ISSN 0968-0446 Bulletin of The Natural History Museum PRESENTED GENERAL LIBRARY 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. 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(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. TUP MATI IRAI DAVID M. WILLIAMS ^ Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD HISTORY MUSEUM 31 JUL1997 CONTENTS PRESENTED GENERAL LIBRARY Introduction Terminology Systematic descriptions Tetracyclus castellum (Ehrenb.) Grunow Tetracyclus pseudocastellum D.M. Williams . References ... 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 which 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 T. castellum has revealed a new fossil species from China, T. 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, T. emarginatus (Ehrenb.) W. Sm. andT.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 a taxon Ehrenberg called Biblarium castellum Ehrenb. (= T. castellum (Ehrenb.) Grunow) and a new fossil species from Inner Mongolia, T. pseudocastellum. I also offer some notes on other taxa possibly confused with T. 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), 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: 41 1 (1862). - Tetracyclus japonicus sensu Lupikina in Nov. Sist. Nizsh. Rast. [1965]: pi. 3, figs 1-3 (1965); Khursevich & Loginova, Iskopaemaya Diatomovaya Flora Belorussii (Sistematicheskii Obzor): pi. 17, fig. 13 (1980); Khursevich in Acta geol. hung. 28: pi. II, fig. 7 (1982). - Tetracyclus stellare sensu J. Y. Li in Bull Inst. Geol. chin. Acad. geol. sci. 5: pi. 1 , fig. 18 (1982); J.Y. Li &Y.Z. Qi inProc. 8th Internal. Dial. Symp.: pi. 2, figs 4, 5 (1986); Valeva & Temniskova-Topalova in Fitologiya 46: pi. Ill, figs 11,12 (1993). - Tetracyclus stellare var. eximia sensu VanLand. in Micropaleontology 31: pi. 1, fig. 9 (1985). - Tetracyclus sp. Tscheremisinova, Diatomovaya Flora Neogenovykh Otlozhenii Pribaikal 'ya (Tunkinskaya Kotolovina): pi. 6, fig. 2 (1973). Figs 1,3,4,7. The Natural History Museum, 1997 D.M. WILLIAMS Figs 1, 3, 4, 7 T. castellum. Fig. 1 : Reproduction of Ehrenberg ( 1 854): pi. 33/2, fig. 1 . Fig. 3: Tetracyclus 'costellatus ' from Temp. & Perag., Dial, monde entier, 2nd ed., slide no. 122, BM 68468, specimen 30 \i long. Fig. 4: Tetracyclus elegans van eximia from Temp. & Perag., Diat. monde entier, 2nd ed., slide no. 122, BM 68468, specimen 25 u long. Fig. 7: T. 'costellatus' var. turris from Temp. & Perag., Diat. monde entier, 2nd ed., slide no. 134, BM 68479, specimen 25 u long. Fig. 2 Biblarium elegans reproduced from Ehrenberg (1854): pi. 33/2, fig. 4a, b. Fig. 5 T. emarginatus var. crassa from Temp. & Perag., Diat. monde entier, 2nd ed., slide no. 122, BM 68468, specimen 40 n long. Fig. 6 T.japonicus from Temp. & Perag., Diat. monde entier, 1st ed., slide no. 79, BM 14331, specimen 55 u long. Fig. 8 T. pseudocastellum. Inner Mongolia, China, BM 81618, specimen 40 [i long. DIATOM SPECIES TETRACYCLUS CASTELLUM Biblarium castellum Ehrenb. in Ber. Akad. Wiss. Berlin [1843]: 47 (1843), nom. nud. -Ehrenb. mBer.Akad. Wiss. Berlin [1845]: 73 (1 845). - Ehrenb. inMikrogeol: pi. 33/2, fig. 1 ( 1 854).Type: 'Ad Bargusinam Sibiriae fossile', Ehrenberg (1843: 47), specimens not located. Iconotype = pi. 33/2, fig. 1 in Ehrenberg (1854). ITetracyclus islandica 0strup in Meddr dansk geol. Foren. 6: 28, pi. 1, fig. 1 (1900). Type: Illagil, Iceland (K 384-holotype?). Tetracyclus costellatus Herib., Diat.foss.Auvergne: 16, pi. 8, fig. 12 (1902), orth. var. , corrected to castellum in Perag. Cat. Dial.: 920 (1903). Type: 'Depot de Celles, Cantal' (BM 68468-isotype). Tetracyclus costellatus var. turris Perag. & Herib. in Herib. Dial. foss. Auvergne: 39, pi. 8, fig. 13 (1902), orth. var., corrected to castellum in Perag., Cat. Dial.: 920 (1903). Type: 'Depot d'Auxillac, Cantal' (BM 68479-3-isotypes). Tetracyclus elegans var. eximia Herib. & Perag. in Herib., Diat.foss. Auvergne: 16, pi. 8, fig. 15 (1902).Type: 'Depot de Celles, Cantal' (BM 68468-isotype). Tetracyclus stellare Herib., Diat.foss. Auvergne: 31, pi. 11, fig. 23 ( 1 903).Type: 'Depot de Joursac, Cantal' (BM 68397-99-isotypes). Tetracyclus stellare var. eximia (Herib.) Hust. in Abh. naturw. Ver. Bremen 23: 98(1914). Tetracyclus lapponicus Tynni in Bull. geol. Surv. Finl. 320: 35, pi. 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, pi. 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 n (n=10) in diameter, each tip curving at its margin (Li & Qi, 1986: pi. 2, fig. 4).Transapical ribs primary (25^5 in 10 |a), 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.l, septum visible in Ehrenberg's illustration; Li & Qi, 1986: pi. 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., Dial, 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- Lager von Bargusina im Gouvernement Irkutzk in Sibirien', Ehrenberg, 1843: 46; 'Ad Bargusinam Sibiriae fossile', Ehrenberg, 1 845: 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, 1 845: 73), he offered only one illustration in the Mikrogeologie (Ehrenberg, 1854: pi. 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: pi. 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 Heribaud 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; Heribaud, 1902: 16, pi. 8, fig. 12). In the same volume, Tempere & Heribaud described the new variety T. costellatus var. turris (Fig. 7; Heribaud, 1902: 39, pi. 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 Heribaud 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 T. costellatus as T. constellatus). VanLandingham acknowledged that Heribaud'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 T. castellum. Isotype material for T. 'costellatus ' and T. 'costellatus ' var. turris are available as part of Tempere & Peragallo's Diatomees 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 van eximina, T. emarginatus van crassa, and T. stellare - Tetracyclus elegans (Ehrenb.) Herib. was based upon another Ehrenberg species, Biblarium elegans, de- scribed from the same Siberian fossil deposit as B. castellum (Ehrenberg, 1854: 90, pi. 33/2, fig. 4a, b). Ehrenberg provided no description and only published illustrations of two specimens, one D.M. WILLIAMS valve and one girdle band (Ehrenberg, 1854, pi. 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). T. rhombus (Ehrenb.) Ralfs in Pritchard has been discussed in more detail in Williams (1996) and T. glans in Williams (1987). Briefly, Ehrenberg's original illustrations of T. rhombus included drawings of specimens from Siberia and a U.S.A. fossil deposit from Columbia River (Ehrenberg, 1854: pi. 33/12, figs 7, 8, pi. 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: pi. 33/2, figs 9, 9*, 10) may indeed belong to T. glans (or some closely related species, e.g. T. pagesi Herib. or T. stella (Ehrenb.) Herib.; cf. Hustedt, 1914: 101, 105; Williams, in prep.) while the Columbia River specimens (Ehrenberg, 1854: pi. 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. Heribaud & Peragallo unnecessarily transf erred Biblarium elegans to Tetracyclus (Heribaud, 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 (Heribaud, 1902: 16, pi. 8, fig. 15). Isotype material is available (Temp. & Perag., Dial, monde entier 2nd ed., slide no. 1 22, BM 68468) and, although only a few specimens were encountered, they are 8-pointed star-shaped valves like T. castellum (Fig. 4). In a later volume of the same study, Heribaud published another new species under the name of Tetracyclus stellare Herib. from the Joursac deposit of Cantal (Fig. 7; Heribaud, 1903: 31, pi. 11, fig. 23). According to Heribaud T. stellare is '. . . intermediate entre le Tetracyclus castellum et le Tetracyclus elegans, dont il nous parait une forme derivee' (Heribaud 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, Heribaud & Peragallo described a new variety of Tetracyclus emarginatus, T. emarginatus var. crassa Herib. & Perag. (Heribaud, 1902: 16, pi. 8, fig. 16; specimens from Temp. & Perag., Dial, monde entier 2nd ed., no. 122, BM 68468; Fig. 5). This taxon only superficially resembles T. castellum and should perhaps be considered in the context of T. emarginatus to which it appears more similar. Tetracyclus islandica, T. lapponicus, and T. chudjakovii - 0strup described the new species Tetracyclus islandica from Illagil in Iceland (0strup, 1900: 28, pi. 1, fig. 1). He made no attempt at a description but drew attention to the unusual shape which he felt made its unique status obvious ('. . . som uden at kraeve naermere Beskrivelse, tydeligt fremgaar af Tab. nost. Fig. 1', 0strup, 1900: 28). There is only one relevant slide of type material present in C of which J.B. Hansen wrote: '0strup 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 0strup'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 T. 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, pi. 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 'T. 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: pi. 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: 1 14, pi. 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 T. 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, T. japonicus sensu stricto; see alsoWilliams, 1989). Other illustrations with different names include Lupikina ( 1 965 : pi . 3, figs 1-3) and Khursevich & Loginova (1980: pi. 17, fig. 13;seealso Khursevich, 1982: pi. 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 (1982&: pi. 1, fig. 18) and Valeva & Temniskova-Topalova (1993: pi. Ill: figs 11, 12) who named speci- mens T. stellare; VanLandingham (1985: pi. 1, fig. 9) who named specimens T. stellare var. eximia; and Tscheremisinova (1973: pi. 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 ( 1 986) which they erroneously called T. peragalli Herib. (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 Internal. Dial. Symp.: pi. 2, figs 3, 6 (1986). Valves with 6 equally spaced points somewhat resembling a 'star', each point rounded at the margin, 20-45 \i (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: pi. 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 n -l (9) IM, China', specimen marked number 5-holotype; IGC-Beijing SZ n -l-01 IMS-isotype. Material examined China. Miocene, Den Hua Jiling Province and Shangdu County of Inner Mongolia, BM 81618, 'No: SZ M -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. I 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. 1 843. Mittheilungen iiber 2 neue asiatische Lager fossiler Infusorien- Erden aus dem russischenTrans-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 osterreichen 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. Heribaud, J. 1902. Les Diatomees fossiles d'Auvergne. 1st Memorie. Paris. 1903. Les Diatomees fossiles d'Auvergne. 2nd Memorie. Paris. H listed t, 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. Iskopaemaya Diatomovaya Flora Belorussii (Sistematicheskii Obzor). Minsk. I -an by, A. 1910. Recherches pale'ophytologiques dans le Massif central. Bull. Serv. Carte geol. detaill. France 20: 1-398. Li, Jia Ying 1982a. Miocene diatom assemblages of Shanwang, Shandong Sheng. Acta hot. sin. 24: 456-467. 1982/J. 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. In M. Ricard (Ed.), Proc. 8th Internal. Dial. Symp.: 699-71 1. Lupikina, E.G. 1 965. 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. 0strup, E. 1900. Diatomeerne i nogle islandiske. Surtarbrand-Lag. II. Meddr dansk geol. Foren. 6: 23-30. Ozornina, S. 1 993. Diatoms and problems of stratigraphy of the Eopleistocene and the late Pleistocene of Central Kamchatka. Doctoral Thesis, Vladivostock. Peragallo, M. 1903. Le catalogue general des Diatomees. 2: 472-973. Clermont- Ferrand. Pritchard, A. 1 86 1 .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 In V.A. Krasilov (Ed.), Paleobotanilca 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. Tempere, J. & Peragallo, H. 1907-1915. Les Diatomees du monde entier. Collection Tempere et Peragallo (2e edition). Arcachon. [See note in R. Ross 1 995 Bull. Br. Mus. not. Hist. (Bot.) 25: 93 concerning the citation of this work.] Tscheremisinova, E.A. 1 97 '3 . Diatomovaya Flora Neogenovykh Otlozhenii Pribaikal 'ya (Tunkinskava 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: 1-40. 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 Meridian (Diatomaceae: Bacillariophyta). Bibl. Dial. 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. Bull. not. Hist. Mm. Land. (Bot.) 27(1): 7-9 Issued 27 June 1997 A new species of Calymperes (Musci: Calymperaceae) from Peninsular Malaysia LEN T. t Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD SYNOPSIS. Calymperes woodii L.T. Ellis, apparently endemic to areas of lowland rainforest in Negeri Sembilan, Peninsular 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, Ungulate 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. Iq, r), superficial cells above hyaline leaf base subquadrate to shortly subrectangular in surface view, mostly 5-15(-22.5) x 7.5-12.5 |im (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 |im, isodiametric to slightly longer than broad, with 4-6 sides or rounded, thick-walled (Fig. li, j), each ventrally drawn out as a subacute to acute protru- sion, dorsally pluripapillose (Fig. Ik, 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. (l-)3-6 cells wide, extending from the leaf base toward the distal end of the hyaline lamina sometimes apparent (Fig.lp), 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. Im-o); in hyaline base unistratose, subentire to irregu- 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. Ip). Gemmiferous leaves often erect and slightly exserted above nongemmiferous leaves, similar to nongemmiferous leaves but up to 5 mm long and sometimes more narrowly Ungulate, 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. Ig). Axillary paraphyses produced in brush-like bunches, filamentous, usually exceeding 0.5 mm long, hyaline, multicellular, normally uniseriate (Fig. Ih). 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. 1 8 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 1 5 1 60 1 , c. 1 00 m, March 1 996, Ellis 960 1 (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 bothCalymperes and Syrrhopodon (the two largest genera in the Calymperaceae). However, more features of C. woodii are Calymperes-\\ke than Syrrhopo don-like. For example, the structure of the proboscis in the gemmiferous leaves is virtually identical to that of several species of Calymperes (Fig. If, g), particularly C. graeffeanum Mull. 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. Ip). The presence of axillary paraphyses (lacking in C. graeffeanum and C. hispidum) is a feature of some closely interre- The Natural History Museum, 1997 L.T. ELLIS n 1mm 200[jm h-r. 50pm 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; i-1: chlorophyllose lamina (i, j: ventral surface, k, 1: in cross-section); m-o: margin above hyaline leaf base (m: ventral surface, n, o: in cross-section); p: margin in hyaline leaf base; q, r: costa at mid-leaf in cross-section, a, b, d, e, g-i, 1-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 MUll. 