Solanaceae Source

A global taxonomic resource for the nightshade family

Solanum pimpinellifolium

Citation author: 
L.
Citation: 
Cent. 8. 1755.
Type: 
Cultivated in Uppsala, Anon. (lectotype, LINN 248.15 [BH neg. 6802], designated by Knapp & Jarvis 1990).
Written by: 
Peralta, I.E., S. Knapp & D.M. Spooner
Habit: 
Annual, biennial, or sometimes perennial herbs, erect initially, later procumbent viny herbs with branches extending to 3 m from center. Stems 8-11 mm in diameter at base, green, usually sparsely (rarely densely velvety) pubescent; trichomes of several types, the most common uniseriate, 1-2-celled, white and thin, ca. 0.5 mm long from a unicellular base, also with sparsely scattered glandular trichomes with unicellular or multicellular heads, occasionally (in northern Peruvian populations) with larger, robust patent uniseriate trichomes to 1 mm long from a multicellular base, these eglandular or with a minute unicellular head, and minute glandular trichomes with unicellular heads.
Sympodial structure: 
Sympodial units 3-foliate; internodes 2-8 cm long.
Leaves: 
Leaves interrupted imparipinnate, 4-12 cm long, 1.5-8 cm wide, green, sparsely to densely pubescent with a mixture of uniseriate glandular and eglandular trichomes, pubescence velvety, more abundant along the veins and abaxially, in northern Peruvian populations with robust patent uniseriate trichomes like those of the stems along the veins abaxially; primary leaflets 2-4 pairs, slightly decreasing in size towards the leaf base, elliptic to broadly elliptic to ovate to broadly ovate, the base cordate or occasionally truncate, oblique and extended basiscopically, the margins entire or round-crenate or slightly dentate in the proximal third (more crenate in putatively hybrid populations in northern part of the species range), the apex somewhat rounded or acute to acuminate; terminal leaflet 2.5-5 cm long, 1-3.5 cm wide, the petiolule 0.5-2 cm long; lateral leaflets (largest) 1.5-3.5 cm long, 1-2 cm wide, the petiolule 0.5-0.7 cm long, with the base usually decurrent basiscopically; secondary leaflets absent; tertiary leaflets absent; interjected leaflets usually few, 1-4 pairs, 0.5-1.2 cm long, 0.2-0.7 cm wide, sessile or with the petiolule to 0.2 cm long, the length between the lateral and interjected leaflets extremely variable, generally 0.5-2 cm long; petiole (0.8) 1.5-5 cm long; pseudostipules absent.
Inflorescences: 
Inflorescences 4-25 cm long, usually simple, occasionally once-branched, with 7-30 flowers, ebracteate, peduncle 2-3 (-5) cm long, pubescent like the stems but with more glandular trichomes with multicellular heads. Pedicels 1-1.3 cm long, the articulation in the distal half. Buds 0.5-1.2 cm long, elongate conical, with the corolla more than 2/3 to 3/4 of the way exserted from the calyx just before anthesis.
Flowers: 
Flowers with the calyx tube 0.05-0.1 mm long, the lobes 0.25-0.5 cm long, 0.1-0.25 cm wide, lanceolate, densely pubescent with uniseriate trichomes like the rest of the inflorescence; corolla 1.2-3 cm in diameter, stellate, pale yellow to bright yellow, the tube 0.1-0.25 cm long, the free portion of the lobes 1-1.5 cm long, 0.2-0.4 cm wide, the tips margins densely and minutely white pubescent abaxially, the lobes strongly reflexed at anthesis; staminal column 0.7-1 cm long, straight, the filaments 0.5-1 mm long, the anthers 0.5-0.7 cm long, the sterile apical appendage 0.3-0.4 cm long; ovary conical to globose, glabrous; style 0.7-1 cm long, ca. 0.5 mm in diameter, densely white pubescent in the basal 1/3, straight, included to usually exserted 0.5-1 mm from the staminal column; stigma capitate, green.
Fruits: 
Fruit to ca. 1 cm in diameter, globose, 2-locular, bright red when ripe, sparsely to densely glandular pubescent with short uniseriate trichomes ca. 0.5 mm long, with multicellular and unicellular heads during maturation, glabrous when mature; fruiting pedicels 1.5-2 cm long, straight or bent in towards the rachis at the articulation point; calyx lobes in fruit 1-1.2 cm long, 0.25-0.4 cm wide, strongly reflexed and parallel with the fruiting pedicel.
Seeds: 
Seeds 2-3 mm long, 1-2.3 mm wide, 0.5-0.8 mm thick, obovate, narrowly winged at the apex and acute at the base, pale brown, pubescent with hair-like outgrowths of the lateral testa cell walls, theseadpressed and giving a silky appearance to the surface or more often shaggy, narrowly winged (0.3-0.4 mm) at the apex and acute at the base.
Distribution: 

Apparently native to coastal areas from northern Peru to central Chile, 0-500 m, although populations are found to central coastal Ecuador (but see below). Solanum pimpinellifolium grows in wet places and on the edges of cultivated fields throughout its native range and has apparently escaped from cultivation in the Galápagos (see Darwin et al. 2003).

