Solanaceae Source

A global taxonomic resource for the nightshade family

Solanum stoloniferum

Citation author: 
D. F. L. Von Schlechtendal
Citation: 
Linnaea 8: 255. 1833.
Type: 
Mexico. Veracruz: Mount Orizaba, Lake Huatulaca, Sep, about 1828-1830. C. J. W. Schiede & F. Deppe 192 p.p. (lectotype, designated by Hawkes [1957: 851] a cultivated plant originally grown in Berlin Botanic Garden, and now at UPS!; isolectotypes: GH!, GOET!, HAL!, W! [F neg. 33133: F!, G!, GH!, LL!, MICH!, MO!, US!]).
Last edited by: 
Spooner, D.M.
Written by: 
Spooner, D.M.
Habit: 
Herbaceous tuber-bearing perennials 0.2-0.7 m tall. Stems 2-6 mm in diameter at base of plant.
Sympodial structure: 
Sympodial units typically 3-6-foliate.
Leaves: 
Pseudostipules 5-16 mm long, lunate. Leaves odd-pinnate, 7.5-22 cm long, 3.5-8 cm wide, strigose, puberulent, or pubescent adaxially and abaxially; petioles 1.5-4 cm long; lateral leaflet pairs 1-4, the size of the lateral leaflets usually diminishing abruptly towards the base of the leaf; most distal lateral leaflets 2.5-3.2 cm long, 1-2.2 cm wide, elliptic to ovate, apex acute to acuminate, base oblique, cuneate to cordate, petiolules 1.5-6 mm long or sometimes the most distal laterals highly decurrent; terminal leaflet 2.8-6 cm long, 2-3.5 cm wide, elliptic to obovate, apex acute to acuminate, base cuneate; interjected leaflets 0-15.
Inflorescences: 
Inflorescence a dichasially branched, ebracteate, monochasial or dichasial cyme, 2-3 forked, generally in the distal half of the plant, with 3-26 flowers, all flowers perfect, peduncle 3.4-10 cm long; pedicels 11-37 mm long, articulate between the proximal ¼ and the distal ¼.
Flowers: 
Flowers with the calyx up to 9-37 mm long, lobes acute to long-attenuate, acumens 1-3 mm long. Corollas 1.8-3.3 cm in diameter, pentagonal to rotate, acumens 1.5-4 mm, edges of corolla flat, not folded dorsally, typically light purple adaxially and darker purple abaxially, or white tinged with purple t the tips, less commonly white throughout. Anthers 3.5-6.5 mm long, connate, yellow, apically poricidally dehiscent and often maturing to a short introrse apical slit, filaments 1-4 mm long. Ovary with style 5-10 mm long, exceeding stamens by 0-4.5 mm, straight, with stigma globose.
Fruits: 
Fruits 0.9-1.7 cm in diameter, globose to lightly ovoid, almost pure white to deep green, sometimes light green with darker green stripes and white spots.
Seeds: 
Seeds from living specimens green-white throughout, ovoid, ca. 2 mm long, with a thick covering of “hair-like” lateral walls of the testal cells that make the seeds mucilaginous when wet. Removal of these hair-like lateral walls by enzyme digestion reveals a honeycomb pattern at their base.
Chromosome number: 

2n = ploidy missing =48 voucher missing = (Spooner & Hijmans 2001)

Distribution: 

Southwestern U.S.A. (Arizona, New Mexico, SW Texas); to Oaxaca, Mexico, (1040-) 1440-3400 (-3700) m; among boulders on steep hillsides, sandy alluvial stream bottoms, in gravel along trails or roadways, in thick leaf mulch under trees, at edges of cultivated or fallow fields, along fencerows and railways, in organic moist soil to dry sandy soils, in grasslands, juniper-pinion scrub, desert tropical deciduous forests, to fir, pine, juniper or oak forests.

Phenology: 
Flowering and fruiting June through September in the north (U.S.A) and July through November farther south in southern Mexico.
Phylogeny: 

Solanum stoloniferum belongs to the potato clade of Solanum (Bohs, in press). Spooner and Sytsma (1992) placed S. stoloniferum in the most derived clade of section Petota based on chloroplast DNA restriction site data. Spooner et al. (2004) erected the Longipedicellata group for S. hjertingii and S. stoloniferum. These two species are polysomic polyploids and 4x(2EBN). There are no clear group-specific morphological characters defining the Longipedicellata group (see discussion in Spooner et al. 2004). Hawkes (1990) placed six species in ser. Longipedicellata: S. fendleri, S. hjertingii, S. matehualae, S. papita, S. polytrichon, and S. stoloniferum. The specimens Hawkes labeled as these names are all tetraploid (2n = 4x = 48) and grow in the southeastern United States (Hawkes’s S. fendleri) and Mexico. Morphological data (Spooner et al. 2001) supported at best only three species in series Longipedicellata: 1) S. polytrichon, 2) S. hjertingii (including S. matehualae), and 3) S. stoloniferum (including S. fendleri and S. papita). Solanum polytrichon was supported as a species only by a canonical variate analysis, but not by a principal components analysis that intermixed S. polytrichon with many other species, and thus was the least supported species.