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. Ib, 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 1 372) 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 |im in surface view. The margins of the leaves in C. woodii are polystratose (Fig. 1 n, o), plane to inflexed, and minutely denticu- late with single-celled teeth; the cells of the chlorophyllose lamina are 6-15 x 6-12.5 \im 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 ofMalesian mosses. 2. London. Ellis, L.T. 1988. Taxonomic notes on Calymperes II. 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. Bull. nat. Hist. Mus. Land. (Bot.) 27(1): 1 1-29 Issued 27 June 1997 A phylogenetic conspectus of the tribe Hyoscyameae (Solanaceae) 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 Introduction 11 History of classification 13 Chemistry and economic botany 14 Chemistry 14 Economic botany 14 Methods 15 Choice of taxa '. 15 Characters 15 Tree construction 16 Results and discussion 16 Tree topology 16 Character analysis 20 Biogeography 24 Generic conspectus 25 Artificial key to the genera of the Hyoscyameae 25 Anisodus Link 25 Atropa L 26 Atropanthe Pascher 26 Hyoscyamus L 26 Mandragora L 26 Physochlaina G. Don 27 Przewalskia Maxim 27 Scopolia Jacq 27 References 28 Appendix I 29 SYNOPSIS. A cladistic analysis of the tribe Hyoscyameae (including Atropa and Mandragora) shows that Atropa and Mandragora are clearly part of the same monophyletic group as the traditional members of the tribe: Hyoscyamus, Physochlaina, Anisodus, Atropanthe, Scopolia, and Przewalskia. The group can further be divided into two main clades, one containing Hyoscyamus and Physochlaina and the other containing the rest of the genera. Characters used in the analysis are discussed and illustrated, and a conspectus of the genera with descriptions and lists of component species is provided. Introductory material includes a review of the nomenclatural history and the economic botany of the Hyoscyameae. INTRODUCTION The Solanaceae are an economically important, cosmopolitan fam- ily with over 2500 species. Members of the family are important to agriculture, with potatoes, tomatoes, peppers, and a host of minor fruit crops cultivated worldwide. Many species are also valuable to medicine, being used in both traditional and pharmaceutical treat- ments. The family is traditionally divided into two subfamilies. The Cestroideae, including petunias, the cestrums, and their relatives, usually have non-compressed, often prismatic seeds and tropane alkaloids. The Solanoideae, which contains the majority of the species in the family, including Solanum and its relatives, have compressed seeds and steroidal alkaloids. This traditional classifica- tion has recently been challenged by cladistic analyses using chloroplast and nuclear DNA data sets, and the family can now be divided into approximately seven monophyletic groups (see Olmstead et al., in press). The family is predominantly tropical in distribution, but the group variously defined as the tribe Hyoscyameae or subtribe Hyo- scyaminae (see p. 13) is exclusively Eurasian and Middle Eastern, with no members in even the subtropics of the northern hemisphere (see Fig. 1 ).The six genera of the traditionally defined Hyoscyaminae (Anisodus, Atropanthe, Hyoscyamus, Physochlaina, Przewalskia, and Scopolia) and the genera of the Atropinae (Atropa and Man- dragora) have often been considered related in the past (see p. 13) but are usually considered separately. As part of on-going studies into the generic phylogeny of the Solanaceae (Persson et al. 1994; The Natural History Museum, 1997 12 A.L. HOARE AND S. KNAPP PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE 13 Knapp et al., in press) we decided to examine the phylogenetic relationships of these eight genera using primarily morphological characters. We have attempted a preliminary generic delimitation, but several potential problems are highlighted in the analysis. HISTORY OF CLASSIFICATION The concept of the family Solanaceae was first used by A.L. de Jussieu in his Genera plantarum (1789) where he included 15 genera in the Solanaceae, all of which are still in current usage, including Atropa, Hyoscyamus, and Mandragora. He divided the family into three groups, the first with capsular fruits, the second having berries, and the third a group of miscellaneous taxa of less certain affinity with the rest of the family. Following de Jussieu 's work and continuing to the present, much effort has been put into the production of a classification system of the family which best reflects the relationships of the genera. G. Don (1838) was the first to recognize tribes within the family. He proposed seven tribes, largely on the basis of the corolla aestivation, characters of the stamens, fruit type, and embryo curvature. These features have been important in most subsequent classifications. Endlicher (1839) produced an alternative classification which was the first to recognize the tribe Hyoscyameae, composed ofHyoscya- mus, Anisodus, and Scopolia. The characters by which he defined this group were a curved embryo, semi-circular cotyledons, and a bilocular capsule with circumscissile dehiscence. Endlicher placed Atropa and Mandragora in the heterogeneous tribe Solaneae, com- posed of genera with baccate fruits. Miers (1849) proposed profound changes to the Solanaceae, with the erection of a new family Atropaceae. He considered this group to be intermediate between the Solanaceae and the Scrophulariaceae on the basis of corolla aestivation and symmetry of floral parts. Thus, the Atropaceae were characterized by having nearly isomer- ous flowers with imbricate or a peculiar aestivation. Within the Atropaceae Miers described 10 tribes, including Hyoscyameae (in- cluding the genera Hyoscyamus, Scopolia, Physoclaena, Cacabus, and Thinogeton) and Atropeae (including the genera Atropa, Nic- andra, Cliocarpus, Anisodus, Mandragora, and Lyciuni). Miers stated that the Hyoscyameae formed a 'very natural tribe', but then contradicted himself by casting some doubt over the affinity of the last four genera with Hyoscyamus. Miers' hesitation over definition of the Hyoscyameae was clearly due to his uncertainty over the exact nature of corolla aestivation in all but Hyoscyamus. The Atropeae he distinguished by their supposedly baccate fruits, and the absence of an epigynous gland. Later (Miers, 1 850), he reclassified/4m.x/s in the Hyoscyameae after seeing its circumscissile fruit and clearly imbricate corolla aestivation. In 1852 Dunal published the account of Solanaceae for de Candolle's Prodromus. This treatment is the last published revision of all the known genera and species in the Solanaceae. It included 60 genera which were placed into two tribes, the Nolaneae and Solaneae. The latter group was subdivided into nine subtribes, including the Hyoscyaminae, defined using the same characters as Endlicher ( 1 839) had used. Dunal recognized only Hyoscyamus and Scopolia in the Hyoscyaminae. Within Scopolia however, he included species now placed in Physochlaina, Hyoscyamus, Anisodus, and Scopolia s.s. which he defined as sections. His rationale for lumping these taxa under Scopolia was that they all differed from Hyoscyamus in the shape of their corolla and calyx. Atropa and Mandragora were placed in the large and heterogeneous subtribe Solanaeae. Within that subtribe the division Atropineae contained many of the genera of the family, including those as diverse as Jaborosa and Discopodium. This group was diagnosed by its 5-parted calyx, campanulate corolla with valvate-plicate aestivation, and its baccate fruit. In the second half of the nineteenth century two further treatments of Solanaceae appeared, those of Bentham (1876) and Wettstein ( 1 895). These were produced at a time when many new species were being discovered as a result of botanical explorations to new regions of the world. Also at this time, Darwin had published his Origin of species ( 1 859) and the theory of evolution by natural selection was beginning to have an impact on taxonomy. Die natiirlichen Pflanzenfamilien, in which Wettstein's system was published, was the first major work to incorporate these ideas, although this had little direct impact on the classification of the Solanaceae. Both Bentham and Wettstein recognized a group centred on Hyoscyamus. Their classifications are summarized in Table 1 . Table 1 Classification of the Hyoscyameae according to Bentham ( 1 876) and Wettstein (1895). Bentham (1876) Wettstein (1895) Solaneae Atropeae: Including - Lycium L. Atropa L. Mandragora Juss. Hyoscyameae: Datura L. Scopolia Jacq. Physochlaina Don. Hyoscyamus L. Cestrineae Salpiglossidae Nicandreae Solaneae: Lyciinae - incl. Atropa L. Hyoscyaminae - Scopolia Jacq. Physochlaena Don. Przewalskia Maxim. Hyoscyamus L. Solaninae Mandragorinae - incl. Mandragora Juss. Datureae Cestreae Salpiglossideae Working extensively on the aestivation of the corolla and calyx, Baehni (1946) suggested pathways for the evolution of morphology in the family and proposed a new classification of the Solanaceae. He recognized five tribes, of which the Atropeae included the subtribe Hyoscyaminae. The genera which made up this subtribe were: Hyoscyamus, Scopolia, Physochlaina, Tunaria, Vestia, Przewalskia, Petunia, and Nierembergia. Atropa and Mandragora were placed in the subtribe Atropinae. The most recent conspectus of the family (D'Arcy, 1979, revised and slightly modified in D'Arcy, 199 1 ) is based on that of Wettstein with some modifications. In this system, 96 genera are included within the Solanaceae, representing about 2300 species. The hyoscyamoid group is recognized at the rank of tribe, and is composed of six genera: Scopolia, Anisodus, Atropanthe, Przewalskia, Physochlaina, and Hyoscyamus. This classification of the tribe Hyoscyameae fol- lows that of Lu & Zhang (1986) who studied the Chinese members of the tribe. They listed the chief characters of the tribe as follows: embryo curved, ovary two-chambered with numerous ovules, con- nective between two anther cells inconspicuous, filaments inserted at the base or near the base of anthers, corolla lobes often imbricate in bud, calyx greatly enlarged after flowering and including the fruit, capsule with circumscissile dehiscence, and plants containing tro- pane alkaloid compounds. Atropa and Mandragora are both placed in the still large and heterogeneous tribe Solaneae, along with 33 other genera. However, some work has cast doubt over this delimitation of the tribes. Tetenyi (1987) examined the biosynthesis of alkaloids and steroids in the Solanaceae, and concluded that the family should be divided into four groups. He recognized the subfamilies Antho- cercidoideae, Cestroideae, Solanoideae, and Atropoideae, based 14 largely on differences in the complexity of alkaloid biosynthetic pathways. Further evidence was taken from geographical and mor- phological data. On this basis, Tetenyi placed the Hyoscyaminae, as defined by Lu & Zhang (1986), together with Atropa and Man- dragora to form the subfamily Atropoideae. The subfamily was defined by its distinctive alkaloid-tropane ester synthesis relation- ship, where tropane esters are dominant forms (see T6t6nyi, 1987), imbricate corolla lobes, and Eurasian distribution. Only Lu & Zhang (1986) have attempted to produce a phylogeny for the whole of the Hyoscyameae. This treatment has been used as the basis for the recent Flora of China account (Zhang et al., 1994). They did not include Atropa and Mandragora in their concept of the tribe because these genera possess baccate fruits. In their classifica- tion, Scopolia and Anisodus were considered the most primitive members of the group, withAtropanthe and Przewalskla being most closely related to them. These relationships were proposed on the basis of whether the genera showed relatively advanced or primitive characters. The primitive features were considered to be solitary flowers, an actinomorphic corolla, stamens inserted at the base of the corolla tube, and inaperturate pollen grains. However, there are a number of reasons to question their results. The first of these is doubt over the pollen descriptions given (see our results below). Secondly, they give no explanation as to how they reached their decisions regarding the evolution of characters, which could be interpreted differently. Finally, Lu & Zhang state themselves that their phylogenetic hypothesis should only be regarded as a basis for further study of the group. Most recently, the Solanaceae have been the subject of molecular studies (Olmstead & Palmer, 1992; Olmstead & Sweere, 1994; Olmstead et al., in press). These chloroplast DNA based phylogenies do not include all the species or even all the genera of the family, but give ideas as to the potential monophyletic groupings of taxa. In cpDNA phylogenies based on the genes rbcL and ndhF (Olmstead & Sweere, 1994), Atropa is grouped with Nolana and Lycium. With restriction sites on the entire chloroplast genome, Atropa and Hyo- scyamus together are sister to Lycium (Olmstead & Palmer, 1992). The most recent and as yet unpublished results of Olmstead et al. (in press) indicate that Hyoscyamus, Physochlaina, Anisodus, and At- ropa (the only genera of the tribe used in the analysis) form a clear, well-defined monophyletic clade. These analyses do not contain enough taxa to draw clear conclusions as to the relationships of the genera, but they do clearly place the Hyoscyameae as defined here and by Tetenyi ( 1 987) together. However, Mandragora is extremely isolated in the cpDNA analysis and possesses many molecular autapomorphies (Olmstead et al., in press). CHEMISTRY AND ECONOMIC BOTANY Chemistry The presence of abundant tropane alkaloids is characteristic of mem- bers of the Hyoscyameae. Tropane alkaloids are characterized by their five-member, nitrogen-containing rings and are found in several families in addition to the Solanaceae, most notably the Erythro- xylaceae (Hegnauer, 1973;Romeike, 1978; Evans, 1979;Lounasmaa, 1988). The range and variety of tropane alkaloids in the Solanaceae arises from the esterification of various acids, such as acetic, propi- onic, and tiglic (see Evans, 1979) with hydroxytropanes derived from amino acids such as ornithine, tryptophan, and phenylalanine. Hyo- scyamine-type alkaloids, derived from phenylalanine, are also found in the Australian genera related to Anthocercis, the Chilean endemic Latua (Schultes & Hofmann, 1980), Acnistus, andSalpichroa, but are A.L. HOARE