Solanum Stipuloideum Rusby, the Correct Name for Solanum Circaeifolium Bitter

Am. J. Potato Res. (2013) 90:301–305 DOI 10.1007/s12230-013-9304-5

Solanum stipuloideum Rusby, the Correct Name for Solanum circaeifolium Bitter

D. M. Spooner & S. Knapp

Published online: 14 March 2013 # Potato Association of America 2013

Abstract Solanum L. section Petota Dumort., which in- silvestres. Todos estos cambios han involucrado sinonimia cludes the cultivated potato (S. tuberosum L.) and its wild de nombres preexistentes, pero nuestra investigación relatives, has been the subject of intensive taxonomic re- intensiva sobre nomenclatura en la mega-diversa Solanum search in the last 25 years. The last comprehensive taxo- (1000+ especies) reveló un nombre, Solanum stipuloideum nomic treatment by Hawkes (1990) recognized seven Rusby,queeselnombreválidoprevioparalaespecie cultivated and 225 wild species, yet the latest estimate is conocida por mucho tiempo como Solanum circaeifolium four cultivated and about 100 wild species. These changes Bitter. Debido a que este último nombre se usa have all involved synonymy of preexisting names, but our frecuentemente en la literatura, aquí presentamos un intensive nomenclatural research in the mega-diverse tratamiento taxonómico de S. stipuloideum con sinonimia Solanum (1000+ species) revealed a name, Solanum completa, a fin de aclarar la circunscripción de stipuloideum Rusby, that is the earlier valid name for the nomenclatura y especies para esta especie común; este specieslongknownasSolanum circaeifolium Bitter. estudio también resalta el valor de tratamientos Because this latter name is frequently used in the literature taxonómicos de todos los géneros. we here present a taxonomic treatment of S. stipuloideum with complete synonymy in order to clarify nomenclature Keywords Potato . Solanum capsicibaccatum . Solanum and species circumscription for this common species; this circaeifolium . Solanum soestii . Solanum stipuloideum . study also highlights the value of genus-wide taxonomic Taxonomy treatments.

Resumen Solanum L. sección Petota Dumort., que incluye Introduction alapapacultivada(S. tuberosum L.) y sus parientes silvestres, ha estado sujeta a investigación taxonómica Solanum is one of the largest genera of flowering plants with intensiva en los últimos 25 años. El último tratamiento over 1000 accepted species, and some 7000 names. The detallado taxonómico por Hawkes (1990) reconocía siete latest comprehensive taxonomic treatment of section especies cultivadas y 225 silvestres, pero la estimación Petota (Hawkes 1990) recognized seven cultivated and más reciente es de cuatro cultivadas y cerca de 100 especies 225 wild species, yet the latest estimate is four cultivated (Ovchinnikova et al. 2011) and about 100 wild species (Spooner 2009). In such large genera complete nomenclatural review is challenging, due in part to allocation of taxa * D. M. Spooner ( ) to species groups to which they do not belong. Such was the Vegetable Crops Research Unit, USDA, Agricultural Research Service, Department of Horticulture, University of Wisconsin, case with the name S. stipuloideum Rusby. H.H. Rusby 1575 Linden Drive, Madison, WI 53706-1590, USA (1907) described S. stipuloideum from a specimen collected e-mail: [email protected] by Miguel Bang from an unspecified locality in Bolivia (Bang 2509) and suggested it was a “Species near S. S. Knapp ” Department of Life Sciences, The Natural History Museum, boerhaavifolium but not having its stems or calyx. Below Cromwell Road, London SW7 5BD, UK we copy Rusby’s description: 302 Am. J. Potato Res. (2013) 90:301–305

Sparsely and very shortly stellate on the lower leaf- acuminate and obtuse at the apex, entire to obscurely surfaces and calyx; a low diffuse or reclining herb; sinuate-dentate, very thin and flaccid, dark-green branches very slender, flexuous, angled, pale; petioles above, pale beneath, the weak venation coarse and 0.5–3 cm long, mostly with one or two small stipuloid irregular; pedicels mostly solitary, 1–3 cm long, fili- leaves at the base; blades 2–4cm.long,1–3cm. form; calyx 5 mm long, the turbinate tube very short, broad, truncate to slightly cordate at the base, short- the herbaceous lobes lance-ovate, obtuse; corolla

Fig. 1 Lectotype specimen of Solanum stipuloideum (A.M. Bang 2509) at the New York Botanical Garden (herbarium code NY) Am. J. Potato Res. (2013) 90:301–305 303

white, nearly 1.5 cm long, not deeply lobed, the lobes acutish; anthers bright-yellow, 5 mm long, broadly oblong, straight, the small pores looking inward and slightly laterally; style 2–3 mm longer than stamens, the stigma broad ([Bang] No 2509).

