A REVISION OF NEPALESE LINDERNIA All. (SCROPHULARIACEAE S.L.)

Heide Maria Baden A thesis presented in partial fulfilment of the degree of M.Sc.

The Biodiversity & Taxonomy of Plants

Royal Botanic Garden Edinburgh &

University of Edinburgh 2005

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ABSTRACT

The Nepalese species of Lindernia (Scrophulariaceae s.l.) were revised. These are at least 13

distinct species with possibly two additional species. For species delimitations, vegetative

and sexual characters, in particular leaf, flower, and fruit morphology, were taken into

account. Numerical taxonomy included leaf length to leaf width ratio and pedicel length to

leaf length ratio. Measurements made allowed a definition of inflorescence type, an

important distinguishing character within Lindernia. An SEM study of Lindernia seed

material was undertaken. Results did not support the traditional groupings into subgenera

proposed 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 is

included.

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CONTENTS

Title page i Abstract ii Contents iii Tables and Figures vi List of Abbreviations viii 1. INTRODUCTION 1 2. AIMS AND OBJECTIVES 3 3. 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

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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 21 3.4. Recent floristic accounts 21

Mukerjee 21

Sivarajan & Mathew 21

Banjeri 21

Philcox 21

Khan & Hassan 22 Diaz Miranda 22

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Cramer 22 Yamazaki 23 Quail Lewis 23

4. 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 40

5. 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

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Testal pattern 46

5.1.1 SEM results and Yamazaki’s recent subgenera 59

5.2 Taxonomic Problems 63

5.2.1 Identity of the type species of the genus 63

5.2.1.1 Identity of the Asian form of the type species 65

5.2.2 Vandellia erecta and Lindernia parviflora 68

. 5.2.3 The identity of L. indet-a 73

5.2.4 Lindernia indet-b from “Dr. Buchanan” 77

5.2.5 The type of L. micrantha D. Don 78

5.2.6 The type of Lindernia oppositifolia (Retz.) Mukerjee 79

6. SUMMARY 80 7. REFERENCES 81 8. 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.

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LIST OF FIGURES AND TABLES

Fig. 1. a – d. Specimens and Illustrations of L. procumbens (Krock.) Philcox 5 Fig. 2. a – b. Roxburgh’s illustrations of Gratiola parviflora Roxb. 9 Fig. 4. 1 a – b. Roots. 25 Fig. 4. 2 a – h. Hairs. 26 Fig. 4. 3 a – d. Stems. 28 Fig. 4. 4 a – g. Lamina shapes. 30 Fig. 4. 5 a – c. Nodes. 31 Fig. 4. 6 a – d. Inflorescence types. 32 Fig. 4. 7 a – e. Calyx types. 33 Fig. 4. 8 a – b. Pressed Corollas. 34 Fig. 4. 8 c – h. Corollas. 36 Fig. 4. 9 a – f. Floral dissections. 37 Fig. 4. 10. a – k. Capsules. 38 Tab. 5 Seed and seed-surface characters from SEM study. 47 Fig. 5. 1 a – p. SEM micrographs of entire seeds. 48 Fig. 5.2 A – P. SEM micrographs of seed coats. 53 Fig. 5.2 α + β. SEM micrographs of appendages. 58 Fig. 5. 3 Classification of Lindernia based on Yamazaki (1981) 59 Fig. 5. 4 a + b. Linnaean specimens 796.3 64 Fig. 5. 4 c – g. Wallich specimens seen by Bentham. 70 Fig. 5. 5 Graph: Manner of inflorescence. 75 Fig. 5. 6 Second unknown specimen from Nepal 76 Fig. 5. 7 Buchanan-Hamilton specimen 77 Fig. 5. 8 a. Lectotype of V. angustifolia Benth. 78 Fig. 5. 8 b + c. L. oppositifolia (Retz.) Mukerjee 78

