A REVISION OF NEPALESE LINDERNIA All.        (SCROPHULARIACEAE S.L.)                     Heide Maria BadenA thesis present...
ABSTRACTThe Nepalese species of Lindernia (Scrophulariaceae s.l.) were revised. These are at least 13distinct species with...
CONTENTSTitle page                                                                  iAbstract                             ...
Sprengel                       13       Buchanan-Hamilton              13       Don (D.)                       14       Ch...
Cramer                                        22            Yamazaki                                      23            Qu...
Testal pattern                                                            46            5.1.1 SEM results and Yamazaki’s r...
LIST OF FIGURES AND TABLESFig. 1. a – d.   Specimens and Illustrations of L. procumbens (Krock.) Philcox   5Fig. 2. a – b....
LIST OF ABBREVIATIONSB            Botanic Garden and Botanical Museum Berlin-DahlemBM           British Museum-Herbarium i...
1. INTRODUCTIONLindernia was named in 1766 by Allioni after von Lindern (1682—1755), who firstillustrated what we now know...
al. 2004) was proposed to encompass Lindernia together with 12 other genera includingTorenia L. and Craterostigma Hochst. ...
2. AIMS AND OBJECTIVESThe goals of this study were to (1) check the taxonomy of Nepalese species, and to revise itwhere ne...
described in Systema Vegetabilium of 1759], and (3) von Lindern’s illustration inTournefortius alsaticus (1728), in which ...
Fig. 1 a: Collections from Herb. Linn. 796.1; mixedsheet of L. dubia L. on left; L. virginiana L. on the        Fig. 1 b: ...
Unfortunately, it has not been possible to verify the types of all the species listed in thesynonymy of the Nepalese speci...
L. antipoda (L.) Alston. See the entry for Alston (1931) on page 14 for a word of cautionconcerning his new combination.Bu...
name G. cordifolia Colsm. even survived Pennell, who failed to synonymise it with his L.anagallis (Burm.f.) Pennell. It ha...
Figure 2 a: Roxburgh’s illustration of Gratiola   Figure 2 b: Roxburgh’s illustration of G. parviflora Roxb. parviflora Ro...
Species Plantarum 17 of 1753. His publication dealt with the errors that occurred in theinterpretation of this species ide...
species. The genera created were to be used for the next century or so, until Pennellsuccessfully argued to have all of th...
but the first description may possibly be of an earlier date (e.g., 1820; there is confusion inthe literature over abbrevi...
grounds of ‘filaments bifid at base or appendaged, and from whence the fertile filamentsare arched over to the upper lip w...
Don (D.) was the herbarium curator and librarian of Lambert. He was “comissioned [byLambert] with the approval of Hamilton...
In Scrophulariae Indicae, Bentham used the genera Vandellia L., Bonnaya Link & Otto(“filaments always abortive, simple, an...
G. Don in his General System (1838) explained that the name Gratiola comes from gratia,grace; on account of the supposed m...
taken in section 5. 2. 2, where the differences between B. parviflora and V. erecta areexplored.Haines in The Botany of Bi...
draw. In Lindernia, the calyx is usually more or less deeply lobed, i.e. the sepals areseparate, whereas in Torenia, the c...
Von Mueller, in Fragmenta Phytographiae Australiae (1882), listed Gratiola pedunculata(Br. Pr. 435) 13 as dispersed throug...
anterior filaments were described as “very small and bumpy [höckerig].” It is here that hementioned L. pyxidaria All. as f...
species, I chose to favour Thunberg’s basionym as the earlier one. During this research, Ireferred to Boldingh’s new combi...
conspecific. In 1968, Philcox revised the Malesian species Lindernia, giving a reference forcytologic work that has been c...
anagallis auctt. (non Burm.f. 1768). Further he commented that “the Lindernia-Ilysanthescomplex, however, does not still s...
Species descriptions and keys were exclusively based on Nepalese material where possible,and missing material was substitu...
4.1. MORPHOLOGICAL CHARACTERSDuration and Habit. The Nepalese species of Lindernia are annual herbs. L. hookeri haspreviou...
of structures (esp. lamina and calyx), and which usually do not exceed a length of 0.1 mm[Fig. 4.2 b], but seldom reach 0....
Fig. 4. 2 e: L. hookeri (C.B.Clarke) Wettst.Lamina margin serrate and ciliate; usually           Fig. 4. 2 f: L. hookeri (...
slender plants; L. anagallis and L. indet-a are more stout. Features such as hairs and colour are most obvious on the angl...
Occasionally, the presence or absence of the petiole has been used as a character, but ingeneral it is not useful as basal...
Fig. 4. 4 b: L. indet-a.Fig. 4. 4 a: L. micrantha (D.Don) Wettst.       Three to five-veined from the base. Lamina sub-Lam...
Nodes. The nodes of the stem in Lindernia are very much influenced by the manner ofattenuation or petiolation of the oppos...
as intermediate to, or same as vegetative part of the plant. The axillary, solitary type oftenproduces only one pedicel pe...
Calyx. The calyx consists of five lobes usually of slightly unequal length. Linderniacrustacea forms one exception in that...
Corolla. The corollae in Lindernia are bilaterally symmetric, semi-closed, and bilabiate.The corolla of L. indet-a is clos...
complex, (1) all 4 stamens bear fertile anthers (e.g., L. crustacea, L. pusilla, L. micrantha,Fig. 4. 9 a, c, f ). In the ...
Fig. 4. 8 c: L. anagallis (Burm.f.) Pennell   Fig. 4. 8 d: L. parviflora (Roxb.) MukerjeeDNEP 2B 191                      ...
Fig. 4. 9 a: Floral dissection of L. crustacea (L.)      Fig. 4. 9 b: Floral dissection of L.F. Muell.; WIL 326           ...
Gynoecium. The gynoecium is bi-carpellate, and the placentation axile. The style ispenicilliform, simple, but often flatte...
Fig. 4. 10 c: L. multiflora (Roxb.) Mukerjee   Fig. 4. 10 d: L. nummulariifolia (D.Don) Wettst.Fig. 4. 10 e: L. anagallis ...
Fig. 4. 10 i: L. antipoda (L.) Alston                 Fig. 4. 10 j: L. parviflora (Roxb.) Mukerjee            Fig. 10 k: L...
on seed characters. The first group is characterized by alveoli, exhibited also byCraterostigma Hochst., parts of Artanema...
5. RESULTS AND DISCUSSIONThe account for the genus in Nepal is attached in the appendix (1). This chapter deals firstwith ...
L. procumbens?’ throughout. A separate, post-SEM study was conducted, comparing seedsof L. parviflora with what is now hyp...
densely rugose appendiculation leaves the shape underneath open for speculation. Here, alatitudinal dissection of the seed...
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).
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A Revision of Nepalese Lindernia All. (Scrophulariaceae s.l./ Linderniaceae).

  1. 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 Plants Royal Botanic Garden Edinburgh & University of Edinburgh 2005
  2. 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. 3. CONTENTSTitle page iAbstract iiContents iiiTables and Figures viList of Abbreviations viii1. INTRODUCTION 12. AIMS AND OBJECTIVES 33. TAXONOMIC HISTORY 3 3.1. Linnaeus, Allioni, Philcox, and the type species of Lindernia 3 3.2. Other first species descriptions 6 Burman 7 Rhumphius 7 Van Rheede 7 Retzius 7 Colsman 7 Willdenow 8 Roxburgh 8 Vahl 8 Maximovicz 9 3.3. Subgeneric treatments – Goodbye to Gratiola 10 Rafinesque 11 Link & Otto 11 Blume 12 Reichenbach 12 iii
  4. 4. Sprengel 13 Buchanan-Hamilton 13 Don (D.) 14 Chamisso 14 Bentham 14 G. Don 16 Dalziel & Gibson 16 Hooker 16 Haines 17 Alston 17 Pennell 17 Borbás 18 Von Mueller 19 Urban 19 Von Wettstein 19 Merrill 20 Boldingh 20 Backer 213.4. Recent floristic accounts 21 Mukerjee 21 Sivarajan & Mathew 21 Banjeri 21 Philcox 21 Khan & Hassan 22 Diaz Miranda 22 iv
  5. 5. Cramer 22 Yamazaki 23 Quail Lewis 234. MATERIALS AND METHODS 23 4.1. Morphological Characters 25 Duration and Habit 25 Roots 25 Hairs and other structures 25 Stems 27 Leaves 28 Nodes 31 Inflorescence 31 Flowering time 32 Calyx 33 Corolla 34 Androecium 34 Gynoecium 38 Fruits 38 Seeds 405. RESULTS AND DISCUSSION 42 5.1 Scanning electron microscopy of seeds 43 Size and shape 43 Colour 44 Longitudinal and latitudinal alveolate rows 44 Appendiculation of testa 45 v
  6. 6. Testal pattern 46 5.1.1 SEM results and Yamazaki’s recent subgenera 59 5.2 Taxonomic Problems 63 5.2.1 Identity of the type species of the genus 63 5.2.1.1 Identity of the Asian form of the type species 65 5.2.2 Vandellia erecta and Lindernia parviflora 68 . 5.2.3 The identity of L. indet-a 73 5.2.4 Lindernia indet-b from “Dr. Buchanan” 77 5.2.5 The type of L. micrantha D. Don 78 5.2.6 The type of Lindernia oppositifolia (Retz.) Mukerjee 796. SUMMARY 807. REFERENCES 818. APPENDICES Appendix (1): Account of the genus for the Flora of Nepal 88 Key to the Species 89 Species descriptions 90 Appendix (2): List of species described in Asia 109 Appendix (3): Bentham’s subgenera from DC X 111 Appendix (4): Krocker’s Flora Silesiaca, p. 398—400 112 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 Roxburgh’s Icones. Use for research purposes only. vi
  7. 7. LIST OF FIGURES AND TABLESFig. 1. a – d. Specimens and Illustrations of L. procumbens (Krock.) Philcox 5Fig. 2. a – b. Roxburgh’s 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 α + β. SEM micrographs of appendages. 58Fig. 5. 3 Classification of Lindernia based on Yamazaki (1981) 59Fig. 5. 4 a + b. Linnaean specimens 796.3 64Fig. 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 78 vii
  8. 8. LIST OF ABBREVIATIONSB Botanic Garden and Botanical Museum Berlin-DahlemBM British Museum-Herbarium in the Museum of Natural HistoryBUC Buchanan-Hamilton abbreviated in SEMBuch.-Ham. Buchanan-Hamilton abbreviated in Herbarium CollectionsC Museum Botanicum Hauniense, CopenhagenDNEP Edinburgh expedition codeE Royal Botanic Garden, EdinburghEMAK, Edinburgh expedition codeICBN International Code of Botanical NomenclatureING Index Nominum GenericorumK Royal Botanic Gardens, KewK-W Honourable East India Company (Wallich) HerbariumLD Botaniska Museet LundLD Nationaal Herbarium LeidenLINN Linnaean Society of LondonNOR NorkettPSW Polunin, Sykes & WilliamsS Staintons.l. sensu latos.str. sensu strictoSSW Stainton, Sykes & WilliamsTI Tokyo Institute viii
  9. 9. 1. INTRODUCTIONLindernia was named in 1766 by Allioni after von Lindern (1682—1755), 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 et1 Appendix 2; the synonymy has not been checked. 1