Recent work in Solanum (Knapp 2013) along with the large scale type digitization project the Global Plants Initiative (see JSTOR Plant Science, http://plants.jstor.org), and research into potatoes (Spooner 2009) has examined the type specimens of Bang 2509 at New York and at Kew; examination of these specimens revealed that they pos- sessed the articulated pedicels of section Petota, and repre- sented the same taxon as S. circaeifolium, described 5 years later by the German botanist Georg Bitter (Bitter 1912). The principle of priority that underpins the Code (McNeill et al. 2012a, b) dictates that the oldest name is that which must be Fig. 2 Close up of the lectotype specimen of Solanum stipuloideum used, so S. stipuloideum must replace the later name S. showing the pseudostipules and pedicel articulation above the base characteristic of Solanum section Petota, and a stellate corolla charac- circaeifolium. We choose not to conserve (McNeill et al. teristic of Solanum circaeifolium 2012a, b)theearliernameS. circaeifolium over S. stipuloideum because final decisions on conservation are not made until the International Botanical Congress (the pubescence, terminal leaflet size and shape, peduncle and next in 2017) and because this species name is being used pedicel length, and corolla diameter as key characters to in upcoming floristic and monographic treatments. distinguish their taxa. Specimens of the typical variety The species known as as Solanum circaeifolium is were identified as narrowly restricted to the extreme one of the most distinctive species in section Petota. northern portion of the species range near Lake Titicaca Unlike most South American species of the section, but in Dept. La Paz, S. soestii to a single locality in southern like many North and Central American diploid species Dept. La Paz, and the other taxa more widespread and (VandenBergetal.2004), it has white, stellate co- overlapping in range (Hawkes and Hjerting 1989; Ochoa rollas and an endosperm balance number of 1. Its 1990). elliptic-conical fruits are somewhat similar to those of Van den Berg and Groendijk-Wilders (1999)evaluat- members of the Conicibaccata group, but they are ed the integrity of these taxa with morphological data narrower and distinctly pointed at the end. Bang 2509 and Van den Berg et al. (2001)withRAPDandAFLP clearly is the same as S. circaeifolium as is evident data. The morphological data documented extensive from a comparison of the description (above), and an overlap of all 26 characters they analyzed, but 18 of examination of the type specimen (Figs. 1 and 2). It them showed statistically significant differences among matches in all aspects specimens we have on loan of taxa (0.05 level). The RAPD data distinguished S. this species, is found in the same generalized locality, soestii but not the other taxa and the AFLP data distin- has the diagnostic medial pedicel articulation and guished all of the taxa recognized by Hawkes (1990). pseudostipules of section Petota, and the stellate co- However, a subsequent AFLP study by Jacobs et al. rollas of S. circaeifolium (Fig. 2). Both Correll (1962) (2008), using a much wider range of species, failed to and Ochoa (1990) cite Bang 2509 at NY as S. separate the numerous accessions used into species- circaeifolium. They do not associate this with Rusby’s specific groups. In addition, our examination of herbar- description, however. ium specimens shows that there is overlap of all char- Solanum stipuloideum as circumscribed here has been acters for reliable discriminations of any of these taxa, treated in various ways (reviewed in Van den Berg and and as a consequence we consider all of these previ- Groendijk-Wilders 1999). The most recent treatment by ously recognized taxa as conspecific. Hawkes (1990) recognized three species and two subspe- cies (S. circaeifolium subsp. circaeifolium and subsp. quimense, S. capsicibaccatum,andS. soestii). Ochoa Taxonomy (1990) recognized two species and two varieties (S. circaeifolium var. circaeifolium and var. capsicibaccatum, Note: the symbol “!” behind a specimen indicates that it was and S. soestii). These authors emphasized leaf and calyx seen and studied by the authors. 304 Am. J. Potato Res. (2013) 90:301–305