AND S. KNAPP in the highest concentrations in members of the Hyoscyameae (Romeike, 1978; Evans, 1979). The biosynthetic pathways leading to the hyoscyamine-type alkaloids are the most complex in the family, and are homologous in all the genera studied (Tetenyi, 1987). The most abundant accumulated end-products in members of the Hyoscyameae are atropine, hyoscyamine, hyoscine (=scopolamine), and tropine, but many other minor tropanes are also found. For complete lists of component alkaloids and a detailed discussion of tropane biosynthesis, see Evans (1979), Romeike (1978), and Tetenyi (1987). Studies into the chemistry of these tropane alkaloids have not only provided data useful for classification, but also have provided insight into the effectiveness of these plants in traditional medicine (Qicheng, 1980; Xiao, 1981; Xiao & He, 1982). Economic botany The use of the Hyoscyameae in medicine has a long history. They have been recorded in the herbals of the ancient Greeks and in the ancient Chinese, Tibetan, and Indian pharmacopoeias (Deb, 1979; Bettolo, 1981, and see references below). Hyoscyamus niger (henbane) has a long history of use over most of its range. The oldest of the Chinese herbals, Shen Nung Pents 'ao Jing (thought to have been written some 2000-3000 years ago) describes the virtues of the seeds of henbane, lang-tang-tze, for curing toothache and for increasing vitality (Xiao & He, 1983). Later Chinese herbals document the use of lang-tang in a variety of illnesses and state that a tonic made from the plant allowed the patient to communicate directly with devils and spirits (Ratsch, 1992). Doctors of the Assyrian empire also used henbane for the cure of toothaches (Press et al., 1989). Dioscorides used henbane to induce sleep and relieve pain, and wrote of its properties in his De materia medica in A.D.77 (Stockwell, 1988). The herbals of the Middle Ages also contain numerous references to narcotic and dangerous properties of henbane (Arber, 1912). One of the best known of the English herbalists, John Gerard (1597), wrote scath- ingly of the fraudulent use of the smoke of henbane seeds to draw worms from the teeth. There is no doubt that the drug eased the pain of toothache, but the worms so miraculously removed by medical practitioners were nothing more than the tiny coiled embryos re- leased from the seeds by the heat (Grieve, 1992). Culpeper, an astrological botanist (see Arber, 1912) writing in the first half of the seventeenth century, recommended an infusion of the leaves for the treatment of gout, swellings, and pains of the joints. He also believed the oil of the seed to be good for deafness and worms in the ears (Culpeper, 1826), thereby perpetuating the worm myth! Several species of Hyoscyamus are used in North Africa for both criminal and medicinal purposes (Boulos, 1983). The Coloured atlas of Tibetan medicine (1704, see Xiao & He, 1983), a commentary on the much earlier complete manual of Tibetan medicine Rygudbzhi of 820, describes the use of the roots of Przewalskia tangutica to relieve pain and reduce swellings. The Atlas included illustrations of the plants used, and that of P. tang- utica is botanically extremely accurate. The roots of Phyoschlaina physaloides are used in Mongolia as a tonic and a cure for asthma, those of P. praealta in Tibet as an analgesic (Xiao & He, 1983) and in India as a drug to dilate the pupils and to cure boils (Sharma & Singh, 1975). Roots of Physochlaina infundibularis were locally regarded as a sort of ginseng in the Chinese provinces of Shanxi and Henan. Several species of Anisodus have a long history of use in Tibetan medicine, as analgesics, anaesthetics, and antispasmodics, but dosages are very small and great care is needed as overdoses are known to cause delirium (Xiao & He, 1983). In addition to their value as herbal remedies, some of these plants PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE 15 gained reputations for their supposed magical properties, and be- came associated with numerous myths. Most notable are those surrounding the mandrake, Mandragora officinarum (Bouquet, 1 936; Moldenke & Moldenke, 1952). The roots of this plant, which sometimes bear a resemblance to the human form, were believed to be the abode of evil spirits. The mandrake was said to scream when pulled from the ground, causing the death of the person uprooting it or anyone who heard the screams. To escape such a fate, people were said to use dogs to pull the plants up, and would drown the cries by blowing loudly on horns. Mandrakes were said to sprout from the sperm of hanged men, and so were to be found growing at the foot of gallows. Gerard (1597) ridiculed these myths as the hoaxes of charlatans, saying '..they are all and everie part of them false and most untrue for I myselfe and my servants also have digged up, planted and replanted very many..'. A mandrake is clearly identifi- able on the thriteenth century Mappa Mundi in Hereford, a map showing the extent of the then-known world and many of its creatures. Numerous images of mandrakes adorn Egyptian tombs and tomb art and it may have been an important drug at that time. In the Old Testament mandrakes are mentioned in two places (in Genesis and in the Song of Solomon) in relation to the procreation of children, and the roots are carried in Eastern Europe as a charm against sterility (Mehra, 1979). Mandragora was also associated with witchcraft, as were hen- bane and deadly nightshade, Atropa belladonna. The magical powers of these plants were believed to enable witches to fly (Stockwell, 1988). The celebrated sixteenth century Spanish phy- sician A.F. de Laguna was probably the first to correlate the use of solanaceous drugs with witchcraft. The ointments used by sup- posed witches were composed of henbane, nightshade, and mandrake and caused hallucinations and delusions. He believed that witches were drug users and that the use of hallucinogenic drugs also increased suggestibility. Thus the confessions wrung from these people represented the delusional speech of deranged minds and were false (Schleiffer, 1979). Accounts of hallucina- tions experienced while using henbane and mandrake almost always involve flying sensations and some have suggested that the urge to move is a hallmark of the intoxication (Schleiffer, 1979). The Hyoscyameae have continued to be employed in medicine right up to the present day. They make an important contribution to both modern and traditional medicine. In traditional medicine the use of these plants is found in Tibet, China, and in the Himalayan countries. They are used to treat swellings, bruises, asthma, to relieve pain, for the treatment of travel sickness, as antidotes to nerve gases, and as anaesthetics and sedatives. Extracts of species of Physochlaina and Mandragora caulescens are also used to make a tonic in Tibet, southern China, and Mongolia, which is administered to replenish weakness and to 'warm the stomach'. Most of the Chinese species in the group are used in medicine in some way (Xiao & He, 1983; Zhang et al., 1994). Hyoscyamus is in the official British Pharmacopoeia (see Grieve, 1992) and has great importance in both theAyurvedic and Unani medicinal systems of India (Dash & Kashyap, 1980;Thakuretal., 1989). The uses of this group of plants reflect the properties of their constituent alkaloids. The tropane alkaloids have been shown to be effective as analgesics, anaesthet- ics, antispasmodics, and to increase the circulation. Today, five species are used commercially as a source of alkaloids in modern Western medicines. Hyoscine, or scopolomine, most important as a sedative, is extracted from Scopolia carniolica. Scopolomine is the infamous 'truth serum' - used unscrupulously to extract information from uncooperative persons. Atropa belladonna is the source of atropine, an antispasmodic used to treat asthma, colic, and eye diseases. Hyoscyamus niger, H. albus, and//, muticus are grown for the extraction of hyoscine, atropine, and hyoscyamine. This brief account highlights the importance of this group of plants for man. However, their beneficial effects are dependent on their wise use. All of these plants are potentially toxic and can cause death. Deadly nightshade, Atropa belladonna, is one of the most poisonous species of the group. This is clearly reflected in both the common and generic names of the plant: in Greek mythology Atropos was the Fate who held the shears to cut the thread of human life. METHODS Choice of taxa Specimens studied were those at The Natural History Museum, London (BM) and the Royal Botanic Gardens, Kew (K). Living plants of some taxa were also examined. Individuals of Anisodus luridus, Mandragora officinarum, Physochlaina orientalis, Atropa belladonna, Hyoscyamus albus, H. niger, and Scopolia carniolica are all grown at Kew, and all but Anisodus and Physochlaina are grown at the Chelsea Physic Garden, London. Scopolia carniolica is also grown in the Harris Garden, Reading. Further details of the specimens studied are given in Appendix I and a complete database of all specimens studied is held at the University of Reading. Genera with more than one species were sampled, with species encompass- ing the range of variation used in the analysis. Taxa used in the analyses were: Lycium chinense Mill., Datura inoxia Mill., Me- an draphysalodes Gaertn., Anisodus luridus Link, Atropa belladonna L., Atropanthe sinensis (Hemsl.) Pascher, Hyoscyamus muticus L., Hyoscyamus niger L., Hyoscyamus senecionis Willd., Mandragora officinarum L., Mandragora caulescens C.B. Clarke, Physochlaina physaloides (L.) G. Don, Physochlaina praealta (Dene.) Miers, Przewalskia tangutica Maxim., and Scopolia carniolica Jacq. Characters The taxa were scored for variation in morphological and palynological characters. The morphological characters used were taken from the flowers, leaves, fruits, and seeds. A list of all the characters is given in Table 2, together with the coding used. Most of the characters are binary and self-explanatory. The data matrix used in the analyses is presented in Table 3. The features of the spermoderm were determined by examining seeds prepared by enzyme etching (Lester & Durrands, 1984). Two seeds of each species were treated with a 1% solution of Driselase for 24 or 48 hours. The longer time was found to be necessary for the seeds of Anisodus luridus, Atropa belladonna, Atropanthe sinensis, Mandragora officinarum, M. caulescens, andPrzewalskia tangutica. The seeds were rinsed with distilled water and allowed to dry before being examined with SEM. Seed measurements were made from 20 seeds when possible, and the average value recorded. Table 2 Character set used in the cladistic analysis. Inflorescence 0. Inflorescence type: raceme 0; solitary flowers 1; cyme 2 1. Flowers: pedicellate 0; sessile 1 Calyx 2. Calyx: tubular 0; campanulate-urceolate 1; cup-shaped 2 3. Calyx: actinomorphic 0; zygomorphic 1 4. Calyx: quite deeply lobed 0; very deeply lobed 1: shallowly lobed 2 5. Apices of calyx lobes: acuminate 0; rounded 1 6. Length of calyx: short (< 1 cm) 0; medium ( 1-3 cm) 1 ; long (> 3 cm) 2 16 A.L. HOARE AND S. KNAPP 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 1 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 1 16. Stamens: regular 0; declinate 1 Stigma 17. Stigma: included in corolla tube 0; exserted 1 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 3 1 . 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 1 4 1 . Tropine: absent 0; present 1 Table 3 Data matrix used in the cladistic analysis. Character Number 0123456789 1111111111 2222222222 3333333333 44 0123456789 0123456789 0123456789 01 Lycium 0000000000 0000000000 0000000000 0000000000 00 Anisodus 1021001 100 001 1 1 1001 1 12100001 10 1000121020 01 Atmpa 002010110000111111000000000121 1101000031 11 Atropanthe 1020001 100 001 1 1 1 1 1 1 1 01 10000100 01 10000030 01 H. muticus 21 1 120101 1 0010000121 021 1010001 0101 10001 1 01 H. niger 21 1 100101 1 0010000121 0210010001 010200001 1 01 H.senecionis 20100010102011000121 0211010001 0102000011 01 M. caulescent 10201 1 1200 201 10100000000201 1 10 11 100171 11 10 M.officinarum 1020101000 201 1 1 10000 0000101 1 12 0000017101 10 P. physaloides 0000000000 00000001 1 1 01 10100001 000202001 1 01 P.praealta 001000000011101001210111100001 100200003101 Przewalskia 100001 1000 00102101 1 1 1210201 100 1001000001 01 Scopolia 1021200000 1 101 1 1001 1 01 100001 10 01 1007101 1 01 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 Tetenyi (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 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 Atmpa 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 andPhysochlaina has been pointed out by previous authors (Lu & Zhang, 1 986). 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 off! 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 1 2, 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 17 17181921223941 HHHHHHHh 0000000 24293338 -H-H 1121 2 121823 i-H-f 1121 1011 143038 I I I 1 I 3 6 821242531 1-H-HHhi 2112011 1013 HHh 43334 1 3 9 i+f I I I 2 1 1 23 5 14202124263033 6 12141527 mm mm 12122111 16323839 2 7133138 MM 2 1 1 I I 1130 172830 t 33536 i+f I I 1 20213134353839 llOlllr 1201220 4 6 7 1011 123032 2001 1001 4 1819212240 -WHHhh 00001 0161728293338 -ilNMI 0112113 -Lycium -Physochlaina physaloides -Physochlaina praealta -Hyoscyamus senedonis -Hyoscyamus muticus -Hyoscyamus niger -Przewalskia -Atropanthe -Anisodus -Scopolia -Atropa 211110 5 7 142432 MOM Mandragora caulescens 12021 729303138 -0-HHH 02000 qffidnarum 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. 18 A.L. HOARE AND S. KNAPP PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE 19 Fig. 4 a) Fruit ofAnisodus tanguticus (photograph courtesy of M. Gilbert), b) Fruit of Atwpa 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. Atwpa and Mandragora cluster together and are nested well within the tribe as a whole. This is in agreement with Tetenyi'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 ( 1 849), Dunal ( 1 852), and Bentham ( 1 876) 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 mSpeciesplantarum ( 1 753), 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, 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. An actinomorphic 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 andScopolia. 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 21 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. 1 4. FILAMENT INSERTION. Lu & Zhang ( 1 986), 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 inPhysochlainapraealta. 1 6. 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 seeds are often associated with capsular fruit in the Solanaceae (Soueges, 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. luridus 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 andPhysochlainaphysaloides, 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. luridus 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 22 A.L. HOARE AND S. KNAPP mmwm?