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


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

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

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  • 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. ABSTRACTThe Nepalese species of Lindernia (Scrophulariaceae s.l.) were revised. These are at least 13distinct species with possibly two additional species. For species delimitations, vegetativeand sexual characters, in particular leaf, flower, and fruit morphology, were taken intoaccount. Numerical taxonomy included leaf length to leaf width ratio and pedicel length toleaf length ratio. Measurements made allowed a definition of inflorescence type, animportant distinguishing character within Lindernia. An SEM study of Lindernia seedmaterial was undertaken. Results did not support the traditional groupings into subgeneraproposed by earlier taxonomists. Taxonomic history of the Nepalese species is summarized,the taxonomic problems discussed, and an account of the genus for the Flora of Nepal isincluded. ii
  • 3. CONTENTSTitle 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. 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. 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. 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 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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
  • 53. rectangular, and framed by crisp latitudinal ridges. Lindernia indet-a is distinct in that itsseed coat is mildly convolute, however it does not have depressions associated with thealveolate rows as in other seeds (Fig. 5.1 h and 5.2 H).Where possible, the total number of depressions observed on one seed was estimated (seeTab. 5; depression number, “DEP#”). In most cases, this number is a function of thenumber of latitudinal rows coinciding with the longitudinal rows, which form the ‘netted’structure across the seed. Here, Lindernia antipoda shows the biggest number, of pits (80—100), L. ruellioides and L. pusilla have the smallest number of pits. The extent of depth(Tab. 5; DEP) is most probably a function of development, structural rigidity, andcircumstances during the sample’s drying. Apparently distinct from the majority of taxaexamined are Lindernia indet-a, which does not show alveoli, and L. parviflora, whosedepressions are elongated rather than circular. Lindernia viscosa also stands out due to itsangulation and the lack of appendages (see following paragraph).Appendiculation of testa. All but two taxa show some form of appendaged tissue attachedto the seed coat. This feature of appendiculation may aid dispersal by fauna (Merrill 1912).Only Lindernia indet-a and L. viscosa lack these. In figures 5.1 h and 5.2 H however, it canbe seen that L. indet-a has an indistinct reticulate pattern, much reduced in comparison withthe strong reticulation exhibited by L. pusilla (Fig. 5.1 n and 5.2 N). Lindernia ruellioidesexhibits the most conspicuous, widest processes (Fig. 5.2 M + α). For NOR 8037 (L.parviflora/ L. procumbens ?), the massive, rugose (=rounded) wall thickenings are dense.When comparing Figs. 5.2 G, I, and O, it becomes conceivable that NOR 8037 may belongto the same (i.e., L. parviflora) taxon, and that G – > O – > I are a series of increasinglymature seeds. When comparing Figs. 5.2 α + β, the differences in appendage dimensionbetween the larger Lindernia ruellioides and the shorter L. anagallis become clear.Appendages on seeds of L. antipoda, L. hookeri, L. ciliata, L. micrantha and L.nummulariifolia, as well as those on Mazus delavayi conform with those of L. anagallisrather than L. ruellioides.Lindernia crustacea and L. multiflora (Figs. 5.2 J + K) have filiform appendages. Whileseveral seeds were sampled of each taxon, and all show this form of appendiculation, it isnot clear whether the origin of this tissue derives from the inside of the capsule, or whetherit is an actual outgrowth from the seed coat. Lindernia pusilla is the most distinct species of 45
  • 54. all Lindernia; its reticulate wall thickening (Barthlott 1981; Canne 1979) is not found inany other seeds in this study, except indistictly in L. indet-a. Distinct from the majorityhere are L. viscosa, L. pusilla, L. indet-a, and L. parviflora.Testal pattern. The seed coats themselves show more or less minute (ca. 1 μm in diameter),tuberculate lumps on almost all seeds. One exception again is Lindernia ruellioides, whosetestal pattern is granular (Fig. 5.2 m + α), reminiscent of a stucco wall. L. anagallis hastuberculate lumps especially in the areas where the appendage attaches to the testal wall;elsewhere the surface appears rather smooth (Fig. 5.2 a +β). Similarly, L. antipoda, L.hookeri, L. micrantha and L. crustacea have a higher density of tuberculation around theappendage. The remaining taxa are evenly tuberculate throughout. Tuberculation in L.viscosa (Fig. 5.2 L) is densest. The nature of tuberculation seems to differ among thespecies, but an even greater magnification would have to be applied to properly analyzethese differences. However, when comparing the general appearance of tuberculation in theLindernia seeds with that of the Mazus seed, a qualitative difference is apparent (Fig.5.2.p), the testal pattern of Mazus appearing more porous. Standing out within this groupare Lindernia ruellioides (granular), L. viscosa (densest), and L. indet-a (partially lackingobvious tuberculation; Fig. 5.2.H). 46
  • 55. Lindernia COLL # LTH WTH SHP ALV LO-R LA-R APP. C. DEP DEP # TESTA SSW 8— anag. 569 0.28 0.22 R Y 7--9 10 W+S Y M 56--90 TUB+SM PSW 10-- 8— 80-- TUB- antip. 5664 0.28 0.24 R Y 12 10 W+S Y M 100 WK SSW hook. 6767 0.4 0.33 R Y 6--8 5—6 W+S Y D 30--48 TUB SSW 10-- cili. 7581 0.28 0.24 A Y 12 7—8 W+S N D 70--96 TUB PSW micr. 5856 0.36 0.22 A--L Y 7--9 6—8 W+S N D 42--72 TUB SSW num. 4018 0.45 0.25 R Y 8--10 8 ca W+S N M 64--80 TUB TI parvi. 1492 0.24 0.12 A--C N 6 ca 6—7 R N S n/a TUB PSW indet-a. 4476 0.38 0.18 A N none 7—8 0 N S n/a SM/ TUB NOR ?? 8037 0.34 0.17 R N n/a n/a R N n/a n/a TUB PSW crust. 5624 0.38 0.29 R Y 8--10 6 F Y D 48--52 TUB BUC multi. 1387 0.22 0.16 R Y 7--9? 6—7 F+W Y D 42--60 TUB SSW TUB- visco. 7056a 0.35 0.26 A N 7--9 6—7 0 N S n/a DEN TI ruell. 1484 0.52 0.4 R n/a n/a n/a W+L N S 32--36 GRAN TI pusi. 1478 0.38 0.39 R--A n/a 7--8 5—6 0 N S 32--36 TUB SSW parvi 8696 0.24 0.14 A--C N 6 ca 6—7 R N S n/a TUB Mazus DNEP 14-- delavayi 2B67 1.6 0.67 R Y 16 n/a W+S N M n/a WKTab. 5: Seed and seed-surface characters from SEM study. Columns show taxa; COLL#= collectors’ codes [see belowfor a legend to the abbreviations]; dimensions, i.e. LTH=length; WTH=width of entire seed; SHP=shape (R=round,not sharply angled, A=angled, L=lanceolate appearance, C=columnar appearance). ALV= presence [Y] or absence [N]of alveolation; in some cases the character could not be scored [n/a]; LO-R= estimated number of longitudinal rows;LA-R= estimated number of latitudinal rows; APP=appendiculation (W=wide, S=short, R=rounded, F=filiform,L=long, 0=absent) C=convolution; DEP=depression (S=shallow, M=medium, D=deep); DEP#=estimated number ofdepressions per seed; TESTA=seed coat pitting (TUB= tuberculate, WK=weakly so, DEN=densely so, SM=smooth,GRAN=granular, i.e. ‘stucco-effect’); Entries in bold signal extreme values in the group of taxa. Abbreviations ofcollectors as follows: SSW = Stainton, Sykes & Williams; PSW = Polunin, Sykes & Williams; TI = Tokyo Institute;NOR = Norkett; BUC = Buchanan-Hamilton; EMAK, DNEP = Edinburgh Expedition Codes; S = Stainton 47
  • 56. Fig. 5.1 a.L. anagallis (Burm.f.) PennellSSW 569Dimensions: 0.28 x 0.22 mm.Longitudinal alveolate rows: 7—9.Latitudinal ridges: 8—10.Surface scrobiculate/ depressed in pitsbetween ridges, ca. 56—90 pits/ seed.Tendency to convolution.Appendages present.Fig. 5.1 b.L. antipoda (L.) AlstonPSW 5664Dimensions: 0.28 x 0.24 mm.Longitudinal alveolate rows: 10—12.Latitudinal ridges: 8—10.Surface scrobiculate/ depressed in pitsbetween ridges, ca. 80—100 pits/ seed.Tendency to convolution.Appendages present.Fig. 5.1 c.L. hookeri (C.B.Clarke) Wettst.SSW 6767Dimensions: 0.4 x 0.33 mm.Longitudinal alveolate rows: 6—8.Latitudinal ridges: 5—6.Surface scrobiculate/ depressed in pitsbetween ridges, ca. 30—48 pits/ seed.Tendency to convolution.Appendages present. 48
  • 57. Fig. 5.1 d.L. ciliata (Colsm.) PennellSSW 7581Dimensions: 0.28 x 0.24 mm.Longitudinal alveolate rows: 10—12.Latitudinal ridges: 7—8.Surface scrobiculate/ depressed in pitsbetween ridges, ca. 70—96 pits/ seed.Raphe visible.Appendages present.Fig. 5.1 e.L. micrantha D.DonPSW 5856Dimensions: 0.36 x 0.22 mm.Longitudinal alveolate rows: 7—9.Latitudinal ridges: 6—8.Surface scrobiculate/ depressed in pitsbetween ridges, ca. 42—72 pits/ seed.Raphe well visible.Appendages present.Fig. 5.2 f.L. nummulariifolia (D.Don) Wettst.SSW 4018Dimensions: 0.45 x 0.25 mm.Longitudinal alveolate rows: 8—10.Latitudinal ridges: ca. 8.Surface scrobiculate/ depressed in pitsbetween ridges, ca. 64—80 pits/ seed.Appendages present. 49
  • 58. Fig. 5.1 g. L. parviflora (Roxb.) Haines TI 1492 Dimensions: 0.24 x 0.12 mm. Longitudinal ridges: 5—8. Latitudinal ridges: ca. 10. Form somewhat columnar— hexagonal. Surface depressions not well visible due to rounded/ rugose processes/ appendages. Fig. 5.1 h. Lindernia indet-a PSW 4476 Dimensions: 0.38 x 0.18 mm. Longitudinal angulation: 5? Latitudinal ridges: 7—8? Surface depressions not obvious.XX Fig. 5.1 i. L. procumbens (Krock.) Philcox? NOR 8037 Dimensions: 0.34 x 0.17 mm. Neither longitudinal nor latitudinal ridges visible due to rounded/ rugose processes /appendages. Surface depressions not obvious. 50
  • 59. Fig. 5.1 j.L. crustacea (L.) F. Muell.PSW 5624Dimensions: 0.38 x 0.29 mm.Longitudinal alveolate rows: 8—10.Latitudinal ridges: ca. 6; not strong.Tendency to convolution/scrobiculation.Estimated number of depressions48—52.Surface depressions well visible.Long hair-like appendages.Fig. 5.1 k.L. multiflora (Roxb.) MukerjeeBUC 1387Dimensions: 0.22 x 0.16 mm.Longitudinal alveolate rows: 7—9?Latitudinal ridges: 6—7; not strong.Estimated number of depressions32.Surface deeply depressed in pitsbetween ridges, ca. 42—63 pits/seed; strongly convolute.Fig. 5.1 l.L. viscosa (Hornem.) MerrillSSW 7056aDimensions: 0.35 x 0.26 mmLongitudinal ridges: 7—9?Latitudinal ridge: 6—7;Surface depressions discernible, butshallow. 51
  • 60. Fig. 5.1 m.L. ruellioides (Thunb.) Bold.TI 1484Dimensions: 0.52 x 0.4 mm.Longitudinal and latitudinal ridgesnot protruding.Conspicuous, branched trichome-like appendages covering shallowdepressions in coat.Estimated number of depressions32—36.Note magnification reduced to 500 x.Fig. 5.1 n.L. pusilla (Thunb.) Bold.TI 1478Dimensions: 0.375 x 0.385 mm.Longitudinal ridges: 7—8.Latitudinal ridges: 5—6.Strongly reticulate seed wallthickenings.Estimated number of depressions32—36.Note magnification reduced to 500 x.Fig. 5.1 o.L. parviflora (Roxb.) HainesSSW 8696.Compare with same taxon (Fig. g.).Dimensions: 0.24 x 0.14 mm.Longitudinal ridges: ca. 6?Latitudinal ridges: 6—7.Estimated number of depressions inseed coat 36—42; not pit-like, butrather laterally elongated. 52
  • 61. Fig. 5. 2 A. L. anagallis (Burm.f.) Pennell SSW 569 Trichome-like appendages. Density of tuberculation variable. Fig. 5. 2 B. L. antipoda (L.) Alston PSW 5664. Trichome-like appendages. Density of tuberculation variable. 100 μm Fig. 5.2 C. L. hookeri (C.B.Clarke) Wettst. SSW 6767 General lack of trichome-like appendages. Density of tuberculation not variable.100 μm 53
  • 62. Fig. 5. 2 D. L. ciliata (Colsm.) Pennell SSW 7581 Trichome-like appendages. Density of tuberculation not variable. 100 μm Fig. 5. 2 E. L. micrantha D.Don PSW 5856 100 μm Trichome-like appendages. 100 μm Density of tuberculation variable. 100 μm Fig. 5.2 F. L. nummulariifolia (D.Don) Wettst. SSW 4018 Trichome-like appendages. Density of tuberculation not variable. 100 μm100 μm 54
  • 63. Fig. 5. 2 G. L. parviflora (Roxb.) Haines TI 1492 Trichome-like appendages rugose/ rounded. 100 μm Density of tuberculation not variable. Fig. 5.2 H. L. indet-a PSW 4476. No appendages. Seed coat relatively smooth; occasionally minutely tuberculate.100 μm Fig. 5.2 I. L. procumbens (Krock.) Philcox?/ L. parviflora (Roxb.) Haines NOR 8037 Trichome-like appendages rugose/ rounded. Tuberculation evenly dense. The differences in appendage extent (Graphs G—I—F) may be due to developmental timing. 55
  • 64. Fig. 5.2 J. L. crustacea (L.) F. Muell. PSW 5624 Filiform cuticular projections. Density of tuberculation variable.100 μm 100 μm 100 μm 100 μm Fig. 5.2 K.. L. multiflora (Roxb.) Mukerjee BUC 1387 Filiform cuticular projections. Density of tuberculation mildly 100 μm variable. Fig. 5.2 L. L. viscosa (Hornem.) Merr. SSW 7056a General lack of cuticular projections. Density of tuberculation mildly 100 μm variable. 56