Solanum stipuloideum Rusby, Bull. New York Bot. Solanum stipuloideum is diploid (2n=2×=24) and pos- Gard 4: 419. 1907. TYPE: BOLIVIA. sin. loc., no date, sesses an endosperm balance number of 1. It flowers and A.M. Bang 2509 (lectotype: NY!; isolectotype: K! fruits mainly from December to April. It is endemic to [K000585637]). Bolivia, from the Department of La Paz south to the Solanum circaeifolium Bitter, Repert. Spec. Nov. Regni Department of Chuquisaca; occuring in a wide range of Veg. 11: 385. 1912—TYPE: BOLIVIA. La Paz: Prov. habitats, in full sun or in partial shade, typically in moist Larecaja, Monte Sorata, Cerro Iminapi, 2650 m, Mar areas, on rocky slopes, among bushes, in scrub or thorn 1860, G. Mandon 400 (holotype: W-0006516! [Correll forests, in or at the edges of forests, and in cultivated fields; neg. 795, BM000881722!, F-1603086!, LL!, MO- from 2000 to 4000 m in elevation. 5594969!, NY-00139095!, UC1152236!; F neg. 12, ThetypelocalityofS. stipuloideum is likely to be in F-1463333!]; isotypes: BM! [BM000603448], F! [fragment the Department of La Paz; the printed locality on the of type but provenance not stated], G [F neg. 23105, F- label of the isotype (“Yungas”) is deliberately crossed 757493!, MO-1691508!, NY!], K! [K000005826, = Correll out, and other Bang specimens with collection numbers neg. 218, BM000881723!, F-1603979!, LL!, NY!, near this one (e.g., Bang 2492, the type of S. dianthum UC1152236!], NY! [NY00139095=Correll neg. 770, Rusby) are from the province of Nor Yungas. It is, BM000881721!, F-1603111!, LL!, NY!, UC1152236!]; P! however, likely that this was caused by re-using existing [Correll neg. 219, BM000881724!, F-1603980!, MO- labels for the specimens they distributed until they were 5594968!, LL!, NY!, UC1152236!]). gone. Solanum capsicibaccatum H. M. Cárdenas, Rev. Agric. We propose stp as the three letter abbreviation of Univ. Cochabamba, Bolivia 2(2): 35. 1944. Solanum Solanum stipuloideum Rusby. circaeifolium var. capsicibaccatum (H.M.Cárdenas)C. M. Ochoa, Phytologia 54: 392. 1983.—TYPE: BOLIVIA. Acknowledgments This research was funded by the National Sci- Cochabamba: Prov. Tarata, between Huaira Pata and Molle ence Foundation Planetary Biodivesity Inventory Program (NSF DEB 0316614). We thank Alan Whittemore and two unnamed reviewers for Ppujru, valley of Río Caine, 2800 m, Mar 1942 [in review. protologue, but April 1942 on specimens], H. Gandarillas 60 (lectotype, designated here: GH! 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Ochoa, Phytologia 50: 181. 1982. Solanum circaeifolium Jacobs, M.M.J., R.G. van den Berg, V.G.A.A. Vleeshouwers, M. var. latifoliolatum (C. M. Ochoa) C. M. Ochoa, Phytologia Visser, R. Mank, M. Sengers, R. Hoekstra, and B. Vosman. — 2008. AFLP analysis reveals a lack of phylogenetic structure 54: 392. 1983. TYPE: BOLIVIA. La Paz: Prov. Inquisivi, within Solanum section Petota. BMC Evolutionary Biology Río Seco, 2900 m, Mar 1978, C. M. Ochoa 11915 (holo- 8(145): 1–12. doi:10.1186/1471-2148-8-145. type: C. M. Ochoa’s personal herbarium; isotypes: C!, US! Knapp, S. 2013. A revision of the Dulcamaroid Clade of Solanum L. [US00037270]). (Solanaceae). PhytoKeys, in press. McNeill, J., F.R. Barrie, W.R. Buck, V. Demoulin, W. Greuter, D.L. Solanum soestii J. G. Hawkes & J. P. Hjerting, Bot. J. Hawksworth, P.S. Herendeen, S. Knapp, K. Marhold, J. Prado, Linn. Soc. 86: 406. 1983.—TYPE: BOLIVIA. La Paz: Prov. W.F. Prud’homme van Reine, G.F. Smith, J.H. 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