^ '^V/*' ///'/ rA'< f '"'- '***" PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE f- -r. 23 Fig. 7 Pollen morphology of the Hyoscyameae. Scale bars beneath photographs, a) Scopolia carniolica, b) Anisodus luridus, d) Atropa belladonna, d) Mandragora caulescent. 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 A.L. HOARE AND S. KNAPP 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 S-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- 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, 198 1 ; 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, 1 994). 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 theTibetan 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 Tetenyi'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 . Fruit a fleshy or juicy berry, white, green, yellowish orange, purplish or black 2 Fruit a dry capsule, usually circumscissile 3 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, actinomorphic 5. Mandragora Berry black and juicy at maturity; plant an erect perennial to 1 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 with more than 5 flowers 7 4. Acaulescent herbs with long, fleshy taproots; leaves sessile; corolla narrowly tubular 7. Przewalskia Erect, often branched perennials with well-developed above-ground stems; leaves variously petiolate; corolla not narrowly tubular 5 5. Corolla slightly zygomorphic, one petal larger than the rest; calyx lobes equal in size; anthers declinate at anthesis 3. Atropanthe Corolla actinomorphic; calyx lobes usually unequal in size; anthers not declinate at anthesis 6 6. Corolla campanulate-urceolate, as wide as long, greenish; calyx lobes rounded at the tips 1 . Anisodus Corolla flaring, longer than wide, usually purplish without; calyx lobes strongly pointed, especially the elongate one 8. Scopolia 7. Inflorescences markedly one-sided, usually unbranched; corolla zygomorphic; calyx lobes spiny at the tips in fruit .... 4. Hyoscyamus Inflorescences not markedly one-sided, usually branched; corolla actinomorphic; calyx lobes not spiny at the tips in fruit 6. Physochlaina 1. Anisodus Link, in Sprengel, Syst. veg. 1: 699 (1825). Type: Anisodus luridus Link. Figs 3d, 4a. Robust subshrubs or perennial herbs, at least sometimes with a massive softly woody rootstock; glabrous or pubescent. Leaves alternate, petiolate, entire or dentate; sympodia difoliate. Inflores- cences of solitary flowers borne in the leaf axils. Flowers 26 subactinomorphic; calyx campanulate-funnelform to funnelform, the lobes unequal in length and variable in shape, usually rounded at the tips; corolla campanulate, longer than or equal in length to the calyx; stamens inserted near the base of the corolla tube; pollen hexapantoporate with slightly elongated pori with rounded ends and scabrate-gemmate ornamentation (Fig. 7b); ovary bilocular, with a disc-like nectary. Fruit a globose or ovoid capsule, with circumscissile dehiscence; fruiting calyx much enlarged, sometimes enclosing the fruit and prominently ribbed, often laterally compressed. Seeds numerous, not markedly compressed. Grasslands and woodland edges, occasionally ruderal around towns and villages; 2800^500 m. LIST OF SPECIES (Zhang et al. 1994). Anisodus acutangulus C.Y. Wu & C. Chen, China; Anisodus carniolicoides (C.Y. Wu & C. Chen) D'Arcy & Zhang (Scopolia carniolicoides C.Y. Wu & C. Chen), S. China; Anisodus luridus Link (Anisodus fischerianus Pascher, A. luridans Link & Otto, A. mairei (H. Lev.) C.Y. Wu & C. Chen, A. stemonifolius G. Don, A. stramonifolius (Wall.) G. Don, Nicandra anomala Link & Otto, Physalis stramonifolia Wall., P. stramonifera Wall., Scopolia anomala (Link & Otto) Airy Shaw, 5. lurida (Link) Dunal, 5. mairei H. Lev., S. stramonifolia (Wall.) Shrestha, Scopolina stramonifolia (Wall.) Kuntze), Whitleya stramonifolia (Wall.) Sweet), India, Bhutan, Nepal, and China; Anisodus tanguticus (Maxim.) Pascher (Scopolia tangutica Maxim.), Nepal, China. 2. Atropa L., Sp. pi. 1: 181 (1753). Type: Atropa belladonna L. Figs 3f, 4b. Perennial herbs; glabrous or slightly pubescent. Leaves alternate, petiolate, simple and entire. Inflorescences of solitary flowers borne in the leaf axils. Flowers actinomorphic; calyx campanulate; corolla tubular-campanulate, twice as long as the calyx, greenish purple or yellow; stamens inserted near the base of the corolla tube, declinate; pollen trizonocolporate with long, distinct colpi with sunken mar- gins and striate-rugulate ornamentation (Fig. 7c); ovary bilocular, with an annular receptacular disc. Fruit a black, juicy berry; fruiting calyx somewhat enlarged but not enclosing the berry. Seeds numer- ous, sublenticular. Woodland and other shady habitats, rocky screes; 0-1800 m. (Hawkes, 1972; Schonbeck-Temesy, 1972). LISTOFSPECIES (Harborne & Khan, 1993,Pojarkova, 1955;Schon- beck-Temesy, 1972; Baytop, 1979). Atropa acuminata Royle ex Lindl., Asia (India, Pakistan, Afghanistan, Mongolia, Iran); Atropa baetica Willk., Spain and Morocco; Atropa belladonna L. (Atropa caucasica Kreyer, A. komarovii Blin. & Schal., A. lutescens Blin. & Schal., A. pallidiflora Schonb.-Tem., A. paschdewiczi Kreyer), widespread in Central Europe and Asia to Iran. 3. Atropanthe Pascher in Oesterr. Bot. Zeitschr. 59: 329 (1909). Type: Atropanthe sinensis (Hemsl.) Pascher (basionym Scopolia sinensis Hemsl.). Subshrubs or perennial herbs; glabrous. Leaves alternate, petiolate, simple and entire. Inflorescences of solitary flowers borne in the leaf axils. Flowers subactinomorphic; calyx tubular-campanulate; co- rolla slightly zygomorphic, with one petal lobe enlarged, tubular-campanulate, twice as long as the calyx; stamens inserted near the base of the corolla tube, declinate; pollen trizonocolporate with short, distinct colpi and striate-rugulate ornamentation (Fig. 6c); ovary bilocular with an annular disc. Fruit a globose capsule, with circumscissile dehiscence; fruiting calyx inflated, abruptly inserted on the pedicel. Seeds rectangular and somewhat com- pressed. Forest and ditches; 1400-3000 m. (Zhang et al. 1994). A.L. HOARE AND S. KNAPP LIST OF SPECIES (Zhang et al. 1994). Atropanthe sinensis (Hemsl.) Pascher (Anisodus sinensis Hemsl.), China. 4. Hyoscyamus L., Sp. pi. 1: 179 (1753). Type: Hyoscyamus niger L. Figs 3b, 4c. Annual, biennial or perennial herbs; variously pubescent. Leaves alternate, sometimes forming a rosette, petiolate and simple, vari- ously sinuate to dentate, rarely entire. Inflorescences of solitary flowers in the leaf axils, condensed to form usually secund, scorpioid cymes. Flowers zygomorphic, sessile or shortly pedicellate; calyx tubular-campanulate or urceolate, the lobes often spine-tipped; co- rolla campanulate or funnelform, the lobes unequal; stamens inserted near the base of the corolla tube; pollen trizonocolporate with long, distinct colpi and weakly striate ornamentation; ovary bilocular with an indistinct disc. Fruit a globose or ovoid capsule, with circumscissile dehiscence; fruiting calyx enlarged, enclosing the fruit, the lobes spine-tipped. Seeds reniform or discoid, strongly compressed. Fields, waysides, and hedges; 0-3600 m. (Zhang etal. 1994; Hawkes, 1972; Al-Musawi, 1979). LIST OF SPECIES (Schonbeck-Temesy, 1972; Al-Musawi, 1979; Feinbrun-Dothan, 1978). Hyoscyamus albus L. (Hyoscyamus arenarius Dunal, H. canariensis Ker-Gawl., H. clusii G. Don, H. major Mill., H. minor Mill., H. varians Vis.), Mediterranean to Iraq and Egypt; Hyoscyamus aureus L., E. Mediterranean to NW Iraq, Sinai, and Egypt; Hyoscyamus flaccidus Wright, Arabia; Hyoscya- mus gallagheri A.G. Mill. & J.A. Biagi, Oman; Hyoscyamus grandiflorus Franch., tropical Africa; Hyoscyamus insanus Stocks (Hyoscyamus angulatus Griff., H. nutans Schonb.-Tem., H. orthocarpus Schonb.-Tem., H. rosularis Schonb.-Tem., H. tenuicaulis Schonb.-Tem.), N. Africa and the Middle East; Hyoscyamus leptocalyx Stapf., W. Iran; Hyoscyamus longipedunculatus Townsend, Iraq; Hyoscyamus malekianus Parsa, Iran; Hyoscyamus muticus L. (Hyoscyamus betaefolius Lam., H. boveanus (Dunal) Ascher & Schweinf., H. datora Forsk., H. falezlez Coss., Scopolia boveana Dunal, S. datora (Forsk.) Dunal, S. mutica (L.) Dunal), N. Africa and the Middle East; Hyoscyamus niger L. (Hyoscyamus agrestis Kit., H. auriculatusTenore, H. bohemicus F.W. Schmidt, H. lethalis Salisb., H. pallidus Waldst. & Kit., H. persicus Boiss. & Buhse, H. pictus Roth, H. syspirensis C. Koch, H. verviensis Lej.), widespread in temperate Eurasia; Hyoscyamus pusillus L. (Hyo- scyamus micranthus Ledeb., H. pungens Griseb.), Egypt to SW and C Asia; Hyoscyamus reticulatus L. (Hyoscyamus afghanicus Pojark., H. arachnoideus Pojark., H. camerarii Fisch. & Mey., H. coelosyriacus Bornmuller, H. issa-sadiqui Parsa, H. kopetdaghi Pojark., H. kotschyanus Pojark., H. kurdicus Bornmuller, H. leucanthera Bornm. & Gauba, H. multicaulis Rech. f. & Edelb., H. pinnatifidus Schldl., H. pojarkovae Schonb.-Tem., H. purpureus Griseb., H. squarrosus Griff.), Egypt to SW Asia; Hyoscyamus senecionis Willd. (Hyoscyamus pinnatisectis Boiss.), Egypt through the Middle East; Hyoscyamus tibesticus Maire (Hyoscyamus cylindrocalyx Rech. f., H. desertorum (Asch.) Ta'ckh.), N. Africa in Sahara to theArabian pemnsu\a\Hyoscyamus turcomanicus Pojark., Trans-Caspian area in Iran, Uzbekistan. 5. Mandragora L., Sp. pi. 1: 180 (1753). Type: Mandragora officinarum L. Figs 3e, 5c, d. Perennial herbs from enlarged taproots; variously pubescent. Leaves alternate, forming a dense basal rosette, very short petiolate or sessile, simple, entire or dentate. Inflorescences of solitary flowers PHYLOGENETIC CONSPECTUS OF HYOSCYAMEAE 27 in the leaf axils. Flowers actinomorphic; calyx flared or cup-shaped, deeply lobed, the lobes long-triangular; corolla flared or cup-shaped, deeply lobed; stamens inserted in proximal part of the corolla tube; pollen cryptaperturate with symmetrical pattern of endoaperture thinnings and gammate-baculate ornamentation (Fig. 7d); ovary bilocular, with an indistinct disc. Fruit a globose or ovoid, fleshy berry, yellow-orange, greenish or white flushed with purple, usually borne beneath the leaves; fruiting calyx slightly enlarged, not en- closing the berry. Seeds reniform, very large. Grasslands, woods, hedges and waysides, stony hillsides and screes; 0-4200 m. (Zhang etal., 1994;Hawkes, 1972). LIST OF SPECIES (Pojarkova, 1955; Schonbeck-Temesy, 1972; Jackson & Berry, 1979; Zhang et al., 1994). Mandragora autumn- alis Bertol. (Mandragora femina Gersault, M. microcarpa Bertol., M. officinalis Moris., M. officinarum Bertol., M. officinarum L. pro parte), Mediterranean; Mandragora caulescens C.B. Clarke (Anisodus caulescens (C.B. Clarke) Diels, A. mariae Pascher, Mairella yunnanensis H. Lev., M. tibetica Grubov.), Himalayas in India, Nepal, Bhutan, and China; Mandragora chinghaiensis Kuang & A.M. Lu, China, restricted to pika warrens (fide M. Gil- bert); Mandragora officinarum L. (Atropa acaulis L., Mandragora acaulis Gaertn., M. haussknechtii Heldr., M. hispanlca Vierhapper, M. mas Gersault, M. neglecta G. Don, M. officinalis Mill., M. praecox Sweet, M. vernalis Bertol.), widespread in Eurasia; Man- dragora turcomanica Mizg., Turkmenistan near Caspian Sea, Russia. 6. Physochlaina G. Don, Gen. Hist. 4: 470 (1837). Type: Physochlaina physaloides (L.) G. Don (basionym Hyoscyamus physaloides L.). Figs 3a, 4d. Perennial herbs; glabrous or variously pubescent. Leaves alternate, petiolate, simple, entire to sinuate. Inflorescences axillary or termi- nal, branched several times. Flowers actinomorphic; calyx tubular-campanulate to tubular-urceolate; corolla campanulate or funnelform, purplish; stamens inserted midway up the corolla tube; pollen either pentacolporate, occasionally tetracolporate, with long, indistinct colpi and scabrate ornamentation or trizonocolporate with long, distinct colpi and striate-rugulate ornamentation (Fig. 6a, b); ovary bilocular, with a fleshy, annular disc. Fruit a globose or oblong capsule, with circumscissile dehiscence; fruiting calyx subcoriaceous and inflated, longer than the capsule. Seeds reniform, compressed. Grasslands and forest edges; 800-4500 m. (Zhang et al., 1994; Pojarkova, 1955). LIST OF SPECIES (Zhang et al., 1994; Kuang & Lu, 1981; Schonbeck-Temesy, 1972; Pojarkova, 1955). Physochlaina capi- tata A.M. Lu, China; Physochlaina infundibularis Kuang, China; Physochlaina macrocalyx Pascher, China; Physochlaina macrophylla Bonati, China; Physochlaina orientalis G. Don (Hyo- scyamus orientalis (G. Don) Bieb., Physochlaina dubia Pascher, Scopolia orientalis(G. Don) Dunal), Caucasus to Iran; Physochlaina physaloides (L.) G. Don (Atropa physaloides Georgi, Hyoscyamus physaloides L., Physochlaena dahurica Miers, Physochlaina physaloides (L.) Miers, P. pseudophysaloides Pascher, Scopolia physaloides (L.) Dunal), China, Mongolia, Russia (Siberia); Physochlaina praealta (Dene.) Miers (Belenia praealta Dene., Hyoscyamus praealtus (Dene.) Walp., Physochlaina grandiflora Hook., P. urceolata Kuang & A.M. Lu, Scopolia praealta (Dene.) Dunal), China, Nepal, Pakistan, and India (Kashmir); Physochlaina semenowii Regel, central Asia. 7. Przewalskia Maxim, in Bull. Acad. Petersb. 27: 507 (1881). Type: Przewalskia tangutica Maxim. Figs 5a, b. Perennial herbs, from an elongate fleshy taproot; pubescent, the trichomes glandular. Leaves alternate, forming a dense basal rosette, simple, entire. Inflorescences of clusters of flowers in the leaf axils. Flowers actinomorphic; calyx tubular-campanulate; corolla tubular- funnelform, greenish yellow or violet; stamens inserted in the distal part of the corolla tube; pollen trizonocolporate with long, distinct colpi and reticulate ornamentation; ovary bilocular, with an annular disc. Fruit a globose capsule, with circumscissile dehiscence; fruit- ing calyx much enlarged and the tube inflated with prominent reticulate veins, completely enclosing the much smaller fruit, the lobes constricted with incurved lobes. Seeds reniform. Open habi- tats on the Qinghai-Tibet plateau, sand dunes, road margins and areas of frost heave; 3200-5000 m. (Zhang et al., 1994). LIST OF SPECIES (Zhang et al. 1 994). Przewalskia tangutica Maxim. (Mandragora shebbearei C. Fischer, Przewlaskia robo-rowskii Batalin, P. shebbearei (C. Fischer) Grubov), China. 