  • 65. Fig. 5.2 M.L. ruellioides (Colsm.) PennellTI 1484.Trichome-like appendagesconspicuously large.Granular tuberculation evenlydense; i.e. ‘stucco – effect.Fig. 5.2 N.L. pusilla (Thunb.) Merr.TI 1478Reticulate appendages.[This shape is reminiscent of somePseudolysimachion seeds describedby Martinez-Ortega & Rico (2001).]Tuberculation evenly dense.Fig. 5.2 O.L. parviflora (Roxb.) HainesSSW 8696Compare with same taxon (Fig. G.).Trichome-like appendages rugose/rounded, appearing reticulate attimes, comparable but muchreduced from 5.3 I.[The shape is reminiscent of someVeronica seeds (Martinez-Ortega &Rico 2001.]Tuberculation consists of evenlydistributed bumps. 57
  • 66. Fig.5 2 P) Mazus delavayi Bonati DNEP 2B 67. Dimensions: 1.6 mm x 0.67 mm Longitudinal alveolate ridges 14 – 16. No apparent latitudinal ridges. Trichome-like appendages. Fig. 5. 2 p) Mazus delavayi Bonati DNEP 2B 67. In contrast to Lindernia spp. this taxon does not show the typical tuberculate pitting. 100 μm However, trichome-like appendages prove to be not genus-dependent.Fig. 5. 2 α. Attachment of appendage in L. Fig. 5. 2 β. Attachment of appendage in L.ruellioides (Colsm.) Pennell. anagallis (Burm.f.) Pennell.Magnification: 4500 x Magnification: 4500 xAppendage stretching over depression in seed coat. Projections much smaller than in α. Seed coatSeed coat coarsely granular/ strongly pitted smooth away from the appendages, with surfacethroughout (like stucko). features more dense closer to the attachments. 58
  • 67. 5.1.1 SEM results and Yamazaki’s recent subgeneraYamazaki included seed characters in his subgeneric classification of Lindernia (1981). Asto the taxa represented here, he grouped the following subgenera:Subgenus I, Lindernia, section (1) Lindernia: L. procumbens (not in the SEM study). (2) Euilysanthes: L. parviflora.Subgenus II, Tittmannia, sect. (1) Tittmannia: L. pusilla, L. viscosa, and L. hookeri. (2) Angustifolia: L. micrantha and L. anagallis. (3) Nummularia: L. nummulariifolia. (4) Torenioides: L. crustacea.Subgenus III, Bonnaya, sect. (1) Bonnaya: L. antipoda, L. ciliata, and L. ruellioides. YAMAZAKI’S SUBGENERA AND SECTIONS FAMILY: SCROPHULARIACEAE GENUS: Lindernia Subgenera: Lindernia Tittmannia BonnayaSection: Lindernia EuilysanthesTittmannia Angustifolia Nummularia Torenioides Bonnaya L. procumbens L. parviflora L. pusilla L. anagallis L. nummulariifolia L.crustacea L. ciliata L. viscosa L. micrantha L. ruellioides L. hookeri L. antipodaFigure 5.3: Classification of Lindernia based on Yamazaki (1981). 59
  • 68. I (1): Seeds of L. procumbens were not examined due to lack of material. They weredescribed by Yamazaki as “oblong, ca. 0.3 x 0.15 mm, reticulate, longitudinally ridged.”I (2): L. parviflora “ellipsoidal, ca. 0.2 x 0.12 mm, reticulate, longitudinally ridged” differsfrom the above in shape and size. This distinction between the above conforms to mypreliminary results.II (1): His description of Lindernia pusilla did not mention the reticulate pattern observedin our sample. However, under 50x microscope, this reticulation is not obvious. Hedescribed the seeds as “shortly cylindric, ca. 0.3 x 0.2 mm, scrobiculate.” Following in thissame section is L. viscosa with exactly the same description. The last species belonging toYamazaki’s group is L. hookeri, but here only the variety L. hookeri var. cochinchinensisBonati is described as “globose or broadly ellipsoidal” (followed by the same dimensionsand testal pattern).II (2): Yamazaki’s grouping of Lindernia anagallis with L. micrantha is sound, but forseed characters alone I would extend it to L. antipoda, L. hookeri, L. ciliata, and L.nummulariifolia. His two descriptions of species, “seeds shortly cylindric, ca. 0.3 x 0.15mm, scrobiculate” for L. micrantha and “seeds cylindric, ca. 0.3 x 0.15 mm, scrobiculate”for L. anagallis differ hardly from those of L. antipoda “shortly cylindric, 0.3 x 0.2 mm,scrobiculate” or L. ciliata “ellipsoidal, ca. 0.25 x 0.15 mm, scrobiculate.”II (3): Lindernia nummulariifolia is distinct from section II (2) because of its size“ellipsoidal, 0.4 x 0.3 mm, scrobiculate.”II (4): L. crustacea, “ellipsoidal, 0.4 x 0.25 mm, scrobiculate,” is distinct due to its filiformappendages. I am not aware of any studies showing whether these appendages are typicalof Torenia. I would include L. multiflora (a taxon that Yamazaki did not discuss).III (1): Yamazaki treated Lindernia antipoda, L. ciliata, and L. ruellioides as distinct, agrouping based on its similar fruits, but which cannot be upheld based on seed charactersalone.Assuming that seed characteristics delimit groups within Lindernia, and further assumingthat the characters observed here are independent from each other (an assumption that maywell be rejected when tested), I propose the following key to groups: 60
  • 69. 1A Granular testal pattern with long hooked appendages.......................... L. ruellioides.1B Tuberculate testal pattern with short, or no appendages............................................22A Alveolate seed coat....................................................................................................32B Non-alveolate seed coat.............................................................................................53A Alveolae with regularly dispersed appendages.........................................................43B Alveolae without regularly dispersed appendages: L. hookeri and L. antipoda.4A Appendages short: L. micrantha, L. anagallis, L. nummulariifolia and L. ciliata.4B Appendages filiform: L. crustacea and L.multiflora.5A Non-alveolae with strong reticulate wall thickening...................................L. pusilla.5B Non-alveolae without or with mild reticulate wall thickening................................66A Projections rugose..................................................................................L. parviflora.6B Projections absent: L. viscosa and L. indet-a.The groups arrived at in the key overlap and do not coincide with those drawn up byYamazaki. A solid key to the species based on seed characters would require a largesample of, e.g., all Linderniaceae s.l. I agree that Lindernia parviflora belongs to a distinctgroup. Further, Yamazaki’s section Bonnaya corresponds in its capsule with sectionAngustifolia, being distinct in their inflorescence types. Their shared alveolate seedcharacteristic suggests that inflorescence type is a weak character, whereas capsule shapemay be a stronger indicator of species affinity. Yamazaki’s series Tittmannia included threespecies that have little in common as far as the seed coat is concerned. In fact, thesubgeneric affinities of Tittmannia and Bonnaya shown in the SEM strongly supportPennell’s choice to unite the two groups into Lindernia.From this small study I interpret two major natural groups of alveolate, and non-alveolatetaxa, as has been found by Rahmanzadeh et al. (2005; he described a third group, that doesnot occur here). These are further divided based on the nature of the appendages. It is 61
  • 70. conceivable, however, that a larger study would reveal a strong connection between alveoliand appendages. Lindernia ruellioides was found distinct based on its granular testalpattern and its large, broad appendages. I further argue for the distinctness of Linderniaparviflora based on its strongly angled, columnar shape also found for L. microcalyx Stehléby Rahmanzadeh et al. (2005). Lindernia pusilla is distinct because of its stronglyreticulate seed coat pattern; L. indet-a’s seed coat lacks projections, is mildly reticulate, butappears smooth in those places where the seed has been exposed to pressure fromneighbouring seeds. Wile a large SEM study may prove to differ, the results from this smallstudy suggest that seed characters alone may not resolve the relationships of taxa withinLindernia. It would, however, be useful to extend the study to include neighbouring taxa inthe Linderniaceae.Regarding the role of morphology in phylogeny reconstruction, it has been argued that it isbetter to include few rigorously studied anatomical characters rather than a large datamatrix with ambiguous characters (Scotland et al. 2003). SEM-based seed characters haveproved informative in other genera 17 of the (ever further disintegrating) Scrophulariaceae,and they are likely to prove informative in the phylogenetic reconstruction of Lindernia(Mill, pers. comm. Aug. 2005). It would be interesting to test, whether the appendagecharacters are useful in predicting affinities within this taxonomic group, and further, to testwhether any of the characters explored above have evolved in other clades also. I suggestto include seed, capsule, and venation characters amongst others in a large morphologicaldata matrix. It would be very useful also to include phytochemical characters in a combinedmorphological and molecular phylogenetic study..A short SEM study on lamina characters was conducted on a few species of Lindernia, i.e.from both sides of a leaf of L. ciliata, L. ruellioides, L. antipoda, and L. multiflora. Thispilot study did not prove to be nearly as informative as the seed characters (at first glance);nevertheless, for completeness, the eight micrographs were included in the appendix (5).17 Agalinis (Canne 1979), Cordylanthus (Chuang & Heckard 1972), Antirrhinae (Elisens 1985), Linaria(Segarra & Mateu 2001). 62
  • 71. 5.2 Taxonomic problemsThis section discusses major and minor problems and open questions concerning thecompleteness of this revision. The following sections are given:(5.2.1) The identity of the type of the genus.(5.2.2) Comparison beween Lindernia parviflora and Vandellia erecta.(5.2.3) The identity of Lindernia indet-a.(5.2.4) Lindernia indet-b of “Dr. Buchanan.”(5.2.5) The type of L. micrantha D.Don.(5.2.6) The question of L. oppositifolia.5.2.1 Identity of the type species of the genusThe state of identity of the type of the genus is problematic, because Allioni did not specifyan official binomial. His publication from 1766 states something along the lines of“Linderniam nomine [..Lindernam..] pyxidariam apellavit.” It took Linnaeus to “activate”this binomial in his Mantissa Plantarum of 1771. There, Linnaeus (re-)created Linderniapyxidaria L., using Allioni’s legitimate genus description indirectly referring to the typespecimen Allioni had in mind for the genus Lindernia. No other Lindernia had beenlegitimately described since 1766. However, because Linnaeus cited in synonymyCapraria gratioloides L., which again is thought to be based on Gratiola dubia L., a taxondistinct from L. pyxidaria, the fact that he also cited von Lindern’s illustration of Allioni’stype did not save him from having made an illegitimate description before the InternationalCode of Botanical Nomenclature, and therefore, in 1965, Philcox published a correct (McNeill, pers. comm. Aug. 2005) lecto-typification [in Taxon (1): 30] when chosingKrocker’s Anagalloides procumbens as the legitimate type of the genus. It is strange,however, that the publishers of Index Nominum Genericorum have not adopted this newtype of the genus in their database, which was created much later than 1965. It seemsunfortunate that this modern lecto-typification, with all the material available today, wasnot connected with an actual herbarium specimen, but in contrast, is based on a copperetching on which not even lamina venation is discernable (recall Fig. 1 d). I am not againstusing illustrations as types, but I believe that a single specimen seen and personallyannotated by Linnaeus could be found superior to Krocker’s illustration (recall Fig. 1 b). 63
  • 72. The type for Linnaeus’ Capraria gratioloides will have to be determined. The scansrequested from the Linnean Society for this project included a specimen entitledgratioloides (Fig. 5.3 a + b), but it is not certain whether this is the type specimen. Oncedetermined, its synonymy will have to be verified, to ultimately reject Linnaeus L.pyxidaria, or to reject Philcox’s typification. It will be helpful to revisit an old Linderniaspecimen ending in “gratioloides” at the BM, curated in the Lindernia genus folders.Further, a type specimen for Allioni’s Lindernia (pyxidaria), e.g., Gagnebin’s specimenfrom 1763 should be found. Last, I suggest that natural variation between all specimensclaimed to represent Allioni’s first description (these are spanning the globe) should becarefully studied, and compared with the type Capraria gratioloides L. I hypothesize thatthe results will either reject synonymy between Vandellia erecta Benth. and Anagalloidesprocumbens Krock., or include Capraria gratioloides L. in it.Fig. 5. 4 a: Herb. Linn. 796.3 bears resemblancewith L. procumbens (Krock.) Philcox. Type Fig. 5. 4 b: Same as a; cut, pasted & enlarged.candidate for Capraria gratioloides L. ? 64
