SBM. IOIO ISSN 0968-0446 Bulletin of The Natural History Museum Taner HISTORY MUSEUM } 51 JUL 1997 PRESENTED | GENERAL LIBRARY Botany Series ad THE NATURAL HISTORY MUSEUM VOLUME 27 NUMBER 1 27 JUNE 1997 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Botany Series is edited in the Museum’s Department of Botany Keeper of Botany: Dr S. Blackmore Editor of Bulletin: Ms M.J. Short Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) © The Natural History Museum, 1997 Botany Series ISSN 0968-0446 Vol. 27, No. 1, pp. 1-73 The Natural History Museum Cromwell Road London SW7 5BD Issued 27 June 1997 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Lond. (Bot.) 27(1): 1-5 Issued 27 June 1997 Notes on the diatom species Tetracyclus castellum (Ehrenb.) Grunow with a description of Tetracyclus pseudocastellum nov. sp. THENATIIRAL DAVID M. WILLIAMS Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD HISTORY MUSEUM | Shes CONTENTS I Gof afore (Tel t (0) eee renee errr ceric tater reece errr eer reer MSrMinOlo py w4s-5. sess cs tesescestacs ate eesetsvesas dastarbet cdi incensuaee Systematic escriptions pescciscestressee eevee etek ee sctseettacsen ts ees Tetracyclus castellum (Ehrenb.) Grunow .........scceeseeeeeeees Tetracyclus pseudocastellum D.M. Williams ..............0006 RELETEN CES aii selec cs sasiectecaccsscttascccesteea dati ssesessevenicesstesievenccereeeee PRESENTED GENERAL LIBRARY | SYNOPSIS. This paper reviews the evidence for retaining the taxon described by Ehrenberg as Biblarium castellum Ehrenb. (transferred to the genus Tetracyclus by Grunow). Consideration of numerous new names proposed since Ehrenberg establishes that T. castellum is a valid taxon hich has been re-described several times during the period 1903-1983. While previously considered to be known only from a few fossil specimens, this paper establishes that it has been recorded as living (from Iceland). In addition, a better understanding of 7: castellum has revealed a new fossil species from China, 7: pseudocastellum. INTRODUCTION The diatom genus Tetracyclus Ehrenb. (Bacillariophyta) comprises at least 30 species, of which only five have been reported living, the remainder occurring exclusively as fossils (Williams, 1987, 1989, 1996). The taxonomy of the genus has more or less relied on the shape and dimensions of the valve as well as the frequency of particular valve characters, such as striae and ribs (Hustedt, 1914; Li, 1982a, b, 1984). While the majority of species are either ellipti- cal or circular in valve outline, there are a few taxa that have more or less star-shaped valves. Two of these species, 7: emarginatus (Ehrenb.) W. Sm. and T. japonicus (Petit) Temp. & H. Perag. have already been described in detail with both light and electron microscopy (Williams 1987, 1989). As a continuation of those studies, this paper describes ataxon Ehrenberg called Biblarium castellum Ehrenb. (=T. castellum (Ehrenb.) Grunow) and a new fossil species from Inner Mongolia, T. pseudocastellum. 1 also offer some notes on other taxa possibly confused with 7. castellum. While this study is limited to light microscopy only, it will serve as a focus for the further study of specimens under the scanning electron microscope if and when appropriate material is discovered and becomes available. TERMINOLOGY A number of papers dealing with the particulars of diatom valve terminology have been published in the last 15 years. For the siliceous parts of the diatom valve and girdle, Anonymous (1975), © The Natural History Museum, 1997 and its updated version Ross et al. (1979), are the standard refer- ences followed in this study. Stosch (1975) presented the first detailed discussion on girdle band morphology and nomenclature. However, since his pioneering effort much has been discovered and some of the conclusions reached in his paper are subject to debate and undoubtedly will be modified in due course; some aspects of possible modifications have been discussed by Mann (1982), Williams (1985), and Round et al. (1990). Additional commentary relevant to Tetracyclus morphology can also be found in Williams (1985, 1987, 1989, 1996). SYSTEMATIC DESCRIPTIONS Tetracyclus castellum (Ehrenb.) Grunow in Verh. zool.-bot. Ges. Wien 12: 411 (1862). — Tetracyclus japonicus sensu Lupikina in Nov. Sist. Nizsh. Rast. [1965]: pl. 3, figs 1-3 (1965); Khursevich & Loginova, Iskopaemaya Diatomovaya Flora Belorussii (Sistematicheskit Obzor): pl. 17, fig. 13 (1980); Khursevich in Acta geol. hung. 28: pl. Il, fig. 7 (1982). — Tetracyclus stellare sensu J.Y. Li in Bull. Inst. Geol. chin. Acad. geol. sci. 5: pl. 1, fig. 18 (1982); J.Y. Li & Y.Z. Qi in Proc. 8th Internat. Diat. Symp.: pl. 2, figs 4, 5 (1986); Valeva & Temniskova-Topalova in Fitologiya 46: pl. III, figs 11, 12 (1993). — Tetracyclus stellare var. eximia sensu VanLand. in Micropaleontology 31: pl. 1, fig. 9 (1985). — Tetracyclus sp. Tscheremisinova, Diatomovaya Flora Neogenovykh Otlozhenii Pribaikal’ya (Tunkinskaya Kotolovina): pl. 6, fig. 2 (1973). Figs 1, 3, 4, 7. XS RN > KS px \y ( 2 D.M. WILLIAMS Figs 1,3,4,7 T. castellum. Fig. 1: Reproduction of Ehrenberg (1854): pl. 33/2, fig. 1. Fig. 3: Tetracyclus ‘costellatus’ from Temp. & Perag., Diat. monde entier, 2nd ed., slide no. 122, BM 68468, specimen 30 yu long. Fig. 4: Tetracyclus elegans var. eximia from Temp. & Perag., Diat. monde entier, 2nd ed., slide no. 122, BM 68468, specimen 25 yp long. Fig. 7: 7: ‘costellatus’ var. turris from Temp. & Perag., Diat. monde entier, 2nd ed., slide no. 134, BM 68479, specimen 25 up long. Fig. 2 Biblarium elegans reproduced from Ehrenberg (1854): pl. 33/2, fig. 4a, b. Fig.5 7. emarginatus var. crassa from Temp. & Perag., Diat. monde entier, 2nd ed., slide no. 122, BM 68468, specimen 40 p long. Fig.6 T. japonicus from Temp. & Perag., Diat. monde entier, \st ed., slide no. 79, BM 14331, specimen 55 p long. Fig. 8 7. pseudocastellum. Inner Mongolia, China, BM 81618, specimen 40 1 long. DIATOM SPECIES TETRACYCLUS CASTELLUM Biblarium castellum Ehrenb. in Ber, Akad. Wiss. Berlin [1843]: 47 (1843), nom. nud. —Ehrenb. in Ber Akad. Wiss. Berlin [1845]: 73 (1845). —Ehrenb. in Mikrogeol.: pl. 33/2, fig. 1 (1854). Type: ‘Ad Bargusinam Sibiriae fossile’, Ehrenberg (1843: 47), specimens not located. Iconotype = pl. 33/2, fig. 1 in Ehrenberg (1854). ?Tetracyclus islandica Ostrup in Meddr dansk geol. Foren. 6: 28, pl. 1, fig. 1 (1900). Type: Ilagil, Iceland (K 384-holotype?). Tetracyclus costellatus Hérib., Diat. foss. Auvergne: 16, pl. 8, fig. 12 (1902), orth. var., corrected to castellum in Perag. Cat. Diat.: 920 (1903). Type: ‘Dépot de Celles, Cantal’ (BM 68468-isotype). Tetracyclus costellatus var. turris Perag. & Hérib. in Hérib. Diat. foss. Auvergne: 39, pl. 8, fig. 13 (1902), orth. var., corrected to castellum in Perag., Cat. Diat.: 920 (1903). Type: ‘Dépot d’ Auxillac, Cantal’ (BM 68479-3-isotypes). Tetracyclus elegans var. eximia Hérib. & Perag. in Hérib., Diat. foss. Auvergne: 16, pl. 8, fig. 15 (1902). Type: “Dépot de Celles, Cantal’ (BM 68468-isotype). Tetracyclus stellare Hérib., Diat. foss. Auvergne: 31, pl. 11, fig. 23 (1903). Type: ‘Dépot de Joursac, Cantal’ (BM 68397—99-isotypes). Tetracyclus stellare var. eximia (Hérib.) Hust. in Abh. naturw. Ver. Bremen 23: 98 (1914). Tetracyclus lapponicus Tynni in Bull. geol. Surv. Finl. 320: 35, pl. 19, figs 10-15 (1982). Type: ‘Gyttya deposit of Sivakkapalo’ (GTL HH/80-holotype, not seen). Tetracyclus chudjakovii Pushkar in Paleobot. Fitostrat Vostoka SSSR: 114, pl. 22, figs 15-17 (1983) (AH CCCP 123/30-79-U- holotype, not seen). Valves with 8 equally spaced points somewhat resembling a ‘star’, 25-45 p (n=10) in diameter, each tip curving at its margin (Li & Qi, 1986: pl. 2, fig. 4). Transapical ribs primary (25-45 in 10 yu), radiate; secondary and tertiary ribs present, extending into each point of the star and meeting at the sternum; striae in equidistant rows. Cingulum consisting of open septate bands. Septum small, not visible in a number of bands (= secondary copulae?) (Fig.1, septum visible in Ehrenberg’s illustration; Li & Qi, 1986: pl. 2, fig. 5). Material examined Living Iceland. ‘Illagil. Tinnardalur, Skagafhordssyssel... (K 384, holotype? of Tetracyclus islandica). Fossil France. Cantal, Joursac, BM 68397-99 (Isotypes of Tetracyclus stellare, Temp. & Perag., Diat. monde entier, 2nd ed., nos 51-53); Cantal, Celles, BM 68468 (Isotype of Tetracyclus costellatus, T.: elegans var. eximia, and T. emarginatus var. crassa, Temp. & Perag., Diat. monde entier, 2nd ed., no. 122); Cantal, Auxillac, BM 68479- 3 (Isotype of T: costellatus var. turris, Temp. & Perag., Diat. monde entier, 2nd ed., nos 133-137). U.S.A. Nevada, Esmeralda Co., SW of Loric Mountain and west of Tonopato Esmeralda formation, USGS 5078 (CAS 382005). What is understood as Tetracyclus castellum is based upon type material from synonymous taxa, as Ehrenberg’s material is unavail- able. In addition five valves were discovered in a fossil deposit from the U.S.A. (Tonopato Esmeralda formation, CAS 382005). As the species is known from so few specimens the synonymy requires further comment. Biblarium castellum — Ehrenberg (1843) described the species Biblarium castellum from a fossil deposit in Siberia (‘Infusorien- 3 Lager von Bargusina im Gouvernement Irkutzk in Sibirien’, Ehrenberg, 1843: 46; ‘Ad Bargusinam Sibiriae fossile’, Ehrenberg, 1845: 73). Although Ehrenberg provided a reasonable (for his time) description (‘B. corpusculorum valvis (intermediis) ovatis obtusis, sinubus marginalibus utrinque quatuor. Laterales valvae nondum observatae’, Ehrenberg, 1845: 73), he offered only one illustration in the Mikrogeologie (Ehrenberg, 1854: pl. 33/2, fig. 1, reproduced here as Fig. 1). The specimen he chose to illustrate is clearly of a girdle band and provides no information on valve structure, of which Ehrenberg appeared to have no knowledge (‘Laterales valvae nondum observatae.’ Ehrenberg, 1845: 73). Ralfs (in Pritchard, 1861: 806) added nothing of significance to the species description, reproduc- ing Ehrenberg’s figure (in Pritchard, 1861: pl. iv, fig. 44) and translating his 1845 text (‘Lateral view of central portion elliptic, with obtuse ends, and four marginal undulations . . . Lateral valves unknown’, Pritchard, 1861: 806). Although Grunow (1862: 411) transferred the species to the genus Tetracyclus, he also appears not to have investigated relevant material and again relied only on Ehrenberg’s description and figure. This approach continued with De Toni referring to Ehrenberg, Grunow, and Ralfs (all of whom used the same single specimen) for his own description of this species for which he was able to provide additional perspective: ‘Valvis late ovatis, obtusis, subrhomboideis, marginibus triundulatus.... (De Toni, 1892: 748). In summary, Tetracyclus castellum has remained a valid name based on Ehrenberg’s one girdle band specimen for which type material has been unavailable for examination, one imagines, since Ehrenberg’s time. However, it is possible to suggest two things from this illustration of a girdle specimen: first, that the valves would be similar in shape, that is like an 8-pointed star; and second, that the girdle has a septum (Fig. 1). Tetracyclus ‘costellatus’ — When Héribaud undertook his study of the fossil diatoms of Auvergne in France he made the new combina- tion Tetracyclus costellatus based on Biblarium costellatum, attributing the specific epithet to Ehrenberg (Fig. 3; Héribaud, 1902: 16, pl. 8, fig. 12). In the same volume, Tempere & Héribaud described the new variety T: costellatus var. turris (Fig. 7; Héribaud, 1902: 39, pl. 8, fig. 15), again with reference to the Biblarium costellatum of Ehrenberg (Lauby [1910: 340], in a study of the same area, also used the name T. costellatum). Ehrenberg never used the name costellatum in connection with the genus Biblarium. That Héribaud made an error with the name was identified by Peragallo (1903: 920) who corrected both names. Mills (1935: 1600) included Tetracyclus costellatus (and the variety turris) in his catalogue as a synonym of T. castellum (to compound confusion Mills misspelt 7: costellatus as T. constellatus). VanLandingham acknowledged that Héribaud’s usage of the name T. costellatus was a misspelling of castellum (‘error? for Biblarium castellum Ehrenberg 1843...’, VanLandingham, 1978: 3981) and included it as an orthographic variant of 7: castellum. Isotype material for 7: ‘costellatus’ and T. ‘costellatus’ var. turris are available as part of Tempére & Peragallo’s Diatomées du monde entier exiccata set (2nd ed., slide no. 122, BM 68468; slide nos 133- 137, BM 68479-83) and although rare, a few specimens have been examined (Figs 3, 7). In each case the valves are 8-pointed stars suggesting that they can be usefully compared with Ehrenberg’s Biblarium castellum. Tetracyclus elegans, T. elegans var. eximina, T. emarginatus var. crassa, and T. stellare — Tetracyclus elegans (Ehrenb.) Hérib. was based upon another Ehrenberg species, Biblarium elegans, de- scribed from the same Siberian fossil deposit as B. castellum (Ehrenberg, 1854: 90, pl. 33/2, fig. 4a, b). Ehrenberg provided no description and only published illustrations of two specimens, one 4 valve and one girdle band (Ehrenberg, 1854, pl. 33/2, fig. 4a, b; reproduced here as Fig. 2). Ralfs transferred the species to Tetracyclus, providing a minimal description (‘Inflations acute’) and noting that ‘Ehrenberg’s figure of this species differs from T. rhombus merely in its more developed inflation’ (Ralfs in Pritchard, 1861: 806-7). De Toni, however, differed from Ralfs and in his view likened T: elegans to T. lacustris Ralfs (= T. glans (Ehrenb.) Mills; see Williams, 1987). 7; rhombus (Ehrenb.) Ralfs in Pritchard has been discussed in more detail in Williams (1996) and 7. glans in Williams (1987). Briefly, Ehrenberg’s original illustrations of 7: rhombus included drawings of specimens from Siberia and a U.S.A. fossil deposit from Columbia River (Ehrenberg, 1854: pl. 33/12, figs 7, 8, pl. 33/2, figs 9, 9*, 10; see Williams, 1996, for notes on the Columbia River deposit). From the illustrations alone, it appears that specimens from Siberia (Ehrenberg, 1854: pl. 33/2, figs 9, 9*, 10) may indeed belong to T. glans (or some closely related species, e.g. T. pagesi Hérib. or T. stella (Ehrenb.) Hérib.; cf. Hustedt, 1914: 101, 105; Williams, in prep.) while the Columbia River specimens (Ehrenberg, 1854: pl. 33/12, figs 7, 8) are probably a small pre-auxospore stage of some elliptical-valved species (see Williams, 1990, 1996). Until Siberian material has been examined these conclusions must be considered unsubstantiated. However, it does explain Ralfs’ and De Toni’s conflicting views noted above. Héribaud & Peragallo unnecessarily transferred Biblarium elegans to Tetracyclus (Héribaud, 1902: 16; Ralfs had already done so). However, part of their reason was to be able to describe further specimens they encountered in the ‘Celles’ deposit as Tetracyclus elegans var. eximia (Héribaud, 1902: 16, pl. 8, fig. 15). Isotype material is available (Temp. & Perag., Diat. monde entier 2nd ed., slide no. 122, BM 68468) and, although only a few specimens were encountered, they are 8-pointed star-shaped valves like 7: castellum (Fig. 4). In a later volume of the same study, Héribaud published another new species under the name of Tetracyclus stellare Hérib. from the Joursac deposit of Cantal (Fig. 7; Héribaud, 1903: 31, pl. 11, fig. 23). According to Héribaud T. stellare is ‘... intermédiaire entre le Tetracyclus castellum et le Tetracyclus elegans, dont il nous parait une forme dérivée’ (Héribaud 1903: 31). This is clearly referring to the shape of the valve outline. Hustedt concluded that Tetracyclus elegans var. eximia was perhaps better understood as a variety of T. stellare and that T: elegans was better understood as a variety of T: lacustris (=T. glans) and transferred both taxa accordingly (Hustedt, 1914, p. 97 for elegans, p. 98 for stellare; unfortunately, Hustedt refers to stellare as stellaris throughout his monograph). There is merit in Hustedt’s decisions but once again, the absence of Ehrenberg’s Siberian material makes judgment difficult. Neverthe- less, inspection of specimens of T: stellare (as well as T. elegans var. eximia) indicates that there is as yet insufficient evidence to relate it most closely to either T. elegans or T. glans and is best considered as a synonym of T: castellum. Finally, Héribaud & Peragallo described a new variety of Tetracyclus emarginatus, T. emarginatus var. crassa Hérib. & Perag. (Héribaud, 1902: 16, pl. 8, fig. 16; specimens from Temp. & Perag., Diat. monde entier 2nd ed., no. 122, BM 68468; Fig. 5). This taxon only superficially resembles 7: castellum and should perhaps be considered in the context of 7. emarginatus to which it appears more similar. Tetracyclus islandica, T. lapponicus, and T. chudjakovii — Ostrup described the new species Tetracyclus islandica from Illagil in Iceland (Ostrup, 1900: 28, pl. 1, fig. 1). He made no attempt at a description but drew attention to the unusual shape which he felt D.M. WILLIAMS made its unique status obvious (*... som uden at kraeve naermere Beskrivelse, tydeligt fremgaar af Tab. nost. Fig. 1’, @strup, 1900: 28). There is only one relevant slide of type material present in C of which J.B. Hansen wrote: ‘Ostrup used to keep raw and cleaned material of everything but in a few cases where the material is scanty there are only slides available. You have got the only material I can find’ (Hansen, pers. comm.).The specimens on this slide were rather rare and too poor to make useful micrographs. However, it was clear that the ‘edges’ of the valve were somewhat more rounded that in @strup’s published illustration, suggesting that it too should be considered a synonym of T. castellum. This is an interesting conclu- sion as it implies that 7: castellum should properly be considered as a sixth (albeit rare) living species of Tetracyclus. Further material needs to be examined, especially using electron microscopy. Tetracyclus lapponicus Tynni was described as a Neogene fossil from the ‘Gyttya deposit in Finland’ (Tynni, 1982: 35, pl. 19, figs 7, 10-15). Tynni suggested that it ‘closely resembles the form T. japonicus described from the Neogene stratum of White Russia (Khursevich & Loginova 1980)’. Khursevich & Loginova’s (1980) specimen is one of T. castellum (see below) and hence T: lapponicus should also be considered a synonym of T. castellum. Tynni remarks that ‘7: ellipticus var. lancea f. subrostrata Hust. — T. lapponicus with their intermediate forms constitute a transitional series. . ., from which it becomes evident that T. ellipticus and lapponicus are closely related forms.’ (Tynni, 1982: 35). Material has not been examined but evidence presented by Tynni (1982: pl. 19, figs 7, 10- 15) does not seem to support his contention and he relies on an unconventional understanding of T. ellipticus var. lancea f. subrostrata (see Williams, 1996). Finally, Pushkar described the new species Tetracyclus chudjakovii Pushkar (1983: 114, pl. 22, figs 15-17) also with an 8-pointed star- shaped valve and again probably a specimen of T. castellum. SUMMARY. Specimens which appear to be 7: castellum (not forget- ting that this taxon was originally based on one illustration of a girdle band) have been described on a number of different occasions after Ehrenberg, from 1903 to 1983, including T: ‘costellatus’, T.: elegans var. eximina, T. stellare, T. islandica, T. lapponicus, and T. chudjakovii. No doubt much of this re-description is due to poor knowledge of genuine T: castellum specimens. To compound mat- ters other errors have crept in, possibly due to peculiarities surrounding its nomenclature, especially an early confusion involv- ing several different usages of the name Tetracyclus japonicus, clearly a different species from T: castellum as it is a has a valve like a 12-pointed star (Fig. 6, 7: japonicus sensu stricto; see alsoWilliams, 1989). Other illustrations with different names include Lupikina (1965: pl. 3, figs 1-3) and Khursevich & Loginova (1980: pl. 17, fig. 13; see also Khursevich, 1982: pl. II, fig. 7; both illustrations are of the same specimen, the latter being turned upside down) who named specimens of this taxon T: japonicus; Li (19825: pl. 1, fig. 18) and Valeva & Temniskova-Topalova (1993: pl. III: figs 11, 12) who named speci- mens T. stellare,; VanLandingham (1985: pl. 1, fig. 9) who named specimens 7, stellare var. eximia; and Tscheremisinova (1973: pl. 6, fig. 2) who named specimens Tetracyclus sp. All these illustrations seem to be of the same taxon and should be considered representatives of T: castellum. More recently additional specimens have been encountered from Kamchatka (Ozornina, 1993 and pers. comm.). One notable exception is the specimens illustrated by Li (1982a) and Li & Qi (1986) which they erroneously called T: peragalli Hérib. (see Williams, 1990). Examination of relevant material from their Chinese deposit reveals specimens that more correctly belong to a new species, a description of which is given below. DIATOM SPECIES TETRACYCLUS CASTELLUM Tetracyclus pseudocastellum D.M. Williams, sp. nov. Fig. 8. Tetracyclus peragalli sensu J.Y. Li & Y.Z. Qi in Proc. 8th Internat. Diat. Symp.: pl. 2, figs 3, 6 (1986). Valves with 6 equally spaced points somewhat resembling a ‘star’, each point rounded at the margin, 20-45 p (n=7) in diameter. Striae in equidistant rows; ribs predominantly primary, with few secondary ribs extending between the points of the star. Cingulum consisting of open septate bands (Li & Qi, 1986: pl. 2, figs 3, 6). Septum small, difficult to observe in a number of bands (those are possibly second- ary copulae). Known only from type material. TYPE. Late Miocene flora of Inner Mongolia, China. BM 81618, ‘No: SZ,,-1 (9) IM, China’, specimen marked number 5-holotype; IGC-Beijing SZ, -1-01 IMS-isotype. Material examined China. Miocene, Den Hua Jiling Province and Shangdu County of Inner Mongolia, BM 81618, ‘No: SZ,,-1 (9) IM, China’. T. pseudocastellum is known only from the type locality and is easily distinguished by the number of points of the valve: 6 for pseudocastellum (Fig. 8), 8 for castellum (Figs 3, 4, 7), and 12 for japonicus (Fig. 6). Only the latter species is known from a detailed study of its morphology (Williams, 1989). Detailed comparison of valve and girdle structure of these species will allow them to be placed in relation to each other as well as other species of Tetracyclus. The (palaeo)biogeographical interpretation of the genus is largely around the Pacific rim, an understanding of the relationships of the species will allow a better understanding of the causes of this distribution (Williams 1996). ACKNOWLEDGEMENTS. [| am especially grateful to Prof. Li for providing material from the Chinese fossil deposit at Shangdu County for further study, Pat Kociolek and CAS for financial assistance to study the United States Geological Survey (USGS) material in their collections, Svetlana Ozornina for material from Kamchatka, Pete York for photomicography, Eileen Cox for reading and commenting on the manuscript and Mats Wedin for translation of relevant Danish text. REFERENCES Anonymous. 1975. Proposals for a standardization of diatom terminology and diag- noses. Beih. nov. Hedwigia 53: 323-354. De Toni, G.B. 1892. Sylloge Algarum 2(2). Patavii. Ehrenberg, G.C. 1843. Mittheilungen iiber 2 neue asiatische Lager fossiler Infusorien- Erden aus dem russischen Trans-Kaukasien (Grusien) und Siberien. Ber. Akad. Wiss. Berlin [1843]: 43-49. 1845. Neue Untersuchungen iiber das kleinste Leben als geologisches Moment. Ber. Akad. Wiss. Berlin [1845]: 53-88. 1854. Mikrogeologie. Leipzig. Grunow, A. 1862. Die dsterreichen Diatomeen nebst Anschluss einiger neuen Arten von andern Lokalitaten und einer kritischen Ubersicht der bisher bekannten Gattungen und Arten. Erste Folge. Epithemieae, Meridioneae, Diatomeae, Entopyleae, Surirelleae, Amphipleureae. Verh. zool.-bot. Ges. Wien 12: 315-472. Héribaud, J. 1902. Les Diatomées fossiles d'Auvergne. \st Memorie. Paris. —— 1903. Les Diatomées fossiles d’Auvergne. 2nd Memorie. Paris. Hustedt, F. 1914. Die Bacillariaceen-Gattung Tetracyclus Ralfs; kritische Studien uber Bau und Systematik der bisher beschriebenen Formen. Abh. naturw. Ver. Bremen 23: 90-107. Khursevich, G.K. 1982. Neogene diatom assemblages from Byelorussia and their stratigraphic significance. Acta geol. hung. 28: 123-134. & Loginova, L.P. 1980. /skopaemaya Diatomovaya Flora Belorussii (Sistematicheskii Obzor). Minsk. Lauby, A. 1910. Recherches paléophytologiques dans le Massif central. Bull. Serv. Carte géol. détaill. France 20: 1-398. Li, Jia Ying 1982a. Miocene diatom assemblages of Shanwang, Shandong Sheng. Acta bot. sin. 24: 456-467. 1982b. Genus Tetracyclus and its stratigraphic significance. Bull. Inst. Geol. chin. Acad. geol. sci. 5: 149-166. 1984. Some new species and varieties of the genus Tetracyclus Ralfs (Bacillariophyta). Acta phytotax. sin. 22: 231-236. ——& Qi Yu Zao 1986. Neogene diatom assemblages in China. Jn M. Ricard (Ed.), Proc. 8th Internat. Diat. Symp.: 699-711. Lupikina, E.G. 1965. Diatomeae novae et curiosae e stratis Ermanicis partis Kamczatkae occidentalis. Nov. Sist. Nizsh. Rast. 1965: 15-22. Mann, D.G. 1982. Structure, life history and systematics of Rhoicosphenia (Bacillariophyta). I. The vegetative cell of Rh. curvata. J. Phycol. 18: 162-176. Mills, F.W. 1935. An index to the genera and species of the Diatomaceae and their synonyms, 1816-1932. 21: 1571-1726. London. @Ostrup, E. 1900. Diatoméerne i nogle islandiske. Surtarbrand-Lag. II. Meddr dansk geol. Foren. 6: 23-30. Ozornina, S. 1993. Diatoms and problems of stratigraphy of the Eopleistocene and the late Pleistocene of Central Kamchatka. Doctoral Thesis, Vladivostock. Peragallo, M. 1903. Le catalogue général des Diatomées. 2: 472-973. Clermont- Ferrand. Pritchard, A. 1861.A history of infusoria including the Desmidiaceae and Diatomaceae, British and foreign. 4th ed. London. Pushkar, V.S. 1983. Novyi vid roda Tetracyclus Rales [sic] (Bacillariophyta) ie miotsena severnogo skhotena alinya Jn V.A. Krasilov (Ed.), Paleobotanika i Fitostratigrafiya Vostoka SSSR: 112-115. Ross, R., Cox, E.J., Karayeva, N.I., Mann, D.G., Paddock, T.B.B., Simonsen, R. & Sims, P.A. 1979. An amended terminology for the siliceous components of the diatom cell. Beih. nov. Hedwigia 64: 513-533. Round, F.E., Crawford, R.M. & Mann, D.G. 1990. The diatoms. Cambridge. Stosch, H. A. von. 1975. An amended terminology of the diatom girdle. Beih. nov. Hedwigia 53: 1-35. Tempére, J. & Peragallo, H. 1907-1915. Les Diatomées du monde entier. Collection Tempere et Peragallo (2e edition). Arcachon. [See note in R. Ross 1995 Bull. Br. Mus. nat. Hist. (Bot.) 25: 93 concerning the citation of this work.] Tscheremisinova, E.A. 1973.Diatomovaya Flora Neogenovykh Otlozhenii Pribaikal’ya (Tunkinskaya Kotolovina). Novosibirsk. Tynni, R. 1982. The reflection of geological evolution in tertiary and interglacial diatoms and silicoflagellates in Finnish Lapland. Bull. geol. Surv. Finl. 320: 140. Valeva, M.T. & Temniskova-Topalova, D.M. 1993. Diatom analysis of Neogene sediments from the Karlov coal basin I. Fitologiya 46: 67-84. VanLandingham, S.L. 1978. Catalogue of the fossil and recent genera and species of diatoms and their synonyms. 7. Vaduz. 1985. Potential Neogene diagnostic diatoms from the Western Snake River Basin, Idaho and Oregon. Micropaleontology 31: 167-174. Williams, D.M. 1985. Morphology, taxonomy and interrelationships of the ribbed araphid diatoms from the genera Diatoma and Meridion (Diatomaceae: Bacillariophyta). Bibl. Diat. 8: 1-228. 1987. Observations on the genus Tetracyclus Ralfs (Bacillariophyta) I. Valve and girdle structure of the extant species. Br. phycol. J. 22: 383-399. 1989. Observations on the genus Tetracyclus Ralfs (Bacillariophyta) II. Morphol- ogy and taxonomy of some species from the genus Stylobiblium. Br. phycol. J. 24: 317-327. —— 1990. Examination of auxospore valves in Tetracyclus from fossil specimens and the establishment of their identity. Diatom Res. 5: 189-194. 1996. Fossil species of the diatom genus Tetracyclus (Bacillariophyta, ‘ellipticus’ group): Morphology, interrelationships and the relevance of morphogenesis to phylogeny. Phil. Trans. R. Soc. Ser. B, 351: 1759-1782. ee _ ae = an Bull. nat. Hist. Mus. Lond. (Bot.) 27(1): 7-9 Issued 27 June 1997 A new species of Calymperes (Musci: Calymperaceae) from Peninsular Malaysia LEN T. ELLIS Department of Botany, ‘The Natural History Museum, Cromwell Road, London SW7 5BD SYNOPSIS. Malaysia, is described and illustrated. During March 1996 fieldwork was undertaken at Pasoh Forest Reserve, an area of lowland rainforest in eastern Negeri Sembilan, Peninsular Malaysia. Two collections from pristine forest within the reserve appear to represent a hitherto undescribed species of Calymperes. These specimens are identical with a collection made by G.H.S. Wood in 1954 from another area of lowland rainforest near the coast in western Negeri Sembilan — Sungei Manyala Forest Reserve. The new species is described here and named in honour of Wood who made the first collection. Calymperes woodii L.T. Ellis, sp. nov. C. subserrato M. Fleisch. affinis, sed foliis dimorphis, spathulatis, margine supra basin hyalinam polystrato. Type: Peninsular Malay- sia, Negeri Sembilan, Sungei Manyala Forest Reserve, 10 miles SE of Port Dickson, FRI [Forest Research Institute] jungle plot 102, c. 18 m, 13 January 1954, G.H.S. Wood 1372 (BM-holotype; BM- K-isotype). Plants reaching 0.5—1.0 cm high, in mats or as scattered shoots. Leaves curled when dry (often in one direction), erect to spreading (sometimes recurved) when moist, dimorphic (gemmiferous and nongemmiferous leaves). Nongemmiferous leaves mostly >3—4 mm long, lingulate to narrowly spathulate, with a calymperoid hyaline basal region; apices subentire to denticulate, broadly obtuse, usually apiculate. Costa ending immediately below leaf apex; in cross- section composed of dorsal and ventral bands of stereids separated by a single row of guide cells, dorsal and ventral surfaces formed by single layers of small chlorophyllose cells (Fig. 1q, r), superficial cells above hyaline leaf base subquadrate to shortly subrectangular in surface view, mostly 5—15(—22.5) x 7.5—12.5 tum (those forming the dorsal surface longer on average than those forming the ventral surface), sometimes smooth, usually with 1—2 blunt papillae, toward leaf apex many protruding subacutely to acutely. Chlorophyllose lamina occupying four-fifths or more of leaf length (above hyaline basal region), unistratose; cells 6-15 x 6—-12.5 tm, isodiametric to slightly longer than broad, with 4—6 sides or rounded, thick-walled (Fig. 1i, j), each ventrally drawn out as a subacute to acute protru- sion, dorsally pluripapillose (Fig. 1k, 1). Hyaline lamina occupying leaf base, usually not sharply defined; composed of large, subquadrate to subrectangular, thin-walled, porose, hyaline cells; an intramarginal, unistratose band of linear, thick-walled cells, c. (1—)3-6 cells wide, extending from the leaf base toward the distal end of the hyaline lamina sometimes apparent (Fig.1p), often obscure or absent. Leaf margin plane to inflexed, from a short distance above the hyaline base to the leaf apex formed by a subentire to denticulate, polystratose rib composed of isodiametric chlorophyllose cells (stereids some- times present internally), most superficial cells protruding as small teeth (Fig. 1m—o); in hyaline base unistratose, subentire to irregu- © The Natural History Museum, 1997 Calymperes woodii L.T. Ellis, apparently endemic to areas of lowland rainforest in Negeri Sembilan, Peninsular larly denticulate, formed by a band of short, broad irregularly polygonal, thin-walled hyaline cells (often with oblique cross- walls), 1—2(—4) cells wide (Fig. 1p). Gemmiferous leaves often erect and slightly exserted above nongemmiferous leaves, similar to nongemmiferous leaves but up to 5 mm long and sometimes more narrowly lingulate, possessing apices modified as gemma-bearing proboscises (Fig. If, g). Proboscis narrowly suboblong to linear, often curved slightly backwards at tip. Costa strong (usually thicker than in nongemmiferous leaves), extending into proboscis, ending below leaf apex. Lamina narrowing abruptly into proboscis and becoming tightly recurved, becoming plane above and forming a narrow margin around the tip of the costa, ending as a rounded to shortly pointed leaf apex. Gemmae arising in a radial mass from ventral surface of the costal apex, fusiform to clavate, multicellular, uniseriate, smooth (Fig. 1g). Axillary paraphyses produced in brush-like bunches, filamentous, usually exceeding 0.5 mm long, hyaline, multicellular, normally uniseriate (Fig. 1h). Rhizoids con- spicuous around base of shoots, papillose, deep purplish red. Gametangia and sporophytes not seen. DISTRIBUTION. Calymperes woodii appears to be endemic to Negeri Sembilan, Peninsular Malaysia. HABITAT. Calymperes woodii has been collected in rainforest at c. 18 m and 100 m above sea-level. Shoots occur in loose mats, or are scattered over rotting logs or soft bark on the trunks of trees in shaded, damp situations. ADDITIONAL SPECIMENS EXAMINED. Peninsular Malaysia, Negeri Sembilan, Pasoh Forest Reserve, 50 Hectare Plot: tree number 151601,c.100 m, March 1996, Ellis 9601 (BM, FRIM); tree number 131666, 100 m, March 1996, Ellis 9602 (BM, FRIM). DISCUSSION. The absence of sporophytes in the type and other specimens of Calymperes woodii makes the generic placement of this species a matter of strong probability, rather than absolute certainty. Features of the gametophyte in C. woodii bear a degree of superficial resemblance to those found in species of both Calymperes and Syrrhopodon (the two largest genera in the Calymperaceae). However, more features of C. woodii are Calymperes-like than Syrrhopodon-like. For example, the structure of the proboscis in the gemmiferous leaves is virtually identical to that of several species of Calymperes (Fig. 1f, g), particularly C. graeffeanum Miill. Hal. and C. hispidum Renauld & Cardot (both illustrated by Ellis, 1988). Another feature, more usually associated with Calymperes than Syrrhopodon, is the possession of an intramarginal rib in the hyaline basal region of the leaf. Although often obscure to the point of absence, such a rib can be demonstrated in some leaves of C. woodii (Fig. lp). The presence of axillary paraphyses (lacking in C. graeffeanum and C. hispidum) is a feature of some closely interre- 8 L.T. ELLIS res ; 1860e° @e WW. VOSS: P k j BGO n Becxe.t De ae ne 2mm Imm 200um 50um Fig. 1 Calymperes woodii L.T. Ellis a: habit (when moist); b—d: nongemmiferous leaf (b, c: in ventral view, d: detail of apex); e—g: gemmiferous leaf (e: in dorsal view, details of apex in f: ventral view, and g: dorsal view; h: apex of axillary paraphysis; il: chlorophyllose lamina (i, j: ventral surface, k, 1: in cross-section); m—o: margin above hyaline leaf base (m: ventral surface, n, 0: in cross-section); p: margin in hyaline leaf base; q, r: costa at mid-leaf in cross-section. a, b, d, e, g—i, I-n, r Drawn from Ellis 9601 (BM). c, f, j, k, o-q Drawn from Wood 1372 (BM). NEW SPECIES OF CALYMPERES lated species of Calymperes, including C. serratum A. Braun ex Miill. Hal., C. subserratum M. Fleisch., and C. subulatum E.B. Bartram (all regarded as conspecific by Eddy (1990) and Menzel & Schultze-Motel (1990), but shown to be distinct by Reese & Streimann (1994)). C. woodii has axillary paraphyses (Fig. 1b, e, h) and shows some other similarities to C. subserratum and its rela- tives, such as the possession of leaves with a poorly defined hyaline base. However, the leaves of these other paraphyses-bearing species are monomorphic (i.e. gemmiferous leaves are unmodified) and narrowly strap-shaped. In contrast, the leaves of C. woodii are strongly dimorphic and mostly narrowly spathulate. The type speci- men of C. woodii (Wood 1372) was originally erroneously identified as C. subserratum. In addition to the features mentioned above, the latter species has entirely unistratose leaf margins which are incurved to involute and largely subentire (toward the leaf apex a few teeth may occur); the cells of the chlorophyllose lamina are <5—10(—12.5) x <5—7.5 um in surface view. The margins of the leaves in C. woodii are polystratose (Fig. In, 0), plane to inflexed, and minutely denticu- 9 late with single-celled teeth; the cells of the chlorophyllose lamina are 6-15 x 6-12.5 um in surface view. ACKNOWLEDGEMENTS. The work at Pasoh was performed under the auspices of the Forestry Research Institute of Malaysia, and with the financial support of the Natural Resources Institute, U. K. I am grateful to these organizations and also thank Dr A.J. Harrington (BM) for his valuable comments on the manuscript for this paper. REFERENCES Eddy, A. 1990. A handbook of Malesian mosses. 