8. Scopolia Jacq., Obs. Bot. 1: 32 ( 1 764). Type: Scopolia carniolica Jacq. Fig. 3c. Perennial herbs; glabrous to minutely pubescent. Leaves alternate, petiolate, simple, entire. Inflorescences of solitary flowers in the leaf axils. Flowers subactinomorphic; calyx cup-shaped with irregular lobes, one usually much longer than the rest; corolla campanulate- funnelform, greenish yellow to reddish purple; stamens inserted near the base of the corolla tube; pollen trizonocolporate with long, indistinct colpi and scabrate ornamentation (Fig. 7a); ovary bilocu- lar, with an annular disc. Fruit a globose capsule, with circumscissile dehiscence; fruiting calyx somewhat enlarged, enclosing the fruit. Seeds subreniform. Woodlands; 500-1500 m. (Lu & Zhang, 1986; Pojarkova, 1955). LIST OF SPECIES (Sandina, 1980; Lu & Zhang, 1986). Scopolia carniolica Jacq. (Hyoscyamus scopolia L., Scopolia atropoides Bercht. & Presl, S. caucasica Kolesn., 5. hladnikiana Fleishm., 5. parviflora (Dunal) Nakai, 5. trichotoma Moench, 5. tubiflora Kreyer, Scopolina atropoides Schult.), Alps, Carpathian mountains, Cauca- sus; Scopolia japonica Maxim., Japan, Korea. ACKNOWLEDGMENTS. This study was in partial fulfilment of the require- ments for an M.Sc. in Pure and Applied Taxonomy at the University of Reading undertaken by A.L.Hoare and supervised by Barbara Pickersgill and S. Knapp. We would like to thank Mike Gilbert for sharing his field knowledge of some of the Chinese species of this group with us and for the use of photographs; Viveca Persson for help with the pollen preparation and description; Johannes Vogel for the use of photographs; Simon Thornton- Wood for help with computing; Malcolm Penn for producing the distribution map in Fig. 1; Mike Gilbert, James Mallet, and Chris Humphries for reviewing the manuscript; the staff of the EM Unit at The Natural History Museum for training and encouragement; the staff of the Photographic Unit at The Natural History Museum for the production of the photographic plates; staff at the Royal Botanic Gardens, Kew and the Chelsea Physic Garden, especially Fiona Crumley, for cultivating living plants for this study; and the staff of the School of Plant Sciences at the University of Reading for help during the M.Sc. course in Pure and Applied Taxonomy undertaken by the senior author. 28 A.L. HOARE AND S. 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Ferguson (Eds), Pollen and spores: form and function: 239-256. London Zhang Zhiyun & LuAnming 1 984. Pollen morphology of the subtribe Hyoscyamineae, (Solanaceae). Acta phytotax. sin. 22: 175-180. & D'Arcy, W.G. 1994. Solanaceae. In Wu Zhengji & P.R. Raven (Co-chairs of the Editorial Committee), Flora of China 17: 300-332. APPENDIX I Specimens examined for pollen analysis (all BM). Anisodus luridus Link - Beer et al. 9410 (Nepal). Atropa belladonna L. - Mohamed 146 (Morocco). Atropanthe sinensis (Hemsl.) Pascher - Wilson 2594 (China, Hupeh). Hyoscyamus muticus L. - Hildebrandt 71 (Egypt). Hyoscyamus niger L. - Davis 52541 (Algeria). Hyoscyamus senecionis L. - Thesiger 189 (Afghanistan). Mandragora caulescens C.B. Clarke - Gardner 479 (Nepal); Polunin et al. 4696 (Nepal). Physochlaina physaloides (L.) G. Don - Heward s.n., July 1847 (India). Physochlaina praealta (Dene.) Miers - Stachey & Winterbotiom s.n. (China, Tibet). Przewalskia tangutica Maxim. - Richardson 56 (China, Tibet). Scopolia carniolica Jacq. - Harris Garden, University of Reading Bull. nat. Hist. Mus. Land. (Bot.) 27( 1 ): 3 1-73 Issued 27 June 1997 A revision of Solanum section Pteroidea: Solanaceae SANDRA KNAPP_ ^ Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD THORUNN HELGASON Department of Biology, University of York, P.O. Box 373, York YO1 5YW CONTENTS Introduction 31 Taxonomic and nomenclatural history 32 History of section Pteroidea 32 Morphology and natural history 33 Stems 33 Leaves 33 Inflorescences 36 Trichomes 36 Flowers 37 Fruits 37 Seeds 37 Cladistics 42 Character coding and tree construction 42 Classification 43 Taxonomic treatment 44 Key to selected groups of Neotropical non-spiny solanums 44 Key to species of Solanum section Pteroidea 44 The Solanum ternatum species group 45 1. Solanum incurvum Ruiz & Pav 45 2. Solanum ternatum Ruiz & Pav 45 The Solanum mite species group 51 3. Solanum anceps Ruiz & Pav 51 4. Solanum angustialatum Bitter 54 5. Solanum chamaepolybotryon Bitter 55 6. Solanum conicum Ruiz & Pav 55 7. Solanum mite Ruiz & Pav 58 8. Solanum savanillense Bitter 62 9. Solanum trizygum Bitter 64 10. Solanum uleanum Bitter 68 Excluded species 69 References '* Exsiccatae ' ' Index 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.l. 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- INTRODUCI [ON toes mo rellas, 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 phytogeny (see D'Arcy, 1991). Taxonomic research effort has been concen- of non-spiny solanums (see Knapp, 1986o; Knapp, 1989; Knapp, The Natural History Museum, 1997 32 S. KNAPP AND T. HELGASON 199 la) we have investigated the small, primarily rainforest species of section Pteroidea 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). It is 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 Solarium mite species group and the Solatium 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 Solarium 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 ( 1 852) and while 900 species were treated in the Prodromus, at least 4000 specific epithets exist for Solarium at present. By convention and for convenience Solarium is usually divided into two main groups, the spiny solanums (subgenus Leptostemonum) and the non-spiny solanums (the rest: subgenera Solarium, 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 Solarium both before and after Dunal (1852) see Knapp (1989, 199 la) and Bohs (1994). Knapp (1989) also provides a list of recent monographs of sections of Solarium, to which can be added a monograph of Solarium section Allophyllum (Bohs, 1990) and the genus Cyphomandra (Bohs, 1994; now with all epithets transferred to Solarium, see Bohs, 1995). Table 1 Characters used to define the major Neotropical subgenera of Solarium (after D'Arcy, 1972). Solarium c. 1 500-2000 species subgenus Solarium subgenus Bassovia subgenus Brevantherum subgenus Potatoe subgenus Leptostemonum 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 History of section Pteroidea The first species of section Pteroidea to be described was Solarium anceps (as Bassovia sylvatica), described by Aublet (1775) from what is now French Guiana. Several more species were described by Ruiz & Pavon (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 naturelle, medicate et economique 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, 1 983). The group composed of the species of section Pteroidea was explicitly given sectional rank ('la section designee 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, 1 986a). 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 ( 1 844) 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 5. 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 Veil., 5. laurinum Dunal, 5. lacteum Veil., 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; bacca globosa 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 33 The first taxonomist to recognize the close relationship between simple and pinnate-leaved taxa of what is now section Pteroldea 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 5. conicum, S. mite, S. trizygum, S. fraxinellum, S. quinquefoliololatum, S. chamaepolybotryon, S. diffusum, S. ternatum, S. pteleifolium, S. conjungens, S. hederiradiculum, S. angustialatum, and 5. 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. la, b). Occasionally some species (e.g. 5. 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 Pteroidea 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 5. 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 is only very weakly developed in 5. mite. Plants range from quite small (a few centimetres in S. conicum and 5. chamaepolybotryon) to more than a metre in height (e.g. 5. 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, 1 989) 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. 5. 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. 5. chamaepolybotryon, S. ternatum) are quite rubbery in texture, often drying quite thick on Fig. 1 a) Lowland forest habitat of 5. anceps, S. mite, S. conicum, S. uleanum: Rio Palcazu valley, Pasco, Peru, b) Cloud forest habitat of 5. trizygum: Cerro Pando, Chiriquf, Panama. REVISION OF SOLANUM SECTION PTEROIDEA 35 36 S. KNAPP AND T. HELGASON Fig. 3 Sympodial structure in Solatium section Pteroidea (modified from Danert, 1967). herbarium sheets. Solatium 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. 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 ofOleria, a diverse group along the eastern Andean slope. Larvae ofOleria 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 5. 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 37 (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. Solatium ternatum has densely pubescent juvenile foliage (see Fig. 1 1 , Knapp & Mallet 6626) but mature specimens are nearly always glabrate suggesting a developmental aspect to pubescence density. Plants described as 5. dendrophilum (here treated as a synonym of 5. ternatum) have dense pubescence more reminiscent of juvenile leaves than other mature individuals of 5. 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 5. 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 5. ternatum species group that in theS. 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 5. mite species group are shorter and stouter in relation to the corolla lobes than those of theS. 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 5. 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 (5. 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 (5. 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. 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 (5. capsiciforme (Domin) G.T.S. Baylis, 5. nigricans M. Martens & Galeotti, 5. aligerum Schldl., 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 5. 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 5. mite to long-pointed in 5. anceps. In 5. 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 5. 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 5. 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 (Soueges, 1907; Lester &Durrands, 1984; Edmonds, 1983; Bohs, 1994). Seed colour is not uniform in section Pteroidea. Seeds of the species in the 5. mite species group tend towards greenish brown, whereas the 5. 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, 2 w C PQ 13 ' Si 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). 40 S. KNAPP AND T. HELGASON I 5 1 I S d REVISION OF SOLANUM SECTION PTEROIDEA 41 Fig. 7 a) Digested testa of 5. 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 5. anceps (Franco et al. 1 42 S. KNAPP AND T. HELGASON 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 30C. 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, 199 la). Two species have projections from the thickened walls, S. incurvum (hair-like, Fig. 6a) and 5. 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 5. 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, 1991/j; 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 5. 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 Table 4 Character set used in the HENNIG86 analysis of Solanum section Pteroidea. 0. Corolla diameter: >10 mm = 0; 5-10 mm = 1 ; <5 mm = 2 1. Corolla texture: papery =0; fleshy = 1 2. Number of flowers per inflorescence: few (<40) = 0; many (>40) = 1 3. Bud shape: globose = 1; elliptic = 4. Corolla in bud: exserted = 0; +/- included = 1 5. Calyx lobe shape: deltate = 0; quadrate = 1 6. Apex of calyx lobes: rounded = 0; apiculate = 1 7. Corolla tube: long (the corolla divided only l /2- 3 A 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 = 1 9. Seed shape: flattened-reniform = 0; ovoid-reniform = 1 10. Seed number per berry: many (>60) = 0; few (<60) = 1 1 1 . Seed colour: brown = 0; reddish = 1 ; green to green-brown = 2 1 2. 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 = 17. Fruit apex: rounded = 0; elongate = 1 18. Fruit texture: smooth = 0; rugose = 1 19. Mature fruit position: nodding = 0; erect = 1 20. Leaf shape: simple = 0; deeply divided (pinnate, pinnatifid or ternate) -1 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 = 1 Table 5 Data matrix used in HENNIG86 analysis of Solanum section Pteroidea. 11111 11111 22222 01234 56789 01234 56789 01234 nudum ternatum incurvum anceps angustialatum chamaepolybotryon conic um mite savanillense trizygum uleanum 10010 00001 20000 1 1000 01 101 01000 110000001001001 10010 000000000001000 1101001011 1101000111 12121 1111001001 2111100011 12121 1111001101 1001000111 !???! 11071 10101 2000001111 10121 10011 11011 1111000111 10121 11001 11101 2000001101 12121 00011 11101 1001000111 12121 10011 11001 10100 11101 10121 01111 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, \99lb). 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 5. 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 r-m 2 1 S.nudum group 1 5 6 13 15 19 20 21 24 i i 1 i i ft i inn t- 03 9 10 23 HHHHHh 00001 S. ternatum 111101100 11 18 S. incurvum i i 7 r , n 17 S.anceps 1.. l.JU. I i 12, 2 4 22 f-l II r ,- r , 8 12 13 14 18 fill! S.angustialatum 2111 21 1 1 1 1 ' 11211 18 ^ S.chamaepolybotryon i ^r^ 01 I 2 S.mite 7 19 20 l * 1 -!.. 11 S.tnzygum 4- 2 s? II S.coniturn 3 6 o i "I"! ~ 5. suvanillense is 22 | 2 2 2 5 16 17 23 1 | r i./jj .... I 1 1 1 la. uleanum 11111 Fig. 8 Cladogram of Solatium 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 5. 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 5. incurvum basal to the clade. Table 6 Classification of Solanum section Pteroidea. Solanum ternatum species group S. ternatum Ruiz & Pav. 5. incurvum Ruiz & Pav. Solanum mite species group Solanum anceps clade S. anceps Ruiz & Pav. 5. angusitalatum Bitter Solanum mite clade 5. mite Ruiz & Pav. 5. chamaepolybotryon Bitter S. trizygum Bitter 5. conicum Ruiz & Pav. S. savanillense Bitter 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 ofWhalen, 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 Solatium section Pteroidea Dunal, Hist. nat. Solanum: 43 (1813). Lectotype species: Solanum mite Ruiz & Pav. (D'Arcy, 1972). Bassovia Aubl., Hist. pi. 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. 1 1 : 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 (Aubl.) Bitter [basionym Bassovia sylvatica Aubl.] (= Solanum anceps Ruiz & Pav.) (Bitter, 1921). S. KNAPP AND T. HELGASON placed in synonymy. Most of these are synonyms of the two most widely distributed species, S. mite and 5. 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 & Pavon 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 & Pavon, 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, 199 la), 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 Index 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 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 5. 