New species of Solanum and Capsicum (Solanaceae) from Bolivia, with clarification of nomenclature in some Bolivian Solanum M I C H A E L N E E , L Y N N BOHS ~, AND SANDRA K N A P P Nee, M. (The New York Botanical Garden, 200th Street & Kazimiroff Boulevard, Bronx, NY 10458-5126, U.S.A.; e-mail: mnee@nybg.org), L. Bohs (Department of Biology, 257 South 1400 East, University of Utah, Salt Lake City, UT 84112-0840, U.S.A.; e-mail: bohs@biology.utah.edu) & S. Knapp (Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom; email: S.Knapp@nhm.ac.uk). New species of Solanum and Capsicum (Solanaceae) from Bolivia, with clarification of nomenclature in some Bolivian Solanum. Brittonia 58: 322-356. 2006.--Nine new species of Solanum and two of Capsicum are described from Bolivia. Notes are provided on some other species, including the complex typification of Solanum aridum. Capsicum eaballeroi, C. ceratoealyx, Solanum chalmersii, S. clandestinum, S. comarapanum, S. eomplectens, S. monanthemon, S. moxosense, S. pedemontanum, S. saturatum, and S. whalenii are described and illustrated, and a new name, S. scuticum, is proposed for the species previously known as S. tabacifolium. Key Words: biodiversity, Bolivia, Capsicum, Solanaceae, Solanum, South America. Bolivian floristic diversity is high, reflecting its great topographic and habitat diversity. Habitats in Bolivia range from seasonally flooded savannas to arid Chaco and high elevation deserts to hyper-humid montane and lowland rainforests. Four areas in the country have been identified as centers of plant diversity and endemism: the Gran Chaco (SA 22 of Davis et al., 1997), southeastern Santa Cruz (SA 23 of Davis et al., 1997), the Llanos de Mojos region (SA 24 of Davis et al., 1997), and the Madidi-Apolo region (SA 36 of Davis et al., 1997). The position of Bolivia at the western edge of the Amazon basin with large areas occupied by the eastern flank of the geologically young Andes makes it a particularly rich region for Solanaceae, whose diversity is largely associated with the Andean slopes (Gentry, 1982; Knapp, 2002a). The genus Solanum in particular is highly diverse in the Andes, but lack of collections for Bolivia may have led to the IAuthor for correspondence country's being relatively neglected as a "hot-spot" of solanaceous diversity. Recent work by a number of institutions, both Bolivian and others, has led to an explosive increase in the number of Bolivian collections available for study. This has uncovered many new, narrowly endemic taxa and has led to the re-examination and clarification of other, previously poorly understood species. Renewed impetus for the study of the Bolivian flora will undoubtably reveal many more new taxa in the future, and the new generation of Bolivian botanists actively working on the flora will accelerate this even further. One of the species described here, Solanum pedemontanum, shows how even widespread and evidently common species have remained nearly unknown until collecting in previously little-explored parts of tropical America greatly increased in the last few decades. Exploration and collecting are also essential for uncovering the existence of rare, local endemics, such as Capsicum caballeroi or S. moxosense. The following notes and new species are Brittonia, 58(4), 2006, pp. 322-356. 9 2006, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. ISSUED: 28 December 2006 2006] NEE ET AL.: SOLANUM AND CAPSICUM (SOLANACEAE) based primarily on 15 years of botanizing in Bolivia by Nee, by a field trip by Nee and Bohs in 1998, and one by all three authors in May of 2001. This last trip was instrumental in solidifying our concepts of a number of new and critical taxa, and provided the opportunity to see them in the field and obtain material for molecular studies. The notes and new species here are in anticipation of the Catalogue of the Vascular Plants of Bolivia by P. JCrgensen et al. (in prep.) of the Missouri Botanical Garden and a number of Bolivian institutions, and are part of the worldwide revision of Solanum being undertaken by the authors in collaboration with Dr. David Spooner of the University of Wisconsin/U.S.D.A, and a host of other contributors. This paper, and one of the most striking new species, is dedicated to the memory of Dr. Michael D. Whalen (1950-1985), the major professor of Knapp at Cornell University and colleague of Nee. Dr. Whalen was travelling in Peru and scheduled to meet with Nee on his first trip to Bolivia in late 1984. It was at this time that Dr. Whalen began to experience the visual problems which cut short his Peruvian trip and which were symptoms of the brain cancer that tragically ended his productive botanical life. He thus never had the chance to visit Bolivia and study its varied and fascinating Solanaceae. He was a professor, mentor, and friend sorely missed even still. Notes on systematic characters The species described here belong to the genera Capsicum and Solanum, both of Solanaceae subfamily Solanoideae. Capsicum is distinguished by its longitudinal anther dehiscence and, in at least the majority of species, its pungent fruits. In the Capsicum species described here, the calyx margin is truncate with five or ten appendages that emerge from below the calyx rim. Similar calyces are found in the related genus Lycianthes Hassl., and they have a different pattern of vasculature than is found in other Solanaceae with a more conventional calyx structure (D'Arcy, 1986). The generic characters distinguishing Solanum are poricidal anther dehiscence and lack of the specialized Lycianthes calyx 323 structure described above. In many Solanum species, particularly in Solanum subgenus Leptostemonum, the anthers are tapered distally and the terminal pores do not enlarge as the flower ages. Other species have oblong anthers with blunt tips and the pores expand into longitudinal slits with age. Most Solanaceae have complex branching patterns on their flowering shoots, and the details of these patterns can be taxonomically useful. Inflorescences are morphologically terminal. Further stem growth occurs by expansion of axillary shoots located below the inflorescence; these, in turn, will terminate in an inflorescence. Thus, the flowering portion of the plant is composed of a series of sympodial units, with number and arrangement of leaves in each sympodial unit of systematic importance. For instance, Solanum section Geminata takes its name from the frequent occurrence of two-leaved sympodia in which the leaves are arranged in pairs (geminate). More information on branching patterns in Solanaceae can be found in Danert (1958, 1967), Child (1979), Bohs (1989), Bell and Dines (1995), and Knapp (2002b). Capsicum L. Bolivia is especially rich in species of Capsicum; eight wild or domesticated taxa are mapped for Bolivia by Eshbaugh (1975). With the two species described below, nine native or naturalized species are now known from the country (Nee, unpubl, data), while several others are widely grown for their pungent fruits. Neither of the new species listed below appears to be close to any of the domesticated species, nor is either known to be gathered in the wild or to enter into commerce. Capsicum caballeroi M. Nee, sp. nov. TYPE: Bolivia. Santa Cruz. Prov. Caballero: Parque Nacional Ambor6, Cerro Bravo, 10 km al N de Comarapa, 17~ 64~ 2400-2500 m, 7-10 Apr 1994 (fl, fr), I. Vargas C. & J.M. Camacho 3118 (HOLOTYPE: U S Z ; ISOTYPES: CORD, MO, NY, US). (Fig. 1) Herba vel frutex, 1-7 m altus. Inflorescentia axillaris, 1-2-flora, pedicellis per anthesin 20-25 mm longis, fructiferis 24-45 mm longis; calyx cupulatus, 2.5 mm 324 BRITTONIA [VOL. 58 FIG. 1. Capsicum caballeroi M, Nee. A. Flowering branch. B. Branch with detail of flower and fruit. C. Flower. D. Corolla spread open to show stamens. E. Gynoecium and section of calyx. E Branch with mature fruit. G. Seed. ( A - E based on Nee et al. 52407, NY; F - G based on Dorr & Barnett 7041, NY.) 2006] N E E ET AL.: SOLANUM AND CAPSICUM (SOLANACEAE) longus, appendicibus 5, per anthesin 0.8-1.8 mm longis, plernmque 5 appendicibus interpositis brevioribus; corolla angusti-campanulata, 10.5-13 • 4~5 mm, lobis 3 x 2 ram, flava; filamenta 4.5 mm longa; antherae 2-2.1 mm longae. Fmctus globosus, 9-11 man diam., vivide tuber, sapore pungenti; semina 5-17, reniformia, 3.84.2 x 3.2 mm. Herb, shrub or treelet, 1-7 m tall; stems glabrous or sparsely pubescent with simple hairs. Sympodial units difoliate and geminate. Leaves 2 - 1 3 x 0.8-4.2 cm, more or less uniform in size and shape, lanceolate, glabrous adaxially and abaxially or sparsely pubescent abaxially along the midrib with simple hairs 0.3 m m long; base acute to attenuate and somewhat oblique; margin slightly revolute; apex attenuate; petioles 2 - 8 mm. Inflorescences axillary, 1-2-flowered; pedicels 2 0 - 2 5 m m in flower, 2 4 - 4 5 m m in fruit, terete, pendulous, slender, 0.6 m m wide at base, 1.4-1.8 m m wide distally, glabrous. Calyx cupulate, 2.5 m m long, the margin truncate, with 5 appendages 0.8-1.8 m m long in flower and with 5 intermediate, slightly shorter ones alternating with these, the appendages (1-) 3 - 5 m m long in fruit, sparsely pubescent with simple hairs; corolla 4 - 6 m m in diameter, 10.5-13 m m long, narrowly campanulate, lemon yellow, shallowly 5lobed, the tube 3 - 6 mm, the lobes ca. 3 x 2 ram, narrowly triangular, acute at apices, glabrous abaxially, papillose at tips of lobes; stamens included; filaments 4.5 mm, attached ca. 1 m m above base o f corolla tube, broadened at base, but without two flaps of tissue on corolla tube above site of insertion; anthers 2-2.1 x 0.8-0.9 mm, oblong, yellow, longitudinally dehiscent; ovary glabrous; style ca. 6 x 0.25 mm, cylindrical to clavate, glabrous; stigma truncate to capitate. Fruit a globose berry, 9-11 m m in diameter, pendent, glabrous, bright red, pungent (or apparently sometimes not); seeds 5-17, 3.8-4.2 x 3.2 mm, reniform, flattened, pale yellow, the surface loosely foveolate. Distribution and e c o l o g y . I K n o w n only f r o m cloud forests (yungas) with Podocarpus spp., Prumnopitys exigua De Laub., Weinmannia spp., Alnus acuminata Kunth subsp. acuminata, and Myrtaceae (including Blepharocalyx salicifoIius O. Berg) between 1880 and 2600 m elevation in Provinces o f Florida and Caballero o f Dept. Santa Cruz and just to the northwest in this same ecolog- 325 ical zone into Dept. Cochabamba, Prov. Carrasco, in the adjacent Parque Nacional Carrasco. P h e n o l o g y . - - F o u n d in flower in April, May, and N o v e m b e r and in fruit in January, March, May, and November; it likely flowers and fruits all year long. E t y m o l o g y . - - N a m e d in honor o f Bolivian botanist Israel Gerardo Vargas Caballero, whose own investigations and those of his students are making wild and domesticated plants of this part o f Bolivia much better known, and incidentally (although incorrect grammatically) for the Province Manuel Maria Caballero, where most o f the specimens have been collected. Additional specimens examined. BOLIVIA. COCHABAMBA.Prov. Carrasco: Serranfa Siberia, 20-35 km W of Comarapa, on the old Cochabamba-Santa Cruz road, ca. 2000 m, 14-15 Jan 1990 (fr), Dorr & Barnett 7041 (NY). SANTACRUZ. Prov. Caballero: Parque Nacional Ambor6, Cerro Bravo, cerca Comarapa, 2600 m, 17 Jun 1995, A. Jardim et al. 1995 (MO, NY); Nee et al. 52407 (LPB, NY, USZ); 50 km N de Mataral (en la carretera Santa Cruz~Comarapa) pasando pot San Juan del Potrero y bajando a la cuenca del alto Rio Ichilo, 23002450 m, 28 May 1989 (fl, fr), Smith et al. 13470 (BOLV, LPB, MO, NY); Siberia-El Empalme, 5 km entrando hacia Khara Huasi, 17~ 64~ 2300 m, 8-9 May 1992 (fr), Vargas & Prado 1282 (MO, NY), 1286 (NY); San Juan del Potrero, Naranjos, 17~ 64~ 2150 m, 12-13 May 1992 (fr), Vargas et al. 1343 (NY); Parque Nacional Ambor6, Cerro Bravo a 10 km N de Comarapa, 17~ 64~ 2400-2500 m, 15 Nov 1995 (fl, fr), Vargas et al. 4151 (NY). Prov. Florida: La Yunga, 7.5 km (linea recta) NE de Mairana, 18~ 63~ 1880 m, 15 Mar 1997 (fr), Saldias 4977 (MO). This species is characterized by its narrowly campanulate lemon yellow corollas, very long pedicels, and stamens with relatively long filaments; the fruits are pendent. The Bolivian species C. eximium and C. cardenasii also have campanulate corollas, but those o f C. caballeroi are much larger and narrower; in addition, C. eximium and C. cardenasii have purple (or rarely whitish) corollas, in contrast to the yellow corollas o f C. caballeroi. The c o m m o n names "ajf de monte" (Vargas & Prado 1282) and "ulupica de yunga" (Vargas et al. 1343) indicate the great similarity to other local species o f Capsicum; in this area the cultivated peppers are called " a f t ' and the wild species (e.g., C. eximium Hunz., from which pungent fruits are col- 326 BRITTONIA lected for local use and the markets) are called "ulupica." Two of the collection labels indicate that the fruits of C. caballeroi are pungent (Vargas & Prado 1282; Vargas et al. 1343), while one indicates they are not (Vargas & Prado 1286). C a p s i c u m