Phenology: 
Solanum pimpinellifolium flowers and fruits throughout the year in response to moisture.
Phylogeny: 

Solanum pimpinellifolium is a member of the Potato clade (sensu Weese & Bohs, 2007); within the tomatoes and wild relatives it is a member of the “Lycopersicon group” and is a member of section Lycopersicon.

Commentary: 

Solanum pimpinellifolium is very closely related to S. lycopersicum and hybridizes with it freely. It differs from cultivated plants of S. lycopersicum in a variety of genetic characteristics, and we feel maintenance of S. pimpinellifolium at the specific rank will better serve both nomenclatural stability and the plant breeding community, who have long used this species in studies of tomato genetics (Frary et al. 2000; van der Knaap & Tanksley 2001). Since the two taxa are so similar, and identification of individual specimens can be problematic owing to putative introgression we use a suite of characters to identify S. pimpinellifolium. Our rule of thumb for identifying individual specimens (especially those that we suspect are of hybrid origin) was to assign to S. pimpinellifolium any specimen that possessed a majority of the characters below.

1. Corollas stellate, deeply (>3/4) divided, the lobes narrowly lanceolate

2. The sterile appendage of the staminal tube longer than the pollen bearing portion, making buds long and thin in comparison to those of S. lycopersicon.

3. Elongate inflorescence (greater than 3 cm) and many (>20) flowers

4. Very small fruits (<1 cm in diameter)

5. Few interjected leaflets

6. Leaflets entire or only slightly crenate in proximal 1/3

7. Terminal leaflet base cordate

In addition, identification can be facilitated by observation of the terminal leaflets, broadly elliptical in S. pimpinellifolium rather than narrowly ovate (S. lycopersicum) and the overall velvety pubescence of very short trichomes in S. pimpinellifolium, without long trichomes (except in putatively introgressed populations). Populations in the northern part of the range of S. pimpinellifolium (coastal Ecuador) and occasionally in higher elevation valleys in northern Peru apparently are the result of introgression with S. lycopersicum and consequently have deeper crenate leaflets and often have a few stout, patent trichomes on the stems and inflorescences. There is great variation and difficulty in identification owing to introgression wherever the two species are in contact.

In the Galápagos Islands, S. pimpinellifolium has been introduced or has only arrived recently, and forms hybrids with the endemic Galápagos tomato S. cheesmaniae where the two species occur in sympatry (Darwin et al. 2003; S.C. Darwin, unpubl.). Some plants identified by Nuez et al. (2004) as “Lycopersicon esculentum var. cerasiforme” we assign to S. pimpinellifolium.

Autogamous populations of S. pimpinellifolium have smaller flowers than outcrossing populations (Georgiady & Lord 2002), caused by heterochronic growth differences in the flower. A single QTL is responsible for four of six flower traits associated with autogamy (Georgiady et al. 2002), but much variation in all these traits exists in both sets of populations. Autogamy is associated with the margins of the range of S. pimpinellifolium (Rick et al. 1977), and studies of variation in nuclear gene sequences (Vac – fruit vacuolar invertase) in northern Peruvian populations (Caicedo & Schaal 2004a) show levels of outcrossing decreasing towards the south (towards the center of the species range). Caicedo & Schaal (2004a, 2004b) showed that genetic variation is high, and that there is no congruence between geographical distribution and genealogy in these populations. They suggest their data shows a northern Peruvian origin for S. pimpinellifolium, but they did not sample across the entire species range (into Chile). The more northerly populations in both their and Rick et al.’s (1977) study may have introgressed with S. lycopersicum.

Solanum pimpinellifolium has been of great importance to tomato breeding for the introduction of important agronomic traits such as disease resistance, and for the investigation of the genetic control of traits such as fruit shape and size (e.g. van der Knaap et al. 2004). Crosses between S. lycopersicum and the TGRC accession S. pimpinellifolium accession LA1589 (an autogamous population from the lower Virú Valley in the Department of La Libertad, Peru) have been the mainstay of studies in fruit development in the tomato (see Morphology above) and LA1589 has recently been used in metabolic profiling (Schauer et al. 2005). Solanum pimpinellifolium had the lowest fruit protein content of any of the species tested, but its leaf metabolite content is very close to that of S. lycopersicum, further underlining their close relationship (Schauer et al. 2005).

One TGRC accession of S. pimpinellifolium (LA1589) has been used to produce backcross recombinant inbred lines with the cultivated tomato. The 126 inbred backcross lines contain introgressed segments of S. pimpinellifolium that covers most of the genome. These lines have been important to produce genetic maps and analysis of quantitative traits (QTL) that are available from the Solanaceae Genome Network (www.sgn.cornell.edu).