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 1 Inflorescences axillary 2 anthers poricidal at the tips, with age splitting longitudinally, bright Inflorescences lateral or leaf-opposed 4 yellow. Fruit a berry, usually green or yellowish green when mature, globose with a smooth surface (5. incurvum, S. ternatum, S. mite) or 2 Plants sma11 trees or shrubs ' branch ; n 8 in a com P lex crown ; inflores - ., ,. ,- ... ,. , ... cences in branch forks; anthers with an enlarged connective variously conical with a rugose surface; fruiting pedicel nodding or Cyphomandra (Solanum section Cyphomandra) erect. Seeds flattened-reniform, many per fruit (S. ternatum, S. incurvum) or ovoid-reniform and few per fruit. Plants wand-like, shrubs, vines, or herbaceous; inflorescences only in leaf axils; anthers without an enlarged connective 3 Section Pteroidea, as here delimited, consists of ten species in two monophyletic clades (see p. 43). The clades are both kept in the 3 Traili "g herbs > rootin g at the nodes ' inflorescences with a single flower; ... - . , . , ,. fruit with smooth surfaces Solanum section Herpystichum section (s.l.) at this time for convenience, despite some doubt as to their degree of relatedness. The 5. ternatum clade, consisting of 5. Herbs (not trailing), slender shrubs or vines, inflorescences with more incurvum and 5. ternatum, has large, lilac or pinkish flowers and than one flower, usually with up to 30 flowers; fruit smooth or rugose globose fruits with many, small, flattened seeds, while the larger 5. Solanum section Pteroidea mite clade, consisting of 5. anceps, S. angustialatum, S. 4 inflorescences internodal; fruit brightly coloured, with thin pericarp chamaepolybotryon, S. conicum, S. mite, S. savanillense, S. trizygum, Solanum section Solanum and 5. uleanum, has smaller, usually greenish flowers, and (with the exception of 5. mite) conical fruits with unusual rugose surfaces, the Inflorescences leaf-opposed or vanously nal; fruit g. 'een .at matu- .,,,., T,, nty, the pericarp not thin Solanum section Gemmata fruits having a few, ovoid, often bright green, seeds. These rugose fruits are unique in Solanum. The section has been accorded ifcy to SU&Aes of Solatium SKC&on Pteroidea subgenenc status by many previous authors (Bitter, 1921; Seithe, 1962;Danert, 1967; D'Arcy, 1972, 1991), on the basis of its extreme 1 Leaves simple 2 morphological difference from the rest of Solanum. We feel, how- ^^ various , y pinnate 4 ever, that until phylogenetic relationships inSolanum are much more clearly resolved, the group (as two clades) should be recognized 2 Climbing herbs; flowers 1.2-1. 4 cm in diameter, purplish; fruit globose, only at the sectional level. the surface smooth; seeds man y ^ fruit (>50) l ' S ' incurvum This monograph is based on herbarium specimens and the exten- Terrestrial herbs or weak subshrubs; flowers 0.4-0.7 cm in diameter, sive field observations of the senior author. The species are delimited white or greenish white; fruit ovoid, beaked, the surface rugose; seeds on morphological grounds, with geographical and ecological prefer- few per fruit (usually 10-40) 3 ences being taken into account where appropriate. More than half of 3 Stem prominently winged; style densdy pub escent along its entire the published names of this species group have as a result, been , ength San Martm? Peru 4. S. angustialatum REVISION OF SOLANUM SECTION PTERO1DEA Stem smooth, terete, not winged; style glabrous or at most papillate in the lower 2/3. Widespread 3. S. anceps 4 Climbing herbs or woody vines 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 smooth 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 rugose 10. S. uleanum 6 Leaves with 5 or fewer leaflets, the leaflets usually obovate, especially the terminal 7 Leaves usually with more than 5 pairs of leaflets, the lateral leaflets lanceolate to elliptic, the terminal leaflet similar in shape, not markedly obovate 10 7 Leaves pubescent on the veins and lamina on both surfaces 8 Leaves glabrous on lamina, occasionally pubescent along the veins and rachis 9 8 Fruit conical; leaf pubescence denser adaxially 8. S. savanillense Fruit globose, smooth; leaves equally pubescent on both surfaces 7. S. mite 9. Fruit conical, the surface rugose; leaves fleshy; plants very small and rooting along the stem 5. S. chamaepolybotryon Fruit globose or at most apically pointed, the surface smooth; leaves membranous; plants often woody at the base and up to 1 m tall 8. S. mite 10 Flowers 5-6 mm in diameter, the petals strongly reflexed at anthesis; fruit globose, smooth 8. S. mite Flowers 9-13 mm in diameter, the petals usually planar or only slightly reflexed at anthesis; fruit conical, rugose 1 1 1 1 Flowers > 1 mm in diameter; leaflets long-petiolulate, the petiolule 3- 17 mm; leaves densely pubescent in a groove along the adaxial side of the rachis 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 9. S. trizygum The Solanum ternatum species group 1. Solanum incurvum Ruiz & Pav., Fl. peruv. 2: 34, fig. 154b (1799). Type: Peru, Huanuco, Muna, August, September, Ruiz & Pavon s.n. (MAMectotype [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 1 2 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, 7527'W, 1023'S, 1 3 July 1984, Smith 7756 (MO, USM); Oxapampa. Rio San Alberto valley E. of Oxapampa, 2700 m, 7522'W, 1027'S, 26 July 1984, Smith & Poetel 8069 (MO). San Martin: Valley of Rio Apisoncho, 30 km above Jucusbamba, 2800 m, 7710'W, 755'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, 5. ternatum, it is reported to have lilac flowers, but variation for flower colour may exist. 2. Solanum ternatum Ruiz & Pav., Fl. peruv. 2: 38, fig. 162b ( 1 799). 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, Huanuco, 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 J 1 Fig. 9 5. incurvum. Habit: Gentry et al. 30871 (MO). Inflorescence: Hamilton & Holligan 1069 (K). (1913). Type: Peru, Huanuco, Muna, May 1863, Pearces.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: 1 42 ( 19 13).Type: Peru, Huanuco, Muna, 10- 11 000 ft, May 1863, Pearce s.n. (K!-holotype). Solanum subquinatum Bitter in Reprium nov. Spec. Regni veg. 12: 1 44 ( 1 9 1 3). Type: Peru, Amazonas, Chachapoyas, 1 835, Matthews s.n. (BMMectotype; K!-isolectotype). Solanum diffusum subsp. miozygum Bitter in Bot. Jb. 54: (Beibl. 1 19): 14 (1916). Type: Peru, Pasco, valley of Rio Pozuzo, tribu- tary of Rio Palcazu, 946'-950'S, 2200 m, 1909-1914, Weberbauer 6783 (B-holotype, destroyed; MOLI-lectotype; F!- isolectotype). Solanum diffusum var. miozygum (Bitter) J.F. Macbr. in Publ. Field Mus. (BoL) 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 PTERO1DEA 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, 9-15x8-12 cm, ternate or with 2-A 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 45 x c. 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. & Melief2952 (MO). Huila: Finca Merenberg, E. of Volcan Purace, near Cauca border, 2300 m, 7602'W, 216'S, 3 April 1986, Gentry et al. 53970 (MO); Finca Merenberg, border with Cauca, E. of Leticia, 2300 m, 7612'W, 216'S, 08 July 1984, Gentry et al. 47779 (MO). Magdalena: Alrededores de Yerbabuena, 2000 m, 26 January 1959, Romero Castaneda 7067 (AAU); Sierra Nevada de Santa Marta, Sierra del Libano, Las Nubes, 1898-1901, Smith 1 162 (BM, BR. F, MA, MO, NY, US, W, WIS). Norte de 48 S. KNAPP AND T. HELGASON Fig. 11 5. ternatum. Habit: Killip & Smith 20235 (GH). Juvenile foliage and flowers: Knapp & Mallet 6626 (US). Fruits: Zaruma et al. 21 A (QCNE). Santander: Pica-Pica Valley, above Tapata (N. of Toledo), 2100-2400 m, 1 March 1927, Killip & Smith 20235 (GH, US). VENEZUELA. Aragua: E. of Colonia Tovar, 7500 ft, 8 April 1854, Fendler 1017 (GOET). Miranda: Colonia Tovar, 1800-2000 m, December 1924,A//arf335(US). ECUADOR. Napo: Carretera Hollin-Loreto, km 25, Centre Challuayacu, en trocha hacia la zona del Guagua Sumaco, 1230 m, 7740'W, 0043'S, 10 November 1988, Hurtado &Alvarado 1121 (MO); Carretera Hollin-Loreto, km 40-50, alrededores de la comunidad Huamam y del Rio Pucuno, 1 200 m, 7736'W,0043'S, 10 October 1988, Hurtado625 (MO). Pasta/a: Capitaine Chiriboga, Rio Pastaza, vicinity of army base, 235 m, 7649'W, 232'S, 21 July 1988, Lewis et al. 13771 (QCNE); 2 km al NE de Mera, Hacienda San Antonio del Bar6n von Humboldt, 1100 m, 7806'W, 0127'S, 18 March 1985, Zaruma 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, 14July \9%6,D'Arcy 16506 (MO); Rio Nangaritza, Pachicutza, camino al hito de Pachicutza, 900-1000 m, 7807'W, 407'S, 18 October 1991, Palacios et al. 8188 (QCNE). PERU. Amazonas: Prov. Chachapoyas, 1 836, Matthew ss.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, V/asshausen & Encarnacion 998 (US); Mendoza, 1600 m, 2 September 1963, Woytkowski 8265 (MO); Bagua, Cordillera Colan SE of La Peca, 2280-2400 m, 7 October 1978,fiarfcowr3829(MO), 1800- 1870m, 17 October 1978, Barbour 4160 (MO). Cajamarca: Cuchero, Dombey s.n. (P [n.v., Morton neg. 8354, F!, MO!, US!]); San Andres de Cutervo, sobre la ruta a las grutas, alN.de SanAndres, 2250 m, 25 June 1 989, Sanchez Vega 4895 (F). Cusco: Dtto. Camanti, Maniri, 8 km W. de Quincemil, a los margenes de la quebrada Garrote, 720 m, 7048'W, 1 3 1 7'S, 20 July 1 990, Timand & Astete 692 (MO); along Rio Pillahuata, 2300-2400 m, 3 May 1925, Pennell 14012 (F); Rio Mapitunuari, c. half way from Luisiana and Rio Apurimac to camp 1 , 800-900 m, 7342'W, 1239'S, 15 June 1968, Dudley 10152 (F). Huanuco: Muna, trail to Tambo de Vaca, 2440 m, 5 June 1923, Macbride 427 (G, F); Huacachi, estacion near Muna, 1980 m, 20 May 1923, Macbride 4698 (F); Muna, May 1863, Pearce 135 (BM); Divisoria, 1600 m, 10 September 1946, Woytkowski 345 1 2 (F, MO); Rupa Rupa, Calpar Bella, Cueva de los Huarinos (margen izquierda del Rio Monzon), 700-900 m, 29 June 1976, Schunke V 9440 (GH, MO); La Divisora, Cordillera Azul near border with Ucayali, 1 620- 1760 m,75 48'W,905'S, lOAugust 1980, Gen/ryetal.29558(MO);Pachitea, Codo de Pozuzo, alluvial fan flood plain of Rfo Pozuzo after it emerges from mountains, trail S. of settlement to main river, 450 m, 7525'W, 940'S, 21 October 1982, Foster9355 (MO); Dtto. HermilioValdizan, La Divisoria, road from Pumahuasi to La Cumbre, 1600-1660 m, 26 June 1978, Plowman & Schunke V. 7394 (MO); Prov. Huanuco, km 452 of Lima-Tingo Maria road, 2500 m, 2 June 198 1 Young & Sullivan 570 (MO); Prov. Leoncio Prado, road between Jingo Maria and Pucallpa, km 35, 1500 m, 7548'W, 9 10'S, 3 June 1981, Sullivan & Young 1 154 (MO); Muna, 1000-1 100 m, 1863, Pearce 144 (BM). Junm: Huatsiroke, 1800 m, 21 February 1960, Woytkowski 5543 (F, MO); Prov. Tarma. Agua Dulce, 1 900 m, 5 March 1 948, Woytkowski 354J6(F, G, MO, US); San Gavan, August 1854, Lechler2440(G, P [n.v. Morton neg. 8252, F,GH, US]); Pichis trail, Dos de Mayo, 1700- 1900m, 2 July \929,Killip & Smith 2581 1 (US). Pasco: San Juan de Cacazu, km 36 on Villa Rica-Pto. Bermudez road, trail behind colegio, 950 m, 7510'W, 1038'S, 13 August 1 984, Knapp & Mallet6626 (BH, K, US); Oxapampa-Cerro de Pasco road, La Suiza to San Gotardo, 2100-2650 m, 7535'W, 1 138'S, 19 May 1983, Smith 4104 (MO); Rio San Alberto valley E. of Oxapampa, slopes of Cordillera Yanachaga, 2400 m, 7522'W, 1034'S, 23 July 1984, Smith & Pretel 7968 (MO); ElTunquiAlto, 57 km from Oxapampa, 1700m,7530'W, 1015'S, 14 May 1982, Smith et al. 1569 (MO); Oxapampa, trail to summit of Cordillera Yanachaga via Rio San Daniel, 2400 m,7527'W, 1023'S, 19 July 1984, Smith et al. 7933 (MO). San Martin: Valley of Rio Apisoncho, 30 km above Jucusbamba, 2800 m, 77 1 0'W, 0755'S, SAugust 1965, Hamilton & Holligan 1078(K);Zepelacio,nearMoyabamba, 1 100m,June 1934,A7wg3665(A,BM, GH, K, MO, US). Ucayali: Rfo Chino al W. del Restaurant Acapulco, 100- 1 100 m, 5 June 1976, Schunke K 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, 6734'W, 1535'S, 31 October 1984, Solomon & Nee 12704 (M, MO); Prov. Sur Yungas, along road 7.0-9.4 km NE of (above) Huancane, 2286-2499 m, 6732'W, 1620'S, 17 May 1990, Luteyn & Dorr 13699 (NY); Prov. Nor Yungas, 4.6 km NE (below) Chuspipata on road to Yolosa, 2800 m, 6747 r W, 1617'S, 8 March 1984, Solomon & Stein 1 1681 (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, 6737'W, 1558'S, 12 February 1983, Solomon 9584 (MO); Hacienda Casana sobre el camino a Tipuani, 50 S. KNAPP AND T. HELGASON Fig. 13 S. anceps. Habit: Allard 22077 (US), (inset circle) S. angustialatum stem from Knapp & Mallet 8567 (F). REVISION OF SOLANUM SECTION PTEROIDEA 51 1400 m, 15 October 1922, Buchtien 7462 (US); Prov. SudYungas, Huancane (cerca Chulumani) 8 kms, 2450 m, 3 1 October 1 98 1 , Beck 488 1 (F); Prov. Nor Yungas, 4.6 km below Yolosa, then 19.1 km on road up the Rio Huar-inilla, 1700 m, 6753'W, 1612'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 5. 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 5. 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., Fl. peruv. 2: 36, fig. 149a (1799). Type: Peru, Huanuco, Cuchero, July, August, Ruiz & Pavon s.n. (MA!-holotype [F neg. 29722, F!, GH!, MO!, US!]). Figs 5c, 13. Bassovia sylvatica Aubl., Hist. pi. Guiane 1: 217, fig. 75 (1775). Type: French Guiana, Aublet s.n. (BMMectotype). Solanum bassovia Dunal in Poir., Encycl. suppl. 3: 754 (1814); Solan, syn.: 22 (1816). nom. nov. for Bassovia sylvatica Aublet. Solanum aubletii Pulle, Enum. vase. pi. Surinam: 411, fig. 16 (1906). nom. nov. for Bassovia sylvatica Aubl. Solanum conjungens Bitter inReprium nov. Spec. Regni veg. 11:12 (1912). Type: Ecuador, Tungarahua, prope Bafios, September 1892, Sodiro 1 14/61 (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 (191 2). Type: Peru, Loreto,Yurimaguas, August 1902, Ule 6276 (B-holotype, destroyed [F neg. 2608, F!, G!, GH!, MO!, US!]; HBGMectotype). Solanum theobromophyllum Bitter inReprium 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. Regniveg. 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 191 1. 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, 1 2 -45 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. Inflorescence 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 1 .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. 650'N, 7518'W, 2000 m, 8 February 1986, Stein & Cogollo 3394 (MO). Boyaca: 130 miles NW of Bogota, 3000 ft, 29 September 1932, Lawrence 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, Quinones 1045 (MO). Putumayo: Orito, Rio Calderas, 300-400 m, 1 1 December 1968, Plowman2l29 (GH). Valle: Cerca a Morales-Cauca, 8 October 1968, Espinal T. & Ramos 2943 (CUVC, F); vereda La Bella, finca Miranda, 1 830 m, 25 January 1983,Fra/icoetal. 