ceratocalyx M. Nee, sp. nov. TYPE: Bolivia. La Paz. Prov. Sud Yungas: 7.5 km (by road) from Huancan6 on road to San Isidro, moist montane forest, 16~ 67~ 2225 m, 10 May 2001 (buds), M. Nee, L. Bohs, S. Knapp & J.M. Mendoza E 51778 (HOLOTYPE: LPB; ISOTYPES: MO, NY, USZ). (Fig. 2) Frutex ad 1.5 m altus. Folia geminata, 5.5-22.5 • 26.5 cm, fete glabra. Inflorescentia axillaris, 6-9-flora, pedicellis per anthesin 9 mm longis, fructiferis 19-23 mm longis, alatis; calyx appendicibus 5, per anthesin 22.5 mm longis, curvatis; corolla flava, intus viridimaculata; antherae longitudinaliter dehiscentes. Fructus baccatus, coccineus, 1 cm diametro. Shrub 1.5 m. tall; young stems minutely puberulent with antrorsely curved simple hairs, older stems nearly glabrous. Sympodial units difoliate and geminate. Leaves 2 22.5 • 0.6-6.5 cm, dimorphic, the minor leaves similar to the majors but about 1/3 the size, elliptic to oblanceolate, nearly glabrous adaxially and abaxially, with minute appressed hairs 0.2-0.3 mm long, mostly along the margin; base attenuate and often oblique; margin slightly revolute; apex long-attenuate; petioles to 3 cm on the largest leaves, mostly less than 8 mm in the flowering portion. Inflorescences axillary to the major leaf, evidently 6-9-flowered from the prominent corky pedicel scars, only 1-4 fruits forming per node; pedicels ca. 9 mm in flower, 19-23 mm in fruit, conspicuously fibbed and winged, erect, 1 mm in diameter at base, gradually ampliate distally to 2.2 m m in diameter, minutely puberulent. Calyx cyathiform, 5-5.5 mm long, the margin truncate to undulate, with 5 incurved appendages 2-2.5 mm long in flower and fruit, these somewhat flattened laterally, glabrous; corolla ca. 0.5 cm in diameter, ca. 6 mm long, broadly campanulate to subrotate, yellow with darker green spots within, deeply 5-lobed, the tube ca. 3 mm, the lobes 3.5 • 1.5 mm, deltate, acute at apices, glabrous except for the papillose infolded margin; stamens included; fila- [VOL. 58 ments ca. 1 mm, inserted low within the calyx tube and apparently without two flaps of tissue on corolla tube above site of insertion; anthers ca. 1.8 x 1.5 mm, ovate, the color unknown, longitudinally dehiscent; gynoecium structure not known with certainty. Fruit a globose berry, ca. 1 cm in diameter, erect, glabrous, bright red, juicy; seeds unknown. Distribution and ecology.--Known only from a few collections in the cloud forests of the Provinces of Nor Yungas and Sud Yungas in the Department of La Paz, Bolivia at 1700-2300 m elevation. Phenology.--Flowering in March, May, and November, fruiting in March, May, and November. E t y m o l o g y . ~ T h e specific epithet refers to the horn-like protuberances on the calyx. Additional specimens examined. BOLIVIA, LA PAz. Prov. Nor Yungas: 4.6 km below Yolosa, then 19. l on road up the Rfo Huarinilla, 16~ 67~ 1700 m, 12 Nov 1982 (fl, fr), Solomon 8844 (MO); 14.3 km SW (above) Yolosaon road to Chuspipata, 16~14'S, 67~ 2000 m, 23 Mar 1984 (fr), Solomon et aL 12086 (MO). Prov. Sud Yungas: Huancan6 6.5 km hacia el Sud, 2280 m, 8 Mar 1980 (fl), Beck 3051 (LPB), same locality, 6~ 67~ 2040 m, 27 May 2001 (fl, fr), Beck et al. 28089 (LPB). Capsicum ceratocalyx is characterized by its conspicuously fibbed and winged pedicels. Unfortunately the stamens and pistil of the few flowers available on the type material seem to be somewhat malformed although the plant was evidently producing fruits. One of the collections (Beck 3051) had been determined by A. Hunziker as C. coccineum (Rusby) Hunz., but this species grows at much lower altitudes in Bolivia and has a scrambling, viny habit. Capsicum coccineum also differs from C. ceratocalyx in its smaller flowers that are uniformly yellow and in its unwinged pedicels. Solanum L. In describing these new species we have used the major clades defined by Bohs (2005), while also including the sectional designations used by both Whalen (1984) and Nee (1999) and other groups in use in recent monographic studies. As phylogenetic studies in Solanum become more species-rich 2006] N E E E T AL." S O L A N U M A N D C A P S I C U M ( S O L A N A C E A E ) 327 FIG. 2. Capsicum ceratocalyx M. Nee. A. Leafy branch. B. Pedicel scars in leaf axils. C. Young bud. D. Partially open bud. E. Anthers, drawn from bud. F. Fruiting calyx. (Based on Nee et al. 51778, NY.) 328 BRITTONIA and robust, these now informal groupings will ultimately be given formal infrageneric names. T H E P O T A T O CLADE Solarium complectens M. Nee & G.J. Anderson, sp. nov. TYPE- Bolivia. Santa Cruz. Prov. Vallegrande: 1 km by road S of Chujllas, 18~ 64~ 2125 m, 26 Dec 1989 (fr), M. Nee 38445 (HOLOTYPE: LPB; ISOTYPES: CONN, CORD, G, MO, NY, US, USZ, UT). (Fig. 3) Herbae scandentes, fruticantes, radiculis adventitiis; rami floriferi herbacei, pubescentes, veteres ad 4 m m diametro, sulcati. Folia plerumque imparipinnata, 1-4juga, raro simplicia; foliola elliptica vel ovata, lateralia 1.8-3.5 x 0.8-1.6 cm, terrninalia ovalia, 2.8-4.1 • 1-2.5 cm; pedicelli 1-4 in apice ramulorum floriferorum; calyx dentibus 5, 2.5 x 0.8 ram; corolla stellata, lobis 4.5 x 1.5 ram; stamina 5; filamenta 0.3 m m longa, pubescentia; antherae oblongae, 1.7 x 0.8 ram; stylus 4.5 m m longus, ad basin pubescens. Bacca globosa, ca. 1 cm diametro; semina compressa, integumento pilifero. Herbaceous vines to several meters; lower stems soft-woody, to at least 4 mm in diameter and sulcate, mostly tightly appressed to tree trunks by clusters of short, fasciculate adventitious roots at the nodes, pubescent when young with weak, simple 3-4-celled hairs 0.4-0.9 mm long, glabrescent when older, unarmed. Sympodial units 3-4- to plurifoliate, not geminate. Leaves 4-12 x 1.5-5 cm, mostly imparipinnate, more or less uniform in size and shape, generally with (3-) 7 (-9) leaflets or rarely simple grading into 3foliolate leaves on some branches, the leaflets opposite to subopposite, membranous to slightly fleshy, sparsely pilose adaxially with 2-4-celled simple hairs ca. 0.8 mm long, the basal cell much the largest, pubescent abaxially with hairs like those of the stem; lateral leaflets 1.8-3.5 x 0.8-1.6 cm, narrowly elliptic to ovate, the bases unequal and obtuse to truncate, the margins revolute, the apices obtuse to acute, the petiolules 12.5 mm long; terminal leaflet 2.8-4.1 • 12.5 cm, always larger than the laterals and usually more broadly ovate, the base acute, the margin revolute, the apex obtuse to acute, the petiolule 9 mm long; petioles 1-4 cm, sparsely pubescent; pseudostipules 3.5-8 ram, foliaceous, cordate. Inflorescence (0-) [VOL. 58 0.5-2 cm, borne on small axillary somewhat leafy short-shoots with bract-like leaves 1-2 mm long along the axis, unbranched, with 14 flowers, all flowers perfect, the axes pubescent to nearly glabrous; peduncle 0.4-2 cm; rachis 0-0.2 cm; pedicels 13-18 mm in flower, to 19 mm in fruit, spaced 0-3 mm apart, articulated at the base. Calyx ca. 3.5 mm long, the tube 1 mm long, the lobes 2.5 x 0.8 mm, lanceolate, acute at apices, pilose; fruiting calyx not accrescent, the lobes 2.5 x 0.8 mm; corolla ca. 1 cm in diameter, 5-7 mm long, stellate, white or white with blue, the tube 1-2 ram, the lobes 4.5 x 1.5 ram, Ianceolate, acute at apices, glabrous abaxially at base, minutely puberulent at tips, glabrous adaxially; filaments pubescent, the free portion ca. 0.3 ram, the filament tube absent; anthers 1.7 x 0.8 ram, oblong, connivent, yellow, the pores broad, directed introrsely, often opening into longitudinal slits with age; ovary glabrous; style ca. 4.5 x 0.20.4 mm, cylindrical, straight, pubescent in the lower half; stigma clavate. Fruit a globose berry, ca. 1 cm in diameter, orange or red, glabrous. Seeds ca. 15 per fruit, ca. 2.5 x 2 ram, flattened, light brown, the entire surface covered by hairlike extensions of the epidermal walls. Distribution and ecology.--Known only from Bolivia in the Departments of Santa Cruz and southeastermost La Paz, in cloud forests along the eastern Andes, with Podocarpus parlatorei Pilg., Prumnopitys exigua De Laub. & Silba (Podocarpaceae), Ceroxylon parvum G. Galeano (Arecaceae), Ternstroemia asymmetrica Rusby (Theaceae), Weinmannia spp. (Cunoniaceae), Blepharocalyx salicifolius O. Berg (Myrtaceae), and the tree fern Dicksonia sellowiana (Presl) Hook. (Dicksoniaceae), from 1800 to 3330 m. Phenology.--Flowering in January, May, and June and fruiting in January, June, May, and December. Etymology.--The specific epithet refers to the habit of embracing or holding fast (Latin complector) to the supporting tree by means of its adventitious roots. Additional specimens examined. B O L I V I A . LA Paz. Prov. Inquisivi: comunidad Choquetanga-Cuchiwasi, bajando Pabellonani a 7 k m NE de Choquetanga, 14~ 67~ 3330 m, 19 Jan 1994 (fl, fr), Salinas 2243 (NY). SANTA CRUZ. Prov. Caballero: 26 k m de Co- 2006] N E E ET AL.: S O L A N U M A N D C A P S I C U M ( S O L A N A C E A E ) 329 FIG. 3. Solanum complectens M. Nee & G.J. Anderson. A. Flowering branch. B. Fruiting branch with detail of leaf pubescence. C. Inflorescence detail. D. Opening bud. E. Flower in longitudinal section. E Flower at anthesis. G. Anthers. (A, C-G based on Nee & Mendoza 52538, NY; B based on Nee 38445, NY.) 330 BRITTONIA marapa, carretera a Cochabamba, 17~ 10"S, 64~ 2598 m, 13 Apr 2003 (fr), Calzadilla et al. 81 (NY); entre 15 y 25 km N de San Juan del Potrero hacia Cerro Bravo, 17~ 64~ 2000-2500 m, 6 Jun 1992 (fl, fr), Killeen & Vargas 4060 (NY); hwy from Epizana to Comarapa, 13 km (by road) E of E1 Churo, 0.4 km W of turnoff to Khara Huasi, 17~ 64~ 2575 m, 24 May 2001 (fr), Nee et al. 51858 (NY); 6 km (by air) N of Comarapa, rd to Cerro Bravo and Tinqui Laguna, 17~ 64~ 2325 m, 3 Aug 2003 (fl), Nee et al. 52447 (NY); hwy Comarapa to Cochabamba, 7.3 km (by road) and 22 km (by road) NW of bridge at Comarapa, 17~ 64~ 2640 m, 6 Aug 2003 (fl), Nee & Mendoza 52538 (LPB, NY, USZ); 50 km N de Mataral (en la carretera Santa Cruz-Comarapa), pasando por San Juan del Potrero y bajando a la cuenca del alto Rfo Ichilo, 2000 m, 25-26 May 1989 (fr), Smith et al. 13385 (MO, NY); Parque Nacional Ambor6, Comarapa, 5-8 km al N pot el Rio Arriba hacia Verdecillo, 17~ 64~ 2300 m, 10 May 1993 (fl, fr), Vargas et al. 2400 (CONN, NY); Parque Nacional Ambor6, Cerro Bravo a 10 km al N de Comarapa, 17~ 64~ 2400-2500 m, 12-16 Nov 1995 (fl, fr), Vargas et al. 4167 (NY); Siberia, 25 km desde Comarapa por la carretera Comarapa-Cochabamba, 17~ 64~ 2550 m, 4~i Nov 2003 (fl), Vargas & Jorddn 7015 (NY). Prov. Florida: Quebrada E1 Durazno, 7 km NE of Mairana, 18~ 63~ 2100 m, 22 Jul 1994 (ster), Nee 45330 (NY); Barrientos, 8 km N de Paredones (Achira Camping), 18~ 63~ 18001900 m, 25 Jun 1996 (fl, fr), Vargas & Soliz 4532 (NY). S o l a n u m c o m p l e c t e n s is a m e m b e r o f S o l a n u m section A n a r r h i c h o m e n u m Bitter, a small and distinctive group o f h e r b a c e o u s scramblers distributed from M e x i c o to Peru. The affinities o f this section are with the potatoes, tomatoes, and relatives (Bohs, 2005). It is significant that neither this species nor any other in sect. A n a r r h i c h o m e n u m has b e e n found in B o l i v i a previously, e s p e c i a l l y not in the central and northern parts o f the Department o f L a Paz w h o s e cloud forests have been the m o s t e x p l o r e d part o f the entire country. This indicates a strong d i s j u c t i o n o f this species f r o m its relatives in northern Peru to M e x i c o (Correll 1962; A n d e r s o n & Jansen, 1998). The o n l y s p e c i m e n o f S o l a n u m c o m p l e c t e n s with s i m p l e or 3-foliolate leaves is from the furthest northwest ( S a l i n a s 2 2 4 3 ) ; in other respects it r e s e m b l e s the rest o f the material available. In Correll (1962), S. c o m p l e c t e n s w o u l d key best to S. c h i m b o r a z e n s e Bitter, k n o w n only from Ecuador, but that species bears green fruits ca. 2 c m in d i a m e ter, has leaves with only three, or less c o m m o n l y five, leaflets, and the filaments are united into a crown. [VOL. 58 THE GEMINATA CLADE Since the p u b l i c a t i o n o f K n a p p (2002b), new species o f section G e m i n a t a s.1. continue to be c o l l e c t e d throughout the neotropics. These forest plants often have narrow, end e m i c distributions and due to their inconspicuous nature (sparse p o p u l a t i o n s o f plants with small white or green flowers and green fruits) are also relatively undercollected. S o l a n u m c h a l m e r s i i S. Knapp, sp. nov. TYPE: Bolivia. L a Paz. Prov. Sud Yungas: 7,5 k m (by road) from Huancan6 on r o a d to San Isidro, m o i s t m o n t a n e forest, 16~ 67~ 2225 m, 10 M a y 2001 (fl, fr), M . N e e , L. B o h s , S. K n a p p & M . M e n d o z a F. 5 1 7 7 7 (HOLOTYPE: L P B ; ~SOTYPES: B M , NY, USZ). (Fig. 4) Species Solano acuminato Ruiz & Pavon similis, sed pubescentia pallida flavovirenti densa, foliis in stato sicco pallide viridibus, gemmis maturis obovoideis, differt. Shrubs or s m a l l trees 2 - 6 m tall; y o u n g stems d e n s e l y white p u b e s c e n t with simple, uniseriate t r i c h o m e s ca. 1 m m long c o m p o s e d o f 2 - 5 cells; o l d e r stems r e m a i n i n g densely white pubescent, occasionally glabrate, u n a r m e d . S y m p o d i a l units unifoliate or difoliate and geminate. L e a v e s 1.5-17 x 1 - 6 cm, simple, dimorphic, the m a j o r leaves 9 - 1 7 x 3 - 6 cm, elliptic to n a r r o w l y elliptic, the m i n o r leaves 1.5-3 x 1 - 2 cm, differing f r o m m a j o r leaves only in size, but occ a s i o n a l l y s o m e w h a t rounder in outline, thin-textured, d r y i n g pale green, evenly pubescent a d a x i a l l y with simple uniseriate trichomes ca. 1 m m long, d e n s e l y p u b e s c e n t a b a x i a l l y with white uniseriate t r i c h o m e s 1 1.5 m m long, the trichomes d e n s e r on the veins; b a s e acute; margin entire; apex acute, r o u n d e d at the very tip; petioles 0 . 