  • 73. Maximovicz’s work would represent a good starting point to find European specimens. Forexample, his reference to Döll’s Flora von Baden (1859) suggested two putative typespecimens for L. pyxidaria All.: Reichenbach 1698 and Schultz 70. This description cited10 further localities with specimens from Germany [also revealing that the leaves areblue—green, ovate-elongate, but obovate at the base of the plant; the seeds germinate evenafter several years of drought, and describing in detail the shape and structure 18 of the seedsand their manner of attachment to the septum].Krocker’s protologue 19 indicates that he described a cleistogamous plant, whose flowersare tiny, hardly two millimeters long, and closed. The German name of the plant“Liegender Bastardhünerdarm” may point to the inbreeding mechanism of the plant ( =bastard). As for the rest of the literal translation, liegender portrays the decumbent habit ofthe plant (= laying down); hünerdarm translated literally means chicken’s intestine. Furtherclues to the identity of the plant are given: The peduncles are longer than the leaves, thefour anthers have no filaments, the style is subulate, the stigma is penicilliform. As to itslocation, the next time I am near the river Oder at the border of Poland and Germany, I willcertainly be on the lookout to re-collect it (although it would not be surprising were it at thebrink of extinction by now 20). Identity of the Asian form of the type speciesThe problem of typification we face is compounded by the difficulty of having acleistogamous European species synonymized with a chasmogamous Asian species. Unlessa lecto-type specimen can be chosen for our as of yet elusive Lindernia pyxidaria L. inAsia, as may be possible for Allioni’s Lindernia (pyxidaria) through careful study of thespecimens he dealt with (e.g., Gagnebin), it remains unclear whether Bentham’s (1835)Asian Vandellia erecta, for which he cited Wallich 3947 (and, initially in 1835, Wallich3943; Figs. 5. 4 a – d) can (emphasis added) be synonymized with Krocker’s Anagalloides18 Döll’s description of the seeds: numerous, very small, long-threadlike, usually with 5 longitudinaledges, which are either latitudianally ribbed or rough and granular; the seeds are usually a bit curved.19 The Latin text from Flora Silesiaca is attached in the appendix (4); crucial words are underlined red.20 L. procumbens is on the Red List in Switzerland. 65
  • 74. procumbens or with Allioni’s Lindernia pyxidaria, the question of illegitimacy set aside fora moment 21.As discussed in section 2.1 of the Taxonomic History, and in section 5.2.1 of the resultsabove Lindernia procumbens (Krock.) Philcox was chosen as the new type species of thegenus Lindernia by Philcox (1965). During this study I was not able to develop a clearunderstanding of this species because of the lack of material from Nepal. The relationshipbetween L. procumbens and L. parviflora (Roxb.) Haines is unclear. Theoretically, thespecies are distinct on the basis of the number of anthers [4 fertile anthers in L. procumbensand 2 fertile anthers in L. parviflora] and the lamina margin [entire in L. procumbens andminutely 1—2-dentate in L. parviflora (Yamazaki 1981). Both of these have been reportedfrom Nepal (e.g., Vandellia erecta Wallich ex Benth.), but I only found materilal of L.parviflora. The specimen determined on the herbarium sheet TI 6303760 (BM) as L.procumbens did not have a corolla; further, the material determined by me to be L.parviflora did not always show the minute dentation on the margin of the lamina. The thirdproblem I have is whether the cleistogamous plants from Europe (i.e. L. procumbens) arethe same species as chasmogamous Nepalese plants (i.e. L. parviflora).The differences between the inbreeding European and outcrossing Asian plants may bedue to the environment. Taking this thought a step further, would it be possible for aspecies to have 4 fertile anthers when flowering early in the year, and 2 fertile antherslater? To find out whether this might be possible, one would have to study thepollination syndrome and foraging behaviours of the insects in the localities andclimates involved (Ronse Decraene pers. comm.).Pollination adaptation in Lindernia has been studied by Maximovicz (1874), Urban (1884),and von Wettstein (1895). These teatments were mostly concerned with the intraspecificadaptations to cleistogamy and chasmogamy. Recently, it has been reported by Lewis(2000) that in typically chasmogamous flowers of L. procumbens and L. dubia, earlyflowers are often cleistogamous, and set fruit by self-pollination. In greenhouse conditions21 And to take the question of illegitimacy back up immediately, would not Bentham’s creation of Vandelliaerecta, in which he cited a wrong synonymy (i.e., Tittmannia trichotoma = L. multiflora), be illegitimate? 66
  • 75. on the other hand, e.g., L. grandiflora Nutt. will not produce capsules, and mustobligatorily outcross. While this evidence makes it more credible that L. procumbens(Krock.) Philcox from Europe may be conspecific with Nepalese material, the geographicvariability and pollinator availability must be considered when searching for the factorsresponsible for a variation in the number of fertile stamens.According to ontogenetic studies (Walker-Larsen & Harder 2000), zygomorphy in theLamiales is often associated with partial loss of the stamen whorl. While generally in theasterids this reduction occurred only seldomly, Lindernia represents one case where such areduction has led to the formation of a didynamous androecium. Walker-Larsen & Harderclaim that the persistence of staminodes after reduction of the stamen whorl occurs in theasterids I clade only in some Solanaceae, and bilabiate-flowered Scrophulariales. Of theantesepalous whorl, the three stamens opposite the lower calyx have moved to the top ofthe upper lip, losing one stamen (probably in order to support the function of the tworemaining stamens). For the two stamens associated with the posterior two calyx lobes, theinsertion remains antesepalous, but the filaments are much extended to allow the anthers tobe connivent below the posterior lip. If these two anterior stamens have lost their primaryfunction of pollen production, it might be to prevent self-pollination. Studies on pollinationsyndromes suggest that such transformations aid pollinator attraction, easily verified by alook at L. antipoda’s corolla with bright yellow staminodes. It is interesting to note that thepartial loss of a stamen whorl is reversible. It is further interesting to note that Blumereported Linderniae with two fertile stamens and two, or sometimes three sterile stamens inJava.It would be very interesting indeed to explore the relationship between L. parviflora withtwo, and L. procumbens with 4 fertile anthers, and to see whether they are actuallyconspecific, and have adaptated differently due to flowering times of the year in differentgeographic areas, and/ or pollinator- availability in temperate zones during separateseasons. 67
  • 76. 5.2.2 Vandellia erecta and Lindernia parvifloraIn this section, first or early historic taxonomic treatments are compared to find evidencefor potential conspecificity of the material that has been treated as distinct since its firstdescription. The question arose upon searching for the type material of (1) L. procumbens(Krock.) Philcox and (2) L. parviflora (Roxb.) Mukherjee. Upon finding that the identitiesof the above are dubious at best, respective treatments and protologues were compared.It is now clear that Krocker’s Anagalloides procumbens is cleistogamous. Philcox’sLindernia procumbens, however, includes both cleistogamous and chasmogamous forms ofL. procumbens, since he synonymized it with Bentham’s Vandellia erecta. However,Philcox never treated L. parviflora, so we do not know whether he would have alsoincluded L. parviflora in synonymy of his [broad] species concept. In search of clueswhether it would be possible that one species has been treated as two distinct species for185 years based on one character alone (although a strong one), it was thought interestingto compare the descriptions of the two according to Bentham, and to see whether othercharacters than number of fertile stamens were used to keep the two in distinct genera.Here I compare Bentham’s historic treatments of the species, starting from 1835’sScrophularineae Indicae with Vandellia erecta (left column), and Bonnaya parviflora(right column), and below moving on to the respective entries in De Candolle’s ProdromusX [DC X] of 1846. Pertaining to V. erecta, Bentham omitted the synonymy of Tittmanniatrichotoma Benth. in his later treatment. He was then able to change “pedunculis axillaribusracemosisve” to “pedunculis omnibus axillaribus.” No characters openly oppose eachother, and throughout the treatments (1835 & 1846), he points out the similarity between V.erecta and Bonnaya / Ilysanthes parviflora, and his incapacity to distinguish them fromeach other by any character other than the stamen number. Especially his later descriptionof I. parviflora points to the typical manner of inflorescence, which does tend to appearsubracemose toward the apex of the plant, with relatively shorter latitudinal branches. 68
  • 77. “V. erecta suberecta, glabra, foliis subsessilibus “B. parviflora (Bentham inovatis oblongisve basi angustatis, pedunculis Wall. Cat. n 3867), humilis,axillaribus racemosisve calyce 2-5 plo ramosa, foliis oblongislongioribus, calycibus ante anthesin 5-partitis vel lanceolatisve, floribusprofunde 5-fidis, laciniis lanceolatis axillaribus subracemosisve,membranaceis capsulas ovatas superantibus.— corollis calyce duploTittmannia trichotoma, Benth. in Wall. Cat. n. longioribus.—Gratiola3943.—Tittmannia erecta, Benth. in Wall. Cat. n. parviflora Roxb. Pl. Corom.3947 (ubi cum Bonnaya parviflora miseitur) 3. 3. t. 203” [see Fig. 2. a)[Fig.].—Valde affinis V. crustaceae, sed satis above].distincta videtur.Hab. in Peninsula, Deyra Dhoon, Munghyr, “Hab. in Hindustania etNapalia, Sillet, Assufghur, Regno Birmannico, Sillet, Wallich, China,&c., Wallich, Royle, &c.” Nelson.” FROM SCROPH. IND. OF 1835 FROM SCROPH. IND. OF 1835In DC X, Bentham continues his treatment 11 years later: “V. erecta; erecta, ramosissima vel “I. parviflora ; ramosissima, gracilis, subdiffusa, glabra, foliis infimis ovatis in petiolum foliis oblongis lanceolatisve, brevem angustatis, inferioribus in petiolum angustatis, floralibus sessilibus vel superioribus sessilibus vel semiamplexicaulibus ovatis semiamplexicaulibus, floribus oblongis lanceolatisve, pedunculis superioribus subracemosis, corolla omnibus axillaribus calyce pluries calyce subduplo longiore, longioribus, calycis segmentis filamentis anticis leviter glandulosis lineari-lanceolatis, capsulis ovato- infra apicem lobulo minutissimo globosis. [..] in India orientali glabro auctis. [...] in India orientali frequens a Peninsula ad Nepal et frequens a peninsula (Wight! herb. n. ripas Irawaddy. Tittmannia erecta 2200) ad Napaliam (Wall.! h. DC.); Benth.! in Wall. cat. n. 3947. Gratiola parviflora Roxb.! Pl. Ilysanthi parviflorae ita similis ut Corom. 3. p. 3. t. 204. Scroph. Ind. p. vix nisi staminibus distingueres. 34. A simillima I. capensi Filamentorum anticorum distinguitur habitu gracili, floribus appendicula longa linearis. (v.s.)” minoribus, filamentorum anticorum ramo glabro, latitudine rami majoris subbreviore. (v. s.) FROM DC X OF 1846 FROM DC X OF 1846 69
  • 78. Fig. 5. 4 d: Wallich 3943; Bentham quoted this specimen in synonymy with V. erecta Benth. in 1835, but excluded it in 1846. It is L. multiflora (Roxb.) Mukerjee Fig. 5. 4 c: Wallich 3947C; Bentham quoted Wallich 3947 as voucher for V. erecta Benth. in 1835 & 1846. Fig. 5. 4 e: Entry of Tittmannia erecta Benth. in the Wallich Catalogue 3943 Fig. 5. 4 g: Wallich s.n., Nepal; determined as Ilysanthes parvifloraAt species level, he writes: Fig. 5. 4 f: Wallich 3947B; another L. procumbens from Nepal. 70