2. London. Ellis, L.T. 1988. Taxonomic notes on Calymperes Il. J. Bryol. 15: 127-140. Menzel, M & Schultze-Motel, W. 1990. The bryophytes of Sabah (North Borneo) with special reference to the BRYOTROP transect of Mount Kinabalu. XI. Calymperaceae (Bryopsida). Willdenowia 19: 475-542. Reese, W.D. & Streimann, H. 1994. Calymperes subserratum (Musci), new to eastern Malesia, with notes on C. serratum and C. subulatum. Bryologist 97(1): 80-82. = > - S , : a - 7 : 7 = = : 7 7 ~ 4 - ees = c = ae - 7 = a - a 7 - - y 7 = a 7 9° 7 a - a — - ~ 7 a 2 7 en 7 =" 2 ~~ 7 4 - : 7 - -" md) > ee att red, oe 7 : 7 - : fs —_ 7 - =) 13 os 7 el islets - ayy inn a ee ere : ar: - oe oy oo —— _ : : oo: : 7 7 : : : > eat = we ——° 64 = > Ga Lao = —— of cece Cpe ee a _ = : : - 7 - vo i % -_ _ : a > =, aha, 5 “—_ew 7) 24 a = @ J Gea _ ee ee a i 7 4 oh 7 = . : - Jie a) 7 a - af 7 - » — 7 o 7 _ a - a oe . nN une ; oe PA a7 ie Oe: > : ce Beno : 7 _ b2& @ waa 7 yy aw _ a ae a ee ok, ea) | nee 7 SO : : - 4 SS 6 : a a 2: a > O eee —. Se © a 7 = ’¢ a Pa’ a a 7 i a : 7 - 7 m 7 : : : : ; —_ re : 7 : - " — ri a & ee oS) -.,. oD ee Jo es oa _s ae ot aes i a f = : ; - 7 7 o : 7 _ Me oy : 7 = - 7 : : . - _ ea - _ ; — Ya == bh ie - aed _ —— = — — A ee 3. _ ag ee = aa: ° 7 _ a 7 - 7 - —— an, : 7 - - co : 7 ——— -*) é a 7 7 a a 7 : a a= > Be > a : oo oS aa a Satyr a > a aS : : = : ; aan a i. : — 7 7 _ ae = 7, ee i - Sa Res . aft, t and = 4 t -. oe a 7 - 7 7 er =) =) 7 7 - 7 ae a - _ a 7 = a = “7 - ? “= : - = - aa 7 an , = = - 7 _ 6 ae ec @ ica eo So 7 a - - Sa - 7 _ yas a > aa oe = - = - 7 a 7 a v 7 => 7 = 2, - - a 7 _ AG 6 I gee ee ee ae — 4 : a =f a a a 7 « Va _ ie w= a == » oe 7 -* ee ™ he - a : 7 = "4 5 7 7 7 i 7 - 7 ‘ a a 7 ' - i 45) 7 . ee S28 Bo > See os OP, ee = - a Pees 7 7 7 : : - al 7 _ s™ : - - a wee a = 7 7 : 7 _- 7 = : 2 wt a Ve _ : - =) — ay > [= 1% —" : 7 Oo an oo - ~ - 7 4 e = _ _ a . 7 a - - a, . 7 a 7 7 538 -4 - OS 7 oc _— 7 7 ee _ 7 7 7~ ee = - _— 7 - : ae ae a DOE i aa iy Se fT Scrat 7 - Oo - ae i 7 7 7 : 7 oa 7 ves 7 : : : 7 7 - - _ 7 = : " ey eee ve — <—eqeee (3 ee eee aa e S a ee —— Pp aca By le 7 - - 7 : - - > - : - = : - : Re eee a er aes a >, =a? ee a a ee eet a _are”6«~ «CS eT oo = 7 P > Gr oT 7 : - 7 =< 7 7 - — - a a © 4) _ 7 - fey : oe a — ie o) aaa iar an ae 7 ? ; - : an .) 7 : ae, ee i? on i y 7 - : - - - : > 7 7 - ro - eo a 7 7 5a 7 7 < a a - a : a _ : = - 7 7 a ; _ fs - i , g i) 7 : - = 7 : - . | = - 7 =e a _ : : - G 7 - : : a iene 7 wor F = - 7 - _ a 2. —- . — og re Ge) Lee _ aan - 7 in : : ~ 7 a 7 a 7; + 5 - 7 7 7 — a 6s) - 7 : _ : : ; ec : ci: 2 ee a - 7 : 4 7 : 7 ta - 7 7 — - 7. 7 a : a a 7 - : - 2 : _ : U : a - : _ a 7 ce ; - ‘. - - i 7 7 7 Baa — | 7 - . ™ 7 a) |< oe ee oe oe , . = ; _ a d ; ; = > & 7 - a - : - | ; * ; | . fe “ 7 _ : 7 : : - - i : : j f 7 en a 7 ; oe - a i < : > a) i Pe a a a : - ee a - Se 7 ae i any, a 7 : 7 = : . . 7 7 - &@ |, a ad - a 7 a .) : o - a 7 - 6 7 . ; 7 i] SS —— Py : 7 7 — : 2 > ac. : a : 7 7 _ a 7 = : ry} - 7 : _ : wy i. oa -< 7 : - _ 7 - - 7 7 7 - 7 5 5 7 = 7 i e 7 a "7 7 " aT) a 7 { a > — - - . - ) - a : - - 7 - - 7 1 [ae - : 3 ; Aa : 7 7 7 =) o : a t a q 7 a 7 . = y : a q 7 : _ 7 7 : : 7 Sw WG 7 : : 5 34% | - ‘ ; y 7 - . n 7 : - 7 7 7 b IC 5 < 5 a, 7 ; 7 - - - : q a — ; 7 - 7 - : - iW a - 7 a 7 7 - : - - 7 5 7 i? : ; i _. oe a 7 i y - rt i Te : r 7 - ; 7 = 5 bel = ; : 7 : 7 : - - , = - : a aan - - 7 7 - 7 - 0 7 - 7 7 5 _ 7 7 7 - _ 7 : 7 - = - - g : o 0 a ai 7 rt - : - - an a 7 . ce. . - _ Ss : a . _ : ° , - ae -_ ee a + oe Ter a 3 ; > tae : : 7: - — “5 ; : a i” 7 7 a : : a a 7 a 7 a - 0 : : 7 — - - on io — al 4 : ; 7 7 + i 7 7 7 7 7 - os 7 4 - S - - : Se : 7 : 7 rast - . = 7 ; Y _ 7 : 7 : ‘ De. 7 7 a 7 7 = : en’) ae", 7 - . ’ _ - a ct ae - - - i. : a a 7 oo ; 7 - 7 7 7 _ 0 oe Se J 7 oe ‘wee [ae | 2 - " = oan as = a 7 A - q - S 7 7 a a > c - = 7 a Z a 7 > ~ ‘ 7 - 7 0 - = — 7 - - a a _ = 7 ~ — : 7 af - bea a a 5 ; a > 6 7 * _ - 7 7 a 7 = 7 = ee ao 4 - - S _ 7 - - ye a 7 a ~ ~~ - . a 7 : aan - : aD tN eas Te _ _— | _.¢ an : — ae Sy ie “any... = | a 7 So 7 7 - - - 7 =) =U - - _ : 7 _ = ; i : 7 - Se 7 = a = = ae 7 sa » a a =< + = ie 5 =. =e : = oo : Th _ — 7 >= ores 064 7 7 = ~ i. 7 6 : : 7 7 : 7 7 7 = 7 - ss ao 7 7 : =_— 7 7 7 7 7 oe ae - a - - - f - _— Ss + a _ a oe os “= - - : 7 - iL ~< y 4 aon cae Var. ; er 7 G <= ‘es ia = 7 7 7 a = 00) Sas - : a a a i = - ; yy oe — @ 7 re : 7 _ - vs Dex : a - - 7 ae - me - . A a - : = eS a - vr _——— = a i. ¢ a : a, a a 7 - 7) : aa ate _ = > 7 - a 2 ; a> oe an - 7 a : Ss A ; 7 G 7 : - : i 2 } 7 10) a _ - - - 7 _ a = a - E - . - ho oo = 7 ae - oe : a 5 7 = an _ ys > 7 : _— + oe - - - 0 - a = _— : - ey a OO a _ - - a7 = -_ a a Gane © — . i 7 = c ; - a 7 ae : a, - 7 7 - 7 7 7 a ee, Se een oe - — ot rT 7 a @ Se - : 7 _ os > 6 a - a : % Bal = . - - a es - a aa e a - = Ga a aa > 4 - : : ay - _ 7 ; —— ; ; - i <« ; — a ; a an = wae. = oo oe 7 = ab Se a 7 cin on a — = ee nw ae a + | sel me a ae le ee ee ee ee Oe eee 2 SF =F 7 - Oo : - . - oo. 7 ae - TT oe 7 7 - 7 . _— _ eee SS ws) sr — = ee ae > = : : : - -_ - , oe my - - = _ 5 7 7 aoe De _ + - a - == # ay 7 - 7 ae {= 7 7 - - : 2. 7 < ORs (see a ae : _ 7 - == vo a . a : 7 - _ 7.-= 7 = = 7 a 7 7 - - os oe f Oo se : : = 7 — 7 - a a -_ in a i * i ae & * < a = en yy 2 eo = a ae - ; ; { fl ae me ae 7 : = “6. a : 7 > a - —_ oa - - _ _ 7 : an . 7. < ; : ; ie - — a a, 7 - -_ -— 7 nt , = _ oe - Oo i 7 ‘a 7 7 7 a — - Oo | : > — 2 es) = 7 a - a ry - - a7. rr _ mt : 7 - - : —» SO a 7 = ” A ne 7 i a hi a - Bull. nat. Hist. Mus. Lond. (Bot.) 27(1): 11-29 Issued 27 June 1997 A phylogenetic conspectus of the tribe Hyoscyameae (Solanaceae) K ¥ a Be $ Wo Lt \ y ALISON L. HOARE Centre for Economic Botany, The Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB SANDRA KNAPP Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD CONTENTS VGu(teova LNeLS C0) tere echt pA ne ere ef CeCe ey ES MS Cer eager oe roo ES co non Roe ere ee 11 Histonyion classifica ttomcscera-cc-sccceeteeececes ccastcotiocexsassoccerssts toc eeetesoce restess suctsetecod svestictscucet desc faeatiety cercer ese ec yecaivestassneeanece tiesto 13 @hemistryiand ECONOMIC DOAN yaa eco casceceaz cers te tere see eat eae eee ace anaes se aod evise na ccoas cscs Fac cs seee ceetusecsesseaseat vesceuttantectscsatectstrt oe 14 CE TMC T AIS Ey teers cree cscs eat ac hee cese oo cs teeee cas acces ae nyt tae RO etn sees eet Seon bree et ac seo sras Tee neee attest aet te recet at resweae seer Tetesacgssusestacees 14 ECOMOMMNG; DOCAN Yates crac tsssrecssrazezcscse rts cron taste ses sceesbe sAcetae este cae scans sete es sonasae actu tnaanenaeepsseneeveot cones at eas soreents testtnteet ecetcczecacaeat 14 Meth odSiarsszsastarecesccetace testassce ts cuts sesovsas esc Sosces sot osc ieee cae sas tuse cea ees ovsaae ceene eater eaes poset haat eat as estcopaces satires ehaes scbeteas tostaticarcaetans JBengets 15 Cy Yo) Celeat 0) By 6:8: HOES eA AR pe i Bee Bree RN ar rR TOT T yD ROP OO ee oY ier Oe Sor Darr ee D 15 Sn arc te rg aoe, srs cca cs sera dees has cen re Sea vcs oe roe Sea ce OSTUSe Tans Tate RUE eT ROTA ou aan Yast tte Seater on Mes TROTS SA GEASS Faasees snipes sfensgatteneces® 15 TPRESICONSUEUCCIOM | isis czscccszesssces sce czececsat snececa ste sae cc ores aoe tene Soe Oe ote n eee e wes Pace sabe codes bua nsssetersrisactisesscacaiyaitsusesns feeateeucccysecesecen soe 16 RESUIES! ani CISCUSSION fs sscaccescosct aca reece eae nee eee ea eee cette SRL ELE E Nee Mean Det gadsaneteMteecs ck iataaeserstet ese 16 IES LODOLO GY sc ccncetsctesceas tase seas sacs saces cose setae ose dat cto te canes Joie Seutana esac eric cate cept ont tan Seats cnde ante teh atoa eae surest eseceesscesstressuesecses> 16 Gharacten anal VSiS ee = 3 cm) 2 Nn khwWN 16 Corolla 7. Corolla: flared 0; campanulate-urceolate 1; cup-shaped 2 8. Corolla: actinomorphic 0; zygomorphic 1 9. Corolla: uniform in colour 0; tube darker than the limb 1 10. Corolla lobes: quite deep 0; shallow 1; very deep 2 11. Apices of corolla lobes: rounded 0; acuminate | 12. Length of corolla: short (< 2 cm) 0; medium (2 cm < x <5 cm) 1; long (> 5 cm) 2 13. Width of corolla tube: narrow (< 1 cm) 0; broad (>1 cm) 1 Stamens 14. Filament insertion on the corolla tube: near middle 0; basal (< 1/3 of way up) 1; apical (> 1/2 way up) 2 15. Stamens: exserted from corolla tube 0; included | 16. Stamens: regular 0; declinate | Stigma 17. Stigma: included in corolla tube 0; exserted | Fruiting calyx 18. Fruiting calyx: cup-shaped 0; ovoid 1; flared-urceolate 2; tubular 3 19. Fruiting calyx: little enlarged 0; inflated 1 20. Fruiting calyx: without prominent ribs 0; with prominent ribs 1 21. Fruiting calyx: membranous 0; subcoriaceous 1; coriaceous to woody 2 Fruit 22. Fruit : berry 0; capsule 1 23. Fruit: globose 0; cylindrical 1 Leaves 24. Leaves: ovate 0; cordate 1; lanceolate 2 25. Leaf margins: entire 0; variously incised 1 26. Leaves: arranged along the stem 0; a basal rosette 1 Seeds 27. Seeds: compressed 0; not compressed 1 28. Seeds: rectangular-subreniform 0; reniform 1; sublenticular 2 29. Seed size: medium (2.5—5 mm) 0; small (< 2.5 mm) 1; large (> 5 mm) 2 30. Seed colour: light brown or mustard yellow 0; dark brown 1 31. Spermoderm cells: deep 0; shallow 1 32. Walls of spermoderm cells: sinuate 0; straight 1 Trichomes 33. Glandular hairs: absent 0; with uniseriate glands only 1; with uni- and multi-seriate glands 2 34. Eglandular hairs: all simple 0; dendritic hairs present 1 Pollen 35. Number of apertures: three 0; none 1; more than three 2 36. Length of apertures: long, almost meeting at the poles 0; short 1 37. Tectum: present 0; absent 1 38. Supratectal ornamentation: absent 0; isolated elements-scabrate 1; scabrate with gemmae 2; reticulate or striate 3 Chemistry 39. 3-tigloyloxytropane: absent 0; present 1 40. Belladonnine: absent 0; present | 41. Tropine: absent 0; present | Table 3 Data matrix used in the cladistic analysis. Character 0123456789 1111111111 2222222222 3333333333 44 Number 0123456789 0123456789 0123456789 01 Lycium 0000000000 0000000000 0000000000 0000000000 00 Anisodus 1021001100 0011110011 1210000110 1000121020 01 Atropa 0020101100 0011111100 0000000121 1101000031 11 Atropanthe 1020001100 0011111111 0110000100 0110000030 01 H. muticus 2111201011 0010000121 0211010001 0101100011 01 H. niger 2111001011 0010000121 0210010001 0102000011 01 2010001010 2011000121 0211010001 0102000011 01 1020111200 2011010000 0000201110 1110017111 10 1020101000 2011110000 0000101112 0000017101 10 H. senecionis M. caulescens M. officinarum P. physaloides — 0000000000 0000000111 0110100001 0002020011 01 P. praealta 0010000000 1110100121 0111100001 1002000031 01 Przewalskia 1000011000 0010210111 1210201100 1001000001 01 Scopolia 1021200000 1101110011 0110000110 0110071011 O01 A.L. HOARE AND S. KNAPP Pollen for this study was taken from herbarium specimens at The Natural History Museum (see Appendix I), except in the case of Scopolia carniolica, in which pollen from live plants was studied. Pollen grains were prepared using the procedure of Erdtmann (1960). Acetolysed pollen grains were studied under the SEM and the light microscope. All chemical characters were taken from the literature, largely from Tétényi (1987) and Romeike (1978). Tree construction Cladistic analyses were undertaken with HENNIG86 (Farris, 1988) using the ie* option (implicit enumeration) with all characters unordered. Tree statistics generated from HENNIG86 include the tree length (L), the ensemble consistency index (CI), and the retention index (RI). The ensemble consistency index (CI) is a measure of character fit on a scale of 0 to 1 and the ensemble retention index (RI) is the fraction of apparent synapomorphy in a character that is retained as synapomorphy on the tree (Farris, 1989). The genus Lycium was chosen as the outgroup for this analysis based on the cpDNA phylogenies produced for the Solanaceae in which Lycium is the sister group to the clade containing Atropa and Hyoscyamus (Olmstead & Palmer, 1992; Olmstead & Sweere, 1994; Olmstead et al., in press). RESULTS AND DISCUSSION Tree topology The HENNIG86 analysis produced a single most parsimonious tree of L=109, CI=52, and RI=58 (Fig. 2). The group can be broadly divided into two clades: the Hyoscyamus + Physochlaina clade and the rest of the genera (see Fig. 2). The Hyoscyamus + Physochlaina clade is defined by the following suite of synapomorphies: cordate leaves (character 24), very small seeds (character 29), glandular hairs with uniseriate and multiseriate glands (character 33), and pollen with isolated scabrate supratectal elements (character 38). A close relationship between Hyoscyamus and Physochlaina has been pointed out by previous authors (Lu & Zhang, 1986). The genus Physochlaina is not, in our tree, a monophyletic group. This could be used as evidence for the suggestion that this genus should be reduced to synonymy within Hyoscyamus. The position of P. praealta does agree with the observations of Zhang & Lu (1984) who suggest this species may be intermediate between the two genera. The characters linking P. praealta to Hyoscyamus include the shape of the calyx and the fruit. However, if the pattern of branching is altered, such that the two species of Physochlaina do form a monophyletic group, then the tree is only one step longer. Clearly, this analysis does not provide convincing evidence either for the lumping of these genera or for keeping them separate. The clade consisting of the rest of the genera in the tribe is defined by the following suite of synapomorphies: a racemose inflorescence (character 0), a medium length calyx (character 6, also present in Hyoscyamus) and corolla (character 12, occurring in many places on the tree), basal filament insertion (character 14), stamens included in the corolla tube (character 15), and non-compressed seeds (charac- ter 27). Within the clade, the only consistent groupings are Scopolia + Anisodus and Atropa + Mandragora (see p. 19). Relationships of the other genera are somewhat ambiguous, although Przewalskia is basal to the clade. Contrasting this with the proposed phylogeny of Lu & Zhang (1986) illustrates an important aspect of cladistics which distinguishes it from other purportedly phylogenetic ap- proaches. Lu & Zhang (1986) hypothesize that Przewalskia is the PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE 1718 1921 223941 TLALA ae NF i ‘Lycium 0000000 35 3 . ’ ¥ Physochlaina physaloides 2 24 29 33 38 10 11 143038 f-§42 BEE se : evden errs Physochlaina praealta . i? ae ae oe 212182 # 3 H4 10 13 pee Hi ‘Hyoscyamus senecionis 21 0 6 821242531 43334 2112011 HH Hyoscyamus muticus 139 arhet Ras gaa He Oe 3B f Hyoscyamus niger 0 $ 142021 24 26 3033 Przewalskia Vi 8 eee ee Ts Tot | 16323839 Bar oe iia ttt Atropanthe BESEESE 1953-0 | eet es lew | 20 21 31 3435 38 39 ‘Anisodus Aerie pe 432% Bn 3 3536 SEaaaE see 2 kal SEaEe PUNT 4 6 71011123032 Scopolia 20011001 172830 ILE UuUaE O11 O 16 17 28 29 33 38 Atropa O81 21-2"hie3 41819212240 S$ 7142432 100001 HHH Mandragora caulescens 102426353741 | 12921 BREBBeBEh SESE EU 2111107 429909138 ‘Mandragora officinarum 02000 Fig. 2 The single most parsimonious cladogram from the HENNIG86 analysis. The characters are discussed in the text, and character states are shown in Table 2. For characters marked on the branches of the cladogram: unshaded marks indicate synapomorphies, stippled marks indicate reversals and parallelisms (homoplasy), and solid marks non-homoplastic synapomorphies. A.L. HOARE AND S. KNAPP PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE 19 Fig. 4 a) Fruit of Anisodus tanguticus (photograph courtesy of M. Gilbert), b) Fruit of Atropa belladonna, Chelsea Physic Garden, c) Fruit of Hyoscyamus niger, Chelsea Physic Garden, d) Fruit of Physochlaina orientalis, RBG Kew. most ‘advanced’ and thus, according to them, derived, taxon of the group. However, in our tree, this genus usually occurs at the base of the clade. This radical difference in placement is most likely due to the large number of autapomorphies (see Fig. 2) which distinguish Przewalskia. Automorphies are not informative about relationships. Atropa and Mandragora cluster together and are nested well within the tribe as a whole. This is in agreement with Tétényi’s (1987) system. Earlier classifications had not included Atropa or Mandragora within the tribe simply on the basis that they bear berries and not capsules. Characters which unite the two genera include those of the fruiting calyx (shape, inflation, and texture), fruit type, and the presence of belladonnine. The close relationship of these two genera was implied in the classifications of some of the early botanists: Miers (1849), Dunal (1852), and Bentham (1876) all placed these two genera in the same tribe or subtribe. This link is also reflected in the naming of some of the species, for example, Linnaeus had described Mandragora officinarum in Species plantarum (1753), but in later editions he changed the name to Atropa acaulis (1762, 1764). Fig. 3 a) Flowers of Physochlaina orientalis, RBG Kew, b) Flowers of Hyoscyamus niger, Chelsea Physic Garden, c) Flowers of Scopolia carniolica, Chelsea Physic Garden, d) Flowers of Anisodus luridus, RBG Kew, e) Flowers of Mandragora autumnalis, RBG Kew, f) Close-up of the flower of Atropa belladonna showing declinate stamens, Chelsea Physic Garden (photograph courtesy of J. Vogel). 20 A.L. HOARE AND S. KNAPP Fig. 5 a) Fruiting plant of Przewalskia tangutica (photograph courtesy of M. Gilbert), b) Fruit of Przewalskia tangutica showing dehiscence (photograph courtesy of M. Gilbert), c) Fruit of Mandragora officinarum, RBG Kew, d) Fruiting plant of Mandragora chinghaiensis (photograph courtesy of M. Gilbert). Character analysis Rather than discussing in detail all of the characters and their distribu- tion on the tree, we have chosen a few to discuss in detail. Some of these are characters that show little homoplasy on the tree, while others are those that surprisingly do not provide any phylogenetic evidence. Morphological characters 3.CALYX SYMMETRY. Anactinomorphic calyx is plesiomorphic in the tribe. This finding is contrary to studies completed for the family as a whole, with the most basal members possessing zygomorphic flowers (see Olmstead & Palmer, 1992; Olmstead et al., in press). Zygomorphy occurs twice on the tree, once in Hyoscyamus and as a synapomorphy uniting Scopolia and Anisodus. In Scopolia and Anisodus the calyces are symmetrical except in their lobing. In Scopolia, one lobe is usually enlarged (Fig. 3c), and in Anisodus the lobing is very irregular (Fig. 3d, 4a). However, in the two species of Hyoscyamus, the asymmetry is manifested differently. In these species the calyx tube is curved, and the lobes show a gradation in size. The zygomorphic calyces of Hyoscyamus should probably not be equated with those of Anisodus and Scopolia. A more meaningful coding of this character may be to represent these as two independ- ent states, or it could be divided into two characters; symmetry of the calyx lobes and of the calyx tube. 7. COROLLA SHAPE. A flared corolla is the plesiomorphic state for the tribe. Within the Hyoscyamus/Physochlaina clade, this character is conservative. In contrast to this, in the rest of the tribe, corolla PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE shape shows much homoplasy. It was difficult to divide shape into discrete states, although the cup-shaped flowers of Mandragora caulescens were quite distinct. Shape may in fact represent several independent characters, and so would be better coded as such. The problem then lies in determining just what these characters should be. 14. FILAMENT INSERTION. Lu & Zhang (1986), in their study of the Chinese members of this tribe, presumed that stamens inserted at the base of the corolla tube were a primitive feature. This analysis suggests that this should actually be regarded as a derived trait, occurring in two places on the tree. Basal stamen insertion is a synapomorphy of the Przewalskia clade (uniting all the genera save Hyoscyamus and Physochlaina) and occurs in Physochlaina praealta. 16. ARRANGEMENT OF THE STAMENS. _ Declinate stamens are found in both Atropa and Atropanthe. Consequently, this character is not indicative of shared ancestry and, in this analysis, is uninformative about generic relationships. It is possible that this feature is an adaptation for improving pollen deposition on insect pollinators. The flowers of these genera are held horizontally or are nodding, and the position of the anthers is such that they would brush the bodies of insects entering them (see Fig. 4f). 19. INFLATION OF THE FRUITING CALYX. An inflated fruiting calyx (Figs 4c, d, 5a) is a synapomorphy of the tribe in this analysis. The non-inflated calyx found in Atropa and Mandragora (Figs 4b, 5c, d) must therefore be considered to be derived within the tribe and is probably related to the method of seed dispersal in these plants. Both these genera produce fleshy or juicy berries. Those of Atropa are black and shiny when ripe, and are eaten by birds. The fruits of Mandragora are whitish, greenish, yellow, orange or purple, and are borne close to the ground, frequently hidden by leaves. These are eaten by small mammals or reptiles. In the other members of the tribe seed dispersal is effected by the seeds being shaken from the capsules. The fruits of Przewalskia become detached and behave as tumble weeds, releasing seeds as they are blown about by the wind (M. Gilbert, pers. comm.). 21. TEXTURE OF THE FRUITING CALYX. A subcoriaceous calyx is plesiomorphic in the tribe. Atropa and Mandragora are united by having membranous calyces. This seems to provide some support for the suggestion that the calyx has a protective function in other members of the tribe. Thickening of the calyx occurs in three places on the tree, in Anisodus, Przewalskia, and as a synapomorphy of Hyoscyamus. 26. LEAF ARRANGEMENT. A rosette arrangement of leaves occurs twice on the tree, in Przewalskia and Mandragora. The rosette habit has been thought to be an adaptation to habitat (Lu & Zhang, 1986), providing protection from wind and grazing animals, and ensuring maximum exposure to the sun. Przewalskia is found at high altitudes in arid grasslands and areas of frost heave (Fig. 5a), while Man- dragora (see Fig. 5d) occurs in a wider range of habitats, which include stony slopes and screes in mountainous regions, grassland, and ruderal habitats. 27. SEED COMPRESSION. Compressed seeds are confined to the Hyoscyamus + Physochlaina clade, and so supports the division of the tribe into two groups. Non-compressed seeds are a synapomorphy of the other clade and are generally uncommon in the family (see Knapp, 1991). This character may be related to the development of the seeds, and possibly to seed dispersal. 29. SEED SIZE. Small seed size is a synapomorphy of the Hyos- cyamus + Physochlaina clade, but is also found in Atropa. Small 21 seeds are often associated with capsular fruit in the Solanaceae (Souéges, 1907), but in the Hyoscyameae this ecological distinction appears not to hold true. 30. SEED COLOUR. In contrast to the genus Solanum and other tribes in the family (see Knapp, 1989; Knapp et al., in press; Knapp & Helgason, in press) seed colour provides little support for the tree topology, and has frequently changed in the Hyoscyameae. The explanation for this may lie in the function of the seed colour, or may reflect the ease with which these changes can occur. 32. SPERMODERM CELL WALLS. Whether the cell walls of the spermoderm are straight or sinuate is uninformative with respect to generic relationships, since straight walls are autapomorphic in both Scopolia and Mandragora caulescens. This is in contrast to the utility of cell wall shape as a character in studies of other groups of Solanaceae (Knapp & Helgason, in press). 34. PRESENCE OF DENDRITIC HAIRS. Dendritic hairs occur in both clades, in Physochlaina and Anisodus. This high level of homoplasy is also found in many other groups of Solanaceae (Knapp, 1991). Trichomes can have a multitude of functions, for example, protec- tion against desiccation, ultra-violet radiation, and insect attack (Metcalfe & Chalk, 1983). Dendritic hairs, in comparison to simple hairs, may have advantages in any of these roles but no evidence exists for an adaptive role. Pollen characters 35. NUMBER OF APERTURES. Triaperturate pollen grains are plesiomorphic in the tribe, as they are thought to be for the family Solanaceae and dicotyledons in general. This is contrary to the suggestion of Lu & Zhang (1986) that inaperturate pollen grains are ancestral in the Hyoscyameae. They (Zhang & Lu, 1984; Lu & Zhang, 1986) identified the pollen grains of Anisodus tanguticus, A. luridus (as A. mairei), A. carniolicoides (as Scopolia carni- olicoides), and A. acutangulus as inaperturate, while our results clearly show (Fig. 7b) the grains of A. /uridus to be porate. The potentially cryptic nature (see Mandragora p. 27, and Diez & Ferguson, 1984) of the apertures in the pollen grains of Anisodus needs further study. The two species which have pollen with more than three apertures, Anisodus luridus and Physochlaina physaloides, are unrelated. Increase in aperture number must therefore have occurred independently in these two taxa. Distribution of apertures in these two taxa differs radically. In A. /uridus the six apertures are distributed randomly on the pollen grain, and the grains are pantoporate (Fig.7b), whereas in P. physaloides the apertures are confined to the equator (Fig. 6a). The principal functions of the pollen aperture are protection, harmomegathic responses (alterations in form accompanying changes in pollen grain hydration, see Blackmore & Barnes, 1986), ion exchange, and pollen tube germination. The significance of the number of apertures to each of these functions is unclear. Increase and irregularities in number of apertures does seem to be related to polyploidy, but cytological information is lacking for both these species (see Table 4). 36. LENGTH OF APERTURES. The presence of pores, rather than colpi, is a synapomorphy which unites Anisodus and Scopolia (see Fig. 7a, b). Reduction of aperture size may have occurred in re- sponse to an increasingly arid environment, reducing the risk of desiccation of the pollen grains. However, the ecology of these genera does not fit in with this, as they are typically plants of moist environments. This highlights the fact that we should be wary of making simplistic explanations about the adaptive significance of i) i A.L. HOARE AND S. KNAPP Fig. 6 Pollen morphology of the Hyoscyameae. Scale bars beneath photographs. a) Physochlaina physaloides, b) Physochlaina praealta, c) Atropanthe sinensis. PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE Fig.7 Pollen morphology of the Hyoscyameae. Scale bars beneath photographs. a) Scopolia carniolica, b) Anisodus luridus, d) Atropa belladonna, d) Mandragora caulescens. characters. Multiple functions of characters considerably compli- cate the issue. This is undoubtedly true for pollen apertures, where complexity of structure and a variety of alternative strategies make simplistic adaptive explanations unrealistic. 37. OCCURRENCE OF A TECTUM. The only taxa in which intectate pollen occurs, are the two species of Mandragora. This type of pollen is believed to be primitive for the angiosperms as a whole (Zavada, 1986). Lack of a tectum is perhaps related to the cryptaperturate condition in Mandragora (Diez & Ferguson, 1984). The structure of the ectexine affects the physical properties of the wall, and this must influence the durability of the pollen and the exchange of materials across the wall. 38. SUPRATECTAL ORNAMENTATION. There is little concurrence between this character and the tree topology. This may be for two reasons. One possibility is that the ornamentation of the pollen grain may not be useful in revealing phylogenetic relationships in this group because of parallel evolution. This analysis does suggest that pollen with a reticulate pattern may have arisen independently in 24 Table 4 Chromosome numbers which have been recorded for the Hyoscyameae. Species Chromosome number Reference Atropa belladonna 2n = 72 Vasudevan, 1975 Hyoscyamus muticus 2n = 28 Al-Musawi, 1979 Hyoscyamus niger 2n = 34 Al-Musawi, 1979 Hyoscyamus senecionis 2n = 34 Al-Musawi, 1979 Mandragora autumnalis 2n = 96 Murin, 1978 2n = 84 Hawkes, 1972 Physochlaina praealta n=41 Vasudevan, 1975 Scopolia carniolica 2n = 48 Vasudevan, 1975 Hawkes, 1972 three lineages. The selection pressures which may have led to this are not known. The function of pollen sculpturing has been thought to be related to the pollen vector (Hemsley & Ferguson, 1985; Ferguson & Pearce, 1986), although in some groups of plants there is no apparent correlation between the vector and ornamentation type (Thanikaimoni, 1986). An alternative explanation is that the delimitation of the character states is not meaningful. This seems likely in view of the difficulty in deciding on character states due to uncertainty about pattern homology. For example, pollen grains which were scabrate and those with isolated granules were coded together, but perhaps these should have been coded separately. Pollen of Anisodus was unique in being scabrate with isolated gemmae (see Fig. 7b). This was coded separately from simply scabrate pollen. The importance of this distinction is unclear. Inves- tigation into the development of sculptural patterns may shed some light on this area, and so help in the interpretation of changes in ornamentation type. Chemical characters 39. PRESENCE OF 3-TIGLOYLOXYTROPANE. Prescence of this com- pound is a synapomorphy for the tribe, but the distribution of the character on the tree is homoplasious. Either the ability to produce this compound has arisen up to four times in the tribe, or there have been reversals in Anisodus and Atropanthe. 40. PRESENCE OF BELLADONNINE. This character is a synapomorphy of Atropa + Mandragora. 41. PRESENCE OF TROPINE. The occurrence of tropine is a synapomorphy of the tribe in this analysis. However, there has been a reversal in this character in Mandragora. Lu & Zhang (1986) identified a number of characters which they considered to be ‘primitive’ for the tribe. Among these were actinomorphic and solitary flowers, stamens inserted at the base of the corolla tube, and inaperturate pollen. They provided no explicit reasoning for their choices, and tried to place taxa on a gradient of ‘advancement’. This analysis suggests that many of these features should be viewed as derived within the tribe. This draws attention to the futility of deciding a priori on criteria of ‘advancement’. Deci- sions based on phylogenetic analyses rather than on intuition can be more easily justified. The diversity of pollen types in the Hyoscyameae means that such characters cannot be used as synapomorphies of the group. Simi- larly, this source of data provides little information about generic relationships in the tribe. The differences in pollen morphology between some of the genera have been used in the past as evidence for their continued recognition (Zhang & Lu, 1984; Sandina & Tarasevich, 1982). Thus, the separation of Scopolia, Atropanthe, and Anisodus is supported by the palynological evidence. However, this is insufficient evidence on its own, because similar levels of varia- A.L. HOARE AND S. KNAPP tion are found within other genera such as Physochlaina (Zhang & Lu, 1984). There is no justification for emphasizing one source of data at the expense of others. Chromosome numbers have not been used in this analysis, but may perhaps be of use in future work on this group. Initial work on the cytology of these plants suggests that this may be informative. The chromosome counts which have been completed are listed in Table 4. A chromosome number of x=12 is widely held to be primitive for the Solanaceae, with aneuploid reduction to x=7 in many lineages (Goodspeed, 1954; Olmstead & Palmer, 1992). Poly- ploidy is common and possibly has been an important factor in the evolution of the tribe. Whether ploidy levels in the Hyoscyameae are due to alloploidy or to simple chromosome doubling is not known. Biogeography The biogeography of the Hyoscyameae is of great interest because it is the only exclusively Eurasian group in the family, the rest of which is largely Gondwanan in distribution (see Symon, 1991).Two theories have been proposed for the origin of the Hyoscyameae. Lu & Zhang (1986) drew attention to the diversity of the tribe in south-western China. Eleven of the forty species of the Hyoscyameae are found here, five of which are endemic to this area (see Fig. 1). On this basis, they concluded that this area was probably ‘the birthplace of hyoscyaminous plants’. However, there is a fundamental flaw in equating the centre of origin of a group with its centre of diversity: centre of origin arguments are often flawed (Humphries & Parenti, 1986) and these dispersal hypotheses always require external, often ad hoc, causes to explain patterns. Linking distribution with the history of the earth has proved a powerful method for understanding the processes that influence the patterns we observe (Nelson & Platnick, 1981; Humphries & Parenti, 1986). An alternative scenario to the centre of origin idea of Lu & Zhang (1986) was proposed by Symon (1991). He considered the distribution of this tribe to be consistent with the ancestral group being rafted north on the Indian plate. On meeting Eurasia, the group evolved in the developing Himalayas, and subsequently spread from there. The geological history of the area occupied by members of the Hyoscyameae is remarkably complex. Although the group is largely Eurasian in distribution at present, the main areas occupied by the genera of the Hyoscyameae were once part of the Gondwana supercontinent. Much of South East Asia, including southern China, consists of terranes rifted from the margins of eastern Gondwana some time during the Jurassic (Hallam, 1994). The southern part of Tibet in the Himalayan mountains is thought to have been the southern margin of the Tethys Sea or still moving to collide with the Laurasian supercontinent in the early to mid Jurassic, while the northern part of the area is more consistent with a non-Gondwanan, Eurasian position. The Lhasa block collided with the other rifted terranes in the late Jurassic to early Cretaceous (Dewey, 1988). Apulia, including Turkey and present day Italy, was connected to Africa during the early Cretaceous, and rotated to collide with Eurasia about 80 million years ago (Hallam, 1994). The Indian plate is thought to have broken away from the Gondwanan land mass last of all, some time near the Cretaceous/Tertiary boundary between 65 and 60 million years ago (Hallam, 1994). The Eurasian-African collision closing the seaway to the Indo-Pacific occurred in the early Miocene, and brought the Middle East into contact with the major land masses of Eurasia. Climate change during the Neogene was probably important to the evolution and distributional patterns of land plants in these areas. The Himalayas have continued to uplift long after the initial collision event between India and Asia (Hallam, 1994) and this may have been a major factor in the general cooling of climate in the Neogene. The general pattern of the break-up of the PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE continents over geological time is thus consistent with the Hyoscyameae being a primarily Gondwanan group which has radi- ated extensively in Eurasia in more recent times. The two main clades within the tribe have broadly overlapping distributions centred in the Himalayan and South China area. Distri- bution patterns within the Hyoscyamus/Physochlaina clade are somewhat confused due to widespread human utilization and distri- bution of these species. The genus Physochlaina is found in much of China, reaching north into Siberia and as far west as the Himalayas. Species of Physochlaina are predominantly plants of montane habi- tats. A possible scenario is that this genus arose with adaptation to high altitudes or to colder climates. This may have occurred during the development of the Himalayas, or alternatively this group might have evolved at low altitudes during later glaciations in the Quater- nary. Hyoscyamus shows a much wider distribution and ecological amplitude. The majority of the species occur in the Middle East and the Mediterranean region, and it has been suggested that its occur- rence in North Africa and northern Europe is a result of human dispersal (Symon, 1991). All species of Hyoscyamus occurring in India occur in Kashmir and the north of the country, perhaps lending support to the idea of a very early origin for the group. Adaptation to more arid and mediterranean climates seems to have occurred in many of the species. This may have arisen as the genus spread into more arid areas or during a period of increased aridity. Such condi- tions are thought to have developed in central Asia following the uplift of the Himalayas and the Tibetan plateau (Manabe & Broccoli, 1990). However, any hypotheses of biogeography and evolutionary history of Hyoscyamus will need to be tested using an in-depth phylogenetic analysis of the entire genus. Within the other clade of the Hyoscyameae, similar ecological factors seem to have been important. The genera in this clade are largely isolated ecologically, growing in different elevational ranges and often in quite different microhabitats. Przewalskia is a narrow endemic from the Qinghai-Xizang Plateau of western China and Tibet, growing between 3200 and 5000 m. The evolution of Przewalskia may have occurred during the uplift of this region (Lu & Zhang, 1986). The timing of the Himalayan orogeny is uncertain. One theory is that the main orogenic events occurred in the Oligocene, at the same time as the Himalayas developed. Alternatively, uplift may have continued well into the Pliocene and palynological evi- dence supports this (Ruddiman et al., 1989). The prolonged uplift of the Himalayas (Ruddiman et al., 1989) and the concomitant expan- sion of grassland habitats at the expense of forests will have had a profound effect on the evolution of plants found in these areas. The widely disjunct distribution of Scopolia (see Fig. 1) suggests that it was once more widespread. This explanation is favoured over one of long-distance dispersal because capsular fruits and small unornamented seeds, which are found in these plants, tend to be locally dispersed (Olmstead & Palmer, 1992). Lu & Zhang (1986) suggest that this genus was widespread during the Tertiary, but became much more restricted with the advance of ice-sheets over the continent during the Quaternary. Atropa and Mandragora are both very widespread genera, occur- ring from southern Europe across to the Himalayas and the mountains of western China. Their animal-dispersed fruits and their wide- spread human use may have helped to expand their ranges considerably. The analysis presented above is a first attempt at a complete phylogenetic classification of the Hyoscyameae. The resultant cladogram shows that two lineages can be identified within the group, one clade consisting of Hyoscyamus + Physochlaina and the other containing Przewalskia + the rest of the genera. Within the Przewalskia clade the relationships of the genera are somewhat 25 ambiguous, although Anisodus + Scopolia and Atropa + Man- dragora always group together. The position of Atropa and Mandragora, clustering well within the tribe, provides evidence for including them in the Hyoscyameae and supports Tétényi’s (1987) grouping. This is further corroborated by their distribution, since they form a phytogeographically coherent group with the Hyoscyameae (Symon, 1991). GENERIC CONSPECTUS This conspectus is intended as an overview of the taxonomy of the genera, but may require revision as more species are studied in detail. The synonymy has been taken in large part from recent floristic or monographic treatments, which are acknowledged and cited as part of each description. Much work remains to be done with the taxonomy and phylogeny of each of these genera and we hope that this conspec- tus will help future workers in these groups. Distributions for each of the species are given in general terms. More complete descriptions, especially for species occurring in China, can be found in the floristic works cited. Artificial key to the genera of the Hyoscyameae 1. Fruita fleshy or juicy berry, white, green, yellowish orange, purplish or 10) Ye) ae cee attra aire hee pei ines pros a Aria canine Raa Rent na Beta nae 2 2. Berry white, green, purplish or yellowish orange at maturity, fleshy, usually held beneath the leaves; acaulescent (occasionally shortly caulescent) herbs with enlarged tap roots; flowers deeply lobed, ACUMOMOLP MIC \ececccesscc-cacesscectuceecescscestccteveseeossasesssvaests 5. Mandragora Berry black and juicy at maturity; plant an erect perennial to | m tall; flowers shallowly lobed, the stamens declinate .................. 