3 - 0 . 7 c m in m a j o r leaves, ca. 0.5 c m in m i n o r leaves, densely pubescent. Inflorescences 1.5-5 cm, o p p o s i t e the leaves or o c c a s i o n a l l y s o m e w h a t internodal, unbranched, with 1 0 - 2 0 flowers, all flowers a p p a r e n t l y perfect, the axes densely w h i t e - p u b e s c e n t with s i m p l e uniseriate t r i c h o m e s 0 . 5 - 1 . 5 ram; p e d u n c l e 1-3 cm; rachis 1 4 . 5 cm; p e d i c e l s 1 0 - 1 2 m m in flower, t a p e r i n g f r o m the abrupt base o f the calyx tube to a slender base 0 . 5 - 0 . 8 m m in diameter, deflexed, 1 5 - 2 2 m m in fruit, 0 . 5 - 1 2006] N E E ET AL.: S O L A N U M A N D C A P S I C U M ( S O L A N A C E A E ) 331 FIG. 4. Solanum chalmersii S. Knapp. A. Flowering branch, with detail of leaf pubescence. B. Inflorescence, with detail of a single simple, uniseriate trichome. C. Opening bud. D. Flower showing petals at anthesis. E. Anthers. F. Infructescence. (Based on Nee et al. 51777, NY.) 332 BRITTONIA mm in diameter at the base, pendent, woody, closely packed, often overlapping, articulated at the base. Buds when very young appearing globose, the corolla soon exerted from the calyx lobes, the buds later becoming obovoid just before anthesis. Calyx 2-2.5 mm long, the tube 1-1.5 mm, the lobes ca. 1 x 1 ram, deltate to broadly triangular, abruptly constricted to an elongate tip ca. 0.5 m m long, densely pubescent with simple, uniseriate trichomes like those of the rest of the inflorescence; fruiting calyx not accrescent, the lobes ca. 1 x 1 ram, brittle and somewhat patent; corolla 1.5-2 cm in diameter, ca. 10 mm long, stellate, white or tinged purplish in some plants, the tube ca. 1 mm, the lobes ca. 1 x 0.5 mm at base, ovate-lanceolate, reflexed at anthesis, acute at apices, densely and evenly pubescent abaxially with simple uniseriate trichomes ca. 0.5 m m long, glabrous adaxially except for the densely papillose margins, the tips of the lobes densely papillose and somewhat cucullate; filaments glabrous, the free portion 0.8-1 ram, the filament tube 1-2 ram, with small teeth arising from the filament tube between the anthers; anthers 4-5 x 1-1.5 turn, oblong, slightly sagittate at base, connivent, yellow, the pores tear-drop shaped, opening into longitudinal slits with age; ovary glabrous; style 6-7 x ca. 0.05 mm, cylindrical, straight, glabrous; stigma capitate. Fruit a globose berry, 1-1.2 cm in diameter, green, glabrous. Seeds numerous (more than 15) per fruit, 3-3.5 x 2 2.5 ram, flattened-reniform, pale yellow in dry material, the surfaces minutely pitted, the margin incrassate and darker yellow. Distribution and e c o l o g y - - I n the understory of montane forest in northern Bolivia, on eastern Andean slopes from 1900-2200 m. Plants of Solanum chalmersii grow both in the forest understory and in disturbed areas along roads and streams, attaining higher population densities in open areas. P h e n o l o g y - - M o s t flowering specimens have been collected in May, but if Solanum chalmersii is like other members of the S. nudum species group, it will flower and fruit year-round, but with pulses at particular seasons (see Knapp, 2002b). E t y m o l o g y - - T h i s species is named in honour of Sir Neil Chalmers, Warden of Wadham College, Oxford and previously Director of [VOL. 58 the Natural History Museum in London. His support of taxonomy made the third author's collecting in Bolivia possible. Additional specimens examined. BOLIVIA. LA PAZ. Prov. Sud Yungas: de Chulumani hacia el N unos 5 km hacia Irupana, entrando hacia Apa Apa, 16~ 67~ 2050 m, 19 Sep 1998, Beck 24480 (NY); Sirupaya vic. de Yanacachi, 2100 m, 16 Nov 1906, Buchtien 315 (NY); along road through primary cloud forest, 7.09.4 km NE of (above) Huancant, 16~ 67~ 1718 May 1990, Luteyn & Dorr 13722 (BM, LPB, NY); 5.2 krn (by road) from Huancan6 on road to San lsidro, moist montane forest, 16~ 67~ 2225 m, 10 May 2001. Nee et al. 51773 (NY, BM, LPB, USZ); road between Unduavi and Puente Villa, around hotel Castillo del Chaco and down to the Rfo Unduavi, humid forest on steep slopes, 16~ 67~ 1900-1930 m, 11 May 2001, Nee et al. 51795 (BM, NY, LPB, USZ); Yanacachi, 3.5 km hacia Chojlla (as "Chajlla"), 4 Apr 1987, Seidel & Richter 851 (LPB, NY); Cant6n Yanacachi, Mina Chollja, camino de acceso de vehfculos a Kacapi, 15~ 68~ 2182 m, 25 Jul 2000, Sihani 261 (LPB); ca. 6 km along road from Huancan6 to San Isidro, 16~ 67~ 1 July 1995, Wood 9955 (K). Solanum chalmersii was mentioned as a probable new species in Knapp (2002b), but at that time material was not sufficient to distinguish it from similar species in the large Solanum nudum species group and good fruiting material was not available in order to place it confidently in that group. Solanum chalmersii possesses the flattened-reniform, pale colored seeds, simple uniseriate trichomes, and more or less closely spaced pedicel scars typical of the S. nudum species group. It is most similar to S. a c u m i n a t u m Ruiz & Pay. of central Peru to Bolivia, with which it is nearly sympatric, but differs in its uniform covering of simple uniseriate trichomes (usually confined to the vein axils in S. acuminatum), its flowers with longer anthers with slightly sagittate bases, and in its longer fruiting pedicels. In Bolivia, S. acuminatum occurs at higher elevations than S. chalmersii. S. Knapp, sp. nov. TreE: Bolivia. Santa Cruz. Prov. Caballero: Parque Nacional Ambor6, Cerro Bravo, pr6xima a l a s juntas del Rfo Alizar y Amparo (20 k m a l NW de San Juan de Potrero), 17~ 64~ 2000 m, 1014 Apr 1994 (fl), I. G. Vargas C., D. Ayala & J. Quiroga 3138 (HOLOTYeE: LPB; 1SOTYPES: BM, NY, USZ). (Fig. 5) Solanum monanthemon 2006] N E E ET A L . : S O L A N U M A N D CAPSICUM ( S O L A N A C E A E ) Species Solano symmetrico Rusby similis, sed foliis axillafibus sparse pubescentibus, inflorescentiis unifioris, peclicellis longis differt. Shrubs to 1.5 m tall; young stems glabrous and somewhat shiny; older stems glabrous, unarmed, Sympodial units difoliate and geminate, the minor leaf often deciduous and so appearing unifoliate. Leaves 2-11 x 1-4 era, simple, dimorphic, the major leaves 7-11 x 2 - 4 cm, ovate to elliptic or narrowly elliptic, the minor leaves 2-5 x 1-2.7 cm, differing from the major ones only in size, thintextured, glabrous and shiny adaxially, glabrous abaxially but with tufts of white simple uniseriate trichomes in the axils of the veins, the trichomes ca. 0.5 ram, very thin and interlaced, arising from the lamina; base acute to slightly attenuate onto the petiole; margin entire; apex acute to long-acuminate and rounded at the very tip; petioles 0.5-1 cm in major leaves, 0.5-0.7 cm in minor leaves, glabrous. Inflorescence consisting of a single flower (occasionally the scar of a second flower apparently present), opposite the leaves or occasionally somewhat internodal, all flowers perfect, the axes glabrous; peduncle and rachis absent or the pedicel appearing jointed 0.6--0.9 cm from the base; pedicels 30-35 m m in flower (to 40 mm if measured to the base of the inflorescence), deflexed, tapering from the base of the calyx tube to a slender base ca. 0.2 m m in diameter, 45-50 m m in fruit, 0.5-1 m m in diameter at the base, ca. 3 m m in diameter at the apex, pendent, slender. Buds when very young appearing globose, the corolla included within the elongate calyx lobes, the buds remaining globose until anthesis. Calyx 4-6.5 m m long, the tube 1-1.5 ram, the lobes 3-5 x ca. 0.5 mm, long-triangular, glabrous; fruiting calyx slightly accrescent, the lobes 5 - 6 x ca. 0.5 ram; corolla 0.8-1 c m in diameter, ca. 5 m m long, stellate, white, the tube minute, less than 0.5 m m long, the lobes ca. 0.5 x 0.3 m m at base, ovate-lanceolate, probably reflexed at anthesis, glabrous abaxially and adaxially except for the minutely papillose tips; filaments glabrous, the free portion ca. 0.5 mm, the filament tube ca. 0.5 ram; anthers 2.5-3 x ca. 1 mm, oblong, slightly sagittate at base, connivent, yellow, the pores tear-drop shaped, opening into longitudinal slits with age; ovary glabrous; style 3 - 4 x ca. 0.1 mm, 333 cylindrical, straight, glabrous; stigma capitate. Fruit a globose berry, 1-1.2 cm in diameter (immature), green, glabrous. Seeds not known from mature fruit. Distribution and ecology.--Only known from two specimens collected in cloud forests in Parque Nacional Ambor6 and Parque Nacional Carrasco on the eastern slopes of the Andes in Bolivia at ca. 2000 m. The cloud forests where S. monathemon grows have Podocarpaceae and Myrtaceae as canopy trees (fide Vargas et al. 3138). Phenology.--Due to the paucity of specimens, data as to flowering and fruiting are not available. Etymology.--From the Greek for single (mono-) flower (anthemon), referring to the unusual single-flowered inflorescences of this species. Additional specimen examined. BOLIVIA. Sehuencas, Rio Fuerte cerca Puente, Parque Nacional Carrasco, 2150 m, 17~ 65~ 30 Nov 1993 fir), P. lbisch & C. Ibisch 93.1477 (LPB). COCI-IA][iAMBA. Prov. Carrasco: Specimens of Solanum monanthemon have occasionally been identified as the more common Bolivian species S. trichoneuron Lillo. It is superficially similar to that species, but differs in its dark bark and single-flowered inflorescences, which look like a single flower on an articulate pedicel arising opposite the geminate leaves. The apparent articulation is actually the joint between the inflorescence axis and the pedicel, as evidenced by some inflorescences with minute second buds that apparently abort early in inflorescence development. Many more collections of S. monanthemon are needed to investigate this phenomenon fully. Mature fruits of S. monanthereon are not known, so placement in one of the species groups of Knapp (2002b) is tentative at present. Overall morphology (i.e., tufts of hairs in the vein axils, short anthers) suggests the S. nudum species group, but this can be misleading (e.g., S. smithii S. Knapp of the S. arenarium species group also possesses these hair tufts; see Knapp, 2002b). Solarium monanthemon can be distinguished from other species with tufts of hairs in the abaxial vein axils by its singleflowered inftorescences and its tree-like habit. It also apparently grows as isolated in- 334 BRITTONIA [VOL. 58 FIG. 5. Solanum monanthemon S. Knapp. A, Fruiting branch. B. Partially open bud, showing the distinctive long-triangular calyx lobes. C. Flower at anthesis. D. Flower in longitudinal section. E. Anthers. (Based on Vargas et al. 3138, NY.) 2006] dividuals in o f the other group that streams and N E E ET AL.; S O L A N U M A N D CAPSICUM ( S O L A N A C E A E ) dense cloud forests, unlike m a n y m e m b e r s o f the S. n u d u m species are plants o f open areas along roads. THE WENDLANDII/ALLOPHYLLUM CLADE S o l a n u m e l a n d e s t i n u m Bohs, sp. nov. TYPE: Bolivia. L a Paz. Prov. N o r Yungas: 2 k m b y road (ca. 1 k m b y air) N E and b e l o w Chuspipata, 16~ 67~ 2950 m, 29 Oct 1984 (fl, fr), M . N e e & J. Solomon 30219 (HOLOTYPE: LPB; ISOTYPE: MO). (Fig. 6) Habitu, foliis et fructibus S. confuso C.V. Morton similis sed pedunculis brevissimis, antheris obtnsis oblongis et seminibus rotundatis differt. Slender shrub 1-2.5 m tall; stems glabrous to d e n s e l y p u b e s c e n t with u n b r a n c h e d hairs, s o m e with b r o a d multiseriate bases giving the y o u n g stems a c o r k y or r o u g h e n e d appearance, unarmed. S y m p o d i a l units difoliate, geminate. L e a v e s 3 - 1 2 x 0 . 5 - 3 cm, simple, dimorphic, the m a j o r leaves 5 - 1 2 x 1-3 cm, narrowly elliptic to lanceolate, the m i n o r leaves 3 - 4 x 0 . 5 - 1 cm, differing f r o m the m a j o r ones o n l y in size, usually subcoriaceous, glabrous adaxially, glabrous to m o d e r ately p u b e s c e n t a b a x i a l l y with u n b r a n c h e d hairs; base cuneate to long-tapered; m a r g i n entire, glabrous to ciliate; apex acute to acuminate; petioles 0 . 5 - 2 cm, 1 - 2 c m in m a j o r leaves, ca. 0.5 c m in m i n o r leaves, glabrous to m o d e r a t e l y p u b e s c e n t in a d a x i a l channel. Inflorescence 0 . 5 - 2 cm, opposite the leaves, u n b r a n c h e d or r a r e l y forked, with 1 - 6 flowers, all flowers perfect, the axes glabrous to sparsely pubescent; p e d u n c l e 0 . 