  • 79. In theory, the type material referred to by Bentham should show a slight difference in (1) habit:erect—suberect versus low-lying and more slender (2) leaf shape: ovate—oblong versus ovate—lanceolate (3) inflorescence: completely axillary versus subracemose toward the apex and (4), if itwere visible from herbarium material, appendage on the anterior filaments versus lack of appendage.His comparison with I. capensis in which he stessed its [more] slender habit and its smaller flowers,should be further explored. It was almost certainly due to the lack of specimens available thatBentham may have had trouble to reach a conclusion about Ilysanthes parviflora and Vandelliaerecta. Below are the specimens Bentham referred to (Fig. 5.4 c – g).Further clues (highlighted in bold) were found in Hooker’s treatment of the Flora ofBritish India, where he distinguished the genera Ilysanthes Rafin. from Vandellia L. “Vandellia, Linn.: Herbs, usually annual, glabrous or pubescent. Leaves opposite, often toothed, penninerved (except in V. erecta). Flowers small, axillary or racemose; pedicels ebracteolate. Sepals 5, free or connate in a subcampanulate calyx, angles not winged. Corolla tube cylindric; upper lip erect, broad, concave, notched or bifid; lower broader, spreading, 3-lobed. Stamens 4, all fertile (except V. molluginoides); 2 posterior usually included; 2 anterior with arched filaments appendaged at or towards the base; anthers touching or cohering in pairs under the upper lip, cells divaricate. Stigma 2-lamellate. Capsule septicidal, valves separating from the placentiferous septum. Seeds rugose. Species about 30, chiefly tropical Asiatic.” “Ilysanthes Rafin.: Annual, glabrous, slender marsh plants. Leaves opposite, entire or toothed, parallel-nerved. Flowers small, axillary, or the uppers racemose; pedicels ebracteolate, often reflexed in fruit. Sepals narrow. Corolla tube cylindric; upper lip erect, broad, concave, notched.; lower larger, spreading, 3-lobed. Stamens, 2 upper alone perfect, subincluded, filaments filiform; anthers meeting, cells divaricate; staminodes 2, 2-lobed, one lobe glandular obtuse, the other glabrous of various shapes. Stigma 2-lamellate. Capsule septicidal; valves entire, separating from the placentiferous septum. Seeds numerous, rugose.— Species 8, N. American, S. African, Indian and one S. American. The staminal characters separating Ilysanthes from Vandellia are not satisfactory; I should prefer that of the nervation of the leaves, which would remove Vandellia erecta (which is hardly recognizable from I. parviflora) to this genus.” 71
  • 80. At species level, he writes: “9. V. erecta, Benth. Scroph. Ind. 36 and in DC X. 415. erect, quite glabrous, leaves sessile elliptic or oblong obtuse 3-5 nerved entire or obscurely crenate, pedicels axillary longer than the leaves and sepals, sepals lanceolate or linear rather shorter than the ovoid-orbicular capsule. Maxim. Diagn. Pl. Nov. Dec. XIX 413. V. pyxidaria Maxim. l.c. 419; [...] Tittmannia erecta Benth in Wall. Cat. 3947. Lindernia pyxidaria, All.; Benth. in DC. Prodr. x. 418; [...] G. integrifolia Roxb. Fl. Ind. i. 137, and Ed. Carey and Wall. i. 138.—Rheede Hort. Mal. ix. t. 57. [...] Branched from the base; branches divaricate, not rooting [...]. Leaves [1.3—1.9 cm]. Pedicels very slender, usually twice as long as the leaves. Sepals [3—4 mm] long, rather obtuse. – Closely resembles Ilysanthes parviflora. So common a plant as this is cannot have escaped the notice of Roxburgh, and I think it is his Gratiola integrifolia 22. Maximovicz has discussed at length the variations exhibited by this plant, and its passage into the northern Lindernia pyxidaria, in which the flowers are sometimes cleistogamous, the filaments straight and sometimes not appendaged. The Indian forms should be carefully studied. 2. I. parviflora, Benth. in DC. Prodr. X 419; erect, simple or diffusely branched, leaves sessile ovate elliptic or linear-oblong usually entire 1—5 nerved, pedicels axillary and in terminal racemes filiform, corolla twice as long as the calyx, fruiting sepals shorter than the ovate-oblong acute capsule. Gratiola parviflora Roxb. Pl. Corom. iii.3, t. 204, and Fl. Ind. i. 140, and Ed. Carey & Wall i. 141. Bonnaya parviflora, Benth in Wall Cat 3867 and Scroph. Ind. 34. Throughout India; in wet places, from the Punjab to Assam, Bengal and the Deccan peninsula (not seen from Ceylon)-Distrib. Siam, Trop Africa. Very near to I. hyssopioides, but a shorter stiffer more branched plant, more opiously flowering, with usually much shorter pedicels, and white or blue flowers not half the size. –This differs from Vandellia erecta in the reduction of the shorter stamens, and in the longer acute capsule. [from Fl. Brit. Ind. CIII Scrophularineae (J.D. Hooker); p. 283/284]Hooker pointed out a second character, i.e. the longer, acute capsule in L. parviflora. Itbecomes clear that Hooker would rather move Vandellia erecta Benth. to the genusIlysanthes, [i.e. Ilysanthes erecta (Benth.) Hooker]. I agree with Bentham that thestaminal characters are not sufficient to distinguish L. parviflora and L. procumbens.However, I must again draw attention to the lack of material available for this study.This is the reason why the following characters to distinguish the two species must beviewed as a hypothesis still in need of testing. Corolla size in L. parviflora is known tovary (Blatter & Hallberg 1918). In comparing some specimens determined as L.procumbens from other areas, I have observed slight differences in capsule shapes.Those of L. procumbens are more ellipsoid. Another character could be the manner in22 A specimen determined Gratiola integrifolia Roxb. was recovered at E. It indeed resembles (and mightbe synonymous with) Vandellia erecta Benth. It is from India. 72
  • 81. which the style breaks off slightly above its base (leaving the first millimeter attached),and forming a white ring at the base, i.e. around the apex of the capsule. The seeds in L.procumbens may prove a good character to separate it from L. parviflora, whose seeds,in my preliminary study, were half the size.Lindernia parviflora (Roxb.) Mukerjee has been treated as distinct from L. procumbens(Krock.) Philcox since it was described (1820). It was, however, explicitly treated asconspecific with L. hyssopioides (L.) Haines by Paul (1978) for Bihar and Orissa, ataxon that appears very distinct from L. parviflora in Indian collections. As discussedabove, it has been impossible to find strong distinguishing characters between L.procumbens and L. parviflora with the material available for this study. It is howeverwell-known, that its synonymy with and affinity to close potential relatives, e.g., L.microcalyx (Pennell) De Wit or Gratiola grandiflora Retz. is extremely doubtful (vander Vlugt 1985). The best solution to the question lies in collecting fresh material, andspecifically, to dissect, e.g., fifty, corollas from several populations to see whetherintraspecific variability occurs.L. procumbens has been included in the key, but the material from which the key wasconstructed was doubtful: During preliminary curation, both species, L. procumbens and L.parviflora were thought to form one group. The specimens of which a flower could beretrieved and dissected all proved to be a typical Ilysanthes sensu Rafinesque with twofertile stamens. The desription included in the key was made from the Wallich specimen atE, but that specimen also did not flower. Interestingly, a publication on the identity of L.pyxidaria (Simon & Geissert 1984) pointed out that the plant illustrated by von Lindernhad its fruits more or less developed, but lacked flowers altogether.5.2.3 The identity of L. indet-aPSW 4476 from (E) and (BM), proved to be distinct from all other specimens (Fig. 5.6).It is represented by one collection of 11 plants from Giri Khola in western Nepal at2500 m, the highest altitude recorded for collections of this genus in Nepal. ThisNepalese species was found to most probably be cleistogamous, since its corolla is 73
  • 82. almost completely closed. It could either be a cleistogamous form 23 of another speciesof Lindernia, its affinities not having been identified, or it could be a separate species. Ihave preliminarily referred to it here as Lindernia indet-a. While it is quite possible thatthis species has been described already, and lack of time during this research is to beblamed for it not having been properly identified, it is as feasable that we are dealingwith a new species. However, a similar individual was collected in Japan (MiyoshiFuruse No. 44991) from 90 m. a. s. l. on 29 Sept. 1966 (K). Lindernia indet-a (Fig. 5.6)was included in the key, because the collection itself consisted of plenty individualplants.If monophyly of Lindernia in Asia was ever tested, it would be interesting to investigatethe question whether Lindernia indet-a shares a most recent common ancestry distinctfrom the remaining Nepalese taxa. As to morphological characters, Lindernia indet-aappears to have strong affinities with the L. dubia complex known from both Americaand Europe. The phenotype of L. indet- a looks similar to, e.g., that of the Claytonspecimen given to Linnaeus from Virginia (Chapter 1.3, Fig.1.a, specimen on the left –L. dubia according to Linnaeus personal copy of Sp. Pl.). Given the facts that (1)Lindernia are weeds, and (2) seed dispersal in Lindernia may very well at times beaffected by migrating aquatic birds, and (3), known disjunctions between Asia and theEastern seaboard of the United States exist (Xiang et al. 1998), the origin of this speciescan only be speculative. Furthermore, several specimens from Europe were found toresemble, but not match Lindernia indet-a. The following species names bearresemblance with L.indet-a: Gratiola inaequalis Walter – America; Lindernia dubia(L.) Pennell sensu lato (1753)—based on Clayton specimen from Virginia; Gratiolaanagallidea Michaux (1803); Lindernia dilatata var. anagallidea H.L. Muehl.; L. dubiavar. anagallidea (Michx.) Cooper; L. attenuata H.L. Muehl.; L. dubia var. dubia (L.)Cooper??; L. pyxidaria ‘var’. major Pursh; Ilysanthes gratioloides (L.) Bentham.Its affinities with the American and the European complex are yet to be elucidated. Forthis process, it would be important to follow up on recent research by Berger & Elisens,who gave a seminar on the “Reevaluation of the Lindernia dubia species complex,” in 23 Fischer (1995) reported a cleistogamous form of L. nummulariifolia (D.Don) Wettst. from Madagascar; it had been described by Bonati as a species of the genus Bryodes, namely B. pierrii. 74
  • 83. which they reported a broad range of morphological variation within the complex, andfound the 4 varieties treated by Lewis (2000) should be treated as one species (Abstractdetail from Botany 2005 Systematics Conference).Since the specimen had initially been determined as L. procumbens (Krock.) Philcox,and seen by the latter, a short comparative morphological study was undertaken,measuring and averaging the pedicel length from three flowering specimens of L.parviflora/L. procumbens, and those of L. indet-a. The following differences could beascertained: The average pedicel length to leaf length ratio in L. parviflora/L.procumbens was 1.7 (ranging between 1.5 and 2); that of L. indet-a was 0.6 (rangingbetween 0.4 and 0.8). Further, measurements were taken to describe the manner ofinflorescence. The obviously stronger decrease in leaf length toward the apex of theplant in L. parviflora/L. procumbens was plotted against that of the other species (Fig.5. 5). In the graph, LL av (p) stands for leaf length average in L. parviflora/L.procumbens, and (a) for that in L. indet-a 25 Leaf length [mm] 20 15 LL av (p) 10 LL av (a) 5 0 1 2 3 4 5 6 7 8 9 10 Node number from Base to ApexFigure 5.5: Depiction of average leaf length (measured from three flowering specimens of the samespecies, respectively) from the base of the plant (left) to the apex (right). LL av (p) denotes leaf lengthaverage of L. parviflora/ L. procumbens? (Krock.) Philcox; LL av (a) denotes leaf length average ofLindernia indet-a. Nodes are numbered from 1 (base) to 9 (apex). 75
  • 84. Fig. 5.6: Lindernia indet-a – BM collection. The geographic locality is 29º 14’ N, 82 º 07’ E, altitude is ca. 2500 m. 76
  • 85. 5.2.4 Lindernia indet-bThe potentially 15th Nepalese species Lindernia indet- b, is represented by a Buchanan-Hamilton collection at BM. It was determined by Tsoong to be L. nummulariifolia, but theleaf shape does not correspond to that known from other Nepalese collections. It is to befound in the L. nummulariifolia folder in BM. A digital image, cropped and pasted, can beviewed in Figure 5. 7 below. Unfortunately, the original handwriting on the bottom was cutoff during photography of the specimen. Figure 5. 7: Specimen Lindernia indet-b. collected in Nepal by Buchanan-Hamilton. It is not sufficiently known to the present author to have included it in the key to the species. The conspicuously attenuate leaf base suggests affinity with L. urticifolia (Hance) Bonati in Fl. Indo-Chine 4: 421.1912. 77
  • 86. 5.2.5 The type of L. micrantha D.DonThe type was not found in the collection at the BM. I chose to stick with the name L.micrantha as proposed by Hara (1980) in Journ. Jap. Bot. 55 (11): 325. In synonymy withthis type he cited Wallich 3951 B as the lectotype of Vandellia angustifolia Benth. (K-W,Figure 5. 8 a). I have reason to believe that Hara understood the nature of Nepalesematerial, and that the type may just be temporarily lost. The last time it was cited from theBM as “Wallich s.n. in type folder” was by Press and Shresta in their Catalogue of Typespecimens from Nepal (2000b). Fig. 5.8 a: Lectotype of V. angustifolia Benth. 78