  10. 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 (Simŏes et al. 1999). Inaddition, herbicide resistance in Lindernia is subject to weed management research(Hamamura et al. 2003, Uchino & Watanabe 2002). Including this ethnobotanical—biochemical 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.2 Rahmanzadeh et al. explain the morphological basis of this tribe, distinguished most importantly by itsovule and seed anatomy: “Intermediate layers of ovule integument are 1—3, endothelial cells are large,transversally elongated, arranged in 6—8 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 5.3 To this day, good potential exists to find Lindernia herbarium specimens erroneously curated inGratiola genus folders.4 Because 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 several genera withinLindernia s.l.. 2
  11. 11. 2. AIMS AND OBJECTIVESThe goals of this study were to (1) check the taxonomy of Nepalese species, and to revise itwhere necessary, (2) revise the synonymy of these species, (3) test the existingclassification using micromorphology, (4) prepare a draft account of the genus for the Floraof Nepal.3. TAXONOMIC HISTORYThis chapter deals chronologically with all the names and synonyms used for Nepalesespecies, starting with a section on recent controversies around the type species of the genus(3.1.), and proceeding to discuss descriptions of the remaining species (3.2). It thencontinues with a section on the treatments that emphasized subgeneric relationships andtaxonomy within (3.3), and ends with (3.4) a section on the more recent regional floralaccounts from adjacent areas, such as India (Sivarajan & Mathew 1983), India and Burma(Mukherjee 1945), China (Deyuan et al. 1998), Japan and Eastern Asia (Yamazaki 1954—1981), Malaysia (Philcox 1968), and Bhutan (Mill 2001).3.1. Linnaeus, Allioni, Philcox, and the type species of LinderniaLindernia was created by Allioni in his Stirpium aliquot descriptiones cum duorumgenerum constitutione, published in Miscellanea Taurinensis 3, describing a speciesformerly known by the polynomial “Alsinoides paludosa foliis anagallidi similibusflosculia monopetalis rubescentibus capsula oblonga,” and referring to a specimencollected by “Gagnebin,” 5 as well as to an American (Virginian) specimen given to him by“Celeb. Linnaeus” (Allioni 1766). He gave both a species description and the genericdescription of Lindernia All., however, did not include a specific epithet in his descriptionof the genus (merely mentioning that von Lindern had called the taxon pyxidariae), nor inthe accompanying figure. Linnaeus published the binomial Lindernia pyxidaria L. inMantissa Plantarum: 252 (1771). Here he referenced (1) Allioni’s illustration, (2) his ownCapraria gratioloides L. from Species Plantarum: 876 [it is actually thought to be5 In the Harvard University Herbarium - Index of Botanists, Abraham Gagnebin holds record number140348; his collector’s ID is 37350. He is known to have collected in Switzerland in the year of 1763.Collections are at BM and P-HA. 3
  12. 12. described in Systema Vegetabilium of 1759], and (3) von Lindern’s illustration inTournefortius alsaticus (1728), in which the plant is thought to be described as pyxidariae.Linnaeus does not cite Gagnebin’s specimen; instead, he wrote on the back of one of hisown “Pyxidaria Allioni” (see Fig. 1. c. for an image).Recently, it had been argued (Philcox 1965) that this name for the type species of the genusis illegitimate under the International Code of Botanical Nomenclature (ICBN), becauseLinnaeus Capraria gratioloides L. was not a synonym of Lindernia pyxidaria L., but thetype of another species. It is not yet clear to me, whether or not C. gratioloides is indeedsynonymous with Gagnebin’s specimen.The database “Index Nominum Genericorum [ING]” lists Lindernia pyxidaria All. as thetype of the genus. It has been verified here that Allioni does not explicitly make a binomialcombination before Linnaeus in 1771 6. Until it has been proven that the Gagnebinspecimen and Capraria gratioloides are distinct, and until it has been verified thatLinnaeus’ description is indeed illegitimate, I suggest not to change the entry in ING. Ifillegitimacy existed through citing a type of another species, as Philcox argues, then theING entry should be changed. The matter needs further investigation, a part of which willfollow in section 5. 2 under taxonomic problems.The legitimate publication to replace Lindernia pyxidaria was made by Krocker in FloraSilesiaca (1790. Fig. 1. c and Appendix 4), in which he described Anagalloidesprocumbens in as great a detail as Allioni described Lindernia (pyxidaria). In contrast toAllioni, Krocker did not cite a type specimen, but only provided an illustration as its type(Fig. 1 d). The illustrations provided by Allioni and Krocker appear very similar, and thiswas the reason why Philcox chose A. procumbens Krock. as the new basionym for theillegitimate precursor. It was not until Philcox made the new combination in 1965, that L.procumbens (Krock.) Philcox 7 was accepted as the type of the genus in all the accounts Iconsulted during this revision.6 It is possible, however, that Allioni made the combination in an as of yet unpublished work.7 For L. procumbens (Krock) Philcox, the latter includes in synonymy, amongst others, Vandellia erectaBenth. Scroph. Ind. 36 (1835), pro maj parte, excl. Wallich 3943, et 3947 pro parte as well as L. erecta(Benth.) Bonati in Lecomte, Fl. Gén. Indo-Chine 4: 420 (1927). 4
  13. 13. Fig. 1 a: Collections from Herb. Linn. 796.1; mixedsheet of L. dubia L. on left; L. virginiana L. on the Fig. 1 b: Allioni’s illustration from Miscellanearight. Wulfen reference in the illustration not found in Taurinensis 3, T5, F.1 (1766); open flower featuresliterature. 4 fertile stamens.Fig. 1 c: Linnean specimen: Herb. Linn. 796.2. On the reverse sideof this sheet, a hardly legible handwriting describes the specimen(from Latin): Pyxidaria Allioni. Calyx 5-partite, equal in length,persistent, erect. Corolla twice as long as the calyx, monopetalous,bilabiate; corolla tube broadening; upper lip rounded, lower lip 3-lobed, central lobe biggest. Androecium 4 in throat, didynamous,filaments very short. [...] ovate, stigma inflexed, bilabiate. Capsule Fig. 1 d: Anagalloides procumbensovate, bivalved, 2-locular, valves elliptic. Krocker. 1790. Flora Silesiaca 2(1), t. 26. 5
  14. 14. Unfortunately, it has not been possible to verify the types of all the species listed in thesynonymy of the Nepalese species. Therefore, dealing with the type of the genus fromEurope, and a potentially conspecific specimen from Virginia, but using this name forAsian material needs further clarification. For further research, it will be necessary toinvestigate the identities of Lindernia pyxidaria Pursh. non Allioni, Capraria gratioloidesL., L. dubia L., and Gratiola anagallidea Michx. For example, one needs to find outwhether Ilysanthes gratioloides Bentham has Capraria gratioloides L. as its (omitted)basionym. This is important, because Bentham (1846) also says that Ilysanthes gratioloidesBenth. = L. pyxidaria Pursh. non Allioni, which however would only give a pointer to thequestion whether Linnaeus synonymy was indeed wrong. A recently published, extensivework by L.S. Olschki (2004, ed.) entitled Le opere minori di Carlo Allioni. Dal “RariorumPedemontii stirpium” all’ “Auctarium ad Floram Pedemontanam” references severalplates of Allioni’s exsiccata and illustrations, including Capraria gratioloides L. [whichlooks very much like L. procumbens (Krock.) Philcox], and should help to shed furtherlight on the matter. It is available at the Kew library, but again, time was too short to delveany deeper into this important matter.Having discussed the problems concerning the type of the genus, I will now proceed inchronological order to describe the authors as well as major treatments in the taxonomichistory of those species of Lindernia that are represented in Nepal. As for L. procumbens(Krock.) Philcox, voucher specimens of its synonym Vandellia erecta Benth. are in theWallich herbarium at Kew (K). The specimens at Edinburgh (E) lack flowering materialand could not be identified.3.2. Other first species descriptionsLinnaeus had described another Lindernia 8 even earlier than L. pyxidaria, i.e. in hisSpecies Plantarum (1753), but under the name Ruellia antipoda L., (BM!, Herb. Hermanncollection, Sloane room). This publication was legitimate and the taxon is now known as8 During revision of first publications I noticed that some species names were attributed to Linnaeus,whereas the actual author was Willdenow, who edited Sp. Pl. after Linnaeus’ death. Further mistakeswere made by citing Willd. as the authority of a name, although he only cited someone else’s work, e.g.,for Gratiola veronicifolia L. in Sp. Pl. ed. Willd., the true author is A. J. Retzius (but see discussion ofWilldenow on page 6 ahead). 6
  15. 15. L. antipoda (L.) Alston. See the entry for Alston (1931) on page 14 for a word of cautionconcerning his new combination.Burman (1707-1780) described Ruellia anagallis Burm.f. [basionym of Linderniaanagallis (Burm.f.) Pennell] in his Flora Indica: 135 (1768), referring to a Kleinhofspecimen from Java named Anagalloides javanica, and an illustration in Rhumphius’Herbarium Amboinense [(5): 460, t. 170, f. 2; 1747], which Burman co-edited. Thisillustration, however, appears to refer to a different taxon (e.g., L. sessiliflora), since thecalyces are depicted as sessile. However, in his voorwoordt (1695), Rumphius disclaimedaccuracy of the figures, explaining that some were lacking flowering or fruiting material atthe time of preparation. He further referred to van Rheede’s Hortus Malabaricus (1689)admitting that some of the plants depicted therein are not known to “us in this particularway [here in Indonesia], however, that one should not be judgemental for the variation ofplants across geographic boundaries.”Retzius (1742-1821), botanist at Lund, described Gratiola grandiflora and Gratiolaveronicifolia in Fasciculus Observationum Botanicarum 4 (1786), separating them on thebasis of flower size. Both types were collected by König, 9 and both species are nowsynonymised with L. antipoda (L.) Alston [one must be aware of any L. grandiflora nonRetzius, however, which may refer to different taxa]. Further, Retzius is the first todescribe Gratiola oppositifolia, whose authority is often cited as Linnaeus or Willdenow,again citing a König specimen. I mention this taxon here despite the fact that a voucherspecimen has not been recovered for Nepal; however, this species is checklisted in theEnumeration of the Flowering Plants of Nepal (Hara et al. 1981); the misidentified voucherspecimen TI 1488 is actually L. micrantha (D.Don) Wettstein.Colsman worked on a thesis of the Gratiolae, entitled Species Gratiolae a D. König initinere orientali collectae (1793), now referred to as Prodromus Descriptiones Gratiolae.Most importantly he described Gratiola ruellioides Colsm. [= L. ruellioides (Colsm.)Pennell], G. ciliata Colsm. [L. ciliata (Colsm.) Pennell], but also G. cordifolia Colsm. [= L.anagallis (Burm.f.) Pennell], and G. verbenaefolia Colsm. [= L. antipoda (L.) Alston]. The9 König’s collections were described by both Colsman and Retzius, and are housed in Copenhagen andLund, respectively. 7