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, 1 August 1960, Tillettei al. 44971 (NY). SURINAM. Nassau Mountains, Marowijne River, forested slopes and summit of plateau A, 430m, 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, 5630'- 634'W. 336'-341'N, 31 July 1963, Maguire et al. 54407 (NY). FRENCH GUIANA. Regina region, E. plateau of MontagueTorte, 1 1 km WNW of Approvague River, 200-450 m, 5222'W, 418'N, 17 June 1988, Feuillet et al. 10178 (NY); Mt. Tortue, 1 1 km WNW of Approvague river, along the road, 200-450 m, 5222'W, 418'N, 16 June 1988, Feuillet et al. 10230 (NY); Saul, Mont Galbao, 17 October 1984, de Foresta 656 (NY); pente NE des Monts Galbao, 10 km au SW de Saul, 500-600 m, 1 1 March 1975, de Granville 2374 (MO, NY); ancienne piste de Saul a Belizon, entre Eau Claire et St. Eloi, 21 August 1981, de Granville 4944 (MO); Saul, trace ORSTROM vers les monts Galbao, sur la Montagne Liane, 19 July 1976, de Granville B5339 (MO); Haul Camopi - Mont Belvedere. 7 December 1984, 52 S. KNAPP AND T. HELGASON Fig. 14 Distribution of 5. anceps (circles) and S. angustlalatum (star in circle). de Granville 7 165 (NY); Montagne Bellevue de 1'Inini, ext. SW versant NW, 550 m, 15 August 1985, de Granville 7502 (NY, US); Montagne Bellevue de 1'Inini, zone centrale, 700-750 m, 20 August 1985, de Granville 7686 (NY); Mont Galbao, secteur E, 600 m, 5317'W, 336'N, 15 January 1986, de Granvilleet al. 8704 (NY); Camp 4, Monpe Soula-Bassin du Hoaut-Marouini, 5 km a 1'Oest, 180 m, 5404'W, 239'N, 3 September 1987, de Granvilleel al. 9975 (NY); MontAtachi Bacca- region de F Inini, centre du plateau sommital, campIV,780m,53 55'W,333'N,21 January 1989,o7809(F,MO).Heredia: Rio Vueltas (upper Rio Patria) on E. slope of Volcan Barba on the Carribean side, 1900 m, 8404'W, 1006'N, 1 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 Sarapiquf, along Q. El Sura between arboretum and station, 9 March 1980, Hammel79^ (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, 1615m, February 1 938, Skutch 36 1 4 (A, K, MO, NY); Cerro de las Caricias, N. of San Isidro, 2000-2400 m, 1 1 March 1926, Standley & Valeria 51987 (F); along the W. fork of the upper Rio Para Blanco beyond the road terminus ofCalleZurqui, 18 March 1974, Utle y & Utley 70 1 (F).Limon: Cordillera de Talamanca, Atlantic slope, canyon of the Rio Sim, 1800-1900 m, 8259'W, 913'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, 8448'W, 1018'N, 18 March 1973, Gentry & Bitrger212\ (F, MO); on and near the Continental Divide about 2-5 kmE. & SE of Monteverde, 1580-1700 m, 1018'N,84 46'W, 17March 1913, Gentry & Burger 217>\ (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 Jose: Along the Rio Para Blanca (Pacific drainage), Cerros de Zuruqui, 1600-1800 m, 1003'N, 8401'W, 6 February 1977, Burger et al. 10250 (F); Cordillera de Talamanca; Chirripo massif, Pacific slope, place along trail known as Abra, 2500m, 2 April 1969, Davidse & Pohl 1529 (F); near Rio Hondura, 1 150 m, 8359'W, 1004'N, 12 August 1972, Lent 2788 (F); vicinity of El 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, 1 500- 1 800 m, 26 December 1 925, Standley & Valeria 44055 (F); Alto de la Palma on Finca Porvenir, c. 5 km N. of San Jeronimo, 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: Robalo 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); Chiriquf border along Continental Divide on Carretera del Oleoductoc. 1 km N. of Quebrada Arena, IHRE Fortuna Hydroelectric project, 1 150 m, 846'N, 8212'W, 1 1 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 Rfos, 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 Fortuna Dam site, 10. 1 miles NW of Los Planes de Hornito, 1250 m, 845'N, 8217'W, 10 April 1980, Croat 50031 (MO); edge of Laguna de Volcan, 9 August 1 972, D 'Arcy & D 'Arcy 6606 (GH, MO); road from Nueva California and Rfo Serano c. 9 miles from Rio Chiriqui Viejo, 1370 m, 7 April 1979, D 'Arcy et al. 1 2988 (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 1 977, Folsom & Collins 1 765 (MO); slope of hill above camp at Fortuna Dam site, 1400-1500 m, 14September 1977, Fo/sometal. 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, 2225m, 13 April 1979, Hammel etz\. 7030 (MO); 1 km N. of Fortuna Lake, 1200 m, 8213'W, 845'N, 3 March 1985, Hampshire & Whitefoord 286 (BM); c. 0.5 km E. of Cerro Pate Macho, headwaters of Rio Palo Alto, 1 800- 2100m,82 2rW,847'N, 1 2 November 1981, Knapp el a\. 2108 (BM, NY); trail to Cerro Pate Macho, headwaters of Rio Palo Alto, above Palo Alto, 1700-2100 m, 8222'W, 847'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). Code: 2 miles N. of Cerro Pilon, 900 m, 16 March 1973, Leisnerl24 (MO, F). Veraguas: E. side of mountain (CerroTute)W. of Escuela (Primer Basica, formerly Agricola) Alto dePiedra, c. 5 miles NW of Santa Fe, 760-850 m, 10 September 1982, D'Arcy 15003 (MO). VENEZUELA, sin. loc., November 1875, Andre 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, 6742'W, 102i;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, Pare Nacional, 1000 m, 1 December 1938, Williams & Alston 139 (BM, NY), Williams 10743 (F). Carabobo:Along Rfo San Gian, al S. de Borburata, arriba de la Plant Electrica, entre Los Tanques y La Toma, 750 m, 27 March 1966, Steyermark & Steyermark 95 1 6 1 (F). Distrito Federal: ?Galipan, 1 846, Linden 1 28 (G); Bosque de Catuche, above Caracas, 1200-1800 m, 9 May 1913, Pittier6\45 (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, 1 500 m, December 1 939, Williams 1 3624 (F). Falcon: Arriba de La Chapa, Sierra de San Luis, 1100m, 18 January 1979, Flora Falcon 2 10 (WIS); Sierra San Luis, ridges around Hotel Parador, c. 1 km S. of Curimagua, 1300-1 350 m,6935'W, 1 PIO'N, 28 September \9U, Knapp & Mallet 6685 (BH, K, MY, VEN); Distrito Bolivar, Sierra de San Luis, Cerro Galicia, around TV antenna at summit, 1500 m, 1 11 1'N, 6942'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 Werff6514 (WIS). Miranda: Quebrada de las Comadres, near las Mostazas, 1 100 m, November 1924, Allart 255 (NY); Campo Experimental Padron - Estacion Experimental de Caucagua, 15 km al E. de Caucagua, 40 m, 22 January 1 976, Berry 1 926 (F); Dtto Paz Castillo, Municipo Reyes Cueta, Los Guayabitos, 1300-1490 m, 1021'16"N, 66 C 38'36"W, 11 December 1987, Castillo & Bocaranda F. 2694 (MO); Quebrada de las Comadres near las Mostazas, 1 100 m, November 1924, Pittier 255 (G); Parque Nacional de Guatopo, headwaters of Rfo Grande, from Quebrada San Antonio to Fila de REVISION OF SOLANUM SECTION PTERO1DEA 67 5 Fig. 22 5. uleanum. Habit: eft Ma//e/ 6524 (K), Schunke V. 3898 (MO). Fruit: Schunke V. 543 1 (MO). Rio Grande, between Santa Teresa and Alatagracio de Orituco, 6.5 km from Rancheria Mi Querencia, 600-700 m, 27 November 1 96 1 , Sleyermark 90 1 05 (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, 13July 1 972, Dwmonfetal. 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, 6241'W, 1040'N, 17 October 1984, Knapp & Mallet 677 1 (BH, F, K, MY, US, YEN); Peninsula de Paria, a lo largo de la Quebrada Nivardo, afluente de Rio Caverna, afluente de Rio Oscuro arriba de Mundo 68 S. KNAPP AND T. HELGASON Nuevo W. de Cerrode Humo, 700-750 m, 7 August 1966, Steyermark & Rabe 96140 (B, NY, US); Peninsula de Paria. cloud forest in tributary headwaters of Rfo Cumana, SW of Cerro de Humo, vicinity of Manacal, 15 km (by air) NW of Irapa, 800 m. 6239'W, 1041'N, 29 November 1979, Steyermark & Liesner 120698 (NY). Solanum trizygum is superficially similar to 5. conicum of the eastern Amazon, and is very closely related to that species (see Fig. 8). It differs from 5. conicum in its more lanceolate leaflets that are very shortly petiolate or sessile, and its more elongate fruit. The fruit of S. trizvgum also resembles that of 5. 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 5. trizygum, occurring in Central America and in the cordillera de la Costa in Venezuela, is quite common in angiosperms (see Knapp, 1 99 1 b) . 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, 1991/7 for a discussion). 10. Solanum uleanum Bitter in Reprium nov. Spec. Regni veg. 12: 139, pi. I (1913). Type: Brazil, Acre, Rio Acre, Porto Carlos, February 1911, Ule s.n. (B-syntype, destroyed; Gl-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 van 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; KMectotype). Solanum uleanum var. gracilescens Bitter in Reprium nov. Spec. Regni veg. 12: 141 (191 3). 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. 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 k'mi' (Vickers 143 - remedy for diarrhoea, plant crushed and mixed with water [Cofan]), 'ahi inta Yko' ( 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 NacionalYasuni, Pozo petrolero Daimi 2, 200 m, 76 1 1 'W, 0055'S, 26 May 1 988, Ceron & Hurtado 4057 (MO, NY, QCNE); Carretera Hollm-Loreto-Coca, km 40, entre Rio Guamani y Rio Pucuno, 1 200 m, 7700'W, 0040'S, 1 1 December 1987, Neill et al. 8 107 (MO, NY), Ceron 2931 (MO, NY), Palacios 2219 (MO); Canton Tena, Rio Blanco community, headwaters of Rio Huambuno, 6 km NNW of Ahuano, 440 m, 7740'W, 0100'S, 19 February 1990, Kohn 1102 (QCNE). Pastaza: Rio Chico, affluent of Rio Pastaza, village of Rio Chico and vicinity, c. 10 km S. of Puyo, 3 km S. of Tarqui, 1000 m, 7755'W, 0103'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, 7621'W, 0015'S, 4 August 1981, Brandbyge et al. 32965 (AAU). PERU. Amazonas: Pongo de Manseriche, Rio Santiago & Rio Maranon, c. 7730'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, 7525'W, 940'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 o 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. 543 1 (MO, WIS).Pasco: Oxapampa, km 28 Reparticion-Iscozacfn (km 86 Villa Rica-Iscozacfn-Pto. Mairo), Rio La Raya near Ameusha com- munity of Laguna, 350 m, 7510'W, 1020'S, 22 August 1984, Knapp & Mallet 6654 (K, NY, US, USM); Oxapampa, trail from Rio Iscozacin to Ameusha community of Villa America, Rio Palcazu Valley, 340 m, 7515'W, 1012'S, 31 August 1984, Knapp & Salick6669 (K, US, USM); Oxapampa, km 15 of Palcazu road, (km 73 Villa Rica-Iscozaciin-Pt. Mairo) along Rio Palcazu, 380 m, 7510'W, 1021'S, 17 August 1984, Knapp & Mallet 6645 (BH, US, USM). San Martin: San Martfn, km 28 of Tarapoto-Yurimaguas road, 650 m, 7615'W, 625'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, 627'S, 23 September 1986, Knapp & A///e/8394 (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 Institute Agropecuario de REVISION OF SOLANUM SECTION PTERO1DEA 69 Fig. 23 Distribution of 5. 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, 7355'W, 802'S, 24 June 1987, Gentry & Diaz 58484 (MO, USM). Solarium 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 5. conicum and 5. mite (both of which are sympatric with 5. 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 5. 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 Veil, (synonym of 5. caavurana Veil. = section Geminata (G. Don) Walp.) 2. Solanum lacteum Veil, (affinities and identification unknown, no type specimen exists and plate in Vellozo resembles no known species of Solanum) 3. Solanum laurinum Dunal (synonym of 5. 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. Nunez of QCNE and P. Wilson for help and support in Ecuador; B. Leon, 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 S. KNAPP AND T. HELGASON 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, GJ. 1977. The variation and evolution of Solatium section Basarthrum. II. Britttmia 29: 116-128. Aublet, J.B.C.F. 1775. Histoire des plantes de la Guiane franfoise. 1-4. Paris. Barboza, G.E. & Hunziker, A.T. 1991. Estudios sobre Solanaceae XXXI. Peculiaridades del androceo de interes taxonomico 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. In P.C. Hoch & A.G. Stephenson (Eds), Experimental and molecular approaches to plant biosystematics: 157-172. St Louis, MO. Bitter, G. 1912. XVI. Weitere Poly botryon- fatten. Reprium nov. Spec. Regni veg. 11: 469-473. 1921. XCV. Aufteilung der Gattung Bassovia (im Dunalischen Sinne) zwischen Solanum, Capsicum und Lycianthex. Reprium nov. Spec. Regni veg. 18: 328-335. Bohs, L. 1990. The systematics of Solanum section Allophyllum (Solanaceae). Ann. Mo. hot. 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. hot. Gdn 74: 359-397. Buchmann, S.L. 1983. Buzz pollination in angiosperms. In C.E. Jones & R.J. Little (Eds), Handbook of experimental polination biology: 73-1 13. 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. 1991. Life form and branching within the Solanaceae. In 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. Wixx. 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. hot. Gdn 59: 262-278. 1991. The Solanaceae since 1976, with a review of its bibliography. In 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. Ill & Brown, K.S., Jr. 1987. Ithomiinae (Lepidoptera: Nymphalidae): summary of known larval food plants. Ann. Mo. hot. Gdn 74: 341- 358. Dunal, M.-F. 1813. Histoire naturelle, medicate el economique de Solanum et des genres qui ont etes confundus avec eux. Montpelier. 1816. Solanorum generumque affmium synopsis. Montpelier. 1852. Solanaceae. In A. P. de Candolle (Ed.), Pmdmmus 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. In W.G. D'Arcy & R.C. Keating (Eds), The anther: form, function and phytogeny. 92-1 10. 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. In PL. 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. Greater, W. (Chairman of the Editorial Committee), Barrie, F.R., Burdet, H.M., Chaloner, W.G., Demoulin, V., Hawksworth, D.L., J0rgensen, 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. Konigstein. Hawkes, J.G. 1 990. 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. 1986. 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. In 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. not. Hist. (Bot.) 19: 63-102. 199 la. A revision of the Solanum sessile species group (section Geminata pro parte: Solanaceae). Bot. J. Linn. Soc. 105: 179-210. 1 99 1 b. A cladistic analysis of iheSolanum 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. PI. 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: 1 29- 131. Nelson, G. 1974. Classification as an expression of phylogenetic relationships. Syxt. 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. In W. Punt & G.C.S. Clarke (Eds), The northwest European pollen flora II: 1-30. Amsterdam. Roe, K. 1971. Terminology of hairs in the genus Solanum. Taxon 20: 501-508. Ruiz Lopez, H. & Pavon, 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 mSolanum. In J.G. Hawkes, R.N. Lester & A.K. Skelding (Eds), The biology and taxonomy of the Solanaceae: 307-319. London. & Anderson, GJ. 1982. Hair morphology and relationships of species in Solanum section Basarthrum. PI. Syxt. Evol. 139: 229-256. Soueges, R. 1 907. Developpment et structure du tegument seminal chez les Solanacees. Annls. Sci. not. 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. 