2 - 2 cm; rachis very short, 1 - 4 m m ; pedicels 1 0 - 2 0 m m in flower, 1 5 - 3 5 m m and slender and curved in fruit, c o n g e s t e d and spaced up to 3 m m apart, articulated at the base. C a l y x 2 - 3 m m long, the tube ca. 1 m m , the lobes 1 - 2 x 1 . 5 - 2 mm, b r o a d l y deltate, almost truncate with a small a c u m e n at tip, glabrous at base, t o m e n t o s e at tips; fruiting c a l y x not accrescent, the lobes 1 - 2 x 1 . 5 - 2 m m ; corolla 1 1.5 c m in diameter, 4 - 7 m m long, stellate, white, the tube 1 - 2 m m , the lobes 3 - 5 x 2 2.5 m m at base, n a r r o w l y triangular to ovate, acute at apices, g l a b r o u s a b a x i a l l y at base, 335 t o m e n t o s e at tips o f lobes, glabrous adaxially; filaments glabrous, the free portion 0 . 5 1 m m , the filament tube absent to 0.5 m m ; anthers 3 - 4 • 1.5-2 m m , oblong, connivent, yellow, the pores broad, directed distally and adaxially, often o p e n i n g into longitudinal slits with age; ovary glabrous; style 4 - 5 • ca. 0.5 mm, cylindrical, straight; glabrous; s t i g m a truncate. Fruit a g l o b o s e berry, 1-1.5 c m in diameter, o r a n g e w h e n ripe, glabrous. Seeds fewer than 15 p e r fruit, ca. 4 • 3 m m , s o m e w h a t flattened, y e l l o w - b r o w n , the surfaces rugulose. Distribution and ecology.--Montane rain and cloud forest, e s p e c i a l l y on slopes or in disturbed areas, o f western B o l i v i a in Yungas de L a Paz, 2 2 0 0 - 3 1 0 0 m. Phenology.--Flowering specimens have b e e n collected in May, June, and A u g u s t through October. F r u i t i n g s p e c i m e n s have b e e n collected in M a r c h , May, June, August, and October. E t y m o l o g y . - - T h e specific epithet, m e a n i n g "hidden," refers to the i n c o n s p i c u o u s appearance o f this species, w h i c h has p r o b a b l y l e d to it being o v e r l o o k e d until now. On a recent B o l i v i a n field trip, three S o l a n u m specialists ate lunch next to the plants without noticing t h e m at first, despite the botanists' interest in collecting this species. W h e n one plant was finally recognized, several others were f o u n d in close proximity. Additional specimens examined: BOLIVIA. LA PAZ. Prov. Nor Yungas: Guanai, 1891 (fr), Bang s.n. (NY); Yolosa, 23 km hacia Chuspipata, 2730 m, 16 Sept 1981 (fl), Beck 4844 (NY); road from Coroico to divide leading to La Paz, 21 May 1980 (fl, fr), D'Arcy & Bejarano 13875 (NY); road from Chuspipata to Yolosa, ca. 68~ 16~ 2000-2700 m, km 20-13, 4/11/1989 (fr), J. E Smith & Beck 1731 (LPB, NY); 2.2 km NE (below) Chuspipata, 16~ 67~ 3000 m, 24 Mar 1982 (fr), Solomon 7296 (MO, NY); 1.6 km NW of Chuspipata, 16~ 67~ 3100 m, 26 Aug 1983 (fl, fr), Solomon 10668 (MO, NY); 0.8 km SE of (below) Chuspipata on road to Chulumani, vic. Chuspipata railroad station, 16~ 67~ 3100 m, 7 Oct 1984 (fl), Solomon & Escobar 12507 (MO); 1.2 km W of Chuspipata, 16~ 67~ 3100 m, 8 June 1985 (fr), Solomon 13851 (MO, NY); 1.2 km E of Cotapata on road between Unduavi and Chuspipata, 16~ 67~ 3100 m, 26 June 1986 (fl), Solomon 15316 (MO), (fr), Solomon 15320 (MO); Saltos de San Juan, 10 km al NE (debajo) de Chuspipata por el camino a Yolosa, 16~ 67~ 2400 m, 28 May 1988 fir), Solomon 18499 (MO). Prov. Sud Yungas: Huancan6, 7 krn hacia el sur, 2400 m, 3/7/1981 (fl, fr), Beck 4713 (NY); 7.5 km (by road) from Huancan6 on road to San 336 BRITTONIA [ V O L . 58 FIG. 6. Solanum clandestinum Bohs. A, B. Flowering branches. C. Inflorescence. D. Bud. E. Flower at anthesis. F. Flower in longitudinal section. G. Anthers. H. Fruiting branch. (A based on Solomon 10668, NY; B-C based on Bohs photos of Nee et al. 51781; D-G based on Beck 4844, NY; H based on Nee et al. 51781, NY.) Isidro, 16~ 67~ 2225 m, 10 May 2001 fir), Nee et al. 51781 (LPB, NY, UT); 9 km de Huancan6 en la carretera hacia San Isidro, 16~ 67~ 2400 m, 2 May 1989 (fr), D.N. Smith & J . E Smith 13061 (LPB, MO, NY). S o l a n u m c l a n d e s t i n u m is u n i q u e in its narr o w l y e l l i p t i c to l a n c e o l a t e and u s u a l l y subcoriaceous leaves, inflorescences with very short p e d u n c l e s and l o n g p e d i c e l s , w h i t e 2006] NEE ET AL.: SOLANUM AND CAPSICUM (SOLANACEAE) corollas, oblong anthers, and conspicuous, orange, few-seeded fruits. It is apparently restricted to the Yungas of La Paz in northwestern Bolivia. Specimens can vary from being nearly glabrous to abundantly pubescent, especially on the young stems. The hairs are unbranched but can have wide multiseriate bases and in some specimens these give the twigs a roughened or corky aspect. Fruiting specimens of Solanum clandestinum can resemble S. confusum C.V. Morton, another species of the Bolivian uplands. Solanum clandestinum is distinctive in its very short inflorescences, with the pedicels clustered and becoming long, slender, and curved in fruit. The seeds of S. clandestinum are not angled like those of S. confusum, and they always lack white pseudohairs on the surface. Flowers of S. clandestinum have white corollas and blunt, oblong anthers, whereas S. confusum flowers usually have violet corollas and tapered anthers. Solanum confusum lacks hairs with multiseriate bases. Parsimony analyses of chloroplast ndhF sequence data place Solanum clandestinum in a clade with S. mapiriense Bitter, S. wendtandii Hook.f, and S. allophyllum (Miers) Standl. (the Wendlandii/Allophyllum clade of Bohs, 2005), but this clade is not well-supported (Bohs, unpubl, data). Nuclear ITS sequences indicate a relationship among S. clandestinum, S. mapiriense, and S. morellifolium Bohs, but this clade receives very little bootstrap support. Data from the nuclear waxy (GBSSI) gene allies S. clandestinum with S. mapiriense with 100% bootstrap support. Thus, three genes suggest an alliance between S. clandestinum and S. mapiriense, the latter another species endemic to the Bolivian Yungas that has been placed in Solanum section Allophyllure (Bohs, 1990). The elliptic leaves of S. mapiriense are much broader than those of S. clandestinum and the anthers are strongly tapered distally, in contrast to the oblong anthers of S. clandestinum. The fruits and seeds of S. mapiriense are incompletely known. T H E LEPTOSTEMONUM CLADE Typification of Solanum aridum Morong and new synonymy- In the arid inter-Andean valleys of southern Bolivia and adjacent Chaco plains to the east grows a colonial 337 herbaceous species of Solanum. It is usually found in unshaded areas, often around temporary puddles or roadsides, on sand or more often on clay. It is also present in the Chaco of western Paraguay and adjacent areas of northwestern Argentina. The correct name for this species has long been in doubt, in part due to the paucity of collections and to the similarity of closely related species in the group. Solanum aridum Morong is now established as the name of this species (annotated by Nee as S. conditum C.V. Morton in herbaria); the lectotype and most of the isotypes are mixed collections, which has led to some confusion. S o l a n u m a r i d u m Morong, Ann. New York Acad. Sci. 7: 173. 1893. TYPE: Paraguay. Presidente Hayes (?): falls of the Rio Pilcomayo, 2 May 1888-1890, T. Morong 1007 (LECTOTYPE: US 48029, nonglandular, nearly unarmed plant with cuneate leaf bases, designated by Morton, 1976; ISOLECTOTYPES: BM 87516 leafy twig, E two twigs, K, M O 2766698, NDGn.y., NY 139054 leafy twigs, P H - 2 sheets, WIS ex WELC 15091 except for plant of Turnera). (Fig. 7) Solanum elaeagnifolium Cav. vat. ovalifolium Kuntze, Revis. Gen. P1. 3, 3: 225. 1898. TYPE: Argentina. Salta: Monte Morro, Nov 1892 (fl, fr), O. Kuntze s.n. (HOLOTYPE" NY 139143; ISOTYPE"US 701686). S. conditum C.V. Morton, Revis. Argent. Solanum 237. 1976. TYPE: Argentina. Santiago de1 Estero: Depto. C. Pellegrini, Estancia E1 Remate, 500 m, 22 Dec 1927 (fl), S. Venturi 5685 (HOLOTYPE: US 1548974; ISOTYPES: NY, SI-2 sheets). Solanum aridum is a member of a group of closely related taxa designated as the Solanum multispinum group by Whalen (1984) and as Solanum subgen. Leptostemonum sect. Melongena subsect. Lathyrocarpum (Dunal) G. Don by Nee (1999). The name Solanum aridum Morong was based on Morong 1007, collected along the Rfo Pilcomayo on the border of Paraguay and Argentina; it was considered by Morton (1976) to be a species perhaps not subsequently collected and certainly not known to him, although, in fact, in the same work he described the same species as new under the name S. conditum C.V. Morton. It appears under this latter name in Nee (1999). 338 BRITTONIA [VOL. 58 Fla. 7. Solanum aridum Morong. Isolectotype specimen Morong 1007 at NY (NY 139054). The majority of the stems on the sheet are S. aridum, but lowermost stem slightly to the left of center with a large globose fruit and cauline spines is S. multispinum N. E. Br. (see text). 2006] NEE ET AL.; SOLANUM AND CAPSICUM (SOLANACEAE) Thomas Morong's collections from Paraguay and preserved at NY are generally good and with copious notes. Morong 1007, however, was a decidedly mixed collection, with at least three different species in two families represented among the various duplicates. Morong collected the plants in May (the dry season) and said in the original description that the plants "had a parched, dried appearance" (Morong & Britton, 1893). It is thus understandable that Morong would collect specimens of one species in flower and fruit (Solanum aridum) and a related species mostly without leaves and in fruit (Solanum multispinum N.E. Br.), although the addition of the flowering specimen of Turnera (Turneraceae) is less explicable. Solanum aridum was lectotypified by C. V. Morton (1976) on the leafy and flowering part of the duplicate US 48029, although Recommendation 9A.3 of the Code (Greuter et al., 2000) would have recommended lectotypification on the specimen NY 139054, which is labelled "type" (Fig. 7). Morong was curator of the Columbia University Herbarium (Britton, 1894), and this herbarium subsequently became the basis of the newly formed New York Botanical Garden herbarium (NY). Morton claimed that his lectotypification was somewhat "arbitrary," but fortunately it was based on the predominant element of Morong 1007, and that which provides the major portion of the original description. The full range of duplicates with their extraneous elements can be explained as follows. The lectotype sheet of Solanum aridum, US 48029, consists of a leafy and flowering portion of S. aridum, along with a very spiny plant with stipitate-glandular stems and with leaf blades cordate at the base. This latter element corresponds to S. multispinum, parts of which are found on other duplicates as well. The specimens BM 87516 and NY 139054 are a mixture of Solanum aridum, plus leafless twigs of S. multispinum with globose fruits, which accounts for the description of a fruit 3 cm in diameter and of a long fruiting pedicel; S. aridum has an ovoid fruit 1.6-1.8 • 1.3-1.7 cm, and the fruiting pedicel is 1.83.1 cm long, but without the bristly hairs present on S. multispinum. The specimen at E 339 consists of two plants of S. aridum, one with leaves and the other with leaves and fruits; in addition there is a flowering plant of S. multispinum. The specimens at K, MO 2766698, and the two sheets at PH consist exclusively of S. aridum. The specimen WIS ex WELC 15091 has a plant with the small, ovoid fruit of S. aridum along with a piece of Turnera of the Turneraceae, which accounts for the "sulphur yellow" corolla in the original description; S. aridum has a white or pale blue corolla. In the type region only two species of Solanum, S. palinacanthum Dunal and S. multispinum N.E. Br., have a fruit similar to that of the ca. 3 cm diameter fruits on Morong 1007. The fruiting pedicels of S. multispinum exactly match those on the fruiting specimen of Morong 1007, while the pedicels of S. palinacanthum are curved and generally much shorter. During his preparation of a treatment of Solanum of Argentina, Morton (1976) annotated many specimens with a name transferring the epithet ovalifolium of S. elaeagnifolium Cav. var. ovalifolium Kuntze to a variety of S. meloncillo Parodi, but this transfer was never published, and instead these plants were named S. conditum C. V. Morton. S o l a n u m m o x o s e n s e M. N e e , sp. nov. TW,E: Bolivia. Beni. Prov. Cercado: Trinidad, 14~ 64~ 200 m, weed in the city, 6 Jan 1989 (ill M. Nee 37519 (HOLOXVeE: LPB; ISOTVPES: G, MO, NY-2 sheets, USZ). (Fig. 8) Herba repens ad nodos radicans, pilis sirnplicibus 23-cellularibus sparse pilosa, aculeis acicularibus 1.7-3.5 mm longibus sparse armata. Folia alterna, ovata, 5-7.5 • 3.5-6 cm, repanda vel non profunde lobata, subtus sparse stellato-tomentosa. Inflorescentia extra-axillaris, racemoso-cymosa, 4--5-florus; calyx inermis; corolla alba, lobis 6.5 • 3.5 mm; antherae lineari-angustatae, 5 • 1 mm; ovarium glabrum. Bacca oblongo-ovoidea, glabra. Herb or vine, creeping and rooting at the nodes, or some stems erect, to 20 cm tall; stems slender, to ca. 