  16. 16. name G. cordifolia Colsm. even survived Pennell, who failed to synonymise it with his L.anagallis (Burm.f.) Pennell. It has since been synonymized by Cramer (1981) in theRevised handbook for the Flora of Ceylon and by Press and Shresta (2000a), who list underScrophulariaceae Gratiola cordifolia Vahl, D. Don, Prodr. Fl. Nep.: 85 (1824). in Nepaliâ.(Specimen not found). = Lindernia anagallis (Burm.f.) PennellWilldenow updated Linnaeus Sp. Pl. (1797) and created two names, Gratiola pusilla[=Lindernia pusilla (Willd.) Bold.], and G. lucida [= L. crustacea (L.) F.Muell.]. Heomitted the reference to the König specimen made by Retzius when citing L. oppositifoliaRetz., which makes one wonder whether it was not clear at that time (1797) how importanttype specimens would become.Roxburgh (1751—1815) was in India, with short breaks, from 1776—1813 (Sealy 1956).He described Gratiola serrata [Lindernia ciliata (Colsm.) Pennell], G. reptans [L.ruellioides (Colsm.) Pennell], and G. parviflora [L. parviflora (Roxb.) Mukherjee] inPlants of the Coast of Coromandel II (1798) and III (1819). The type for G. parviflora isoften cited as an illustration from Pl. Corom. [sometimes as 203 (Fig. 2 a), or as 204, whichactually depicts G. rotundifolia). While 203 matches his description, his Icones 524 [Fig. 2b] gives a much better depiction of the venation, which is 3—5-veined from the base. Theillustration of an open flower makes the plate very useful compared with the specimenalone. Further, his illustrations of G. serrata (Icones 1515) and G. reptans [Icones 1516]are outstanding; his depiction of L. oppositifolia (Icones 526) is equally excellent, and allcan readily be used in identification (in addition to the König specimens). His research issummarized in Flora Indica of 1820 and Flora Indica ed. Carey of 1832. Roxburgh’s mostin-depth observations give extremely valuable references to the understanding of the AsianLindernia species.Vahl described Gratiola cordifolia Colsman in Enumeratio Plantarum 1:95 (1804) withoutreference to Colsman, but referring to his König type. Hence the species G. cordifolia Vahlreferred to in the literature is a synonym. Both names are now sunk into synonymy withLindernia anagallis (Burm.f.) Pennell. 8
  17. 17. Figure 2 a: Roxburgh’s illustration of Gratiola Figure 2 b: Roxburgh’s illustration of G. parviflora Roxb. parviflora Roxb. Roxburgh’s Icones 524, with detail; notice difference in Pl. Corom. 3: 3, t. 203 (1819); with detail. venation compared with Fig. 2 a.Maximovicz (1827-1891) treated several species in Decas 19 of Diagnoses PlantarumNovarum Japoniae et Mandshuriae. Here he stated that Linnaeus’ Capraria gratioloidesdescribed in Systema Naturae 10 (2): 1117 of 1759 equalled Lindernia dubia L. from 9
  18. 18. Species Plantarum 17 of 1753. His publication dealt with the errors that occurred in theinterpretation of this species identity; this extensive chapter on Lindernia All. promiseshelp to clarify the issues surrounding Allioni’s and Linnaeus’ descriptions. It citednumerous works, which again cited numerous specimens with their European localities ofLindernia pyxidaria L. He then explained “Japoniae dubia civis, nam Miquel (Prol. 356.) habuit tantum specc. fructifera, forsan ad V[andellia] erecta referenda. Planta wolgensis, quam solam examinavit Ledebour (Fl. Ross. III. 225) pariter pertinet ad Vandelliae erectae formam ad Linderniam vergentem, quam supra scripsit. In meridionalibus Asiae, ubi genuina Vandellia erecta crescit, fortassis etiam non deest, sed a collectoribus negligitur, quia, cum forma corollis apertis ante oculos, hanc formae flore clauso praeferunt.”In the above abstract, Maximovicz described the distribution of Lindernia pyxidaria L. inAsia. He stated that the latter, and Vandellia erecta Benth. are not conspecific, but atransition exists between the two (as discussed in Fl. Ross). He further explained that acleistogamous form of L. procumbens probably exists, however, that its small, closedcorolla enticed plant collectors not to collect it, but rather to collect those plants with opencorollas instead. This work elaborated on the occurrence of Lindernia pyxidaria L. andVandellia erecta Benth. in Asia, which have both been sunk into synonymy for L.procumbens (Krock.) Philcox by Philcox. Because the identity of this taxon is not clear andits affinities need to be elucidated, and because the new species from Nepal with a closedcorolla needs a name, Maximovicz’s work on the Asian Lindernia could prove importantduring further research.Maximovicz further treated Ilysanthes hyssopioides Benth. (not reported for Nepal),Bonnaya brachiata Link & Otto [= L. ciliata (Colsm.) Pennell], and synonymized B.veronicaefolia (Retz.) Spreng. with B. verbenaefolia (Retz.) Spreng. [= L. antipoda (L.)Alston].3.3. Subgeneric treatments – Goodbye to GratiolaFollowing the first century of descriptions and treatments, several authors attempted toarrange the Lindernia complex into distinct genera. With increasing material, it hadbecome obvious that many of the species described as Gratiola were too distinct from theother Gratiola spp. Genera had to be found to place these mainly tropical semi-aquatic 10
  19. 19. species. The genera created were to be used for the next century or so, until Pennellsuccessfully argued to have all of them merged into one genus, Lindernia (1935). The thengeneric, now sub-generic distinctions were mainly based on androecium and length of fruit(siliquosae versus brachycarpae), but dehiscence (Bentham 1835) and venation (Hooker1884) were also suggested. For a while, Willdenow’s genus Hornemannia was used forsome short-fruited Lindernia [e.g., L. viscosa (Hornem.) Merr.] by Link and Otto (1820) Irecommend Stearn’s article “The generic name Hornemannia and its diverse applications”for further information on why this genus cannot be used now (1972).Rafinesque described the new genus of Ilysanthes in Annals of Nature 13 (1820), where hemade the important distinction “[...] stamina two fertile under the upper lip; anthersunilocular; two sterile filaments under the lower lip. Ovary oblong; style compressedabove, bilamellate [...]. This genus differs from Gratiola by the calix, corolla, and capsul.The name means mud flower. Habit of Gratiola and Lindernia; leaves opposite sessile,flowers axillary.” He then went on to describe a species (Ilysanthes riparia) from Ohio,where it “grows in the mud or even in the water”. The genus Ilysanthes is spotted easilyamongst a pile of Lindernia specimens; these plants are decisively more slender than therest, and their lamina is distinctly three to five-veined from the base.Link & Otto created Bonnaya in 1828; they described B. brachiata as the type of thegenus [L. ciliata (Colsm.) Pennell], emphasizing among other characters the unique long-aristate margin which itself is framed in white, the narrow bracts subtending the – withheight of the plant shortening – pedicels, and interestingly, described the capsule as almostquadrangular, and the septum as having a green thread running up the middle of either side,to which the seeds are attached. This type is illustrated in Icones Plantarum SelectarumHorti Regii Botanici Berolinensis: 25, t. 11. Here they stated that the seed material wasbrought from Manila by “Herr[n] Dr. von Chamisso.” The material did not withstand theGerman climate but, planted in loose, sandy soil, seeded in the shelter of glasshouses. Theydistinguished this genus from its ally Hornemannia on the basis of capsule, androecium,and calyx. It was named in honour of the keen botanist “Herrn Marquis de Bonnay,”French ambassador to the Royal Prussian Court in Berlin.Link and Otto also published a synonym for our L. crustacea (L.) F. Muell., i.e.,Hornemannia ovata in this illustrated list of selected plants of the Berlin Botanic Garden, 11
  20. 20. but the first description may possibly be of an earlier date (e.g., 1820; there is confusion inthe literature over abbreviations like Icon. Pl. Select. and Icon. Hort. Bot. Berol., which donot refer to the same publication). According to Stearn (1972), this description ofHornemannia ovata has been regarded as a combined description of a generic and specificprotologue by, e.g., Index Kewensis (as was the case in Bonnaya brachiata above), butLink and Otto intended to only add another species to Willdenow’s Hornemannia genus.Blume described the Scrophularinae of Buitenzorg/ Bataviam [Java] in Bijdragen tot deFlora van Nederlandsch Indie(1825) and in it several species of Lindernia important to thisdiscussion. His treatment included Linnaeus’ Gratiola veronicaefolia, and Roxburgh’s G.reptans and G. serrata. He distinguished these Gratiola L. from Diceros Lour., underwhich he included Diceros glanduliferus [a synonym of G. viscosa Hornemann, i.e. L.viscosa (Hornem.) Boldingh. The distinction was based on the corolla being tubular andbilabiate (“labio superiore bilobo aut emarginato; inferiore trilobo”); the stamens havingtwo anthers with divergent locules, and two or three (! – this is the first time a botanistobserved the third, usually lost, anther) sterile anthers in Gratiola Linn.; in contrast, thecorolla being funnel-shaped, (“limbo subbilabiato, interdum subaequali”) and the stamensdidynamous, i.e. all 4 fertile in Diceros Lour. Blume, however, misunderstood Loureiro, asBentham (1846) in DC X points out, Diceros Bl., non Lour.)He further distinguished the genus Mimulus L., in which he divided M. javanicus Bl. [asynonym for L. crustacea (L.) F.Muell]. Distinguishing characters here were the tubularcalyx being 5-dentate (“quinquedentatis”) rather than 5-lobed (“quinquepartitus”), theanther lobes divaricate, and the calyx completely covering the capsule (all these charactersmaking it more akin to Torenia, so Blume says). Lastly, he distinguished Torenia obtusaBl., which is a synonym for L. anagallis (Burm.f.) Pennell. This distinction was supportedbased on the corolla character of “labio superiore obtuso aut retuso, ” and the anthercharacter of “antherarum lobis divaricatis, interdum effoetis. ” He admitted hereafter thatthis genus is hardly distinguishable from Mazus.Reichenbach made the combinations Tittmannia ovata (non Benth.)/ Tittmannia viscosafrom Hornemannia ovata Link & Otto and from Gratiola viscosa Hornemann inIconographia Botanica Exotica (1823/1824), herewith creating a new genus. According toBentham (1835), Reichenbach separated Tittmannia from all the other genera on the 12