2. Atropa 3. Flowers solitary or in short inflorescences of 2-3 flowers .............. 4 Flowers in elongate or branched inflorescences, the inflorescence usu- ally withimore*than:S TOWEDS Pei cn = a : Bt 7 _ _ - 6 _ a - _ a a ; ; : : : ; : 7 a : 7 a - 7 - oo 7 = 0 - - : 7 3 i 7 - - -— m~ ~¢co-a me > : a : : ee ey ea —_ — _ a re ee a a ‘es *) > ae oe : — —— ce a c= - ~G . aa — : o : : _ - a ‘ 7 : : 7 S ; 7 : 53 ae - SS See ee ee ee ( _ _ _ a _ - i‘ 7 7 -_ ea 7 : a _ a eo ¥ 7 _ — on: s, ‘2 a _ ae ih aay al ia are nn ome ~~ | =f. —— | (ne ee Se, Re 5 cn i os ne > : ’ onary : x aes 8 - _ aa s 2} a) 7 = - 7 7 Y Z - > ia _ : ar 6 (om oo - a en 7k ea =e ° — 7 7 7 - = = ot - af : oA Bit : . . y —s* 7. ™= oo - be as oe 7 vo c) 7 rl 7 > ; > = 7 7 7 7 ; 7 : _- _ : - - <> _ : ve a- = 5. b= oe 7 = = a os ¢ o <2 Gg e — > Xe) Some ae oe Se a = oe. + Watt 8 . ee hed — ee 7 a _ a - - Q > : a 7 : a. : —_ =~ i. fe ‘eo : : - I - : ar y —_ os a ——— 7 o 7 Uae ‘ - a 7 | - TJ . a i, 0 a ae — : —<_ | “Tomes “20 a 7 a 7 - a oa Ae ~~ = 7 SS wall 7 ‘ za a 7 : . 7 : . - rA ——- 7 - 7 ¢ - - : : 2 A rr) Ve Pa > ee OE _.. - re se a 7 ~ : : OG ee i> ee © oo «a 2 = a 7 ‘es. ; - 7 7 . : : : : ; aa.” CA sc. , - 7 ¢ @- =2- ? : - : - 7 —= ie - 7 — : oes = —: as - a 7 7 is a 57 ig . 7 7 : : 7 7 ea : 7 - i 7 = 7 = = _ 3 - ak 7 - = = 4 7 - 2 ~~ P= 7 ~~ a 7 7 a aaa _ : - 7 iy 4 a on - : iy oon 7 ; Oo _ = - Bs ——— 7 > a _ ee oa 7 <2 7 a a 7) : Ph i : 7 i 7 = - - = - 7 ead - mie TT 7 7 AG os ven 4 =. i >, — 5 7 7 - a ~ J " a a : . - i 7 i. ere : = 7 a = : nian _ : ; uly woe aA s33 an opus > © = - _ Po eee se = 0 >. 7 me _ SS am er ai = -e a - Bull. nat. Hist. Mus. Lond. (Bot.) 27(1): 31-73 A revision of Solanum section Pteroidea: Solanaceae Issued 27 June 1997 ohare e — SIN Rae) ‘9 \ Be SANDRA KNAPP. Department of Botany, The Natural History Museum, Cromwell Road, London SW7 SBD THORUNN HELGASON Department of Biology, University of York, PO, Box 373, York YOI 5YW CONTENTS Mm trO duct OM ssese Ase cde cesscete stu soccte ces eadaeaaea sae asbontcu'tinsacasccscpcnstecetsscsscsyscecseaisenteds daseass bets banated edsesstesassatoiesasasisesatassedssspassashacseeeses 31 slaxonomicrand:nomenclaturalthistory cee sccccta certs ce sccetceetae scarcer rete tnss setae ctet ners erteetaserse car siciewtstresnsvstsesticersistsetit esereserestes 32 HIStoryiOf SCCHON PICTON CC weccs care cance crccescistectty aceite deat acti Aieven sve vas ceriesevinsates tases tas voiav che saioes teri pstes eras tovss¥ contosesestsnssinetittes 32 Morpholopy andinaturall istry cccseccececscsces votes coos cs tatccttesstctretes cots versetsbeowessaencetessepreyinecteras-sicesstecsuscctstseneatsuraserietesisoteterssressst 33 SCE MMIS eases cree e erase pape tae ee es apa cere coe cn aes ne te coer ae een Cae she sac saire ss chats ep Ser tacestazeatet ent satcassstatancaasusrssstesse stedteqiseeeesttiaete 33 |r er Noh are gee pet ete re pero rreo erensoe eV aerurperar nme tiger iis Carer coo tthircaee aera eeteea cerca irenccn cc Penrna reer are rodcs 33 Mnf] OVeSCOMCESH Sosa cecsnshe snes tees aecisac nee oS SOE Rae soc Siegen ck Sette Aes eae oe DSR TOT Cae anon t Sec O ee MEA ee neU cen cTSEG Teas os aac ea daooes cveseses soe eweeeet 36 SETUCHOINIES axis Soeeescrsstas ces sees cdots ccevasasoecuse uate taceen can tascete eae De daad neat oUt estat ote kcs Mas canes: sete aroaesusmbeeaiotn ce ae taaobereseta ass Lessee tats 36 RTO WLS ass he oe se oars coe setae paecs fucvon eae tas ss opetic ea an ae cu cease coe ees cP s eases an ae nse hs Uane austere UMNG Saat sane tecWostes sneer ieioees santa aaraateeaees 37. |Sy DU Seopa eRe eee te Pre rars eatery serene eh marc Penn Re oe ce eee ce each eoere aes ea ae center cone Canton treo 37 SCC S re error acca senerete ce or etna neat ma mectasa cc caeste det cr wnat cress an sretencasaeencan feeau eer cere Suess Sic manasa aht os oa cant tase nig raeetavertie es 37 CG) Ee Sine een oar ere ners aoe ann on eRe een ene er eet eran Sete ack soe arena peer eeoreerocrre a eecos Terre La en EET 42 @haractercoding and tree COnSUUCtONesycsvescesscs cece eSees eecs vce: Sac eve sessaetucsysocesccaaestass ct tgrsessoactassectocsecttesacenaucducsae soeansancenee 42 GT aS Sift catia veces eerset ccs tees reece aren ecw eee erro ahs Nar ena eta eas whee es nctanet tests ceseas Ue cre ast caesssigbucsnacsercedeerssdeteceeeseee 43 TRAX OMOMIC treatemenitixessces sae ee aera acta aa este sea ceas ta Sard oot ca MOAI RDS NSE RET suse ssees guaaeessisseee sper esr rrese teats aecas 44 Key to selected groups of Neotropical non-spiny SOlanuMs ..................csscscscsscssssssssssscssssncsssnsssensetsorsrenenssaresacessassseoosereess 44 Key to species of Solelirn Section PICIOId eG Lit sac cc vances. sssarestvascovicoseskcpstaysnc tervasninessvesaenesteresedcraqusysuyqsseveessansnencacgesceorseqrrss 44 Whe SOlanumiternatumiSPECleS|OTOUP sce: recseres se saeseh acess sa cetaeee cesonsscttesceese tease veces tsctes ttsecerecs carers sspeartstontstcst-catses cesensrcnecdcees: 45 LER SOLENULINSEPCUNVILINIRULZ OA AV a eastern eee iam eee eerie a: see W ae Teme eee coer eakars sas eek sdctet se wkaseeysGSeece tsanrovecsteesscssaetestaatesa ers 45 DESO LANUNTLETTITUITIARUIZ (SC AV eee eae eee arene ast eco cea cre cece ey rete raes scene essa tue oMe cba tes suceecie lon sists since sued wtesect zeeseer= 45 Phe Solanum mite species, QUO wy is. co css 2 2ee ls haak 1Oskig ucsan ade cost peasant yeotgeunsesddstsvedanteterksis+xearsnedthesatdnersseoatearcnartateeendeowoss ntecrers Si Be Solanum anceps: RUIZ) SxR averse vtec cose tevesss testa cc etege snes ta ston Sesto eeeers pea eoeene wc eneacec ote artussasassescestscsrsagvacrsencnstcscevac= 51 4 SOlanurni GN ees tical aterm Be Deen ceccgecss cote crss coe tec aetese stn tei sarsees tae coceecane on cnescest aves soedscassesuctcursectntervure Seve cnratsacansiostns 54 5. Solanum chamaepolybotryon Bitter .........:..ccccccsssssssscsssecssetessenecsrsesseoretsesenescenensossssvoneseceesonsossasossssnsssesssssenesseteeeesaeates Ss) 6 SS OLANUM CONIC RUZ SCAN s Sacece cssairecscues Fee ce tate ea cek Sendo cacea ts Saas e atc as cent an ova sate eae sa ete TACT Sank see ne ores Scene sane sateen Sada aD Te SOlARUM MITE RUIZ GE PAVE cercece sree Haak aes Da ae ote seca tear oho saat ceeeae anion cSocees aaccesceneiders ssa acond secteneysecenteenaae= sactatastonss 58 RT COLANIMMESAVGNIMENSE Ble listarec cores eect stent cotee see peer trance cee sree edocs tira ne toaat cen s terse ceectoeet sence tetas ont canessates wostncstecste-ss 62 9. Solanum trizy gum Bitter .........scecscoosssseseseoosesonssssanssasassavtsautecsstscecrenscsacnsetensnasdecsnrseatnaccsososnesesosestosassaevesssoosseeneseseageaneees 64 VO Seolcantand Tale QUA BRET ve cnc cse ce ccass .doscspsco< cee oxcacausc cee see teasece otra ses tats tes teas vesn taeee cee sus cutasennestrsecressacepccsseisteeastsaeess3 68 Excluded SPeci€ ........ccsssecscecseererssseterssvescnssossscsesuseascnseeasarsecnsoeauesnssecssususoeseresnsnsssnsecsoasseuassssesuanesessesensnenenesnesucseseneasegensorsreetents 69 RRELETCTICES reese a eee eee eg TESS ee ore Rogan Shean nace e sea nde asa ensSev evs Trttae for auteTase eneticesrenet mane resaesatesesy 70 EXSIC CALC reer aN ee GT Ta oes Ba sea pate tee sae Sao Coney Facet Savee Sareea tant taet su oneenaas cease aveaneeser’ 71 J Fe Pe: ea ee Pp Se RD Rr eS rr eee a Cee ee cr TEP EY Oy eee Tc cece cre 1/5} SYNOPSIS. Solanum section Pteroidea is a small group of ten species of Neotropical primary forest herbs and vines. The group is treated in this monograph as a unit for convenience, but cladistic analysis shows that it is almost certainly not monophyletic. The ten species are therefore placed in two monophyletic species groups: the Solanum ternatum species group, defined by its woody vining habit and large flowers, and the Solanum mite species group, defined by its conical rugose fruits and ovoid-reniform seeds with distinctive testal morphology. The history of nomenclature and composition of section Pteroidea s.1. are discussed. Illustrations and distribution maps are provided and photographs of several of the species show characters of the flowers, fruits, and seeds. trated on groups of economic importance, such as potatoes, toma- INTRODUCTION toes, morellas, and the spiny solanums. The genus is diverse, with some 1000 or more valid species (D’Arcy, 1991), but monographs Although it is one of the five or six largest genera of flowering plants, do not exist for the majority of species groups in Solanum. As part of little monographic work has been done in Solanum L. (Solanaceae) an ongoing research programme into the taxonomy and phylogeny (see D'Arcy, 1991). Taxonomic research effort has been concen- _ of non-spiny solanums (see Knapp, 1986a; Knapp, 1989; Knapp, © The Natural History Museum, 1997 52 199 1a) we have investigated the small, primarily rainforest species of section Preroidea with the aim of determining the monophyly of the group and the species boundaries within it. The section, whose members are characterized by a scorpioid cyme inflorescence which is axillary in position, is quite heterogeneous, and has apparently no close relatives (see p.33). Itis clear from our analyses that the section as treated here is not strictly monophyletic and can be divided into two groups. We have called these groups the Solanum mite species group and the Solanum ternatum species group, following the convention of Whalen (1984). The true nature of the relationships between these two monophyletic lineages will only become clear with a large scale analysis of all non-spiny solanums. Several potential sister groups have been identified; these will be treated in future monographs, and larger scale relationships tested as more monophyletic groups are identified. TAXONOMIC AND NOMENCLATURAL HISTORY Solanum is most species rich in the New World tropics and sub- tropics, and thus many of the taxa have been described relatively recently. The last comprehensive treatment of the genus was by Dunal (1852) and while 900 species were treated in the Prodromus, at least 4000 specific epithets exist for Solanum at present. By convention and for convenience Solanum is usually divided into two main groups, the spiny solanums (subgenus Leptostemonum) and the non-spiny solanums (the rest: subgenera Solanum, Brevantherum, Bassovia, and Potatoe — D’ Arcy, 1972, see Table 1). Taxonomy of non-spiny solanums has long been confused, and there is consider- able disagreement as to monophyly within that portion of the genus. For a detailed history of the taxonomy of Solanum both before and after Dunal (1852) see Knapp (1989, 1991a) and Bohs (1994). Knapp (1989) also provides a list of recent monographs of sections of Solanum, to which can be added a monograph of Solanum section Allophyllum (Bohs, 1990) and the genus Cyphomandra (Bohs, 1994; now with all epithets transferred to Solanum, see Bohs, 1995). Table 1 Characters used to define the major Neotropical subgenera of Solanum (after D’ Arcy, 1972). Solanum c. 1500-2000 species subgenus Solanum subgenus Bassovia Stout anthers, simple hairs, no spines Stout anthers, simple hairs, pinnate leaves, axillary inflorescences, pointed fruits Stout anthers, entire leaves, dendritic or stellate hairs Scandent species or herbs, pinnate leaves usually with interstitial leaflets, lateral inflorescences, articulated pedicels Tapering anthers, stellate hairs, almost always with prickles subgenus Brevantherum subgenus Potatoe subgenus Leptostemonum History of section Pteroidea The first species of section Pteroidea to be described was Solanum anceps (as Bassovia sylvatica), described by Aublet (1775) from what is now French Guiana. Several more species were described by Ruiz & Pavoén (1799) from collections made in Peru (S. anceps, S. conicum, S. diffusum, S. incurvum, S. mite, S. ternatum). Ruiz & Pavon noted the similarity between these taxa, and commented upon it in the Flora peruviana et chilensis (1799). In his Histoire S. KNAPP AND T. HELGASON naturelle, médicale et économique de Solanum, Dunal (1813) at- tempted to treat taxonomically all known species of Solanum in a hierarchical fashion. He divided the genus into a series of nested groups, marked by different symbols (for a discussion of these and their significance to sectional nomenclature in Solanum see Knapp, 1983). The group composed of the species of section Pteroidea was explicitly given sectional rank (‘la section designée sous le nom Pteroidea’) by Dunal, one of the few groups of taxa to be assigned rank in his 1813 monograph. In his section Pteroidea Dunal (1813) grouped together species sharing the following characters: ‘Foliis impari-pinnatis; foliolis integerrimis acuminatis; pedunculis axillaribus aggregatis, petiolis brevioribus. PTEROIDEA. The sim- ple-leaved species (see Morphology for a discussion of the nature of leaf division in section Pteroidea) were not considered related to the pinnate-leaved species by Dunal (1813), and were placed in a heterogeneous group, with species now placed in either the genus Lycianthes or Solanum section Geminata (sensu lato, see Knapp, 1986a). In 1816, Dunal again grouped the pinnate-leaved species of section Pteroidea together, adding S. seaforthianum (now recog- nized as a member of section Jasminosolanum) to the group. He did, however, recognize the similarities of the simple-leaved taxa, and put them in a group of their own, but without rank. In his General system of gardening and botany, one of the best compendia of flowering plants known at the time, George Don (1838) basically followed Dunal in separating the simple and pinnate species, but he put the pinnate taxa in his subsection Potatoe with the potatoes and their relatives, while the simple-leaved taxa were placed in subsec- tion Holophylla, a large and very heterogeneous group of species. Walpers (1844) followed Don’s system, but elevated Don’s subsec- tions to the rank of section. He retained the separation of the taxa based on leaf morphology and kept them in the groups where Don had placed them. In the Prodromus (1852) Dunal attempted a worldwide revision of all known species of Solanum — the last time this has been so done. In this work he radically re-organized his system of classification, creating an explicit hierarchical structure. Here, Dunal separated pinnate and simple-leaved species of section Pteroidea, putting the former in the group Polybotryon in subsec- tion Dulcamara and the latter in the group Bassovioides in subsection Micranthes (see Table 2). He described no new species of either group, but included S. pteleifolium Sendtn. (as_ S. pteleaefolium, see species treatment of S. mite) with the pinnate taxa and a group of little-known ambiguous simple-leaved species in the group Bassovioides (S. cormanthum Vell., S. laurinum Dunal, S. lacteum Vell., see Excluded Taxa for correct identification and placement of these taxa). Table 2 Classification of the species of section Pteroidea in Dunal (1852). Sectio I. Pachystemonum Subsectio III. Dulcamara. — Cymis terminalibus, dein lateralibus alaribus axillaribusque; corollis 5-angulato-plicatis, 5-fidis, 5-partitisve, coeruleis vel albis; baccis globosis ovatisque. ** Polybotryon. — Foliis impari-pinnatisectis, segmentis, integerrimis, saepius acuminatis vel simplicibus indivisis; cymis subaxillaribus, pluribus, aggregatis vel solitariis, nonnunquam radicibus oppositis; corollis 5-fidis vel 5-partitis. Subsectio IV. Micranthes. — Frutices suffruticesque; foliis integris, glabris, pilosis, tomentosis vel hispidis; calyce 5-fido, 1-2 lin. diam.; corolla duplo triplove calyce longiore; baccé globos4 ovataque, cerasi vel olivae parvae magnitudine. § 3. Bassovioides. — Cymis subaxillaribus intrafoliaceis aut suboppositifoliis; foliis brevioribus 2, 3, 4 aggregatis vel subsolitariis. REVISION OF SOLANUM SECTION PTEROIDEA The first taxonomist to recognize the close relationship between simple and pinnate-leaved taxa of what is now section Pteroidea was Georg Bitter (1912). In describing section Polybotryon, he clearly separated the taxa included in Dunal’s ‘Artengruppe Polybotryon’ into those with axillary inflorescences and those with leaf-opposed or lateral inflorescences. Bitter explicitly grouped the pinnate and simple-leaved taxa together in his new section Polybotryon, stating that the axillary inflorescence was the grouping character. In the section he included S. conicum, S. mite, S. trizygum, S. fraxinellum, S. quinquefoliololatum, S. chamaepolybotryon, S. diffusum, S. ternatum, S. pteleifolium, S. conjungens, S. hederiradiculum, S. angustialatum, and S. theobromophyllum, most of which he de- scribed in the same paper. In 1921, Bitter united all of Dunal’s (1852) various ambiguous grades (excluding the species he recog- nized as the segregate genus Lycianthes) possessing axillary inflorescences and elevated the group to subgeneric rank, as subgenus Bassovia (Aubl.) Bitter. He based its elevation in rank solely on the peculiar axillary inflorescence possessed by all species in the group. Subsequent authors have for the most part followed Bitter in placing these species in a group of subgeneric rank diagnosed by possession of an axillary inflorescence (Seithe, 1962; Danert, 1967, 1970; Gilli, 1970; D’Arcy, 1972; D’Arcy, 1991). No attempts have been made to determine relationships with other groups of solanums. D’Arcy (1991) however, did include section Pteroidea as part of subgenus Solanum in his review of taxonomy of the Solanaceae. Child (1991) is the only author to place the section in subgenus Potatoe (G. Don) D’ Arcy, but he did not explain clearly his reasons for doing so. From his introduction, it seems to be largely due to the herbaceous habit of many members of section Pteroidea, and per- haps due to their pinnate leaves. Recent cpDNA analyses of the Solanaceae (Olmstead & Palmer, 1991; Spooner et al., 1993) have not included members of section Pteroidea, thus it is still largely perceived as an isolated and morphologically very distinct group. MORPHOLOGY AND NATURAL HISTORY Species in section Pteroidea are all forest understory plants. Mem- bers of the group range from being herbs or semi-woody shrubs to 3 m to woody climbers up to 10 m in length. They occur in a wide range of elevations, but always in the deep shade of the forest understory (see Fig. 1a, b). Occasionally some species (e.g. S. mite) are found growing along roadsides or streams. Solanum anceps occurs at low elevations (at or near sea level) in the Amazon basin and S. incurvum to 3000 m in the Peruvian and Ecuadorian Andes. Most species are relatively rare in the habitats in which they occur, but some species (e.g. S. chamaepolybotryon) form what appear to be clonal groups. Stems Members of section Preroidea are usually slender, single-stemmed shrubs (Fig. 2a) or herbs or are variously climbing. Solanum mite has occasionally been been described on labels as a branching shrub, but this is not the common growth form for any species in the section. Most of the species will root along the stem; plants of S. conicum are apparently weak-stemmed, often falling over and rooting in that fashion. Other species in the group (e.g. S. uleanum) are trunk climbers, adhering to the substrate with small, adventi- tious roots (see Fig. 2b). The two species that we segregate as the S. ternatum species group (S. ternatum, S. incurvum) are quite woody climbers with lower stems up to 3 cm in diameter in some plants. Amongst the species of the S. mite species group, woodiness 33 is only very weakly developed in S. mite. Plants range from quite small (a few centimetres in S. conicum and S. chamaepolybotryon) to more than a metre in height (e.g. S. savanillense, S. mite) to several metres long for some of the vining taxa. In all Solanum species the young non-reproductive stem is monopodial with the leaves arranged in a 2/5 phyllotaxic spiral. When a stem begins its reproductive phase, sympodial growth begins (Danert, 1958; Bell & Dines, 1995). Each inflorescence is terminal and shoot continuation is initiated in the axil of the leaf subtending the inflorescence. A single lateral continuation of the shoot produces a monochasial growth pattern, a double one a dichasial pattern. In some species these two patterns occur in a single plant (Bell & Dines, 1995). Bell & Dines (1995) arrange species within the family along a continuum from monochasial to dichasial branching. The determining factor for pattern expression is dor- mancy of axillary buds in any given sympodial unit. Sympodial units in Solanum consist of leaves along each shoot terminating in an inflorescence. In the genus these units can vary from plurifoliate (members of section Brevantherum, section Holophylla, the S. niti- dum species group, see Knapp, 1989) to unifoliate (section Geminata, see Knapp, 1986a). All members of section Pteroidea have what appear to be unifoliate sympodial units (see Fig. 3 for our working hypothesis of stem structure in the group). Danert (1967) was unsure whether the vegetative axis in section Pteroidea was monopodial or monochasial (as in the rest of Solanum), and urged further ontoge- netic studies. Whether or not the unifoliate sympodia of section Pteroidea and those found elsewhere in the genus are homologous can really only be determined by such detailed ontogenetic studies. Leaves The leaves of members of section Pteroidea are generally petiolate (the petiole can be very short or absent in some species, most notably Solanum angustialatum), with pinnate, brochiodromous venation, and entire margins. Leaf shape has been used widely in section Pteroidea for determining relationships (see p. 32). The compound leaves have usually been described as imparipinnate, but are more strictly pinnatifid or deeply pinnately lobed, as thin wings of leaf tissue remain along the midrib or rachis. For the purposes of this treatment, these leaves will be referred to as pinnate, and the divisions will be described as leaflets. The petiole-like constriction at the base of the leaflets will be described as a petiolule. There are seven pinnate and three simple-leaved species in this section. The simple-leaved species, S. incurvum, S. angustialatum, andS. anceps, have entire margins; and in S. anceps leaf size, and to some extent leaf shape, is highly variable. Pinnate leaves are generally ternate to 9-jugate, with the terminal leaflet larger, and usually of a somewhat different shape than the paired leaflets. Leaflet numbers vary consid- erably within and between taxa, and exact numbers of leaflet pairs are generally not good distinguishing characteristics of species, although general trends to more or fewer leaflet pairs are good characters. Leaflet pairs are often not perfectly opposite and are occasionally markedly oblique at the base (e.g. S. conicum). The leaves of members of section Pteroidea are often very dark green, a common trait in understory plants. Several species (e.g. Solanum anceps, S. savanillense, S. uleanum) develop deep purple leaf undersides in certain conditions. Populations are often highly polymorphic for this character, differing in plants growing side by side. Whether this is due to genetics or environment is unclear. Leaf texture is membranous, as is usually the case in forest understory plants, but the leaves of some species (e.g. S. chamaepolybotryon, S. ternatum) are quite rubbery in texture, often drying quite thick on a Fig. 1 a) Lowland forest habitat of S. anceps, S. mite, S. conicum, S. uleanum: Rio Palcazu valley, Pasco, Peru, b) Cloud forest habitat of S. trizy: Cerro Pando, Chiriqui, Panama. ‘(nog ‘une ues ‘pZS9 1aj]DW = ddvuy) wunupajn °s Jo Nqey SuIquT|D (q ‘(nIVg ‘OOSND ‘g¢PO 1a/]DW YP ddouy) wngIuOd ’s Jo Viqey qn4ys snosdeqIaH (&@ 7 “BI KF e ’ , As # F] - < a S ge 4 MN Qa Z. © QO sa DY = ~ ‘ S wm eg e) a © 2 > ea a4 36 rN Fig. 3 Sympodial structure in Solanum section Pteroidea (modified from Danert, 1967). herbarium sheets. Solanum uleanum has very thin and delicate leaves, particularly the juvenile plants. Plants in the family Solanaceae are widely known for their toxic qualities derived from a diverse array of alkaloids, steroids, and phenolic glycosides (see Brown, 1987 for a review). Herbivorous insects found on the leaves of these plants are often restricted to the family, and many host-specific relationships have evolved. Among the most specialized herbivores on leaves of Solanaceae are the caterpillars of ithomiine butterflies (Nymphalidae: Ithomiinae). The adults of these butterflies are aposematic (Brown, 1987) and have evolved a wide array of colour patterns along the eastern slopes of the Andes. Host specificity of ithomiine larvae is common at species level in Solanum (Drummond & Brown, 1987). Very few host Table 3 Ithomiine larval records from members of Solanum section Pteroidea. Butterfly Host plant Country Reference Oleria vicina (Salvin) _ S. trizygum Costa Rica Drummond & Brown, 1989 Oleria makrena S. trizygum Venezuela Drummond & (Hewitson) Brown, 1989 Oleria agarista Solanum sp. section Ecuador Drummond & (Felder) Pteroidea Brown, 1989 Oleria janarilla S. anceps Peru Drummond & (Hewitson) Brown, 1989 (based on S.K. record) Oleria agarista S. mite Ecuador __ Beccaloni,1995 agarista (Felder) Oleria agarista S. anceps Ecuador __ Beccaloni,1995 agarista (Felder) Oleria sp. S. anceps Ecuador _ S.K. pers. obs. S. KNAPP AND T. HELGASON records exist for members of section Pteroidea, perhaps due to their understory habitat, or to their small size. The only larvae reared (see Table 3) from members of section Pteroidea are species of Oleria, a diverse group along the eastern Andean slope. Larvae of Oleria feed on a wide variety of other solanums (and on the genus Lycianthes) so they are probably not specific to members of the section. Their oviposition behaviour is unusual in that most host-specific lepidoptera oviposit directly on the host plant itself. This behaviour may account for the paucity of records. The senior author has observed Oleria females (in Ecuador) testing plants of S. anceps, then ovipositing on a stick or another non-solanaceous plant some metres away. This may be a form of parasitoid avoidance behaviour, but detailed ecological field studies need to be carried out. Inflorescences The inflorescence of members of section Pteroidea is a scorpioid cyme with the flowers arranged in two rows along the axis. This inflorescence type is common to most species of Solanum and has been variously misinterpreted as a raceme by earlier authors (Dunal, 1852). The position of the inflorescence is probably morphologi- cally terminal (see p. 33), but due to shoot and inflorescence rachis concaulescence and subsequent shortening of internodes (Danert, 1967) it is apparently axillary (Fig. 3). The growth of renewal shoots from axillary buds below the inflorescence causes axillary inflores- cences in Cyphomandra (Bohs, 1994), but the situation in section Pteroidea needs detailed anatomical study to ascertain whether the axillary inflorescence is homologous in these two groups. In several species of section Pteroidea multiple cymes appear to emerge from each leaf axil. This has been attributed to insertion of a subsidiary shoot on the pleiochasial inflorescence some distance from other subsidiary inflorescences (Child, 1979), but no detailed anatomical work has been done to verify this. The nature of the inflorescence in section Pteroidea has been largely responsible for its problematic phylogenetic position and its separation as an isolated subgenus by previous authors (see above). Inflorescence length is taken from the base to apex, including both the peduncle (length from base to first pedicel) and the rachis (the axis bearing pedicels). Generally in section Pteroidea the flowers occur only in the distal half to one third of the usually unbranched inflorescence. The pedicels are articulated at the base, never leaving pegs or prominent scars (see Anderson, 1977; Hawkes, 1990; Bohs, 1994). In any given inflorescence only a few, usually up to three, flowers will be open at a time, but the number of flowers per inflorescence can be determined by counting the number of pedicel scars or remnants. Pubescence of the inflorescence generally paral- lels that of the rest of the plant and hair types in the inflorescence do not differ from those found on leaves and stems. Trichomes Trichomes have traditionally provided many useful characters in Solanum taxonomy (Seithe, 1962; Roe, 1971; Seithe, 1979; Edmonds, 1982; Seithe & Anderson, 1982; Whalen, 1984; Knapp, 1991a). In section Pteroidea, however, the trichomes of all species are simple, uni- or multicellular, and uniseriate. Thus they have not been par- ticularly useful taxonomic characters in the group. Both S. anceps and S. mite have variable degrees of pubescence, from densely pubescent to completely glabrous. When analysed as separate taxa during cladistic analysis, however, the pubescent and glabrous plants always are grouped as sister taxa, suggesting that pubescence is polymorphic, as in other groups of Solanum (Knapp, 1989). Whether degree of pubescence is developmental or genetic is not known in section Pteroidea, but in some groups of spiny solanums REVISION OF SOLANUM SECTION PTEROIDEA (see Whalen et al., 1981; pers. obs. in Ecuador by Leo Roth of S. marginatum) lack of prickles is a single gene trait. The juvenile foliage and young leaves and shoots of most of the variably pubes- cent taxa are much more densely pubescent than mature leaves. Solanum ternatum has densely pubescent juvenile foliage (see Fig. 11, Knapp & Mallet 6626) but mature specimens are nearly always glabrate suggesting a developmental aspect to pubescence density. Plants described as S. dendrophilum (here treated as a synonym of S. ternatum) have dense pubescence more reminiscent of juvenile leaves than other mature individuals of S. ternatum. Flowers All species have actinomorphic, pentamerous flowers. The calyx is synsepalous and the corolla is sympetalous, although the floral tube is usually very short. The calyx lobes are usually much smaller than the corolla lobes and vary from broadly deltate to almost subulate in some collections of Solanum anceps. Pubescence of floral parts parallels that of the rest of the plant, but corolla lobes are generally glabrous except along the tips and margins. There are basically two types of corollas in section Pteroidea. In the S. ternatum species group the corolla is quite large (12-20 mm in diameter) and fleshy with the apices of the lobes usually planar at anthesis and somewhat cucullate (Fig. 4a). In the S. mite species group the corolla is much smaller, usually 5-10 mm (occasionally to 13 mm in S. conicum), with more membranous lobes that are usually strongly reflexed at anthesis (see Figs 4, 5). The corolla in both groups can be either white or pale pinkish purple, but many more collections record pigmented flowers in the S. ternatum species group that in the S. mite species group. All species have five yellow stamens of equal length inserted at the base of the corolla. The filament bases occasionally form a minute tube, but we are unconvinced of its value as a taxonomic character as emphasized by previous authors (Barboza & Hunziker, 1991). Considerable variation exists as to length or even presence of the tube, and its size is variable enough within species to not warrant its use as a taxonomic character. Anthers in members of the S. mite species group are shorter and stouter in relation to the corolla lobes than those of the S. ternatum species group.Anthers are poricidal at the tips (as in all solanums, e.g. Barboza & Hunziker, 1991) and the pore lengthens to a slit with age (see Barboza & Hunziker, 1991; Endress, 1996). Flowers of Solanum species are usually ‘buzz-pollinated’ by bees (vibratile pollination) (Buchmann, 1983; Knapp 1986a, b; Bohs, 1994). One of us (S.K.) has seen meliponiine bees (probably the genus Melipona) visiting the flowers of S. uleanum in San Martin, Peru, but flower visitors to other species have not been observed or recorded in the literature. The conical ovary is bilocular with axile placentation and there are from few (S. savanillense) to many ovules (S. ternatum). The style is straight, glabrous, papillose to densely pubescent, and usually is exserted from the anther cone. In some species however (S. conicum, S. mite, S. anceps) short-styled flowers do occur in most inflores- cences (see Fig. 4). Whether this is indicative of a derived andromonoecious breeding system (Whalen & Costich, 1986; Knapp et al., in press) is not known. The stigma on long-styled flowers is generally small and capitate, and in live plants often bright green. Stigmas of short-styled flowers are poorly developed, as is common in other species of Solanum (see Whalen & Costich, 1986). Pollen grains of members of section Pteroidea are tricolporate with a granular exine as are all other members of the genus Solanum (Anderson, 1977; Punt & Monna-Brands, 1980; Bohs, 1994). Fruits Fruits of members of section Pteroidea are unusual in Solanum. 