2.5 rnm in diameter, nearly glabrous to sparsely pilose with simple, 2-3-celled hairs, sparsely armed with prickles 1.7-3.5 m m long, slightly reflexed, nearly straight or slightly curved. Sympodial units 2- to 3-foliate, not geminate. Leaves 5 7.5 • 3.5-6 cm, simple, more or less uniform 340 BRITTONIA [VOL. 58 FIG. 8. Solanum moxosense M. Nee. A. Habit. B. Leafy branch with inflorescence and detail of prickles. C. Close-up of leaf base with detail of stellate hairs. D. Partially open flower. E. Anther. F. Fruits. (A-E based on Nee 37519, NY; F based on Nee 34261, NY.) in size and shape, ovate, thin-textured, glabrous adaxially, very sparsely pubescent abaxially with sessile 4-rayed stellae 0.7-1 mm wide, the midpoints shorter than the lateral rays, sparsely armed above and below with straight acicular prickles 1-3.5 mm long on the major veins; base truncate to subcordate, nearly equal to very unequal and with one side offset by up to 1 cm from the other; margin repand to shallowly lobed with 3-4 rounded to obtuse lobes per side; apex obtuse; petioles 0.7-3 cm, with hairs and prickles like those of the stems. Inflorescence 5-6 cm, extra-axillary or opposite the leaves, unbranched, with 4-5 flowers, the plants probably andromonecious and only the lowest one or two flowers fertile, the axes nearly glabrous to sparsely pilose with simple or stellate hairs and with 1 or 2 acicular prickles ca. 2 mm long; peduncle 3-4 cm; rachis ca. 2 2006] NEE ET AL.: SOLANUM AND CAPSICUM (SOLANACEAE) cm; pedicels 9-11 m m in flower, to 15 mm and somewhat thickened in fruit, expanded distally to 1.5 mm in diameter, recurved, widely spaced, 7-14 mm apart, articulated at the base. Calyx 6-7 mm long, the tube 2-3 mm, the lobes in anthesis to 4 x 1.4 mm including the ca. 1 mm long caudate tip, oblong-ovate, with a few sessile stellate hairs, unarmed; fruiting calyx somewhat accrescent, the lobes to ca. 6.5 x 3.2 mm; corolla ca. 1.3 cm in diameter, 12 mm long, stellate to stellate-pentagonal, white, lobed about half way, the tube 6 mm, the lobes ca. 6.5 x 3.5 mm, triangular, tomentose abaxially with small sessile stellae, glabrous adaxially; filaments glabrous, the free portion ca. 2 mm, the filament tube absent; anthers ca. 5 x 1 ram, linear-tapered, not connivent, yellow, the pores minute and directed distally, not opening into longitudinal slits with age; ovary glabrous; style ca. 8 x 0.5 mm, cylindrical, straight, glabrous; stigma capitate. Fruit (immature) an oblong-ovoid berry, ca. 1 cm in diameter, the color when ripe unknown, glabrous. Seeds unknown. Distribution and ecology.--The few collections to date show Solanum moxosense is adapted to weedy or disturbed areas around cattle pens, but the original adaptation is probably to natural disturbances of the seasonally inundated savanna in which Trinidad is situated. So far this species is only known from the immediate vicinity of the city at ca. 200 m in elevation. It is a weed in the city of Trinidad and was also seen commonly on dirt roads and near cattle yards near Puerto A1macrn, 7 km SW of the city, but not yet blooming there, so no specimens were collected. No collections have been seen from the Estaci6n Biol6gica del Beni, ca. 150 km to the west, one of the few areas of the Beni plains that has been thoroughly explored botanically. Phenology.--Flowering in January and August. Etymology.--The name refers to the Llanos de Moxos, one of the largest seasonally flooded savannas in the world. This part of Bolivia is a perfectly flat alluvial plain between the pre-Cambrian shield of crystalline rocks to the east and the Andes to the west, which provide the recent sediments from their numerous rivers flowing northwards to 341 the Madeira and then the Amazon. The region has been identified as a center of diversity and endemism and a priority for conservation (Beck & Moraes-R., 1997). Additional specimens examined. BOLIVIA. BERYL Prov. Cercado: Trinidad, on savanna, 200 m, Aug 1944 (fl), Cdrdenas 3525 (MO, US); Trinidad, 14~ 64~ 150 m, weedy areas in the town, 25 Feb 1987 (fl, fr), Nee 34261 (MO, NY). Solanum moxosense is a much more nearly glabrous plant than related species in the Solanum multispinum species group (sensu Whalen, 1984) or sect. Melongena subsect. Lathyrocarpum G. Don (sensu Nee, 1999), most of which are found further south in southern Bolivia (from about Santa Cruz de la Sierra south) into northern Argentina and especially Paraguay. It appears most similar to species such as S. hieronymi Kuntze from dry areas of southern Bolivia, northwestern Argentina and Paraguay, or S. comptum C.V. Morton of eastern Paraguay and northeastern Argentina, but these species are more robust in all respects, more densely stellatetomentose, and display the very spiny fruiting calyx typical of the group. In habit, Solanum moxosense resembles the Cuban endemic Solanum chamaeacanthum Griseb., but that species appears not to root along the nodes, the leaves are fleshier, smaller (1.5--4 x 1.2-2 cm), with more regular and more acute lobes, and the plant is more densely armed with longer prickles (to 6.5 mm long). Solanum moxosense is also similar to the creeping species S. flagellare Sendtn. and S. reineckii Briquet of coastal southeastern Brazil, but differs again by more profuse rooting at the nodes and a suite of other characters. S o l a n u m p e d e m o n t a n u m M. Nee, sp. nov. TYPE: Ecuador. Napo: Afiangu, Parque Nacional Yasunf, 0~ 76~ 30 May-21 Jun 1982 (fr), B. Ollgaard et al. 39285 (HOLOTYPE:QCA; ISOTYPE:NY). Fig. 9 Liana lignosa, aculeis solum recurvis 1.5-3 mm longis armata. Folia plerumque geminata, altero minore, folia longiora anguste eliptica, 10-18 x 2.5-6.5 cm, integravel angulati-lobata, supra inermia, subtus costa aculeis recurvis armata. Inflorescentiae extra-axillares, in- 342 BRITTONIA [VOL. 58 FIG. 9. Solanum pedemontanum M. Nee. A. Flowering branch with detail of adaxial leaf surface with prickles and hairs. B. Inflorescence and flower. C. Flower at anthesis. D. Anthers. E. Gynoecium. F. Infructescence. G. Dry fruit. (A, B, D - G based on Balslev & Balseca 4721, NY; C based on Skutch 4545, NY.) 2006] N E E ET AL.: SOLANUM A N D CAPSICUM (SOLANACEAE) errnes, racemosae; calyx 3 mm longus, fere truncatus; corolla eburnea, profunde lobata, lobis linearilanceolatis, 12-20 x 2-2.5 mm; antherae attenuatae, 1112 x 1.5 mm, vel nonnunquam 6-7.5 mm longae. Baccae nitidae, aurantiacae vel rubrae, 1.5-2 cm diametro; semina complanata, reniformia, 3.4-3.7 x 2.8-3.2 ram. Scrambling vine; stem closely tomentose with ferruginous sessile (or nearly) 7 - 8 rayed stellae, 0.4 m m wide, with obsolete midpoints, eventually glabrescent, armed with uniform small recurved prickles 1.5-3 m m long from enlarged flattened bases. Sympodial units unifoliate or usually difoliate, rarely 3-foliate, geminate. Leaves 5 - 1 8 x 1.3-6.5 cm, simple, dimorphic or not, the major leaves 10-18 x 2.5-6.5 cm, narrowly elliptic, the minor leaves 5 - 1 8 x 1.3-6.5 cm, differing from the major ones only in size, thin-textured, sparsely to densely tomentose adaxially with sessile 4 - 8 - r a y e d stellae 0 . 3 0.5 m m wide, the midpoints absent to 1.5 m m long, the trichomes not obscuring the surface, tomentose abaxially with sessile or very short-stipitate stellae 0.6-0.7 m m wide, somewhat larger than those above but with obsolete midpoints, the surface visible, the leaf undersides with recurved prickles on the midrib similar to those o f the stem; base unequal (oblique); margin entire or very shallowly angulate-lobed; apex acute to attenuate; petioles 0,8-2.5 cm, armed with hairs and recurved prickles like those of the stem. Inflorescence 3-5.5 cm, extra-axillary, unbranched, with 5 - 1 5 flowers, all flowers apparently perfect, the axes stellate-tomentose, unarmed; peduncle 0.8-1.1 cm; rachis 2 . 5 4.5 cm; pedicels 8 - 9 m m in flower, 1 6 - 2 0 m m in fruit, thick and enlarged at the apex, slightly curved, spaced 1-1.5 m m apart, articulated at the base. Calyx ca. 3 m m long, the tube ca. 2.5-3 mm, the margin almost truncate and lobes nearly absent, with an apicular extension of the midrib up to 1 m m long, stellate-tomentose abaxially, glabrous adaxially; fruiting calyx accrescent, to ca. 6 m m long, splitting nearly to the base, eventually breaking off irregularly, the remaining calyx tube flattened and somewhat w o o d y ; corolla 2.5-5 c m in diameter, 18-20 m m long, stellate, attenuate and slightly curved in bud, creamy white, the tube 2 mm, the lobes 1 2 - 2 0 x 2-2.5 ram, linear-lanceolate, tomentose abaxially with weak whitish stellae, 343 glabrous adaxially; filaments obsolete; anthers (6-) 11-12 x 1.5 mm, very slender, attenuate, not connivent, yellow, the pores minute and directed distally, not opening into longitudinal slits with age; ovary sparsely tomentose at apex, glabrescent; style ca. 12 x 0.3-0.4 mm, exceeding the anthers by about 1.5 mm, clavate, straight to slightly curved, stellate-tomentose near the base; stigma truncate to capitate. Fruit a globose berry, 1.5-2 c m in diameter, shiny bright orange or red, glabrous. Seeds numerous (more than 15 per fruit), 3.4-3.7 x 2.8-3.2 mm, flattened, reniform, light brown or yellowish, the surfaces minutely pitted. Distribution and e c o l o g y . - - H u m i d evergreen forests of the A m a z o n basin near the eastern base of the Andes from Colombia to the southern limit of the tropical forest just northwest of the city o f Santa Cruz, Bolivia, also more rarely on the western Andean slopes in Ecuador, at 9 0 - 1 8 0 0 m elevation. P h e n o l o g y . - - F l o w e r i n g and fruiting all year. E t y m o l o g y . - - T h e epithet reflects the distribution along the piedmont, the lower eastern slopes of the Andes and immediately adjacent A m a z o n lowlands, and the western Andean slopes in Ecuador. C o m m o n n a m e s . - - E c u a d o r : "cocona del monte" (Lawesson et al. 39532B); Peru: "ayac mull~ca" (bitter fruit, Mexia 6485), "chirapahuasca" ( Schunke 77). U s e s . - - T h e seeds are said to be used for "skin spots; caustic" (Mexia 6485). Additional specimens examined. COLOMBIA. AMAZONAS: Municipio Leticia, Parque Nacional Natural Amacayacu, sector de Mata-matg, a la orilla de la quebrada Bacaba, 3~ 70~ 100 m, 16 Apr 1992 (fr), Rudas & Prieto 4278 (BM); Municipio Leticia, brazo del Rio Amazonas, lado de la estaci6n principal del Parque Nacional Natural Amacayacu, 14 Jun 1992 (fl), Rueda 556 (BM); Loretoyacu River, 100 m, Mar 1946 (fr), Schultes 7130 (GH). E C U A D O R . ESMERALDAS: San Josr, kin. 321 along railroad from Ibarra to San Lorenzo, I~ 78~ 350 m, 4 May 1982 (fr), Boom 1364 (MO); Cantrn San Lorenzo, Reserva Indigena Aw~i,Cation del Rio Mira, l0 km W de Alto Tambo, Comunidad "La Unirn", 1~ 78~ 250 m, 16-26 Mar 1991 (fr), Rubio et al. 1112 (NY). MORONA-SANTIAGO: Pozo petrolero "Garza" de TENNECO, 35 kin NE de Montalvo, 1~ 76~ 260 m, 2-12 Jul 1989 (fr), Zak & Espinoza 4376 (MO, NY), 4448 (BM, MO, NY). NAPo: Yasun/ Forest Reserve, 1-3 km E of Pontificia Universidad Catrlica de Ecuador Science Station, by Tiputini River, 0~ 344 BRITTONIA [ V O L . 58 76~ 240 m, 16 Jun 1995 (fr), Acevedo-Rodrfguez & Cedeho 7339 (MO); Cant6n Archidona, S del Volc~in 1~ 75~ 200 m, 30 May 1980 (fr), Brandbyge & Asanza 31260 (AAU, BM); Pastaza Cant6n, pozo Sumaco, carretera Hollfn-Loreto, km. 31, Comuna Challua Yacu, 0~ 77~ 15-17 Nov 1988 (fr), A. A1varado 53 (NY); Orellana Cant6n, Parque Nacional Yasunf, carretera y oleoducto de Maxus en construcci6n, km. 54-58, 00~ 76~ 250 m, 26-30 Sep 1993 (ft, fr), Aulestia & Andi 760 (BM, MO, QCNE); Aguarico, Reserva Etnica Huaorani, carretera y oleoducto de Maxus en construcci6n, km 61, S del rio Tivacuno, 0~ 76~ 250 m, 26-30 Oct 1993 (fr), Aulestia et al. 1041 (MO, NY); Rio Wai si ay~i, 1 km upstream from the outlet in Rio Aguarico, 1~ 76~ 6 Aug 1981 (fr), Brandbyge et al. 33229 (NY, QCA); road between Baeza and Lago Agria, 72.5 km W of Lago Agria, 1166 m, 19 Dec 1979 (fr), Croat 49532 (MO, NY); Cant6n La Joya de los Sachas, Parque Nacional Yasunf, carretera de Maxus, km. 45, 0~ 76~ 230 m, 8-15 Aug 1993 (fr), Dik 96 (QCNE); Orellana, Parque Nacional Yasunf, carretera y oleoducto de Maxus en construcci6n, 13-16 Sep 1993 (fl), Dik 449 (BM); Aguarico, Reserva Etnica Huorani, carretera Maxus, km. 72-75, 0~ 76~ 270 m, 23-31 Jan 199- (fr), Dik & Andi 931 (MO, QCNE); Cant6n Aguarico, Reserva Etnica Huorani, carr. Maxus, km 77-78, 0~ 76~ 250 m, 24-28 Feb 1994 (fr), Dik & Enomenga 1089 (MO, QCNE); 3.54.8 km E of Rfo Conejo on rd to Lago Agrio (7.8-9.1 km W of Lago Agrio), 1 Apr 1972 (fr), Dwyer & Macbryde 9811 (MO); 17 km W of Lumbaque (70 km W of Lago Agrio), 1130 m, 6 Nov 1974 (fl, fr), Gentry 12573 (MO); Cant6n Orellana, Sector Huashito, 20 km N de Coca, propiedad de PALMORIENTE, 0~ 77~ 250 m, 3-21 Nov 1989 (fr), Gudi~o 150 (MO, NY); road Lago Agrio--E1 Conejo, between E1 Conejo and Proyecto San Miguel, 380 m, 15 Feb 1980 (fl, fr), Harling & Andersson 16534 (MO); road Coca-Auca oilfields, 3 km along the road to Yucca, 0~ 76~ 400 m, 20 Aug 1979 (ft), HolmNielsen et al. 19635 (AAU); Rfo Aguarico, Tangoy, 1 