  21. 21. grounds of ‘filaments bifid at base or appendaged, and from whence the fertile filamentsare arched over to the upper lip with connivent anthers’ by the short, not plicate calyx, anda globular fruit [which were considered by Chamisso as the true Toreniae (and the oneswith a siliquose fruit and short calyx as the Vandelliae)]. The genus Tittmannia wasmaintained by Bentham for several of his nomina nuda, but Bentham changed its status tosubgeneric level, replacing Vandellia at generic level in later treatments (Scroph. Ind. &DC X). For our purposes, Reichenbach should mainly be kept in mind as being the authorof one T. ovata, whereas the other is Bentham, whose Wallich type forms one basis ofLindernia viscosa. It should also be noted that the basionym reference by Reichenbach toG. viscosa Hornem. Hort. Hafn.: 19 is wrong, (and that to this day I have not seen theprotologue for it).Sprengel made new combinations in Systema Vegetabilium 1:41 (1824) and Syst. Veg. 2:803 (1825), moving Retzius’ Gratiola cordifolia, G. verbenaefolia, and G. veronicifoliainto Bonnaya.Buchanan-Hamilton was the first botanist to collect in Nepal, i.e. 1802—1803. He was aphysician, who “spent 14 months in and around Kathmandu, recording information on allforms of natural resources” (Press & Shresta 2000a). His botanical collection, which hegave to Lambert (in London), formed the basis for Don’s Prodromus Flora Nepalensis (seebelow). He published a few names in Lindernia, but none of them persist as basionyms. 10He moved Willdenow’s Gratiola lucida to Torenia lucida (which is L. crustacea), again amove supported by both Blume, and Chamisso & Schlechtendahl, and also considered byBentham. He then created Torenia alba (already described as Capraria crustacea L.; hencein one year (1831) he gave this taxon two different names, as can be traced in Wallich’sCatalogue. An interesting specimen from Buchanan-Hamilton’s Nepal collection wasfound in the BM general collection. It was later determined as L. nummulariifolia (D.Don)Wettst., but it bears little resemblance to the other specimens from Nepal and mayrepresent a distinct species (see section 5.1.4).10 The reason for a lack of names authored by Buchanan-Hamilton may very well be due to the fact thathe gave a lot of his work to fellow contemporaries (e.g., Don, Wallich), who took over the publicationswithout paying the tribute that we would expect today, and to the fact that a lot of his work remainsunknown as of this day (Fraser-Jenskins, in preparation). 13
  22. 22. Don (D.) was the herbarium curator and librarian of Lambert. He was “comissioned [byLambert] with the approval of Hamilton” (Fraser-Jenkins, in preparation) to write anaccount of the not yet studied Nepalese collections. Including Hamilton’s and Wallich’scollections, but excluding Smith’s herbarium (to which Don did not have access) resultedin this publication being based on ca. 2000 specimens from Nepal (Press & Shresta 2000a).In this Prodromus Florae Nepalensis (1825), Don described L. micrantha and V.nummulariifolia. In this first account for Nepal, Don referred to the genera Lindernia andVandellia in the sense of Linnaeus (Mant. 89) and Jussieu (Gen. 122). [Jussieu, in thisGenera Plantarum (1789) stressed the entire superior lip of the corolla, the bifid stigma,and the capsule unilocular in Vandellia versus Lindernia having a corolla with shortlyemarginate superior lip, an emarginate stigma and a bilocular capsule.] Unfortunately, Donomitted most of the information that came with Buchanan-Hamilton’s detailed collectionnotes; therefore, no specimen localities within Nepal can be traced through D. Don’sProdromus, nor through the Wallich Catalogue. While Don’s publication reads 1825, theactual distribution of the material occurred before the close of the year 1824. This dateshould be used when citing names from this Prodromus Flora Nepalensis (Fraser-Jenkins,in preparation).Chamisso (1781 - 1838) moved Linnaeus’ Capraria crustacea into Torenia crustacea (L.)Cham. & Schlecht. in 1827. This is an interesting move, as Lindernia crustacea is the onlytaxon within the Lindernia complex that has a tubular calyx with short lobes and withseams (remnants of wings?) along the tube of the calyx. The move is supported by Blume,who while not placing it in Torenia per se, at least had placed this taxon into the Toreniae.Bentham (1800-1884) published Scrophulariae Indicae in 1835. Preceding this treatment,in or around 1831 he created several names that, following the Code of BotanicalNomenclature, are nomina nuda; all specimens are in K-Wallich. They include Tittmanniagrandiflora Benth., (K-Wall. Cat. 3949!), now synonymized under L. anagallis (Burm.f.)Pennell; Tittmannia angustifolia Benth., (K-Wall. Cat. 3951!), which had already beendescribed as L. micrantha D.Don; Tittmannia erecta Benth., (K-Wall. Cat. 3947!), now L.procumbens (Krock.) Philcox; and Tittmannia ovata Benth. non Reichenb., (K-Wall. Cat.3942!), now L. viscosa (Hornem.) Bold. 14
  23. 23. In Scrophulariae Indicae, Bentham used the genera Vandellia L., Bonnaya Link & Otto(“filaments always abortive, simple, and club-shaped”) in the tribe Gratiolae to describemost species of this treatment on Nepalese Lindernia, except Gratiola multiflora Roxb. andVandellia hookeri C. B. Clarke. In the beginning pages, elaborating the distinguishingcharacters of all treated Scrophularineae, he commented (100 years ahead of Pennell) that“perhaps it might be better to consider these three genera [Lindernia, Vandellia, andBonnaya] as forming but one”.It was in De Candolle’s Prodromus X (1846), that Bentham first used Lindernia as a genus,herewith adopting the European-based taxon concept as a distinct subtribe in his tribeGratiolae (in 1835 solely encompassing Asian species). He called this subtribe theLindernieae (= at least posterior stamens fertile) 11, and herewith excluded them from thegenus Gratiola on the basis of the latter having a globose, 4-valved capsule. Gratiola wasplaced within the subtribe Eugratioleae (together with, e.g., Mimulus, Mazus, and Bacopa).All our species were represented in the subtribe Lindernieae, in which he included BonnayaLink & Otto, Vandellia L., Lindernia All. (which he knew only from Europe) togetherwith, e.g., Torenia L. (calyx tubular), Curanga Juss. (calyx 4-partite), and Ilysanthes Raf.Vandellia crustacea was again the sole member of the section Torenioides (howeverincluded in Vandellia together with V. multiflora, V. hirsuta (= L. pusilla), V. scabra, andV. glandulifera Bl. (both = viscosa), V. erecta (but L. pyxidaria was excluded into the onlyLindernia). He further listed V. nummulariifolia, V. pedunculata and V. angustifolia.Treated as distinct from these were Ilysanthes gratioloides, I. parviflora, as well asBonnaya brachiata, B. reptans, B. veronici- and verbenaefolia, B. grandiflora, and B.oppositifolia.It becomes quite clear that Bentham worked mainly from herbarium material. Manymultiple entries treat the same taxon; had he had more material at hand, he might haverealized the synonymy between several of his listed species. Still, his comments, especiallythose on the similarities between ‘species,’ are useful. For example, he treated Linderniapyxidaria, but explained that it looks like Vandellia erecta 12, Ilysanthes gratioloides, I.capensis, and I. parviflora, all of which may indeed prove to form a tightly allied complex.11 A cladogram of Bentham’s taxonomic concept as understood from DC X (1846) is attached in theappendix 3.12 See section 5. 2. 2 15
  24. 24. G. Don in his General System (1838) explained that the name Gratiola comes from gratia,grace; on account of the supposed medicinal good qualities. His section on the Gratioleae(Scrophulariaceae) included the genus Bonnaya, where he treated Bonnaya brachiata Link& Otto, B. repens Spreng., B. veronicaefolia Spreng., and B. verbenaefolia Spreng. as partof one complex on account of their long siliquose fruits, and (1) their racemoseinflorescence (most definitely erroneously including B. pusilla auctt.? in this list) – theother part (2) consisting of B. grandiflora (Retz.) Spreng., B. peduncularis Benth., and B.oppositifolia (Retz.) Spreng. with axillary flowers. The second complex was distinguishedby its short fruits, i.e. Brachycarpae, where he included B. parviflora (Roxb.) Benth. (Wall.Cat. 3867; this specimen was lectotypified by Forman (1997).Vandellia was treated much later in the publication. He again separated the Brachycarpaeon account of the capsule being shorter than the calyx ( V. crustacea, V. alba, V. erecta, V.hirsuta, and V. scabra) from those with the capsule being “ovate, a little longer than thecalyx”, here listing V. nummulariifolia D. Don. He then treated the Siliquosae (with thelong pod), including V. diffusa, V. pedunculata, and V. angustifolia, ending with a fewpoorly known species (V. cordifolia, V. roxburghii, and V. multiflora). Lastly he treated thegenus Lindernia, where he included L. pyxidara L.Generally, his treatment closely followed Bentham’s Scrophularineae Indicae of 1835, butincluded interesting facts about the eponomy of the genera. One nomenclatural questionbegs to be resolved as well: G. Don distinguished B. peduncularis and V. pedunculata.These are distinct species and should not be confused.Dalziel & Gibson compiled The Bombay Flora in 1861. These Short descriptions of allindigenous plants included Bonnaya Link & Otto; Ilysanthes, Rafin. and Vandellia L.Hooker in the Flora of British India did not describe any new Nepalese species, but hepointed out that if Roxburgh’s Gratiola lucida was conspecific with Vandellia crustacea,then Roxburgh’s omission of the appendages on the filaments would be faulty. A more in-depth look at his concept of Vandellia erecta Benth.[= Lindernia procumbens (Krock.)Philcox] and Ilysanthes parviflora (Roxb.) Benth. [Lindernia parviflora (Roxb.) Haines] is 16