37 Fruits in Solanum are generally smooth globose berries, but in section Pteroidea they are globose to pointed apically and smooth to markedly rugose or warty (see Fig. 5). A few other isolated species of Solanum have variously conic berries (S. capsiciforme (Domin) G.T.S. Baylis, S. nigricans M. Martens & Galeotti, S. aligerum Schidl., members of section Petota series Conicibaccata Bitter, and some members of section Cyphomandra), but none of them has the markedly rugose surface found in the berries of section Pteroidea. Pointed berries in section Pteroidea are of two kinds: conic, where the apex is full of seeds, and apiculate, where the apex is empty of seeds and is prolonged into an occasionally elongate beak. Solanum trizygum, S. chamaepolybotryon, S. savanillense, and S. conicum all have conic fruits. Solanum uleanum, this group’s closest relative, has a round fruit with a truncate apex empty of seeds that is conspicuously membranous and flattened in dried specimens. In the other species, the fruits are basically globose to ovoid, and often apiculate, varying from slightly apiculate in immature berries of S. mite to long-pointed inS. anceps. In S. anceps, variants in fruit shape are geographically coherent, suggesting that there may be discrete morphological clusters within the species as currently delimited. All fruits within section Pteroidea are green, and held erect in most species. An exception to this is S. mite, where fruits are nodding at maturity. Fruits and seeds provide many of the best characters for distinguishing species in this group. The identification of non- fruiting specimens of some taxa is difficult, and collectors are urged to record fruit characteristics in field notes. Nothing is known about the fruit or seed dispersal in Solanum section Pteroidea. All species produce green fruits that remain green at maturity, although mature fruits are quite soft and juicy. Solanum trizygum fruits (observed by S.K. in Monteverde, Costa Rica) apparently fall to the forest floor rather than being taken by birds or bats as is so common in other Solanum species. It is possible that they are eaten and thus dispersed by small ground-dwelling rodents. Fruits of the herbaceous species may be too close to the ground to be taken by bats, but those of the climbing species may be dispersed by bats. Mature fruits of S. trizygum smell strongly of wintergreen, but this has not been observed or noted by collectors for any other species in the section. Seeds The seeds of members of section Pteroidea are typical for Solanum and essentially reniform in outline. They differ however from the more typical solanum seed in being plump and somewhat ellipsoid (ovoid-reniform) rather than flattened. As in most of the studied species of Solanum the lateral epidermal cell walls are thickened and lignified (Souéges, 1907; Lester & Durrands, 1984; Edmonds, 1983; Bohs, 1994). Seed colour is not uniform in section Pteroidea. Seeds of the species in the S. mite species group tend towards greenish brown, whereas the S. ternatum species group has reddish to orange- brown seeds. Seed colours have been coded as they appear in dried specimens, but in the case of the three or four taxa that have only one specimen with mature seed, this should be treated with some cau- tion. Seed colour is an inconsistent character in the phylogenetic analysis (see p. 43) and is occasionally dependent on whether or not the specimen has been air-dried, dried over very hot driers or preserved in alcohol before drying. The fine structure of seeds has been useful for resolving the relationships among species where morphological characters ex- hibit complex patterns of variation. In Solanaceae, lateral cell wall structure can be seen after enzymatic digestion of the outer cell wall (Lester & Durrands, 1984). In order to examine cell wall structures, seeds were collected from herbarium specimens (BM, F, GH, MO, ‘(topenog ‘eloy ‘Cp6 ‘[e 10 ddpuy) asuazpiupans ‘¢§ (p ‘(nag ‘uNIReP URS ‘p7ZSO jajyjvW 2 ddvuy) wnuvajn ‘s (9 “(nag ‘ue URS ‘29¢8 Ja1]DW 3 ddpuy) wninjpYsnsun °s§ (q ‘(NJOg ‘OdSeg ‘9799 12a]]DW 3p ddpuy) wnjousay ‘s (ep “BLY p | 2 Eine Fig.5 a) S. mite (Knapp 8012, San Martin, Peru), b) S. savanillense (Knapp et al. 9044, Loja, Ecuador), c) S. anceps (Knapp & Mallet 6396, Cuzco, Peru), d) S. angustialatum (Knapp & Mallet 8567, San Martin, Peru). S. KNAPP AND T. HELGASON 40 (OW ‘16L8 Yowojos) unjvUusa] *§ JO bYSO} polsosiq (2 (OW ‘IL80€ ‘Te 19 Kuan) unainoul “¢ JO BSI) pajsasiq (q ‘e 9 “31 REVISION OF SOLANUM SECTION PTEROIDEA 41 Fig.7 a) Digested testa of S. savanillense (Knapp et al. 9044, QCNE), b) Digested testa of S. conicum (Plowman & Davis 4806, GH), c) Digested testa of S. conicum, close-up (Knapp & Mallet 6452, F), d) Digested testa of S. anceps (Franco et al. 1876, MO). 42 NY) and washed in a 20% v/v solution of sodium hypochlorite. They were then incubated in a 1% w/v solution of Driselase (SIGMA) in Sorensen’s buffer at pH 5.5 for 24 hrs at 30°C. Prepared seeds were washed in distilled water, air dried, and mounted on aluminium stubs using epoxy resin. These were then coated in a Gold-Palladium mixture, and photographed using a Hitachi S-2500 scanning elec- tron microscope. The most striking feature of the seed structures revealed by enzymatic digestion is the absence in eight of the ten taxa of projections from the cell wall thickenings (Fig. 7). Of the few species of Solanum that have been treated, to our knowledge, in this way, none have been found within the genus that have the combina- tion of thickened walls without projections (see e.g. Edmonds, 1983; Knapp, 1991a). Two species have projections from the thickened walls, S. incurvum (hair-like, Fig. 6a) and S. ternatum (flap-like, Fig. 6b), both species with larger flowers. There also appears to be variation among the species in cell size, though this would have to be confirmed using additional samples. In S. anceps, S. mite, and S. trizygum (the only species for which samples were available from more than one specimen) cell size, shape, and structure is consistent within a species and over a wide geographic range. The other striking character of the seed coat is the highly convoluted cell shape in most taxa (Fig. 7). This may be unusual even within the family, where most specimens analysed have cells that have a more or less regular shape, even where the cell walls are sinuous, e.g. S. ternatum (Fig. 6c). Seed coat characters shown in the SEM study are congru- ent with other seed and fruit characters, and provide many of the supporting characters for the two species group clades revealed by the cladistic analysis (see p. 43). It cannot be said with certainty, however, whether these are plesiomorphic or a synapomorphic characters without a more comprehensive investigation of this char- acter throughout the genus. CLADISTICS Few explicit morphologically based cladistic treatments for groups of Solanum exist. Increased interest in the use of molecular charac- ters has meant an increase in the use of parsimony analyses, but to date only a few groups of solanums have been studied (Knapp, 1989, 1991b; Spooner et al., 1993). In part the difficulty in attempting character analyses in Solanum lies in its extreme diversity and in the choice of appropriate outgroups. Choosing a range of outgroups (Watrous & Wheeler, 1981) has been thought to increase the likeli- hood of obtaining an accurately rooted tree. Recent work, however (Nixon & Carpenter, 1993), has shown that multiple outgroups perform no better at ‘polarizing’ ingroup nodes, but that multiple outgroups might improve inference. Character coding and tree construction Most of the characters used in the analysis are binary, and were polarized with reference to the outgroup, the S. nudum species group (see below). Most characters are self-explanatory but details on the variation and distribution of morphological characteristics in the species of section Pteroidea can be found in the section on morphol- ogy. Table 4 lists the characters used and their states and the data matrix is presented in Table 5. The cladistic analyses were undertaken using the computer pro- gramme HENNIG86 (Farris, 1988) using the ie* option (implicit enumeration) with all characters unordered. The ensemble consist- ency index (CI) is a measure of consistency in the entire data set with respect to the fit of characters to the tree . When the fit of a character S. KNAPP AND T. HELGASON Table 4 Character set used in the HENNIG86 analysis of Solanum section Pteroidea. Corolla diameter: >10 mm = 0; 5-10 mm = 1; <5 mm = 2 Corolla texture: papery =0; fleshy = | Number of flowers per inflorescence: few (<40) = 0; many (>40) = 1 Bud shape: globose = 1; elliptic = 0 Corolla in bud: exserted = 0; +/— included = | Calyx lobe shape: deltate = 0; quadrate = 1 Apex of calyx lobes: rounded = 0; apiculate = | Corolla tube: long (the corolla divided only %—% of the way to the base) = 0; short (divided almost to base) = 1 8. Corolla lobes at anthesis: planar or nearly so = 0; strongly reflexed = | 9. Seed shape: flattened-reniform = 0; ovoid-reniform = | 10. Seed number per berry: many (>60) = 0; few (<60) = 1 11. Seed colour: brown = 0; reddish = 1; green to green-brown = 2 12. Projections from testal cell walls : present = 0; absent = 1 13. Projections from testal cell walls: hair-like = 0; flap-like = 1; absent = 2 14. Testal cell shape: regular = 0; convoluted = 1 15. Cell wall sinuousity: 1.6-2 mm = 0; 2-3 mm = 1; 3-4 mm = 2 16. Fruit shape: round = 1; conic = 0 17. Fruit apex: rounded = 0; elongate = | 18. Fruit texture: smooth = 0; rugose = | 19. Mature fruit position: nodding = 0; erect = 1 20. Leaf shape: simple = 0; deeply divided (pinnate, pinnatifid or ternate) =] 21. Leaf texture: fleshy = 0; membranous = 1 22. Leaf petioles: not winged = 0; winged = 1 23. Plant habit: erect = 0; climbing = 1 24. Pedicel scars: flush with rachis surface = 0; raised = | NAUWAWNHHS Table 5 Data matrix used in HENNIG86 analysis of Solanum section Pteroidea. 11111 11111 22222 01234 56789 01234 56789 01234 nudum 10010 00001 20000 11000 01101 ternatum 01000 11000 00010 01001 10010 incurvum 00000 00000 01000 11010 01011 anceps 11010 00111 12121 1111001001 angustialatum 21111 00011 12121 1111001101 chamaepolybotryon 10010 00111 17771 11071 10101 conicum 20000 01111 10121 10011 11011 mite 11110 00111 10121 11001 11101 savanillense 20000 01101 12121 00011 11101 trizygum 10010 00111 12121 10011 11001 uleanum 10100 11101 10121 01111 11111 is perfect (with no parallelisms or reversals) then the consistency index is 1. The ensemble retention index (RI) is the fraction of apparent synapomorphy in all characters retained as synapomor- phies on the tree (Farris, 1989). The Solanum nudum species group (section Geminata) was selected as the outgroup as its species are of somewhat generalized morphology and are thought to be basal among non-spiny solanums (Bohs, pers. comm.; Knapp, 1989, 1991b). Cyphomandra was initially also used, but produced very low resolution in the tree, especially with unordered characters. This reflects the difficulty in classifying Cyphomandra and perhaps its problematic position in the genus Solanum (Bohs, 1994; also see below). The analysis produced three most parsimonious trees of length = 55 steps, CI = 0.52 and RI = 0.52, one of which has exactly the same topology as the strict consensus tree (Fig. 8). The other two tree topologies differed in the placement of Solanum mite relative to the rest of the mite clade: in the first S. mite was basal to the clade (mite + [chamaepolybotryon + the rest]), while in the second S. mite and REVISION OF SOLANUM SECTION PTEROIDEA 43 10 22 Boe Be S.nudum group 1 5 6 13 15 19 20 21 24 S.ternatum a ue Slt 4 6 44 Oe ©0001 | 118 t S.incurvum hei 7 aaa : S.anceps | a | a &eEd Hed cli Mee eee oe ; ree S.angustialatum y a) es Pe | 8 12 13 14 18 21 tt eo ieee S. chamaepolybotryon PSP 2-51 31 A fs 0 CG a es er rt S.mite 7 19 20 11 Bos g ee | 11 he es t S.trizygum 2 16 0 23 : i#+t S.conicum a3 7 a —iHi- 0 11 01 —i-t——- S. savanillense 8:15:22 22 oo1] 2 516175 S.uleanum y HS aces ales es | Fig.8 Cladogram of Solanum section Pteroidea. L = 55, CI = 0.52, RI = 0.52. For characters marked on the branches of the cladogram: unshaded marks indicate synapomorphies, stippled marks indicate reversals and parallelisms (homoplasy), and solid marks non-homoplastic synapomorphies. S. chamaepolybotyron were sister to the rest of the clade ([mite + chamaepolybotryon] + [the rest]). Analysis of the changes in each character suggest strongly that these tree topologies are largely defined by the suite of seed characters (see Fig. 8), with the exception of seed colour. These characters separate the ternatum-incurvum clade, whose members have small ‘hairy’ seeds and many seeded fruits, from the S. mite species group, whose members have ovoid- reniform seeds without projections and fewer seeds per fruit. This shows clearly that pinnate leaves are a derived character of the group, and that it has arisen twice, once in theS. mite species group, and once in S. ternatum. The simple-leaved taxa do not form a separate clade. These trees provide clear support for the treatment of section Pteroidea as two distinct monophyletic species groups (see Table 6), and we suggest that in any treatments of the genus Solanum at a group level, these clades should be treated as separate monophyletic groups. Section Pteroidea as a whole is clearly not a monophyletic group (see p. 32) and although treated as a unit for the purposes of this monograph, should not be lumped in further cladistic analyses. It may be that other taxa, if included in the analysis, would be placed as sister groups to either of these clades — a possibility hinted at by the very low resolution of the tree when S. diploconos (Mart.) Bohs (as Cyphomandra) was used as an outgroup, and confirmed by the fact that when added to the matrix presented here, it was the sister taxon to S. ternatum, with S. incurvum basal to the clade. Table 6 Classification of Solanum section Pteroidea. Solanum ternatum species group S. ternatum Ruiz & Pav. S. incurvum Ruiz & Pav. Solanum mite species group Solanum anceps clade S. anceps Ruiz & Pav. S. angusitalatum Bitter Solanum mite clade S. mite Ruiz & Pav. chamaepolybotryon Bitter trizygum Bitter conicum Ruiz & Pav. savanillense Bitter ay S: S: S: S. uleanum Bitter Classification We prefer not to assign ranks or formal names to the groups found in these analyses until further cladistic studies are done more widely in the genus Solanum. The monophyletic clades identified here, how- ever, have been given informal species group names (following the convention of Whalen, 1984) and their classification is summarized in Table 6. Groups of equal ‘rank’ are indented equally and the sequencing convention (Nelson, 1974; Forey, 1992) has been used. 44 TAXONOMIC TREATMENT Solanum section Pteroidea Dunal, Hist. nat. Solanum: 43 (1813). Lectotype species: Solanum mite Ruiz & Pav. (D’ Arcy, 1972). Bassovia Aubl., Hist. pl. Guiane 1: 217, t. 5 (1775). Lectotype species: Solanum sylvaticum (Aubl.) Bitter [basionym Bassovia sylvatica Aubl.] (= Solanum anceps Ruiz & Pav.) (D’Arcy, 1972). Solanum grad. ambig. Polybotryon Dunal in DC., Prodr. 13(1): 28, 66 (1852), pro parte. Lectotype species: Solanum mite Ruiz & Pav. (D’ Arcy, 1972). Solanum section Polybotryon Bitter in Reprium nov. Spec. Regni veg. 11: 469 (1912). Lectotype species: Solanum mite Ruiz & Pav. (D’Arcy, 1972). Solanum subgenus Bassovia (Aubl.) Bitter in Reprium nov. Spec. Regni veg. 17: 329 (1920/1?). Lectotype species: Solanum sylvaticum (AubI.) Bitter [basionym Bassovia sylvatica Aubl.] (= Solanum anceps Ruiz & Pav.) (Bitter, 1921). Slender wand-like shrubs, herbs or woody high-climbing lianas; young stems and leaves pubescent or glabrous, the trichomes if present simple and uniseriate. Leaves simple or pinnate (pinnatisect), fleshy or membranous, often very dark green in live plants, leaf undersides often dark purple or reddish. Inflorescence a scorpioid cyme borne in the axil of the leaf, unbranched, usually bearing 5-30 flowers; pedicel scars not raised. Buds usually rounded to ellipsoid, strongly exserted from the minute calyx tube. Flowers sympetalous, stellate, the tube very short; corolla 5-20 mm in diameter, fleshy or membranous, in the S. ternatum species group the lobes planar at anthesis and usually cucullate, in the S. mite species group the lobes usually reflexed at anthesis, sometimes strongly so; stamens five, the anthers poricidal at the tips, with age splitting longitudinally, bright yellow. Fruit a berry, usually green or yellowish green when mature, globose with a smooth surface (S. incurvum, S. ternatum, S. mite) or variously conical with a rugose surface; fruiting pedicel nodding or erect. Seeds flattened-reniform, many per fruit (S. ternatum, S. incurvum) or ovoid-reniform and few per fruit. Section Pteroidea, as here delimited, consists of ten species in two monophyletic clades (see p. 43). The clades are both kept in the section (s.].) at this time for convenience, despite some doubt as to their degree of relatedness. The S. ternatum clade, consisting of S. incurvum and S. ternatum, has large, lilac or pinkish flowers and globose fruits with many, small, flattened seeds, while the larger S. mite clade, consisting of S. anceps, S. angustialatum, S. chamaepolybotryon, S. conicum, S. mite, S. savanillense, S. trizygum, and S. uleanum, has smaller, usually greenish flowers, and (with the exception of S. mite) conical fruits with unusual rugose surfaces, the fruits having a few, ovoid, often bright green, seeds. These rugose fruits are unique in Solanum. The section has been accorded subgeneric status by many previous authors (Bitter, 1921; Seithe, 1962; Danert, 1967; D’ Arcy, 1972, 1991), on the basis of its extreme morphological difference from the rest of Solanum. We feel, how- ever, that until phylogenetic relationships in Solanum are much more clearly resolved, the group (as two clades) should be recognized only at the sectional level. This monograph is based on herbarium specimens and the exten- sive field observations of the senior author. The species are delimited on morphological grounds, with geographical and ecological prefer- ences being taken into account where appropriate. More than half of the published names of this species group have as a result, been S. KNAPP AND T. HELGASON placed in synonymy. Most of these are synonyms of the two most widely distributed species, S. mite and S. anceps. Solanum mite is relatively homogeneous (excluding variation in pubescence) over its range, whereas S. anceps has a number of forms that are somewhat geographically coherent. This variation is described in the species account. Section Pteroidea is a poorly collected group, and the material on which many of these descriptions are based is somewhat limited. A general comment here on the lectotypification, particularly of Ruiz & Pavén names, will save repetition in the species accounts. We have lectotypified all of these names using specimens from MA matched, if possible, to plates in Flora peruviana et chilensis (Ruiz & Pavén, 1799). In most cases the choice was straightforward, but when not, we have chosen the best specimen. Other lectotypes have been chosen with an eye to the wide distribution of isolectotypes. When this was not possible, the best preserved specimen was selected. Any lectotype not directly attributed to another author is designated by us here. Photographs of type specimens are cited in the recommended manner (see Knapp, 1989, 1991a), with the negative number cited in square brackets. Herbaria in possession of prints of that negative are also included in the brackets. Copies of these negatives are generally available from the institutions where they are housed: F for F negatives and US for Morton negatives. Herbaria are cited using the acronyms in /ndex herbariorum (Holmgren et al., 1990) and types seen are indicated by an exclama- tion mark (!). All non-type specimens cited in the species accounts have been seen by the authors, unless otherwise indicated. Key to selected groups of Neotropical non-spiny solanums I” Minflorescencessaxillary 22... 2;..:01-c0--20sc-cessovesssessscssntse-svesecnceusstersnseecnns 2 Inflorescences lateral or leaf-opposed ...............csseccsesssserssserersesenees 4 2 Plants small trees or shrubs, branching in a complex crown; inflores- cences in branch forks; anthers with an enlarged connective ............. Senucaseactacsssnesiastscs Cyphomandra (Solanum section Cyphomandra) Plants wand-like, shrubs, vines, or herbaceous; inflorescences only in leaf axils; anthers without an enlarged CONNECTIVE ............::cseeeeees 3 3 Trailing herbs, rooting at the nodes, inflorescences with a single flower; fruit with smooth surfaces ............. Solanum section Herpystichum Herbs (not trailing), slender shrubs or vines, inflorescences with more than one flower, usually with up to 30 flowers; fruit smooth or rugose So Set Pe rede Ana ie Os, UE ree Solanum section Pteroidea 4 Inflorescences internodal; fruit brightly coloured, with thin pericarp Soe Crd aenReeoten vain reat estenai tees tsia totasestwssteetsts Solanum section Solanum Inflorescences leaf-opposed or variously terminal; fruit green at matu- rity, the pericarp notithin .......:5.4...:.2:8%5 Solanum section Geminata Key to species of Solanum section Pteroidea [ve eaves’ sitn ple areseteccssecreteectsseccactserseecess Oe fovscncestresceasroreresecargcanes 2 Leaves variously pimmate ..................ssscsssssssscesssscssrsecssssesseseressosescees 4 2 Climbing herbs; flowers 1.2—1.4 cm in diameter, purplish; fruit globose, the surface smooth; seeds many per fruit (>50) ........ 1. S. incurvum Terrestrial herbs or weak subshrubs; flowers 0.4—0.7 cm in diameter, white or greenish white; fruit ovoid, beaked, the surface rugose; seeds few per fruit (usually 10—40) ..........cceesseeseeeeeeeeeteeeereeneeetsseeseeteenees 3 3. Stem prominently winged; style densely pubescent along its entire length. San Martin, Pert <.......:020.sc.-ssrcoesearseseaeeee 4. S..angustialatum REVISION OF SOLANUM SECTION PTEROIDEA Stem smooth, terete, not winged; style glabrous or at most papillate in the lower 2/3. Widespread ..........c.csesseceeee 3. S. ANCEPS a> Aclimbing herbs OF WOGGY: VINES 25.5 inc. sss cae eatsocricez Seance reicees 5 Terrestrial herbs or wand-like subshrubs, occasionally in large colonies 6 5 Woody vines, the basal stems often to several cm in diameter; flowers 1.6—2 cm in diameter, the petals planar, fleshy, cucullate; fruit globose, the surface’smooths .8 2 ne caress ace ee 2. S. ternatum Herbaceous vines; flowers 0.6-1 cm in diameter, the petals strongly reflexed, not fleshy or markedly cucullate; fruit conical, the surface EUS OSE rs ase esse ett canst seSteccenvsscectitees snes eee coon ta 10. S. uleanum 6 Leaves with 5 or fewer leaflets, the leaflets usually obovate, especially Cea aL CELT LLY) Wveaiey PAS et tan SEA Ec ce eR ae i Leaves usually with more than 5 pairs of leaflets, the lateral leaflets lanceolate to elliptic, the terminal leaflet similar in shape, not markedly ODOVALE Faeccecsatsetseye fasta setsatates Pera Pere eke 10 7 Leaves pubescent on the veins and lamina on both surfaces ........... 8 Leaves glabrous on lamina, occasionally pubescent along the veins and ACIS of Sore sion eri ccttecrre eos aoe are co pate en oo wae eee 9 8 Fruit conical; leaf pubescence denser adaxially ..... 8. S. savanillense Fruit globose, smooth; leaves equally pubescent on both surfaces. .... Per rt eV ay ptr terranes Sree rit core erece ere od ec eee eee 7. S. mite 9. Fruit conical, the surface rugose; leaves fleshy; plants very small and rooting along the Stem 2-sc<-s<.secseses estas 5. S. chamaepolybotryon Fruit globose or at most apically pointed, the surface smooth; leaves membranous; plants often woody at the base and up to | m tall ........ ey atassnevedassustclive vetassstucesscusrvutsasesTeeae reins Gietes Sozthc taze mean tree 8. S. mite 10 Flowers 5-6 mm in diameter, the petals strongly reflexed at anthesis; {ruitislobOse;SinGothe eesti ee eee eee 8. S. mite Flowers 9-13 mm in diameter, the petals usually planar or only slightly reflexed at anthesis; fruit conical, rugose ................0cscss 10 mm in diameter; leaflets long-petiolulate, the petiolule 3— 17 mm; leaves densely pubescent in a groove along the adaxial side of tS ACHES eecesyecaces ces cecc freee e ieee este ee sess es 6. S. conicum Flowers < 10 mm in diameter; leaflets short-petiolulate, the petiolule c. 1 mm long; leaves only sparsely pubescent if at all and then only with a few scattered trichomes abaxially ..............c:ccceeees 9. S. trizygum The Solanum ternatum species group 1. Solanum incurvum Ruiz & Pav., Fl. peruv. 2: 34, fig. 154b (1799). Type: Peru, Hudnuco, Mufia, August, September, Ruiz & Pavon s.n. (MA!-lectotype [F neg. 29716, F!]). Fig. 9. Climbing herb, up to 2 m in length at maturity, often trailing along the forest floor. Stems c. 8 mm in diameter, minutely to densely pub- escent with simple uniseriate trichomes 0.3—1.0 mm long. Leaves simple, 6-20 x 3-9 cm, elliptic to ovate, with c. (5)6—7 pairs of primary veins, glabrous to somewhat densely pubescent with simple uniseriate trichomes, denser along the veins both abaxially and adaxially, the base acuminate, the apex acute; petiole 1.5—9 cm long, glabrous to pubescent with simple uniseriate trichomes, glabrate. Inflorescence to 12 cm long, axillary, 1-3 inflorescences per leaf axil, bearing 3-6 open flowers at a time, with up to 12 scars, glabrous to sparsely pubescent with simple uniseriate trichomes. Buds rounded, becoming ellipsoid, strongly exserted from the calyx tube. Pedicels at anthesis 0.8—1.8 cm long, 1-2 mm in diameter, quite soft and lax, 45 pendent, glabrous to sparsely pubescent like the rest of the inflores- cence. Flowers with the calyx tubec. 0.5 mm long, conical, the sides very straight, the lobes 1-2 x 1-2 mm, acute to slightly obtuse and spreading, glabrous to sparsely pubescent with a few scattered uniseriate trichomes; corolla 12-14 mm in diameter, reddish violet to purple, the tube c. 2 mm long, the lobes 5—6 mm long, planar to very slightly reflexed, sparsely pubescent abaxially, the tips minutely papillate; anthers 3-4 x 1-1.2 mm; free portion of the filaments minute, the filament tube minute; ovary globose to bottle-shaped, glabrous; style c. 6 mm long, straight, glabrous; stigma minutely capitate. Fruit a globose (somewhat conical when immature) berry, 1-1.3 x 1-1.3 cm, green at maturity, drying black, the surface smooth; fruiting pedicel 1-1.5 cm long, nodding. Seeds c. 80-100 per fruit, 1-2 mm, flattened-reniform, orange-brown; epidermal cells regular, rectangular, with long hair-like projections, especially at the margins. COMMON NAMES AND USES. None recorded. DISTRIBUTION. Eastern slopes of the Andes from S. Ecuador to S. Peru, montane forest and forest edges, 1540-3000 m. (Fig. 10). SPECIMENS EXAMINED ECUADOR. Morona-Santiago: 9-10 km SE of San Juan Bosco, 1540- 1600 m, 27 January 1981, Gentry et al. 30871 (MO). PERU. Huancavelica: Choimacota Valley, Huanta, 2800-2900 m, 28 February 1926, Weberbauer 7570 (F, MOL). Huanuco: Playapampa, 2750 m, 16 June 1923, Macbride 4491 (F). Pasco: Oxapampa, trail to summit of Cordillera Yanachaga via Rio San Daniel, 3000 m, 75°27'W, 10°23'S, 13 July 1984, Smith 7756 (MO, USM); Oxapampa. Rio San Alberto valley E. of Oxapampa, 2700 m, 75°22'W, 10°27'S, 26 July 1984, Smith & Poetel 8069 (MO). San Martin: Valley of Rio Apisoncho, 30 km above Jucusbamba, 2800 m, 77°10'W, 7°55'S, 6 August 1965, Hamilton & Holligan 1069 (K). Solanum incurvum is one of the most poorly collected species in section Pteroidea. It grows at the highest elevations, and is appar- ently not at all common where it occurs. Considerable variation in pubescence exists among the few specimens examined, which ap- parently is not correlated with elevation or any other discernible ecological factor. Like its close relative, S. ternatum, it is reported to have lilac flowers, but variation for flower colour may exist. 2. Solanum ternatum Ruiz & Pav., Fi. peruv. 2: 38, fig. 162b (1799). Type: Peru, Huanuco, Cuchero, June, July, Ruiz & Pavon s.n. (MA!-holotype; B, destroyed [F neg. 2639, F!, MO], F!- isotypes). Figs 4a, 11. Solanum diffusum Ruiz & Pav., Fl. peruv. 2: 37, fig. 161b (1799). Type: Peru, Hudnuco, sin loc., June, July, August, Ruiz & Pavon s.n. (MA!-holotype, fragment F!). F neg. 12996 is of an obvious isotype of Solanum diffusum, but some confusion exists as to the labelling: prints with negative number 12296 (F!, MO!, NY!) are said to have been taken at B, but photographs without a negative number of the same sheet (F!, GH!, US!) are said to have been taken at MA. Solanum semievectum Bitter in Reprium nov. Spec. Regni veg. 11: 542 (1913). Type: Peru, sin loc., Poeppig s.n. (B-holotype, de- stroyed [F neg. 2638 — F!]). Solanum moritzianum Bitter in Reprium nov. Spec. Regni veg. 11: 565 (1913). Type: ‘Nouvelle Grenade’, either Venezuela or Co- lombia, sin loc., December 1852, Moritz 1028 (P-lectotype [F neg. 39192, G!, US!]; BM!, HBG!, K!, P [Morton neg. 8357, F!, GH!, US!)). Solanum feddei Bitter in Reprium nov. Spec. Regni veg. 12: 67 46 S. KNAPP AND T. HELGASON Dom Fig.9 S. incurvum. Habit: Gentry et al. 30871 (MO). Inflorescence: Hamilton & Holligan 1069 (K). (1913). Type: Peru, Hudnuco, Mufia, May 1863, Pearce s.n. (K!- holotype). Solanum dendrophilum Bitter in Reprium nov. Spec. Regni veg. 12: 143 (1913). Type: Peru, San Martin, Cerro Campana, Spruce 4385 (K!-holotype; K!-isotype). Solanum semiscandens Bitter in Reprium nov. Spec. Regni veg. 12: 142 (1913). Type: Peru, Hudnuco, Muna, 10—11000 ft, May 1863, Pearce s.n. (K!-holotype). Solanum subquinatum Bitter in Reprium nov. Spec. Regni veg. 12: 144 (1913).Type: Peru, Amazonas, Chachapoyas, 1835, Matthews s.n. (BM!-lectotype; K!-isolectotype). Solanum diffusum subsp. miozygum Bitter in Bot. Jb. 54: (Beibl. 119): 14 (1916). Type: Peru, Pasco, valley of Rio Pozuzo, tribu- tary of Rio Palcazu, 9°46’-9°50'S, 2200 m, 1909-1914, Weberbauer 6783 (B-holotype, destroyed; MOL!-lectotype; F!