The type specimen of S. pimpinellifolium in the Linnaean herbarium in London (see Knapp & Jarvis 1990) is a close match for collections from around Lima (e.g., Mexia 8328) in central Peru. These populations are comprised of glabrous plants with entire leaflets with cordate bases, elongate inflorescences, and very narrow petals. Seed of the type specimen of S. peruvianum was sent to Linnaeus by Bernard de Jussieu (Knapp & Jarvis 1990), whose brother Joseph was in the Lima area (see discussion under S. peruvianum). Linnaeus indicated no provenance for S. pimpinellifolium, but perhaps seeds also came to him via Jussieu along with S. peruvianum. Miller (1768) mentioned that he received seeds of both species from B. de Jussieu.

Luckwill (1943a) misunderstood Lycopersicon pissisi and equated it with S. peruvianum, possibly due to its habitat in coastal deserts. The protologue clearly states that the species has red fruit and no inflorescence bracts. The type specimen also has 3 leaves per sympodial unit, making it unmistakably a member of the “Lycopersicon” species group.

Luckwill (1943a: 24) described his L. esculentum subsp. intermedium citing a “well-defined and pure breeding form … which has been under cultivation for scientific purposes at the John Innes Horticultural Institute, Merton, for a number of years and has been widely employed for research work under the name of L. racemigerum Lange.” From his description of this taxon and from several UK cultivated specimens collected during that time (but none from John Innes found), we believe this is a hybrid between S. lycopersicum and S. pimpinellifolium, as is the type of L. racemigerum itself. These hybrids apparently form easily in cultivation, the reason that Brezhnev (1958) recognized S. pimpinellifolium only at the subspecific rank (see synonymy above).

Lycopersicon pimpinellifolium (L.) Mill. subsp. intermedium Luckwill is a misprint in Index Kewensis for L. esculentum Mill. subsp. intermedium Luckwill, but the subspecific epithet was indeed transferred to L. pimpinellifolium later (see synonymy above) by Prokhanov (1950) in the treatment of Lycopersicon for the Flora of the USSR (see also discussion under S. lycopersicum).

References: 

Miller, P. 1768. The gardener’s dictionary, Abridged 8th ed
London.

Luckwill, L.C. 1943. The genus Lycopersicon: an historical, biological, and taxonomical survey of the wild and cultivated tomatoes.
Aberdeen Univ. Stud. 120: 1-44.

Brezhnev, D.D. 1958. Tomat in Ovoshchnye paslenovye (Tomat, Baklazhan, Chernyi Paslen, Dynnaia Grusha, Perets, Fizalis, Mandragora).
In Kul’turnaia Flora SSSR 20, ed. P. M. Zhukovskii, 7-278. Moscow-Leningrad: State Printing House.

Rick, C.M., J.F. Fobes, & M. Holle. 1977. Genetic variation in Lycopersicon pimpinellifolium: evidence of evolutionary change in mating systems.
Pl. Syst. Evol. 127: 139-170.

Knapp, S. & C.E. Jarvis 1990. The typification of the names of New World Solanum species described by Linnaeus.
J. Linn. Soc., Bot. 104: 325-367.

Frary, A., T.C. Nesbitt, S. Grandillo, E. van der Knaap, B. Cong, J. Liu, J. Meller, R. Elber, K.B. Alpert, & S.D. Tanksley 2000. fw2.2: a quantitative trait locus key to the evolution of tomato fruit size.
Science 289: 85-88.

Van der Knaap, E., & S.D. Tanksley 2001. Identification and characterization of a novel locus controlling early fruit development in tomato.
Theor. Appl. Genet. 103: 353-358.

Georgiady, M.S. & E.M. Lord 2002. Evolution of the inbred flower form in the currant tomato, Lycopersicon pimpinellifolium.
Int. J. Pl. Sci. 163: 531-541.

Darwin, S.C., S. Knapp, & I.E. Peralta 2003. Taxonomy of tomatoes in the Galápagos Islands: native and introduced species of Solanum section Lycopersicon (Solanaceae).
Syst. Biodiv. 1(1): 29-53.

Van der Knaap, E., A. Sanyal, S.A. Jackson, & S.D. Tanksley 2004. High-resolution fine mapping and fluorescence in situ hybridization analysis of sun, a locus controlling tomato fruit shape, reveals a region of the tomato genome prone to DNA rearrangements.
Genetics 168: 2127-2140.

Nuez, F., J. Prohens, & J.M. Blanca 2004. Relationships, origin, and diversity of Galápagos tomatoes: implications for the conservation of natural populations.
Amer. J. Bot. 91: 86-99.

Caicedo, A.L. & B.A. Schaal 2004. Heterogenous evolutionary processes affect R gene diversity in natural populations of Solanum piminellifolium.
Proc. Nat. Acad. Sci., USA 101(50): 17444-17449.

Caicedo, A.L. & B.A. Schaal 2004. Population structure and phylogeography of Solanum pimpinellifolium inferred from a nuclear gene.
Mol. Ecol. 13: 1871-1882.

Schauer, N., D. Zamir, & A.R. Fernie 2005. Metabolic profiling of leaves and fruits of wild species tomato: a survey of the Solanum lycopersicum complex.
J. Exp. Bot. 56: 297-307.

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