  • 87. 5.2.6 The type of Lindernia oppositifolia (Retz.) MukherjeeLindernia oppositifolia (Retz.) Mukerjee is cited by Hara in the Enumeration of theFlowering Plants of Nepal [EFPN], but no specimen was recovered from Nepal duringthis revision. It is not clear whether the citation made by Yamazaki (1977) in Journ. Jap.Bot 52 (8) for Nepal is based on the misidentified specimen from Tokyo Institute cited inthe EFPN, or whether other specimens have been collected in Nepal and been depositedelsewhere than the BM, Kew, or E. The König type of the species is from Tranquebar (Fig.5.8 b). The specimen from K. M. Matthew P 282 (1981) is described on the sheet as“leaves green, flowers white with pink centre. Locally abundant. On marshy ground withinscrub jungles” (Fig. 5. 8 c). Fig. 5.8 c: Lindernia oppositifolia Retz. K.M.Matthew P282 (1981); India. Fig. 5.8 b: Type of Gratiola oppositifolia Retz. (LD) Digital image of photograph. 79
  • 88. 6. SUMMARYThis project has only scratched the surface of work still waiting to be done on this genus.However, the account created for the Flora of Nepal is an immediate result that should notbe underestimated, despite missing data. The revision demonstrated the importance ofalpha taxonomy and its rewards to science: The use of correctly typified nomenclature isfundamental for plant science on all levels.To understand Lindernia in its full geographical extent, a combined morphological andmolecular phylogenetic study would be very useful. The morphological charactersevaluated in this study should be coded rigorously. This analysis should includemorphometric coding of the corolla shape, a character for which the current availability ofherbarium material will not suffice. A research study should be designed to test thevariability of corolla and androecium particularly to resolve the questions pertaining to theLindernia parviflora/ L. procumbens complex. This would involve sampling of severalpopulations across the whole of several flowering seasons across national boundaries.Further, the knowledge about phytochemical compounds in this group should be extendedthrough, e.g., further chromatography of flavonoids, to include these characters in thephylogenetic analysis, because its application has proven useful in delimiting taxa (DiazMiranda 1977). While it would be favourable to include all geographic regions in thephylogenetic study, a recollection of specimens known from Nepalese herbarium materialwith the simultaneous production of detailled line-drawings may be treated as a priority todirectly benefit completion of the Lindernia section in the Flora of Nepal. 80
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  • 92. _____. 1771. Mantissa Plantarum: 251-252 In Carl Linnaeus Mantissa Plantarum 1767 & 1771, ed. J. Cramer. 1961: 87-88. Codicote/Herts, New York: Wheldon & Wesley, Ltd. and Hafner Publishing.Martinez-Ortega, M. M. & E. Rico. 2001. Seed morphology in Veronica. Plant Systematics and Evolution 227 (3-4): 15—32.Maximovicz, C. J. I. 1874. Diagnoses Plantarum Novarum Japoniae et Mandshuriae 19: 407—422.Merrill, E. D. 1912. Notes on the Flora of Manila with special reference to the introduced element. The Philippine Journal of Science 7 (3): 145—209._____. 1918. Scrophulariaceae. In Species Blanconae: 347—349.Mill, R.R. 2001. Scrophulariaceae. In Flora of Bhutan, ed. Grierson 2 (3): 1122—1125.Mueller, F. v. 1867—1868. Fragmenta phytographiae Australiae 6: 101—105.Mukerjee, S. K. 1945. A revision of the Indo-Burmese species of Lindernia Allioni. The Journal of the Indian Botanical Society (Formerly the J. of Ind. Bot.) 24: 127—134.Myase, T., T. Andoh & A. Ueno. 1995. Linderniosides A and B, oleanane saponins from Lindernia pyxidaria. Phytochemistry 40 (5): 1499—1502.Olschki, L. S. 2004. (ed). Le opere minori di Carlo Allioni. Dal “Rariorum Pedemontii stirpium” all’ “Auctarium ad Floram Pedemontanam.” A cura di R. Caramiello e G. Forneris. 24Oxelman, B., P. Kornhall, R. G. Olmstead, B. Bremer. 2005. Further disintegration of the Scrophulariaceae. Taxon 54 (2): 411—425.Paul, S. R. 1878. The genus Lindernia All. (Scrophulariaceae) in Bihar and Orissa. J. Ind. Bot. Soc. 57. Supplement: 57, X.2.Pennell, F. W. 1935. Scrophulariaceae of eastern temperate North America. Monogr. Acad. Nat. Sci. Philadelphia 1: 137._____. 1936. New and Noteworthy Papuan Scophulariaceae. Brittonia 2: 181—182._____. 1939. A summary of the Scrophulariaceae of New Guinea. J. Arn. Arb. 20: 75—84._____. 1943a. A second summary of the Scrophulariaceae of New Guinea. J. Arn. Arb. 24 (3): 243—256._____. 1943b. The Scrophulariaceae of the Western Himalayas. The Academy of Natural Sciences of Philadelphia. Monographs 5: 20—32. 24 This book is in Italian. It is available for loan at Kew. The publishing house gives the following English description: [...] The present volume reprints four studies which Allioni published before the appearance of the Flora [pedemontana] and a fifth which came later. [...] demonstrate his early acceptance of Linnaean binomial nomenclature, his interests in teaching, his belief in the importance of botanical gardens and herbaria, and his interest in artistic depiction as an aid to scientific description. 84
  • 93. Philcox, D. 1963—1964. Typification of Ruellia antipoda Linn. and clarification of its synonymy. Kew Bulletin 17: 481—484._____. 1965. A new name for the generitype of Lindernia. Taxon 14 (1): 30._____. 1968. Revision of the Malesian species Lindernia All. Kew Bulletin 22 (1): 1—72.Press, J. R. & K. K. Shresta. 2000a. Collections of flowering plants by Francis Buchanan- Hamilton from Nepal, 1802 – 1803. Bull. Nat. Hist. Mus. Lond. (Bot.) 30 (2): 101—130._____ & _____. 2000b. Catalogue of type specimens from Nepal. The Natural History Museum: London._____, _____ & D. A. Sutton. 2000. Annotated checklist of the flowering plants of Nepal. The Natural History Museum: London.Rafinesque. 1820. Annals of Nature: 13.Rahmanzadeh, R., K. Müller, E. Fischer, D. Bartels, and T. Borsch. 2004. The Linderniaceae and Gratiolaceae are further lineages distinct from the Scrophulariaceae (Lamiales). Plant Biol. 7: 67—78.Reichenbach. 1824. Iconographia botanica exotica: 1: 26—27.Retzius, A. J. 1790. Fasciculus Observationum Botanicarum 4: 8.Rheede van, H. 1689. Hortus Malabaricus 9.Roxburgh, W. 1798. Plants of the Coast of Coromandel 2: 29—32._____. 1819. Pl. Corom. 3: 3._____. 1820. Flora Indica 1: 136 – 143._____. 1832. Flora Indica, ed. Carey. 3: 96.Rumphius, G. E. 1747. Herbarium Amboinense (5): 460 t. 170 f. 2.Scotland, R. W., R. G. Olmstead, J. R. Bennett. 2003. Phylogeny reconstruction: The role of morphology. Syst. Biol. 52 (4): 539—458.Sealy, J. R. 1956. The Roxburgh Flora Indica Drawings at Kew. Kew Bull. 11 (2): 297 —348.Segarra, J. G. & L. Mateu. 2001. Seed morphology of Linaria species from eastern Spain - identification of species and taxonomic implications. Bot. J. Linn. Soc. 135: 375 —389.Simŏes, C. M. O., M. Amoros, and L. Girre. 1999. Mechanismal antiviral activity of triterpenoid saponins. Phytotherapy research 13 (4): 323—328.Simon, M. & F. Geissert. 1984. Lindernia procumbens (=L. pyxidaria), une plante alsacienne. Bull. Soc. Bot. Centre-Ouest 15: 27—34. 85
  • 94. Sivarajan, V. V. and P. Mathew. 1983. The genus Lindernia All. (Scrophulariaceae) in India. J. Bomb. Nat. Hist. Soc. 80 (1): 131—140.Sprengel, K. 1824. Caroli Linnaei Systema Vegetabilium 1: 41—42_____. 1825. Caroli Linnaei Systema Vegetabilium 2: 800—801.Stearn, W. T. 1972. The generic name Hornemannia and its diverse applications. Taxon 21 (1): 105—111._____. 1992. Botanical Latin (4th ed.) Devon, Newton Abbott.Thunberg, C. P. 1794. Prodromus Plantarum Capensicum (12): 177.Uchino, A. & Watanabe, H. 2002. Mutations in the acetolactate synthase genes of sulfonylurea-resistant biotypes of Lindernia spp.. Weed Biology and Management 2 (2): 104—109.Urban. 1884. Studien ueber die Scrophulariaceen-Gattungen Ilysanthes, Bonnaya, Vandellia und Lindernia. In Berichte der deutschen botanischen Gesellschaft 2: 429—442.Vahl, M. 1804. Enumeratio Plantarum 1: 88—100.Van der Vlugt, P. J. 1985. Lindernia. Aquarium:172—176.Walker-Larsen, J. and L. D. Harder. 2000. The evolution of staminodes in angiosperms: patterns of stamen reduction, loss, and functional re-invention. Am. J. Bot. 87 (10): 1367—1384.Wettstein von, R. 1891. Scrophulariaceae. In Die natürlichen Pflanzenfamilien. 1895. ed. A. Engler and K. Prantl. 4 (3B): 69—80.Willdenow, C. L. 1797. Species Plantarum: 103—105.Xiang, Q-Y., D. E. Soltis & P. S. Soltis. 1998. The Eastern Asian and Eastern and Western North American Floristic disjunction: Congruent phylogenetic patterns in seven diverse genera. Molecular phylogenetics and evolution 10 (2): 178—190.Yamazaki, T. 1954. Notes on Lindernia, Vandellia, Torenia and their allied genera in Eastern Asia 1. Journ. Jap. Bot. 29: 299—306._____. 1955. Notes on Lindernia, Vandellia, Torenia and their allied genera in Eastern Asia 2. Botanical Magazine Tokyo 68: 14—24._____. 1955. Notes on Lindernia, Vandellia, Torenia and their allied genera in Eastern Asia 3. Journ. Jap. Bot. 30 (6):_____. 1966. Scrophulariaceae. In The Flora of Eastern Himalaya, ed. H. Hara: 285—295._____. 1977. Lindernia in Japan and Taiwan. Journ. Jap. Bot. 52 (8): 253—256. 86
  • 95. _____. 1978. New or Noteworthy plants of the Scrophulariaceae from Indo-China (1). Journ. Jap. Bot. Vol. 53 (1): 1—11._____. 1978. New or Noteworthy plants of the Scrophulariaceae from Indo-China (2). Journ. Jap. Bot. Vol. 53 (4): 97- 106._____. 1980. New and Noteworthy Plants of Scrophulariaceae from Indo-China (6). Journ. Jap. Bot. Vol. 55(11):328 – 336 (contd. from. 55: 204 – 208)._____. 1981. Revision of the Indo-Chinese species of Lindernia All. (Scrophulariaceae). Journal of the Faculty of Science: University of Tokyo._____. 1983. New or Noteworthy plants of the Scrophulariaceae from Indo-China (7). Journ. Jap. Bot. Vol. 58 (1):14- 17._____. 1983. New or noteworthy plants of the Scrophulariaceae from Indo-China (8). Journ. Jap. Bot. Vol. 58 (2):46 – 50._____. 1985. Revision of the Genera Limnophila and Torenia. J. Fac. Sci. Univ. Tokyo 3 (13): 603—625._____. Scrophulariaceae. In Flora of Thailand 5 (2): 178—202. 87
  • 96. APPENDIX (1): ACCOUNT OF THE GENUS FOR THE FLORA OF NEPALLindernia All., Misc. Taur. 3 (1): 178—185, t. 5, f. 1. 1766.Vandellia P. Browne ex L., Syst. Nat. ed. 12 (2): 384, 422. 1767; Mant. (12): 89. 1767.Anagalloides Krock., Fl. Siles 2 (1): 398. 1790.Ilysanthes Rafin., Annals of Nature: 13. 1820.Bonnaya Link & Otto, Icones Pl. Select. 10: 25, t. 11. 1828.Tittmannia Rchb., Icon. Bot. Exot. 1: 26, t. 38. 1824 (nom. rej.).Annual herbs, stems quadrangular, erect or procumbent, sometimes rooting at nodes,glabrous or hairy on angles. Leaves opposite; lamina sessile or petiolate, linear to orbicular,obscurely crenate to serrate, sessile or petiolate. Flowers solitary in the axils of the leaves,or in terminal racemes sometimes appearing very condensed; occasionally inflorescenceintermediate. Pedicels slender or robust, shorter or longer than leaves, extending in fruit.Calyx persistent, glabrous or hairy, more or less equally 5-lobed to 5-dentate or unequally5-lobed and bilabiate; lobes lanceolate, imbricate in bud,; calyx lobes 1—5-veined, marginssmooth or ciliate, apex acute or obtuse, smooth or ciliate. Corolla 2-lipped, lower lip 3-lobed, longer, wider; upper lip 2-lobed, smaller, much more narrow, about same size as oneof the three anterior lobes. Androecium didynamous, anterior filaments longer, arching toposterior side, with club-shaped geniculum, posterior filaments straight, withoutgeniculum; anthers connivent or free, either all fertile, or posterior fertile and anteriorsterile or anterior reduced to staminodes; stigma enlarged, flattened, sometimes 2-lobed.Capsule unilocular, bi-valved, globose, ovoid, or linear, enclosed by or to 2.5 times as longas the calyx, septicidal. Seeds numerous, alveolate or not, appendaged or not.Notes: In the following species descriptions, I refer to the anterior and posterior pairs ofanthers; each pair always consists of two, adding up to four anthers. If all four are fertile, Iwrite “all stamens fertile.” All here refers to the number “4.” If only the posterior pair isfertile, I write “anterior stamens sterile” or “anterior staminodes,” and “posterior stamensfertile,” herewith saving to repeat the superfluous number “2.” The species distributionlisted below may not be complete. Additional voucher specimens recovered in the futureshould be added. 88