- isolectotype). Solanum diffusum var. miozygum (Bitter) J.F. Macbr. in Publ. Field Mus. (Bot.) 8: 111 (1930). Basionym: Solanum diffusum subsp. miozygum Bitter. Woody, high climbing vine, to 6—7 m (or more) long. Stems c. 0.5 cm in diameter, quite stout and woody at the base and somewhat four- REVISION OF SOLANUM SECTION PTEROIDEA 47 Fig. 10 Distribution of S. incurvum. lobed, greenish, not conspicuously white-lenticellate, glabrous to sparsely to densely (type of S. dendrophilum) pubescent with sim- ple, uniseriate trichomes 2-3 mm long, these drying white and cateniforme. Leaves pinnate, 9-15 x 8-12 cm, ternate or with 2-4 pairs of leaflets, somewhat fleshy, pubescent with scattered to dense simple uniseriate trichomes along the veins abaxially, glabrous to densely pubescent adaxially, the trichomes 5—10-celled, c. 2-3 mm long; petiole 2-6 cm long; lateral leaflets 2-6 x 1-3 cm, lanceolate or narrowly elliptic to elliptic or obovate, if the leaf more than ternate the leaflets usually narrower, with 4-6 pairs of primary veins, the base attenuate, oblique, enlarged basiscopically, the apex acute; petiolule 0.5—1 cm; basal leaflets smaller than the laterals if the leaf more than ternate; terminal leaflet 2-10 x 1-3 cm, slightly more obovate, the base attenuate, the apex acute to occasionally acumi- nate; petiolule c. 0.5 cm. Inflorescence axillary, 1.5—6 cm long, bearing flowers only in the distal 1/3, simple, occasionally 2 per axil, with 2-4 flowers open at a time, the pedicel scars raised, widely spaced, up to 24 per inflorescence. Buds elliptic, c. 6 x 3 mm, strongly exserted from the calyx tube. Pedicels at anthesis 1—1.3 cm long, c. 0.5 mm in diameter, erect to horizontal. Flowers with the calyx tube very open, almost flat, c. 2 mm long, the lobes 1.5—2 x 2 mm, quadrate with a distinct apical lobe, glabrous to sparsely pubescent with simple uniseriate trichomes like the rest of the plant; corolla 16-20 mm in diameter, white to greenish to pink, lobed c. 3/ 4 of the way to the base, the lobes cucullate, planar at anthesis, minutely papillate at the tips and along the margins; anthers 4-5 xc. 2 mm, slightly sagittate at the base, poricidal at the tips; free portion of the filaments c. 0.5 mm long, the filament tube absent; ovary conical, glabrous; style c. 8 mm long, glabrous; stigma capitate to clavate. Fruit a globose to slightly apically pointed, green berry, 1— 1.2 cm in diameter, 1-1.5 cm long, the surface smooth; fruiting pedicel 1.5—1.7 cm long, fleshy, c. 2 mm in diameter at the apex, pendent. Seeds 80-140 per berry, 1.2-2 x 1.2-1.8 mm, flattened, almost round, reddish brown; epidermal cells more or less regular, with flap-like thickenings. COMMON NAMES AND USES. None recorded. DISTRIBUTION. Tropical wet forest to humid cloud forest, in deep shade or forest edges from 100-2800 m. In the Andean region from Colombia and Venezuela to Bolivia. (Fig. 12). SPECIMENS EXAMINED COLOMBIA. Cundinamarca: Municipio de San Bernardo; Vereda Santa Marta, alrededores de la Laguna La Chorrera, 2300-2350 m, 20 July 1981, Diaz P. & Melief 2952 (MO). Huila: Finca Merenberg, E. of Volcan Purace, near Cauca border, 2300 m, 76°02'W, 2°16'S, 3 April 1986, Gentry et al. 53970 (MO); Finca Merenberg, border with Cauca, E. of Leticia, 2300 m, 76°12'W, 2°16'S, 08 July 1984, Gentry et al. 47779 (MO). Magdalena: Alrededores de Yerbabuena, 2000 m, 26 January 1959, Romero Castafieda 7067 (AAU); Sierra Nevada de Santa Marta, Sierra del Libano, Las Nubes, 1898-1901, Smith 1162 (BM, BR, F, MA, MO, NY, US, W, WIS). Norte de 48 S. KNAPP AND T. HELGASON Fig. 11S. ternatum. Habit: Killip & Smith 20235 (GH). Juvenile foliage and flowers: Knapp & Mallet 6626 (US). Fruits: Zaruma et al. 21A (QCNE). Santander: Pica-Pica Valley, above Tapata (N. of Toledo), 2100-2400 m, 1 November 1988, Hurtado & Alvarado 1121 (MO); Carretera Hollin-Loreto, March 1927, Killip & Smith 20235 (GH, US). km 40-50, alrededores de la comunidad Huamant y del Rio Pucuno, 1200 m, VENEZUELA. Aragua: E. of Colonia Tovar, 7500 ft, 8 April 1854, 77°36'W, 00°43'S, 10 October 1988, Hurtado 625 (MO). Pastaza: Capitaine Fendler 1017 (GOET). Miranda: Colonia Tovar, 1800-2000 m, December Chiriboga, Rio Pastaza, vicinity of army base, 235 m, 76°49'W, 2°32'S, 21 1924, Allart 335 (US). July 1988, Lewis et al. 13771 (QCNE); 2 km al NE de Mera, Hacienda San ECUADOR. Napo: Carretera Hollin-Loreto, km 25, Centro Challuayacu, Antonio del Barén von Humboldt, 1100 m, 78°06'W, 01°27'S, 18 March en trocha hacia la zona del Guagua Sumaco, 1230 m, 77°40'W, 00°43'S, 10 1985, Zarwna et al. 21A (AAU, MO, QCNE). Zamora-Chinchipe: REVISION OF SOLANUM SECTION PTEROIDEA 49 Fig. 12 Distribution of S. ternatum. Road from Loja to Zamora, 14 July 1986, D’Arcy 16506 (MO); Rio Nangaritza, Pachicutza, camino al hito de Pachicutza, 900-1000 m, 78°07'W, 4°07'S, 18 October 1991, Palacios et al. 8188 (QCNE). PERU. Amazonas: Prov. Chachapoyas, 1836, Matthewss.n. (BM, K); hills NW of Pomacocha, 2300-2700 m, 19 June 1962, Wurdack 940 (K, US); between Molinopampa & Mendoza, 10 km E. of Molinopampa, 2400 m, 23 February 1978, Wasshausen & Encarnacién 998 (US); Mendoza, 1600 m, 2 September 1963, Woytkowski 8265 (MO); Bagua, Cordillera Colan SE of La Peca, 2280-2400 m, 7 October 1978, Barbour 3829 (MO), 1800-1870 m, 17 October 1978, Barbour 4160 (MO). Cajamarca: Cuchero, Dombey s.n. (P [n.v., Morton neg. 8354, F!, MO!, US!]); SanAndrés de Cutervo, sobre la ruta alas grutas, al N. de SanAndrés, 2250 m, 25 June 1989, SdnchezVega 4895 (F). Cusco: Dtto. Camanti, Maniri, 8 km W. de Quincemil, a los margenes de la quebrada Garrote, 720 m, 70°48'W, 13°17'S, 20 July 1990, Timand & Astete 692 (MO); along Rio Pillahuata, 2300-2400 m, 3 May 1925, Pennell 14012 (F); Rio Mapitunuari, c. half way from Luisiana and RioApurimac to camp 1, 800-900 m, 73°42'W, 12°39'S, 15 June 1968, Dudley 10152 (F). Huanuco: Muifia, trail to Tambo de Vaca, 2440 m, 5 June 1923, Macbride 427 (G, F); Huacachi, estaci6n near Mufia, 1980 m, 20 May 1923, Macbride 4698 (F); Mufia, May 1863, Pearce 135 (BM); Divisoria, 1600 m, 10 September 1946, Woytkowski 34512 (F, MO); Rupa Rupa, Calpar Bella, Cueva de los Huarifios (margen izquierda del Rio Monzén), 700-900 m, 29 June 1976, Schunke V. 9440 (GH, MO); La Divisora, Cordillera Azul near border with Ucayali, 1620— 1760 m, 75°48'W, 9°05'S, 1 0August 1980, Gentry etal. 29558 (MO); Pachitea, Codo de Pozuzo, alluvial fan flood plain of Rio Pozuzo after it emerges from mountains, trail S. of settlement to main river, 450 m, 75°25'W, 9°40'S, 21 October 1982, Foster9355 (MO); Dtto. Hermilio Valdizan, La Divisoria, road from Pumahuasi to La Cumbre, 1600-1660 m, 26 June 1978, Plowman & Schunke V. 7394 (MO); Prov. Hudnuco, km 452 of Lima-Tingo Maria road, 2500 m, 2 June 1981 Young & Sullivan 570 (MO); Prov. Leoncio Prado, road between Tingo Maria and Pucallpa, km 35, 1500 m, 75°48'W, 9°10'S, 3 June 1981, Sullivan & Young 1154 (MO); Mufia, 1000-1100 m, 1863, Pearce 144 (BM). Junin: Huatsiroke, 1800 m, 21 February 1960, Woytkowski 5543 (F, MO); Prov. Tarma, Agua Dulce, 1900 m, 5 March 1948, Woytkowski 35416(F, G, MO, US); San Gavan, August 1854, Lechler 2440 (G, P [n.v. Morton neg. 8252, F, GH, US]); Pichis trail, Dos de Mayo, 1700-1900 m, 2 July 1929, Killip & Smith 25811 (US). Pasco: San Juan de Cacazu, km 36 on Villa Rica-Pto. Bermudez road, trail behind colegio, 950 m, 75°10'W, 10°38'S, 13 August 1984, Knapp & Mallet6626 (BH, K, US); Oxapampa-Cerro de Pasco road, La Suiza to San Gotardo, 2100-2650 m, 75°35'W, 11°38'S, 19 May 1983, Smith 4104 (MO); Rio San Alberto valley E. of Oxapampa, slopes of Cordillera Yanachaga, 2400 m, 75°22'W, 10°34'S, 23 July 1984, Smith & Pretel 7968 (MO); El TunquiAlto, 57 km from Oxapampa, 1700 m, 75°30'W, 10°15'S, 14 May 1982, Smith et al. 1569 (MO); Oxapampa, trail to summit of Cordillera Yanachaga via Rio San Daniel, 2400 m, 75°27'W, 10°23'S, 19 July 1984, Smith et al. 7933 (MO). San Martin: Valley of Rio Apisoncho, 30 km above Jucusbamba, 2800 m, 77°10'W, 07°55'S, 8August 1965, Hamilton & Holligan 1078 (K); Zepelacio, near Moyabamba, | 100m, June 1934, Klug3665(A, BM, GH, K, MO, US). Ucayali: Rio Chino al W. del Restaurant Acapulco, 100- 1100 m, 5 June 1976, Schunke V. 9144 (MO); La Divisoria cerca a Rio Chino, 1400-1600 m, 12 June 1976, Schunke V. 9241 (MO). BOLIVIA. La Paz: Prov. Nor Yungas, Serrania de Bella Vista, 16 km N. of Carrasco (37 km N. of Caranavi) on road to Palos Blancos, 1500 m, 67°34'W, 15°35'S, 31 October 1984, Solomon & Nee 12704 (M, MO); Prov. Sur Yungas, along road 7.0-9.4 km NE of (above) Huancané, 2286-2499 m, 67°32'W, 16°20'S, 17 May 1990, Luteyn & Dorr 13699 (NY); Prov. Nor Yungas, 4.6 km NE (below) Chuspipata on road to Yolosa, 2800 m, 67°47'W, 16°17'S, 8 March 1984, Solomon & Stein 11681 (MO); Prov. Nor Yungas, 13.7 km NW of San Pedro on road through Inchuara-Mejillones, and along trail to 12 de Octubre, 1500 m, 67°37'W, 15°58'S, 12 February 1983, Solomon 9584 (MO); Hacienda Casana sobre el camino a Tipuani, 50 Lom Fig. 13S. anceps. Habit: Allard 22077 (US), (inset circle) S. angustialatum stem from Knapp & Mallet 8567 (F). S. KNAPP AND T. HELGASON REVISION OF SOLANUM SECTION PTEROIDEA 1400 m, 15 October 1922, Buchtien 7462 (US); Prov. Sud Yungas, Huancané (cerca Chulumani) 8 kms, 2450 m, 31 October 1981, Beck 4881 (F); Prov. Nor Yungas, 4.6 km below Yolosa, then 19.1 km on road up the Rio Huar-inilla, 1700 m, 67°53'W, 16°12'S, 12 November 1982, Solomon 8791 (MO). Solanum ternatum can be a very large woody liana, with lower stems up to 2 cm in diameter. In cross-section these woody stems are in the shape of an ‘8’. Like many of the members of the section, consider- able variation in pubsecence exists within the species, with densely pubescent specimens having been described as S. dendrophilum. The degree of fleshiness of the leaves of S. ternatum has also led to the description of many synonyms, but this character is unrelated to geography or habitat, and seems to vary at random throughout the range of the species. Polymorphism in flower colour is common throughout the species range, and unlike members of the S. mite species group, purple flower colour does not co-occur with purple leaf undersides (see S. anceps and S. savanillense). The Solanum mite species group 3. Solanum anceps Ruiz & Pav., Fi. peruv. 2: 36, fig. 149a (1799). Type: Peru, Hudnuco, Cuchero, July, August, Ruiz & Pavon s.n. (MA!-holotype [F neg. 29722, F!, GH!, MO!, US!]). Figs 5c, 13. Bassovia sylvatica Aubl., Hist. pl. Guiane 1: 217, fig. 75 (1775). Type: French Guiana, Aublet s.n. (BM!-lectotype). Solanum bassovia Dunal in Poir., Encycl. suppl. 3: 754 (1814); Solan. syn.: 22 (1816). nom. nov. for Bassovia sylvatica Aublet. Solanum aubletii Pulle, Enum. vasc. pl. Surinam: 411, fig. 16 (1906). nom. nov. for Bassovia sylvatica Aubl. Solanum conjungens Bitter in Reprium nov. Spec. Regni veg. 11: 12 (1912). Type: Ecuador, Tungarahua, prope Bafios, September 1892, Sodiro 114/61 (B-holotype, destroyed [F neg. 2656, F!, G!, GH!, MO!, NY!]; possible lectotype to be found in the Sodiro herbarium in Ecuador which is held privately in the monastery where he was resident). Solanum hederiradiculum Bitter in Reprium nov. Spec. Regni veg. 11: 12 (1912). Type: Peru, Loreto, Yurimaguas, August 1902, Ule 6276 (B-holotype, destroyed [F neg. 2608, F!, G!, GH!, MO!, US!]; HBG!-lectotype). Solanum theobromophyllum Bitter in Reprium nov. Spec. Regni veg. 11: 472 (1912). Type: Brazil, Amazonas, Rio Jurua, Cachoeira Miry, May 1901, Ule 5490 (W!-holotype; G!, HBG!-isotypes). Solanum theobromophyllum var. procerius Bitter in Reprium nov. Spec. Regni veg. 12: 145 (1913). Type: Brazil, Acre, Estella, 1912, Ule s.n. (no herbarium cited). Bitter cited no herbarium when he described this variety, and specifically cited the date of the collection as 1912. However, a Ule collection (at G!, K!) labelled ‘Rio Acre, Seringal Auristella, E. Ule 9735’ could be type mate- rial. The sheet at G is dated March 1911 and the K sheet is dated April 1911. In the 1913 publication, Bitter cited many K collec- tions, but the K sheet is only annotated ‘Solanum theobromophyllum’ in Bitter’s hand and dated 1914. The location of the type of this variety remains obscure. Solanum sylvaticum (Aubl.) Bitter in Reprium nov. Spec. Regni veg. 17: 330 (1921). non Solanum sylvaticum Dunal, Solan. syn.: 24. (1816). (= Lycianthes sylvatica (Dunal) Bitter, a synonym of Lycianthes geminata (Vahl) Bitter). Slender, single-stemmed shrub, to 2 m tall. Stems c. 4 mm in diameter, green, conspicuously white-lenticellate, glabrous to minutely red-papillate on new growth to densely pubescent with simple uniseriate trichomes c. 0.5 mm long. Leaves simple, 12 —45 51 x (3—)5—15 cm, very variable in size, elliptic to obovate, with 10-15 pairs of primary veins, glabrous to densely pubescent with simple uniseriate trichomes c. 0.5—1 mm long, these soon deciduous on the lamina and remaining only sparsely along the veins, the base acute to attenuate (truncate in isolated populations near Iquitos), the apex acute to acuminate; petiole 1—5 cm long. /nflorescence axillary, 1-3 cm long, c. 2-4 per axil, simple, bearing flowers c. 1 cm from the base, with 3-4 flowers open at a time, c. 40-60 pedicel scars, glabrous or if the plant pubescent then with scattered uniseriate trichomes. Buds globose, c. 2 mm in diameter, c. 1/2 included in the calyx tube. Pedicels at anthesis 5—7 mm long, c. 0.5 mm in diameter, nodding. Flowers with the calyx tube c. 1 mm long, broadly conical, the lobes broadly deltate, 0.5—1 x 1—1.5 mm, glabrous or sparsely pubescent with uniseriate trichomes; corolla white, 5-7 mm in diameter, lobed nearly to the base, the lobes reflexed at anthesis, densely papillose at the tips and along the margins; anthers |.5—2 x c. 1 mm, poricidal at the tips, free portion of the filaments c. 0.05 mm, the filament tube c. 0.05 mm; ovary conical, glabrous; style 4— 5 mm long, minutely papillate in lower 2/3 or glabrous; stigma clavate. Fruit a conical, green berry, 1—1.2 cm in diameter, 1—2.3 cm long, the beak 2-8 mm, occasionally breaking off and appearing absent, the surface rugose, the raised portions white; fruiting pedicel 0.8—1.8 cm long, erect. Seeds 2—3.5 x 1.5—2.2 mm, greenish brown, flattened, round to ovoid-reniform, c. 40 seeds per fruit; epidermal cells highly sinuous and irregular, with anticlinal thickenings but without projections. COMMON NAMES AND USES. Peru: ‘ullcu panga’ (Williams 7322). DISTRIBUTION. Colombia to Bolivia and into Brazil, from 100- nearly 3000 m, in a wide range of wet forest habitats. (Fig. 14). SPECIMENS EXAMINED COLOMBIA. sin loc., Goudot 136 (K). Antioquia: 8 km S. of Angostura on road to Represa Miraflores, c. 6°50'N, 75°18'W, 2000 m, 8 February 1986, Stein & Cogollo 3394 (MO). Boyaca: 130 miles NW of Bogotd, 3000 ft, 29 September 1932, Lawrance 345 (MO); 130 miles N. of Bogota, 3500 ft, 3 March 1933, Lawrance 645 (GH). Meta: Sierra de la Macarena, Cano Entrada, 550 m, 23 January 1950, Philipson et al. 2205 (BM, GH); Guamal Municipio, 9 March 1987, Quifiones 1045 (MO). Putumayo: Orito, Rio Calderas, 300-400 m, 1 1 December 1968, Plowman2129 (GH). Valle: Cerca a Morales-Cauca, 8 October 1968, Espinal T. & Ramos 2943 (CUVC, F); vereda La Bella, finca Miranda, 1830 m, 25 January 1983, Francoet al. 1876 (MO); Cerro La Horqueta (San Antonio), Cordillera Occidental vertiente oriental, c. km 17 de carretera Cali-Buenaventura, 2050 m, 25 November 1983, Silverstone-Sopkin 1487 (MO), 1910 m, 6 January 1986, Silverstone- Sopkin & Rodriguez 2095 (MO). GUYANA. Southern Pakaraima Mountains, escarpment to foot of Kopinang Falls, 2750 ft, 2 September 1961, Maguire et al. 46080A (NY); Upper Mazaruni River basin, NE side of Mt. Ayanganna, 800-900 m, | August 1960, Tillett et al. 44971 (NY). SURINAM. Nassau Mountains, Marowijne River, forested slopes and summit of plateau A, 430 m, 31 December 1954, Cowan & Lindeman 39020 (NY); Lely Mts, SW plateaus, along E. road on plateau 1, 550-710 m, 29 September 1975, Lindeman et al. 535 (C, F, K, MO, NY, WIS); Wilhelmina gebergte, Frederick Top, 2.5 km SE of Juliana Top, 500 m, 56°30’— 6°34'W, 3°36’-3°41'N, 31 July 1963, Maguire et al. 54407 (NY). FRENCH GUIANA. Regina region, E. plateau of Montague Torte, 1 1 km WNW of Approvague River, 200-450 m, 52°22'W, 4°18'N, 17 June 1988, Feuillet et al. 10178 (NY); Mt. Tortue, 11 km WNW of Approvague river, along the road, 200-450 m, 52°22'W, 4°18'N, 16 June 1988, Feuillet et al. 10230 (NY); Saiil, Mont Galbao, 17 October 1984, de Foresta 656 (NY); pente NE des Monts Galbao, 10 km au SW de Saiil, 500-600 m, 11 March 1975, de Granville 2374 (MO, NY); ancienne piste de Saiil a Belizon, entre Eau Claire et St. Eloi, 21 August 1981, de Granville 4944 (MO); Saiil, trace ORSTROM vers les monts Galbao, sur la Montagne Liane, 19 July 1976, de Granville B5339 (MO); Haut Camopi — Mont Belvedere, 7 December 1984, 52 S. KNAPP AND T. HELGASON Fig. 14 Distribution of S. anceps (circles) and S. angustialatum (star in circle). de Granville 7165 (NY); Montagne Bellevue de I’ Inini, ext. SW versant NW, 550 m, 15 August 1985, de Granville 7502 (NY, US); Montagne Bellevue de I’ Inini, zone centrale, 700-750 m, 20 August 1985, de Granville 7686 (NY); Mont Galbao, secteur E, 600 m, 53°17'W, 3°36'N, 15 January 1986, de Granvilleet al. 8704 (NY); Camp 4, Monpé Soula-Bassin du Hoaut-Marouini, 5 km al’Oest, 180 m, 54°04'W, 2°39'N, 3 September 1987, de Granville et al. 9975 (NY); MontAtachi Bacca-— region de1’Inini, centre du plateau sommital, camp IV, 780 m, 53°55'W, 3°33'N, 21 January 1989, de Granville et al. 10842 (NY); sin. loc., 1859, Leprieur s.n. (G); Saiil, Batard d’Eau, 15 September 1978, Prévost 304 (MO); Crique Cacao — bassin de la Haute Camopi, 54°12'W, 2°20'N, 10 May 1987, Prévost & Sabatier 2422 (NY); Saiil region, trail to Crique Limonade, S. of airfield at Saiil, 200-210 m, 53°12'W, 3°36'N, 10 November 1986, Skog et al. 7380 (NY, US). ECUADOR. Morona-Santiago: Taisha, c. 5 km NNW of the military camp, 500 m, 77°30'W, 2°23'S, 14 June 1980, Brandbyge & Asanza C. 31824 (AAU, NY);Taisha, 34km ESE of the military camp, 450 m, 77°30'W, 2°23'S, 15 June 1980, Brandbyge & Asanza C. 31873 (AAU, NY); Taisha, 8-10 km NNW of military camp, 650-700 m, 77°31'W, 2°21'S, 16 June 1980, Brandbyge & Asanza C. 31927 (AAU); Pumpuentza, SSW of village, 250 m, 77°20'W, 2°25'S, 29 June 1980, Brandbyge & Asanza C. 32365 (AAU, NY); end of road construction into Cordillera del Condor from Guisme, |2 km past Rio Zamora, 900 m, 78°27'W, 3°37'S, Brandbyge & Balslev 42280 (AAU); along Rio Metzera grande on Hacienda Sangay (plantation of Compafiia Ecuatoriana del Té C.A.) near Palora, c. 950 m, 77°58'W, 1°40'S, 15 February 1984, Knapp & Mallet 6279 (BH, K, QCA, QCNE, US); along new road Méndez-Morona, km 55-62, 800 m, 23 August 1989, van derWerff & Gudino 11400 (MO, QCNE); pozo petrolero Garza de TENNECO, c. 35 km NE de Montalvo, 260 m, 76°42'W, 1°49'S, 2-12 July 1989, Zak & Espinoza 4358 (QCNE), Zak & Espinoza 4629 (MO, NY, QCNE). Napo: Estacion de INIAP, San Carlos, 6 km SE de Los Sachas, 250 m, 19 April 1985, Baker & Trushell 6099 (NY); Comunidad de Chiro Isla, on Rio Napo, 200-275 m, 75°52’30'W, 0°36’06'S, 15 April 1990, Bensman 148 (MO); Estacién Biolégica Jatun Sacha, Rio Napo, 8 km al E. de Misahualli, 450 m, 77°36'W, 1°04'S, 22 October 1988, Cerén M. & Iguago 5430 (MO, NY, QCNE), 400 m, 10August 1989, Cerén M. 7378 (MO, NY, QCNE); Cerro Antisana, 2 miles SE of Borja, 5700 ft, 3 August 1960, Grubb et al. 1210 (K); via Hollin-Loreto, entre Rio Guamani y Rio Pucuno, km 40, 1200 m, 12 December 1987, Palacios 2222 (MO, NY, QCNE). Pastaza: Hacienda SanAntonio del Baron von Humboldt, 2 km al NE de Mera, 1300 m, 78°06'W, 1°27'S, 27 February—19 March 1985, Baker et al. 5651 (NY); Lorocachi, 24 km SSE of military camp, 200 m, 75°58'W, 1°38'S, 24 May 1980, Brandbyge & Asanza C. 30829 (AAU, NY); Ceilan, path from Ceilan to Rio Cononaco on S. side of Rio Curaray, 200 m, 75°40'W, 1°36'S, 7 June 1980, Brandbyge &Asanza C. 31783 (AAU, F, MO); along road between Puyo & Macas at km 19 S. of Puyo, 1200 m, 77°53'W, 1°37'S, 9 October 1980, Croat 50575 (MO); pozo petrolero Moretecocha de ARCO, 75 km al E. de Puyo, 580 m, 77°25'W, 1°34'S, 4-21 October 1990, Gudino et al. 1008 (MO, NY, QCNE); 17 km N. of Palora, c. 2 km N. of Tashapi (Rio Pastaza crossing), 46 km S. of Puyo on Puyo-Palora road, c. 900 m, 77°52'W, 1°42'S, 17 February 1984, Knapp & Mallet 6303 (BH, QCA, QCNE, US); Kapasi (Amuntai), Rio Pastaza, 235 m, 76°48'W, 2°31'S, 14-20 July 1988, Lewis et al. 13738 (QCNE); Captaine Chiriboga, Rio Pastaza, vicinity of army base, 235 m, 76°49'W, 2°32'S, 25-29 July 1988, Lewis et al. 13898 (QCNE); pozo petrolero Villano 2, 100 m del Rio Lliquino, 360 m, 77°27'W, 1°29'S, 24 July 1992, Palacios 10299 (QCNE); vicinity of Puyo, 750-1000 m, August 1939, Skutch 4466 (K); pozo petrolero Villano 2 de ARCO, entre los rios Iquino y Villano, 350 m, 77°27'W, 1°29'S, Tirado et al. 189 (QCNE). Sucumbios: Along road from Puerto Carmen de Putumayo, (on Colombian frontier) and Lago Agrio, vicinity of Tarapoa, 76 km E. of Lago Agrio, 240 m, 76°23'W, 0°07'S, 27 April 1984, Croat 58622 (MO, NY); 4.2-7.5 km W. of Lago Agrio, near Lago Agrio-Baeza road, c. 340 m, 31 March 1972, MacBryde & Dwyer 1367 (MO, US). Tungurahua: Along Rio Topo (Rio Toro on maps) above village of Rio Negro, on Bafios-Mera road, REVISION OF SOLANUM SECTION PTEROIDEA 1200-1400 m, 78°13'W, 1°22'S, 22 January 1984, Knapp & Mallet 6183 (BH, QCA, QCNE); Zamora-Chinchipe: Above Valladolid on road to Yangana, 2300 m, | February 1985, Harling & Andersson 21373 (GB, NY); Parque Nacional Podocarpus, Quebrada San Francisco, along Loja-Zamora road, 2040-2250 m, 79°05'W, 3°58'S, 23 June 1988, Aligaard 74954 (AAU, QCNE). PERU. San Gavan, August 1854, Lechler 2464 (K); Casapi, Matthews 1967 (K); Amazonas: Alrededor de la comunidad Kusu, Rfo Numpatkin, 1100-1300 ft, 10 March 1973, Kayap 536 (MO); Quebrada Huampami lugar Tsaesim, 7200 ft, 4 April 1973, Kayap 575 (MO); Huampami, 800-850 ft, 29 July 1974, Kayap 1347 (M); Bongara, 4 km N. of Pomacochas on road to Rioja, trail down gorge to W. of road, 2150-2200 m, 77°22'W, 5°40'S, 2 June 1986, Knapp et al. 7506 (MO); Bongara, Shillac, N. by trail from Pedro Ruiz, 2300 m, 78°O1'W, 5°49'S, 31 August—2 September 1983, Smith & Vasquez S. 4899 (MO, NY); Bongara, Sipabamba, Shilla, c. 1850-1900 m, 6 May 1981, Young & Eisenberg 375 (F, MO, NY). Cajamarca: Cutervo, San Andrés de Cutervo, carretera entre San Andrés y Santo Tomas, km 15 a 20, 15 March 1989, Diaz & Beltran 3335 (NY); Colasay, 2500 m, 30 October 1961, Woytkowski 7000 (MO). Cusco: Atalaya, near junction of Rfo Carbon & Rio Alto Madre de Dios, 31 July 1973, Foster 2411 (K, MO); Limonchayoe, c. 1 km from Cuzco-Pto. Maldonado road at Huayhumbe, c. 16 km E. of Quincemil, 400-500 m, 70°40'W, 13°15'S, 25-26 April 1984, Knapp & Mallet 6396 (BH, US, USM); Kosiiipata, Quitacalzon (Quebrada Sta. Ali- cia), c. km 163 Lucre-Paucartambo-Shintuya road, 1100-1200 m, 71°15'W, 13°07'S, 11 May 1984, Knapp & Mallet 6427 (BH, US, USM); near Pilcopata, road from Pilcopata to Patria, 6 February 1975, Plowman & Davis 5006 (GH); Kosfiipata valley, Rio Tono, first foothill ridge on road N. of Patria, 750-850 m, 71°12'W, 13°07'S, 27 November 1985, Wachter 81 (F).Huanuco: Tingo Maria, valley of Rio Huallaga, c. 7000 ft, 11-14 July 1937, Belshaw 3089 (US); Tingo Maria-Pucallpa, 1510 m, 15°WNW, 5 January 1971, Ellenberg 3889 (MO); Pachitea, Codo de Pozuzo, floodplain of Rio Pozuzo after emerges from mountains, trail N. of settlement to Rio Mashoca, 500 m, 75°25'W, 9°37'S, 19 October 1982, Foster 9298 (MO); La Divisoria, Tingo Maria-Pucullpa, near Loreto border, 1150-1350 m, 29 March 1977, Gentry et al. 18876 (F, MO); Rio Huallaga canyon below Rio Santo Domingo, c. 4000 ft, 3 June 1923, Macbride 4243 (F); Leoncio Prado, Dist. Emilio Valdizan, along old road to La Divisoria, 1380 m, 16 April 1976, Plowman 5906 (GH); Cuchero, 1830, Poeppig s.n. (K, W), 1625 (W); Pampayaco, October 1829, Poeppig 1469 (F, W); between Acomayo & Carpish Divide, 8500 ft, October 1945, Sandeman 5270 (K); Bosque Nacional de Iparia, a lo largo del Rio Pachitea cerca del campamento Miel de Abeja, | km arriba de Tournevista o unos 20 kms arriba de la confluencia con el Rio Ucayali, 300— 400 m, 26 December 1966, Schunke V. 1414 (F, MO); Bosque Nacional de Iparia, a lo largo del Rio Pachitea cerca del campamento Miel de Abeja, | km arriba de Tournevista 0 unos 20 kms arriba de la confluencia con el Rio Ucayali, W. de caserfo La Paz, 23 May 1967, Schunke V. 1981 (F, GH, K); Calpar Bella, cueva de las Huarifios, margen izquierda del Rio Monz6n, 700— 900 m, 29 July 1976, Schunke V. 9454 (MO); E. de Tingo Maria, cerca al Cerro Quemado, 672-800 m, 21 February 1978, Schunke V. 9914 (MO), 2 May 1978, Schunke V. 10108 (MO); Divisoria, 1700 m, 26 September 1946, Woytkowskiet al. 560 (F). Junin: E. of Quimiri bridge, near La Merced, 800- 1300 m, 1-3 June 1929, Killip & Smith 23939 (F, NY, US); Puerto Yessup, c. 400 m, 10-12 July 1929, Killip & Smith 26221 (US), Killip & Smith 26239 (NY, US). Loreto: Rio Tigre, caserio Nuevo Canaan, Lago Lamas Tipishca, 15 December 1979, Ayala et al. 2543 (MO, NY); Pefia Negra, 25 km SW of Iquitos, | August 1972, Croat 18651 (F, MO, NY); Mishana, Rio Nanay, c. 130 m, 20 September 1978, Diaz & Jaramillo 576 (MO); Jenaro Hererra, margen derecha Rio Ucayali, 2 September 1982, Encarnacién 26268 (MO, US); Jenaro Hererra, Rio Ucayali, 7 December 1977, Gentry et al. 21185 (MO); Andoas, Rfo Pastaza near Ecuador border, 210 m, 76°28'W, 2°48'S, 15 August 1980, Gentry et al. 29790 (MO); Iquitos, c. 100 m, 2-8 August 1929, Killip & Smith 27329 (F, NY, US); Soledad on Rio Itaya, c. 110 m, 20-22 September 1929, Killip & Smith 29584 (F, MA, NY, US); Yurimaguas, lower Rio Huallaga, c. 135 m, 22 August—2 September 1929, Killip & Smith 29076 (NY, US); San Antonio, on Rio Itaya, c. 110 m, 18 September 1929, Killip & Smith 29420 (NY, US), Killip & Smith 29493 (F, NY, US); Balsapuerto, c. 220 m, January 1933, Klug 2864 (F, NY, US); Gamitanacocha, Rio Mazan, 100— 125 m, 18 February 1935, Schunke V. 280 (F, GH, NY, US); Santa Marfa de Nanay, NW del Rio Nanay, 130 m, 27 February 1968, Schunke V. 2432 (F); Alpahuayo, field station of IIAP, c. 26 km along Iquitos-Nauta road, 130 m, 53 23 August 1988, van der Werff et al. 10219 (MO, NY); Jenaro Herrera, c. 170 m, 73°45'W, 4°50'S, 3 July 1981, Vasquezet al. 2151 (MO); carretera Nauta- Iquitos, c. 150 m, 73°45'W, 4°29'S, 17 July 1981, Vasquez 2243 (MO); Pto. Almendras, 122 m, 73°15'W, 3°45'S, 19 October 1981, Vasquez & Jaramillo 2584 (MO), 7 December 1982, Vasquez & Jaramillo 3499 (MO); Recreo, Manatf, 110 m, 72°50'W, 3°42'S, 17 October 1983, Vasquez & Jaramillo 4475 (AAU, MO, NY); Nauta, 160 m, 73°35'W, 4°32'S, 3 June 1984, Vasquez & Jaramillo 5097 (F, MO, NY); Pto.Almendras, Rio Nanay, 122 m, 73°25'W, 3°48'S, 15 August 1984, Vasquez & Jaramillo 5471 (MO, NY); Alpahuayo, estaci6n ITAP, 150 m, 73°30'W, 4°10'S, 6 June 1985, Vasquez et al. 6559 (MO); Iquitos, Puerto Almendras-Rfo Nanay, 122 m, 73°25'W, 3°45'S, 29 December 1986, Vasquez & Jaramillo 8680 (MO); Iquitos, PuertoAlmendras, 122 m, 73°25'W, 3°48'S, 11 April 1988, Vasquez & Jaramillo 10533 (MO); Iquitos, km 4 carretera Iquitos-Nauta, terrenos de CRI, 150 m, 73°20'W, 4°10'S, 14 March 1989, Vasquez et al. 11923 (MO, NY); La Victoria on the Amazon River, 6 September 1929, Williams 3137 (F). Madre de Dios: Parque Nacional de Manu, Cocha Cashu Biological Station, c. 400 m, 71°22'W, 11°52'S, 24 September 1982, Emmons 81 (MO), 13 October 1982, Emmons 132 (MO), 12 November 1982, Emmons 146 (MO); Cocha Cashu, vicinity ox-bow lake of Rio Mant, between Panagua & Tayakome, 17-24 August 1974, Foster et al. 3312 (F); Rio Manu, Cocha Cashu station, 400 m, 9 February 1977, Foster & Terborgh 6071 (F), 14 March 1977, Foster & Terborgh 6222 (F); Aguas Calientes, across and downriver from Shintuya on RioAlto Madre de Dios, 400-500 m, 71°15'W, 12°40'S, 13 May 1984, Knapp & Mallet 6436 (BH, K, US, USM); Manu park, Cocha Cashu uplands, 400 m, 71°04'W, 11°45'S, 18 August 1986, Niifiez 5770 (MO); Tambopata, Cuzco Amazonico tourist lodge, 69°03'W, 12°33'S, 20 May 1989, Niifiez & Phillips 10464 (MO). Pasco: Pichis valley, Santa Rosa de Chivis, Rio Nochos, 9 km SW of Puerto Bermudez on new highway, 300-400 m, 74°58'W, 10°20'S, 7 September 1982, Foster 8592 (MO); Pichis valley, San Matias ridge, 10-12 km SW of Puerto Bermudez above Santa Rosa de Chivis trail to Loma Linda, 1000 m, 75°00'W, 10°20'S, 29 September 1982, Foster et al. 8969 (MO); Misericordia trail, Lanturachi-Santa Barbara, 2300-3300 m, 75°40'W, 10°20'S, 3 July 1985, Fosteret al. 10481 (F, NY); San Juan de Cacazu, km 36 of Villa Rica-Pto. Bermtidez road, along Rio Chivis, c. 950 m, 75°10'W, 10°38'S, 14 August 1984, Knapp & Mallet 6629 (BH, US, USM); km 15 of Palcazu road, km 73 Villa Rica-Iscozacin-Pto. Mairo, along Rio Palcazu, c. 380 m, 75°10'W, 10°21'S, 17-18August 1984, Knapp & Mallet 6639 (BH, K, MO, NY, US, USM), Knapp & Mallet 6644 (BH, K, US, USM); km 28 Repartition-Iscocazin, km 86 Villa Rica-Iscozacin-Pto. Mairo, along Rio La Raya near Amuesha community of Laguna, c. 350 m, 75°10'W, 10°20'S, 22- 23 August 1984, Knapp & Mallet 6655 (BH, US, USM); Iscozacin, forests near PEPP (Proyecto Especial Pichis-Palcazu) camp, Rio Iscozacin, tributary of Rio Palcazu, c. 320 m, 75°13'W, 10°12'S, 27 August 1984, Knapp & Mallet 6658 (BH, K, US, USM); Palcazu valley, Rio San José in the Rfo Chucurras drainage, 400-500 m, 75°20'W, 10°09'S, 14 May 1983, Smith 4035 (MO); 5 km SE of Oxapampa, 1850 m, 75°23'W, 10°36'S, 24 December 1983, Smith 5346 (MO); around Villa Rica, 1400 m, 26 February 1986, van der Werff et al. 8281 (MO, NY); Iscozacin, 7 October 1984, Whalen & Salick 862 (BH, NY). San Martin: Tingo Maria, 625-1100 m, 30 October 1949-19 February 1950, Allard 20850 (F); Boquerén pass, 92 km from Tingo Maria on highway to Pucallpa, c. 400 m, 16 December 1949-5 January 1950, Allard 22077, 22116 (US); Tingo Maria, 625-1100 m, 30 October 1949-19 February 1950, Allard 22522 (US); along road between Tocache Nuevo & Juanjuf, c. 965 km N. of Tocache Nuevo, 84 km S. of Juanjui, 14.1 km beyond bridge over Rio Pulcache, c. 600 m, 76°40'W, 7°41'S, 8 April 1984, Croat 58022A (MO); along Tocache Nuevo-Juanjuf road, valley of Rio Huallaga, 5 km S. of Cachaco, 42 km N. of Tocache Nuevo, 330 m, 76°38'W, 7°58'S, 8 April 1984, Croat 58041 (MO); Venceremos, near Amazonas border, km 291 on Rioja- Pomacocha road, 1850 m, 77°40'W, 5°45'S, 11 February 1984, Gentry et al. 45399 (MO); Zepelacio, near Moyobamba, c. 1100 m, August 1934, Klug 3757 (BM, F, GH, K, MO, NY, WIS); Cufiumbuque-Sisas road, c. | hr driving time from Cufiumbuque, c. 1/3 of way to Sisas, c. 850 m, 76°39'W, 6°35'S, 5 June 1984, Knapp & Mallet 6476 (BH, K, MO, NY, US, USM); km 436 of Carretera Marginal, c. 10-15 km E. of Naranjo, 180 kmW. ofTarapoto, c. 850 m, 77°20'W, 5°53'S, 2 July 1984, Knapp & Mallet 6555 (BH, F, K, MO, NY, US, USM); Cataratas de Ahuashiyacu, km 15 of Tarapoto- Yurimaguas road, 700 m, 76°21'W, 6°28'S, 29 July 1986, Knapp 7857 (MO, USM); Rio Cafiuto, Curarelandia, property of J. Schunke V. near km 23 of Tochache Nuevo- Puerto Pizana road, 475 m, 76°36'W, 8°06'S, 19 December 1981, Plowman & 54 Schunke V. 11509 (F, MO, NY); Tocache Nuevo, Quebrada Huaquisha, margen derecha del Rio Huallaga, 19 February 1970, Schunke V. 3813 (F, G, NY, US); San Juan de Pacayzapa, al E. del Puente (carretera a Moyobamba), 900 m, 7 April 1973, Schunke V. 5864 (F, NY, US); Quebrada Luis Salas, 5 km NE de Puerto Pizana, 350-370 m, 1 August 1973, Schunke V. 6612 (C, MO); Quebrada de Huaquisha, margen derecha del Rio Huallaga, 400-450 m, 3 July 1974, Schunke V. 7143 (C, MO); camino a Shunté, E. de Puente de Palo Blanco, 500-800 m, 14 July 1974, Schunke V. 7394 (MO); E. del Puente del Rio Uchiza, 400 m, 24 July 1974, Schunke V. 7745 (MO); camino al Roque, 3-4 km de San Juan de Pacaizapa, 1000-1050 m, 3 July 1977, Schunke V. 9765 (MO); Cerro Campana, November 1855, Spruce 4377 (K, W); San Roque, 1350-1500 m, 6 January 1930, Williams 6929 (F), 12 January 1930, Williams 7322 (F), 3 February 1930, Williams 7689 (F). Ucayali: Bosque Nacional von Humboldt, km 86 Pucallpa-Tingo Maria road, 270 m, 75°00'W, 8°40'S, 8 August 1980, Gentry & Horna 29521 (MO); Bosque Nacional von Humboldt, km 86 Pucallpa-Tingo Maria road, 270 m, 75°00'W, 8°40'S, 6 February 1981, Gentry et al. 31046 (MO); Bosque Nacional von Humboldt, km 88 Pucallpa-Tingo Maria road, 270 m, 75°02'W, 8°45'S, 15 March 1982, Gentry et al. 36396 (MO); Rio Chino al W. del Restuarant Acapulco, 1000- 1100 m, 6 June 1976, Schunke V. 9165 (MO); Bosque Nacional de Iparia, a lo largo del Rio Ucayali cerca del pueblo de Iparia, unos 80 km arriba del confluencia con el Rio Pachitea, 27 August 1968, Schunke V. 2712 (F, G, NY, US); Cinchona, carretera antigua a Pucallpa, 1200-1300 m, 9 May 1978, Schunke V. 10139 (F, MO); Bosque Nacional von Humboldt, km 86 Pucallpa- Tingo Maria road, Arboretum, 330 m, 75°05S'W, 8°45'S, 4 April 1982, Smith et al. 1184 (MO); Bosque Nacional A.V. Humboldt, 74°45'W, 8°40'S, 1 March 1983, Vasquez 3876 (MO, NY); Bosque Nacional von Humboldt, Quebrada Tahuahillo, c. 200 m, 20 June 1981, Young 967 (MO). BOLIVIA. sin loc., Bang 2513 (NY), Bang 2526 (F, GH, K, NY, US); Espirito Santo, 1891, Bang 1210 (NY). Beni: Prov. Ballividn, Serrania de Pilon Lajas, 15.8 km N. of the bridge over the Rio Quiquibey on road to San Borja, 1100 m, 67°11'W, 15°24'S, 19 November 1985, Solomon 14806 (MO). Cochabamba: Prov. Chapare, along road from Cochabamba to Villa Tunari, N. of Cochabamba, 20 November 1980, Croat 51262 (MO); La Paz: Nor Yungas, Coroico- Yolosa, subiendo el Rio San Juan a 10 kms, 2100 m, | April 1982, Beck 7498 (F); San Carlos, Mapiri region, 850 m, 26 January 1927, Buchtien 1287 (G, GH, US); Mapiri region, San Carlos, bei Sarampiuni, 600 m, 31 December 1926, Buchtien 1288 (US); Mapiri region, San Carlos, 850 m, 11 December 1926, Buchtien 1289 (NY, US); Hacienda Casana sobre el camino a Tipuani, 1400 m, 13 December 1922, Buchtien 7470 (US); Prov. Inquisivi, N. of Cajuata betwen Turculi & Loma Linda, 1970 m, 67°15'W, 16°49'S, 26 December 1989, Dorr et al. 6816 (AAU, NY); Prov. Sud Yungas, Yanacachi, 2200 m, 3 January 1981, Liberman 262 (F); Yungas, 6000 ft, 1885, Rusby 766 (NY); Prov. NorYungas, 4.6 km below Yolosa, then 19.1 km on road up Rio Huarinilla, 1700 m, 67°53'W, 16°12'S, 12 November 1982, Solomon 8821 (MO, NY); Prov. Nor Yungas, Serrania de Bella Vista, 16 km N. of Carrasco, 37 km N. of Caranavi on road to Palos Blancos, 1500 m, 67°34'W, 15°35'S, 31 October 1984, Solomon & Nee 12674 (MO); Prov. Larecaja, 19 km al SW de Guanay por el camino aTipuani, 1200 m, 67°59'W, 15°34'S, 23 January 1988, Solomon 17675 (NY). Pando: Prov. Nicolas Suarez, en la zona de Campoana, junto a la barraca San José, hacia las riberas del Narueda, 290 m, 15 January 1983, Ferndndez Casas & Susanna 8299 (MO, NY). BRAZIL. Acre: Rio Branco, 33 km NNE of Rio Branco on road to Porto Acre (AC-10) then several km E. on Ramal de Canindé, 67°37'W, 9°45'S, 18— 28 July 1989, Daly et al. 6118 (NY); Placido de Castro, Seringal Triunfo, c. 17 km WNW of Placido de Castro on road to Rio Branco (AC-040), c. 67°15'W, 10°19'S, 30 July—1 August 1989, Daly et al. 6133 (NY); 45 km from Rio Branco on Rio Branco-Brasileia road, 2 October 1980, Lowrie et al. 331 (K, NY); 40 km from Rio Branco on Rio Branco-Santa Rosa road, 8 October 1980, Lowrie et al. 441 (NY); Rio Jurua between Mundurucus & Tatajuba, 13 May 1971, Maas et al. P12903 (NY); Rio Branco, proxima a Col6nia Penal, 10 July 1965, Pires 10062 (US); km 18 road Cruzeiro do Sul to Japiim, 26 October 1966, Prance et al. 2833 (NY); Rio Moa 8 km above Cachoeira Grande, 27 April 1971, Prance et al. 12573 (NY). Amazonas: E. bank of Rio Madeira, 1 km N. of Humaitd, 2 December 1966, Prance et al. 3541 (NY). Amapa: Rio Oiopoque, | kmW. of Cachoiera Utussansain, near frontier with French Guiana, 52°55'W, 2°08'N, 8 September 1960, /rwinet al. 48077 (NY). Mato Grosso: source of the Jatuarana River, Machado River region, Decem- ber 1931, Krukoff 1599 (G, NY). Rond6nia: Sao Lourenco cassiterite mine, S. KNAPP AND T. HELGASON c. 20 km NW of Rio Madeira across from Mutumparan4, c. 7 km N. of S. Lourengo on road to A Macisa mine, 15 July 1979, Calderon et al. 2855 (K, US); Presidente Medici, BR 364 (Cuiaba-Porto Vehlo, km 300, estrada para Alvorada do Oeste, km 24, linha 110, 62°63'W, 11°12'S, 28 June 1984, Cid et al. 4829 (K, NY, US); Rio Javari, behind Estirao de Equador, 10 August 1973, Lleras et al. P17286 (NY); c. 2 km E. of Mineracaéo at Campo Novo, 300 m, 63°55'W, 10°34'S, 22 April 1987, Nee 34977 (K, NY, US). Solanum anceps is one of the most widespread and variable of the species of section Pteroidea. It is basically an Amazonian lowland species, but is found in the Andes in Peru and Bolivia up to elevations of about 2000 m. There is little discernible difference between the high elevation and low elevation forms of S. anceps as so often occurs in other groups of Solanum (see Knapp, 1986a, 1991a). Considerable geographical variation in pubescence exists within S. anceps, with some densely hairy populations being found through- out the species range. New leaves are always more densely pubescent than mature leaves and there may be a developmental or genetic component to pubescence in S. anceps. Leaf size also varies a great deal in S. anceps, with some leaves attaining very large sizes in shady habitats. Purple leaf undersides are common in S. anceps and are noted on many herbarium labels. Unlike in S. savanillense, the purple colour of leaves is not also found in flowers (see discussion of S. savanillense). The senior author has seen plants with and without purple leaves growing side by side, so perhaps there is a genetic rather than environmental cause for this characteristic. In lowland Peru (departments of Loreto & Ucayali), a series of specimens appear quite distinct. These plants (given a manuscript name honouring A.C. Smith by C.V. Morton) have longer petioles, more truncate leaf bases, and somewhat more pointed fruits, but a series of well-marked intermediates occur and thus we prefer to retain this variant within S. anceps. Some specimens from Bolivia have more strongly decurrent leaf bases than those from farther north, but again, a complete range of intermediates exist. The degree to which these forms are reproductively isolated and may be in the process of speciation or differentation is worth further study. Other characters, such as DNA sequence or leaf chemistry, may differ in concert with the morphological ones observed here. 4. Solanum angustialatum Bitter in Reprium nov. Spec. Regni veg. 11: 471 (1912). Type: Peru, San Martin, near Tarapoto, Spruce 4849 (W!-holotype [F neg. 33045, F!, MO!, US!]; BR!, G!, K!- isotypes). Figs 4b, 5d, 13. Small single-stemmed shrub or herb, toc. 1 m tall. Stems c. 5 mm in diameter, completely glabrous, green or purple, conspicuously white- lenticellate, conspicuously winged, the wingc. 