1 98 1 . Taxonomy of Solanum section Lasiocarpa. Gentes Herb. 12:41-129. REVISION OF SOLANUM SECTION PTEROIDEA 71 EXSICCATAE Agostini, G. 48 (trizygum ). Allard, H.A. 6236 (mite); 20481 (mite); 20850 (anceps); 22077 (anceps); 221 16 (anceps); 22522 (anceps). Allan, A. 255 (trizygum); 335 (ternatum). Allen, P.M. 4953 (trizygum); 16520 (trizygum). Andre, 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 el 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, CM. 3089 (anceps); 3109 (mite). Bensman, R. 148 (anceps). Berry, P.E. 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. 1 168 (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. & Ceron, M. 4604 (mite). Ceron, C. E. & Hurtado, F. 4057 (uleanum). Ceron, C.E. & Iguago, C. 5430 (anceps). Chacon, LA. 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); 51 156 (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. 61 18 (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. & Beltrdn, 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). Dryer, V.J. 1069 (trizygum); 1 194 (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 Falcon 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. 241 1 (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. 1 165 (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. 1 8876 (anceps); 2 1 1 85 (anceps); 23086 (mite); 29558 (terna- tum); 29790 (anceps); 3087 1 (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). Hading, G. &Andersson, L. 21373 (anceps). Hart,J. 134 (mite). Hurtado, F. 625 (ternatum). Hurtado, F. & Alvarado, A. 1121 (ternatum). Iltis, H.H. & Iltis, CM. 284 (mite). Irwin, H.S. et al. 48077 (anceps). hern, 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); 2581 1 (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. & Alcom, 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 S. KNAPP AND T. HELGASON (anceps); 6452 (conicum); 6456 (conicutn); 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 (tematum); 6629 (anceps); 663 1 (conicum); 6632 (mite); 6639 (anceps); 6644 (anceps); 6645 (uleanum); 6654 (uleanum); 6655 (anceps); 6658 (anceps); 6664 (conicum); 6685 (trizygum); 6771 (trizygum); 6852 (trizygum); 693 1 (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. 1 102 (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. etal. 13646 (conicum); 13738 (anceps); 13771 (ternatum); 13898 (anceps). Liberman, M. 262 (anceps). Lindeman, J.C. 535 (anceps). Linden, J.J. 1 28 (trizygum). Linhart, Y.B. 155 (trizygum). Lleras, E. et al. PI 7286 (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. PI 2838 (mite); PI 2903 (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); 381 19 (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). Nunez, P. 5770 (anceps); 6473 (mite). Nunez, P. & Phillips, O. 10464 (anceps). Nunez, P. et al. 10555 (mite). 0llgaard, B. 74539 (savanillense); 74630 (savanillense); 74954 (anceps); 98451 (mite). Ollgaard, 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). Plowman, T. & Ramirez R., M. 11212 (mite). Plowman, T. & Schunke V, J. 7394 (ternatum); 1 1509 (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). Prevost, M.F. 304 (anceps). Prevost, 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 Castaheda, 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, F.A. 1487 (anceps). Silverstone-Sopkin, F.A. & 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. & Vdsquez, R. 4899 (anceps). Smith, D.N. et al. 1 184 (anceps); 1569 (ternatum); 7933 (ternatum). Smith, H.H. 1 162 (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. 1 1681 (ternatum). Soukup,J. 22 10 (mite). Spichiger, R. & Encarnacion, F. 8440 (mite). Spruce, R. 3882 (mite); 4377 (anceps); 4385 (ternatum); 4432 (chamaepolybotryon); 4462 (uleanum); 4849 (angustialatum). Standley, PC. 68905 (trizygum); 8671 1 (trizygum); 90563 (trizygum). Standley, PC. & Valeria, 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. & Morton, O.B. 9536 (mite). Sullivan, G. & Young, K. 1 154 (ternatum). REVISION OF SOLANUM SECTION PTEROIDEA 73 Tessmann, G. 3890 (uleanum). Tillett, 5.5. 673-226 (conicum). Tillett, 5.5. et al. 44971 (anceps). Timand, M. & Astete, H. 692 (ternatum). Tirado, G. et al. 189 (anceps). Tyson, E.L. 7144 (trizygum). Vie, E. s.n. (anceps); 2608 (anceps); 5201 (mite); 5490 (anceps); 6276 (anceps); 6922 (mite); 973 1 (mite). Utley, J. & Utley, K. 701 (trizygum); 2902 (trizygum). Vdsquez, R. 2243 (anceps); 3876 (anceps). Vdsquez, R. & Jaramillo, N. 2584 (anceps); 3499 (anceps); 4475 (anceps); 5097 (anceps); 5471 (anceps); 6370 (mite); 8287 (mite); 8362 (mite); 8680 (anceps); 10533 (anceps); 1 1699 (mite). Vdsquez, R. et al. 2151 (anceps); 6559 (anceps); 1 1923 (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. 1 1400 (anceps). Werff, H. van deret al. 8281 (anceps); 8357 (conicum); 10219 (anceps); 13 122 (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, LI. 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. 21 A (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 Solarium alatibaccatum Bitter 55 Solanum aligerum Schldl. 37 Solatium anceps Ruiz & Pav. 50*, 51, 52* Solanum angustialatum Bitter 50*, 52*, 54 Solanum apiculatibaccatum Bitter 58 Solanum aubletii Pulle 5 1 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 Veil. 69 Solanum dendrophilum Bitter 46 Solanum diffusum Ruiz & Pav. 45 Solanum diffusum subsp. miozygum Bitter 46 Solanum diffusum var. 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 Veil. 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 var. procerius Bitter 5 1 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 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. Volume 2 No. 1 New Himalayan species of Pedicularis with special reference to those from the eastern Himalaya. P.C. Tsoong. 1955. Pp. 1- 34. Facsimile edition. 4.35 No. 2 Mosses of Dominica, British West Indies. E.B. Bartram. 1955. Pp. 37^*9. Mosses of the Ecuadorian Andes collected by P.R. Bell. E.B. Bartram. 1955. Pp. 51-64. Facsimile edition. 4.00 No. 3 Novitates Himalaicae-1 . F. Ludlow & W.T. Stearn. 1956. Pp. 65-8 1 , 8 plates, 1 1 figs. Facsimile edition. 5.25 No. 4 Saxifraga of the Himalaya 1 . Section Kabschia. H. Smith. 1958. Pp. 83-129, 14 figs. 9.65 No. 5 The Polypodiaceae and Grammitidaceae of Ceylon. W.A. Sledge. 1960. Pp. 131-158, 4 figs. Facsimile edition. 3.90 No. 6 Allium and Milula in the central and eastern Himalaya. W.T. Stearn. 1960. Pp. 159-191, 4 plates, 10 figs. Facsimile edition. 4.35 No. 7 The identity of hopyrum aquilegioides. L.G. de Beer & W.T. Stearn. 1960. Pp. 193-202, 3 figs. Facsimile edition. 3.15 No. 8 On the geographical relationships of the angiosperm flora of New Guinea. R. Good. 1960. Pp. 203-226, 1 fig. Facsimile edition. 3.60 No. 9 Saxifraga of the Himalaya II. Some new species. H. Smith. 1960. Pp. 227-260. 9 plates, 17 figs. Facsimile edition. 5.65 No. 10 New species of Taraxacum from the Himalayan region. J.L. van Soest. 1961. Pp. 261-273, 8 plates. Facsimile edition. 6.15 No. 1 1 The athyriod ferns of Ceylon. W.A. Sledge. 1962. Pp. 275-323. 3 plates. Facsimile edition. 5.25 No. 12 The genus Epilobium in the Himalayan region. PH. Raven. 1962. Pp. 325-382, 7 plates, 13 figs. Facsimile edition. 5.90 Volume 3 No. 1 A revision of the genera Buchenavia and Ramatuella. A. W. Exell & C.A. Stace. 1963. Pp. 1^46, 5 figs. Facsimile edition. 5.15 No. 2 The diatom genus Capartogramma and the identity of Schiiostauron. R. Ross. 1963. Pp. 47-92, 2 plates, 38 figs. Facsimile edition. 5.15 No. 3 Angiosperms of the Cambridge Annobon Island Expedition. A.W. Exel. 1963. Pp. 93-1 18, 10 plates. Facsimile edition. 3.75 No. 4 A revision of the genus Petrorhagia. P.W. Ball & V.H. Hey wood. 1964. Pp. 1 19-172, 3 plates, 22 figs. Facsimile edition. 4.90 No. 5 Marine algae of Gough Island. Y.M. Chamberlain. 1965. Pp. 1 73-232, 4 plates, 80 figs. 12.50 No. 6 The Ceylon species of Asplenium. W.A. Sledge. 1965. Pp. 233- 277, 1 plate, 3 figs. Facsimile edition. 5.15 Volume 4 No. 1 Cuticular studies as an aid to plant taxonomy. C.A. Stace. 1965. Pp. 1-78, 5 plates, 10 figs. Facsimile edition. 7.20 No. 2 The genus Elaphoglossum in the Indian peninsula and Ceylon. W.A. Sledge. 1967. Pp. 79-96. Facsimile edition. 3.25 No. 3 No. 4 No. 5 No. 6 No. 7 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. Tome, and Annobon). A.W. Exell. 1973. Pp. 325^11. 12.50 Volume 5 No. 1 The dryopteroid ferns of Ceylon. W.A. Sledge. 1973. Pp. 4 figs. 6.90 No. 2 New Himalayan and Tibetan species of Corydalis (Papaveraceae). F. Ludlow & W.T. Stearn. 1975. Pp. 45-69, 15 plates, 14 figs. 8.40 No. 3 The marine algae of Trinidad, West Indies. W.D. Richardson. 1975. Pp. 71-143, 12 plates, 2 figs. 13.45 No. 4 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 No. 5 Frank Ludlow ( 1 885-1972) 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 No. 6 Studies in the genus Hypericum L. (Guttiferae). 1 . Infrageneric classification. N.K.B. Robson. 1977. Pp. 291-355, 9 figs. 14.20 No. 7 Sphagnales of tropical Asia. A.Eddy. 1977. Pp. 357^45, 4 plates, 17 maps, 25 figs. 17.80 Volume 6 No. 1 The handwriting of Joseph Banks, his scientific staff and amanuenses. J.B. Marshall. 1978. Pp. 1-85, 62 figs. 18.30 No. 2 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, 1 fig. 24.40 No. 3 The lichenicolous Hyphomycetes. D.L. Hawksworth. 1979. Pp. 183-300, 47 figs. 24.40 No. 4 The species of Chisocheton (Meliaceae). D.J. Mabberley. 1979. Pp. 301-386, 3 plates, 10 figs. 24.40 Volume 7 No. 1 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 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, 1 fig. 5.40 No. 3 A revision of the genus Anacyclus L. (Compositae: Anthemideae). C.J. Humphries. 1979. Pp. 83-142, 27 figs. 15.60 Volume 8 No. 1 The Thelypteridaceae of Ceylon. W.A. Sledge. Pp. 1-54, 5 figs. 1981. 15.15 No. 2 Studies in the genus Hypericum L. (Guttiferae) 2. Characters of the genus. N.K.B. Robson. 1981. Pp. 55-226, 73 figs. 33.55 No. 3 A revision of the lichen family Thelotremataceae in Sri Lanka. M.E. Hale, Jr. 1981. Pp. 227-332, 20 figs. 24.80 No. 4 Vascular plant collections from the Tristan da Cunha group of islands. E.W. Groves. Pp. 333^20, 33 figs. 21.40 Volume 9 No. 1 The lichenicolous Coelomycetes. D.L. Hawksworth. 1981. Pp. 1-98, 36 figs. 22.70 No. 2 The genus Callithamnion (Rhodophyta: Ceramiaceae) in the British Isles. PS. Dixon & J.H. Price. 1981. Pp. 99-141,5 figs. 12.50 No. 3 Parmelia subgenus Amphigymnia (lichens) in East Africa. H. Krog & T.F.V. Swinscow. 1981. Pp. 143-231, 31 figs. 21.05 No. 4 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 Volume 10 No. 1 Taxonomic studies in the Labiatae tribe Pogostemoneae. J.R. Press. 1982. Pp. 1-89, 33 figs. 21.85 No. 2 The typification of Hudson's algae: a taxonomic and nomenclatural reappraisal. L.M. Irvine & P.S. Dixon. 1982. Pp. 91-105. 5.40 No. 3 Seaweeds of the Faroes. Various authors. 1982. Pp. 107-225, 13 figs. 27.15 No. 4 The lichen genus Steinera. A.M. Henssen & P.W. James. 1982. Pp. 227-256, 24 figs. 9.70 Volume 11 No. 1 The algae of Lightfoot's Flora scotica. P.S. Dixon. 1983. Pp. 1-15, 2 figs. 5.55 No. 2 A taxonomic study of the lichen genus Micarea in Europe. B.J. Coppins. 1983. Pp. 17-214, 57 figs, 28 maps. 37.75 No. 3 The hepatics of Sierra Leone and Ghana. E.W. Jones & A.J. Harrington. 1983. Pp. 215-289, 8 figs. 18.15 No. 4 Studies in the Corallinaceae with special reference to Fosliella and Pneophyllum in the British Isles. Y.M. Chamberlain. 1983. Pp. 291-^63, 89 figs. 33.45 Volume 12 No. 1 A revision of the Morinaceae (Magnoliophyta-Dipsacales). M.J. Cannon & J.F.M. Cannon. 1984. Pp. 1-35, 9 figs. 11.40 No. 2 An introduction to fern genera of the Indian subcontinent. C.R. Fraser-Jenkins. 1984. Pp. 37-76, 1 fig. 12.50 No. 3 A revision of African Sphagnales. A. Eddy. 1985. Pp. 77-162, 47 figs. 23.05 No. 4 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. 1 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 & PA. Sims. 1985. Pp. 277-381, 33 plates. 28.75 Volume 14 No. 1 Cytological observations on Indian subcontinent and Chinese Dryopteris and Polystichum (Pteridophyta: Dryopteridaceae). M. Gibby. 1985. Pp. 1^2, 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. 1 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. 1 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. 269-31 1, 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, 1 fig. 25.10 No. 4 Some Cretaceous and Palaeogene Trinacria (diatom) species. PA. 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 Corydalis (Papaveraceae: Fumarioideae) in Nepal. M. Liden. 1 989. Pp. 479-538, 26 figs. 19.35 Studies on the Cretan flora 2. The Dianthusjuniperinus complex (Caryophyllaceae). N.J. Turland. 1992. Pp. 165-169. 41.25 Volume 19 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- 6 1,36 figs. A revision of the Solanum nitidum group (section Holophylla pro pane): Solanaceae. S. Knapp. 1989. Pp. 63-102, 21 figs. Six new species of Solanum sect. Geminata from South America. S. Knapp. 1989. Pp. 103-1 12, 8 figs. The application of names of some Indian species of Ocimum and Geniosporum (Labiatae). J.R. Press & V.V. Sivarajan. 1989. Pp. 11 3- 11 6, 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. 1 17-158, 41 figs. Facsimile edition. 52.45 Volume 20 No. 1 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29. Brathys (part 2) and 30. Trigynobrathys. N.K.B. Robson. 1990. Pp. 1-15 1 , 22 figs, 46 maps. Facsimile edition. 49.25 No. 2 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 Volume 21 No. 1 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 No. 2 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 Volume 22 No. 1 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 No. 2 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 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. 1 fig. 43.25 No. 2 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 Volume 24 No. 1 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. 35^8. 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. 9 1-100. 43.25 No. 2 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. 101-1 14. Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical species (excluding Australia). D.J. Galloway. Pp. 1 15-160. 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 Volume 25 No. 1 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 No. 2 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 Alpine's De plantis exoticis, with special emphasis on the flora of Crete. N.J. Turland. Pp. 127-159. Book review. M.G. Gilbert. P. 1 6 1 . 43.40 Volume 26 No. 1 A morphological study of Chaetoceros species (Bacillariophyta) from the plankton of the Pacific ocean of Mexico. D.U. Hemandez-Becerril. 1996. Pp. 1-73. 43.40 No. 2 Studies in the genus Hypericum L. (Guttiferae) 6. Sections 20. Myriandra to 28. Elodes. N.K.B. Robson. 1996. Pp. 75-217. 43.40 CONTENTS 11 31 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 LT. 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 Bulletin of The Nat BOTANY SERIES Vol.27, No. 1, June 1997