  • 97. KEY TO THE SPECIES OF LINDERNIA1A Calyx with lobes at least 1/3 of total calyx length or bilabiate .......................….21B Calyx cup-like with teeth up to 1/4 of total calyx length…….….…2. L. crustacea2A Inflorescence a terminal raceme, sometimes very condensed and appearingumbellate; upper bracts differently shaped than leaves below, marking an abrupt transitionto the flowering part of the plant..........................................................................................32B Flowers solitary, axillary; upper leaves subtending pedicels similarly shaped toleaves below and, if smaller, decreasing in size gradually toward the apex of the plant........................................................................................................….………...…..............83A Capsules ≥ 10 mm, cylindrical, much longer than the calyx.................................43B Capsules ≤ 5 mm, ovoid, about the length of the calyx...……......…....................64A Lamina serrations usually exceeding 14 per side, except on round leaves, where itcould be as few as 10 ...........................................................................................................54B Lamina serrations usually not more than 6--8 per side, except on very long, linearleaves, where it could be as many as 10.............................................................1. L. antipoda5A Leaves sessile or subsessile, marginal serrations extended into a long bristle; plantnot rooting at the nodes...........................................................…………...………5. L. ciliata5B Leaves distinctly petiolate, marginal serrations not extended into a long bristle;plant rooting at the nodes ................................................................................6. L. ruellioides6A Lamina hairy above and at least on veins below; calyx hairy.............................76B Lamina and calyx glabrous...............................................................11. L. multiflora7A Plant erect, not rooting at the nodes; length: width ratio of lamina > 2.2, bracts ≥ 3mm, calyx > 4 mm, 2 stamens fertile, 2 sterile................................................13. L. hookeri7B Plant diffusely branching, rooting at the nodes; length: width ratio of lamina < 2,bracts ≤ 2 mm, calyx < 4 mm, all 4 stamens fertile..............................................8. L. viscosa 89
  • 98. [inflorescence solitary, axillary]8A Stem densely hairy...............................................................................................98B Stem glabrous ....................................................................................................109A Lamina hairy, entire or with 3—4 serrations per side; leaf size markedly decreasingtoward apex of plant; capsule shorter than calyx...............................7.L. pusilla9B Lamina glabrous, 6—10 distinct serrations per side; leaf-size not markedlydecreasing toward apex of plant; capsule longer than calyx................12. L. nummulariifolia10A Lamina venation pinnate or uni-veined.............................................................1110B Lamina three to five –veined from the base......................................................1211A Lamina uni-veined, length: width ratio > 5, all 4 stamens fertile.....10.L. micrantha11B Lamina venation pinnate, ratio <5, anterior stamens sterile................3. L. anagallis12A Leaves subtending pedicels markedly decreasing in size toward the apex of theplant, pedicels 1.5—2 times the length of the leaf .........................................................1312B Leaves subtending pedicels not decreasing in size much toward the apex of theplant, pedicels more or less the length of the leaves.......................................14. L. indet-a13A Stamens 4, all fertile.......................................................................4. L. procumbens13B Stamens 2 posterior fertile, 2 anterior sterile....................................9. L. parvifloraSPECIES DESCRIPTIONS1. Lindernia antipoda (L.) Alston, in Trimen, Handb. Fl. Ceylon 6 Suppl: 214. 1931.Holotype BM: Sloane room; Hermann collection!Ruellia antipoda L., Sp. Pl. 2: 635. 1753.—Lectotype: Hermann 235, Ceylon (BM!).Gratiola grandiflora Retz., Obs. B. 4: 8. 1786.—Holotype: König s.n. (LD!).Gratiola veronicifolia Retz., Obs. B. 4: 8. 1786.—Holotype: König s.n. (LD!).Gratiola verbenaefolia Colsm., Prodr. Desc. Grat.: 8. 1786.—Holotype: König s.n.: Habitatin India orientali. (LD!).Bonnaya veronicaefolia (Retz.) Spreng., Syst. Veg.1:41. 1824.Bonnaya veronicifolia var. verbenaefolia (Colsm.) Hook.f., Fl. Br. Ind. 4: 285. 1884.Bonnaya veronicifolia var. grandiflora (Retz.) Hook.f., Fl. Br. Ind. 4: 285. 1884. 90
  • 99. Ilysanthes veronicifolia (Retz.) Urb., Ber. Deutsch. Bot. Ges. 2:436. 1884.Lindernia verbenaefolia (Colsm.) Pennell, Monogr. Acad. Sci. Philad. 5:31. 1943b.Lindernia anagallis var. grandiflora (Retz.) Mukerjee, J. Ind. B. S. 24: 133. 1945.Vandellia antipoda (L.) Yamazaki, J. Jap. B. 30: 177. 1955.Hooker in Fl. Br. Ind. 4: 285 (1884); Hara et al. in Enum. Fl. Pl. Nep. 3: 116 (1982);Deyuan et. al. in Fl. Chin. 18: 36 (1998); Press et al. in Ann. Check. Fl. Pl. Nep. 294(2000); Mill in Fl. Bhutan 2(3): 1125 (2001).Annual to 30 cm, stem erect or occasionally decumbent and rooting at nodes, glabrous, to 2mm diameter. Lamina uni-veined, sessile, narrowly elliptic to oblanceolate, to 5.5 X 1 cm(length: width ratio 3—5), glabrous, base attenuate, apex acute, lamina margin serrate,ciliate. Inflorescence a terminal raceme, bracts ca. 5—6 X 1 mm; pedicels shorter thanleaves. Calyx glabrous, to 6 mm in flower, to 7 mm in fruit; calyx lobes to 5 mm in flower,to 6 mm in fruit, calyx lobe margin ciliate. Corolla white, blue, or pale purple, 10—12 mm,central lobe of lower lip ca. 1.5 X 4.5—8 mm, upper lip ca. 2 mm; anterior staminodesgeniculate, bright yellow; posterior filaments 1.3—1.5 mm, anthers fertile, 1.1—1.2 mm.Ovary ca. 1.2—1.5 mm, style 5—6 mm. Capsule green or purplish, elongate, 13—16 X1—1.5 mm. Seeds 0.3 x 0.2 mm, alveolate.Flowering Time: March—December.Fruiting Time: March—December.Habitats: On (sandy) seasonally flooded plains and river banks.Altitudinal Range: 100—2300 m.Distribution: W, C, and E Nepal; [Deccan; Tibetan Plateau; W. Himalaya; E.Himalaya; Assam-Burma; E. Asia; SE. Asia].Notes: The uni-veined lamina distinguishes this species from the at times(due to variation in lamina margin serration) similar appearing L. ruellioides.Specimens seen:DNEP2 A50 (E) Gardner, P.C. 157 (BM)DNEP2 A55 (E) Hara et al. 1486 (BM)DNEP2 B40 (E) Kanai et al. 1456 (BM)DNEP2 B44 (E) D.H.Nicolson 2824 (BM)Gardner, P.C. 156 (BM) Nicolson, D.H. 3060 p. p. (mixed) (BM) 91
  • 100. Norkett 8066 (BM) SSW 5664 (E; BM)Norkett 9165 (BM) SSW 5831 (BM)Noshiro et al. 9760068 (BM) SSW 7045 (BM)B.D. Pande 23 (BM) SSW 7181 (BM)PSW 3664 (BM) Wallich 3857F (K-W)PSW 5664 (E) Wallich 3858C (K-W)J.D.A. Stainton 1538 (E) K. Wesche s.n. (E)SSW 37 (BM) Wraber 34396 (BM)SSW 2778 (BM)2. Lindernia crustacea (L.) F. Muell., Syst. Cens. Austral. Pl. 1: 97. 1882. Lectotype: LINN785.3!Capraria crustacea L., Mant. 1: 87. 1767.Antirrhinum hexandrum Forst.f., Prodr.: 43. 1786.—Type: Tahiti: Forster s.n. (K)Gratiola lucida Willd., Sp. Pl. 1: 103. 1797.—Holotype: (B!)Torenia flaccida R. Br., Prodr.: 440. 1810.—Type: Australia. Banks & Solander s.n. (BM!)Hornemannia ovata Link & Otto, Ic. Pl. Select. 9: t. 3. 1820.—Type: Illustration, l.c.Gratiola aspera Roth., Nov. Pl. 2. 1821.—Type: ex India orientalis Heyne (K);Tittmannia ovata (Link & Otto) Rchb., Iconogr. Pl. Exot. 1: 27. 1823.—Type: Illustration.Mimulus javanicus Bl., Cat. Gew. Buit.: 83. 1823.—Type: (L!)Torenia minuta Bl., Bijdr.: 755. 1825.—Holotype: Herb. Hort. Bot. Bog. [L0003521]Blume 758, Java, Buitenzorg [..] Graze (L!); Isotype: Herb. Hort. Bot. Bog. [L0003520]Archip. Ind. (L!)Morgania lucida (Willd.) Spreng., Syst. Veg. 2: 803. 1825.Morgania aspera (Roth) Spreng., Syst. Veg. 2: 803. 1825.Torenia crustacea (L.) Cham. & Schlechtend., Linnaea 2: 570. 1827.Torenia alba Buch-Ham. ex Wall. Cat. 3961, nom. nud. 1831.—Type: Buch-Ham. (E!)Torenia lucida (Willd.) Buch-Ham. ex Wall. Cat 3962, nom. nud. 1831. (K-W!)Torenia varians Roxb., Fl. Ind., ed Carey 3: 96. 1832.—Holotype: Roxburgh Icones 1269(K!).Vandellia crustacea (L.) Benth., Scroph. Ind.: 35. 1835.Vandellia varians (Roxb.) G. Don, Gen. Syst. 4: 549. 1838.Vandellia alba Benth., Scroph. Ind.: 35. 1835.—Type: Wallich 3961 (K-W!).Vandellia minuta (Bl.) Miq., F. Ind. Bat.2: 691. 1857.Pyxidaria crustacea (L.) Kuntze, Rev. Gen.2: 464. 1891. 92
  • 101. Lindernia minuta (Bl.) Koord., Exk. Fl. Java 3: 178. 1912.Vandellia crustacea Benth. ex Hook.f. in Fl. Br. Ind. 4: 279 (1884); Hara et al. in Enum.Fl. Pl. Nep. 3: 116 (1982); Deyuan et. al. in Fl. Chin. 18: 32 (1998); Press et al. in Ann.Check. Fl. Pl. Nep. 294 (2000); Mill in Fl. Bhutan 2(3): 1124 (2001).Annual to 20 cm, stem winged, erect or occasionally procumbent and rooting at nodes,branching, occasionally sparsely pubescent at angles, to 0.8 mm diameter. Laminapinnately nerved, petiolate, ovate, ovate--oblong, or broadly ovate, to 15 X 9 mm, uppersurface glabrous, lower surface with sparse short hairs on protruding midrib, basesubtruncate or rounded, apex obtuse, margin sparsely ciliate. Inflorescence solitary andaxillary, or subracemose; pedicels 8 to 30 mm, longer than leaves; bracts to 8 X 5 mm.Calyx 3—4 mm in flower, 3.5—4.5 mm in fruit, shortly hairy; calyx lobes short, to 1 mmin flower, to 2 mm in fruit. Corolla blue or purple, ca. 6 mm, pubescent inside. Anteriorfilaments geniculate, stamens all fertile. Capsule ovoid, ca. 3 X 2 mm. Seeds ca. 0.4 X 0.25mm, alveolate, filiform appendage.Flowering Time: June—December.Fruiting Time: June—October.Habitats: On grassy or stony, dry or moist places, terraced banks, in sunand shade.Altitudinal Range: 200—1900 m.Distribution: W, C, and E Nepal; [Deccan; Tibetan Plateau; W. Himalaya; E.Himalaya; Assam-Burma; E. Asia; SE. Asia].Notes: This is so far the only Nepalese Lindernia I have seen with acalyx so shortly lobed, it would better fit in with Torenia, was it not for its much smallercorolla. According to “Checklist of medicinal plants in Southeast Asia”, used “againstboils, and itchiness. In Indonesia, cures scable: kudis.”Chromosome number: 2n = 42 (Darlington & Wylie 1955)Specimens seen:Buch-Ham 1390 (Wall.Cat. 3961) PSW 5624 (BM)Norkett 8082 (BM) SSW 4103 (E; BM)Noshiro et al. 9760521 (BM) SSW 6454 (E; BM)Noshiro et al. 9760067 (BM) SSW 7055 (E; BM) 93
  • 102. SSW 7578 (BM) L.H.J.Williams 326 (BM)Wallich 3945C (K-W) T. Wraber 36770 (BM)Wallich 3962 (E) T. Wraber 36785 (BM)Wesche No. 7/ 92 (BM)3. Lindernia anagallis (Burm.f.) Pennell, Journ. Arn. Arb. 24: 252. 1943. Holotype: Java,D. Kleinhof s.n.1759 (L)!Ruellia anagallis Burm.f., Fl. Ind. 135. 1768.—Holotype: Java, D. Kleinhof s.n.1759 (L)!Gratiola cordifolia Colsm., Prodr. Descr. Grat. 15: 1793.—Holotype: ‘in graminosishumidusculis Zeylon[a]e’ König s.n. (C!)Gratiola cordifolia Vahl, Enum.1: 97. 1804.—Holotype: ‘in graminosis humidusculisZeylon[a]e’ König s.n. (C!)Gratiola pedunculata R. Br., Prodr. Fl. Nov. Holl. 435. 1810.Bonnaya cordifolia (Colsm.) Spreng., Syst. Veg. 1: 42.1824.Torenia obtusa Bl., Bijdr.: 754. 1826.—Holotype: Java, locis humidis Provinciarumoccidentalium Javae Blume 1986 (L!)Tittmannia grandiflora Wall. ex Benth. 1831. Wall. Cat. No. 3949, nom. nud. (K!)Vandellia pedunculata Benth. Scroph. Ind. 1835.—Holotype: East Bengal Sylhet, Wall.Cat. No. 3949G (K!); isotype (K-W!)Vandellia cordifolia (Colsm.) G.Don, Gen. Syst. 4: 549. 1838.Vandellia roxburghii G. Don, Gen. Syst. 4: 549. 1838.Lindernia obtusa (Bl.) Koord., Exc. Fl. Java 3: 178. 1912.Lindernia cordifolia (Colsm.) Merr., Enum. Born., Pl.: 524. 1921.Vandellia pedunculata Benth. ex Hook.f. in Fl. Br. Ind. 4: 282 (1884); Hara et al. in Enum.Fl. Pl. Nep. 3: 116 (1982); Deyuan et. al. in Fl. Chin. 18: 35 (1998); Press et al. in Ann.Check. Fl. Pl. Nep. 294 (2000); Mill in Fl. Bhutan 2(3): 1123 (2001).Annual to 25 cm, stem simple or branched, to 1.4 mm diameter, glabrous. Lamina sessile tosubpetiolate, ovate to broadly deltoid, 10--25 X 6--12 mm, glabrous base truncate toattenuate, apex acute, margin distantly, ciliate or fused into rim. Inflorescence solitary,axillary, one or two flowers per node, pedicels 15—22 mm, often longer than leaves. Calyx5—6.5 mm in flower and fruit, glabrous, calyx lobes 4—5 mm in flower and fruit, marginand apex ciliate [or apex a white tip]. Corolla purplish pink and white, to 13 mm, central 94
