Solanaceae Source

A global taxonomic resource for the nightshade family

Solanum acaule

Citation author: 
Bitter
Citation: 
Repert. Spec. Nov. Regni Veg. 11: 391. 1912.
Type: 
Bolivia. Puna Putanca, 8 Jan 1904, K. Fiebrig 3429 (holotype, B, destroyed; lectotype, S-042822 [upper left hand plant] designated by Hawkes and Hjerting, 1969: 250; isotypes, G00074132 [Correll neg. 317, F-1603729, LL, NY, UC1152154], JE, LD [Correll neg. 911, F-1603007, LL, NY, UC1152154], M0100317, NY00139040 [Correll neg. 318, F-1603728, LL, MO-5588146, NY, UC1152154], SI, U0006802, W-1739 [Correll neg. 625, F-1603325, LL, NY, UC1152153], Z [Correll neg. 626, F-1603326, LL, NY, UC1152154]).
Last edited by: 
Spooner, D.M.
Written by: 
Spooner, D.M. & A. Clausen
Habit: 
Herbs 0.1-0.3 m tall, rosette forming to erect. Stems 2-4 mm in diameter at base of plant, green, sometimes purple or green and purple mottled, unwinged, subglabrous to moderately pubescent with short multicellular hairs; tubers typically placed one at the end of each stolon.
Sympodial structure: 
Sympodial units tri- to plurifoliate, not geminate.
Leaves: 
Leaves odd-pinnate, the blades 9-24.5 x 2.2-6 cm, green, membranous to chartaceous, subglabrous to moderately pubescent adaxially and abaxially with hairs like those of the stems; lateral leaflet pairs 4-6, subequal except for the most proximal 1-2 pairs that are greatly reduced in size, to unequal with the most distal pairs clearly larger; most distal lateral leaflets 1.6-4.4 x 0.8-2.4 cm, broadly ovate to broadly elliptic, the apex obtuse, the base typically oblique to cuneate, sessile and broadly decurrent to petiolulate with petiolules up to 2 mm long; terminal leaflet 2.4-7 x 1.3-4.2 cm, orbicular to very broadly obovate, the apex rounded to obtuse, the base truncate to cuneate; interjected leaflets 0-7, sessile to short petiolulate, ovate to orbicular, petiolules 0-5 mm; petioles 2.2-6.5 cm, pubescent as the stems. Pseudostipules absent to minute and scale-like to 1 mm long, pubescent with hairs like those of the stem.
Inflorescences: 
Inflorescences 3-7 cm, distinct in this species, typically a monochasium with an "extra" flower at its base (see discussion), often near the base but sometimes in the distal half of the plant, unbranched, with 1-4 flowers, with all flowers apparently perfect, the axes pubescent with hairs like those of the stem; peduncle highly contracted and often single flowered, 1-4 mm long; pedicels 13-35 mm long in flower and fruit, spaced 1-10 mm apart, articulation absent.
Flowers: 
Flowers homostylous, 5-merous. Calyx 3-5 mm long, the tube 1-2 mm, the lobes 1-4 mm, acute to acuminate, the acumens 1-3 mm long, with hairs like those of the stem. Corolla 1.4-2.5 cm in diameter, rotate, blue to more rarely white or blue and white streaked adaxially and abaxially, the tube 1-2 mm long, the acumens 0-1 mm long, the corolla edges flat, not folded dorsally, glabrous adaxially, minutely puberulent abaxially, especially along the midribs, ciliate at the margins, especially at the tips of the corollas. Stamens with the filaments 1-2 mm long; anthers 2-3 mm long, lanceolate, connivent, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style ca. 3.5 mm x ca. 1 mm, exceeding stamens by ca. 1 mm, straight, glabrous; stigma clavate to capitate.
Fruits: 
Fruit a globose to slightly ovoid berry, 1-2 cm wide, 1-2.2 cm long, green to green tinged with purple when ripe, glabrous.
Seeds: 
Seeds from living specimens ovoid and ca. 2 mm long, whitish to greenish in fresh condition and drying brownish, with a thick covering of “hair-like” lateral walls of the testal cells that make the seeds mucilaginous when wet, green-white throughout; testal cells honeycomb-shaped when lateral walls removed by enzyme digestion.
Chromosome number: 

2n = 4x = 48 voucher: Ochoa & Salas 14945 (CIP) (Hijmans, et al. 2007)

Distribution: 

Solanum acaule is widespread and common in upland habitats from northern Peru (Dept. Cajamarca), south through Bolivia to northern Argentina (Prov. San Juan), and with one record in northern Chile (Antofagasta Region), on dry rocky hillsides, high puna, among herbs, spiny shrubs and low woods, along streamsides, dry river beds and alluvial cones, (2000) 2400-4700 m in elevation.