hour upstream from Zancudo, 0~ 75~ 300 m, 29 Aug 1979 (fr), Holm-Nielsen 20092 (AAU, BM); 3 km E de Caserfo de Huamanf, N de la carretera HollinLoreto, 0~ 77~ 1200 m, 17 Sep 1988 (fr), Hurtado & Alvarado 310 (MO, NY); vfa Lago Agrio-Rfo San Miguel (frontera con Colombia), 0~ 76~ 700 m, 11 Feb 1980 (fr), Jaramillo 2232 (MO, QCA); Coca-Rfo Payamino-Cooperativa de la Comuna, Rio Payamino, 23 Feb 1981 (fr), Jaramillo & Coello 4141 (QCA, QCNE); Parque Nacional Yasunf, Bloque 16, CONOCO, localidad Daimi 1, 0~ 76~ 14 Sep 1989 (fr), Jaramillo et al. 10941 (NY, QCA); Afiangu, Rio Napo, 0~ 76~ 28 Apr--6 May 1983 (fr), Lawesson et al. 39532B (AAU, QCA); Coca, 7 Sep 1983 (fl, fr), Lescure 2029 (NY, QCA); 4.2-7.5 km W of Lago Agrio (5-8.2 km E of Rfo Conejo) near Lago AgrioBaeza road, 340 m, 31 Mar 1972 (fr), MacBryde & Dwyer 1366 (MO); carretera Hollfn-Loreto, km 32, 8 km W de Guamani, arriba del Rfo Guamani, 0~ 77~ 1200 m, 20 Sep 1988 (fl), Neill et al. 8609 (MO, NY); Prov. Orellana, Yasunf National Park, Maxus road and pipeline construction project, km 21, 0~ 76~ 250 m, 24 Jul 1994 (fl, fr), Pitman 644 (BM, MO, QCNE). NAPO-PASTAZA: vic. Puyo, 750-1000 m, Sep 1939 (fl), Skutch 4545 (MO). PASTAZA: Lorocachi, 5 km upriver Rio Curaray from the military camp, petrolero "Moretecocha" de ARCO, 75 km E de Puyo, 1~ 77~ 580 m, 4-21 Oct 1990 (fl), Gudi~o et al. 854 (BM, MO, NY); Rfo Papayacu at Rfo Curaray, 1~ 76~ 235 m, 23 Mar 1980 (fl), Holm-Nielsen et al. 22639 (AAU); Montalvo, Rfo Bobonaza, 29 Dec 1976 (fl, fr), McElroy 251 (CM, QCA). PICHINCHA: Carretera Quito-Puerto Quito, kin. 113, 10 kmal N de la carretera principal, 0~ 79~ 27-29 Dec 1983 (fl, fr), Balslev & Balseca 4721 (NY, QCA); Reserva Forestal ENDESA, Rio Silanche, Corp. Forestal Juan Manuel Durini, km 113 de la carretera Quito-Puerto Quito, 10 km N de la carretera principal, 00~ 79~ 650-700 m, 18 Feb 1984 (fl), Jaramillo 6360 (MO). SUCUMniOS: Reserva Faunfstica Cuyabeno, Laguna Grande, 0~ 76 ~10'W, 265 m, 11 Mar 1990 (fr), Balslev et al. 97395 (QCNE); Cantrn Gonzalo Pizarro, Parroquia E1 Reventador, 2 km SO del caserio El Reventador, 0~ 77~ 1800 m, 31 Jul 1991 fir), Yrnez & Bonilla 339 (QCA). PERU. HU~NUCO: Pachitea, Codo de Pozuzo, Rfo Pozuzo, 9~ 75~ 450 m, 18 Oct 1982 (fl, fr), Foster 9259 (BM, MO); Hu~inuco to Pamayacu, 13 Jan 1927 (fr), Kanehira 189 (F). LORETO: Prov. Alto Amazonas, Dtto. Pastaza, Rfo Pastaza, Andoas, Capahuari Norte, 12 Nov 1979 (fr), Ayala 2252 (NY); Maynas, Dtto. Iquitos, Rio Maroon, tributary of Rio Nanay, 150 m, 10 Dec 1976 (fl), Davidson & Revilla 5376 (F, MO, NY); Prov. Maynas, Rfo Itaya, 10 minutos arriba de San Juan de Muniches, 3~ 73~ 130 m, 21 Nov 1978 (fr), Dfaz et al. 644 (MO, NY); Prov. Alto Amazonas, Quebrada Nucuray (tributary del Rfo Marafi6n), a media hora arriba de la quebrada, 4~ 75~ 23 Jan 1979 (fr), Dfaz & Ruiz 897 (MO, NY); Prov. Alto Amazonas, Rfo Pastaza, una hora arriba de la boca del Lago Rimachi, 4~ 76~ 200 m, 25 Jan 1979 (fr), D:az & Ruiz 918 (MO, NY); Prov. Alto Amazonas, orillas del Rfo Pastaza, entre Rimachi y Rfo Witoyacu, 4~ 76~ 31 Jul 1979 (fr), Dfaz et al. 1298 (MO, NY); Maynas, mouth of Rfo Nanay below Iquitos, 120 m, 18 Mar 1979 (fl, fr), Gentry et al. 25818 (MO, NY); Prov. Alto Amazonas, Capihuari, 5 km NE of Andoas on Rfo Capihuari, near Ecuador border, 240 m, 17 Nov 1979 (fl, fr), Gentry & Dfaz S. 28184 (MO, NY); Prov. Alto Amazonas, Andoas, Rfo Pastaza near Ecuador border, 2~ 76~ 14 Aug 1980 (fr), Gentry et al. 29691 (MO, NY); Prov. Maynas, Explorama Camp, Quebrada Sucusari, Rio Napo, 3~ 72~ 130 m, 30 May 1991 (fl), Gentry 74303 (BM, MO, NY); lower R/o Mom6n, tributary of Rfo Nanay, near Iquitos, 8 Dec 1979 (fl), Jones & Davidson 9718 (NY); along Rfo Marafirn, near mouth of Rfo Tigre, 115 m, 19 Aug 1929 (fr), Killip & Smith 27519 (NY); Puerto Arturo, lower Rfo Huallaga below Yurimaguas, 135 m, 24-25 Aug 1929 (fl, fr), Killip & Smith 27753 (NY, US); Rfo Itaya, above Iquitos, 110 m, 17-22 Sep 1929 (fl), Killip & Smith 29389 (NY); Mishuyacu, near Iquitos, 100 m, Feb-Mar 1930 (fl), Klug 910 (E NY, US); Maynas, Dtto. Iquitos, Rfo Itaya, barredera de Pefia Negra, 110 m, 6 Mar 1973 (fl, fr), MeDaniel 16879 (MO); Maynas, Dtto. Punchana, Rfo Mom6n, mouth to Porvenir, 90 m, 10 Jan 1994 (fl, fr), McDaniel & Rimachi Y 32105 (MO); Distrito Iquitos, Creek Itaya, 100 m, 6 Feb 1932 (fl), Mexia 6485 (F, MO, NY, PH, UC); Rio Javari, below mouth of 2006] NEE ET AL.: SOLANUM AND CAPSICUM (SOLANACEAE) Rio Curaqa, 27 Oct 1976 (fr), Prance et al. 24160 (MO, NY); Gamitanacoeha, Rio Maz~in, 100-125 m, 18 Jan 1935 (fl), Schunke 77 (E NY, UC); Maynas Prov., Las Amazonas, rivera de la Quebrada Sucusari, Explor Napo Camp, 3~ 72~ 140 m, 18 Apr 1991 (fr), Vdsquez & Jaramillo 16131 (MO); Caballo-Cocha on the Amazon River, 14 Aug 1929 (fl), Ll. Williams 2479 (F); Alto Rfo Itaya, 145 m, 10 Sep 1929 (fl), L1. Williams 3448 (F). PASCO: Prov. Oxapampa, Palcazu valley, Cabeza de Mono, 5-6 km W of Iscosacin, 10~ 75~ 325 m, 17-20 Apt 1983 (fr), D. N. Smith 3825 (NY). BRAZIL. ACRE: Municipio Rio Branco, estrada Rio Branco/Puerto Acre km 33, a 2 km da margem, 12 Oct 1980 (fr), Cid & Nelson 2883 (NY); Municipio Bujad, basin of Rio Purus, Rio Antimari, Floresta Estadual do Antimari, 9~ 68~ 8 Mar 1997 (fl, fr), Daly et al. 9352 (MO); Boca do Acre-Rio Branco road, 14 km from Rio Branco, 27 Sep 1980 (fr), Lowrie et al. 228 (R); Proj. RADAM, sub-base de Cruzeiro do Sul, Ponto 2, SB-18-ZB, 16 Feb 1976 (fl, fr), Marinho 206 (NY); Municipio Sena Madureira, Fazenda Nova Olinda, 10~ 69~ 8 Sep 1995 (fr), Oliveira et al. 644 (NY); Mun. Bujarf, Riozinho do Andir~i, 24 Mar 1995 (fl, fr), M. de Pardo et al. 60 (MO); Mait~i, Rio Moa, 26 Oct 1966 (fl, fr), Prance et al. 2893 (NY, WIS); Mun. Sena Madureira, bacia do Rio Purus, Faz. Nova Olinda, Rio Iaco, Carreador do S~o Bento II, ca. 10 km da Sede, 10~ 69~ 29 Oct 1993 (fr), Silveira et al. 686 (MO, NY); RBO-AC, EE.A., Rio Antimari, 26 Feb 1992 (fr), Sothers & Santos 83 (MO, NY). AMAZONAS:Mun. Sgo Panlo de Oliven~a, Palmares, 11 Sep-Oct 1936 (fl), Krukoff 8421 (F). BOLIVIA. COCNABAMBA. Prov. Carrasco: Puerto Villarroel, 16~ 64~ 200 m, 8 Jun 2000 (fr), Villavicencio et al. 1780 (NY). LA PAZ. Prov. Iturralde: Luisita, W del Rio Beni, zona inundada del Rfo Muqui, 13~ 67~ 180 m, 29 Feb 1984 (fl, fr), Beck & Haase 10142 (MO). SANTACRUZ. Prov. Ichilo: [Rfo] Yapacanf, 400 m, Jan 1892 (fl), Kuntze s.n. (NY); Parque Nacional Ambor6, Rfo Saguayo, 17~ 63~ 500 m, 19 Jan 1988 (fr), Nee 35991 (NY, USZ); La Vfbora, ca. al puente sobre el Rfo Chore, 17~ 63~ 275 m, 9 Nov 1990 (fr), Saldias et al. 1274 (MO, NY). belongs to a closely related a s s e m b l a g e of woody vines that scramble by m e a n s o f recurved prickles. Four species are f o u n d in Mexico and Central A m e r i c a (Nee, 1993), but m a n y more are f o u n d in South America, i n c l u d i n g this surprisingly widespread species that has not yet received a name. The group has been called the S o l a n u m l a n c e i f o l i u m group by W h a l e n (1984) and section M i c r a c a n t h a D u n a l b y Nee (1999). Evidently some flowers are functionally male, as heterostyly is c o m m o n in all collections seen. However, a study of the living plants is needed to fully understand the sexual expression of the flowers. Solanum pedemontanum 345 The young parts are ferruginoustomentose, but the general aspect is m u c h paler and more yellowish than in S o l a n u m a t u r e n s e Dunal, which is partly sympatric in the northern part of the range of S. p e d e m o n t a n u m . The underside o f the leaf, in particular, has both pale stellate hairs and a partially visible pale surface. T h e very close tomenturn of this species is a good visual character. A few of the specimens resemble S. l e u c o p o g o n Huber with very short trichome midpoints instead of the long midpoints that gives a very pilose appearance to the latter species. Part of this material was reported for Peru by Macbride (1962) as S o l a n u m h e t e r o p h y l l u m Lam., a s y n o n y m o f S. s u b i n e r m e Jacq., which differs by b e i n g an upright shrub with violet corollas and decidedly curved anthers. Solanum pedemontanum resembles the description of S. t e r n i f o l i u m Werderm., a species described in the mid-20th century (Werdermann, 1940) from a single Ecuadorian specimen collected at M e r a ( S h u l z e R h o n h o f 2 8 5 8 ) and lost from the Berlin h e r b a r i u m during the Second World War. T h e description of S. t e r n i f o l i u m is detailed but not quite definitive; the n a m e is m u c h more likely a s y n o n y m of S. a t u r e n s e Dunal. O n l y the local phase of S. a t u r e n s e , but not S. p e d e m o n t a n u m , would be described as having " p e d u n c u l o . . . stellato-piloso," "calyx stellato-pilosus," and " c o r o l l a . . . extus dense stellato-pilosis." Solanum c o m a r a p a n u m M. Nee, sp. nov. TYPE: Bolivia. Santa Cruz. Prov. Caballero: Abra Catalina, along hwy. from C o m a r a p a to C o c h a b a m b a , 8 k m W of Comarapa, 17~ 64~ 2425 m, 10 M a r 1988 (fl, fr), M . N e e & J. S o l o m o n 36599 (HOLOTYPE: LPB; ISOTYPES: AD, BH, CORD, K, MO, NY, USZ, WIS). (Fig. 10) Frutex 1.5-3 m altus; rami juniores aculeis rectis vel parum recurvatis ad 8 mm longis armati, vetustiores saepe inermes; caules dense stellato-tomentosi; folia ramorum floriferorum plerumque inermia, ovata, 7-20 • 3-10 cm, ambitu variabilia, integra, repanda vel lobulata. Inflorescentiae 2-4-ramosae; calyx tubo 2 mm longo, lobis lanceolatis ad fere linearibus; corolla rotato-quinquangula, 2-2.5 cm lata; antherae 4.5-5 mm longae, f e r e lineares; ovarium dense stellatotomentosum. Baccae dense stellato-tomentosae, 1.9 cm 346 BRITTONIA [VOL. 58 FIG. 10. Solanum comarapanum M. Nee. A. Juvenile leaf. B. Branch with flowers and fruits. C. Flower at anthesis, abaxial and adaxial views. D. Anthers. E. Gynoecium. F. Infructescence. (A, C - E based on Nee & Solomon 34035, NY; B based on Nee et al. 37413, NY; F based on Nee & Solomon 36599, NY.) 2006] NEE ET AL." SOLANUM AND CAPSICUM (SOLANACEAE) diametro; semina complanata, reniformia, 3.6 • 2.9 mm. Shrubs 1.5-3 m tall; stems densely stellate-tomentose, ferruginous when young, paler when mature, with 6 - 9 - r a y e d stellae ca. 0.6 m m broad, the midpoints ca. 0.2 m m long, y o u n g plants and sprouts normally armed with broad-based prickles to 8 m m long, these mostly straight, some slightly reflexed or recurved, the base usually densely tomentose, older stems mostly unarmed. Sympodial units difoliate, not geminate. Leaves 7 - 2 0 • 3 - 1 0 cm, simple, more or less uniform in size and shape on flowering branches, but those o f y o u n g plants or sprouts much larger (to 35 • 23 cm), ovate, thin-textured, pubescent adaxially with stipitate stellae 0.3-0.6 m m broad, the rays 6 - 8 , the midpoints ca. 1 m m long, the stipes variable in length, up to 0.2 m m long, more densely stellate-tomentose abaxially and with the stellae lighter in color, unarmed except rarely with a few stout prickles on major veins on undersides o f sprout leaves; base truncate to rounded, oblique; margin highly variable in outline, with leaves o f flowering stems often entire or irregularly repand or lobed near the base, the juvenile leaves with three lobes per side, the sinuses to reaching within 2 c m of the midrib and the lobes again pinnately lobed, the lobes rounded to rarely acute; apex attenuate to sometimes rounded; petioles to 5 cm, densely stellatetomentose, unarmed except on sprouts, then with prickles 1.5-2.5 (-4) c m long like those o f the stem. Inflorescences 4 - 8 cm, extraaxillary, 2 - 4 branched, with few to 26 flowers, all flowers evidently perfect, the axes densely stellate-tomentose with hairs like those o f the stem, unarmed; peduncle 1-2 cm; rachis 2 - 6 cm; pedicels ca. 10 m m in flower, 17-19 m m in fruit, sulcate and slightly clavate, spaced 2 - 4 m m apart, articulated at the base; calyx 3 - 7 m m long, the tube ca. 2 mm, the lobes 4 . 5 - 8 • ca. 1.5 mm, lanceolate to nearly linear, densely stellatetomentose abaxially; fruiting calyx not accrescent, the lobes 4 . 5 - 8 • ca. 1.5 mm; corolla 2-2.5 cm in diameter, 8 - 1 1 ( - 1 6 ) m m long, rotate-pentagonal, white, the tube 1 mm, the lobes 7 . 5 - 1 1 ( - 1 5 ) • 2.5-5 mm, deltate, with ample interpetalar tissue, densely stellate-tomentose abaxially on the 347 midribs with slightly smaller and finer stellae than on the stems, glabrous adaxially; filaments glabrous, the free part ca. 1.5 mm, the filament tube absent; anthers 4 . 5 - 5 • 1-1.3 mm, nearly linear, connivent, yellow, the pores minute and directed distally, not opening into lognitudinal slits with age; ovary densely stellate-tomentose; style ca. 7 • 0 . 4 0.5 ram, cylindrical, straight or slightly curved, with a few stellae in the basal third; stigma capitate. Fruit a globose berry, to 1.9 c m in diameter, shiny, green when immature, the color when mature not known, densely stellate-tomentose with sessile stellae, tardily glabrescent. Seeds ca. 25 per berry, ca. 3.6 • 2.9 mm, flattened, reniform, yellowish tan, the surfaces minutely foveolate. Distribution and ecology.