Commentary: 

Solanum stoloniferum is one of the most common, widespread, and polymorphic species of wild potatoes from North and Central America. It is very similar to some populations of S. demissum, S. hjertingii, and S. verrucosum. Solanum stoloniferum has pedicel articulation below the distal ¼ of the pedicel, flat corolla margins, and straight styles exserted only 0.4.5 mm beyond the tip of the anthers (Spooner et al. 2001). Solanum demissum has pedicel articulation in the upper ¼ of the pedicel, S. hjertingii has curved styles exserted 2-8 mm beyond the tip of the anthers, and S. verrucosum has corollas with the edges inrolled dorsally.

Van den Berg et al. (2002) showed S. polytrichon to be completely intermixed with S. stoloniferum (including S. fendleri and S. papita) in an analysis of AFLP data. A reexamination by Spooner et al. (2004) of living accessions previously identified as S. polytrichon at Sturgeon Bay Wisconsin in 2001 showed putative diagnostic characters of spreading pubescence, enlarged terminal leaflets, and white corollas to vary tremendously in this species. For example, Graham 250, 256, 315, 358; Hawkes 1468; Ochoa 14178, possessed a clear expression of dense spreading pubescence, enlarged terminal leaflets, and white corollas. Other accessions, however, are impossible to clearly and consistently distinguish as S. polytrichon and were supported as species in the canonical variate analysis only by widely overlapping character states. For example, Tarn et al. 235, Rodríguez 2558, 2571; Spooner et al. 4109, have only few spreading hairs, and Tarn et al. 212 and Spooner et al. 4118 have purple corollas. Accessions identified as S. papita (Correll 20075 and Tarn et al. 114) have white corollas and spreading hairs. There is such an intergradation of these “species-specific” character as to make S. polytrichon untenable to recognize.

There is a tendency for S. stoloniferum to be of shorter stature with fewer interjected leaves in the northern than in the southern part of its range, and these northern populations have been named S. fendleri. There are so many exceptions to these characters however (e.g., Blumer 1566, type of S. fendleri subsp. arizonicum, that is taller; or Blumer 1579 or Lemmon 2845 from Arizona with up to four sets of interjected leaflets) that height alone fails to distinguish these formerly recognized species.

Solanum stoloniferum can easily be confused with some landraces of S. tuberosum. Solanum tuberosum often has wider leaflets and longer peduncles than S. stoloniferum, and some landraces have pink corollas (never pink in S. stoloniferum). Solanum tuberosum is so variable, however, (including cultivars with narrower leaflets, shorter peduncles, and white or blue or purple corollas; e.g., Mexia 2658 at E, RSA) that the two taxa may be impossible to distinguish consistently and reliably on herbarium sheets.

Hawkes (1990) recognized S. ×vallis-mexici as a triploid nothospecies, formed by hybridization between S. stoloniferum and S. verrucosum. Spooner et al. (2004) had difficulty distinguishing herbarium specimens of S. ×vallis-mexici from S. stoloniferum, and poorly preserved specimens of S. verrucosum could also be confused with either of them. Spooner et al. (2004) maintained this nothospecies based on some populations documented to be triploid and specimens with a generalized aspect of S. ×vallis-mexici.

Spooner et al. (2004) illustrated some of this variability in their Figs. 32 (the type of S. fendleri, illustrating a short plant with few lateral leaflets and an enlarged terminal leaf), Fig. 33 (the type of S. polytrichon, illustrating a densely pubescent plant with few lateral leaflets and enlarged terminal leaflet), and Fig. 34 (a common morphotype of S. stoloniferum, a taller plant with more, and subequal-sized lateral and terminal leaflets). Correll (1962) also was confounded by tremendous variation in S. stoloniferum, but recognized names Spooner et al. (2004) placed under S. stoloniferum and suggested even the need to segregate species from S. stoloniferum. This broad synonymy in S. stoloniferum runs counter to the supposition of Correll (1962: 374):

“It is quite probable that when lengthy detailed field studies are made a different, more strict, interpretation of this species will result in the segregation of several rather well defined entities.”

Schlechtendal first published S. stoloniferum in Linnaea 8: 255 [May or June] 1833, and he and Bouché reported on this species in detail a few months later in Verh. Ver. Preuss. Gartenb. 9: 319, tab. 2 [September] 1833. Hawkes (1957) reasoned that the lack of a clear description and confusion in location of the type invalidated the first publication, but this is incorrect because the first publication satisfied the requirements of the Code. Hawkes (1957) chose a lectotype of S. stoloniferum, a cultivated specimen from the Berlin Botanic Gardens, grown from tubers collected in the wild, now deposited at UPS. Hawkes’s lectotypification must be followed, but his opinion that the name was not validly published in Linnaea is rejected.