Phenology: 
Flowering and fruiting mainly from December to April.
Phylogeny: 

Solanum acaule is a member of Solanum sect. Petota Dumort., the tuber-bearing cultivated and wild potatoes. Within sect. Petota, Solanum acaule is a member of a very diverse clade related to the cultivated potato. On a higher taxonomic level, it is a member of the informally-named Potato Clade, a group of perhaps 200-300 species that also includes the tomato and its wild relatives (Bohs, 2005).

Commentary: 

Correll (1962), Hawkes (1990), and Ochoa (1999) placed S. acaule (tetraploid) in series Acaulia Juz., and related it to S. albicans (Ochoa) Ochoa, a hexaploid species from Ecuador and northern Peru. Many studies have made it clear, however, that both S. acaule and S. albicans are also closely related to S. demissum Lindl., a species from Mexico and Guatemala that Correll (1962) and Hawkes (1990), placed in the separate series Demissa Bukasov. The relationships of S. acaule, S. albicans, and S. demissum were shown by Bonierbale et al. (1990) and Debener et al. (1990) with single- to low-copy nuclear restriction fragment polymorphisms, by Spooner et al. (1995) and Kardolus (1999) with morphological data, by Kardolus (1998) and Jacobs et al. (2008) with Amplified Fragment Length Polymorphism (AFLP) data, and by Nakagawa and Hosaka (2002) with plastid and nuclear RFLPs. Based on these data Spooner et al. (2004) extracted S. demissum from its other North and Central American hexaploid relatives in series Demissa Bukasov, and classified S. acaule, S. albicans, and S. demissum in an informal Acaulia Group.

The taxonomy of S. acaule and S. albicans has been a state of flux, with changing ideas on the status of S. albicans, and with varying treatments of infraspecific categories within S. acaule at the subspecies, varietal, and form ranks. We do not partition S. acaule into infraspecific categories as do Correll (1962), Hawkes (1990), and Ochoa (1999).

Hosaka and Spooner (1992) used 94 probe-enzyme combinations of nuclear RFLPs to examine 105 accessions of all three subspecies (sensu Hawkes, 1990) of S. acaule (subsp. acaule, aemulans, punae) and of S. albicans throughout their distribution area, from northern Peru to northern Argentina. They were able to support the recognition of S. albicans and all subspecies of S. acaule except subsp. acaule and subsp. punae that were combined into one. In the present treatment we recognize S. acaule, S. albicans, and S. ×aemulans at the species level with no infraspecific categories, and treat subsp. punae as a synonym of S. acaule.

Kardolus (1999) used phenetic analysis of morphological data to distinguish most accessions of subsp. acaule and subsp. punae, and he ascribed the mixture of the two in the study of Hosaka and Spooner (1992) to be due to misidentifications. McGregor et al. (2002) likewise supported both subsp. acaule and subsp. punae with AFLP data, but the molecular distinction between the two was slight, and again, reidentifications were needed to make their AFLP and morphological data fit each other. Taxonomists have distinguished these two taxa by overlapping character states of habit and pubescence, with subsp. punae very rosette-like and with long-spreading hairs on the stem, petiole, and leaf, and subsp. acaule with less flat rosettes and shorter appressed hairs. We find these characters to be so variable as to be impossible to use.

More difficult to treat is S. acaule subsp. palmirense. Kardolus (1998) described this as the first known hexaploid cytotype of S. acaule, based on a single collection from Chimborazo Province, Ecuador (Spooner et al. 5070). At that time, this was the only known Ecuadorian collection of either S. acaule or S. albicans, but our treatment found another collection of S. albicans from the same province (Ochoa 13395). Kardolus (1998) classified subsp. palmirense in S. acaule, rather than in S. albicans, based on AFLP affinities of the former, despite the fact that its ploidy and morphology was a near match for S. albicans. A subsequent AFLP study by McGregor et al. (2002) also grouped subsp. palmirense with S. acaule. Solanum albicans is considered to be a hybrid of S. acaule and another species (Hawkes, 1990), and its AFLP distinction from other populations of S. albicans may be due to another hybrid origin. While Kardolus’s (1998) placement of subsp. palmirense into S. acaule may be correct, we consider this issue unsolved from lack of sufficient comparative material. We synonymize this subspecies into S. albicans because of its morphological and ploidy affinities with that species.

Solanum acaule is in the pedigree of the hybrid cultivated potato species S. juzepczukii Bukasov (a triploid hybrid of diploid clones of S. tuberosum L. and S. acaule) and S. curtilobum Juz. and Bukasov (a pentaploid hybrid of tetraploid clones of S. tuberosum and S. juzepczukii) (Hawkes, 1962; Schmiediche et al., 1980, 1982). It is also a parent in the hybrid wild species S. ×indunii K. A. Okada and A. M. Clausen (S. acaule × S. boliviense Dunal) (Okada and Clausen, 1982), and S. ×viirsooi K. A. Okada and A. M. Clausen (S. acaule × S. infundibuliforme Phil.) (Okada and Clausen, 1985).