--Known only from a small area of the mountains at the border of the Departments of C o c h a b a m b a and Santa Cruz at ( 1 3 0 0 - ) 2 5 0 0 - 2 8 0 0 m elevation. It apparently also occurs in the Department of Tarija, but the identification o f the single collection is still somewhat doubtful. It occurs in weedy disturbed areas such as along roadsides in the dry forests just below the cloud forest zone but does not descend into the semi-arid valleys, which are dominated by cacti and Schinopsis h a e n k e a n a Engl. (Anacardiaceae). Many o f the collections are f r o m the highway to C o c h a b a m b a between C o m a r a p a and the cloud forest area known as "Siberia." P h e n o l o g y . - - F l o w e r i n g in March, June, and December; fruiting in February, March, June, and December. E t y m o l o g y . - - T h e species is named after the town of Comarapa, the site o f many collections of this species. Additional specimens examined. BOLIVIA. COCHABAMBA.Prov. Carrasco: Siberia, 2800 m, Dec 1959 (fl, fr), M. Cdrdenas 5779 (K). SANTACRUZ. Prov. Caballero: 12.5 km by road NW of Comarapa on road to Cochabamba, near Torrecillas, 2450 m, 10 Feb 1987 (fr), Nee & Solomon 34035 (LPB, NY); along road from Comarapa to Cochabamba, 15 km NW of Comarapa, 6.5 km NW of Torrecillas, 2600 m, 10 Mar 1988 (fr), Nee & Solomon 36605 (LPB, NY); hwy. from Comarapa to Cochabamba, 4 km (by air) NW of Comarapa, 17~ 64~ 2100 m, 11 Dec 1992 (fl), Nee 43056 (LPB, NY, USZ). Prov. Florida: Hierba Buena, 1300 m, 8 Jun 1966 (fl, fr), R. F. Steinbach 271 (F, GH, MO, NY, UC, WIS). Prov. Vallegrande: 6 km E of Guadalupe, 2450 m, 5 Feb 1988 (fr), Nee & Saldias P. 36242 (LPB, NY); between Mataralcito and E1 Palmar on road from Valle- 348 BRITTONIA grande to Tierras Nuevas, 17 km ESE of Vallegrande, 2150 m, 29 Dec 1988 (fl, fr), Nee et al. 37413 (LPB, NY). TARIJA. Prov. O'Connor: camino de Junacas a Narvfiez, 1950 m, 8 Nov 1974 (fl), Tiirpe et al. 5047 (W). Most species of spiny solanums, and perhaps most especially those of this group (the S o l a n u m t o r v u m group of Whalen, 1984 or section Torva Nees of Nee, 1999), are characterized by the leaves becoming progressively smaller and less lobed as the plant matures. Saplings and sprouts have deeply lobed, spiny leaves and often the flowering branches of older plants have much smaller, entire, and unarmed leaves, sometimes being easy to confuse with the unarmed subgenus S o l a n u m . However, each species has a particular range of variation of leaf lobing. S o l a n u m c o m a r a p a n u m has very tomentose and unusually prominently lobed leaves except on the oldest or least vigorously growing material, and at least some of the lobes tend to be undulate. The herbarium specimens available have a widely divergent aspect because of this leaf lobing, but observations in the field and collections from individual plants demonstrate that they all belong to a single species. The very pubescent fruits are useful for identification, but apparently this is variable, as some collections, noticeably N e e & Saldias 3 6 2 4 2 and N e e et al. 3 7 4 1 3 , have glabrescent fruits; fully mature fruits of this species are not yet known. The ferruginous color of the young growth, partly maintained in older plants, is obscured on many of the specimens as they were collected at the edge of the gravel highway and the dense tomenturn has become covered with road dust. Solanum saturatum M. Nee, sp. nov. TYPE: Bolivia. Santa Cruz. Prov. Caballero: 1.5 km down from E1 Empalme (on Comarapa-Cochabamba hwy) on road to Khara Huasi, 17~ 64~ 2475 m, 2 Aug 2003 (fl), M. N e e et al. 5 2 4 1 3 (HOLOTYPE: USZ; ISOTYPES:LPB, NY). (Fig. 11) Frutex 1-4 m altus; caulis ad 4 cm diametro; rami juniores aculeis parum recurvatis ad 7 mm longis armati, vetustiores fere semper inermes; caules dense stellatotomentosi; folia ramorum floriferorum inermia, ovatolanceolata, 7-15 • 3-10 cm, integra vel lobata. Inflorescentiae simplices vel 2-ramosae; calyx lobis triangulariter apiculatis; corolla rotato-quinquangula, 4- [VOL. 58 4.5 cm lata; antherae 9-11 x 2 ram, attenuatae; ovarium glabrum. Baccae glabrae, 1.2-1.4 cm diametro; semina complanata, reniformia, 2.5 x 2 ram, Shrubs or treelets, 1-4 m tall; stems sparsely tomentose with sessile or stipitate 7 8-rayed stellate hairs 0.7-0.8 mm wide, the midpoints absent, grading into small prickles to at least 2 mm long with a stellate hair on the tip, fertile branches densely stellatetomentose with sessile or short-stalked stellae, the younger parts ferruginous-tomentose, the lower part of the main stem and sprouts heavily armed with thick-based, slightly recurved prickles to 7 x 3 mm at the base, with sessile stellae on the bottom 1/3; fertile branches unarmed, or only rarely with a few short prickles. Sympodial units unifoliate or the uppermost ones difoliate and geminate. Leaves 7-15 x 3-10 cm, more or less uniform in size and shape, but younger plants or sprouts with much larger leaves (to 22 x 16 cm), ovate-lanceolate, thin-textured, ferruginous-tomentose on both sides with sessile or short-stalked stellae, scabrous adaxially with sessile 5-7-rayed stellae 0.50.6 mm wide, the midpoint nearly obsolete to 0.3 mm long, the leaf surface partially visible, more densely and loosely tomentose abaxially with lighter colored short-stalked stellae obscuring the leaf surface, unarmed even on vigorous sprout leaves (except on very young plants or sprouts); base truncate to cuneate; margins of blades on fertile branches usually entire, or sometimes with 2 shallow lobes per side; apex acute; petioles 1-2.5 (-5.5) cm, densely stellate-tomentose, unarmed. Inflorescence 2-5.5 cm, extraaxillary, usually appearing terminal, eventually overtopped and lateral, unbranched or forked, with 3-10 flowers, often with one flower at the base and the rest more clustered toward the tip, all flowers perfect, the axes ferruginous-tomentose with stellate hairs, unarmed; peduncle 0-0.8 cm; rachis 2-3.5 cm; pedicels 13-15 mm in flower, up to 20 mm in fruit, spaced 3-8 mm apart, articulated at the base. Flowers with a thick sweet fragrance, the calyx ca. 5 m m long, the lobes ca. 3 x 2.5 mm, triangular, apiculate at tips, densely stellate-tomentose; fruiting calyx not accrescent, the lobes ca. 3 x 2.5 mm; corolla 4-4.5 cm in diameter, 25 mm long, rotate-stellate, pentagonal in bud, white, the tube 6 mm, the 2006] NEE ET AL.; SOLANUM AND CAPSICUM (SOLANACEAE) 349 FIG. 11. Solanum saturatum M. Nee. A. Stem detail. B. Juvenile leaf. C. Flowering branch. D. Bud with detail of stellate hairs. E. Flower at anthesis. F. Anthers. G. Style and stigma. (A based on Steinbach 8343, NY; B-G based on Nee et al. 52413, NY.) 350 BRITTONIA lobes ca. 10 x 15 ram, ovate-lanceolate, with ample glabrous interpetalar tissue, acute at apices, abaxially pubescent with brownish hairs on the midrib, glabrous adaxially; filaments glabrous, the free part 2 - 3 mm, the filament tube absent; anthers 9-11 x ca. 2 ram, tapered, not connivent, yellow, the pores small and directed distally, not opening into longitudinal slits with age; ovary glabrous; style ca. 11 • 0.5 ram, cylindrical, straight or slightly curved, glabrous; stigma capitate. Fruit a globose berry, 1.2-I .4 cm in diameter, green (perhaps not fully ripe?), glabrous. Seeds numerous (one count of 17), ca. 2.5 • 2 ram, flattened-reniform, yellow, minutely foveolate. Distribution and ecology.--Found in the cloud forest along the "old" highway from Santa Cruz to Cochabamba, in the Departments o f C o c h a b a m b a and Santa Cruz, Bolivia at 2 2 0 0 - 3 0 0 0 m elevation. Phenology.--Flowering in February, May, June, August, September, and October, and fruiting in February, August, and October. It will probably be found to flower and fruit throughout the year. Etymology.--The epithet s a t u r a t u m refers to the thick sweet fragrance of the flowers, an odor rare elsewhere in S o l a r i u m , but also found in related species of the S. t o r v u m group from Department La Paz, Bolivia into Peru. Additional specimens examined. BOLIVIA. COeHAnAMnA OR SANTA CRUZ: hwy. CochabambaSanta Cruz, km. 205, 9000 ft, 18 Sep 1964 (fl), Badcock 381 (K, NY). COCHABAMUA.Prov. Carrasco: Siberia Este, Monte Hotel, 17~ 64~ 2700 m, 22 Sep 2003 (fl), Duran et aL 92 (MO); hwy. Epizana-Comarapa, 15 km (by road) E of highway bridge near Pojo, 9 km W of El Churo, 17~ 64~ 2675 m, 24 May 2001 (ft), Nee et al. 51856 (NY); road from Comarapa to Cochabamba, 4 km W of border with Depto. Santa Cruz, 20 km (by air) and 28 km (by road) NW of Comarapa, 17~ 64~ 2525 m, 10 Feb 1987 (fr), Nee & Solomon 34054 (LPB, NY). SAr~TACRUZ. Prov. Caballero: S of Ambor6 National Park, Cerro Bravo area, 4-10 km N of Comarapa, 17~ 64~ 2300-3000 m, 23 Jun 1995 (fl), Abbott & Jardim 17206 (NY); Siberia Sud, Astilleros, 17~ 64~ 2850 m, 20 Sep 2003 (fl), E, Ferndndez et al. 31 (MO); Astillero, 17~ 64~ 2550 m, 13 Aug 2003 (fl, fr), E. Fernrndez et al. 2002 (MO); between Capitla and Siberia on the old Cochabamba hwy, 17~ 64~ 2500 m, 18 Aug 2000 (fl), Kuroiwa & Maeda 2205 (NY); 6.5 krn (by air) N of Comarapa, Cerro Bravo area, 17~ 64~ 2380 m, 3 Aug 2003 (ill Nee et al. 52442 (NY); Empalme, 17~ [VOL. 58 64~ 2600 m, 6 Feb 2004 (fl, fr), Rivera et aL 52 (MO); Bergwald yon Comarapa, 2800 m, 20 Oct 1928 (fl, fr), J. Steinbach 8343 (BM, E, F, GH, MO, NY, PH). Prov. Florida: 7 km (by air) NE of Mairana, 18~ 63~ 2200 m, 2 Jun 1991 (fl), Nee 40657 (NY). Solanum saturatum is a m e m b e r o f Whalen's (1984) S o l a n u m t o r v u m group and Nee's (1999) section Torva. Before more material b e c a m e available and fieldwork demonstrated the difference of the two species, N e e & S o l o m o n 3 4 0 5 4 and J. S t e i n b a c h 8 3 4 3 were considered to belong to S. c o m a r a p a n u m . The two are nearly sympatric, but S. s a t u r a t u m grows in clearings in cloud forest, whereas S. e o m a r a p a n u m is more likely to be found on shrubby roadsides in drier w o o d land just below this cloud forest zone. The leaves of S. s a t u r a t u m never develop the complex lobing found in S. c o m a r a p a n u m . Solanum s a t u r a t u m is morphologically somewhat similar to the local expression of S. a s p e r o l a n a t u m Ruiz & P a v . (often annotated as its s y n o n y m S o l a n u m h i s p i d u m Pers.), but this species in the Siberia area has long-stipitate stellate hairs giving it a distinctly bristly appearance, and its inflorescence does not begin branching near the base, but instead in the more distal half o f the inflorescence. s c u t i c u m M. Nee, nora. nov. Replaced name: S o l a n u m t a b a c i f o l i u m Dunal, in A. DC., Prodr. 13(1): 261. 1852. nora. illeg. TYPE: Brazil, Bahia, 1830, S a l z m a n 3 8 9 (r~OLOTYPE: G - D C [F neg. 6808, photos: BH, F, MO, NY, US]; probable isotypes (without number but "nov. sp."): K-2 sheets, W). N o n S. t a b a c c i f o l i u m Vell., 1825. (Fig, 12) Solanum Shrubs 1.5-3 (-4) m tall; flowering stems densely ferruginous-tomentose with 7 - 8 rayed stellae 0.4-0.8 m m broad, with stipes of various lengths, to 1 m m long, tardily and irregularly glabrescent, unarmed or often with a few broad-based prickles to 4 m m long. Sympodial units difoliate, geminate. Leaves simple, dimorphic, the major leaves 15-23 x 7 - 1 2 cm, ovate, the m i n o r leaves 8.5-14 • 5 - 9 . 5 cm, about the same shape as the majors, thin-textured, moderately and evenly ferruginous-tomentose adaxially with 4 - 8 - r a y e d stellae 0.3-0.5 m m wide, the hairs 2006] N E E ET A L . : S O L A N U M A N D C A P S I C U M ( S O L A N A C E A E ) FIG. 12. Representative specimen of Solanum scuticum M. Nee (Nee 33378, NY). 