Chapter 9. Solanum Sect. Lycopersicon
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Chittaranjan Kole Editor Wild Crop Relatives: Genomic and Breeding Resources Vegetables [email protected] Editor Prof. Chittaranjan Kole Director of Research Institute of Nutraceutical Research Clemson University 109 Jordan Hall Clemson, SC 29634 [email protected] ISBN 978-3-642-20449-4 e-ISBN 978-3-642-20450-0 DOI 10.1007/978-3-642-20450-0 Springer Heidelberg Dordrecht London New York Library of Congress Control Number: 2011922649 # Springer-Verlag Berlin Heidelberg 2011 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer. Violations are liable to prosecution under the German Copyright Law. The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover design: deblik, Berlin Printed on acid-free paper Springer is part of Springer Science+Business Media (www.springer.com) [email protected] Chapter 9 Solanum sect. Lycopersicon Silvana Grandillo, Roger Chetelat, Sandra Knapp, David Spooner, Iris Peralta, Maria Cammareri, Olga Perez, Pasquale Termolino, Pasquale Tripodi, Maria Luisa Chiusano, Maria Raffaella Ercolano, Luigi Frusciante, Luigi Monti, and Domenico Pignone 9.1 Introduction such as their bright yellow flowers and pinnatifid, non-prickly leaves. ’The plant group Solanum sect. Lycopersicon con- Tomatoes belong to the large and diverse family sists of 13 closely related species or subspecies: the Solanaceae, which includes more than 3,000 species, cultivated tomato, Solanum lycopersicum (formerly occupying a wide variety of habitats (Knapp 2002). Lycopersicon esculentum), which includes the domes- Recent taxonomic revision of the Solanaceae has rein- ticated tomato and wild or weedy forms of the tegrated Lycopersicon into the genus Solanum with cherry tomato (S. lycopersicum ‘cerasiforme’) (Peralta a revised new nomenclature (Peralta and Spooner et al. 2008), and the wild species Solanum arcanum, 2001; Spooner et al. 2005; Peralta et al. 2008). The S. cheesmaniae, S. chilense, S. chmielewskii, S. corne- majority of taxonomists as well as most plant breeders liomulleri, S. galapagense, S. habrochaites, S. huay- and other users have accepted the reintegration of lasense, S. neorickii, S. pennellii, S. peruvianum, tomatoes to Solanum (e.g., Caicedo and Schaal 2004; S. pimpinellifolium) (Tables 9.1 and 9.2; Peralta et al. Fridman et al. 2004; Schauer et al. 2005; Mueller et al. 2005; Spooner et al. 2005). Four species have been 2009; see also http://tgrc.ucdavis.edu/key.html). segregated from the green-fruited species S. peruvia- Morphological characters, phylogenetic relation- num sensu lato (s.l.); two of them S. arcanum and ships, and geographical distribution have demon- S. huaylasense have been described as new species strated that tomatoes (Solanum sect. Lycopersicon (Peralta et al. 2005) from Peru, while the other two (Mill.) Wettst.) and their immediate outgroups in Sola- S. peruvianum and S. corneliomulleri had already been num sect. Lycopersicoides (A. Child) Peralta and sect. named by Linnaeus (1753) and MacBride (1962), Juglandifolia (Rydb.) A. Child form a sister clade to respectively. In addition, S. galapagense, another yel- potatoes (sect. Petota Dumort.), with Solanum sect. low-to orange-fruited species, was segregated from Etuberosum (Bukasov & Kameraz) A. Child being S. cheesmaniae; both species are endemic to the Gala´- sister to potatoes + tomatoes (Spooner et al. 1993). pagos Islands (Darwin et al. 2003; Knapp and Darwin Analyses of multiple datasets from a variety of genes 2007). All members of sect. Lycopersicon are closely unambiguously establish tomatoes to be deeply related diploid species (2n ¼ 24) (Peralta and Spooner nested in Solanum (Bohs and Olmstead 1997, 1999; 2001; Nesbitt and Tanksley 2002) and are character- Olmstead and Palmer 1997; Olmstead et al. 1999; ized by a high degree of genomic synteny (Chetelat Bohs 2005). However, tomatoes and their close rela- and Ji 2007; Stack et al. 2009) and are to some degree tives can be easily distinguished from any other group intercrossable (Taylor 1986). The group Solanum of Solanum species on the basis of shared features sect. Juglandifolia contains the two woody tomato- like nightshades S. ochranthum and S. juglandifolium. These two species are partially sympatric and they are S. Grandillo (*) morphologically similar, both being woody perennials CNR-IGV, Institute of Plant Genetics, Division of Portici, with rampant, liana-like stems up to 30 m in