  25. 25. taken in section 5. 2. 2, where the differences between B. parviflora and V. erecta areexplored.Haines in The Botany of Bihar and Orissa (1922) reduced the four genera to two unitingBonnaya with Vandellia and Ilysanthes with Lindernia based on stamen characters.Alston described his L. antipoda (L.) as synonymous with Ruellia anagallis Burm.f.(instead of non Burm.f.) in his contribution to the Scrophulariaceae of Trimen’s Handbookto the Flora of Ceylon (1931). This mistake in synonymy almost certainly must have had todo with Alston not seeing Burman’s type from Java, as it could not have escaped Alstonthat the Java specimen is synonymous not with R. antipoda, but with the Linderniacordifolia (Colsm.) Merr. that he described in the same account. This speculation was alsomade by Philcox (1968). Alston’s fault was recalled and changed 50 years later by Cramerin A revised handbook to the Flora of Ceylon 3 (1981). In the meantime, wrongdeterminations of herbarium material were frequent.Pennell in his Scrophulariaceae of eastern temperate North America merged all generadescribed above into Lindernia (1935). He justified his decision as follows: “By the union of the four-anthered Lindernia All. and Vandellia L. with the two-anthered Ilysanthes Raf. and Bonnaya Link & Otto is formed a large and clearly natural genus. It is characterized by the remarkably uniform corolla (with narrow posterior lip much shorter than the widely spreading anterior lip), by similar curiously recurving anterior filaments (the proximal portion of each projecting as if it were an appendage and the filament forked, although actually the process is formed by the sharp inbending of the filament), and by similar septicidal dehiscence of the capsule (that nearly always leaves the entire septum persisting as a median plate).”He explained (Schlechter had brought it to his attention) that nothing but the anthercharacter [four anthers in Lindernia and two anthers in Ilysanthes] separates L. pyxidaria ofEurope from I. dubia (L.) Barnh. and I. anagallidea (Michx) Raf. of North America (1936).He concluded that “the loss of the anterior anther represents not a racial distinction so muchas an ultimate stage of evolutionary change, actual kinship being in other characters.” Hetook part in the Archbold expeditions to Papua and New-Guinea, reported in Brittonia 2(1936), and in the Journal of the Arnold Arboretum 20 and 24 (1939 & 1943a). In Brittoniahe argued that Torenia and Lindernia are separate, though a clear distinction is not easy to 17
  26. 26. draw. In Lindernia, the calyx is usually more or less deeply lobed, i.e. the sepals areseparate, whereas in Torenia, the calyx is united, with raised ridges above the five centralveins. However, there are exceptions to the rule, since L. crustacea has united sepals. Asecond distinguishing character is Torenia’s larger corolla with an angular, open throat, andLindernia’s smaller corolla with a flattened throat. The capsule in Torenia is usuallycompletely covered by the calyx, whose apices are connivent, whereas in Lindernia thecapsule usually exceeds the calyx in length, and the calyx lobe apices point outwards. Asfor similarities, Torenia also has a tendency to suppress the anterior stamens. It is that yearof 1936 that Pennell moved Gratiola ruellioides Colsm. to Lindernia ruellioides (Colsm.)Pennell and G. ciliata Colsm. to Lindernia ciliata (Colsm.) Pennell. In 1939, he reported anew species, Lindernia crenata Pennell from this expedition.Pennell did not treat L. anagallis and L. cordifolia as conspecific – he said that L. cordifoliahas anterior filaments which are anther bearing, as opposed to L. anagallis without anterioranthers (maybe that is the case in New Guinea). His description of L. cordifolia hints to amix up in his description in Brittonia 2: 182 (1936) with this one and L. angustifolia(Benth.) Wettst. He also rectified in 1943a his citation of some L. anagallis specimens as L.veronicifolia and L. antipoda in Brittonia and Arn. Arb. 20: 81 (1939).Borbás (1844—1905) has often been referred to as the combiner of Krocker’s L.procumbens, (e.g., by Yamazaki in Flora of Thailand, by Hara et al. in the Enumeration ofthe Flowering Plants of Nepal, and by Mill in Flora of Bhutan). It could not be verified byme that the Hungarian botanist made this new combination. Bekesvarmegye floraja [= Theflora of Bekes county] is cited for the location of the description, but in this to Scottishbotanists slightly obscure Journal [also cited as Magyar Tudomanyos Akademia.Ertekesezek a Termeszettudomanyok Korebol. Kot. 11. szam 18 (1881)], Borbás wroteGratiola officinalis L. [...] Pyxidaria procumbens (Krock.) (Lindernia pyxidaria All.),herewith showing the connection between the genera Pyxidaria, Anagalloides, andLindernia, but not officially combining Lindernia procumbens (Krock.) Borbás. 18
  27. 27. Von Mueller, in Fragmenta Phytographiae Australiae (1882), listed Gratiola pedunculata(Br. Pr. 435) 13 as dispersed throughout the whole of “Australiam extratropicam” but wasnot aware that G. pedunculata had already been described as Ruellia anagallis by Burman.In 1882 he combined Capraria crustacea L. [= Lindernia crustacea (L.) F. Muell.] as wellas G. serrata Roxb. (1793) into Lindernia, not aware that G. ciliata had already beendescribed by Colsman. I am fond of the name pedunculata because L. anagallis has thelongest peduncles of all the species of Lindernia examined in the course of this study.Urban (1884) wrote Studien ueber die Scrophulariaceen-Gattungen Ilysanthes, Bonnaya,Vandellia und Lindernia. Here, he described the corollae, specifically the shape and extentof division of the upper lips, then the nature of the lower lips in great detail. He then wenton to describe the two forms of corollae, the closed kind and the opened kind, whichaccording to Urban can occur on the same plant, i.e., chasmogamy and cleistogamy seemsto depend on geographic location, but added that a closed corolla may have to do withwater-shortage. Von Wettstein and Urban having been contemporary botanists in Berlin,there is little doubt that they conversed on this topic. They seemed to agree that the Asianspecies of Lindernia pyxidaria are more adapted to pollination, while the European speciestend to inbreed. While Urban found anther characters good to distinguish the genera, helater went on to hypothesize sliding scales rather than clear distinctions between the generalisted in his title, and that at least Bonnaya and Ilysanthes should be united.Von Wettstein published three sections of Lindernia for our Nepalese species in Engler &Prantl’s Natürliche Pflanzenfamilien IV (3b) of 1895.Sect I Vandellia L. included Lindernia nummulariifolia (D.Don) Wettst. from theHimalaya [as well as L. sessiliflora (Benth.) Wettst.] as those with clayx 5 toothed, afterflowering 5-parted with L. diffusa (L.) Wettst. from Africa and the Neotropics, as well as L.pedunculata (Benth.) Wettst. [i.e. L. anagallis] and L. angustifolia (Benth.) Wettst. [i.e. L.micrantha D. Don) from East-India to China and Japan etc.Sect. II Eulindernia Wettst. featured Lindernia hirsuta (Benth.) Wettst. [L. viscosa] fromthe whole of Southeast Asia, and L. scabra (Benth.) Wettst. [L. pusilla, in SE-Asia as wellas S. Africa and Madagascar] as those with a capsule not longer than calyx. Appendages on13 I am fond of the name Gratiola pedunculata, because this species has the longest peduncles of allNepalese Lindernia. 19
  28. 28. anterior filaments were described as “very small and bumpy [höckerig].” It is here that hementioned L. pyxidaria All. as flowering mostly cleistogamously in temperate Asia (= L.pyxidaria All. et Aut.), and flowering only chasmogamously (= Vandellia erecta Benth.) inEast-India, and that both forms occur in localities between.Sect. III Hornemannia Link et Otto Calyx 5-toothed. Capsule not longer than calyxincluded L. crustacea (L.) F. Muell., L. molluginoides (Benth.) Wettst. and L. hookeri(C.B.Clarke) Wettst. from East-India. This entry forms one of the cases whereHornemannia was attributed to Link and Otto, whereas its true authority was Willdenow.Merrill’s Species Blanconae (1918) pointed to the identity of a species which hitherto hadbeen sunk into synonymy with L. antipoda: Vandellia grandiflora (Retz.) Merr., whichaccording to Philcox is conspecific with L. philippinensis. Merrill was the first author towrite about the ecology of Lindernia (1912). In these non-systematic Notes on the Flora ofManila with special reference to the introduced element he spoke about the geographicalorigin of species, stating that “in most treatments of tropical floras, whether of the easternor the western hemispere [...] species [such as Oxalis repens Thunb., Sida cordifolia L., andEvolvus alsinoides L., i.e. various grasses and sedges] are usually listed as native ones. It isfar more probable that some have originated in one hemisphere [...] they have beenaccidentally distributed by man within the past 400 years [...]and in most cases it isprobable that their original homes will never definitely be known.” He went on to explainthat “a considerable number are aquatic species or those that grow in swampy places,having minute seeds that might readily be transported by adhering in mud to the feet orfeathers of migratory wading or swimming birds.” A last quote, “while it is only reasonableto suspect that most have been introduced by man (by sticking to clothing, to the hair ofanimals) especially the rice paddy forms, and such genera as [...] Polygala, Salomania,Hydrolea, Lindenbergia, Bacopa, Mazus, Vandellia, Torenia, Bonnaya, Dopatrium,Utricularia owe their presence to natural causes (migratory birds, winds, etc).” Merrill listsVandellia crustacea Benth. as accidentally introduced, and of Oriental or European origin.Boldingh in Zakflora voor de Landbouwstreken op Java (1916) made the combinations forL. pusilla and L. viscosa. Since he omitted the basionyms, it is not clear whether Boldinghreferred to Thunberg’s Selago pusilla (1794) or to Willdenow’s Gratiola pusilla (1797), orto another species altogether. Because the description of S. pusilla did not contradict the 20