Hawkes (1944) designated Balls et al. 4864 as the type of S. malinchense, and Balls et al. 5097 as the type of S. longipedicellatum var. longimucronatum, but more than one specimen of these numbers exist, necessitating that Spooner et al. (2004) choose a lectotype. Also, K has two sheets each labeled with the same numbers as these type collections, but thesr specimens are taken from plants cultivated later and thus are not type material.

Bitter (1912) based S. schizostigma on A. Aschenborn 306, from an unknown locality in Mexico. The holotype was destroyed and no duplicates have been located. The proximal part of the plant is unknown. The pseudostipules and medial pedicel articulation clearly identify it as a member of sect. Petota. Despite an excellent description, Correll (1952, 1962) and Hawkes (1990) could not assign this name to any accepted species, although Hawkes suggested it may apply to S. cardiophyllum or S. hintonii. The anthers 5-5 1/2 mm long preclude S. cardiophyllum, and the subrotate corollas preclude any 2x(1EBN) diploid species. The white subrotate corollas, styles 8-9 mm long and ovoid fruits identify S. schizostigma as S. stoloniferum.

Spooner et al. (2004) were not able to locate the types of S. orbiculatibaccatum or S. coriaceifoliolatum despite searches at WIR. Correll (1952, 1962) and Hawkes (1990) both ally both of them with the “collective species” S. stoloniferum. The description of S. orbiculatibaccatum falls entirely within the range of morphology of S. stoloniferum. The description of S. coriaceifoliolatum is ambiguous and Spooner et al. (2004) placed it in doubtful and excluded names.

References: 

Bitter, G. 1912. Solana nova vel minus cognita, V, VI, IX, X, XI.
Repert. Spec. Nov. Regni Veg. 11: 349-394, 431-473, 12: 1-10, 49-90, 136-162.

Hawkes, J.G. 1944. Potato collecting expeditions in Mexico and South America. II.
Systematic classification of the collections, 1-142. Cambridge: Imp. Bur. Pl. Breed. Genet, Imp. Agric. Bur.

Correll, D.S. 1952. Section Tuberarium of the genus Solanum of North America and Central America.
U.S.D.A. Agric. Monogr. 11: 1-243.

Hawkes, J.G. 1957. On the lectotypes of Solanum stoloniferum Schlechtendal et Bouché, S. oxycarpum Schiede and S. verrucosum Schlechtendal.
Wiss. Z. Martin-Luther-Univ. Halle-Wittenberg 6(5): 849-854.

Correll, D.S. 1962. The potato and its wild relatives.
Contr. Texas Res. Found., Bot. Stud. 4: 1-606.

Hawkes, J.G. 1990. The potato: evolution, biodiversity and genetic resources.
Oxford: Belhaven Press.

Spooner, D.M. & K.J. Sytsma 1992. Reexamination of series relationships of Mexican and Central American wild potatoes (Solanum sect. Petota): evidence from chloroplast DNA restriction site variation.
Syst. Bot. 17:432-448.

Spooner, D.M., R.G. van den Berg, & J.T. Miller 2001. Species and series boundaries of Solanum series Longipedicellata (Solanaceae) and phenetically similar species in ser. Demissa and ser. Tuberosa: implications for a practical taxonomy of sect. Petota.
Amer. J. Bot. 88: 113-130.

Spooner, D.M. & R.J. Hijmans 2001. Potato systematics and germplasm collecting, 1989-2000.
Amer. J. Potato Res. 78:237-268; 395.

Van den Berg, R.G., G. Bryan, A. del Rio, & D.M. Spooner 2002. Reduction of species in the wild potato Solanum section Petota series Longipedicellata: AFLP, RAPD and chloroplast SSR data.
Theor. Appl. Genet. 105:1109-1114.

Spooner, D.M., R.G. van den Berg, A. Rodríguez, J. Bamberg, R.J. Hijmans, & S.I. Lara-Cabrera 2004. Wild potatoes (Solanum section Petota; Solanaceae) of North and Central America.
Syst. Bot. Monog. 68: 1-209 + 9 plates.

Bohs, L. 2005. Major clades in Solanum based on ndhF sequences.
Pp. 27-49 in R. C. Keating, V. C. Hollowell, & T. B. Croat (eds.), A festschrift for William G. D’Arcy: the legacy of a taxonomist. Monographs in Systematic Botany from the Missouri Botanical Garden, Vol. 104. Missouri Botanical Garden Press, St. Louis.

Genetics: 

Chloroplast DNA restriction site data available in: Spooner and Sytsma (1992). AFLP, RAPD, and microsatellite data listed in: Van den Berg et al. (2002).

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Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith