A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae)

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M.Sc. Thesis - Royal Botanic Garden Edinburgh & University of Edinburgh, UK. 2005.

Text of A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae)

  • 1. A REVISION OF NEPALESE LINDERNIA All.(SCROPHULARIACEAE S.L.) Heide Maria BadenA thesis presented in partial fulfilment of the degree of M.Sc. The Biodiversity & Taxonomy of PlantsRoyal Botanic Garden Edinburgh& University of Edinburgh 2005

2. ABSTRACTThe Nepalese species of Lindernia (Scrophulariaceae s.l.) were revised. These are at least 13distinct species with possibly two additional species. For species delimitations, vegetativeand sexual characters, in particular leaf, flower, and fruit morphology, were taken intoaccount. Numerical taxonomy included leaf length to leaf width ratio and pedicel length toleaf length ratio. Measurements made allowed a definition of inflorescence type, animportant distinguishing character within Lindernia. An SEM study of Lindernia seedmaterial was undertaken. Results did not support the traditional groupings into subgeneraproposed by earlier taxonomists. Taxonomic history of the Nepalese species is summarized,the taxonomic problems discussed, and an account of the genus for the Flora of Nepal isincluded.ii 3. CONTENTSTitle pageiAbstractiiContentsiiiTables and FiguresviList of Abbreviations viii1. INTRODUCTION 12. AIMS AND OBJECTIVES33. TAXONOMIC HISTORY3 3.1. Linnaeus, Allioni, Philcox, and the type species of Lindernia 3 3.2. Other first species descriptions6Burman7Rhumphius 7Van Rheede7Retzius 7Colsman 7Willdenow 8Roxburgh8Vahl8Maximovicz9 3.3. Subgeneric treatments Goodbye to Gratiola 10Rafinesque11Link & Otto 11Blume 12Reichenbach 12 iii 4. Sprengel 13 Buchanan-Hamilton13 Don (D.) 14 Chamisso 14 Bentham14 G. Don 16 Dalziel & Gibson 16 Hooker 16 Haines 17 Alston 17 Pennell17 Borbs 18 Von Mueller19 Urban19 Von Wettstein19 Merrill20 Boldingh 20 Backer 213.4. Recent floristic accounts21 Mukerjee 21 Sivarajan & Mathew 21 Banjeri21 Philcox21 Khan & Hassan22 Diaz Miranda 22 iv 5. Cramer22Yamazaki23Quail Lewis 234. MATERIALS AND METHODS23 4.1. Morphological Characters25Duration and Habit25Roots 25Hairs and other structures25Stems 27Leaves28Nodes 31Inflorescence 31Flowering time32Calyx 33Corolla 34Androecium34Gynoecium 38Fruits38Seeds 405. RESULTS AND DISCUSSION 42 5.1 Scanning electron microscopy of seeds43Size and shape43Colour44Longitudinal and latitudinal alveolate rows 44Appendiculation of testa45 v 6. Testal pattern465.1.1 SEM results and Yamazakis recent subgenera 59 5.2 Taxonomic Problems 635.2.1 Identity of the type species of the genus 63 5.2.1.1 Identity of the Asian form of the type species 655.2.2 Vandellia erecta and Lindernia parviflora 68 .5.2.3 The identity of L. indet-a735.2.4 Lindernia indet-b from Dr. Buchanan 775.2.5 The type of L. micrantha D. Don 785.2.6 The type of Lindernia oppositifolia (Retz.) Mukerjee796. SUMMARY807. REFERENCES 818. APPENDICES Appendix (1): Account of the genus for the Flora of Nepal88 Key to the Species 89 Species descriptions 90 Appendix (2): List of species described in Asia109 Appendix (3): Benthams subgenera from DC X111 Appendix (4): Krockers Flora Silesiaca, p. 398400112 Appendix (5): SEM micrographs of stomata on lamina 115 Appendix (6): CD-ROM with morphology matrices, herbarium specimen, and digital images of selected literature including Roxburghs Icones. Use for research purposes only. vi 7. LIST OF FIGURES AND TABLESFig. 1. a d. Specimens and Illustrations of L. procumbens (Krock.) Philcox 5Fig. 2. a b. Roxburghs illustrations of Gratiola parviflora Roxb. 9Fig. 4. 1 a b. Roots.25Fig. 4. 2 a h. Hairs.26Fig. 4. 3 a d. Stems.28Fig. 4. 4 a g. Lamina shapes.30Fig. 4. 5 a c. Nodes.31Fig. 4. 6 a d. Inflorescence types.32Fig. 4. 7 a e. Calyx types.33Fig. 4. 8 a b. Pressed Corollas. 34Fig. 4. 8 c h. Corollas. 36Fig. 4. 9 a f. Floral dissections. 37Fig. 4. 10. a k. Capsules. 38Tab. 5 Seed and seed-surface characters from SEM study.47Fig. 5. 1 a p. SEM micrographs of entire seeds.48Fig. 5.2 A P. SEM micrographs of seed coats. 53Fig. 5.2 + . SEM micrographs of appendages. 58Fig. 5. 3 Classification of Lindernia based on Yamazaki (1981) 59Fig. 5. 4 a + b. Linnaean specimens 796.364Fig. 5. 4 c g. Wallich specimens seen by Bentham.70Fig. 5. 5 Graph: Manner of inflorescence.75Fig. 5. 6 Second unknown specimen from Nepal 76Fig. 5. 7 Buchanan-Hamilton specimen 77Fig. 5. 8 a. Lectotype of V. angustifolia Benth. 78Fig. 5. 8 b + c. L. oppositifolia (Retz.) Mukerjee 78vii 8. LIST OF ABBREVIATIONSBBotanic Garden and Botanical Museum Berlin-DahlemBM British Museum-Herbarium in the Museum of Natural HistoryBUCBuchanan-Hamilton abbreviated in SEMBuch.-Ham. Buchanan-Hamilton abbreviated in Herbarium CollectionsCMuseum Botanicum Hauniense, CopenhagenDNEP Edinburgh expedition codeERoyal Botanic Garden, EdinburghEMAK,Edinburgh expedition codeICBN International Code of Botanical NomenclatureINGIndex Nominum GenericorumKRoyal Botanic Gardens, KewK-WHonourable East India Company (Wallich) HerbariumLD Botaniska Museet LundLD Nationaal Herbarium LeidenLINN Linnaean Society of LondonNORNorkettPSWPolunin, Sykes & WilliamsSStaintons.l. sensu latos.str. sensu strictoSSWStainton, Sykes & WilliamsTI Tokyo Instituteviii 9. 1. INTRODUCTIONLindernia was named in 1766 by Allioni after von Lindern (16821755), who firstillustrated what we now know as Lindernia pyxidaria L. in Tournefortius Alsaticus (1728).In this Miscellanea Taurinensis: Stirpium aliquot descriptiones cum duorum generumconstitutione, Allioni did not explicitly include a specific epithet when describing thisspecies from Alsace-Lorraine in France, but cited a specimen collected by Gagnebin inSwitzerland, and provided an illustration (Fig. 1. b; p. 5). In 1965, Philcox concluded thatL. pyxidaria was illegitimate. He also decided that it was conspecific with Anagalloidesprocumbens Krocker (Fig. 1. d; p. 5) and he described the new combination L. procumbens(Krock.) Philcox, which he designated as the type species. This replacement has not beenfollowed through by Index Nominum Genericorum, which still maintains Linderniapyxidaria All. as the (problematic) first species descripion of the genus.With ca. 100 species, Lindernia was placed in the Scrophulariaceae and is distinguishedfrom other genera in the Scrophulariaceae sensu lato by the following charactercombination: (1) absence of bracteoles, (2) presence of a 5-angular, relativelyactinomorphic calyx, (3) a conspicuously 2-lipped corolla, and (4) a persistent septum inthe bisepticidally dehiscent capsule (Pennell 1943a, Deyuan et. al. 1998, Lewis 2000, Mill2001). The group was treated in its widest sense by Pennell, who in 1935 moved the generaVandellia L., Bonnaya Link & Otto, and Ilysanthes Raf. into the genus Lindernia.In Lindernia, the largest species diversity occurs in both tropical and subtropical Asia (ca.60 spp.) and Africa (ca. 40 spp.). For Southeast Asia ca. 47 species have been recorded.Philcox reported 23 from Malesia, Pennell reported 10 for Western Himalaya, Papua, NewGuinea, Deyuan treated 29 for China; Mill treated 10 for Bhutan, Yamazaki treated 30 forThailand, 47 for Indochina, Japan, Taiwan. Bentham treated 27 for India (excluding thesynonymous ones, as far as my research went). 1Recently, the Scrophulariaceae have been subject to extensive phylogenetic analysis(Albach et al. 2005, Oxelman et al. 2005), and the family Linderniaceae (Rahmanzadeh et1Appendix 2; the synonymy has not been checked.1 10. al. 2004) was proposed to encompass Lindernia together with 12 other genera includingTorenia L. and Craterostigma Hochst. With ca. 100 species, Lindernia is the largest genusof the new family. This study split the traditional tribe Gratiolae 2, created by Bentham in1835, into two families: the Linderniaceae and the Gratiolaceae. Early classification ofLindernia spp (Linnaeus 1771; Willdenow 1779; Retzius 1786; Colsmann 1793; Vahl1804; Roxburgh 1819) stressed links with the genus Gratiola. 3It is impossible to judge the monophyly of Lindernia as of yet; the estimated phylogeniesso far (Rahmanzadeh et al. 2004) suggest Lindernia s.l. and Lindernia s.str 4. Thenomenclature of this group is of yet an unfinished puzzle, especially the complexsurrounding the type species of the genus. These questions should be sorted out before, orin conjunction with more phylogenetic studies, for the latter to be of value to ourunderstanding of the evolution of Lindernia.Medicinal properties of Lindernia have long been studied, with references to vires medicasnihil certi (Allioni 1765), and Usus latet (Krocker 1790) reflecting the connection betweenhumans and this plant. Miyase et. al. (1995) have isolated the oleanane saponinsLinderniosides A and B from Lindernia pyxidaria. Saponins exhibit important detergent orsurfactant, as well as antiviral properties important to medicine (Simes et al. 1999). Inaddition, herbicide resistance in Lindernia is subject to weed management research(Hamamura et al. 2003, Uchino & Watanabe 2002). Including this ethnobotanicalbiochemical aspect into our study of Asian Lindernia [as has been done by Diaz Mirandafor South America (1977)] could provide supplementary clues to clearing up questions ofaffinity within this group.2Rahmanzadeh et al. explain the morphological basis of this tribe, distinguished most importantly by itsovule and seed anatomy: Intermediate layers of ovule integument are 13, endothelial cells are large,transversally elongated, arranged in 68 longitudinal rows, and thickened only towards the endosperm.;the endosperm of mature seeds is smooth or furrowed, the seeds have longitudinal ridges, and the testacells show hook-like wall thickenings (2005). Compare with results of seed anatomy in Chapter 5.3To this day, good potential exists to find Lindernia herbarium specimens erroneously curated inGratiola genus folders.4Because only few taxa of Lindernia were sampled during this study (only six), it is probably too early toassign species into the Lindernia s.str. and s.l. We might in fact be dealing with sever