  29. 29. species, I chose to favour Thunberg’s basionym as the earlier one. During this research, Ireferred to Boldingh’s new combinations as Lindernia pusilla (Thunb.) Bold. and L.viscosa (Hornem.) Bold. The latter separated the two by their inflorescence type (solitary,axillary versus racemose). It had been argued that Boldingh’s combinations wereillegitimate due to the fact that they were published in a descriptive key; the ICBNhowever, appears to allow publications in descriptive keys pre-dating 1925, henceBoldingh’s combinations can be accepted for the Flora of Nepal.Backer in Onkruidflora der Javasche Suikerrietgronden (1931 – 34) continued Boldingh’swork on Javanese Lindernia. His contributions are summarized in 1968’s Flora of Java III;Scrophulariaceae, and 1973’s Atlas of 220 Weeds of sugar-cane fields in Java (ed. vanSteenis). Here he included several illustrations, maintaining a narrow-leaved form of L.anagallis var. angustifolia.3.4 Recent floristic accountsMukerjee (1945) in his revision of the Indo-Burmese species of Lindernia listed 28 speciesfor this region. It is in this work that Mukerjee made the new combinations Linderniamultiflora (Roxb.) Mukerjee and L. oppositifolia (L.) Mukerjee; the basionym authority ofthe latter should be Retzius, not Linnaeus. In synonymy with L. pyxidaria he citedVandellia erecta Benth. and Gratiola integrifolia Roxb. This revision lacks speciesdescriptions and references to type material. Sivarajan & Mathew revised the genus(1983) with 22 species for India, including species descriptions, which were lacking inMukerjee’s previous treatment.Banjeri (1958) gave a detailed description of his expedition to Eastern Nepal; in his list ofcollections are Lindernia anagallis with light blue flowers at 1220 m (B 1125), and L.ruellioides with light purple flowers at 1527 m (B 1091). This interesting narrative includesa map and description of the flora along his route.Philcox (1963 – 64) lectotypified Ruellia antipoda L. as the type of L. antipoda (L.)Alston, and clarified that Ruellia antipoda L. (Type in Hermann collection) and Gratiolaruellioides Colsm. (Type in Copenhagen) are distinct on account of the projection on theteeth of the lamina. He further found that G. ruelliodes Colsm. and G. reptans Roxb. are 21
  30. 30. conspecific. In 1968, Philcox revised the Malesian species Lindernia, giving a reference forcytologic work that has been carried out for L. crustacea (2n = 42), L. antipoda (2n = 18),and L. viscosa (2n = 42). The last two findings, however, had not been published by thattime. Philcox’s determination slips date among the latest found on herbarium sheets, andseveral herbaria have adopted his revision to arrange the taxa within genus folders (e.g.,Edinburgh, Kew, Leiden). He decided to treat L. anagallis and L. angustifolia Benth. [= L.micrantha] as synonyms, a lumping not sensible for Nepalese material.Khan & Hassan treated Lindernia All. from Bangladesh, and conducted experimentsconcerning pollination and propagation, and concluded that for L. antipoda, L. ciliata, L.crustacea, L. multiflora, L. pusilla and L. viscosa the possibility of cross-pollination isremote, further concluding that capsules can form through self-pollination, and in somecases can produce parthenocarpically.Diaz Miranda (1977) included in his morphological account of Lindernia in SouthAmerica a phytochemical survey of leaf flavonoids research. The characters 14 were foundto mirror the four sections Torenioides, Nummularia, Lindernia, and Brachycarpae. Hetreated L. crustacea, placing it in Pennell’s section Torenioides, secondly L. diffusa (withalmost sessile calyces), thirdly Lindernia procumbens (Krock.) Philcox, synonymizing itwith L. pyxidaria L, Gratiola inundata Kitaibel ex Schultes, and G. integrifolia Roxburgh.He treated L. microcalyx Pennell, Stehle & Quentin, and lastly L. dubia (L.) Pennell, whichhe synonymized with Capraria gratioloides L. and Ilysanthes riparia Rafin. Diaz Mirandaalso mentioned unclarity in synonymy of the Lindernia dubia sensu lato complex.Cramer in A revised handbook to the Flora of Ceylon (1981) described Lindernia as apantropic genus of about 80 species predominantly palaeotropic. He pointed out that untilPhilcox traced Burman’s type of Ruellia anagallis, this taxon had been treated conspecificwith Ruellia antipoda L. With this revision of the Flora of Ceylon, he clarified Alston’scombination of Lindernia antipoda (L.) Alston, which the latter had synonymized with L.14 Combining chemical and morphological data appears to be a useful approach in Lindernia, and should beextended beyond the five taxa treated here. 22
  31. 31. anagallis auctt. (non Burm.f. 1768). Further he commented that “the Lindernia-Ilysanthescomplex, however, does not still seem to be clear and needs further investigation.”Yamazaki initially followed the treatment in Haines’ Botany of Bihar and Orissa and keptVandellia separate fron Lindernia in the Flora of Eastern Himalaya (1966). However, healso sunk Lindernia ruellioides in synonymy with Vandellia antipoda (L.) Yamazaki. Thismove meant a lot of confusion over the identity of these two taxa, which can be seen inherbarium folders to this day (meaning lots of countries may find themselves with a “new”species L. ruellioides, once curation has caught up). By 1977, he adopted Pennell’ssynonymy of genera. His treatments of Lindernia spanned many years of consecutivesections in New and noteworthy plants of Scrophulariaceae from Indo-China, all of whichwere supplied with high-quality line drawings (Yamazaki 1978, 1980, 1983). He wrote theRevision of the Indo-Chinese species of Lindernia All. (Scrophulariaceae) in 1981. A laterchapter will refer to his sub-generic classification of Lindernia.Quail Lewis 15 (2000) revised the New World Species of Lindernia, treating the L. dubiacomplex as four distinct varieties, and treated an overall 12 distinct species. Althoughdealing with New World taxa alone, the treatment is useful for Nepal in its overview of themorphological characters, its chapter of the generic affinities, and its references toglasshouse flowering experiments.4. MATERIALS AND METHODSThe study used of herbarium specimens from Edinburgh (E, some of which included digitalimages of live plants before collection), Kew (K), the Museum of Natural History (BM),Nationaal Herbarium Nederland at Leiden (L). Specimens were also observed at theLinnean Society (LINN). Digital images of type material were obtained from theUniversity of Copenhagen (C), the Botanic Garden and Botanical Museum Berlin-Dahlem(B), and the University of Lund (LD) provided. Protologues were mainly held in the libraryof RBGE, with copies of additional literature obtained from RBG Kew.15 I have included her publication under Lewis in the references in chapter 7, because I have seen otherworks citing her as Lewis 2000. I am not sure whether some of her work appears elsewhere under QuailLewis. 23
  32. 32. Species descriptions and keys were exclusively based on Nepalese material where possible,and missing material was substituted from India or Malaya. A total of 105 herbarium sheetswas included for Nepal, of which at least three specimens per species (data permitting)were measured to cover morphological variation, and hence to achieve representativeranges for each Nepalese species. As mentioned by Davis and Heywood (1973),“classification based primarily on herbarium material should not be pushed too far, unlessfield notes are exceptionally complete, or we have studied the group in the field ourselves.”The lack of information about colours in most species is regrettable, a fault that has begunto change with increased digital imaging of live plants. Colour images of live plants wereused in this section wherever possible. Sixty five characters found useful for identificationwere scored in an Excel spreadsheet, 16 a synthesis of which was used to create adichotomous, artificial 13 couplet-long key to the species. The descriptions were limited toca. 150 words, again focussing on characters aiding identification.A Leo Supra 55VP Scanning electron microscope (SEM) was employed to investigate andcompare seed morphology of all thirteen taxa. Additionally, a pilot study to examinedifferences in lamina structure was carried out. Since the herbarium specimens werealready dry, the untreated seeds were mounted on Agar Scientific Carbon tabs andsputtercoated with platinum in a peltier cooled EMITECH K 575X. One advantage of nothydrating and critical-point drying the seed material was that characteristcs actually changewhen hydrating the seeds, because the epidermis may obscure the alveoli (Juan et al. 1997).An obvious drawback is the convolution of some seeds, which does not allow for accuratemeasurement of seed dimensions.The results were used to investigate the value of SEM study for species delimitation, aswell as their coherence with subgeneric delimitations established in previous literature onthis taxonomic group.16 Appendix (6; CD-ROM). 24
  33. 33. 4.1. MORPHOLOGICAL CHARACTERSDuration and Habit. The Nepalese species of Lindernia are annual herbs. L. hookeri haspreviously been described as a perennial herb in, e.g., the Flora of Bhutan, but no evidencefor this was found in Nepalese material. D.Don describes L. micrantha as a perennial (signinterpreted in Stearn), but again the material available in this study did not exhibit thischaracter. The species of Lindernia are either erect, spreading, creeping, or prostrate. Theirheight generally ranges from 4 to 25, seldom to 30 cm.Roots. Root types in this genus vary from fibrous (Fig. 4.1 a) as in Lindernia indet-a toshort and slender (Fig. 4.1 b) as in most other species. Some decumbent species alwaysroot at the nodes (e.g., L. ruellioides) while others do not exhibit this character at all. Fig. 4.1 a: Roots of Lindernia a. Fig. 4.1 b: Roots of L. procumbensHairs and other structures. In this study, the use of this character was mainly based onpresence and absence. Only a few species of Lindernia are hairy. However, on glabrousplants, particular parts exhibit small, short teeth, which ‘embroider’ the margins and apices 25
  34. 34. of structures (esp. lamina and calyx), and which usually do not exceed a length of 0.1 mm[Fig. 4.2 b], but seldom reach 0.2 mm (e.g., on young leaves of L. nummulariifolia). Otherliterature has referred to this character as ciliate (Lewis 2000), and I adopt this term here,but caution not to confuse this character with the conspicuous, long-aristate margin in L.ciliata, which itself is smooth and not ciliate despite its epithet.Usually, all species exhibiting the character lamina hairy have a hairy calyx. Further, thestem can be hairy but the lamina be glabrous (except when young). This is the case in L.nummulariifolia; here also, length of scabous structures on lamina margin is inverselyproportional to leaf size. Lastly, care should be taken when defining the state of the stem: Itmay appear densely hairy at the nodes, but in fact perceived hairiness at the nodes is almostalways an increased hairiness of the decurrent lamina base margin. ciliate margin ▲ Fig. 4. 2 a: L. micrantha (D.Don) Wettst. Fig. 4. 2 b: L.antipoda.(L.) Alston Lamina margin distantly, obscurely serrate, and Lamina margin serrate and ciliate. Not to be smooth. confused with c., L. ciliata, which is not ciliate. Also visible here are the sunken stomata. Fig. 4. 2 c: L. ciliata (Colsm.) Pennell Fig. 4. 2 d: L. parviflora (Roxb.) Mukerjee Lamina margin long-aristate serrate. Lamina margin entire to obscurely 2—3 dentate, and smooth. 26