4 mm wide and often purple, the internodes usually short. Leaves simple, 13-35 x 4-15 cm, elliptic, with 18-20 pairs of primary veins, completely glabrous, occasionally purple abaxially, the base narrowing c. 1-3 cm from the stem to a winged petiole, the apex acuminate, occasionally abruptly; petiole 1-3 cm, strongly winged, the wings continuing onto the stem. Inflorescence axillary, 1-3 cm long, occasionally 2 per axil, simple, with c. 2-3 flowers at a time, c. 30 pedicel scars, minutely papillate. Buds globose, 1.5—2 mm in diameter, exserted from the calyx tube. Pedicels at anthesis 7-8 mm long, c. 0.5 mm in diameter, horizontal with a marked kink, so the flowers are nodding. Flowers with the calyx tube 0.5—1 mm long, broadly conical, the lobes c. 0.5 x 1 mm, broadly deltate to rounded, glabrous; corolla white, 4-5 mm in diameter, lobed nearly 3/4 of the way to the base, nodding at anthesis, the lobes very strongly reflexed at anthesis, densely papil- late on both surfaces; anthers c. 2 x 1.5 mm, poricidal at the tips, free REVISION OF SOLANUM SECTION PTEROIDEA portion of the filaments less than 0.05 mm, the filament tube absent; ovary conical, densely red-papillose; style 2-3 mm long, densely white-pubescent with white papillae c. 0.05 mm long along its entire length; stigma clavate, often markedly bifid. Fruit a conical, green berry, c. 1 cm in diameter, 1.5—2 cm long, the beak 2-4 mm long, the surface rugose, the raised portions white; fruiting pedicel 7-8 mm long, erect, enlarged at the apex. Seeds 2—3.5 x 1.5—2.2 mm, greenish brown, flattened-round to ovoid-reniform, c. 20 seeds per fruit; epidermal cells highly sinuous and irregular, with anticlinal thickenings but without projections. COMMON NAMES AND USES. None recorded. DISTRIBUTION. Middle elevation premontane forests in NE Peru, 700-1200 m, only known from the Maynesian Andes (see Spruce, 1908). (Fig. 14). SPECIMENS EXAMINED PERU. San Martin: Trail to television antenna, Cerro de la Escalera, km 17.5 of Tarapoto- Yurimaguas road (2.5 km N. of Cataratas de Ahuashiyacu), 850-1200 m, 76°21'W, 6°27'S, 7 September 1986, Knapp 8277 (MO, USM); Cataratas deAhuashiyacu, km 15Tarapoto- Yurimaguas road, 700 m, 76°21'W, 6°29'S, 19 August 1986, Knapp & Alcorn 7792 (MO, USM); trail to televi- sion antenna, Cerro de La Escalera, km 17.5 of Tarapoto- Yurimaguas road (2.5 km N. of Cataratas de Ahuachiyacu), 850-1200 m, 76°21'W, 6°27'S, 7 August 1986, Knapp 7905 (F, MO, NY, USM); trail to television antenna, Cerro de La Escalera, km 17.5 of Tarapoto- Yurimaguas road (2.5 km N. of Cataratas de Ahuashiyacu), 1200 m, 76°21'W, 6°27'S, 24 January 1987, Knapp & Mallet 8567 (K, MO, NY, US, USM). Solanum angustialatum is only known from the area above Tarapoto (Departmento San Martin, Peru) in wet premontane forest. It is sympatric with S. anceps and is perhaps derived from that species. The eastern slopes of the Andes are an area of very high diversity, with many species of extremely local distribution. The type of S. angustialatum was collected by Spruce in what he called the Maynesian Andes (Spruce, 1908), an isolated chain running between approximately Pucallpa (Ucayali) and Bagua on the Rio Marafion (Departmento Amazonas). Many of the plants collected by Spruce were only known from their types until recent collecting in the area of Tarapoto and the mountains behind it (S.K. in 1986) added substantially to holdings of these plants. Further collecting in other parts of this small but quite distinct mountain range may reveal range extensions for many of these apparent narrow endemics. The broadly winged stem of S. angustialatum is sometimes difficult to see on herbarium specimens, but the strongly decurrent leaves are quite distinctive and unlike those occurring in S. anceps. 5. Solanum chamaepolybotryon Bitter in Reprium nov. Spec. Regni veg. 11: 471 (1912). Type: Peru, San Martin, propeTarapoto, 1855-56, Spruce 4432 (W!-holotype [F neg. 33057, F!,G!, MO!, US!]; K!-isotype). Small fleshy herb, 10—30cm tall. Stems slender, glabrous or witha few simple uniseriate trichomes in a distinct line along one side, not conspicuously lenticellate, green or purplish. Leaves pinnate, c. 20 x 10 cm, elliptic to obovate, with c. 2 pairs of leaflets, glabrous or pubescent with simple uniseriate trichomesc. 0.5 mm long, along the veins, rachis and petiole; petiole c. 3 cm long; lateral leaflets 3.5—7 x 1-2.5 cm, withc. 5 pairs of primary veins, the base attenuate, the apex acute to acuminate; petiolule c. 0.5 cm, lightly winged; basal leaflets equal to the laterals in size and shape; terminal leaflet more obovate, 4.5-8.5 x 1—2.5 cm, the base long-attenuate onto the rachis, the apex acuminate; petiolule 0.5—1 cm long, often winged and not differenti- ated from the rachis. /nflorescence axillary, 0.5—1 cm long, simple, bearing flowers only at the tip, with c. 2 flowers open at a time, c. 5 35 scars, glabrous. Buds globose, c. 1 mm in diameter, exserted from the calyx tube. Pedicels at anthesis 3—S mm long, filiform, horizontal or nodding. Flowers with the calyx tube conical, 0.5—1 mm long, the lobes 0.5—1 x 0.5—-1 mm, deltate, glabrous or with a few scattered simple, uniseriate trichomes; corolla greenish yellow or purple, 7-8 mm in diameter, lobed nearly to the base, the lobes reflexed (7?) at anthesis, minutely papillate at the tips and along the margins; anthers c. 1.50.5 mm, poricidal at the tips, the free portion of the filaments c. 0.5 mm long, the filament tube c. 0.5 mm long; ovary conical, glabrous; style 2—2.5 mm long, glabrous; stigma minutely capitate. Fruit (immature) a conical, green berry, c. 4 mm in diameter, c. 1 cm long, the beak c. 3 mm long, surface unknown, but appearing somewhat rugose; fruiting pedicelc. 1 cm long, erect. Seeds not seen. COMMON NAMES AND USES. None recorded. DISTRIBUTION. In N. Peru in the Maynensian Andes of Spruce (Spruce, 1908). (Fig. 15). SPECIMENS EXAMINED PERU. San Martin: Rioja, near km 398 of Carretera Marginal between Pomacochas & Rioja, trail to Quebrada Venceremos and Rio Serranoyacu, 1300-1400 m, 10 July 1984, Knapp & Mallet 6590 (BH, K, US, USM). Solanum chamaepolybotryon is the most diminutive species of the S. mite species group and apparently only grows as a small, fleshy herb in middle elevation cloud forest in NE Peru. It has only been collected a few times, but is apparently clonal and grows in large groups (Knapp & Mallet 6590). This species also occurs in the Maynesian Andes of Spruce (1908, see discussion under S. angustialatum) and may be derived from the more widespread S. mite or S. conicum. 6. Solanum conicum Ruiz & Pav., Fl. peruv. 2: 88, fig. 162b (1799). Type: Peru, Hudnuco, Chinchao et Cuchero, August, September, Ruiz & Pavoén s.n. (MA!-lectotype; B [F neg 2602, F!, G!, GH!, MO!, NY!] destroyed, F!-isolectotypes). Figs 2a, 16. Solanum alatibaccatum Bitter in Reprium nov. Spec. Regni veg. 12: 68 (1913). Type: Ecuador, Loja, Palandra, 22 October 1906, André s.n. (K!-holotype). Erect or climbing (scrambling) herb, to 2 m in length (height). Stems c. 0.5 cm in diameter, glabrous, greenish, conspicuously white- lenticellate when dry. Leaves (10—)15—25(-38) x (6—)12—17(—20) cm, pinnate, elliptic, with (3—)4—5(-9) pairs of leaflets, often un- evenly paired and not perfectly opposite, glabrous abaxially, densely pubescent adaxially along the midribs of the leaflets and extending to the rachis, a few scattered trichomes on the lamina, the trichomes c. 0.25 mm long, simple, uniseriate, composed of 5—6 cells, drying white; petiole 2-9 cm long; rachis densely pubescent adaxially in a groove with simple uniseriate trichomes c. 0.25 mm long; lateral leaflets 6-12 x 1-2.8 cm, lanceolate to narrowly elliptic, with 8-9 pairs of veins, the base truncate, somewhat oblique, the apex long- acuminate, petiolule 0.3-1.5 cm; basal pair of leaflets markedly smaller than the laterals, 1.5-6 x 0.7—2 cm, lanceolate to narrowly elliptic, the base truncate, oblique, the apex long acuminate, petiolule 0.3-1.7 cm; terminal leaflet 7-12 x 1.2-3 cm, elliptic, the base truncate to acute and somewhat decurrent onto the rachis, the apex long acuminate, petiolule 0.3-1.5 cm long. Inflorescence 0.7-2.5 cm long, simple or occasionally once-branched, with 3-6 flowers open at a time, with c. 10-16 scars, finely and densely pubescent with simple uniseriate trichomes less than 0.25 mm long or with whitish papillae. Buds c. 0.5 mm in diameter, globose to obovate, exserted from the calyx tube. Pedicel at anthesis 0.5—0.8 cm long, c. 56 S. KNAPP AND T. HELGASON Fig. 15 Distribution of S$. chamaepolybotryon (star in circle) and S. conicum (circles). 0.5 mm in diameter, nodding, finely pubescent like the rest of the inflorescence. Flowers with the calyx tube 0.5—1 mm long, conical, the lobes 0.5-1 x c. 0.5 mm, broadly deltate to triangular with an apical projection, papillate to finely pubescent like the rest of the inflorescence; corolla 10-13 mm in diameter, greenish white to white, lobed nearly to the base, the lobes more or less reflexed at anthesis, densely papillate at the tips and margins; anthers c. 2 x 1.5 mm, poricidal at the tips, free portion of the filaments 1-1.5 mm long, the filament tube absent; ovary conical, glabrous; stylec. 5 mm long, glabrous; stigma minutely capitate. Fruit a long-conical, sharply pointed, green berry, 1.5-1.8 x c. 1 cm, the surface rugose, the rugosities white; fruiting pedicel 0.8-1 cm long, horizontal or nodding. Seeds c. 2.0 x 2.0 mm, flattened-round, brown, c. 40 seeds per fruit; epidermal cells sinuous and irregular, with anticlinal thickenings but without projections. COMMON NAMES AND USES. Ecuador: ‘palu rugu’ (Shemluck 303 — stems and leaves as a tea for snakebite). DISTRIBUTION. Premontane and montane forests, 200—2000 m, S. Ecuador to SE Peru. (Fig. 15). SPECIMENS EXAMINED ECUADOR. Pastaza: Kapawi (Amuntai), Rio Pastaza, 235 m, 76°48'W, 2°31'S, 14-20 July 1988, Lewis et al. 13646 (QCNE); village of Rio Chico (8 km from Puyo), near chacra of Reuben Santi, 3 km from village, 1000 m, 21 July 1980, Shemluck 303 (F). Zamora-Chinchipe: Nangaritza, Canton Shaime, en la union de los Rios Nangaritza & Numpatakaime, 1000 m, 78°40'W, 4°20'S, 7 December 1990, Palacios 6607 (QCNE); hill about 1 km upstream from Shaime along Rio Nangaritza, 900-1100 m, 16 February 1994, van der Werff et al. 13122 (BM). PERU. Amazonas: Prov. Bongaraé, Yambrasbamba, 1860-2000 m, 2 March 1967, Tillet673—226 (GH). Ayacucho: Estrella, betwen Huanta & Rio Apurimac, c. 500 m, 8, 14 May 1929, Killip & Smith 23055 (NY, US). Cusco: Paucartambo, vicinity of village of Pilcopata along Rio Pilcopata, 700-800 m, 71°10'W, 13°05'S, 10 May 1984, Knapp & Mallet 6425 (BH, G, GH, K, NY, US, USM); along Rio Carbén near Atalaya, junction of Rio Carbon & RioAlto Madre de Dios, 500-600 m, 71°07'W, 13°00'S, 15 May 1984, Knapp & Mallet 6452 (BH, K, US, USM); Kosjfiipata, Quitacalzon (Quebrada Sta. Alicia), c. km 163 of Lucre-Paucartambo-Shintuya road, 1100-1200 m, 71°15'W, 13°07'S, 16 May 1984, Knapp & Mallet 6456 (BH, K, US, USM); La Convencién, Valle de Santa Ana, above Quillabamba, 5800 ft, 20 January 1975, Plowman & Davis 4806 (GH). Huanuco: Huacachi, estacién near Mufia, 1980 m, 20 May 1923, Macbride 4134 (F); cumbre de Divisoria, 1600 m, 20 May 1978, Schunke V. 10185 (MO). Madre de Dios: Manu, Aguas Calientes, across and downriver from Shintuya on Rio Alto Madre de Dios, 400-500 m, 71°15'W, 12°40'S, 13 May 1984, Knapp & Mallet 6435 (BH, K, US, USM). Pasco: Oxapampa, Iscozacin, forests near PEPP (Proyecto Especial Pichis-Palcazu) camp, Rfo Iscozacin, tributary of the Rio Palcazu, 320 m, 75°13'W, 10°12'S, 27 August 1984, Knapp & Mallet6664 (G, K, MO, NY, US); Oxapampa, trail from Rio Iscozacin to Ameusha community of VillaAmerica, Rfo Palcazu Valley, 340 m, 75°15'W, 10°12'S, 31 August 1984, Knapp & Salick 6667 (US); 5 km SE of Oxapampa, Oswaldo Miller property, 1850 m, 75°23'W, 10°36'S, 9 December 1982, Smith 2905 (NY), 23 May 1983, Smith 4170 (K, NY); surroundings of Oxapampa, 1800 m, 4 March 1986, van der Werff et al. 8357 (MO, NY); San Juan de Cacazu, km 36 of Villa Rica-Pto. Bermudez road, along Rfo Chivis, 950 m, 75°10'W, 10°38'S, 14 August 1984, Knapp & Mallet 6631 (BH, K, US, USM). San Martin: Prov. Rioja, km 436 of Carretera Marginal, c. 10-15 km E. of Naranjo, 180 km W. of Tarapoto, 850 m, 77°20'W, 5°53'S, 2 July 1984, Knapp & Mallet 6553 (F, K, US, USM). REVISION OF SOLANUM SECTION PTEROIDEA Fig. 16S. conicum. Habit: Smith 4170 (NY) and Knapp & Mallet 6456 (K). Fruit: Smith 4170 (NY). 57 58 Solanum conicum is superficially very similar to S. mite in the vegetative state. Differences include the larger numbers of lanceo- late to elliptic (rather than obovate) leaflets with long petiolules, its habit of becoming scandent and rooting along the stem, and its very conical fruit with a pointed apex and rugose surface texture. Leaflets are more elliptic than those of either S. uleanum or S. mite, and generally have strongly oblique, somewhat truncate bases. Solanum conicum can be confused with larger leaved specimens of S. uleanum, but the leaflets are not so decurrent on the rachis nor so rounded looking as those of the latter. In southern Peru S. conicum more closely resembles S. mite than in other parts of its range and non- fruiting specimens are hard to tell apart. Like most of the other species in section Pteroidea great variation in pubescence occurs throughout the species range, with some very densely specimens occuring in all areas. 7. Solanum mite Ruiz & Pav., Fl. peruv. 2: 38, fig. 163a (1799). Type: Peru, Junin, Pozuzo et Chinchao, August, September, Ruiz & Pavons.n. (MA!-lectotype; B-isolectotype [F neg. 2625 -F!]). Figs 5a, 17. Solanum pteleifolium Sendtn. in Mart., Fl. bras. 10: 15 (1846). Type: Brazil, Amazonas, trajectu Puraque-Goara, R. Negro, June— October, Martius s.n. (M!-lectotype [F neg. 6538 — F!, NY!, US!]; M!-isolectotypes). The original spelling pteleaefolium is correctable under Article 60.8 of the Code (Greuter et al., 1994) to pteleifolium. Solanum mite subsp. hexazygum Bitter in Reprium nov Spec. Regni veg. 11: 10 (1912). Type: Bolivia, La Paz, San Carlos bei Mapiri, 15°S, 700 m, August 1909, Buchtien 1438 (US!-lectotype; NY!- isotype). Bitter cited two herbaria in his original description — Herb. Buchtein, now housed at US and Herb. Vratisl. (either WRSL or BRA, both of which Bitter could have visited) from which we have not been able to obtain a specimen. Solanum quinquefoliolatum Bitter in Reprium nov. Spec. Regni veg. 11: 11 (1912). Type: Brazil, Amazonas, Marary, Rio Jurua, Ule 5201 (B-syntype, destroyed; G!-lectotype [F neg. 23148, F!, MO!, NY!, US!]). Solanum huallagense Bitter in Reprium nov. Spec. Regni veg. 12: 138 (1913). Type: Peru, Loreto, Yurimaguas, Rio Huallaga, May 1885, Spruce 3882 (K!-lectotype; BM!, BR!, W! [F neg. 33079, G!, MO!, US!]-isolectotypes). In the original description of this species, Bitter cites specimens at K, BM, and W. The sheet at K, selected here at the lectotype, is from the first set of Spruce’s collections and is annotated in Bitter’s hand. Solanum apiculatibaccatum Bitter in Reprium nov. Spec. Regni veg. 12: 141 (1913). Type: Brazil, Acre, Cobija, January 1912, Ule 9731 (No herbarium cited by Bitter, but F neg. 2705 [F!, G!, GH!] is of a sheet at B that is now no longer extant which may perhaps have been the type). Small single-stemmed shrub to | m tall, often growing in large colonies in open places. Stems c. 1 cm in diameter, green, white- lenticellate, very woody at the base, when dry usually hollow, extremely variable in pubescence, from glabrous to densely pubes- cent with simple uniseriate trichomes 1—1.5 mm long, these drying white and cateniforme. Leaves pinnate, 10—S0 x 7-25 cm, elliptic to obovate, with 2-5 pairs of leaflets, the leaflets not always perfectly opposite, the pubescence reflecting that of the entire plant, glabrous to densely pubescent both adaxially and abaxially with simple uniseriate trichomes on the veins and the lamina, the trichomes 1-2 mm long, white, leaves glabrescent with age, but not markedly so, leaves rarely purple abaxially; petiole S—15 cm long; lateral leaflets S. KNAPP AND T. HELGASON elliptic to obovate, 7.5—15 x 2-3 cm, with c. 12-14 pairs of primary veins, the base attenuate, markedly oblique and enlarged basiscopically, the apex acute to acuminate; petiolule 2-3 mm; basal leaflets usually somewhat smaller than the laterals, but similar in shape; terminal leaflet obovate, usually much wider than the laterals, 9-15 x 3.5—8(-—10) cm, the base attenuate, usually winged and decurrent onto the rachis, the apex acute to acuminate; petiolule winged onto the rachis. Inflorescence axillary, 0.5—S cm, 1-8 aris- ing from an axil, occasionally once-branched, bearing 5—10 flowers at anthesis, with up to 100 scars, pubescence as the rest of the plant, glabrous to densely pubescent. Buds globose, c. 2 x 2 mm, exserted from the calyx tube. Pedicels at anthesis 4-6 mm long, filiform, nodding. Flowers with the calyx tube 1—-1.5 mm, broadly conical, abruptly narrowing to the pedicel, the lobes 1—1.5 xc. 1 mm, deltate, glabrous to sparsely pubescent with simple uniseriate trichomes like the rest of the inflorescence; corolla S—6 mm in diameter, greenish white, lobed nearly to the base, the lobes strongly reflexed at anthesis, the tips and margins minutely papillate, occasionally in pubescent plants with a few simple uniseriate trichomes abaxially; anthers 1—1.5 x 1—1.5 mm, poricidal at the tips, papillose abaxially, the free portion of the filaments 0.5—1 mm long, the filament tube 0.05 mm; ovary conical, glabrous; style 3-4 mm long, glabrous or minutely papillose in the lower half, some flowers short-styled and the style included in the anther cone; stigma capitate to slightly clavate. Fruit a globose, occasionally somewhat apically pointed, green berry, 0.8—1.2 cm in diameter, 1—-1.2 cm long, the surface smooth; fruiting pedicel 0.8—1 cm long, nodding. Seeds 2.3-3.0 x 1.6—2.3 mm, ovoid-reniform, brown, 35-60 seeds per fruit; epider- mal cells sinuous and irregular, with anticlinal thickenings but without projections. COMMON NAMES AND USES. Peru: ‘arco sacha blanco’ (Martin et al. 1619). . DISTRIBUTION. Throughout theAmazon basin from Colombia and the E. slopes of the Andes in Peru and Bolivia to the mouth of the Amazon in Brazil, from nearly sea level to 1500 m (Fig. 18). SPECIMENS EXAMINED COLOMBIA. Putumayo: Selva higrofila entre Quebrada de la Hormiga y San Antonio de Giiamaes, 330 m, 18 December 1940, Cuatrecasas 11151 (US); Rio Caqueta, downriver from Puerto Limén, 300-350 m, 20 December 1968, Plowman 2185 (GH). ECUADOR. Napo: Estacié6n Biolégica Jatun Sacha, 450 m, 77°36'W, 1°04'S, 24 August 1988, Cerén & Cerdén 4604 (MO, NY, QCNE); Estacion Biolédgica Jatun Sacha, Rio Napo, 8 km al E. de Misahualli, 400 m, 77°36'W, 1°04'S, 11-14April 1989, Céron 6389 (MO, NY, QCNE); Reserva Faunistica Cuyabeno, Rio Aguarico, Zancudo detras del Campamento militar, 230 m, 75°32'W, 0°29'S, 25 September 1991, Palacios et al. 7684 (QCNE); Estacion Bioldgica Jatun Sacha, S. floodplain of Rio Napo, 375-400 m, 77°36-37'W, 1°04'S, 30 July 1990, Webster 28483 (QCNE). Pastaza: Lorocachi, pica a Lagatococha a | hora siguiendo margen derecha del Rio Curaray, 200 m, 75°59'W, 1°39'S, | June 1980, Jaramilloet al. 31719 (AAU, F, NY). Zamora- Chinchipe: Taisha, 1500 ft, 5 February 1962, Cazalet & Pennington 7676 (B); Shaime, at junction of Rio Nangaritza & Rio Numpatakai, 100-1080 m, 78°42'W, 4°22'S, 7 December 1990, Mligaard 98451 (AAU, QCNE). PERU. Amazonas: Bagua, Cordillera Colan SE of Peca, 7500-7900 ft, 7 October 1978, Barbour 3831 (MO); trail E. from La Peca into Serrania de Bagua, 100-1400 m, 15 June 1978, Gentry et al. 23086 (F, MO); Bongara, c. 7 km above Pedro Ruiz on road to Pomacochas, c. 1500 m, 77°57'W, 5°58'S, 3 July 1984, Knapp & Mallet 6561 (BH, US, USM); 8 km above Pedro Ruiz (Jazan) on road to Pomacochas, 1500—1600 m, 77°53'W, 5°55'S, 3 June 1986, Knapp & Alcorn 7541 (C, MO, USM). Huanuco: Tingo Maria, 625—1100 m, 30 October 1949-19 February 1950, Allard 20481 (F, US); Tingo Maria, 7 July 1940, Asplund 12043 (US); highway Tingo Maria-La Oroya, 15.5 kmW. of Tingo Maria, March 1977, Boeke 1200 (NY); at Rio Haullaga at Tingo Maria, 4 October 1972, Croat 21039 (F, GH, MO); on route 16, near km 39 REVISION OF SOLANUM SECTION PTEROIDEA Fig. 17S. mite. Habit: Knapp et al. 6324 (K), Buchtien 1249 (US). Fruit: Knapp & Mallet 7027 (NY). 59 60 N. of Tingo Maria, Cordillera Azul, 10 November 1975, Davidson 3487 (F); Hda. Shapigilla, cerca a Tingo Marfa, 700-800 m, 10 August 1946, Ferreyra 880 (GH, US, USM), 25 February 1947, Ferreyra 1575 (NY, US, USM); Tulumayo, entre Tingo Maria y Divisoria, carretera Hudnuco- Pucallpa, 600-700 m, 5 August 1947, Ferreyra 2138 (US, USM); Pachitea, Codo de Pozuzo, floodplain of Rio Pozuzo as it emerges from the mountains, 450 m, 75°25'W, 9°40'S, 16 October 1982, Foster 9198 (NY); Tingo Maria, forest reserve behind University, 780-900 m, 28 March 1977, Gentry & Daly 18773 (F); Tingo Maria, c. 600 m, | June 1977, Hart 596 (A); Mujia, c. 7000 ft, 23 May—4 June 1923, Macbride 4001 (F); R. bank of Rio Huallaga near Cayumba, 790 m, 20 October 1936, Mexia 8326a (GH, K); Jardin Botanico de Tingo Maria, Vda. Pimentel 358, 670 m, 8 December 1981, Plowman & Ramirez R. 11212 (F, K); Tingo Maria, February 1944, Soukup 2210 (US); Bosque Nacional de Iparia, a lo largo del Rio Pachitea cerca del campamento Miel de Abeja, | km arriba del pueblo de Tournevista o unos 20 km arriba de la confluencia con el Rio Ucayali, 300-400 m, 28 November 1966, Schunke V. 1280 (US); 10 km downstream from Tingo Maria, 630 m, 30 October 1938, Stork & Horton 9536 (F, K). Junin: valley of Rio Ulcumayo, 4-10 km W. of San Ramén, 800-1100 m, 31 November 1962, Iltis & Iltis 284 (K, NY, WIS); Chanchamayo, 14 October 1863, Jsern 2241 (F); La Merced, c. 700 m, 29 May—4 June 1929, Killip & Smith 23561, 24066 (F, NY, US); Rio Paucartambo valley, near Perene bridge, 700 m, 19 June 1929, Killip & Smith 25331 (F, GH, US); Pichis trail, 625 m, 28 June— 8 July 1929, Killip & Smith 26140 (NY, US); Chanchamayo, along Rio Colorado, N. bank, W. of Puente Colorado, 12 km N. of La Merced, c. 850 m, 22 March 1984, Knapp et al. 6324 (BH, F, GH, K, NY, US, USM); La Merced, c. 2000 ft, 10-24 August 1923, Macbride 5267 (F); La Merced, 2500 ft, August 1945, Sandeman 5043 (K); 2 km W. of San Ramon, along river, 8 October 1984, Whalen & Salick 864 (BH, NY, USM). Loreto: Yurimaguas-Tarapoto road, 15 km SW of Yurimaguas, 180 m, 76°13'W, 5°59'S, 10 October 1985, Gentry et al. 52221 (MO); Explorama Tourist Camp Yanamono, halfway between Indiana and mouth of Rio Napo, 120 m, 72°50'W, 3°28'S, 20 February 1988, Gentry et al. 61713 (NY); Yurimaguas, lower Rio Huallaga, c. 135 m, 23 August-7 September 1929, Killip & Smith 27614 (BM, F, NY, US); Puerto Arturo, lower Rio Huallaga below Yurimaguas, c. 135 m, 24-25 August 1929, Killip & Smith 27760 (F, NY, US); between Yurimaguas & Balsapuerto, lower Rio Huallaga basin, 135— 150 m, 26-31 August 1929, Killip & Smith 28108 (NY, US); Balsapuerto, c. 220 m, January 1933, Klug 2872 (A, BM, F, G, GH, K, MO, NY, US); Yanamono, Explorama Tourist Camp on Rio Amazonas between Indiana & mouth of Rio Napo, c. 80 km N. of Iquitos, c. 100 m, 72°48'W, 3°28'S, 22 July 1984, Knapp 6592 (BH, K, US, USM); Iquitos and vicinity, along Rio Amazonas, July 1967,Martinet al. 1619 (K); Isla Escabino near Santa Maria, c. 100 m, 14 March 1974, McDaniel & Rimachi Y. 18383 (IBE, US), 1 July 1974, McDaniel & Rimachi Y. 18903 (IBE, NY); Rio Amazonas, Isla Rondifia, opposite Leticia, 18 March 1977, Plowman et al. 6401 (GH); carretera Iquitos a Santa Maria, 6 September 1973, Rimachi Y. 507 (F, IBE, US); Isla Iquitos, Santa Martha, 18 February 1974, Rimachi Y. 876 (IBE, NY, US); Yanamono, campamento Explorama lodgel, 106 m, 72°50'W, 3°30'S, 17 April 1985, Vasquez & Jaramillo 6370 (MO), 10 October 1986, Vasquez & Jaramillo 8287 (MO, NY); Iquitos, Buena Suerte, Rio Itaya, 130 m, 73°30'W, 4°10'S, 15 November 1986, Vasquez & Jaramillo 8362 (F, NY); Indiana, Yanamono, 106 m, 72°50'W, 3°30'S, 19 February 1989, Vdsquez & Jaramillo 11699 (NY); La Victoria on the Amazon River, August-September 1929, Williams 2829, 2923 (F); Fortaleza, Yurimaguas, lower Rio Huallaga, 155-210 m, October-November 1929, Williams 4264 (F); Sapoto-yacu, Santa Rosa, lower Rio Huallaga, 155-210 m, October— November 1929, Williams 4905 (F); Puerto Arturo, Yurimaguas, lower Rio Huallaga, 155-210 m, October-November 1929, Williams 5351 (F). Madre de Dios: Tambopata, c. 30 air km or 70-80 river km SSW of Puerto Maldonado at Rio La Torre (Rio d’Orbigny)/Rio Tambopata (SE Bank), Tambopata Nature Reserve, c. 260 m, 69°17'W, 12°49'S, 16 April 1980, Barbour 4800 (MO), 31 May 1980, Barbour 5461 (F); Tambopata Reserved Zone, 5.1 km down main trail from Explorer’s Inn, near Laguna Cocococha, 69°17'W, 12°50'S, 6 March 1988, Bell & Wiser 88-8 (NY, US); primary floodplain of Rio La Torre, La Torre trail in Tambopata Reserved Zone, 69°17'W, 12°50'S, 7 March 1988, Bell & Wiser 88-40 (AAU, G, US); Tambopata, 280 m, 69°18'W, 12°50'S, 26 February 1984, Gentry et al. 46254 (MO); Rio Tambopata, near Puerto Maldonado, 280 m, 9 November 1986, Nufiez 6473 (F, NY); Cuzco Amazonico, trail 1, 15 km NE of Puerto S. KNAPP AND T. HELGASON Maldonado, 200 m, 69°03'W, 12°33'S, 27 May 1989, Miifiez et al. 10555 (MO); Explorer’s Inn at confluence of Rio Tambopata & Rfo La Torre, 39 km SW of Puerto Maldonado, Rio La Torre trail, 69°20'W, 12°50'S, 22 September 1984, Smith & Shuler 177 (F, US), 13 October 1985, Smith et al. 713 (K, NY, US), 17 October 1985, Smith et al. 788 (NY, US); Explorer’s Inn, near the confluence of Rio Tambopata & Rio La Torre, 39 km SW of Pto. Maldonado, Laguna Chica trail, 69°20'W, 2°50'S, 17 January 1989, Smith et al. 1355 (NY, US); Tambopata Wildlife Reserve, 30 km S. of Puerto Maldonado, 260 m, 69°17'W, 12°15'S, 11 October 1984, Young & Stratton 24 (NY); Tambopata Reserve, junction Rio Tambopata & Rio La Torre, 250 m, 16 March 1981, Young 126 (NY), 18 March 1981, Young 134 (F, NY). Pasco: Enefias-Alto Yurinaki-La Florida road, c. 9 km E. of Villa Rica, 1250-1400 m, 75°15'W, 10°50'S, 12 August 1984, Knapp & Mallet 6623 (BH, K, NY, US); c. 1 km from division of Villa Rica-Pto. Bermtidez road and Villa Rica-Palcazu road, on Palcazu branch, along small tributary to Rfo Cacazu, c. 500 m, 75°10'W, 10°30'S, 15 August 1984, Knapp & Mallet 6632 (BH, K, NY, US); Pozuzo, c. 2000 ft, 20-22 June 1923, Macbride 4676 (F). San Martin: Vicinity of Uchisa, 17-18 July 1937, Belshaw 3109 (GH, US); Pueblo Mantencién, property of Hernan Ortiz, c. 10 km S. of Tocache Nuevo, 400-700 m, 26 April 1983, Bohs & Schunke V. 2168 (F, GH); Tocache Nuevo, Rio de la Plata, 550-600 m, 76°25'W, 8°10'S, 2 November 1980, Croat 51014 (F, MO); along road betwen Tarapoto & Moyobamba, c. 10 km NW of Tabalosos, c. 500 m, 76°43'W, 6°15'S, 7 November 1980, Croat 51156 (MO); Zepelacio, near Moyobamba, c. 1100 m, June 1934, Klug 3686 (A, BM, F, GH, K, MO, NY, US); Juanjui, Alto Rio Huallaga, 400-800 m, February 1936, Klug 4252 (BM, F, GH, K, NY, US); on road between Cufiumbuque & Sisas, c. | hr driving time from Cufiumbuque, 1/3 of way to Sisas, c. 850 m, 76°39'W, 6°35'S, 5 June 1984, Knapp et al. 6473 (BH, K, US, USM); c. 5 km N. of Tarapoto along Rio Shilcayo, c. 400 m, 76°22'W, 6°30'S, 7 June 1984, Knapp & Mallet 6484 (BH, F, K, NY, US), Knapp & Mallet 6486 (BH, K, US, USM); km 28 of Tarapoto- Yurimaguas road, 650 m, 76°15'W, 6°25'S, 20 June 1984, Knapp & Mallet 6526 (BH, K, US, USM); Lamas, c. km 62 of Tarapoto-Yurimaguas road, along Rio Yuracyacu, c. 260 m, 76°18'W, 6°18'S, 23 June 1984, Knapp & Mallet 6538 (BH, K, MO, NY, US, USM); km 436 of carretera Marginal, c. 10-15 km E. of Naranjo, 180 km W. of Tarapoto, c. 850 m, 77°20'W, 5°53'S, 2 July 1984, Knapp & Mallet 6554 (BH, F, K, US, USM); Naranjal, trail to Jorge Chavez, km 85 of Tarapoto-Yurimaguas road, c. 200 m, 76°17'W, 6°15'S, 2 April 1986, Knapp & Mallet 6931 (AAU, F, MO, NY, USM); 5-15 km E. of Shapaja on road to Chazuta, 200-300 m, 76°10'W, 6°36'S, 9 April 1986, Knapp & Mallet 7027 (NY, USM), Knapp & Mallet 7033 (MO, NY, USM), Knapp & Mallet 7036 (MO, NY, USM); Toma del Shilcayo, along Rio Shilcayo N. of Tarapoto, 380-400 m, 76°22'W, 6°30'S, 14April 1986, Knapp & Mallet 7065 (NY, USM), Knapp & Mallet 7068 (MO, USM); Convento, trail to Tioyacu & Neuvo Lamas (then to Rio Shanusi), km 68 of Tarapoto- Yurimaguas road, c. 200 m, 76°17'W, 6°16'S, 19 April 1986, Knapp & Mallet 7086 (NY, USM), Knapp & Mallet 7087 (F, NY, USM), 24 April 1986, Knapp & Mallet 7212 (F, MO, NY, USM), 26 April 1986, Knapp et al. 7218 (MO, NY, USM); trail to Boca Toma del Shilcayo, along Rio Shilcayo N. of Tarapoto, 400 m, 76°22'W, 6°30'S, 20-21 May 1986, Knapp & Alcorn 7331 (F, USM), Knapp & Alcorn 7332 (MO, NY, USM), 12 August 1986, Knapp 8012 (MO, NY, USM); km 54 of Tarapoto-Yurimaguas road, 350 m, 76°18'W, 6°23'S, 3 September 1986, Knapp 8264 (MO, NY, USM); Caserio El Progreso, km 30 of Tarapoto-Yurimaguas road, 700 m, 76°19'W, 6°25’5”S, 25 September 1986, Knapp & Mallet 8440 (MO, NY, USM); San Juan de Pacayzapa, E. del puente (carretera a Moyobamba), 900 m, 5 April 1973, Schunke V. 5837 (K, NY, US); camino a Roque, 8 km de San Juan de Pacayzapa, 800-900 m, 2 May 1973, Schunke V. 6169 (F, NY, US); Quebrada de Santiago, al E. de Puerto Pizana, 350-380 m, 29 July 1973, Schunke V. 6507 (C); Quebrada de Santa Rosa de Cachiyacu, carretera a Progreso, 500- 700 m, 19 July 1974, Schunke V. 7595 (F, G, MO, NY, US); Pucayacu, Tarapoto, 750 m, 11 December 1929, Williams 6045 (F); San Roque, 1350- 1500 m, 6 January 1930, Williams 6956 (F), 7 January 1930, Williams 7035 (F). Ucayali: Becerro Isla, abajo de Jenaro Herrera, 17 November 1981, Spichiger & Encarnacién 1065 (G, MO, NY); Cordillera Azul, km 15 on Tingo Maria-Pucallpa road, dirt road near Puente Pumahuasi, 700 m, 5 June 1981, Young & Sullivan 715 (F, NY, USM). BOLIVIA. sin loc., Bang 2248 (GH, NY, US). Cochabamba: Villa Tunari 34 kms hacia Cochabamba, 670 m, 25 November 1981, Beck 7361 (F, NY); Prov. Chapare, Todos Santos, 300 m, 22 October 1966, Steinbach REVISION OF SOLANUM SECTION PTEROIDEA ° - poet - or fe ° 61 Fig. 18 Distribution of S. mite. 424 (F, NY, WIS); Locotal, Prov. Chapare, 1500 m, 2 February 1929, Steinbach 9020 (GH). La Paz: Rurrenabaque, 1000 ft, 25 November 1921, Cardenas 1168, 2046 (NY); Mapiri region, San Carlos, 600 m, 18 December 1926, Buchtien 1249 (NY, US); San Carlos, Mapiri, 700 m, 15°S, 3 August 1907, Buchtien 1438 (NY, US); Bopi Rover, 3000 ft, 11 October 1922, Rusby 578 (NY); Guanai, 2000 ft, May 1886, Rusby 800 (NY); near La Paz, 10,000 ft, April 1885, Rusby 813 (NY-left-hand specimen). Pando: along Rio Madre de Dios, upstream and from 22 km WSW of Florencia, 135 m, 67°34'W, 11°30'S, 23 August 1985, Nee 31504 (NY); Loma Alta, Rio Madre de Dios, 110 m, 65°58'W, 10°47'S, 18 June 1987, Solomon 17159 (NY). Santa Cruz: Prov. Ichilo, Buena Vista, 370 m, 63°40'W, 17°27'S, 2 August 1987, Nee 35480 (NY); Estancia San Rafael de Amboro, | kmW. toward Rio Surutu, 15 km SSE of Buena Vista, 375 m, 63°37'W, 17°35'S, 29 July 1987, Nee et al. 35433 (NY); Parque Nacional Amboro, along Rio Saguayo, | km NE of entrance into first Andean foothills, 400 m, 63°43'W, 17°39'S, 21 January 1988, Nee 36036 (NY); c. 3-4 km S. of San Rafael & 0.5 km N. of San Salvador, 11 km by air SW of Villa German Busch, 600-650 m, 63°56'W, 17°29'S, 19 November 1988, Nee & Saldias P. 36888 (NY); SW side of Buena Vista, 360 m, 63°40'W, 17°28'S, 15 December 1988, Nee 37200 (NY); Parque Nacional Amboro, along Rio Saguayo, 1.5-3 km NE of entrance into first Andean foothills, 375 m, 63°43'W, 17°38-39'S, 21 De- cember 1988, Nee 37315 (NY); Parque Nacional Amboro, along Rio Saguayo, slopes along Quebrada Yapoje, above confluence with Rio Saguayo, 400 m, 63°44'W, 17°34'S, 13 December 1989, Nee 38119 (NY); Parque Nacional Amboro, along Rio Isama (Rio Pitasama on maps), 450 m, 63°37'W, 17°41'S, 12 October 1990, Nee 39259 (NY); Parque Nacional Amboro, 5 km SWS of Buena Vista, W. side of Rio Surutu, 320 m, 63°40°30”"W, 17°29°30”S, 20 October 1990, Nee 39355 (NY); Parque Nacional Amboro, W. side of Rfo Surutu, 2 km NE of El Carmen on trail to river crossing, 320 m, 63°41'W, 17°31'S, 29 October 1990, Nee 39570 (NY); E. side of Rio Yapacani at junction with Rio Surutu, 0.5 km upstream and S. from highway bridge over Rio Yapacani at Villa Yapacani, 285 m, 63°50'W, 17°24'S, 30 October 1990, Nee 39603 (NY); Rio Surutu, 400 m, | July 1924, Steinbach 6080 (A). BRAZIL. Acre: Maas et al. P12838 (NY); Cruzeiro do Sul, Rio Jurua, Rio Moa, 29 October 1966, Prance et al. 2955 (NY, US, WIS); opposite Cruzeiro do Sul, N. bank of Rio Jurua, 27 October 1966, Pranceet al. 2908 (K, NY, US, WIS); Santa Maria de Marmellos, Madeira, Ule 6922 (HBG, B[destroyed, syntype of S. guinquefoliolatum]). Amazonas: near mouth of Rio Embira, tributary of Rio Tarauaca, 70°15'W, 7°30'S, 3 June 1933, Krukoff 4642 (A, NY); Rio SolimGdes & Rio Javari, Ilha Aramaga, opposite Tabatinga, 23 July 1973, Prance et al. 16698 (NY). Para: estrada Santarém-Cuiaba (BR 163) km 780 de Cuiaba, 430 m, 54°54'W, 9°22'S, 29 May 1983, Silva 159 (MO, NY, US). Rond6nia: Costa Marques, Chapada dos Pareis, dist. Alto Floresta, estrada P-56, km 17, 62°63'W, 11°12'S, 15 June 1984, Cid et al. 4568 (NY); E. bank of Rio Madeira near junction of Rio Abuna, 21 July 1968, Prance et al. 6236 (NY). Solanum mite is the most common of the species of section Ptero- idea, forming large thickets in treefall gaps in the primary and secondary forest and along streams and roads in partial shade. Like S. anceps, it is basically an Amazonian species, but unlike S. anceps, S. mite occurs only in the southern part of the Amazon basin, not extending far north of the N. bank of the Rio Amazonas. Solanum mite is superficially similar to both S. conicum and S. uleanum, but can be differentiated easily from those species by its rounded fruit, pendent at maturity. Solanum mite can be hard to distinguish from S. conicum in flower, but the latter generally has larger flowers with petals held planar at anthesis, while S. mite has tiny ones with reflexed petals. Other differences from S. conicum are discussed with that species. Numbers of leaflets and size of leaves are extremely variable in S. mite, but leaflet shape is consistently obovate, with the terminal leaflet usually much larger and more enlarged in the distal 62 —-—..__. ————_—_—+ Qum Fig. 19S. savanillense. Habit, flowers, and fruit: Madsen & Elleman 75239, Madsen 85749, 85898 (AAU). third. The type specimen of S. pteleifolium (Martius s.n.) has ternate leaves with very large, broad leaflets. There exists however a range ofintermediates in both leaflet numbers and size: Maas et al. P12838 from Acre, Brazil and Plowman et al. 6401 from near Leticia on the Colombia/Peru border approach Martius’s collection in their broader leaflets, but given the range of variability in S. mite, we prefer to take a broad concept of the species. Many of the minor variants have been described as separate species by Bitter (see synonymy), but the range of variation in S. mite as recognized here encompasses all of these. Huge variability in leaf pubescence of collections made by one of us (S.K.) in Departmento San Martin, Peru, shows that pubescence S. KNAPP AND T. HELGASON density and occurrence is quite variable within populations of S. mite. In these collections, made in the Tarapoto area in 1986, no differences in phenology or other ecological characteristics were observed, and no morphological differences other than pubescence were seen. The nature of inheritance of this character is not known, but is likely to be relatively simple. 8. Solanum savanillense Bitter in Reprium nov. Spec. Regni veg. 12: 66 (1912). Type: Ecuador, Zamora-Chinchipe, Tambo de Savanilla, 18 December 1876, André 4565 (K!