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LIST OF ABBREVIATIONS

B Botanic Garden and Botanical Museum Berlin-Dahlem BM British Museum-Herbarium in the Museum of Natural History BUC Buchanan-Hamilton abbreviated in SEM Buch.-Ham. Buchanan-Hamilton abbreviated in Herbarium Collections C Museum Botanicum Hauniense, Copenhagen DNEP Edinburgh expedition code E Royal Botanic Garden, Edinburgh EMAK, Edinburgh expedition code ICBN International Code of Botanical Nomenclature ING Index Nominum Genericorum K Royal Botanic Gardens, Kew K-W Honourable East India Company (Wallich) Herbarium LD Botaniska Museet Lund LD Nationaal Herbarium Leiden LINN Linnaean Society of London NOR Norkett PSW Polunin, Sykes & Williams S Stainton s.l. sensu lato s.str. sensu stricto SSW Stainton, Sykes & Williams TI Tokyo Institute

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1. INTRODUCTION

Lindernia was named in 1766 by Allioni after von Lindern (1682—1755), who first

illustrated what we now know as Lindernia pyxidaria L. in Tournefortius Alsaticus (1728).

In this Miscellanea Taurinensis: Stirpium aliquot descriptiones cum duorum generum

constitutione, Allioni did not explicitly include a specific epithet when describing this

species from Alsace-Lorraine in France, but cited a specimen collected by Gagnebin in

Switzerland, and provided an illustration (Fig. 1. b; p. 5). In 1965, Philcox concluded that

L. pyxidaria was illegitimate. He also decided that it was conspecific with Anagalloides

procumbens 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 been

followed through by Index Nominum Genericorum, which still maintains Lindernia

pyxidaria All. as the (problematic) first species descripion of the genus.

With ca. 100 species, Lindernia was placed in the Scrophulariaceae and is distinguished

from other genera in the Scrophulariaceae sensu lato by the following character

combination: (1) absence of bracteoles, (2) presence of a 5-angular, relatively

actinomorphic calyx, (3) a conspicuously 2-lipped corolla, and (4) a persistent septum in

the bisepticidally dehiscent capsule (Pennell 1943a, Deyuan et. al. 1998, Lewis 2000, Mill

2001). The group was treated in its widest sense by Pennell, who in 1935 moved the genera

Vandellia 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, New

Guinea, Deyuan treated 29 for China; Mill treated 10 for Bhutan, Yamazaki treated 30 for

Thailand, 47 for Indochina, Japan, Taiwan. Bentham treated 27 for India (excluding the

synonymous ones, as far as my research went).1

Recently, the Scrophulariaceae have been subject to extensive phylogenetic analysis

(Albach et al. 2005, Oxelman et al. 2005), and the family Linderniaceae (Rahmanzadeh et

1 Appendix 2; the synonymy has not been checked.

2

al. 2004) was proposed to encompass Lindernia together with 12 other genera including

Torenia L. and Craterostigma Hochst. With ca. 100 species, Lindernia is the largest genus

of the new family. This study split the traditional tribe “Gratiolae2,” created by Bentham in

1835, into two families: the Linderniaceae and the Gratiolaceae. Early classification of

Lindernia spp (Linnaeus 1771; Willdenow 1779; Retzius 1786; Colsmann 1793; Vahl

1804; Roxburgh 1819) stressed links with the genus Gratiola.3

It is impossible to judge the monophyly of Lindernia as of yet; the estimated phylogenies

so far (Rahmanzadeh et al. 2004) suggest Lindernia s.l. and Lindernia s.str4

. The

nomenclature of this group is of yet an unfinished puzzle, especially the complex

surrounding the type species of the genus. These questions should be sorted out before, or

in conjunction with more phylogenetic studies, for the latter to be of value to our

understanding of the evolution of Lindernia.

Medicinal properties of Lindernia have long been studied, with references to vires medicas

nihil certi (Allioni 1765), and Usus latet (Krocker 1790) reflecting the connection between

humans and this plant. Miyase et. al. (1995) have isolated the oleanane saponins

Linderniosides A and B from Lindernia pyxidaria. Saponins exhibit important detergent or

surfactant, as well as antiviral properties important to medicine (Simŏes et al. 1999). In

addition, 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 Miranda

for South America (1977)] could provide supplementary clues to clearing up questions of

affinity within this group.