  • 103. anterior lip lobe to 3.5 X 6 mm wide, posterior lip 2 mm, throat internally, tube externallyhairy; ovary 2 mm, style ca. 5 mm. Anterior filaments ca. 3 mm, geniculate, posteriorsubsessile in throat; anterior stamens sterile, subconnivent, anthers ca. 1 mm; posteriorstamens fertile. Capsule golden brown, elliptic to elongate, 7—10 X 1.5 mm . Seedsalveolate, appendaged.Flowering Time: March—late November.Fruiting Time: April—November.Habitats: Base of path, mixed sedge and herb vegetation. Moist terracesnear river, moist swampy fields, boggy ground; irrigated fields; by stream through fields.Altitudinal Range: 300 – 1300 m.Distribution: W, C, E; [Tibetan Plateau; W. Himalaya; E. Himalaya; Assam-Burma; E. Asia; SE Asia; Australasia].Notes: Margin serrate with 6 to 7 incisions of ca. 0.2 mm depth per side;trichomes on corollas only observed once. It is not certain whether this character existsthroughout the species.Specimens seen:DNEP 2B191 (E) SSW 2777 (E; BM)PSW 1833 (BM) SSW 5216 (E; BM)PSW 3648 (BM) SSW 7180 (BM)J.D.A.Stainton 65 (E; BM) Wallich 3949F (E; K-W)SSW 569 (E; BM) Wallich 3949 (K);4. Lindernia procumbens (Krock.) Philcox, Taxon 14(1): 30. 1965. Holotype: Fl. Siles. 2(1): t. 26. 1790. Illustration (E!).Anagalloides procumbens Krock., Fl. Siles. 2 (1): 398. 1790.—Holotype: Fl. Siles. 2: tab.26. 1790. Illustration (E!).Vandellia erecta Benth. Scroph. Ind. 36. 1835.—Type: Wallich 3947 (K-W!).Vandellia erecta Benth. ex Hooker in Fl. Br. Ind. 4: 282 (1884); Hara et al. in Fl. E. Him.288 (1966); Hara et al. in Enum. Fl. Pl. Nep. 3: 117 (1982); Deyuan et. al. in Fl. Chin. 18:35 (1998); Press et al. in Ann. Check. Fl. Pl. Nep. 294 (2000); Mill in Fl. Bhutan 2(3):1123 (2001). 95
  • 104. Annual, to 15 cm, stem winged, 0.5--1 mm diameter, glabrous. Leaves sessile, lamina 3—5-veined from the base, lanceolate, 8—16 X 3—7 mm, glabrous; base rounded or claspingthe stem, apex acute—acuminate, margin subentire to distantly serrate. Inflorescencesolitary, axillary, or subracemose; pedicels winged, 5—14 mm, often 1.5—2 times longerthan lamina; leaves subtending pedicels 3—10 mm long, 1—4 mm wide. Calyx to 3 mm infruit, glabrous; calyx lobes ca. 1.5 in fruit. Corolla not seen. Stamens all fertile. Capsulestriate, 2.5—3.X 1.8—2.2 mm wide, callous style base at apex of non-dehisced capsule.Seeds ca. 0.5 X 0.3 mm.Flowering Time: ??Fruiting Time: ??Habitats: ??Altitudinal Range: ??Distribution: Nepal. [E. Asia; China: Prov. Kirinensis (Mandshuria); Assam-Burma].This species has been reported from Nepal only through old collections, which didnot include any geographic information. It looks similar to L. parviflora, but has 4 fertileanthers. Corolla not seen; it may fall off shortly after collection (I would advise to make adrawing of it).Specimens seen:Nepal D. Wallich 1821 (K; E) Benth. in Wall. Cat 3947 B+C (K-W).5. Lindernia cilata (Colsm.) Pennell, Brittonia 2: 182. 1936. Holotype: König s.n., Java(C!).Gratiola ciliata Colsm., Prodr. Desc. Grat.: 14. 1793. Holotype: König s.n., Java (C!).Bonnaya brachiata Link & Otto, Ic. Pl. Select.: 25, t.11. 1828. Illustration (E!).Gratiola serrata Roxb., Fl. Ind., ed Carey & Wall. 1: 139. 1832.—Holotype: Icones 1515 (K!).Kyrtandra personata Blanco, Fl. Filip.:18. 1837. Type not seen.Bonnaya personata (Blanco) Hassk., Flora 47: 56. 1864. Type not seen.Lindernia serrata (Roxb.) F. Muell., Syst. Cens. Austral. Pl.1: 97. 1882.I. serrata (Roxb.) Urban, Ber. Deutsch. Bot. Ges. 2: 436.1884.Ilysanthes ciliata (Colsm.) Kuntze, Rev. Gen. Pl. 2: 461. 1891.B. aristo-cerrata Hayata, Ic. Pl. Formos.9: 79. 1920. 96
  • 105. Vandellia brachiata (Link & Otto) Biswas, Spec. Publ. Bot. Soc. Beng. 1: 36. 1950.Vandellia ciliata (Colsm.) Yamazaki, J. Jap. Bot. 30: 176. 1955.Bonnaya brachiata Link & Otto ex Hooker in Fl. Br. Ind. 4: 284 (1884); Hara et al. inEnum. Fl. Pl. Nep. 3: 116 (1982); Deyuan et. al. in Fl. Chin. 18: 36 (1998); Press et al. inAnn. Check. Fl. Pl. Nep. 294 (2000); Mill in Fl. Bhutan 2(3): 1125 (2001).Annual, to 10 cm, stem simple or diffusely branching, not rooting at nodes, to 1 mmdiameter, glabrous or shortly hairy on angles. Lamina sessile, elliptic to lanceolate, to 3—4X 1 cm, glabrous to shortly hairy, base attenuate, apex acute—acuminate, margin withusually 14 or more, long aristate teeth per side, not ciliate. Inflorescence a terminal raceme,bracts ca. 3—3.5 X 0.5 mm wide; pedicels shorter than lamina; calyx ca. 4.5 mm in flowerand fruit, calyx lobes ca. 1.5—2 mm in flower, ca. 2.5—3 mm in fruit. Corolla white orpink, deeper pink in throat, yellow, or lilac, ca. 8—10 mm. Capsule dark brown and striate,ca. 9—11 X 1 mm. Seeds ca. 0.25 x 0.3 mm, alveolate, appendaged.Flowering Time: August—December.Fruiting Time: August—December.Habitats: Sal forest. Terai forest and grassy clearings. On steep rocky orsandy banks and moist terraces near river. In moist swampy fields.Altitudinal Range: 500—1900 m.Distribution: W, C, E Nepal. [Deccan; Tibetan Plateau; W. Himalaya; E.Himalaya; Assam-Burma; SE. Asia]Notes: This species is easily identified by its lamina with a single midriband the long-aristate margin.Specimens seen:SSW 7581 (BM) O. Polunin 1928A (BM)EMAK 9 (E) O. Polunin 1961B (BM)K. Wesche No. 7/91 (1995)(BM) P.C. Gardner 1550 (BM)Norkett 6788 (BM) A.H. Norkett 8080 (BM)T. Wraber 187 (BM) PSW 5813 (BM)K. N. Sharma E192 (BM) SSW 6850 (BM) 97
  • 106. 6. Lindernia ruellioides (Colsm.) Pennell, Brittonia 2: 182. 1936. Holotype: König s.n.,Java et India orientali (C!).Gratiola ruellioides Colsm., Prodr. Desc. Grat.: 12. 1793.—Holotype: König s.n., Java etIndia orientali (C!).Gratiola reptans Roxb., Fl. Ind. 1: 140. 1820.—Holotype: Icones 1516 (K!).Bonnaya reptans (Roxb.) Spreng., Syst. Veg.1: 41. 1824.Bonnaya reptans (Roxb.) Spreng. described by Hooker in Fl. Br. Ind. 4: 284 (1884); Haraet al. in Enum. Fl. Pl. Nep. 3: 117 (1982); Deyuan et. al. in Fl. Chin. 18: 36 (1998); Press etal. in Ann. Check. Fl. Pl. Nep. 294 (2000); Mill in Fl. Bhutan 2(3): 1125 (2001).Annual, to 10 cm, stem stoloniferous, simple, rooting at nodes, to 1 mm diameter, glabrous.Leaves petiolate 3—7 mm, lamina pinnately veined, ovate to elliptic on flowering shoots,to 25 mm X 13 mm, orbicular on non-flowering shoots, to 15 X 10 mm, glabrous, baseattenuate, apex round, margin with ca. 14 or more serrations per side, mucronate, non-aristate, ciliate. Inflorescence a terminal raceme; pedicels 15—22 mm, shorter than lamina,bracts ca. 4 X 1 mm. Calyx 4.5--5 mm in flower, 7—8 mm in fruit, glabrous, calyx lobesnarrow, 3—4 mm in flower, 6—6.5 mm in fruit, margin and apex glabrous. Corolla blue,pink, or cream, 9--10 mm, central lower lip lobe to 1.5 X 9 mm; upper lip externallyorange, 4--5 mm; anterior filaments ca. 5 mm, free (not connivent), unappendaged;posterior filaments ca. 2 mm; anterior anthers sterile, ca. 0.3 mm; posterior anthers fertile,ca. 1 mm. Ovary 2 mm, style ca. 6.5 mm. Capsule greenish—beige or light brown, linear,to 15 mm X 1.5 mm. Seeds ca. 0.5 X 0.4 mm, conspicuously (100 μm) appendaged.Flowering Time: August—November.Fruiting Time: August—November.Habitats: Moist terraces near river, moist swampy fields, on argillaciousearth, in sal forest, prostrate in short grass.Altitudinal Range: 400—1500 m.Distribution: W, C, E Nepal. [Tibetan Plateau; W. Himalaya; E. Himalaya; E.Asia; SE. Asia].Notes: This species is at first sight often confused with L. ciliata, but itlacks the long-aristate teeth on the lamina margin. Also, it roots profusely at the nodes; the 98
  • 107. leaves on the non-flowering shoots are roundish, on the flowering shoots elongate, andwider than in L. ciliata.Specimens seen:SSW 7054 (E) Wraber 79 (BM)SSW 7045a (BM) TI 1484 (BM)W&S 8251 (BM) SSW 6798 (BM)Wraber 243 (BM) Wesche 5/ 178 (BM)SSW 9272 (E; BM) Wraber 79 (BM) D (K-W) Wallich 38567. Lindernia pusilla (Thunb.) Bold., Zakfl. Java 165. 1916. Type not seen.Selago pusilla Thunb., Prodr. Pl. Capens. 12: 99. 1794. Type not seen.Diceros caespitosa Bl., Bijdr. 753. 1826. Holotype: Herb. Lugd. Batav. 764 Buitenzorg(L!); Isotype: Herb. Hort. Bot. Bog. [L0003516] Archip. Ind. Java Bogor Tjikaniki; (L!).Tittmannia ovata Benth. non Reichenb. nom. nud. Wall. Cat. 3942 (K-W!).Vandellia scabra Benth., Scroph. Ind. 36. 1835. Wall. Cat. 3945 (K-W!).Tittmannia pusilla Benth. in Wall. Cat 3945C (K-W!).L. pusilla (Thunb.) Bold., Zakfl. Landb. Java 165. 1916.Torenia hirta Cham. et Schlecht. in Linnaea 2: 571. Type not seen.Vandellia scabra Benth. described by Hooker in Fl. Br. Ind. 4: 281 (1884); Hara et al. inEnum. Fl. Pl. Nep. 3: 117 (1982); Deyuan et. al. in Fl. Chin. 18: 33 (1998); Press et al. inAnn. Check. Fl. Pl. Nep. 294 (2000).Annuals, to 20 cm. Stems procumbent, ca. 1 mm diameter, rooting and reddish at nodes.Leaves sessile, lamina cordate, 3—5-veined from base, 8—13 X 6—10 mm, sparsely,coarsely hairy on both sides, veins protruding below, lamina margins undulate—serrate.Inflorescence axillary, solitary, lamina size decreasing toward the apex of the plant,pedicels longer than leaves. Calyx to 3.5 mm in flower, to 4.5 mm in fruit, hairy, calyxlobes ca. 2 mm in flower, ca. 3 mm in fruit, margins ciliate. Corolla white or purple, ca.4—6 mm; upper lip slightly bifid, lower lip deeply tri-lobed, bilobed yellow spot in middleof lower lip, ca. 5—6 mm wide, central lower lip lobe ca. 3 mm. Stamens all fertile orposterior fertile. Capsule green, ovoid—globose, ca. 2.5—3.5 X 3 mm. Style not persistentin fruit. Seeds ca. 0.4 x 0.4 mm, shortly octagonally cylindric, reticulation protruding ca. 5μm . 99
  • 108. Flowering Time: May—October.Fruiting Time: September—November.Habitats: Mixed sedge and herb vegetation.Altitudinal Range: Occurs at least at 500 m.Distribution: E Nepal; [W. Himalaya; E. Himalaya; SE. Asia].Notes: The combination of general hairiness, conspicuous corolla andglobose capsule with non-persistent style make this Lindernia easy to spot. We need morecollections.Specimens seen:TI 1478 (1963, BM)Buch-Ham. 1381 (E)Buch-Ham. 1395 (E)Wall. Cat.3945F (K-W)DNEP 2B—192 (E) 100
  • 109. 8. Lindernia viscosa (Hornem.) Bold., Zakfl. Java 165. 1916. Type not seen.Gratiola viscosa Hornem.?? 1807. Protologue not seen. Type: C!Hornemannia viscosa (Hornem.) Willd. Enum. Hort. Berol.: 654. 1809.Diceros glanduliferus Bl., Bijdr.: 752. 1826.—Holotype: Herb. Lugd. Batav. [L0003530],Java (L!).Vandellia hirsuta Ham. ex Benth. Scroph. Ind. 36. 1835.—Type: Habitat in Camrupa (?)orientalis humidis. Buch.-Ham. 1380 (E!).Lindernia hirsuta (Ham. ex. Benth.) Wettstein. Type not seen.Vandellia viscosa (Hornem.) Merrill, Phillipp. Journ. Sci. Bot.7:246. 1912.Lindernia glandulifera (Bl.) Backer, Onkruidfl. Java Suiker.: 616. 1931.Vandellia hirsuta Benth. described by Hooker in Fl. Br. Ind. 4: 284 (1884); Hara et al. inEnum. Fl. Pl. Nep. 3: 117 (1982); Deyuan et. al. in Fl. Chin. 18: 33 (1998); Press et al. inAnn. Check. Fl. Pl. Nep. 294 (2000); Mill in Fl. Bhutan 2(3): 1122 (2001).Annuals, to 15 cm. Stems procumbent, ca. 1 mm diameter, rooting at nodes, hairy. Leavessessile to 5 mm petiolate. Lamina pinnately veined, ovate to lanceolate, 15—40 X 12—22mm, sparsely, coarsely hairy on both sides, increasingly so on veins below, hairs adpressed,lamina margins dentate or undulate, shortly ciliate, longer on younger leaves. Inflorescenceracemose, bracts to 1.5 mm, pedicels shorter than leaves. Calyx hairy, to 3 mm in flower, to4 mm in fruit, calyx lobes 1.75 mm in flower, 2.25 mm in fruit, margins ciliate. Corolla 4—5.5 mm, white or blue; ovary ca. 1.25 mm, style ca. 2 mm; stamens all fertile. Capsuleovoid, almost as long as calyx. Seeds ca. 0.3 X 0.4 mm, ellipsoid—oblong, slightlytapering towards point of attachment to septum.Flowering Time: May—September.Fruiting Time: June—November.Habitats: Moist terraces, in sal forest, on edge of cultivation.Altitudinal Range: 200—900 m.Distribution: C, E Nepal. [Deccan; W. Himalaya; E. Himalaya; E. Asia; SE.Asia]. 101
  • 110. Specimens seen:L.H.J.Williams 325 (BM) SSW7056a (1954, BM)Norkett 8084a (BM)9. L. parviflora (Roxb.) Haines, Bot. Bihar & Orissa: 635. Holotype: Roxburgh, Pl.Coromand. 3:3, t. 203. 1819. Illustration (E!).Gratiola parviflora Roxb. Pl. Coromand. 3: 3, t. 203 (1819). Holotype: Roxburgh, Pl.Coromand. 3:3, t. 203. 1819. Illustration (E!).Hooker in Fl. Br. Ind. 4 (1884); Hara et al. in Enum. Fl. Pl. Nep. 3: 117 (1982); Press et Ann. Check. Fl. Pl. Nep. 294 (2000).Annual, to 15 cm, stem winged, 0.5--1 mm diameter, not rooting at nodes, glabrous. Leavessessile, lamina 3—5-veined from base, lanceolate, 8—16 X 3—7 mm, glabrous; baserounded or clasping the stem, apex acute—acuminate, margin subentire to distantly serrate.Inflorescence solitary, axillary, appearing subracemose; pedicels winged, 5—14 mm long,often 1.5—2 times longer than lamina; leaves subtending pedicels 3—10 X 1—4 mm;leaves toward apex at least half the size as at base of plant. Calyx to 3 mm in flower, to 3mm in fruit, glabrous; calyx lobes ca. 1.5 in flower and fruit, sparsely ciliate on apex.Corolla white to yellowish white, 4—5.5 mm, upper lip slightly bifid; anterior stamenssterile, posterior stamens fertile; ovary 1.5—2 mm, style 3—4 mm, stigma broadening.Capsule ovoid, occasionally tapering toward the apex, 2.5—3.5 X 1.8—2.2 mm. Seedscylindrical to cylindrically hexagonal, ca. 0.25 X 0.15 mm.Flowering Time: November—December.Fruiting Time: November—December.Habitats: Fallow paddy fields.Altitudinal Range: Sea-level—800 m.Distribution: C, E Nepal. [W. Himalaya].Notes: This species is hard to distinguish from L. procumbens, except for its occasionallyobscurely toothed lamina margin and its anterior pair of anthers being sterile. A thirdcharacter might be the callous style base that remains on the non-dehisced capsule of L.procumbens, but not on L. parviflora. 102