-holotype). Figs 4d, 5b, 19. REVISION OF SOLANUM SECTION PTEROIDEA 63 Fig. 20 Distribution of S. savanillense (star in circle) and S. trizygum (circles). Slender, wand-like shrub to 1.2 m tall. Stems c. 7 mm in diameter, fleshy and somewhat translucent, completely glabrous or with a few scattered uniseriate trichomes near the somewhat swollen nodes, the nodes occasionally dark purple, usually the stems green without conspicuous lenticels when dry. Leaves 10-17 x 7-16 cm, pinnate, with 1-2 pairs of leaflets, occasionally purple abaxially, sparsely pubescent abaxially with simple uniseriate trichomes c. 0.5 mm long, 4-5-celled, these denser along the veins, more densely pubes- cent adaxially, the trichomes c. | mm long and 8—10-celled; petiole 3.54 cm long; rachis of leaf minutely winged, sparsely pubescent with simple, uniseriate trichomes c. 0.05 mm long; lateral leaflets obovate, 10—12.5 x 3—5.5 cm, the base attenuate, the apex acumi- nate; petiolule 0.5—0.8 cm long; basal leaflets obovate, smaller, 5-10.5 x 1.7-4.2 cm, the lamina often narrower on the acroscopic side of the leaflet, the blade attenuate, the apex acuminate; petiolule c. 0.5 cm; terminal leaflet obovate, broader than any of the laterals, 6-16 x 3-7 cm, the base attenuate, the apex acuminate; petiolule c. 1 cm long. Inflorescence 1-3 cm long, simple, up to 3 rachis arising from a single leaf axil, bearing flowers only in the distal half, with only | or 2 flowers open at a time, but with c. 10-15 scars, sparsely pubescent with simple, uniseriate trichomes like those of the leaves. Buds c. 0.5 cm in diameter, globose to ovate, strongly exserted from the calyx tube. Pedicels at anthesis 0.5—0.7 cm long, c. 0.5 mm in diameter, nodding, glabrous or minutely pubescent. Flowers with the calyx tube 1.5—2 mm long, conical, the lobes c. 0.5—1 x 0.5 mm, deltate, splitting irregularly at the sinuses, with a prominent terminal projection, sparsely pubescent with trichomes like those of the inflorescence, the trichomes denser on the tip of the lobes; corolla 10-12 mm in diameter, white or purple (see discussion), lobed nearly to the base, the lobes planar at anthesis, densely papillate at the tips and margins; anthers 2.5—3 x c. | mm, poricidal at the tips, free portion of the filaments c. 0.5 mm long, the filament tube minute; ovary conical, glabrous; style 5—6 mm long, densely short- pubescent in the lower half; stigma clavate. Fruit an elongate, conical green berry, |—2 xc. | cm, the surface lightly rugose; fruiting pedicel 1—1.2 cm long, erect. Seeds c. 3.0 x 2.5mm, ovoid-reniform, bright green, c. 20 per fruit; epidermal cells highly sinuous and irregular, with anticlinal thickenings but without projections. COMMON NAMES AND USES. None recorded. DISTRIBUTION. S. Ecuador in cloud forest, from 2300-3000 m. (Fig. 20). SPECIMENS EXAMINED ECUADOR. Loja: Sin loc., 1876, André K694 (K); Parque Nacional Podocarpus, Nudo de Cajanuma; sendero del Centro de Informacion a las Lagunas del Compadre, 2830-2880 m, 21 November 1990, Gavilanes et al. 381 (AAU); Parque Nacional Podocarpus, Cajanuma, trail to El Mirador, 2800-3200 m, 4°07'S, 79°10'W, 20 October 1994, Knapp et al. 9044, 9045 (QCNE); Parque Nacional Podocarpus, above Nudo de Cajanuma around Centro de Informacién, 2800-3000 m, 79°10'W, 4°05'S, 6 September 1988, Madsen & Elleman75238 (AAU, LOJA); Parque Nacional Podocarpus, above Nudo de Cajanuma around Centro de Informacién, 2800-3000 m, 79°10'W, 4°05'S, 6 September 1988, Madsen & Elleman75239 (AAU); Parque Nacional Podocarpus, E. of Nudo de Cajanuma, just N. of Centro de Informacién, 2900 m, 79°10'W, 4°05'S, 31 January 1989, Madsen 85749 (AAU, LOJA, QCNE); Parque Nacional Podocarpus, above Nudo de Cajanuma around Centro de Informacién, 2800-3000 m, 79°10'W, 04°05'S, 14 May 1988, Allgaard et al. 74105 (AAU); Parque Nacional Podocarpus, at Cajanuma, S. of Loja, at Centro de Informaci6n, 2900 m, 79°10'W, 4°05'S, 31 May 1988, @llgaard 74539 (AAU, LOJA); Parque Nacional Podocarpus, E. of Nudo de Cajanuma, trail E. of Centro de Informaci6n, tocreston trail to Lagunas de Compadre, 2850-3050 m, 79°10'W, 4°05'S, 7 June 1988, Aligaard 74630 (AAU, LOJA). Zamora- Chinchipe: Pass between Loja and Zamora and along trail toward Zamora, 2360-2800 m, 29 July 1982, Clemants et al. 2252 (NY); Parque Nacional Podocarpus, road Loja-Zamora, just E. of pass, 2800 m, 79°07'W, 3°58'S, 15 March 1989, Madsen 85898 (AAU). 64 Solanum savanillense is superficially similar to many of the other pinnate-leaved members of section Pteroidea. It can be distin- guished easily, however, by its larger flowers, more elliptic, slightly less rugose fruits and its tall stature. This last character is impossible to ascertain from herbarium sheets, but in the field S. savanillense is quite distinct from S. mite or S. conicum, the only other members of this group to attain such sizes. The leaves of S. savanillense are more pubescent adaxially than abaxially, the reverse of the pattern in other members of the group. In 5-foliolate leaves of S. savanillense, the basal pair of leaflets is conspicuously smaller than the other pair, a characteristic not found elsewhere in the section. In Parque Nacional Podocarpus in southern Ecuador S. savanillense is polymorphic for both flower and stem colour but the leaves are monomorphic. In populations collected ascending the Nudo de Cajanuma, groups of plants were either green-stemmed and white-flowered or with purple nodes and purple flowers. Leaf undersides of neither morph were purple as occurs in S. anceps (see discussion under that species), perhaps indicating that the colour variation in S. savanillense is purely genetic in origin. Solanum savanillense grows in the primary forest understory in deep shade. Bitter, in describing S. savanillense, stated that the type was collected in Costa Rica. André never collected in Costa Rica, but was in southern Ecuador in December of 1876. The type specimen does not bear any annotation stating Costa Rica, so it is likely that Bitter made a mistake in transcribing label data or just confused the localities of specimens he saw at K. André’s locality Tambo de Savanilla probably corresponds to the present-day village of Sabanilla in the province of Zamora-Chinchipe or to the pass on the Loja to Zamora road (3°38'S, 79°05'W), which lies within the Parque Nacional Podocarpus. In Ecuador, nudo means a pass and tambo a stopping place, usually a small town. 9. Solanum trizygum Bitter in Reprium nov. Spec. Regni veg. 11: 470 (1912). Type: Venezuela, Distrito Federal, Colonia Tovar, Moritz 1644 (B-holotype, destroyed [F neg. 2702, F!, G!, GH!, MO!, NY!, US!, WIS!], fragment at F!; HBG!-lectotype; BM!, HBG!-isolectotypes). A Moritz collection (BM!, F!, GH!, W [F neg. 33118, F!, MO!, US!, WIS!]) distributed from W could also be type material, but as it is labelled “Colombia, leg. Moritz’ we have excluded it from consideration. Fig. 21. Solanum fraxinellum Bitter in Reprium nov. Spec. Regni veg. 11: 469 (1912). Type: Mexico, Veracruz, prope Mirador, 3000-3800 ft, Sartorius s.n. (W!-holotype [F neg. 33075, F!, G!]; G!-isotype [Morton neg. 8516, F!, GH!, NY!]). Solanum trizygum var. tetrazygum Bitter in Reprium nov. Spec. Regni veg. 11: 471 (1912). Type: Venezuela, sin loc., 27 June 1891, Eggers 13223 (C!-holotype). Solanum quinquejugum Bitter in Reprium nov. Spec. Regni veg. 11: 564 (1912). Type: Mexico, Puebla, Teziutlan, July 1866, Hahn s.n. (P!-holotype [F neg. 39200, G!]). Solanum pittieri Bitter in Reprium nov. Spec. Regni veg. 12: 66 (1912). Type: Costa Rica, Heredia, Alto del Roble, 2000 m, 1888, Pittier 18 (G!-holotype). Somewhat woody shrub to 1 m. Stems c. 5 mm in diameter, usually bright green and fleshy, completely glabrous, but occasionally with sparse reddish papillae (when dry) or simple uniseriate trichomes scattered near the nodes. Leaves 10-40 x 5-10 cm, pinnate, elliptic, with 3-6 (most commonly 4) pairs of leaflets, glabrous or minutely papillose adaxially, occasionally with a few uniseriate trichomes along the veins abaxially, occasionally purple abaxially and along the rachis; petiole 2-5 cm long; lateral leaflets lanceolate to elliptic, S. KNAPP AND T. HELGASON 4—15(-25) x 2-4 cm, with 6-8 pairs of primary veins, the base attenuate, only very occasionally oblique, the apex acuminate; petiolule 0.1 cm long; basal leaflets either equal to the laterals or somewhat smaller; terminal leaflet similar in shape to the laterals, 6— 20 x 3-6 cm, the base attenuate, the apex acuminate; petiolule c. 1 cm or less. Inflorescence axillary, 1-4(-9) cm long, simple, c. 2-4 arising from a single axil, bearing 2—3 flowers at a time, with 20-30 pedicel scars, glabrous or minutely papillate at the tip. Buds elliptic, the calyx strongly 5-ridged (in dry specimens) in early bud, c. 3 x 2 mm, strongly exserted from the calyx tube. Pedicels at anthesis 1— 1.5 cm, c. 0.5 mm in diameter at the tip, nodding. Flowers with the calyx tube 1—1.5 mm, open-conical, the lobes deltate to broadly triangular, 0.5—1 x 0.5—1 mm, glabrous; corolla 9-10 mm, greenish white or white, lobed nearly to the base, the lobes reflexed at anthesis, minutely papillose at the margins and tips; anthers 2—2.5 x 1—1.5 mm, poricidal at the tips, the free portion of the filaments 0.5— 1 mm long, the filament tube absent; ovary conical, glabrous; style 5—6 mm long, minutely papillose in the lower half; stigma minutely capitate, occasionally somewhat clavate. Fruit a conical, green berry, the beak not abruptly narrowed, c. 1—1.2 cm in diameter, |.2— 2.5 cm long, the surface rugose, the raised portions white, when ripe smelling distinctly of wintergreen; fruiting pedicels 1.5—2 mm long, c. 3 mm in diameter at the apex, erect. Seeds 3.0 x 2.0-2.5mm, ovoid-reniform to elliptic, greenish brown, c. 40 seeds per fruit; epidermal cells sinuous and irregular, with anticlinal thickenings but without projections. COMMON NAMES AND USES. Guatemala: ‘candelaria’ (Steyermark 33815, 35135, 37209, 37732, 48735, 51729); Venezuela: ‘ajicillo’ (Berry 1926). DISTRIBUTION. Montane and premontane forest from Mexico to the Cordillera de la Costa in Venezuela, absent from the Andes, from c. 600-3200 m. (Fig. 20). SPECIMENS EXAMINED MEXICO. ?Teotalcingo (Chinantla) Montes, 914m, June 184?, Galeottil 165 (BR). Chiapas: Mun. La Concordia, ElTriunfo Reserve, trail WSW from Palo Gordo towards Finca Catarina, 1850 m, 15°40'N, 92°51'W, 25 February 1990, Hampshire et al. 697 (BM). Oaxaca: Cerro Mirador, 15 km al NNW de Valle Nacional, 1000-1200 m, 96°22'W, 17°93'N, 16 October 1992, del Castillo 1520 (BM); Cerro Mirador, 15 km al NNW de Valle Nacional, 1000—1200 m, 96°22'W, 17°53'N, 27 April 1993, Manriquezet al. 3819 (BM). GUATEMALA. Alta Verapaz: Mountains along road between Tactic & the divide on road to the Tamahu, 1500-1600 m, 1 May 1941, Standley 90563 (F); Baja Verapaz: Nifio Perdido in San Jose road, N. 6 km, bordering Arroyo El] Caracol, 24 May 1977, Lundell & Contreras 20973 (F, MO); mountain of Purulha between La Union & Purulha, 1600 m, | October 1972, Molina R. & Molina 27734 (F). Huehuetenango: Vicinity of Maxbal, about 17 miles N. of Barillas, Sierra de los Cuchumatanes, 1500 m, 15 July 1942, Steyermark 48735 (F); Cerro Negro, 2 miles E. of Las Palmas, Sierra de los Cuchumatanes, 1600-2000 m, 31 August 1942, Steyermark 51729 (F); Quezaltenango: Along old road between Finca Pirineos and Patzulin, 1200-— 1400 m, 9 February 1941, Standley 86711 (F); lower S. facing slopes of Volcan Santa Maria, between Finca Pirineos and Los Positos, between Santa Maria de Jestis and Calahuaché, 1300-1500 m, 8 January 1940, Steyermark 33815 (F); W. slopes of Volcan Zunil, opposite Santa Maria de Jesus, 1500 m, 21 January 1940, Steyermark 35135 (F). San Marcos: | mile above Africa, c. 3.3 miles above Finca Armenia above San Rafael, 1600 m, 13 July 1977, Croat 40937 (NY); Finca Vergel, near Rodeo, 900 m, 15 March 1939, Standley 68905 (F); above Finca El Porvenir on “Todos Santos Chiquitos’, lower S. facing slopes of Volcan Tajumulco, 1300-1500 m, 7 March 1940, Steyermark 37209 (F); Between Finca El Porvenir and Loma Corona, 9 miles NW of El Porvenir, SW-facing slopes of Volcan Tajumulco, 1300-2000 m, 14 March 1940, Steyermark 37732 (F). HONDURAS. Santa Barbara: 10 kms W. de Lago Yojoa, 1500-2000 m, 88°0S'W, 14°55'N, 28 April 1973, Clewell & Hazlett 3859 (MO). REVISION OF SOLANUM SECTION PTEROIDEA Fig. 21S. trizygum. Habit: Skutch 3166 (A). Fruit: Lent 2788 (F). 65 66 COSTA RICA. Alajuela: La Palma de San Ramon, 1300 m, 23 October 1922, Brenes 3719 (F, NY); SanAntonio de San Ram6n, 850 m, 15 July 1927, Brenes 5625 (F); Cerro de La Muralla de San Ramén (El Socorro), 2 September 1927, Brenes 5704 (F); Santiago de San Ramon, 1150 m, 29 July 1937, Brenes 22613 (F); Cordillera Central, 7 miles N. of Carrizal, between Volcan Poas & Volcan Barba, 1850 m, 25 May 1976, Croat 35489 (MO); Monteverde, Cordillera de Tilaran, Reserva Vert. Atlantico, 1500-1580 m, 14 December 1976, Dryer 1069 (F); above Rio Gorrién, Bajos del Toro, 1550 m, 84°18'W, 10°13'N, 20 January 1974, Lent 3776 (MO); region of Zarcero, 1680 m, 29 September 1937, Smith A456 (F); Pueblo Nuevo, Cantén San Carlos, 1100 m, 15 April 1939, Smith 1900 (F); Stefano Ruiz, Cantén Llano Barito, 1650 m, 9 June 1941, Smith 2750 (F). Cartago: Near El Copey, in cloud forest area, Cordillera de Talamanca, 1800 m, 23 April 1949, Allen 16520 (F); hillside overlooking Rio Grande de Orosf about 3 km SE of Tapanti, 1400 m, 16 April 1967, Lent 822 (F); Tausito, 1400 m, 83°46'W, 10°46'N, 16 February 1974, Lent 3819 (F); 12 km S. of Turrialba by air, 4 km SE of Pejibaye along Rio Gato, 700 m, 83°42'W, 9°48'N, 16 April 1983, Liesner 14374 (MO). Guanacaste: SW slopes of Volcan Rincon de la Vieja &Volcén Santa Maria along the trail from Hda. Guachipelin, 1000 m, 85°21'W, 10°48'N, 30 July 1971, Burger & Pohl 7809 (F, MO). Heredia: Rio Vueltas (upper Rio Patria) on E. slope of Volcan Barba on the Carribean side, 1900 m, 84°04'W, 10°06'N, | April 1973, Gentry & Burger 2863 (F, MO, NY): Braulio Carrillo Park, Zuruqui, 1700-2000 m, March 1983, Gomez 20172 (MO); Finca La Selva, the OTS field station on the Rio Puerto Viejo just E. of its junction with the Rio Sarapiqui, along Q. El Sura between arboretum and station, 9 March 1980, Hammel 7983 (F, MO); Alto de Roble, 2000 m, May 1888, Pittier 18 (G); Vara Blanca de Sarapiqui, N. slope of Central Cordillera, 1500-1750 m, July-September 1937, Skutch 3166 (A, GH, K, MO, NY); Vara Blanca de Sarapiqui, N. slope of Central Cordillera, 1615 m, February 1938, Skutch 3614 (A, K, MO, NY); Cerro de las Caricias, N. of San Isidro, 2000-2400 m, 11 March 1926, Standley & Valerio 51987 (F); along the W. fork of the upper Rio Para Blanco beyond the road terminus of Calle Zurqui, 18 March 1974, Utley & Utley 701 (F). Lim6én: Cordillera de Talamanca, Atlantic slope, canyon of the Rio Sini, 1800-1900 m, 82°59'W, 9°13'N, 15 September 1984, Davidse & Herrera Ch. 29142 (MO). Puntarenas: Las Tablas, Rio Cotoncito, 10 December 1983, Chacon et al. 1811 (MO); Monteverde, Cordillera de Tilaran, 1520-1580 m, 12 February 1977, Dryer 1194 (F); about 2 km SE of Monteverde, on the Pacific watershed, 1500-1550 m, 84°48'W, 10°18'N, 18 March 1973, Gentry & Burger 2721 (F, MO); on and near the Continental Divide about 2-5 km E. & SE of Monteverde, 1580—1700 m, 10°18'N, 84°46'W, 17 March 1973, Gentry & Burger 2731 (F); Monteverde Cloud Forest Reserve, Cordillera de Tilaran (Pacific slope), 1500-1620 m, 20 January 1984, Linhart 155 (MO), 31 May 1985, Pounds 501 (MO), 13 July 1984, Pounds 274 (MO), 26 March 1984, Pounds 196 (MO). San José: Along the Rio Para Blanca (Pacific drainage), Cerros de Zuruqui, 1600-1800 m, 10°03'N, 84°O1'W, 6 February 1977, Burger et al. 10250 (F); Cordillera de Talamanca; Chirrip6 massif, Pacific slope, place along trail known asAbra, 2500 m, 2 April 1969, Davidse & Pohl 1529 (F); near Rio Hondura, 1150 m, 83°59'W, 10°04'N, 12 August 1972, Lent 2788 (F); vicinity of E] General, 880 m, August 1936, Skutch 2789 (GH, K, MO, NY), 1035 m, February 1939, Skutch 4147 (K, MO, NY); vicinity of Santa Maria de Dota, 1500-1800 m, 26 December 1925, Standley & Valerio 44055 (F); Alto de la Palma on Finca Porvenir, c. 5 km N. of San Jer6nimo, 1500 m, 18 August 1975, Utley & Utley 2902 (F, MO); Cordillera de Talamanca, about 25 km N. of San Isidro de El General along Pan American Highway, 3200 m, 29 January 1965, Williams et al. 25580 (F). PANAMA. Bocas del Toro: Rébalo trail, N. slopes of Cerro Horqueta, 1830-2130 m, 5 August 1947, Allen 4953 (F, G, MO); vicinity of Fortuna Dam, 1300-1400 m, 6 February 1987, Bohs & McPherson 2307 (GH); 7.2 miles beyond Campamento Chami (12+12 miles from Rio San Felix), 1500 m, 20 June 1986, D’Arcy 16328 (MO), D’Arcy 16343 (MO); Chiriqui border along Continental Divide on Carretera del Oleoductoc. 1 km N. of Quebrada Arena, IHRE Fortuna Hydroelectric project, 1150 m, 8°46'N, 82°12'W, 11 May 1982, Knapp 5064 (MO). Chiriqui: Along road in vicinity of branch in road to Cerro Colorado and Escopeta; above Rio San Felix near town of San Felix (c. 13 miles N. of Rio San Felix Bridge), 800-1200 m, 15 March 1976, Croat 33456 (MO); vicinity of Monte Azul, 1.4 miles N. of Entre Rios, on E. slopes of Cerro Punta, 3 miles by road from town of Cerro Punta, 2250 m, 25 November 1979, Croat 48589 (MO); along road between Gualaca and S. KNAPP AND T. HELGASON Fortuna Dam site, 10.1 miles NW of Los Planes de Hornito, 1250 m, 8°45'N, 82°17'W, 10 April 1980, Croat 50031 (MO); edge of Laguna de Volcan, 9 August 1972, D’Arcy & D’Arcy 6606 (GH, MO); road from Nueva California and Rio Serano c. 9 miles from Rfo Chiriqui Viejo, 1370 m, 7 April 1979, D'Arcy et al. 12988 (MO); between Palo Alto and top of ridge (divide) near Cerro Pate Macho, above Rio Palo Alto, 1640-2160 m, 18 March 1979, D'Arcy et al. 12647 (MO), D’Arcy 12672 (MO); Bajo Chorro, Boquete District, 1830 m, 6 February 1938, Davidson 63 (F, MO); Boquete, Finca Collins, 1520 m, 7 August 1967, Dwyer & Hayden 7661 (MO), Dwyer & Hayden 7670 (MO); Cerro Colorado, Bocas Road, 1500 m, 17 February 1977, Folsom & Collins 1765 (MO); slope of hill above camp at Fortuna Dam site, 1400-1500 m, 14 September 1977, Folsom et al. 5486 (MO); along trail from end of Rio Palo Alto road to Chiriqui border with Bocas del Toro Province near peak of Cerro Pate Macho, 2070 m, 20 November 1978, Hammel 5804 (BM, MO, NY); 9 miles from Rio Chiriqui Viejo bridge near Nueva California on road to Rio Sereno, 7 April 1979, Hammel et al. 6848 (MO); trail from Paso Respingo to Bajo Chorro, Cerro Punta to Boquete, 2225 m, 13 April 1979, Hammel et al. 7030 (MO); | km N. of Fortuna Lake, 1200 m, 82°13'W, 8°45'N, 3 March 1985, Hampshire & Whitefoord 286 (BM); c. 0.5 km E. of Cerro Pate Macho, headwaters of Rio Palo Alto, 1800— 2100 m, 82°21'W, 8°47'N, 12 November 1981, Knapp et al. 2108 (BM, NY); trail to Cerro Pate Macho, headwaters of Rio Palo Alto, above Palo Alto, 1700-2100 m, 82°22'W, 8°47'N, 15 March 1982, Knapp et al. 4260 (MO); Finca Collins, c. km 9.5 on the Quiel road above Boquete, 1830 m, 15 May 1971, Proctor 31944 (F); 6 miles above Cerro Punta on the Boquete Trail, 2300 m, 7 March 1974, Tyson 7144 (MO). Coclé: 2 miles N. of Cerro Pilén, 900 m, 16 March 1973, Leisner 724 (MO, F). Veraguas: E. side of mountain (Cerro Tute) W. of Escuela (Primer Basica, formerly Agricola) Alto de Piedra, c. 5 miles NW of Santa Fé, 760-850 m, 10 September 1982, D’Arcy 15003 (MO). VENEZUELA. sin. loc., November 1875, André K693 (K); sin. loc., Moritz s.n. (US). Aragua: Rancho Grande, pica detras del Hotel, Parque Nacional ‘H. Pittier’, May 1962, Agostini 48 (US); prope coloniam Tovar, 1854, Fendler 1016 (BR, G, GH, NY); Parque Nacional between Rancho Grande & Dos Riitos, 900 m, 19 May 1943, Killip & Lasser 37758 (A, US); Parque Nacional ‘Henri Pittier’ (Rancho Grande), trail to Pico Guacomayo, behind station, 1 100-1400 m, 67°42'W, 10°21'N, 27 October 1984, Knapp & Mallet 6852 (BH, MY, US, VEN); Parque Nacional Henri Pittier, Rancho Grande, trail to Toma, 1300 m, 4 October 1968, Plowman 1931 (GH, K); Rancho Grande, Parque Nacional H. Pittier, 3 February 1968, Walter & Walter 472 (B); in the forest of Rancho Grande, Parc Nacional, 1000 m, | December 1938, Williams & Alston 139 (BM, NY), Williams 10743 (F). Carabobo:Along Rio San Gian, al S. de Borburata, arriba de la Plant Eléctrica, entre Los Tanques y La Toma, 750 m, 27 March 1966, Steyermark & Steyermark 95161 (F). Distrito Federal: ?Galipan, 1846, Linden 128 (G); Bosque de Catuche, above Caracas, 1200-1800 m, 9 May 1913, Pittier6145 (US); Chacarito Gorge, around Caracas, 800-1000 m, 8 May 1921, Pittier 9508 (GH, NY, US); Catuche wood, 1200-1300 m, 22 January 1922, Pittier 10092 (GH, US, NY); Cerro Naiguata, laderas pendientes de lado del March que miran hacia el N. arriba del pueblo de Naiguata, vecinidad de Quebrada Frontina, 5 km al SW de los tanques de la Electricidad de Caracas (Cocuizal), 900-1100 m, 2 November 1963, Steyermark 91851 (F); Agua Negra, or above Caracas, 1500 m, December 1939, Williams 13624 (F). Falcén: Arriba de La Chapa, Sierra de San Luis, 1100 m, 18 January 1979, Flora Falc6n210 (WIS); Sierra San Luis, ridges around Hotel Parador, c. 7 km S. of Curimagua, 1300-1350 m, 69°35'W, 11°10'N, 28 September 1984, Knapp & Mallet6685 (BH, K, MY, VEN); Distrito Bolivar, Sierra de San Luis, Cerro Galicia, around TV antenna at summit, 1500 m, 11°11'N, 69°42'W, 29 March 1984, Plowman et al. 13440 (F, NY); Sierra de San Luis, vicinity of Hotel Parador, S. of La Tabla, 1450 m, 16 July 1967, Steyermark 98915 (US); Sierra San Luis, arriba del Hotel Parador, 1500 m, 25 August 1978, Wingfield & van der Werff 6574 (WIS). Miranda: Quebrada de las Comadres, near las Mostazas, 1100 m, November 1924, Allart 255 (NY); Campo Experimental Padrén — Estacién Experimental de Caucagua, 15 km al E. de Caucagua, 40 m, 22 January 1976, Berry 1926 (F); Dtto Paz Castillo, Municipo Reyes Cueta, Los Guayabitos, 1300-1490 m, 10°21°16”N, 66°38°36”W, 11 December 1987, Castillo & Bocaranda F. 2694 (MO); Quebrada de las Comadres near las Mostazas, 1100 m, November 1924, Pittier 255 (G); Parque Nacional de Guatopo, headwaters of Rio Grande, from Quebrada San Antonio to Fila de REVISION OF SOLANUM SECTION PTEROIDEA 5 om 67 Fig. 22 S. uleanum. Habit: Knapp & Mallet 6524 (K), Schunke V. 3898 (MO). Fruit: Schunke V. 5431 (MO). Rio Grande, between Santa Teresa and Alatagracio de Orituco, 6.5 km from Rancheria Mi Querencia, 600-700 m, 27 November 1961, Steyermark 90105 (US). Monagas: El Paramo, NE of Las Delicias, NE of Caripe, 1200-1450 m, 13 April 1945, Steyemark 62034 (F). Sucre: Peninsula de Paria, en el camino entre Los Pocitos de Santa Isabel a Roma, 10-15 km NW de Irapa, 700-1060 m, 13 July 1972, Dumontet al. VE-7649 (NY); Peninsula de Paria, Cumbre La Estrella, W. of Manacal (turnoff 13.2 km W. of Irapa) N. of El Paujil, 800-850 m, 62°41'W, 10°40'N, 17 October 1984, Knapp & Mallet 6771 (BH, F, K, MY, US, VEN); Peninsula de Paria, a lo largo de la Quebrada Nivardo, afluente de Rio Caverna, afluente de Rio Oscuro arriba de Mundo 68 Nuevo W. de Cerro de Humo, 700-750 m, 7 August 1966, Steyermark & Rabe 96140 (B, NY, US); Peninsula de Paria, cloud forest in tributary headwaters of Rio Cumana, SW of Cerro de Humo, vicinity of Manacal, 15 km (by air) NW of Irapa, 800 m, 62°39'W, 10°41'N, 29 November 1979, Stevermark & Liesner 120698 (NY). Solanum trizygum is superficially similar to S. conicum of the eastern Amazon, and is very closely related to that species (see Fig. 8). It differs from S. conicum in its more lanceolate leaflets that are very shortly petiolate or sessile, and its more elongate fruit. The fruit of S. trizygum also resembles that of S. savanillense, but is longer and has a more pronounced beak. Solanum trizygum is quite com- mon locally in the cloud forests of Central America. The distribution pattern of S. trizygum, occurring in Central America and in the cordillera de la Costa in Venezuela, is quite common in angiosperms (see Knapp, 1991). This may be indicative of some dispersal in the past, but alternatively may support geologi- cal hypotheses linking the Cordillera de la Costa with the Pacific plate (see Knapp, 19915 for a discussion). 10. Solanum uleanum Bitter in Reprium nov. Spec. Regni veg. 12: 139, pl. I (1913). Type: Brazil, Acre, Rio Acre, Porto Carlos, February 1911, Ule s.n. (B-syntype, destroyed; G!-lectotype). Both of the syntypes (Ule s.n. from Porto Carlos and Ule s.n. from San Franscico) cited by Bitter were destroyed at B. The collection from Porto Carlos is represented in the herbarium at G by a duplicate annotated in Bitter’s hand that matches the plate accompanying the original description. The second syntype, Ule s.n. collected in June 1911 at San Franscico may be the same specimen as the type of var. unipedunculatum below. The num- bering and dating of Ule’s collections is occasionally somewhat confused. Figs 2b, 4c, 22. Solanum uleanum var. unipedunculatum Bitter in Reprium nov. Spec. Regni veg. 12: 140 (1913). Type: Brazil, Acre, San Fran- cisco, May 1911, Ule 9756 (B holotype?, destroyed; K!-lectotype). Solanum uleanum var. gracilescens Bitter in Reprium nov. Spec. Regni veg. 12: 141 (1913).Type: Peru, San Martin, Cerro Campana, December 1855, Spruce 4462 (K!-holotype). Creeping herb, often tightly adhering to tree trunks and fallen logs, attaining up to 6 or 7 m in length. Stems c. 5 mm in diameter, copiously rooting at and between the nodes, pale greenish white, sparsely to densely pubescent with simple, uniseriate 5—6-celled trichomes 0.5—1 mm long, drying cateniforme. Leaves 3-15 x 2.5- 10 cm, pinnate, elliptic, with 3-7 pairs of leaflets, the petiole 0.8-6 cm long; rachis of leaf minutely winged, especially between the terminal leaflet and the ultimate pair, sparsely to densely pubescent with trichomes like those of the stem; lateral leaflets elliptic, 1.5—6 x 0.4-2 cm, sparsely to densely pubescent with simple uniseriate trichomes like those of the stems, these denser adaxially, especially along the veins, the base attenuate, winged onto the rachis, petiolule c. 2 mm long, the apex obtuse to rounded; basal pair of leaflets smaller than the laterals, the apex more rounded; terminal leaflet equal in size to the laterals, elliptic to obovate, strongly winged onto the rachis. Inflorescence 1-10 cm long, axillary, occasionally 2-3 separate rachis arising from a single axil, occasionally branched, with 3-4 open flowers at a time, with up to 100 scars unevenly spaced c. 0.5 mm apart, sparsely to densely pubescent with simple uniseriate trichomes 0.5—1 mm long, drying white. Buds c. 3 mm in diameter, globose soon becoming ellipsoid, strongly exserted from the minute calyx tube. Pedicels at anthesis 0.5—0.7 cm long, filiform, nodding, sparsely pubescent like the rest of the inflorescence. S. KNAPP AND T. HELGASON Flowers with the calyx tube c. 0.5 mm long, conical, the lobes 0.5 x 0.5—0.75 mm, quadrate with an apical projection, sparsely to densely pubescent with simple uniseriate trichomes, these denser on the apical projection; corolla 6-10 mm in diameter, greenish white, lobed nearly to the base, the lobes somewhat cucullate and slightly reflexed at anthesis, minutely papillose at the tips and along the margins; anthers 1.5—2.5 x 1—1.2 mm, poricidal at the tips, the pores lengthening to slits, free portion of the filaments c. 0.5 mm long, the filament tube minute and glabrous; ovary glabrous, conical; style 3— 4 mm long, straight, densely long-papillose in the lower 1/2; stigma capitate. Fruit a conical, green berry, 1—1.2 x 1.5—1.6 cm, the beak c. 5 mm long and not containing seeds, truncate at the tip; fruiting pedicel 0.8—1 cm long, hanging. Seeds c. 20 per berry, 3—3.5 x 1.5— 2.5 mm ovoid-reniform, greyish green to grey-brown; epidermal cell walls sinuous, thickened but without projections. COMMON NAMES AND USES. ‘pupu huasca’ (Kohn 1102 — used medicinally by mother to prevent bleeding from umbilical cord of baby), ‘yana barabacha panga’ (Shemluck & Ness 174 — mashed leaves applied to wound like hydrogen peroxide, juice also used [Quechua]), ‘ofa kthi’ (Vickers 143 — remedy for diarrhoea, plant crushed and mixed with water [Cofan]), ‘ahi inta 1k6’ (Vickers 273 — for stomach ache [Siona]). DISTRIBUTION. Eastern slopes of the Andes from central Ecuador to central Peru, from 200-1200 m elevation, usually growing in primary forest or at the edges of clearings (Fig. 23). SPECIMENS EXAMINED ECUADOR. Napo: Parque Nacional Yasunf, Pozo petrolero Daimi 2, 200 m, 76°11'W, 00°55'S, 26 May 1988, Cerén & Hurtado 4057 (MO, NY, QCNE); Carretera Hollin-Loreto-Coca, km 40, entre Rio Guamani y Rio Pucuno, 1200 m, 77°00'W, 00°40'S, 11 December 1987, Neill et al. 8107 (MO, NY), Ceron 2931 (MO, NY), Palacios 2219 (MO); Cant6n Tena, Rio Blanco community, headwaters of Rio Huambuno, 6 km NNW of Ahuano, 440 m, 77°40'W, 01°00'S, 19 February 1990, Kohn 1102 (QCNE). Pastaza: Rio Chic6, affluent of Rio Pastaza, village of Rio Chicé and vicinity, c. 10 km S. of Puyo, 3 km S. of Tarqui, 1000 m, 77°55'W, 01°03'S, August 1979, Shemluck & Ness 174 (F). Sucumbios: Rio Aguarico, Shushufindi, 23 February 1975, Vickers 143 (F); Shushufindi, 18 July 1979, Vickers 273 (F); San Pablo de los Secoyas, 300 m, 76°21'W, 00°15'S, 4 August 1981, Brandbyge et al. 32965 (AAU). PERU. Amazonas: Pongo de Manseriche, Rio Santiago & Rio Marafion, c. 77°30'W, anno 1924, Tessmann 3890 (G, NY). Huanuco: Pachitea, Codo de Pozuzo, alluvial fan floodplain of Rio Pozuzo after it emerges from mountains, trail to NW behind settlement, 450 m, 75°25'W, 9°40'S, 18 October 1982, Foster 9269 (MO); camino a Ayamiria a 2 km de Miel de Abeja, 300-400 m, 20 January 1967, Schunke V. 1538 (F); Bosque Nacional de Iparia, a lo largo del Rio Pachitea cerca del Campamento Miel de Abeja, 1 km arriba del pueblo de Tournevista 0 unos 20 km arriba del confluencia con el Rio Ucayali, 300-400 m, 28 February 1968, Schunke V. 1696 (F, GH, K, NY, US). Loreto: Prov. Coronel Portillo, Padre Abad, Granja del Sr Barrera, NE de la chacra de Cesar Vela (Aguaytia), 295 m, 22 October 1972, Schunke V. 5431 (MO, WIS). Pasco: Oxapampa, km 28 Repartici6n-Iscozacin (km 86 Villa Rica-Iscozacin-Pto. Mairo), Rio La Raya near Ameusha com- munity of Laguna, 350 m, 75°10'W, 10°20'S, 22 August 1984, Knapp & Mallet 6654 (K, NY, US, USM); Oxapampa, trail from Rio Iscozacin to Ameusha community of Villa América, Rio Palcazu Valley, 340 m, 75°15'W, 10°12'S, 31 August 1984, Knapp & Salick 6669 (K, US, USM); Oxapampa, km 15 of Palcazu road, (km 73 Villa Rica-Iscozaciin-Pt. Mairo) along Rio Palcazu, 380 m, 75°10'W, 10°21'S, 17 August 1984, Knapp & Mallet 6645 (BH, US, USM). San Martin: San Martin, km 28 of Tarapoto-Yurimaguas road, 650 m, 76°15'W, 6°25'S, 20 June 1984, Knapp & Mallet 6524 (F, K, MO, NY, US, USM); km 28 of Tarapoto-Yurimaguas road, 750-800 m, 76°19'W, 6°27'S, 23 September 1986, Knapp & Mallet8394 (MO); Quebrada de Ishichimi cerca a Tocache, 400 m, 12 March 1978, Schunke V. 10020 (MO); road by Rio Tocache, Dtto Tocache Nuevo, 12 April 1970, Schunke V. 3898 (F, G, NY); al W. del Vivero del Instituto Agropecuario de REVISION OF SOLANUM SECTION PTEROIDEA 69 Fig. 23 Distribution of S. uleanum. Tocache, 400 m, 10 November 1969, Schunke V, 3602, (F, NY, US); Cerro Monte, nr. Tarapoto, 1855, Spruce 4462 (K). Ucayali: Trail from Quebrada Shesha, (tributary of Rio Abujao) to base of Cerro Las Cachoeiras, c. 70 km NE of Pucallpa, 300-400 m, 73°55'W, 8°02'S, 24 June 1987, Gentry & Diaz 58484 (MO, USM). Solanum uleanum is certainly one of the most beautiful of the species of section Pteroidea. Its small size and peculiar (but found elsewhere in the section, see S. anceps) creeping habit make it conspicuously different from the other pinnate-leaved species. Even so, it can be difficult to distinguish on the herbarium sheet. Solanum uleanum differs from S. conicum and S. mite (both of which are sympatric with S. uleanum) in its more rounded, smaller leaflets which are more winged onto the rachis (i.e. without a petiolule) than in other pinnate species and greenish flowers. Fruiting specimens of S. uleanum are very rare, but its fruit, with an elongate non-seed- bearing beak is distinctive. As with almost all other members of the section, Solanum ule- anum possesses great variability in pubescence density. The type specimen (a photograph in the original publication and the lectotype at G) is densely hairy, while other collections are almost glabrous. There appear to be no environmental factors influencing this, but more detailed field studies could help clarify the situation. Solanum uleanum often grows up trees at the edge of gaps or clearings in the forest, and individuals growing in the deep shade of the understory have much thinner, more membranous leaves. EXCLUDED SPECIES 1. Solanum cormanthum Vell. (synonym of S. caavurana Vell. = section Geminata (G. Don) Walp.) 2. Solanum lacteum Vell. (affinities and identification unknown, no type specimen exists and plate in Vellozo resembles no known species of Solanum) 3. Solanum laurinum Dunal (synonym of S. decorticans Sendtn. = section Lysiphellos Bitter) 4. Solanum loxophyllum Bitter (= section Anarrhicomenum Bit- ter) 5. Solanum marantifolium Bitter (= section Geminata (G. Don) Walp.) 6. Solanum pentaphyllum Bitter (= section Herpystichum Bitter) 7. Solanum robustifrons Bitter (= section Geminata (G. Don) Walp.) ACKNOWLEDGMENTS. Many people have helped us in the course of this study, but we would particularly like to thank the following: both the late M.D. Whalen and the late T. Plowman for their inspiration in the study of solanums; D. Neill and T. Nufiez of QCNE and P. Wilson for help and support in Ecuador; B. Leén, G. Lamas and the people of Tarapoto for help and support in Peru; S. France for the illustrations; the staff of the EM Unit at The Natural History Museum for technical help; the staff at the Photographic Unit of The Natural History Museum, especially C. Keates and D. Adams; V. Persson and J.R. Press of the Botany Department of The Natural History 70 Museum for help with illustrations; K. Adamson, A. Hume, and E. Smith for help with labelling and returning loans; J. Wheeler for carefully checking the exsiccatae; N. Turland for help with the databases; and J. Mallet for being the senior author’s constant field companion. We would also like to thank the curators of the herbaria cited in the text for loan of specimens, and for permission to sample seeds from the specimens in their care. REFERENCES Anderson, G.J. 1977. The variation and evolution of Solanum section Basarthrum. II. Brittonia 29: 116-128. Aublet, J.B.C.F. 1775. Histoire des plantes de la Guiane frangoise. 1-4, Paris. Barboza, G.E. & Hunziker, A.T. 1991. Estudios sobre Solanaceae XXXI. Peculiaridades del androceo de interés taxondémico en Solanum. Kurtziana 21: 185- 194. Beccaloni, G.W. 1995. Studies on the ecology and evolution of Neotropical ithomiine butterflies (Nymphalidae: Ithomiinae). Unpublished Ph.D. thesis, University of London. Bell, A.D. & Dines, T.D. 1995. Branching patterns in the Solanaceae. /n P.C. Hoch & A.G. Stephenson (Eds), Experimental and molecular approaches to plant biosystematics: 157-172. St Louis, MO. Bitter, G. 1912. XVI. Weitere Polybotryon-Arten. Reprium nov. Spec. Regni veg. 11: 469473. 1921. XCV. Aufteilung der Gattung Bassovia (im Dunalischen Sinne) zwischen Solanum, Capsicum und Lycianthes. Reprium nov. Spec. Regni veg. 18: 328-335. Bohs, L. 1990. The systematics of Solanum section Allophyllum (Solanaceae). Ann. Mo. bot. Gdn 77: 398-409. 1994. Cyphomandra (Solanaceae). Flora Neotropica 63: 1-175. — 1995. Transfer of Cyphomandra (Solanaceae) and its species to Solanum. Taxon 44: 583-587. Brown, K.S., Jr. 1987. Chemistry at the Solanaceae/Ithomiinae interface. Ann. Mo. bot. Gdn 74: 359-397. Buchmann, S.L. 1983. Buzz pollination in angiosperms. Jn C.E. Jones & R.J. Little (Eds), Handbook of experimental polination biology: 73-113. New York. Child, A. 1979. A review of branching patterns in the Solanaceae. In J.G. Hawkes, R.N. Lester & A.K. Skelding (Eds), The biology and taxonomy of the Solanaceae: 345— 356. London. — 199]. Life form and branching within the Solanaceae. /n J.G. Hawkes, R.N. Lester, M. Nee & N. Estrada R. (Eds), Solanaceae III; taxonomy, chemistry, evolu- tion: 151-160. Kew, Richmond. Danert, S. 1958. Die Verzweigung der Solanaceen im reproduktiven Bereich. Abh. dt. Akad. Wiss. Berl. 1957(6): 1-183. 1967. Die Verzweigung als infragenerisches Gruppenmarkel in der Gattung Solanum L. Kulturpflanze 15: 275-292. 1970. Infragenerische Taxa der Gattung Solanum L. Kulturpflanze 18: 253-297. D’Arcy, W.G. 1972. Solanaceae studies II: Typification of subdivisions of Solanum. Ann. Mo. bot. Gdn 59: 262-278. 1991. The Solanaceae since 1976, with a review of its bibliography. Jn J.G. Hawkes, R.N. Lester, M. Nee & N. Estrada R. (Eds), Solanaceae III: taxonomy, chemistry, evolution: 75-138. Kew, Richmond. Don, G. 1838. Solanum. A general system of gardening and botany 4: 397-442. Drummond, B.A. III & Brown, K.S., Jr. 1987. Ithomiinae (Lepidoptera: Nymphalidae): summary of known larval food plants. Ann. Mo. bot. Gdn 74: 341— 358. Dunal, M.-F. 1813. Histoire naturelle, médicale et economique de Solanum et des genres qui ont étés confundus avec eux. Montpelier. 1816. Solanorum generumque affinium synopsis. Montpelier. 1852. Solanaceae. In A.P. de Candolle (Ed.), Prodromus systematis naturalis regni vegetabilis 13(1): 1-690. Edmonds, J.M. 1982. Epidermal hair morphology in Solanum section Solanum. Bot. J. Linn. Soc. 85: 153-167. 1983. Seed coat structure and development in Solanum section Solanum. Bot. J. Linn. Soc. 87: 229-246. Endress, P.K. 1996. Diversity and evolutionary trends in angiosperm anthers. /n W.G. D’Arcy & R.C. Keating (Eds), The anther: form, function and phylogeny: 92-110. S. KNAPP AND T. HELGASON Cambridge. Farris, J.S. 1988. Hennig86: version 1.5. Published by the author, Jamaica Plains. — 1989. The retention index and the rescaled consistency index. Cladistics 5: 417- 419. Forey, P.L. 1992. Formal classification. /n P.L. Forey, C.J. Humphries, I-L. Kitching, R.W. Scotland, D.J. Seibert & D.M. Williams (Eds), Cladistics: a practical course in systematics: 160-169. Oxford. Gilli, A. 1970. Bestimmungsschiissel der Subgenera und Sektionen der Gattung Solanum. Reprium Spec. nov. Regni veg. 81: 429-435. Greuter, W. (Chairman of the Editorial Committee), Barrie, FR., Burdet, H.M., Chaloner, W.G., Demoulin, V., Hawksworth, D.L., Jorgensen, P.M., Nicolson, D.H., Silva, P.C., Trehane & McNeill, J. (Secretary to the Editorial Committee). 