer et al. 2012; Cericola herbarium specimens from 283 indivual collection et al. 2013) and non-overlapping medicinal uses events (ca. 400 specimens) from 22 herbaria (A, BH, (Meyer et al. 2014). Despite this relationship and its BM, CAS, DS, E, F, G, G-DC, GH, HIFP, K, L, LE, high potential for eggplant breeding, S. insanum has MA, MO, NY, P, S, TAN, U, US; all herbarium remained largely underutilized, despite being the acronyms follow Index herbariorum, http:// progenitor and thus the only species included in the sweetgum.nybg.org/science/ih/); all data for these primary genepool of eggplant (Syfert et al. 2016). specimens is available on the website Solanaceae Crop wild relatives are likely to play a significant Source and on the Natural History Museum Data role in securing 21st century food security (Dempewolf Portal (http://dx.doi.org/10.5519/0001289). We et al. 2014). This is due to their potential uses in plant complemented information from dried specimens breeding to produce crop varieties which withstand with the evaluation of morphological traits of S. adverse impacts of climate change, increasing scarcity insanum plants growing in their native environment of nutrients, water and other inputs, and new pest and in Sri Lanka and in plants grown at the Universitat diseases (Hopkins and Maxted 2010).
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