351 352 BRITTONIA mostly sessile and not overlapping, but with denser and more stipitate hairs on the major veins, densely pale stellate-tomentose abaxially with mixed sessile and stipitate stellae forming several layers, the stipitate stellae like those of the stem, usually unarmed abaxially or with a few acicular prickles up to 9 m m long on the major veins beneath; base acute to truncate or subcordate, unequal; margin usually entire, but often irregularly repand to undulate or with 1-3 shallow lobes per side; apex acute to attenuate; petioles 11.5 (-3.5) cm, stellate-tomentose like the stem, unarmed or rarely with a few prickles like those of the leaf underside. Inflorescence 0.5-3 cm, opposite the leaves, branched, many- (more than 10-) flowered, all flowers perfect, the axes densely ferruginoustomentose, the stellae 6-8-rayed, the midpoints slightly shorter than the rays, the stipes of various lengths, to 0.6 m m long; peduncle 0-0.7 cm; rachis 1.5-4.5 cm; pedicels ca. 8 m m in flower, to 11-20 m m long in fruit, spaced 1-2 m m apart, articulated at the base. Calyx 3 m m long, the tube 2 ram, the lobes ca. 1 x 1.5-2.5 ram, triangular, attenuate-apiculate at tips, densely stellatetomentose abaxially; fruiting calyx not accrescent, the lobes ca. 1 x 1.5-2.5 ram; corolla ca. 2 cm in diameter, 14-16 m m long, stellate-pentagonal, white, the tube 2-3 mm, the lobes 9 x 4.5 mm, deltate, with ample interpetalar tissue, acute at apices, closely and finely stellate-tomentose abaxially on the midrib with stellae smaller than on the rest of the inflorescence, glabrous adaxially; filaments glabrous, the free part ca. 1 ram, the filament tube absent; anthers 6-7.5 x l m m , tapered, free, yellow, the pores minute and directed distally, not opening into longitudinal slits with age; ovary densely stellate, glabrescent; style of hermaphrodite flowers ca. 8 x 0.3-0.5 mm, cylindrical, straight, stellate in the lower part; stigma capitate. Fruit a globose berry, 0-1.4 cm in diameter, green, turning yellow-orange, glabrous. Seeds numerous, 90-265 per fruit, 1.6-2.5 ram, flattened-reniform, yellow, the surface minutely foveolate. Distribution and e c o l o g y . - - O n l y the Bolivian and Paraguayan collections are listed here; there are very numerous collections from Brazil from the states of Acre, Bahia, [VOL. 58 Distrito Federal, Espfrito Santo, Goi~is, Mato Grosso, Minas Gerais, Paran~i, Rio de Janeiro, Rondrnia, Santa Catarina and S~o Paulo, where it is a common, somewhat weedy species of open areas and forest edges. The distribution nearly coincides with the geological formation of pre-Cambrian crystalline rocks, known as the Brazilian shield. In Bolivia the known distribution is also confined to the pre-Cambrian shield area of eastern Department Santa Cruz. Etyrnology.--The epithet scuticum is derived from the Latin scutum, a shield, in figurative reference to the geological Brazilian shield. Additional specimens examined (outside Brazil). BOLIVIA. SANTACRUZ. Prov. Iquflode Ch~ivez:E of San Javier on road to Concepci6n, 16~ 62~ 450 rn, 22 Feb 1995 (fl, fr), Abbott 16345 (SI); road to dam on Rio Zapocoz, 0-2 km NW of Concepci6n, 480 m, 28 Dec 1986 (fl, fr), Nee 33378 (LPB, NY); 6 km E of San Javier on road to Concepci6n, 16~ 62~ 540 m, 1 Dec 1990 fir), Nee 40142 (LPB, MO, NY). PARAGUAY. AMAMBAYIUpper Rio Apa, Jan 191213 (fl), Hassler 11252 (BM, GH, K, MO, S, UC, W). This c o m m o n species, a m e m b e r of section Torva sensu Nee (1999) or the Solanum torvum group sensu Whalen (1984), is here documented from Bolivia; it has long needed a name, but none has been found among the several thousand epithets that have been proposed in Solanum. The species was known to Sendtner (1846) who treated it under a very heterogeneous concept of Solanum torvum Sw., in which he included several other species from Mexico to Brazil in a complicated hierarchy of infraspecific taxa at indeterminate ranks, rendering all his infraspecific epithets invalid. These names will be treated in more detail in a revision of section Torva (Whalen & Nee, in prep.). The similar S. torvum, found sparingly along coastal eastern Brazil from Bahia to Sao Paulo and now spread throughout the tropics as an abundant weed, differs most obviously by its sparsely glandular-tomentose pedicels; Solanum scuticurn lacks glandular hairs on the pedicels. Solanum tabacifoIium Dunal is a later homonym of S. tabaccifolium Veil. and thus illegitimate (Greuter et al., 2000; Article 53.1.). Solarium tabaccifolium Veil. is probably a synonym of S. mauritianum Scop., a 2006] NEE ET AL.: SOLANUM AND CAPSICUM (SOLANACEAE) widespread member of the Brevantherum clade sensu Bohs (2005). A Brazilian specimen, Lhotsky s.n. (G-DC), was one of the three syntypes of S. daturif o l i u m Dunal; it belongs to Solanum scuticum. However, from among the syntypes, Schulz (1909) chose Sieber 67 (G) from Martinique as the lectotype for S. daturifolium Dunal. Sieber 67 belongs to the widespread species S. torvum Sw., making the name S. daturifolium a synonym of S. torvum. Solanum scuticum is similar to S. rudepann u m Dunal from Mexico, Central America, Colombia, and Ecuador in having very ferruginous inflorescences, stems, and leaves. However, in S. scuticum the calyx in bud stage is much shorter than that of S. rudepann u m and the lobes are very shallow. In late bud stage and especially by the time the plant is in fruit, the calyx lobes split apart and become oblong, often with an acuminate tip. Many herbarium specimens of S. scuticum had been previously identified as S. asperolanatum Ruiz & Pay., but this is a very different, more robust species from midelevations in the Andes from Bolivia to Venezuela, with a larger inflorescence which normally has a peduncle several cm long. S o l a n u m whalenii M. Nee, sp. nov. TyPe: Bolivia. La Paz. Prov. Nor Yungas: 3.2 km S of and below Chuspipata on road to Chulumani, 16~ 67~ 2900-3000 m, 28 Sep 1985 (fl), M. Nee & J. Solomon 31958 (HOLOTYPE: LPB; ISOTYPES: AD, BH, CORD, G, K, MO, NY, P, US, WIS). (Fig. 13) Frutex vel arbor parva, 2-8 m altus; caules aculeis muniti. Folia ovata, integra vel leviter lobata, (12-) 1840 x (6-) 10-20 cm, valde bicoloria, super glabrescentia viridiaque, inferne dense et persistenter albidotomentosa. Inflorescentia extra-axillaris,dense albidotomentosa, (7-) 10-25 cm, 2-9-ramosa, pedunculo 1.5-3.5 cm longo; pedicelli floriferi 2-2.5 cm longi, fructiferi 2.4-3.1 cm longi, 5-angulati; corolla alba, 4 cm diametro, profunde 5-1obata; antherae 8.5 x 2.5 mm. Fructus glaber, 1.2-1.5 diametro, viridis; semina plurrima, 2 x 1.6 mm. Solano albido Dunal valde simile sed robustius. Shrub or tree, 2-8 m tall; stems densely appressed-tomentose with sessile, multirayed stellae so dense as to be individually difficult to distinguish, flowering stems usually unarmed but the lower stem with stout prickles, younger stems conspicuously striate 353 when dry. Sympodial units difoliate, geminate. Leaves (12-) 1 8 4 0 x (6-) 10-20 cm, simple, more or less uniform in size and shape, ovate on flowering stems, strongly bicolorous, thin-textured, initially pubescent adaxially with sessile whitish stellae 0.4 m m broad and with ca. 16 rays, the midpoints very short or obsolete, soon glabrate-green, densely white-tomentose abaxially with sessile and short-stipitate stellae in interwoven layers, with about 16 rays, the midpoints obsolete, unarmed; base asymmetrically obtuse to sub-cordate, rarely acute; margin entire or obscurely sinuate-repand; apex acuminate; petioles of major leaves 4-5 cm, strongly striate or canaliculate when dry, this striation continuing onto the lower part of the midvein and onto the stem, densely whitish tomentose like the stem, unarmed. Inflorescence (7-) 10-25 cm, up to ca. 30 cm in fruit, extraaxillary, 2-9-branched, with many (up to more than 100) flowers, all flowers apparently perfect, the axes densely stellatetomentose; peduncle 1.5-3.5 cm; rachis 4 - 1 6 cm, up to 26 cm in fruit; pedicels 20-25 m m in flower, to 24-31 m m in fruit, spaced 3-5 m m apart, straight, relatively stout, gradually enlarged toward apex, 5-angled; calyx 6-10 m m long, the tube ca. 2 m m long, 5-costate, the lobes ca. 2 x 4 mm, deltoid, apiculate at tips, densely stellate-tomentose abaxially; fruiting calyx somewhat accrescent, often splitting in the sinuses and the lobes then appearing longer, the lobes 6-7 x 3-4 mm; corolla ca. 4 cm in diameter, 21-25 m m long, rotate-stellate, white, the tube 1.5-2 mm, the lobes 19-23 x 5 - 6 mm, ovate-lanceolate, with ample glabrous interpetalar tissue, acute at apices, densely stellate-tomentose abaxially with stellae like those of the inflorescence, glabrous adaxially except for a few stellate hairs distally along the midrib; filaments glabrous, the free part ca. 2 mm, the filament tube absent; anthers 8.5 x 2.5 mm, slightly tapered, not connivent, Yellow, the pores small, directed distally, not opening into longitudinal slits with age; ovary glabrous or minutely stipitate-glandular at apex; style 11-13 x 0.8-0.9 m m and wellexserted, cylindrical, straight or somewhat curved, glabrous or minutely stipitateglandular at base; stigma capitate. Fruit a globose berry, 1.2-1.5 cm in diameter, dark 354 BRITTONIA [VOL. 58 FIG. 13. Solanum whalenii M. Nee, A. Leafy branch with inflorescences. B. Bud with detail of inflorescence trichomes. C. Open flower. D. Anthers. E. Inflorescence with immature berries. (A based on Nee et at. 5J765; B - E based on Nee et aL 51806). F. Inflorescence of Solanum albidum Dunal for comparison (based on Nee 36826, NY). 2006] N E E ET AL." S O L A N U M A N D C A P S I C U M ( S O L A N A C E A E ) gre en w h e n immature, the c o l o r when fully ripe unknown, glabrous. S e e d s numerous (ca. 120 per fruit), ca. 2 x 1.6 mm, flattenedreniform, y e l l o w - b r o w n , the surface minutely foveolate. Distribution.--Andean slopes in western and central B o l i v i a in Departments o f C o c h a b a m b a and La Paz at 1 8 0 0 - 3 0 0 0 m elevation. Phenology.--Flowering in May, September, and October, and fruiting in January, March, May, and October. Further collecting will show whether f l o w e r i n g is confined to the dry season. Etymology.--The epithet honors M i c h a e l D e n n i s Whalen, an expert on the Solanaceae, w h o s e work on S o l a n u m was cut short m u c h too soon. Additional specimens examined. BOLIVIA. COCHABAMBA. Prov. Carrasco: Sehuencas, despu~s pasar el Rfo Fuerte, 17~ 65~ 2100 m, 1 May 1993 (ft, fr), P. Ibisch & C. lbisch 194 (LPB); narrow canyon of Rio Monte Puncu, 5 km NE of Monte Puncu, l0 km (by air) NW of Epizana, 17~ 65~ 2700-2750 m, 10 Mar 1988 (fr), Nee & Solomon 36625 (MO), 36634 (MO). Prov. Chapare: camino de Cochabamba a Villa Tunari, Villa Tunari, 23 Sep 1982 (fl), Cabrera & Guti6rrez 33722 (LPB, SI); Incacorral, 1900 m, Mar 1941, C6rdenas 2246 (US); km. 120, Cochabamba to Chapare, 1800 m, Apr 1961, Cdrdenas 5974 (K, US); 23.8 km N of Colomi 0unction of the road to Candelaria) on road to the Chapare, then 5 km NW (left) on side road, upper Rio Cayani, 17~ 65~ 2600 m, 19 Oct 1985 (fl), Solomon 14404 (MO); Incachaca, 2500 m, 16 Sep 1921 (fl), J. Steinbach 5786 (SI); Nordosthange der Sierra de Cochabamba, Umgebung von Incachaca, 2500 m, Jul 1926, Werdermann 2063 (MO); Incacorral, forest opening, 2400 m, 10 Jun 1929, J. Steinbach 9820 (BM, E, F, MO, NY, PH, US). Prov. Mizque: Mizque, 2000 m, Jun 1940, Cdrdenas 2133 & 2134 (US). LA PAZ. Prov. Larecaja: Quiabaya, Sorata, Mandon 425 (G-DC [F photo 23130; a specimen of this number at G seems to be S. albidum]. Prov. Murillo: Valle de Zongo, vic. Escuela Cambaya, 28.3 km N of (below) La Cumbre, along Rio Zongo, 16~ 68~ 2560-2800 m, 10 May 1990 (fl), Luteyn & D o r r 13616 (MO); Rfo Zongo valley, 22.5 km below dam at Lago Zongo, 16~ 68~ 3000 m, 9 Oct 1982 (fl, fr), Solomon 8432 (MO); Zongo valley, 25.2 km below the dam at Lago Zongo, 16~ 68~ 2700 m, 19 Jan 1985 (fr), Solomon 13102 (MO). Prov. Nor Yungas: road from La Paz to Coroico, S of divide, ca. 20 km from La Paz, 4000-4700 m, 21 May 1980 (fl), D'Arcy & Bejarano 13833 (MO, NY); 9 km by road (ca. 4 km by air) down from and NE of Chuspipata, 16~ 67~ 2450 m, 29 Oct 1984 (fl, fr), Nee & Solomon 30227 (NY). Prov. Sud Yungas: Chuspipata 6.5 km hacia Chulumani, 2570 m, 13 Sep 1981 (fl), Beck4780 (LPB). 355 This species is similar to the m o r e widespread S o l a n u m a l b i d u m , but is m o r e robust in nearly every part and is usually found at h i g h e r altitudes. It is m o r e restricted ecologically; in the area w h er e they are sympatric in Bolivia, S. a l b i d u m g r o w s f r o m 270 to 2600 m, but is generally f o u n d b el o w 2000 m, w h er eas S. w h a l e n i i is restricted to 1 8 0 0 3000 m. Th e co l o r o f the fully mature fruits is not known; they are h e l d erect on erect inflorescences and i f they r e m a i n dark green, it m a y indicate that they are dispersed by bats. S o l a n u m w h a l e n i i is o n e o f the most spectacular species o f section T o r v a ( S o l a n u m t o r v u m group sensu W h a l e n , 1984). A photograph o f the flower appears on the co v er o f a t h e m e issue of the P h i l o s o p h i c a l T r a n s a c t i o n s o f the R o y a l S o c i e t y , B i o l o g i c a l S c i e n c e s dedicated to " T a x o n o m y for the twenty-first century" (Godfray & Knapp, 2004). Acknowledgments We thank N S F for funding this w o r k through the Planetary B i o d i v e r s i t y Inventory program, award DEB 0316614 'PBI S o l a n u m - a w o r l d w i d e t r e a t m e n t ' ; Bo b b i A n gell for preparing the illustrations; K i m Watson, S h an n o n Crouch, and H e a t h e r Rolen for help with the manuscript and specimen i m ages; R M. JCrgensen and an a n o n y m o u s rev i e w e r for helpful c o m m e n t s on the manuscript; the curators o f the herbaria m e n t i o n e d in the text for loan o f s p e c i m e n s in their care; and G. Mois6s M e n d o z a F. and Israel Vargas C. for their valuable help in collecting these and other B o l i v i a n plants in the field. Literature Cited Anderson, G.J. & R.K. Jansen. 1998. Biosystematic and molecular systematic studies of Solanum section Basarthrum and the origin and relationships of the pepino dulce (5:. muricatum). Monographs in Systematic Botany from the Missouri Botanical Garden 68: 17-32. 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