length National Research Council, Via Universita` 133, 80055 Portici, Naples, Italy (Correll 1962; Rick 1988). Based on evidence from e-mail: [email protected] molecular sequence data sect. Juglandifolia is the C. Kole (ed.), Wild Crop Relatives: Genomic and Breeding Resources, Vegetables, 129 DOI 10.1007/978-3-642-20450-0_9, # Springer-Verlag Berlin Heidelberg 2011 [email protected] Table 9.1 Principal ecological, botanical, and reproductive features of the wild tomatoes (Solanum sect. Lycopersicon) and related Solanum species 130 Species Geographic distribution Habitat Mating systema Crossability Distinguishing morphological featuresc to tomatob S. lycopersicum Adventive worldwide in the tropics and Usually mesic sites, often SC-autogamous BC Plants semi-erect to sprawling; fruits red, “cerasiforme” subtropics (near sea level – 2,400 m); feral or weedy 1.5–2.5 cm perhaps native in Andean region S. cheesmaniae Endemic to Gala´pagos Islands Arid, rocky slopes, prefers SC-autogamous BC Plants semi-erect to sprawling, flowers (sea level – 1,500 m) shaded, cooler sites very small, pale; fruit purple, greenish- yellow, or orange, 0.5–1.5 cm S. galapagense Endemic to Gala´pagos Islands Arid, rocky outcrops and SC-autogamous BC Plants erect; leaves highly subdivided; (sea level – 650 m) slopes, sometimes near internodes short; flowers small, pale, shoreline fruit orange (0.5–1 cm) S. pimpinellifolium Lowland Ecuador and coastal Peru Arid, sandy places, often near SC-facultative BC Plants semi-erect to sprawling, flower (sea level – 500 m) sources of water or on the small-large; fruit red (0.5–1 cm) edges of farm fields S. chmielewskii Inter-Andean valleys of central and Rather moist, well-drained, SC-facultative UI Plant sprawling or trailing; flowers small, [email protected] southern Peru (1,600–3,100 m) rocky slopes pale; fruit green (1–1.5 cm) S. neorickii Inter-Andean valleys from Cusco to Rather moist, well-drained, SC-autogamous UI Plants sprawling or trailing; flowers tiny, central Ecuador (1,500–2,500 m) rocky slopes pale; fruit green; seeds tiny S. arcanum Northern Peru, coastal and inter-Andean Varied, but generally dry, Mostly SI, rarely UI, EL Plants erect to prostrate, reduced leaflet valleys, middle watershed of Maran˜on rocky slopes SC-facultative no.; flowers mostly straight anther (500–3,000 m) tubes and undivided inflorescences; fruit whitish-green with dark stripe S. chilense Southern Peru, northern Chile Very arid and sometimes saline, SI UI, EL Plants erect; leaves finely pubescent; (50–3,500 m) rocky slopes or washes anthers straight; inflorescences compound; peduncles long; fruit purplish-green S. peruvianum Mostly coastal central/southern Peru and Arid, sandy, or rocky dry Mosty SI, rarely UI, EL Plants procumbent; anthers bent; northern Chile (sea level – 2,500 m) washes, sometimes near SC-facultative inflorescence simple; fruit purplish- agricultural fields green S. corneliomulleri Western Andes of central/southern Peru Rocky or sandy slopes and SI UI, EL Erect to decumbent; leaves glandular (1,000–3,000 m) dry washes pubescent; fruit purplish-green S. huaylasense Limited to Callejon de Huaylas, and Rı´o Rocky slopes and waste places SI UI, EL Spreading, anthers straight, inflorescence Fortaleza, Peru (1,000–2,900 m) compound; fruit purplish-green S. habrochaites Northwestern and western central Peru, Varied, but generally mesic Mostly SI, some UI Sprawling shrub or vine; densely western and southern Ecuador slopes or stream banks SC-facultative pubescent; flowers large; anthers (40–3,300 m) straight; fruit green with dark stripe, hairy S. Grandillo et al. S. pennellii Coastal valleys of central to southern Very arid, sandy or rocky Mostly SI, some UI Spreading shrub; 2 leaves per Peru (near sea level to 1,920 m) slopes, or dry washes SC-facultative sympodiumd; leaflets broad, round; foliage sticky; anthers poricidal; pedicel usually articulated at base 9 S. juglandifolium Temperate rainforests of Columbia and Mesic slopes and SI UI (no Woody vine or rampant shrub; 8–10 leaves Solanum Ecuador (1,200–3,100 m) stream banks hybrids) per sympodiumd; leaves rough, rugose; anthers orange-yellow, poricidal; flowers scented; fruit green (to 2 cm); seeds winged sect. S. lycopersicoides Restricted to narrow range in southern Arid rocky slopes, usually SI UI, EL, HS Woody, erect to sprawling shrub; anthers Lycopersicon and S. ochranthum Peru and northern Chile south-facing white, poricidal; style hooked; flowers