1994. International code of botanical nomenclature. Regnum vegetabile 131. K6nigstein. Hawkes, J.G. 1990. The Potato: evolution, biodiversity and genetic resources. London. Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (Eds) 1990. Index herbariorum. 8th ed. Part 1. Regnum veg. 120. Den Haag. Knapp, S. 1983. Sectional nomenclature in Solanum (Solanaceae). Taxon 32: 635- 636. — 1986a. A revision of Solanum section Geminata (G. Don) Walpers. Unpublished Ph.D. thesis, Cornell University. —— 1986b. Reproductive biology of Solanum section Geminata in a Costa Rican cloud forest. Jn W.G. D’ Arcy (Ed.), Solanaceae: biology and systematics: 253-263. New York. 1989. A revision of the Solanum nitidum group (section Holophylla pro parte): Solanaceae. Bull. Br, Mus. nat. Hist. (Bot.) 19: 63-102. 1991a. A revision of the Solanum sessile species group (section Geminata pro parte: Solanaceae). Bot. J. Linn. Soc. 105: 179-210. 1991b. Acladistic analysis of the Solanum sessile species group (section Geminata pro parte: Solanaceae). Bot. J. Linn. Soc. 106: 73-89. Persson, V. & Blackmore, S. [In press]. Pollen morphology and evolution of dioecy in Solanum. Pl. Syst. Evol. Lester, R.N. & Durrands, P. 1984. Enzyme treatment as an aid in the study of seed surface structures of Solanum species. Ann. Bot. 53: 129-131. Nelson, G. 1974. Classification as an expression of phylogenetic relationships. Syst. Zool. 22: 344-359. Nixon, K. & Carpenter, J. 1993. On outgroups. Cladistics 9(4): 413-426. Olmstead, R. & Palmer, J. 1991. Chloroplast DNA and systematics of the Solanaceae. In J.G. Hawkes, R.N. Lester, M. Nee & N. Estrada R. (Eds), Solanaceae III: taxonomy, chemistry, evolution: 161-168. Kew, Richmond. Punt, W. & Monna-Brands, M. 1980. The northwest European pollen flora, 8. Solanaceae. /n W. Punt & G.C.S. Clarke (Eds), The northwest European pollen flora IT. \-30. Amsterdam. Roe, K. 1971. Terminology of hairs in the genus Solanum. Taxon 20: 501-508. Ruiz Lopez, H. & Pavén, J. 1799. Pentandria monogynia. Solanum. Flora peruviana et chilensis 2: 31-41. Madrid. Seithe, A. 1962. Die Haararten der Gattung Solanum L. und ihre taxonomische Verwertung. Bot. Jb. 81: 261-336. 1979. Hair types as taxonomic characters in Solanum. In J.G. Hawkes, R.N. Lester & A.K. Skelding (Eds), The biology and taxonomy of the Solanaceae: 307-319. London. & Anderson, G.J. 1982. Hair morphology and relationships of species in Solanum section Basarthrum. Pl. Syst. Evol. 139: 229-256. Souéges, R. 1907. Développment et structure du tégument seminal chez les Solanacées. Annis. Sci. nat. Botanique ix, 6: 1-124. Spooner, D.M., Anderson, G.J. & Jansen, R.K. 1993. Chloroplast DNA evidence for the interrelationships of tomatoes, potatoes, and pepinos (Solanaceae). Amer. J. Bot. 80: 676-688 Spruce, R. 1908. Notes of a botanist on the Amazon and Andes. 2. (A.R. Wallace, ed.). London. Walpers, W.G. 1844. Solanaceae. Repertorium botanices systematicae 3: 38-100. Watrous, L. & Wheeler, Q.C. 1981. The outgroup method of phylogeny reconstruc- tion. Syst. Zool. 30: 1-21. Whalen, M.D. 1984. Conspectus of species groups in Solanum subgenus Leptostemonum. Gentes Herb. 12: 179-282. — & Costich, D.E. 1986. Andromonoecy in Solanum. In W.G. D’Arcy (Ed.), Solanaceae: biology and systematics: 284-302. New York. & Heiser, C.B., Jr. 1981. Taxonomy of Solanum section Lasiocarpa. Gentes Herb. 12: 41-129. REVISION OF SOLANUM SECTION PTEROIDEA EXSICCATAE Agostini, G. 48 (trizygum ). Allard, H.A. 6236 (mite); 20481 (mite); 20850 (anceps); 22077 (anceps); 22116 (anceps); 22522 (anceps). Allart, A. 255 (trizygum); 335 (ternatum). Allen, PH. 4953 (trizygum); 16520 (trizygum). André, E. s.n. (conicum); K693 (trizygum); K694 (savanillense); 4565 (savanillense). Asplund, E. 12043 (mite). Aublet, J.B.C.F. s.n. (anceps). Ayala, F. et al. 2543 (anceps). Baker & Trushell 6099 (anceps). Baker et al. 5651 (anceps). Bang, M. 2248 (mite); 2513 (anceps); 2526 (anceps). Barbour, P. 3831 (mite); 3829 (ternatum); 4160 (ternatum); 4800 (mite); 5461 (mite). Beck, St.G. 4881 (ternatum); 7361 (mite); 7498 (anceps). Bell, D. & Wiser, S. 88-8 (mite); 88-40 (mite). Belshaw, C.M. 3089 (anceps); 3109 (mite). Bensman, R. 148 (anceps). Berry, PE. 1926 (trizygum). Boeke, J.D. 1200 (mite). Bohs, L. & McPherson, G. 2307 (trizygum). Bohs, L. & Schunke V., J. 2168 (mite). Brandbyge, J. & Asanza C., E. 30829 (anceps); 31783 (anceps); 31824 (anceps); 31873 (anceps); 31927 (anceps); 32365 (anceps). Brandbyge, J. & Balslev, H. 42280 (anceps). Brandbyge, J. et al. 32965 (uleanum). Brenes, A.M. 3719 (trizygum); 5625 (trizygum); 5704 (trizygum); 22613 (trizygum ). Britton, N.L. & Rusby, H.H. 1210 (anceps); 2513 (anceps); 2526 (anceps). Buchtien, O. 1249 (mite); 1287 (anceps); 1288 (anceps); 1289 (anceps); 1438 (mite); 7462 (ternatum); 7470 (anceps). Burger, W.C. & Pohl, R.W. 7809 (trizygum). Burger, W.C. et al. 10250 (trizygum). Calderon, C.E. et al. 2855 (anceps). Cardenas, M. 1168 (mite); 2046 (mite). Castillo, A & Bocaranda, F: 2694 (trizygum). Castillo, M. del 1520 (trizygum). Cazalet, P-C.D. & Pennington, T.D. 7676 (mite). Ceron, C.E. 2931 (uleanum); 7378 (anceps); 6389 (mite). Ceron, C.E. & Ceroén, M. 4604 (mite). Ceron, C. E. & Hurtado, F. 4057 (uleanum). Ceron, C.E. & Iguago, C. 5430 (anceps). Chacon, I.A. et al. 1811 (trizygum). Cid, A.C. et al. 4568 (mite); 4829 (anceps). Clemants, S.E. et al. 2252 (savanillense). Clewell, A. & Hazlett, D. 3859 (trizygum). Cowan, R.S. & Lindeman, J.C. 39020 (anceps). Croat, T.B. 18651 (anceps); 21039 (mite); 33456 (trizygum); 35489 (trizygum); 40937 (trizygum); 48589 (trizygum); 50031 (trizygum); 50575 (anceps); 51014 (mite); 51156 (mite); 51262 (anceps); 58022A (anceps); 58041 (anceps); 58622 (anceps); Cuatrecasas, J. 813 (mite). D'Arcy, W.G. 12672 (trizygum); 15003 (trizygum); 16328 (trizygum); 16343 (trizygum); 16506 (ternatum). D'Arcy, W.G. & D’Arcy, J.J. 6606 (trizygum). D'Arcy, W.G. et al. 12647 (trizygum); 12988 (trizygum). Daly, D.C. et al. 6118 (anceps); 6133 (anceps). Davidse, G. & Herrera Ch., G. 29142 (trizygum). Davidse, G & Pohl, R.W. 1529 (trizygum). Davidson, C. 3487 (mite). Davidson, M.E. 63 (trizygum). Diaz, C. & Beltran, 3335 (anceps). Diaz, C. & Jaramillo, N. 576 (anceps). Diaz P. & Melief, B. 2952 (ternatum). Dombey, P. s.n. (ternatum). Dorr, L.J. et al. 6816 (anceps). at Dryer, V.J. 1069 (trizygum); 1194 (trizygum). Dudley, T.R. 10152 (ternatum). Dumont, K. et al. VE-7649 (trizygum). Dwyer, J.D. & Hayden, M.V. 7661 (trizygum); 7670 (trizygum). Eggers, H.F-A. Baron von 13223 (trizygum). Ellenburgh, H. 3889 (anceps). Emmons, L. 81 (anceps); 132 (anceps); 146 (anceps). Encarnacion, F. 26268 (anceps). Espinal T., S. & Ramos, J. 2943 (anceps). Fendler, A. 1016 (trizygum); 1017 (ternatum). Fernandez C., J. 8299 (anceps). Ferreyra, R. 800 (mite); 1575 (mite); 2138 (mite). Feuillet, C. et al. 10178 (anceps); 10230 (anceps). Flora Falcén 210 (trizygum). Folsom, J.P. & Collins, L. 1765 (trizygum). Folsom, J.P. et al. 5486 (trizygum). Foresta, H. de. H.F 656 (anceps). Foster, R.B. 2411 (anceps); 8592 (anceps); 9198 (mite); 9269 (uleanum); 9298 (anceps). 9355 (ternatum). Foster, R.B. & Terborgh, J. 6071 (anceps); 6222 (anceps). Foster, R.B. et al. 3312 (anceps); 8969 (anceps); 10481 (anceps). Franco, P. et al. 1876 (anceps). Galeotti, H. 1165 (trizygum). Gavilanes et al. 381 (savanillense). Gentry, A. 31046 (anceps). Gentry, A. & Daly, D. 18773 (mite). Gentry, A. & Diaz, C. 58484 (uleanum). Gentry, A. & Horna, M. 29521 (anceps). Gentry, A. et al. 18876 (anceps); 21185 (anceps); 23086 (mite); 29558 (terna- tum); 29790 (anceps); 30871 (incurvum); 36396 (anceps); 45399 (anceps); 46254 (mite); 47779 (ternatum); 53970 (ternatum); 61713 (mite). Gentry, J.L. & Burger, W.C. 2721 (trizygum); 2731 (trizygum); 2863 (trizygum). Gomez, L.D. 20172 (trizygum);. Goudot 136 (anceps). Granville, J. de 2374 (anceps); 4944 (anceps); B5339 (anceps); 7165 (anceps); 7686 (anceps). Granville, J. de et al. 7502 (anceps); 8704 (anceps); 9975 (anceps); 10842 (anceps). Grubb, P.J. et al. 1210 (anceps). Gudino, E. et al. 1008 (anceps). Hahn, L. s.n. (trizygum). Hamilton, A.C. & Holligan, P.M. 1069 (incurvum); 1078 (ternatum). Hammel, B. 5804 (trizygum); 7983 (trizygum). Hammel, B. et al. 6848 (trizygum); 7030 (trizygum). Hampshire, R. & Whitefoord, C. 286 (trizygum). Hampshire, R. et al. 697 (trizygum). Harling, G. & Andersson, L. 21373 (anceps). Hart, J. 134 (mite). Hurtado, F. 625 (ternatum). Hurtado, F. & Alvarado, A. 1121 (ternatum). Iltis, H.H. & Iltis, C.M. 284 (mite). Irwin, H.S. et al. 48077 (anceps). Isern, J. 2241 (mite). Jaramillo, J. et al. 31719 (mite). Kayap, 536 (anceps); 575 (anceps); 1347 (anceps). Killip, E.P. & Lasser, T. 37758 (trizygum). Killip, E.P. & Smith, A.C. 20235 (ternatum); 23055 (conicum); 23561 (mite); 23839 (anceps); 25331 (mite); 25811 (ternatum); 26140 (mite); 26221 (anceps); 26261 (anceps); 23939 (anceps); 26239 (anceps); 27329 (anceps); 27614 (mite); 27760 (mite); 28103 (mite);29584 (anceps); 29076 (anceps); 29420 (anceps); 29493 (anceps). Klug, G. 2864 (anceps); 2872 (mite); 3665 (ternatum); 3686 (mite); 3757 (anceps); 4252 (mite). Knapp, S. 5064 (trizygum); 6592 (mite); 7857 (anceps); 7905 (angustialatum); 8012 (mite); 8264 (mite); 8277 (angustialatum). Knapp, S. & Alcorn, P. 7331 (mite); 7332 (mite); 7541 (mite); 7792 (angustialatum). Knapp, S. & Mallet, J. 6183 (anceps); 6279 (anceps); 6303 (anceps); 6396 (anceps); 6425 (conicum); 6427 (anceps); 6435 (conicum); 6436 72 (anceps); 6452 (conicum); 6456 (conicum); 6476 (anceps); 6484 (mite); 6486 (mite); 6524 (uleanum);6526 (mite); 6538 (mite); 6553 (conicum); 6554 (mite); 6555 (anceps); 6561 (mite); 6590 (chamaepolybotryon); 6623 (mite); 6626 (ternatum); 6629 (anceps); 6631 (conicum); 6632 (mite); 6639 (anceps); 6644 (anceps); 6645 (uleanum); 6654 (uleanum),; 6655 (anceps); 6658 (anceps); 6664 (conicum); 6685 (trizygum); 6771 (trizygum); 6852 (trizygum); 6931 (mite); 7027 (mite); 7036 (mite); 7065 (mite); 7068 (mite); 7086 (mite); 7087 (mite); 7212 (mite); 8394 (uleanum); 8440 (mite); 8567 (angustialatum). Knapp, S & Salick, M.J. 6667 (conicum); 6669 (uleanum). Knapp, S. et al. 2108 (trizygum); 4260 (trizygum); 6324 (mite); 6473 (mite); 7218 (mite); 7506 (anceps). Kohn, E. 1102 (uleanum); 9198 (mite). Krukoff, B.A. 1599 (anceps); 4642 (mite). Lawrance, A.E. 345 (anceps); 485 (anceps); 645 (anceps). Lechler, W. 2440 (ternatum); 2464 (anceps). Leisner, R. 724 (trizygum); 14374 (trizygum). Lent, R.W. 822 (trizygum); 2788 (trizygum); 3776 (trizygum); 3819 (trizygum). Leprieur, M. s.n. anno 1859 (anceps). Lewis, W.H. et al. 13646 (conicum); 13738 (anceps); 13771 (ternatum); 13898 (anceps). Liberman, M. 262 (anceps). Lindeman, J.C. 535 (anceps). Linden, J.J. 128 (trizygum). Linhart, Y.B. 155 (trizygum). Lleras, E. et al. P17286 (anceps). Lowrie, S.R. et al. 331 (anceps); 441 (anceps). Lundell, C.L. & Contreras, E. 20973 (trizygum). Luteyn, J.L. & Dorr, L.J. 13699 (ternatum). Maas, P.J.M. et al. P12838 (mite); P12903 (anceps). Macbride, J.F. 427 (ternatum); 4001 (mite); 4134 (conicum); 4243 (anceps); 4279 (ternatum); 4491 (incurvum); 4676 (mite); 4698 (ternatum); 5267 (mite); 29722 (anceps). MacBryde, B. & Dwyer, J.D. 1367 (anceps). Madsen, J.E. 75238 (savanillense); 85749 (savanillense); 85898 (savanillense). Madsen, J.E. & Elleman, L. 75239 (savanillense). Maguire, B. et al. 46080A (anceps); 54407 (anceps). Manriquez, G.I. et al. 3819 (trizygum). Martin, R. et al. 1619 (mite). Martius, K.F-P. von s.n. (mite). Matthews, A. s.n. (ternatum); 1967 (anceps). McDaniel, S. & Rimachi Y., M. 18383 (mite); 18903 (mite). Mexia, Y. 8326a (mite). Molina R., A & Molina, A.R. 27734 (trizygum). Moritz, J.W.K. s.n. (trizygum); 1028 (ternatum); 1644 (trizygum). Nee, M. 31504 (mite); 34977 (anceps); 35480 (mite); 36036 (mite); 36603 (mite); 37200 (mite); 37315 (mite); 38119 (mite); 39259 (mite); 39355 (mite); 39570 (mite). Nee, M. & Saldias P., M. 36888 (mite). Nee, M. et al. 35433 (mite). Neill, D. et al. 8107 (uleanum). Nifiez, P. 5770 (anceps); 6473 (mite). Nifnez, P. & Phillips, O. 10464 (anceps). Nufiéz, P. et al. 10555 (mite). Allgaard, B. 74539 (savanillense); 74630 (savanillense); 74954 (anceps); 98451 (mite). Allgaard, B. et al. 74105 (savanillense). Palacios, W. 2219 (uleanum); 2222 (anceps); 16607 (conicum); 0299 (anceps). Palacios, W. et al. 8188 (ternatum); 7684 (mite). Pearce, R. s.n. (ternatum); 135 (ternatum); 144 (ternatum). Pennell, F-W. 14012 (ternatum). Philipson, W.R. et al. 2205 (anceps). Pires, J.M. 10062 (anceps). Pittier, H. 18 (trizygum); 255 (trizygum); 6145 (trizygum); 9508 (trizygum); 10092 (trizygum). Plowman, T. 1931 (trizygum); 2129 (anceps); 3831 (mite); 5906 (anceps). Plowman, T. & Davis, E.W. 4806 (conicum); 5006 (anceps). S. KNAPP AND T. HELGASON Plowman, T. & Ramirez R., M. 11212 (mite). Plowman, T. & Schunke V., J. 7394 (ternatum); 11509 (anceps). Plowman, T. et al. 6401 (mite); 13440 (trizygum). Poeppig, s.n. (anceps); 1469 (anceps). Pounds, W.Z. 196 (trizygum); 274 (trizygum); 501 (trizygum). Prance, G.T. et al. 2833 (anceps); 2908 (mite); 2955 (mite); 3541 (anceps); 6236 (mite); 12573 (anceps); 16698 (mite). Prévost, M.F. 304 (anceps). Prévost, M.F. & Sabatier, D. 2422 (anceps). Proctor, G.R. 31944 (trizygum). Quinones, L. 1045 (anceps). Ramos, J.E. 2943 (anceps). Rierm. 919 (anceps). Rimachi Y., M. 507 (mite); 876 (mite). Romero Castaneda, R. 7067 (ternatum). Ruiz, H. & Pavon, J. s.n. (ternatum); s.n. (anceps); s.n. (mite); s.n. (conicum). Rusby, H.H. 578 (mite); 766 (anceps); 800 (mite); 813 (mite). Sanchez Vega, I. 4895 (ternatum). Sandeman, C. 5043 (mite); 5270 (anceps). Sartorius, C.C.W. s.n. (trizygum). Schunke, J.M. 280 (anceps); 2432 (anceps). Schunke V., J. 1280 (mite); 1414 (anceps); 1538 (uleanum); 1696 (uleanum); 1981 (anceps); 2712 (anceps); 3602 (uleanum); 3813 (anceps); 3898 (uleanum); 5431 (uleanum); 5837 (mite); 5864 (anceps); 6169 (mite); 6507 (mite); 6612 (anceps); 7143 (anceps); 7394 (anceps); 7595 (mite);7745 (anceps); 9144 (ternatum); 9165 (anceps); 9440 (ternatum); 9454 (anceps); 9765 (anceps); 9914 (anceps); 10020 (uleanum); 9241 (ternatum); 10108 (anceps); 10139 (anceps); 10185 (conicum). Shemluck, M. 303 (conicum). Shemluck, M. & Ness, F: 174 (uleanum). Silva, M.N. 159 (mite). Silverstone-Sopkin, FA. 1487 (anceps). Silverstone-Sopkin, FA. & Rodriguez, A. 2095 (anceps). Skog, L. et al. 7380 (anceps). Skutch, A.F. 2789 (trizygum); 3166 (trizygum); 3614 (trizygum); 4147 (trizygum); 4466 (anceps). Smith, A. A456 (trizygum); 1900 (trizygum); 2750 (trizygum). Smith, D.N. 2905 (conicum); 4035 (anceps); 4104 (ternatum); 4170 (conicum); 5346 (anceps); 7756 (incurvum). Smith, D.N. & Pretel, A. 7968 (ternatum); 8069 (incurvum). Smith, D.N. & Vasquez, R. 4899 (anceps). Smith, D.N. et al. 1184 (anceps); 1569 (ternatum); 7933 (ternatum). Smith, H.H. 1162 (ternatum). Smith, S.F. & Shuhler, A.M. 177 (mite). Smith, S.F. et al. 1355 (mite); 6713 (mite). Sodiro, A. 114/61 (anceps). Solomon, J.C. 8791 (ternatum); 8821 (anceps); 9584 (ternatum); 12674 (anceps); 14806 (anceps); 17159 (mite); 17675 (anceps). Solomon, J.C. & Nee, M. 12674 (anceps); 12704 (ternatum). Solomon, J.C. & Stein, B.A. 11681 (ternatum). Soukup, J. 2210 (mite). Spichiger, R. & Encarnacion, F: 8440 (mite). Spruce, R. 3882 (mite); 4377 (anceps); 4385 (ternatum); 4432 (chamaepolybotryon); 4462 (uleanum); 4849 (angustialatum). Standley, P.C. 68905 (trizygum); 86711 (trizygum); 90563 (trizygum). Standley, P.C. & Valerio, J. 44055 (trizygum); 51987 (trizygum). Stein, B.A. & Cogollo, A. 3394 (anceps). Steinbach, J. 6080 (mite); 9020 (mite). Steinbach, R.F. 424 (mite). Steward, W.C. et al. P12903 (anceps). Steyermark, J.A. 33815 (trizygum); 35135 (trizygum); 37209 (trizygum); 37732 (trizygum); 48735 (trizygum); 51729 (trizygum); 62034 (trizygum); 90105 (trizygum); 91851 (trizygum); 98915 (trizygum). Steyermark, J.A. & Liesner, R. 120698 (trizygum). Steyermark, J.A. & Rabe, M. 96140 (trizygum). Steyermark, J.A. & Steyermark, C. 95161 (trizygum). Stork, H.E. & Horton, O.B. 9536 (mite). Sullivan, G. & Young, K. 1154 (ternatum). REVISION OF SOLANUM SECTION PTEROIDEA Tessmann, G. 3890 (uleanum). Tillett, S.S. 673-226 (conicum). Tillett, S.S. et al. 44971 (anceps). Timand, M. & Astete, H. 692 (ternatum). Tirado, G. et al. 189 (anceps). Tyson, E.L. 7144 (trizygum). Ule, E. s.n. (anceps); 2608 (anceps); 5201 (mite); 5490 (anceps); 6276 (anceps); 6922 (mite); 9731 (mite). Utley, J. & Utley, K. 701 (trizygum); 2902 (trizygum). Vasquez, R. 2243 (anceps); 3876 (anceps). Vasquez, R. & Jaramillo, N. 2584 (anceps); 3499 (anceps); 4475 (anceps); 5097 (anceps); 5471 (anceps); 6370 (mite); 8287 (mite); 8362 (mite); 8680 (anceps); 10533 (anceps); 11699 (mite). Vasquez, R. et al. 2151 (anceps); 6559 (anceps); 11923 (anceps). Vickers, W. 143 (uleanum); 273 (uleanum). Wachter, T.S. 81 (anceps). Walter, H. & Walter, E. 472 (trizygum). Wasshausen, D.C. & Encarnacion, F. 998 (ternatum). Weberbauer, A. 6783 (ternatum); 7570 (incurvum). Webster, G.L. 28483 (mite). Werff, H. van der & Gudino, E. 11400 (anceps). re Werff, H. van der et al. 8281 (anceps); 8357 (conicum); 10219 (anceps); 13122 (conicum). Whalen, M.D. & Salick, J. 862 (anceps); 864 (mite). White, G.E. 7033 (mite). Williams, L.O. 10743 (trizygum); 13624 (trizygum). Williams, L.O. & Alston, H.G. 139 (trizygum). Williams, L.O. et al. 25580 (trizygum). Williams, Ll. 3137 (anceps); 2829 (mite); 2923 (mite); 4264 (mite); 4905 (mite); 5351 (mite); 6045 (mite); 6929 (anceps); 6956 (mite); 7035 (mite); 7322 (anceps); 7689 (anceps); Wingfield, R. & Werff, H. van der 6574 (trizygum). Woytkowski, F. 5543 (ternatum); 7000 (anceps); 8265 (ternatum); 34512 (ternatum); 34560 (anceps); 35416 (ternatum). Woytkowski, F. et al. 560 (anceps). Wurdack, J.J. 940 (ternatum). Young, H.J. & Stratton, D.A. 24 (mite). Young, K. 126 (mite); 134 (mite); 967 (anceps). Young, K. & Eisenberg, M. 375 (anceps). Young, K. & Sullivan, G. 570 (ternatum); 715 (mite). Zak, V. & Espinosa, R. 4358 (anceps); 4629 (anceps). Zaruma, J. et al. 21A (ternatum). INDEX Principal references are in bold, whilst synonyms are in italics. An asterisk denotes a figure. Bassovia sylvatica Aubl. 44, 51 Cyphomandra Sendtn. 36, 37, 42 Lycianthes Bitter 32, 36 Solanum alatibaccatum Bitter 55 Solanum aligerum Schldl. 37 Solanum anceps Ruiz & Pav. 50*, 51, 52* Solanum angustialatum Bitter 50*, 52*, 54 Solanum apiculatibaccatum Bitter 58 Solanum aubletii Pulle 51 Solanum bassovia Dunal 51 Solanum capsicforme (Domin) G.T.S. Baylis 37 Solanum chamaepolybotryon Bitter 55, 56* Solanum conjungens Bitter 51 Solanum conicum Ruiz & Pav. 55, 56*, 57* Solanum cormanthum Vell. 69 Solanum dendrophilum Bitter 46 Solanum diffusum Ruiz & Pav. 45 Solanum diffusum subsp. miozygum Bitter 46 Solanum diffusum vat. miozygum (Bitter) J.F. Macbr. 46 Solanum diploconos (Mart.) Bohs 43 Solanum feddei Bitter 45 Solanum fraxinellum Bitter 64 Solanum hederiradiculum Bitter 51 Solanum huallagense Bitter 58 Solanum incurvum Ruiz & Pav. 45, 46*, 47* Solanum lacteum Vell. 69 Solanum laurinum Dunal 69 Solanum loxophyllum Bitter 69 Solanum marantifolium Bitter 69 Solanum marginatum L.f. 37 Solanum mite Ruiz & Pav. 58, 59*, 61* Solanum mite subsp. hexazygum Bitter 58 Solanum moritzianum Bitter 45 Solanum nigricans M. Martens & Galeotti 37 Solanum nudum Dunal 42 Solanum pentaphyllum Bitter 69 Solanum pittieri Bitter 64 Solanum pteleifolium Sendtn. 58 Solanum quinquefoliolatum Bitter 58 Solanum quinquejugum Bitter 64 Solanum robustifrons Bitter 69 Solanum savanillense Bitter 62*, 63* Solanum seaforthianum Andrews 32 Solanum semievectum Bitter 45 Solanum semiscandens Bitter 46 Solanum subquinatum Bitter 46 Solanum sylvaticum (Aubl.) Bitter 44, 51 Solanum ternatum Ruiz & Pav. 45, 48*, 49* Solanum theobromophyllum Bitter 51 Solanum theobromophyllum vat. procerius Bitter 51 Solanum trizygum Bitter 63*, 64, 65* Solanum trizygum var. tetrazygum Bitter 64 Solanum uleanum Bitter 67*, 68, 69* Solanum uleanum var. gracilescens Bitter 68 Solanum uleanum var. unipedunculatum Bitter 68 Volume 2 No. | No. 10 Volume 4 No. | No. 2 Bulletin of The Natural History Museum Botany Series Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. New Himalayan species of Pedicularis with special reference to those from the eastern Himalaya. P.C. Tsoong. 1955. Pp. I- 34. Facsimile edition. £4.35 Mosses of Dominica, British West Indies. E.B. Bartram. 1955. Pp. 37-49. Mosses of the Ecuadorian Andes collected by P.R. Bell. E.B. Bartram. 1955. Pp. 51-64. Facsimile edition. £4.00 Novitates Himalaicae—1. F. Ludlow & W.T. Stearn. 1956. Pp. 65-81, 8 plates, 11 figs. Facsimile edition. £5.25 Saxifraga of the Himalaya 1. Section Kabschia. H. Smith. 1958. Pp. 83-129, 14 figs. £9.65 The Polypodiaceae and Grammitidaceae of Ceylon. W.A. Sledge. 1960. Pp. 131-158, 4 figs. Facsimile edition. £3.90 Allium and Milula in the central and eastern Himalaya. W.T. Stearn. 1960. Pp. 159-191, 4 plates, 10 figs. Facsimile edition. £4.35 The identity of Jsopyrum aquilegioides. L.G. de Beer & W.T. Stearn. 1960. Pp. 193-202, 3 figs. Facsimile edition. PAS On the geographical relationships of the angiosperm flora of New Guinea. R. Good. 1960. Pp. 203-226, 1 fig. Facsimile edition. £3.60 Saxifraga of the Himalaya II. Some new species. H. Smith. 1960. Pp. 227-260. 9 plates, 17 figs. Facsimile edition. £5.65 New species of Taraxacum from the Himalayan region. J.L. van Soest. 1961. Pp. 261-273, 8 plates. Facsimile edition. £6.15 The athyriod ferns of Ceylon. W.A. Sledge. 1962. Pp. 275-323. 3 plates. Facsimile edition. £5.25 The genus Epilobium in the Himalayan region. P.H. Raven. 1962. Pp. 325-382, 7 plates, 13 figs. Facsimile edition. £5.90 A revision of the genera Buchenavia and Ramatuella. A.W. Exell & C.A. Stace. 1963. Pp. 1-46, 5 figs. Facsimile edition. £5.15 The diatom genus Capartogramma and the identity of Schizostauron. R. Ross. 1963. Pp. 47-92, 2 plates, 38 figs. Facsimile edition. £5.15 Angiosperms of the Cambridge Annobon Island Expedition. A.W. Exel. 1963. Pp. 93-118, 10 plates. Facsimile edition. £3375 A revision of the genus Petrorhagia. P.W. Ball & V.H. Heywood. 1964. Pp. 119-172, 3 plates, 22 figs. Facsimile edition. £4.90 Marine algae of Gough Island. Y.M. Chamberlain. 1965. Pp. 173-232, 4 plates, 80 figs. £12.50 The Ceylon species of Asplenium. W.A. Sledge. 1965. Pp. 233— 277, 1 plate, 3 figs. Facsimile edition. £5.15 Cuticular studies as an aid to plant taxonomy. C.A. Stace. 1965. Pp. 1-78, 5 plates, 10 figs. Facsimile edition. £7.20 The genus Elaphoglossum in the Indian peninsula and Ceylon. W.A. Sledge. 1967. Pp. 79-96. Facsimile edition. £3.25 No. 8 Volume 5 No. | No. 2 No. 6 No. 7 No. 3 No. 4 Volume 7 No. 1 Fungi of recent Nepal expeditions. F.L. Balfour-Browne. 1968. Pp. 97-141, 4 figs. Facsimile edition. £3.75 A synopsis of Jamaican Myrsinaceae. W.T. Stearn. 1969. Pp. 143-178, 8 plates, 25 figs. £4.55 The Jamaican species of Columnea and Alloplectus (Gesneriaceae). W.T.Stearn. 1969. Pp. 179-236, 8 plates, 29 figs. £6.40 New or little known Himalayan species of Swertia and Veratrilla (Gentianaceae). H. Smith. 1970. Pp. 237-258, 16 plates, 7 figs. £8.25 A survey of the tropical genera Oplonia and Psilanthele (Acanthaceae). W.T. Stearn. 1971. Pp. 259-323, 10 plates, 18 figs. £12.40 Angiosperms of the islands of the Gulf of Guinea (Fernando Po, Principe, S. Tomé, and Annobon). A.W. Exell. 1973. Pp. 325-411. £12.50 The dryopteroid ferns of Ceylon. W.A. Sledge. 1973. Pp. 1-43, 4 figs. £6.90 New Himalayan and Tibetan species of Corydalis (Papaveraceae). F. Ludlow & W.T. Stearn. 1975. Pp. 45-69, 15 plates, 14 figs. £8.40 The marine algae of Trinidad, West Indies. W.D. Richardson. 1975. Pp. 71-143, 12 plates, 2 figs. £13.45 A revision of the Macaronesian genus Argyranthemum Webb ex Schultz Bip. (Compositae-Anthemideae).C.J. Humphries. 1976. Pp. 145-240, 2 plates, 26 figs. £14.20 Frank Ludlow (1885-1972) and the Ludlow-Sherriff expedi- tions to Bhutan and south-eastern Tibet of 1933-1950. W.T. Stearn. 1976. Pp. 243-268, 1 fig. Reliquiae botanicae himalaicae. F. Ludlow. 1976. Pp. 269-289, 7 plates, 8 figs. Facsimile edition. £11.10 Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric classification. N.K.B. Robson. 1977. Pp. 291-355, 9 figs. £14.20 Sphagnales of tropical Asia. A-Eddy. 1977. Pp. 357-445, 4 plates, 17 maps, 25 figs. £17.80 The handwriting of Joseph Banks, his scientific staff and amanuenses. J.B. Marshall. 1978. Pp. 1-85, 62 figs. £18.30 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. II. Phaeophyta. J.H. Price, D.M. John & G.W. Lawson. 1978. Pp. 87-182, | fig. £24.40 The lichenicolous Hyphomycetes. D.L. Hawksworth. 1979. Pp. 183-300, 47 figs. £24.40 The species of Chisocheton (Meliaceae). D.J. Mabberley. 1979. Pp. 301-386, 3 plates, 10 figs. £24.40 The distribution of Padina pavonica (L.) Lamour. (Phaeophyta: Dictyotales) on British and adjacent European shores. J.H. Price, I. Tittley & W.D. Richardson. 1979. Pp. 1-67, 3 plates, 2 figs. £17.40 No. 2 No. 3 Volume 8 No. | No. 2 No. 3 No. 4 Volume 9 No. 1 No. 2 No. 3 No. 4 Volume 10 No. | No. 2 No. 3 No. 4 Volume 11 No. | No. 2 No. 3 No. 4 Volume 12 No. 1 No. 2 No. 3 No. 4 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. 111. Rhodophyta (Bangiophyceae). D.M. John, J.H. Price, C.A. Maggs, G.W. Lawson. 1979. Pp. 69-82, | fig. £5. A revision of the genus Anacyclus L. (Compositae: Anthemideae). C.J. Humphries. 1979. Pp. 83-142, 27 figs. £15.60 The Thelypteridaceae of Ceylon. W.A. Sledge. Pp. 1-54, 5 figs. 1981. £15.15 Studies in the genus Hypericum L. (Guttiferae) 2. Characters of the genus. N.K.B. Robson. 1981. Pp. 55-226, 73 figs. £33.55 A revision of the lichen family Thelotremataceae in Sri Lanka. M.E. Hale, Jr. 1981. Pp. 227-332, 20 figs. £24.80 Vascular plant collections from the Tristan da Cunha group of islands. E.W. Groves. Pp. 333-420, 33 figs. £21.40 The lichenicolous Coelomycetes. D.L. Hawksworth. 1981. Pp. 1-98, 36 figs. £22.70 The genus Callithamnion (Rhodophyta: Ceramiaceae) in the British Isles. P.S. Dixon & J.H. Price. 1981. Pp. 99-141,5 figs. £12.50 Parmelia subgenus Amphigymnia (lichens) in East Africa. H. Krog & T.F.V. Swinscow. 1981. Pp. 143-231, 31 figs. £21.05 The genus Selaginella in tropical South America. A.H.G. Alston, A.C. Jermy & J.M. Rankin. 1981. Pp. 233-330, 18 figs. £23.05 Taxonomic studies in the Labiatae tribe Pogostemoneae. J.R. Press. 1982. Pp. 1-89, 33 figs. £21.85 The typification of Hudson’s algae: a taxonomic and nomenclatural reappraisal. L.M. Irvine & P.S. Dixon. 1982. Pp. 91-105. £5.40 Seaweeds of the Faroes. Various authors. 1982. Pp. 107-225, 13 figs. £27.15 The lichen genus Steinera. A.M. Henssen & P.W. James. 1982. Pp. 227-256, 24 figs. £9.70 The algae of Lightfoot’s Flora scotica. P.S. Dixon. 1983. Pp. 1-15, 2 figs. £5.55 A taxonomic study of the lichen genus Micarea in Europe. B.J. Coppins. 1983. Pp. 17-214, 57 figs, 28 maps. £37.75 The hepatics of Sierra Leone and Ghana. E.W. Jones & A.J. Harrington. 1983. Pp. 215-289, 8 figs. £18.15 Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. Y.M. Chamberlain. 1983. Pp. 291-463, 89 figs. £33.45 A revision of the Morinaceae (Magnoliophyta-Dipsacales). M.J. Cannon & J.E.M. Cannon. 1984. Pp. 1-35, 9 figs. £11.40 An introduction to fern genera of the Indian subcontinent. C.R. Fraser-Jenkins. 1984. Pp. 37-76, | fig. £12.50 A revision of African Sphagnales. A. Eddy. 1985. Pp. 77-162, 47 figs. £23.05 Studies in the genus Hypericum L. (Guttiferae) 3. Sections 1. Campylosporus to 6a. Umbraculoides. N.K.B. Robson. 1985. Pp. 163-325, 24 figs, 34 maps. £39.20 Volume 13 No. | The lichen genus Usnea subgenus Neuropogon. F.J. Walker. 1985. Pp. 1-130, 39 figs. £31.85 No. 2 Cytotaxonomic studies of the ferns of Trinidad. A.C. Jermy & T.G. Walker. 1985. Pp. 131-276, 69 figs. £31.50 No. 3 Some genera of the Biddulphiaceae (diatoms) with interlocking linking spines. R. Ross & P.A. Sims. 1985. Pp. 277-381, 33 plates. £28.75 Volume 14 No. | Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae). M. Gibby. 1985. Pp. 1-42, 78 figs. £12.50 No. 2 A redisposition of the species referred to the ascomycete genus Microthelia. D.L. Hawksworth. 1985. Pp. 43-181, 73 figs. £34.25 No. 3 A classification of the genus Dryopteris (Pteridophyta: Dryopteridaceae). C.R. Fraser-Jenkins. 1986. Pp. 183-218, 4 figs. £11.75 No. 4 Evolutionary cladistics of marattialean ferns. C.R. Hill & J.M. Camus. 1986. Pp. 219-300, 27 figs. £24.65 Volume 15 No. | Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 1. Genera A-F. J.H. Price, D.M. John & G.W. Lawson. 1986. Pp. 1-122, 1 fig. £33.60 No. 2 Cytology of the fern flora of Madeira. I. Manton, J.D. Lovis, G. Vida & M. Gibby. 1986. Pp. 123-161, 12 plates. £14.70 No. 3 A revision of the lichen genus Xanthoparmelia in Australasia. J.A. Elix, J. Johnston & P.M. Armstrong. 1986. Pp. 163-362, 42 figs, 117 maps. £42.90 Volume 16 No. | Studies in the genus Hypericum L. (Guttiferae) 7. Section 29. Brathys (part 1). N.K.B. Robson. 1987. Pp. 1-106, 14 figs, 25 maps. £28.75 No. 2 The lichen genus Ramalina in Australia. G.N. Stevens. 1987. Pp. 107-233, 15 plates, 31 figs. £32.20 No. 3 An annotated list of vascular plants collected in the valleys south of Mt Everest. G. Miehe. 1987. Pp. 225-268, 4 figs. £16.50 No. 4 Further genera of the Biddulphiaceae (diatoms) with interlock- ing linking spines. R. Ross & A.Sims. 1987. Pp. 269-311, 13 plates. £15.60 Volume 17 No. 1 Studies in Pseudocyphellaria (lichens) 1. The New Zealand species. D.J. Galloway. 1988. Pp. 1-267, 124 figs. £55.65 Volume 18 No. 1 An illustrated catalogue of the type specimens in the Greville diatom herbarium. D.M. Williams. 1988. Pp. 1-148, 74 plates. £38.00 No. 2 Erik Acharius and his influence on English lichenology. D.J. Galloway. 1988. Pp. 149-194, 18 figs. £17.05 No. 3 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 2. Genera G. J.H. Price, D.M. John & G.W. Lawson. 1988. Pp. 195-273, | fig. £25.10 No. 4 Some Cretaceous and Palaeogene Trinacria (diatom) species. P.A. Sims & R. Ross. 1988. Pp. 275-322, 13 plates. £17.55 No. 5 A monograph of Dryopteris (Pteridophyta: Dryopteridaceae) in the Indian subcontinent. C.R. Fraser-Jenkins. 1989. Pp. 323— 477, 79 figs. £34.10 No. 6 Volume 19 Volume 20 No. 1 No. 2 Volume 21 No. 1 No. 2 Volume 22 No. 1 No. 2 Corydalis (Papaveraceae: Fumarioideae) in Nepal. M. Lidén. 1989. Pp. 479-538, 26 figs. £19.35 A new species of Maytenus (Celastraceae) in Ethiopia. Sebsebe Demissew. 1989. Pp. 1-3, 1 fig. Central American Araliaceae — a precursory study for the Flora Mesoamericana. M.J. Cannon & J.F.M.Cannon. 1989. Pp. 5— 61, 36 figs. A revision of the Solanum nitidum group (section Holophylla pro parte): Solanaceae. S. Knapp. 1989. Pp. 63-102, 21 figs. Six new species of Solanum sect. Geminata from South America. S. Knapp. 1989. Pp. 103-112, 8 figs. The application of names of some Indian species of Ocimum and Geniosporum (Labiatae). J.R. Press & V.V. Sivarajan. 1989. Pp. 113-116, 4 figs. Revision of Piper (Piperaceae) in the New World 1. Review of characters and taxonomy of Piper section Macrostachys. M.C. Tebbs. 1989. Pp. 117-158, 41 figs. Facsimile edition. £52.45 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29. Brathys (part 2) and 30. Trigynobrathys. N.K.B. Robson. 1990. Pp. 1-151, 22 figs, 46 maps. Facsimile edition. £49.25 The marine algal flora of Namibia: its distributions and affinities. G.W. Lawson, R.H. Simons and W.E. Isaac. 1990. Pp. 153-168, 1 fig, 7 plates. The infrageneric classification of Gentiana (Gentianaceae). T.- N. Ho and S.-W. Liu. 1990. Pp. 169-192, 13 figs. Revision of Piper (Piperaceae) in the New World. 2. The taxonomy of Piper section Churumayu. M.C. Tebbs. 1990. Pp. 193-236, 49 figs. £34.10 Historical and taxonomic studies in the genus Titanoderma (Rhodophyta, Corallinales) in the British Isles. Y.M. Chamber- lain. 1991. Pp. 1-80, 247 figs. £42.35 Early collections of the Holy Thorn (Crataegus monogyna cv. Biflora). A.R. Vickery. 1991. Pp. 81-83, 1 fig. A taxonomic study of the species referred to the ascomycete genus Leptorhaphis. B. Aguirre-Hudson. 1991. Pp. 85-192, 76 figs. The typification and identification of Calymperes crassilimbatum Renauld & Cardot (Musci: Calymperaceae). L.T. Ellis. 1991. Pp. 193-194, 1 fig. £42.35 An account of southern Australian species of Lithophyllum (Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J. Campbell. 1992. Pp. 1-107, 63 figs. £41.25 Palynological evidence for the generic delimitation of Sechium (Cucurbitaceae) and its allies. J.L. Alvarado, R. Lira-Saade & J. Caballero. 1992. Pp. 109-121. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 3. Genera H-K. J.H. Price, D.M. John & G.W. Lawson. 1992. pp. 123-146. Two new species of Solanum section Geminata (Solanaceae) from Cerro del Torr in western Colombia. S. Knapp. 1992. Pp. 147-152. Fissidens ceylonensis Dozy & Molkenb. (Musci: Fissidentaceae) and some allied taxa from southern India. L.T. Ellis. 1992. Pp. 153-156, 2 figs. New species of Piper (Piperaceae) from Central America. M. Tebbs. 1992. Pp. 157-158. Studies on the Cretan flora 1. Floristic notes. N.J. Turland. 1992. Pp. 159-164. Volume 23 No. 1 No. 2 Volume 24 No. 1 No. 2 Volume 25 No. 1 No. 2 Volume 26 No. 1 No. 2 Studies on the Cretan flora 2. The Dianthus juniperinus complex (Caryophyllaceae). N.J. Turland. 1992. Pp. 165-169. £41.25 Revision of Piper (Piperaceae) in the New World 3. The taxonomy of Piper sections Lepianthes and Radula. M.C. Tebbs. 1993. Pp. 1-50, 18 figs. Mounting techniques for the preservation and analysis of diatoms. S.J. Russell. 1993. Pp. 51-54. | fig. £43.25 New taxa of Gentiana (Gentianaceae) from Western China and the Himalayan region. T.-N. Ho and S.-W. Liu. 1993. Pp. 55— 60. 2 figs. New combinations, names and taxonomic notes on Gentianella (Gentianaceae) from South America and New Zealand. T.-N. Ho and S.-W. Liu. 1993. Pp. 61-66. Studies in Hypericum: validation of new names. N.K.B. Robson. 1993. Pp. 67-70. Generic monograph of the Asteraceae—Anthemideae. K. Bremer and C.J. Humphries. 1993. Pp. 71-177. 12 figs. £43.25 Pre-Linnaean references for the Macaronesian flora found in Leonard Plukenet’s works and collections. J. Francisco-Ortega, A. Santos-Guerra and C.E. Jarvis. Pp. 1-34. Studies on the lichen genus Sticta (Schreber) Ach.: II. Typification of taxa from Swartz’s Prodromus of 1788. D.J. Galloway. Pp. 35-48. Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 4. Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F. Prud’homme van Reine and W.J. Woelkerling. Pp. 49-90. Studies on the Cretan flora 3. Additions to the flora of Karpathos. N.J. Turland and L. Chilton. Pp.91-100. £43.25 Observations on the benthic marine algal flora of South Georgia: a floristic and ecological analysis. D.M. John, P.J.A. Pugh and I. Tittley. Pp. 101-114. Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical species (excluding Australia). D.J. Galloway. Pp. 115-160. Morphology and ecology of seedlings, fruits and seeds of Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. Pp. 161-172. A study of Bixa (Bixaceae), with particular reference to the leaf undersurface indumentum as a diagnostic character. R.E. Dempsey and N.C. Garwood. Pp. 173-180. £43.40 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. Pp. 1-94. William Roxburgh’s St Helena plants. Q.C.B. Cronk. Pp. 95-98. £43.40 Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae) 5. Genera P. G.W. Lawson, W.J. Woelkerling, J.H. Price, W.F. Prud’ homme Van Reine and D.M. John. Pp. 99-122. A new species of Odontorrhynchos (Orchidaceae, Spiranthinae) from Boliva. D.L. Szlachetko. Pp. 123-125. Linnaeus’s interpretation of Prospero Alpino’s De plantis exoticis, with special emphasis on the flora of Crete. N.J. Turland. Pp. 127-159. Book review. M.G. Gilbert. P. 161. £43.40 A morphological study of Chaetoceros species (Bacillariophyta) from the plankton of the Pacific ocean of Mexico. D.U. Hernandez-Becerril. 1996. Pp. 1-73. £43.40 Studies in the genus Hypericum L. (Guttiferae) 6. Sections 20. Myriandra to 28. Elodes. N.K.B. Robson. 1996. Pp. 75-217. £43.40 Notes on the diatom species Tetracyclus castellum (Ehrenb.) Grunow with a description of Tetracyclus pseudocastellum nov. sp. D.M. Williams A new species of Calymperes (Musci: Calymperaceae) from Peninsular Malaysia L.T. Ellis A phylogenetic conspectus of the tribe Hyoscyameae (Solanaceae) A.L. Hoare and S. Knapp A revision of Solanum section Pteroidea: Solanaceae S. Knapp and T. Helgason BOTANY SERIES Vol. 27, No. 1, June 1997