2 Rahmanzadeh et al. explain the morphological basis of this tribe, distinguished most importantly by its ovule 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 testa cells 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 in Gratiola genus folders. 4 Because only few taxa of Lindernia were sampled during this study (only six), it is probably too early to assign species into the Lindernia s.str. and s.l. We might in fact be dealing with several genera within Lindernia s.l..

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2. AIMS AND OBJECTIVES

The goals of this study were to (1) check the taxonomy of Nepalese species, and to revise it

where necessary, (2) revise the synonymy of these species, (3) test the existing

classification using micromorphology, (4) prepare a draft account of the genus for the Flora

of Nepal.

3. TAXONOMIC HISTORY

This chapter deals chronologically with all the names and synonyms used for Nepalese

species, 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 then

continues with a section on the treatments that emphasized subgeneric relationships and

taxonomy within (3.3), and ends with (3.4) a section on the more recent regional floral

accounts 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 Lindernia

Lindernia was created by Allioni in his Stirpium aliquot descriptiones cum duorum

generum constitutione, published in Miscellanea Taurinensis 3, describing a species

formerly known by the polynomial “Alsinoides paludosa foliis anagallidi similibus

flosculia monopetalis rubescentibus capsula oblonga,” and referring to a specimen

collected by “Gagnebin,”5

5 In the Harvard University Herbarium - Index of Botanists, Abraham Gagnebin holds record number 140348; 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.

as well as to an American (Virginian) specimen given to him by

“Celeb. Linnaeus” (Allioni 1766). He gave both a species description and the generic

description of Lindernia All., however, did not include a specific epithet in his description

of the genus (merely mentioning that von Lindern had called the taxon pyxidariae), nor in

the accompanying figure. Linnaeus published the binomial Lindernia pyxidaria L. in

Mantissa Plantarum: 252 (1771). Here he referenced (1) Allioni’s illustration, (2) his own

Capraria gratioloides L. from Species Plantarum: 876 [it is actually thought to be

4

described in Systema Vegetabilium of 1759], and (3) von Lindern’s illustration in

Tournefortius 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 his

own “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 genus

is illegitimate under the International Code of Botanical Nomenclature (ICBN), because

Linnaeus Capraria gratioloides L. was not a synonym of Lindernia pyxidaria L., but the

type of another species. It is not yet clear to me, whether or not C. gratioloides is indeed

synonymous with Gagnebin’s specimen.

The database “Index Nominum Genericorum [ING]” lists Lindernia pyxidaria All. as the

type of the genus. It has been verified here that Allioni does not explicitly make a binomial

combination before Linnaeus in 17716

. Until it has been proven that the Gagnebin

specimen and Capraria gratioloides are distinct, and until it has been verified that

Linnaeus’ description is indeed illegitimate, I suggest not to change the entry in ING. If

illegitimacy existed through citing a type of another species, as Philcox argues, then the

ING entry should be changed. The matter needs further investigation, a part of which will

follow in section 5. 2 under taxonomic problems.

The legitimate publication to replace Lindernia pyxidaria was made by Krocker in Flora

Silesiaca (1790. Fig. 1. c and Appendix 4), in which he described Anagalloides

procumbens in as great a detail as Allioni described Lindernia (pyxidaria). In contrast to

Allioni, 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 this

was the reason why Philcox chose A. procumbens Krock. as the new basionym for the

illegitimate precursor. It was not until Philcox made the new combination in 1965, that L.

procumbens (Krock.) Philcox7

6 It is possible, however, that Allioni made the combination in an as of yet unpublished work.

was accepted as the type of the genus in all the accounts I

consulted during this revision.

7 For L. procumbens (Krock) Philcox, the latter includes in synonymy, amongst others, Vandellia erecta Benth. 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).