  35. 35. Fig. 4. 2 e: L. hookeri (C.B.Clarke) Wettst.Lamina margin serrate and ciliate; usually Fig. 4. 2 f: L. hookeri (C.B.Clarke) Wettst.bearing a longer protrusion on the projecting part Lamina base is often densely beset with long hairs, whereas the apex has none [see 4. 1 e]. Thisof the margin. phenomenon is shared by a few Nepalese species. Calyx (inset) also strongly hairy. ciliate margin ▲Fig. 4. 2 g: L. anagallis (Burm.f.) PennellLamina margin serrate, but ciliate often only onthe projecting part of the margin. A further Fig. 4. 2 h: L. ruellioides (Colsm.) Pennellcharacter exhibited by many species of Lindernia Lamina margin deeply serrate, and ciliate.(see also b + f) is the red colour of the margin.Stems. All stems are quadrangular, and are winged to some extent. The most extremeexamples of a winged stem can be found in L. hookeri (Fig. 4. 3 c), L. pusilla (Fig. 4. 3a), and L. multiflora. Stem diameters were measured near the base of the plant as thewidth of one of the four sides of the quadrangle; they ranged between 0.5—2 mm. Thetaxa often referred to as of the subgenus Ilysanthes (i.e. L. parviflora) are the most 27
  36. 36. slender plants; L. anagallis and L. indet-a are more stout. Features such as hairs and colour are most obvious on the angles of the stem. Fig. 4. 3 a: L. pusilla (Thunb.) Bold. Stem strongly winged. Also visible are the Fig. 4. 3 b: L. anagallis (Burm.f.) Pennell flattened hairs attached to the angles. Stem winged., not hairy. Also visible is the reddening characteristic of this taxon. Fig. 4. 3 d: L. parviflora (Roxb.) Mukerjee Stem winged, not hairy. Fig. 4. 3 c: L. hookeri (C.B.Clarke) Wettst. Stem winged, sparsely hairy on angles.Leaves. The leaves in Lindernia are highly variable in shape (base, margin, apex) as well asvenation. They can be almost circular (L. nummulariifolia, L. pusilla) or linear and verynarrow (L. micrantha), almost rhombic (L. indet-a), or lanceolate (L. hookeri), and shapecan even vary within an individual plant. As Roxburgh (1832) pointed out, the shape of theleaves in L. ruellioides varies between round on non-flowering and oval on floweringshoots on the stoloniferous plants. 28
  37. 37. Occasionally, the presence or absence of the petiole has been used as a character, but ingeneral it is not useful as basal leaves may be petiolate and upper leaves sessile.Furthermore, definition of petiole length may be arbitrary: by the time the lamina of L.micrantha reaches the stem, it is so narrow that it is impossible to define the base of thelamina and the beginning of a petiole. A more important character state to distinguish, e.g.,L. micrantha (Fig. 4. 4 a) from L. anagallis (Fig. 4. 5 b) is the lamina length: width ratio. Inthe Nepalese species, the ratio ranges between 1 and 12, but is relatively constant (+/- 1)within one species.Lamina margin of Nepalese Lindernia ranges from entire-undulate (Fig. 4. 4 b + d) tomoderately (Fig. 4. 4 a + c) to sharply serrate (Fig. 4. 4 e + f). It is long-aristate in L. ciliata(Fig. 4. 4 e), but not aristate in L. ruellioides (Fig. 4. 4 f). It has been found useful inseveral cases to count the number of incisions per side of the lamina. The numbers aresurprisingly constant, although care must be taken when counting the number for L.ruellioides – as mentioned above, leaf shape, and hence number of serrations per side of thelamina varies, but a solution to this was adopted in the key.The lamina can be three to five-veined from the base (Fig. 4. 4 b + d), uni-veined (Fig. 4. 4a + e), or pinnately veined (Fig. 4. 4 c, f + g). Just like the angles of the stem, veins tend tobe coloured reddish in a few species. 29
  38. 38. Fig. 4. 4 b: L. indet-a.Fig. 4. 4 a: L. micrantha (D.Don) Wettst. Three to five-veined from the base. Lamina sub-Lamina obscurely serrate. Leaf length to leaf entire to undulate.width ratio 7:1. Petiole not easy to define.Fig. 4. 4 c: L. crustacea (L.) F.Muell. Fig. 4. 4 d: L. parviflora (Roxb.) Mukerjee Lamina 3-veined from the base. Sunken stomata visible. Margin obscurely dentate near apex.Fig. 4. 4 e: L. ciliata (Colsm.) Pennell Fig. 4. 4 f: L. ruellioides (Colsm.) PennellLamina uni-veined 30
  39. 39. Nodes. The nodes of the stem in Lindernia are very much influenced by the manner ofattenuation or petiolation of the opposite leaves. As mentioned above, the hairiness oftenassociated with the base margins of ciliate laminas tend to make the nodes appear hairy; theorigin of the hair, however, is the leaf and not the stem. Increased hairiness in these areasseems to be associated with the fusion of tissue.Fig. 4. 5 a: Lamina base and node of Fig. 4. 5 c: Lamina base and nodeL. muliflora (Roxb.) Mukerjee of L. anagallis (Burm.f.) Pennell Fig. 4. 5 b: L. anagallis (Burm.f.) PennellInflorescence. Two main types of inflorescence were recorded: (A) About half the specieshave solitary flowers in the axils of the leaves. These are frequently supported by pedicelsas long, sometimes much longer than the leaves. Included in the definition of axillary,solitary flowers are those taxa, whose leaves gradually decrease in size toward the apex ofthe plant. These taxa, namely Lindernia parviflora and L. procumbens may appearsubracemose at times. The important distinction between “flowers in the axils of leaves”and a “terminal, racemose inflorescence” (Fig. 4. 6 a) is the gradual decrease in size of theleaves toward the apex of the plant, as well as the maintenance of their shape (Figs. 4. 6 b,c + d). In contrast, (B) a terminal raceme has flowers subtended by bracts (Fig. 4. 6 a)These bracts are never exactly the same shape as the leaves, except possibly wheresubtending the first pair of pedicels. Instead, they are small and shaped differently than theleaves on the lower, vegetative part of the plant; the terminal raceme was defined as thepresence of a marked, abrupt change in leaf size from the vegetative to the flowering partof the plant, neglecting the location of the first flowering internode, which often classified 31
  40. 40. as intermediate to, or same as vegetative part of the plant. The axillary, solitary type oftenproduces only one pedicel per node, and sometimes a shoot out of the opposite axil (4.6 cand d).Flowering time. Flowering times were estimated using the dates of material from Nepal. Itis possible that the flowering season may be slightly longer. The season spans roughly fromMarch to November. Fig. 4. 6 a: Terminal raceme of L. multiflora Fig. 4. 6 b: Solitary, axillary inflorescence of L. pusilla Pedicels slightly reflexed. Fig. 4. 6 c: Solitary, axillary inflorescence of L. Fig. 4. 6 d: Solitary, axillary inflorescence of L. anagallis parviflora 32
  41. 41. Calyx. The calyx consists of five lobes usually of slightly unequal length. Linderniacrustacea forms one exception in that its calyx is dentate rather than lobed (Fig. 4.7 a); L.hookeri forms a second exception, in that its calyx is bilabiate (Fig. 4.7 b). Usually,however, the calyx is deeply lobed (Fig. 4.7 c). Venation is either single, or three to five-veined from the base of the lobe. All calyces are persistent. Hairs are usually longest on thecalyx, but glabrous taxa have glabrous calyces. If the calyx itself is glabrous, its calyx lobemargins can be ciliate. The distinction between hairy (hairs longer than 0.3 mm; Fig. 4.7 e)and ciliate (hairs ca. 0.1 mm or less) as discussed above is important, because calyces mayappear hairy due to ciliate margins (Fig. 4.7 d). Fig. 4. 7 a: 5-dentate calyx of L. Fig. 4. 7 c: 5-partite calyx of Fig. 4. 7 b: bilabiate calyx of L. L.antipoda crustacea hookeri Fig. 4. 7 d: Glabrous calyx with Fig. 4. 7 e: Hairy calyx without ciliate margin on apex of lobe (L. ciliate margin (L.pusilla) anagallis) 33
  42. 42. Corolla. The corollae in Lindernia are bilaterally symmetric, semi-closed, and bilabiate.The corolla of L. indet-a is closed, revealing cleistogamy. While accurate measurementscan only be made from exquisitely well-pressed herbarium material (Diaz Miranda 1977),the numbers referred to in this study can be used as guidance to live material. The lower(anterior) lip is always in three parts, and wider (ca. 3—8 mm) than the upper (posterior)lip, which in size approximates one of the three lower lip lobes (Figs. 4. 8 e—g). The upperlip is shallowly 2- lobed, or not lobed at all, and ca. 2—4 mm. Corolla length ranges from 4to 12 mm. The central lower lip lobe length can reach to 4 mm. Corolla colours range fromwhite to yellowish to blue to purple, but are never red because they are pollinated by smallinsects, probably bees, not birds or bats. Comparison of the corollas indicates the closerelationship between L. ruellioides and L. antipoda. Their similarities can be acknowledgedby comparison of well-pressed specimens (Figs. 4. 8 a + b below). Fig. 4. 8 a: L. antipoda Fig. 4. 8 b: L. ruellioidesAndroecium.All species of Lindernia are didynamous (Judd & Olmstead 2004), i.e. the length of theanterior pair of filaments is distinct from that of the posterior pair. Within the genus,however, three general distinctions mark a more or less stable subgeneric difference. In one 34
  43. 43. complex, (1) all 4 stamens bear fertile anthers (e.g., L. crustacea, L. pusilla, L. micrantha,Fig. 4. 9 a, c, f ). In the next, (2) only the posterior pair bears fertile anthers, whereas theanterior pair bears sterile ones (e.g., L. hookeri Fig. 4. 9 e). The last case features (3) aposterior pair with fertile anthers, but the anterior pair reduced to staminodes (e.g., L.ruellioides, Fig. 4. 9 d). In this last case, a geniculum at some level along the filaments isoften present.It should be made clear that the terms anterior and posterior refer to the place of insertion,not the place where the actual anther is located. Mukerjee (1945) stated that “in Ilysanthes,the fertile stamens are posterior and the staminodes anterior, while in Bonnaya thearrangement is just the reverse,” but this arrangement was not observed in Nepalesematerial. Often both pairs are connivent, sometimes the anterior staminodes are notconnivent, but parallel. 35
  44. 44. Fig. 4. 8 c: L. anagallis (Burm.f.) Pennell Fig. 4. 8 d: L. parviflora (Roxb.) MukerjeeDNEP 2B 191 DNEP 2B 125Fig. 4. 8 e: L. antipoda (L.) Alston Fig. 4. 8 f: L. nummulariifolia (D.Don) Wettst.Fig. 4. 8 g: L. pusilla (Thunb.) Merrill Fig. 4. 8 h: L. micrantha D.Don Herbarium Specimen EMAK soaked in OT2 36
  45. 45. Fig. 4. 9 a: Floral dissection of L. crustacea (L.) Fig. 4. 9 b: Floral dissection of L.F. Muell.; WIL 326 nummulariifolia (D.Don) Wettst.; STA 4018. Scale bar denotes millimeters. Fig. 4. 9 d: Floral dissection of L. ruellioidesFig. 4. 9 c: Floral dissection of L. pusilla (Willd.) (Colsm.) Pennell; SSW 9272. Anterior stamensBold.; as no flower for Nepal was available, a rudimentary.specimen from area 6 was substituted. ant. anther ► post. anther ► filament appendage ▲ Fig. 4. 9 f: Floral dissection of L. micranthaFig. 4. 9 e: Floral dissection of L. hookeri (D.Don) Wettst.; EMAK 174. Soaked in 6 (10 g(C.B.Clarke) Wettst.; SSW 6767 aerosol OT2 / 100 g H2O) / 1 (acetone). Note upper lip removed; anthers dorsifixed; appendage club-shaped. 37