  • 111. Specimens seen:Norkett 8084 (BM) Norkett 8037 (BM)SSW 8696 (E) DNEP 2B 125 (E)TI 1490 (K)10. Lindernia micrantha D. Don, Prodr. Fl. Nep. 85. 1824. Type: BM, not seen.Tittmannia angustifolia Benth. nom. nud.—Type in Wall. Cat. 3951 (K-W!).Vandellia angustifolia Benth., Scoph. Ind. 37. 1835.—Type in Wall. Cat. 3951 (K-W!).Lindernia angustifolia (Benth.) Wettst. in Engler, Pflfam. IV - 3b: 79. 1891.Vandellia angustifolia Benth. described by Hooker in Fl. Br. Ind. 4: 282 (1884); Hara et Enum. Fl. Pl. Nep. 3:116 (1982); Deyuan et. al. in Fl. Chin. 18: 35 (1998); Press et al. inAnn. Check. Fl. Pl. Nep. 294 (2000).Annual to 20 cm, stem simple or branching, erect, not rooting at nodes, 0.5—1.3 mmdiameter, slightly winged, glabrous. Leaves sessile to 2mm petiolate, lamina uni-veined,linear, entire to obscurely serrate, 10—27 X 1.4—4.2 mm, glabrous; base attenuate, apexrounded, margin entire to obscurely incised, glabrous or ciliate. Inflorescence solitary,axillary, leaves subtending pedicels same size as leaves below, smaller when subtendingpedicels of immature capsules; pedicels 4—15 mm in flower, 10—28 mm in fruit. Calyx3.5—4.5 in flower, 3.5—5 mm in fruit, glabrous; calyx lobes 2.7—3.6 mm in flower, ca. 4mm in fruit, margins and/ or apex ciliate. Corolla white—pink, 6—9 mm, lower lip ca. 8mm wide, central anterior lip lobe ca. 3 mm wide; posterior lip ca. 5 mm wide; anterior 2filaments ca. 5 mm, appendaged; posterior 2 filaments ca. 1 mm, stamens all fertile, anthers0.8—1.3 mm. Ovary 2.2—2.7 mm, style 3.5—4 mm. Capsule golden brown, striate,ellipsoid to linear, 9—15 X 1.2—2 mm. Seeds lanceoloid, alveolate 3.5 x 2.5 mm.Flowering Time: September—October.Fruiting Time: September—October.Habitats: Moist terraces; growing on bank of paddy field, in paddy field.mud.Altitudinal Range: 700—1300 m. 103
  • 112. Distribution: C Nepal; [W. Himalaya; E. Asia; SE. Asia].Notes: This species is easily recognized by its narrow, linear leaves; as tothis character, however, it is posible that we are dealing with L. oppositifolia, whoseexistence in Nepal has been reported (but without voucher specimens). The inflorescenceof L. oppositifolia are alternating, axillary, solitary peduncles, one per node, whereas L.micrantha usually has two.Specimens seen:EMAK 174 (E) J.H. de Haas 2675 (BM)PSW 5856 (E; BM) SSW 7096 (E; BM)Wall Cat 3951 B (E, K-W)11. Lindernia multiflora (Roxb.) Mukerjee, J. Ind. Bot. Soc. 24: 131. Holotype:Roxburgh, Icones 1267 (K!)Torenia multiflora Roxb., Fl. Ind., ed. Carey, 3: 96. 1832.—Holotype: Roxburgh, Icones 1267(K!)Vandellia multiflora (Roxb.) G. Don, Gen. Syst. 4: 549. 1838.Pyxidaria trichotoma (Benth.) Kuntze, Rev. Gen. 2: 464. 1891. nom. illegit.Lindernia papuana Pennell, J. Arn. Arb. 24: 251. 1943.—Isotype: Clemens 10907, Papua,Kajabit Mission [L0003525] (L!).Vandellia trichotoma Benth. nom. nud. Wallich 3943 (K-W!).Tittmannia trichotoma Benth. nom. nud. Wallich 3943 (K-W!).Vandellia multiflora (Roxb.) G. Don. described by Hooker in Fl. Br. Ind. 4: 282 (1884);Hara et al. in Enum. Fl. Pl. Nep. 3: 116 (1982); Press et al. in Ann. Check. Fl. Pl. Nep. 294(2000).Annuals to 15 cm, stem 0.8—1.25 mm diameter, simple or branched, not rooting at nodes,winged, glabrous. Leaves sessile to 8 mm petiolate, lamina pinnately veined, 20—35 X 8—14 mm, glabrous; base attenuate, apex acute or rounded, margin entire to serrate, glabrousor ciliate. Inflorescence a terminal raceme, bracts minute, ca. 1 mm X 0.8 mm; pedicels ca.3.5 mm in flower, 5—8 mm in fruit, shorter than leaves, slightly reflexed in fruit. Calyx ca.2 mm in flower, 2.75—4 mm in fruit, glabrous; calyx lobes ca. 1.25 mm in flower, 2—3.5 104
  • 113. mm in fruit, glabrous, margins ciliate. Corolla purplish—white, 3.5—4 mm, central lobe oflower lip ca. 0.6 mm. Stamens all fertile, ca. 0.3 mm. Capsule ovoid, ca. 3 x 2 mm, apexacute. Seeds alveolate, filiform appendages.Flowering Time: ??Fruiting Time: ??Habitats: Moist terraces.Altitudinal Range: ??Distribution: [Deccan; W. Himalaya; E. Himalaya; SE. Asia]Notes: Only one Nepalese specimen was available for this description. Theplant resembles L. viscosa, but is glabrous. The slightly reflexed pedicels in fruit and therelatively acute apex of the capsule are good distinguishing characters.Specimens seen:Banerji, Shresta & Upadhya 2530 (BM).12. Lindernia nummulariifolia (D. Don) Wettst., in Engl., Pflfam. IV - 3b: 79. 1891.Lectotype: Wallich ex. Herb. Hookerianum s.n., Nepal, 1821 (K!).Vandellia nummularifolia D.Don, Prodr. Fl. Nep. 86. 1825.—Lectotype: Wallich ex. Herb.Hookerianum s.n., Nepal, 1821 (K!).Hara et al. in Enum. Fl. Pl. Nep. 3: 117 (1982); Deyuan et. al. in Fl. Chin. 18: 31 (1998);Press et al. in Ann. Check. Fl. Pl. Nep. 294 (2000); Mill in Fl. Bhutan 2(3): 1124 (2001).Annual, to 12 cm, erect, stem simple or branching at base, to 1 mm diameter, not rooting atnodes, hairy to 0.2 mm. Leaves sessile or subsessile, lamina pinnately 3—5-veined (butseemingly palmate), ovate to suborbicular, 10—15 X 10—15 mm, glabrous and glossyabove, sparsely ciliate on veins below; base rounded, apex rounded or obtuse, margin to10-dentate, 0.4 mm deep serrate to serrulate, ciliate to shortly hairy. Inflorescence solitary,axillary, occasionally subracemose and 3 pedicels per node; pedicels 1—4 cm, but usuallyabout equal the lamina length, glabrous, but shortly hairy toward the apex; bracts 0.7—1.3mm. Calyx 2—3 mm in flower, 2.5—3.5 mm in fruit, glabrous; calyx lobe margins ciliateon apex. Corolla pink and white, 5—7 mm, central lower lobe ca. 1--4 mm, upper lip ca. 3 105
  • 114. mm X 2.5 mm. Anterior filaments ca. 4 mm, geniculum to 0.15 mm, densely papillose,connivent; posterior filaments ca. 1.5 mm, connivent; anterior anthers sterile, posterioranthers fertile. Capsule linear—cylindric, acuminate, 5—6 X 2 mm wide, 1.5—2 timeslength of calyx. Seeds ca. 0.4 mm long, ca. 0.3 mm wide, ellipsoid, alveolate.Flowering Time: July—August.Fruiting Time: August—October.Habitats: Stoney wayside track; Moist terraces, in shade, in rice fields.Altitudinal Range: 1100—2800 m.Distribution: W, C, E Nepal. [Deccan; Tibetan Plateau; W. Himalaya; E.Himalaya; Assam-Burma].Notes: The usually hairy stem is contrasted by a glossy upper lamina.The capsules are acutely-tipped, and angled. The seeds are large compared to the otherLindernia.Specimens seen:SSW 4018 (BM) SSW 6438 (BM)SSW 6782 (BM) TI 9760019 (BM)TI 9760206 (BM) Sharma E258 (BM)Dhwoj 580 (E) H. Flatt 79 (BM)J. H. de Haas 2042 (BM) Shresta 1025 (BM)Polunin 1929 (BM) PSW 5169 (E) (BM)Polunin 1998 (BM) PSW 5622 (BM)13. Lindernia hookeri (C. B. Clarke) Wettst., in Engl., Pflfam. IV - 3b: 79. 1895. Type:Herb. Ind. Or. Hook. fil. & Thompson, Khasia (E!).V. hookeri C.B.Clarke ex Hook. F., Fl. Brit. Ind. 4: 280. 1884. Type: Herb. Ind. Or. Hook.fil. & Thompson, Khasia (E!).Vandellia hookeri Clarke mss. ex Hooker in Fl. Br. Ind. 4: 280 (1884); Hara et al. in Enum.Fl. Pl. Nep. 3: 116 (1982); Press et al. in Ann. Check. Fl. Pl. Nep. 294 (2000); Mill in Fl.Bhutan 2(3): 1123 (2001). 106
  • 115. Annual to 15 cm. Stem erect, to 1.5 mm diameter, sparsely pubescent at angles, hairs to 1mm long. Leaves shortly to 6 mm petiolate, lamina pinnately veined, ovate or lanceolate, to25 X 10 mm, lamina base attenuate to cuneate, apex acuminate, margin serrate, with 8 to10 incisions per side, less deep towards base. Upper lamina surface with hairs to 0.7 mm,lower surface glabrous between veins, hairs to 1.3 mm on midribs. Inflorescence a terminalcyme and/ or solitary, axillary. Pedicels approximately as long as leaves. Subtending bractsapproximate leaf size at base of inflorescence, much smaller, bract-like at apex. Calyxbilabiate, 5-lobed, lobes subequal, divided to middle in flower, +/- to base in fruit and fullycovering the capsule, to 10 mm, strongly hairy (to 1.5 mm); lobes with strong central veins.Corolla blue – purple, to 10 mm. Anterior filaments ca. 3 mm, posterior filaments shorter,with basal appendage; anterior anthers ca. 0.5 mm, sterile, posterior anthers fertile. Capsulelight brown to golden, ellipsoid to ovoid, to 6 X 4 mm. Seeds to 0.4 x 0.35 mm, alveolate,not appendaged.Flowering Time: August—September.Fruiting Time: August—September.Habitats: Moist terraces.Altitudinal Range: 900—1700 m.Distribution: C, W Nepal; [E. Himalaya].Notes: This plant is easily recognized by its bilabiate, hairy calyx and itsfat capsules.Specimens seen:SSW 6767 (E; BM) SSW 7056 (BM)Banjeri, Shresta & Upadhyay 2528 (BM)14. Lindernia indet-aAnnual to 15 cm, stem erect or occasionally procumbent and rooting at nodes, to 2 mmdiameter, glabrous. Lamina 3—5-veined, sessile, broadly elliptic to rhombic, to 25 X 12mm, glabrous, base decurrent, apex obtuse, lamina margin subentire—undulate.Inflorescence solitary, axillary, leaves subtending pedicels 14—22 X 5—12 mm; pedicelsto 10 mm, shorter than leaves. Calyx to 4 mm in fruit, glabrous; calyx lobes to 3 mm in 107
  • 116. fruit, calyx lobe margin smooth. Corolla white, closed. Filaments absent, anteriorstaminodes, posterior stamens fertile. Capsule ca. 3 x 2 mm, ovoid, striate, glabrous, onlybase of style (to 1 mm) persitent on capsule. Seeds ca. 0.35 x 0.2 mm, not alveolate, notappendaged, seed coat smooth.Flowering Time: at least JulyFruiting Time: at least JulyHabitats: In mud of drained paddy-field.Altitude: At least at 2400 m.Distribution: W Nepal. Possibly endemic.Notes: The description above stems from a single collection (Polunin, Sykes, and Williams4476); the 10 plants collected varied little. A good character may be that duringinflorescence, one side develops a peduncle, while the opposing side develops a furthershoot that carries two leaves. Next node up, the reverse.Specimens seen: PSW 4476 (E; BM). 108
  • 117. APPENDIX (2) Indo- Indo- W. Sri Malesia Bhutan Thailand China Nepal Burma Himal China Lanka JavaL. anagallis (Burm.f.) Pennell 1 1 1 1 1 1 1 1L. antipoda (L.) Alston 1 1 1 1 1 1 1 1L. blancoi (Merr.) Philcox 1L. brevipedunculata Migo 1L. cambodgiana (Bonati) Philcox 1L. celebica Philcox 1L. cephalantha T. Yamaz. 1 1L. cerastioides T. Yamaz. 1L. ciliata (Colsm.) Pennell 1 1 1 1 1 1 1L. crenata Pennell 1L. crustacea (L.) F. Muell. 1 1 1 1 1 1 1 1L. cyrtotricha P. C. Tsoong 1L. delicatula P. C. Tsoong 1L. dictyophora Tsoong 1 1 1L. dubia (L.)Pennell 1L. elata (Benth.) Wettst. 1 1L. glabra Philcox 1L. hookeri (C.B.Clarke) Wettst. 1 1 1 2L. hyssopioides (L.) Haines 1 1 1L. jiuhuanica X. H. Guo & X. L. Liu 1L. junciformis (Bonati) T. Yamaz. 1 1L. kiangsiensis P. C. Tsoong 1L. kerrii T. Yamaz. 1 1L. khaoyaiensis T. Yamaz. 1L. laotica Bonati 1L. latifolia ? 1 1L. maxwellii T. Yamaz. 1 1L. macrobotrys P. C. Tsoong 1L. megaphylla P. C. Tsoong 1L. micrantha D. Don 1 1 1 1 1 1L. mollis (Benth.) Wettst. 1 1 1 1L. molluginoides Wettst. 1
  • 118. L. Montana ? 1L. multiflora (Roxb.) Mukerjee 1 1 1L. nummulariifolia (D.Don) Wettst. 1 1 1 1 1 1L. oblonga (Benth.) Merr. & Chun 1L. parviflora (Roxb.) Haines 1 1 1L. philippinensis Philcox 1L. pierreanoides T. Yamaz. 1 1L. procumbens (Krock.) Philcox 1 1 1 (1?) 1 1 1L. pusilla (Willd.) Bold. 1 1 1 1 1 1L. rivularis Kerr ex Barnett 1L. rotundifolia ? 1L. ruellioides (Colsm.) Pennell 1 1 1 1 1L. satakei T. Yamaz. 1L. scapigera R. Br. 1L. scutellariiformis T. Yamazaki 1L. sessiliflora (Benth.) Wettst. 1L. setulosa (Maxim.) Tuyama ex. Hara 1L. srilankana Cramer & Philcox 1L. stemodioides (Miq.) Kerr 1L. stolonifera T. Yamaz. 1L. stricta P. C. Tsoong & T. C. Ku 1L. subconnivens Philcox 1L. succosa (Kerr. Ex Barnett) Philcox 1L. subulata R. Br. 1L. taishanensis F.Z.Li 1L. tenuifolia (Colsm.) Alston 1 1 1L. thorelii Bonati 1L. udawnensis T. Yamaz. 1 1L. umbellata T. Yamaz. 1L. urticifolia (Hance)Bonati 1L. viatica (Kerr. Ex Barnett) PhilcoxL. viscosa (Hornem.) Bold. 1 1 1 1 1 1 1L. yaoshanensis P. C. Tsoong 1L. indet-a 1L. indet-b (1?) Philcox Mill Yamazaki Yamazaki (Baden) Mukerjee Pennell Deyuan et al. Haines Backer 110
  • 119. BENTHAM’S GENERA (1846) Tribe VIII: Gratiolae Subtribe: LindernieaeGenus: Vandellia Lindernia Ilysanthes BonnayaSection Ellobum Nummularia Ilyogeton Tittmannia Torenioides V. angustifolia L. pyxidaria I. parviflora B. brachiata V. pedunculata B. reptans V. nummulariifolia V. erecta B. veronicaefolia V. scabra V. hirsuta V. crustacea V. multiflora APPENDIX (3) 111
  • 120. Appendix (4) Krocker’s protologue of Anagalloides procumbens nostra _______________________________________ 112
  • 121. _____________________________________________________
  • 122. 114
  • 123. APPENDIX (5): Micrographs of lamina surfaces at magnification x 2000. Column to left = upper; to right = lower surface; L. ciliata, L. multiflora, L. antipoda, L. ruelliodes.5 _______ 115
  • 124. 116