Kardolus and Groendijk-Wilders (1998) studied the inflorescence architecture of members of series Acaulia, series Demissa, and series Megistacroloba. They described five types of inflorescence architecture, including a type not recognized previously of a monochasium with an "extra" flower at its base in series Acaulia and S. demissum. None of these were species-specific and even varied on individual plants, but showed species-specific tendencies towards one type.

References: 

Vellozo, J.M. C. 1829. Flora fluminensis. Volume 2.
Rio de Janeiro, Brazil.

Hawkes, J.G. 1962. The origin of Solanum juzepczukii Buk. and S. curtilobum Juz. et Buk.
Zeitschrift für Pflanzenzüchtung 47: 1–14.

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

Schmiediche, P.E., J.G. Hawkes & C.M. Ochoa 1980. Solanum ×juzepczukii Buk. and S. ×curtilobum Juz. et Buk. I. A study of the natural variation of Solanum ×juzepczukii, S. ×curtilobum and their wild progenitor, S. acaule.
Bitt. Euphytica 29: 685–704.

Schmiediche, P.E., J.G. Hawkes & C.M. Ochoa 1982. Solanum ×juzepczukii Buk. and S. ×curtilobum Juz. et Buk. II. The resynthesis of Solanum ×juzepczukii and S. ×curtilobum.
Euphytica 31: 395–707.

Okada, K.A. & A.M. Clausen 1982. Natural hybridization between Solanum acaule Bitt. and S. megistacrolobum Bitt. in the Province of Jujuy, Salta.
Euphytica 31: 817-835.

Okada, K.A. & A.M. Clausen 1985. Natural triploid hybrids between Solanum acaule Bitter and S. infundibuliforme Philippi in the Province of Jujuy, Argentina
Euphytica 34: 219-231.

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

Bonierbale, M., W. Ganal & S.D. Tanksley 1990. Applications of restriction fragment length polymorphisms and genetic mapping in potato breeding and molecular genetics.
In: Vayda, M. E. and W. D. Park (eds.), The molecular and cellular biology of the potato. CAB International, Wallingford, U.K. pp. 13-24.

Hosaka, K. & D.M. Spooner 1992. RFLP analysis of the wild potato species, Solanum acaule Bitter (Solanum sect. Petota).
Theor Appl Genet 84: 851-858.

Spooner, D.M., R.G. van den Berg, & J.B. Bamberg 1995. Examination of species boundaries of Solanum series Demissa and potentially related species in series Acaulia and series Tuberosa (sect. Petota).
Syst. Bot. 20: 295-314.

Kardolus, J.P., & N. Groendijk-Wilders 1998. The inflorescence architecture of Solanum acaule and related taxa of Solanum section Petota (Solanaceae).
Acta Bot. Neerl. 47: 195-207.

Kardolus, J.P. 1998. A biosystematic analysis of Solanum acaule.
Wageningen Agricultural University, Wageningen, The Netherlands.

Ochoa, C.M. 1999. Las papas de sudamerica: Peru (Parte I).
Lima, Peru: International Potato Center.

Kardolus, J.P. 1999. Morphological variation within series Acaulia Juz. (Solanum sect. Petota).
In Solanaceae IV: Advances in Biology and Utilization, ed. M. Nee, D. E. Symon, R. N. Lester, and J. P. Jessop, 257-274. Kew, U.K.: Royal Botanic Gardens, Kew.

Nakagawa, M., & K. Hosaka. 2002. Species relationships between a wild tetraploid potato species, Solanum acaule Bitter, and its related species as revealed by RFLPs of chloroplast and nuclear DNA.
Amer. J. Potato Res. 79: 85-98.

McGregor, C.E., R. van Treuren, R. Hoekstra, & Th. J.L. van Hintum. 2002. Analysis of the wild potato germplasm of the series Acaulia with AFLPs: implications for ex situ conservation.
Theor. Appl. Genet. 104: 146–156.

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.

Hijmans, R., T. Gavrilenko, S. Stephenson, J. Bamberg, A. Salas & D.M. Spooner 2007. Geographic and environmental range expansion through polyploidy in wild potatoes (Solanum section Petota).
Global Ecol. Biogeogr. 16: 485-495.

Jacobs, M.J., R.G. van den Berg, V.G.A.A. Vleeshouwers, M. Visser, R. Mank, M. Sengers, R. Hoekstra & B. Vosman 2008. AFLP analysis reveals lack of phylogenetic structure within Solanum section Petota.
BMC Evol. Biol. 8, 145: 2-12.

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