  46. 46. Gynoecium. The gynoecium is bi-carpellate, and the placentation axile. The style ispenicilliform, simple, but often flattened into two stigmatic lobes at the apex. In somespecies, the style is persistent until dehiscence of the capsule (Fig. 4. 10 e, f + j); in otherspecies, it leaves a ring on the capsule where it was attached, but falls off before the fruitripens. (In Pennell’s words (1943b) “[the capsule of Lindernia procumbens is] tipped by awhite callose style-base”).Fruits. All fruits are septicidally dehiscent capsules. Shapes range from long, linear,subulate (ca. 1 x 12 mm) [Fig. 4. 10 b + i] to almost globose (2 x 2.2 mm) [Fig. 4. 10 g].The style is generally persistent in L. anagallis, L. viscosa, L. crustacea, and L. parviflora[Fig. 4. 10. e, f, h, j]. The texture of the outer capsule wall is usually striate. Fig. 4. 10 a: Dehisced capsule of L. hookeri Fig. 4. 10 b: Dehisced - and closed capsule of L. ciliata (C.B.Clarke) Wettst. (Colsm.) Pennell 38
  47. 47. Fig. 4. 10 c: L. multiflora (Roxb.) Mukerjee Fig. 4. 10 d: L. nummulariifolia (D.Don) Wettst.Fig. 4. 10 e: L. anagallis (Burm.f.) Pennell Fig. 4. 10 f: L. viscosa (Hornem.) MerrillFig. 4. 10 g: L. pusilla (Thunb.) Bold. Fig. 4. 10 h: L. crustacea (L.) F. Muell. 39
  48. 48. Fig. 4. 10 i: L. antipoda (L.) Alston Fig. 4. 10 j: L. parviflora (Roxb.) Mukerjee Fig. 10 k: L. ciliata: Capsule with seedsSeeds. Seed size ranges from ca. 0.2 x 0.1 mm (Lindernia parviflora; L. multiflora) to ca.0.5 x 0.4 mm (L. pusilla; L. ruellioides). Shape varies from the generally round seeds,which can be ellipsoid to elongate (L. nummulariifolia) or quadrangularly rounded (L.pusilla) to generally 5—6 angled seeds with flattened sides, which can be either straight (L.viscosa; L. parviflora) or curved (L. indet-a). Seed colour is usually an orange-red, but canbe green during different stages of maturity (Fig. 10 k above). Most seeds of NepaleseLindernia are alveolated, i.e., the seed coat is depressed between protruding ridges.According to Rahmanzadeh, three goups can be distinguished within Linderniaceae based 40
  49. 49. on seed characters. The first group is characterized by alveoli, exhibited also byCraterostigma Hochst., parts of Artanema and parts of Torenia. The second group ischaracterized by those taxa not exhibiting alveolated seeds, part of which are the Lindernias.str., represented by, e.g., L. parviflora. The third group of the Linderniaceae,characterized by aulacospermous seeds (Rahmanzadeh et al. 2005) was not represented byNepalese Lindernia. Alveolation is strongly associated with appendages, or hook-likethickenings attached to the centre of the depression (see Table 5.1. ALV and APP).Appendage or projection shape is usually short and thick (e.g., L. anagallis), sometimeslong and filiform (L. crustacea; L. multifora), and in L. parviflora consists of dense twisted,round or rugose tissue. Some seeds show a strong tendancy to convolution due toappression to the other seeds in the often tightly-packed capsule. The testal patterns showdifferent densities of tuberculation. 41
  50. 50. 5. RESULTS AND DISCUSSIONThe account for the genus in Nepal is attached in the appendix (1). This chapter deals firstwith results from the SEM seed study (5.1), followed by a comparison of Yamazaki’ssubgeneric limits with those subgeneric distinctions apparent in seed characteristics (5.1.2).Secondly, this chapter discusses specific taxonomic problems and the research needed tosolve them (5.2.1—5.2.6).5.1 Scanning electron microscopy of seeds.According to Lewis (2000), all Lindernia [of the New World] are strongly angled orribbed. The seed coat is alveolate or pitted (e.g., L. nummulariifolia), coarsely or irregularlyribbed (L. hookeri), or areolate (L. procumbens). In this study, four sections could bedistinguished.The seeds of the Nepalese species of Lindernia show considerable variation and a roughgrouping of species was possible by comparison of features such as their alveolate ridges,appendages, and tuberculate seed coats. The comparisons in this chapter, summarized inTable 5 below, are made under the assumptions that like was compared with like, but insome cases the results can but guide further research. Further, care must be taken whengeneralizing from a small study like this one: the seeds used most certainly were collectedat different stages in the life cycle, and were most certainly exposed to different dryingmethods. While it might be considered a characteristic feature of, e.g., Lindernia anagallis,to deflate the seeds as soon as death sets in, it might just have been a process unique to thisparticular sample. It is known, however, that scrophulariaceous seeds are often influencedby tight packing within the capsule (Elisens & Tomb 1983; Lewis 2000). This studyconfirmed that an in-depth SEM study within this group would prove interesting indeed.Because L. procumbens was poorly known at the time of the SEM study, it is only nowclear that the taxon initially determined as L. procumbens is actually L. parviflora, andtherefore L. parviflora was examined twice, from two distinct collections. The un-identified collection, NOR 8037 is included in this analysis, and was named ‘L. parviflora / 42
  51. 51. L. procumbens?’ throughout. A separate, post-SEM study was conducted, comparing seedsof L. parviflora with what is now hypothesized to represent L. procumbens. While thisstudy needs further investigation, and L. procumbens, once identified, should be includedin a follow-up SEM study, preliminary results show a stark difference in size, with L.procumbens seeds being about twice the size, and differently shaped than L. parviflora.Comparison of the seeds made it very clear that L. indet-a and the L. procumbens/ L.parviflora complex are not conspecific (Figs. 5. 2 h and 5. 2 i). Lindernia indet-a had beentreated synonymously with L. procumbens by Philcox and Hara. Summarized in Table 5,the following characters were analyzed:Size and shape. Seed size ranges from the very small (0.24 x 0.12 mm) seed of Linderniaparviflora and L. multiflora (0.22 x 0.16 mm) to the more than twice as large (0.52 x 0.4mm) seeds of L. ruellioides (note reduced magnification in micrograph). Although seedsize within one accession did not vary much, samples in which the seeds were convolutedshowed that the extent to which convolution occurred can vary within one capsule. Further,shapes and sizes of seeds could not be verified in those samples that were stronglyconvoluted (L. anagallis, L. antipoda, L. hookeri, L. crustacea, and L. multiflora). Thedimensions given in Table 5 and in the captions of Figs. 5. 2 a—p are taken from themicrographs; it is obvious that the width of the strongly flattened L. anagallis seed, forexample, is smaller than the width measured in the micrograph. These convoluted seedswould benefit from the soaking and critical point drying method applied to material beforemounting and sputtercoating (e.g., Juan et al. 1997).Shapes, where discernable, vary from the ellipsoid seeds of Lindernia nummulariifolia(Fig. 5. 2 f) to the quadrangular seeds of L. pusilla (Fig. 5. 2 n), which are almost widerthan long. The shape of L. micrantha’s seeds could be described as slightly lanceoloid,whereas those of L. parviflora are strongly angled and columnar. The seed shape of L.indet-a, with its relatively strong angles and one rounded side curving towards the hilum,i.e., the attachment to the free central placenta, makes it easy to picture the placement ofthe embryo within the anatropous ovule. In contrast to L. indet-a, but in concert with L.parviflora, the form represented by L. viscosa shows no curvature, but regular angles thatframe six relatively flat sides. As to the taxon represented by Norkett 8037 (Fig. 5.2 i), the 43
  52. 52. densely rugose appendiculation leaves the shape underneath open for speculation. Here, alatitudinal dissection of the seed would be of great value.Colour. The colours of the seeds ranged from yellowish-orange to red to bright green. Thecolour of the seed was not used to distinguish taxa; it is likely to vary with maturity of theseed. More important are differences in angularity and size (Mill pers.comm. Aug. 2005).Longitudinal and latitudinal alveolate rows. Longitudinal alveolate rows are known fromother genera in the Scophulariaceae, e.g., Bryodes Benth. and Psammetes Hepper (Fischer& Hepper 1997). In this study, almost all seeds were characterized by this structure. Taxacould be grouped into those whose seeds exhibit alveolate rows (Lindernia s.l.), and thosewhich do not (Lindernia s. str., Rahmanzadeh et al. 2005). Where they do, the longitudinaland latitudinal rows form a structural network between which the seed coat is depressed byapproximately 20 μm. Yamazaki (1981) used the term ‘scrobiculate’ to describe thischaracter state in Lindernia. Another term is bothrospermous (Rahmanzadeh et al. 2005).Those pits or depressions are well-preserved in L. ciliata, L. micrantha, and L.nummulariifolia (Figs. 5. 2 d—e). They are also present in L. anagallis, L. antipoda, L.hookeri, L. crustacea, and L. multiflora. Lindernia viscosa (Fig. 5. 2 l) has been termed‘scrobiculate’ by Yamazaki (1981), and a few depressions are visible, but they areshallower than in those seeds mentioned above. The same phenomenon arose in theinterpretation of L. ruellioides, for example, which seems to have but obscure longitudinalrows and depressions (hence the entry in Tab. 5: n/a). This difficulty in discerning thenumber of rows on a seed may be due to the absence of convolution. Therefore, countingthe number of rows was attempted only for those seeds, whose rows were three-dimensionally apparent. A distinction was made between the use of the ‘row’ character andthat of mere ‘angulation.’A row was defined as housing depressions of the seed coat. Angulation was defined as asimple structural line not associated with a depression in the seed coat. This distinction wasalso made by Lewis (2000), who described the seeds as ‘faintly to strongly angled orribbed). Lindernia antipoda and L. ciliata had the highest estimated number (10—12) oflongitudinal alveolate rows (Juan et al. 1997), Lindernia parviflora’s number is half this.For this taxon, the nature of depressions differs in that their shape is not round, but rather 44

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