5

Fig. 1 a: Collections from Herb. Linn. 796.1; mixed sheet of L. dubia L. on left; L. virginiana L. on the right. Wulfen reference in the illustration not found in literature.

Fig. 1 b: Allioni’s illustration from Miscellanea Taurinensis 3, T5, F.1 (1766); open flower features 4 fertile stamens.

Fig. 1 c: Linnean specimen: Herb. Linn. 796.2. On the reverse side of 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 ovate, bivalved, 2-locular, valves elliptic.

Fig. 1 d: Anagalloides procumbens Krocker. 1790. Flora Silesiaca 2(1), t. 26.

6

Unfortunately, it has not been possible to verify the types of all the species listed in the

synonymy of the Nepalese species. Therefore, dealing with the type of the genus from

Europe, and a potentially conspecific specimen from Virginia, but using this name for

Asian material needs further clarification. For further research, it will be necessary to

investigate the identities of Lindernia pyxidaria Pursh. non Allioni, Capraria gratioloides

L., L. dubia L., and Gratiola anagallidea Michx. For example, one needs to find out

whether Ilysanthes gratioloides Bentham has Capraria gratioloides L. as its (omitted)

basionym. This is important, because Bentham (1846) also says that Ilysanthes gratioloides

Benth. = L. pyxidaria Pursh. non Allioni, which however would only give a pointer to the

question whether Linnaeus synonymy was indeed wrong. A recently published, extensive

work by L.S. Olschki (2004, ed.) entitled Le opere minori di Carlo Allioni. Dal “Rariorum

Pedemontii stirpium” all’ “Auctarium ad Floram Pedemontanam” references several

plates of Allioni’s exsiccata and illustrations, including Capraria gratioloides L. [which

looks very much like L. procumbens (Krock.) Philcox], and should help to shed further

light on the matter. It is available at the Kew library, but again, time was too short to delve

any deeper into this important matter.

Having discussed the problems concerning the type of the genus, I will now proceed in

chronological order to describe the authors as well as major treatments in the taxonomic

history 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 the

Wallich herbarium at Kew (K). The specimens at Edinburgh (E) lack flowering material

and could not be identified.

3.2. Other first species descriptions

Linnaeus had described another Lindernia8

8 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 mistakes were 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 of Willdenow on page 6 ahead).

even earlier than L. pyxidaria, i.e. in his

Species Plantarum (1753), but under the name Ruellia antipoda L., (BM!, Herb. Hermann

collection, Sloane room). This publication was legitimate and the taxon is now known as

7

L. antipoda (L.) Alston. See the entry for Alston (1931) on page 14 for a word of caution

concerning his new combination.

Burman (1707-1780) described Ruellia anagallis Burm.f. [basionym of Lindernia

anagallis (Burm.f.) Pennell] in his Flora Indica: 135 (1768), referring to a Kleinhof

specimen from Java named Anagalloides javanica, and an illustration in Rhumphius’

Herbarium Amboinense [(5): 460, t. 170, f. 2; 1747], which Burman co-edited. This

illustration, however, appears to refer to a different taxon (e.g., L. sessiliflora), since the

calyces are depicted as sessile. However, in his voorwoordt (1695), Rumphius disclaimed

accuracy of the figures, explaining that some were lacking flowering or fruiting material at

the 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 particular

way [here in Indonesia], however, that one should not be judgemental for the variation of

plants across geographic boundaries.”

Retzius (1742-1821), botanist at Lund, described Gratiola grandiflora and Gratiola

veronicifolia in Fasciculus Observationum Botanicarum 4 (1786), separating them on the

basis of flower size. Both types were collected by König,9

and both species are now

synonymised with L. antipoda (L.) Alston [one must be aware of any L. grandiflora non

Retzius, however, which may refer to different taxa]. Further, Retzius is the first to

describe 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 voucher

specimen has not been recovered for Nepal; however, this species is checklisted in the

Enumeration of the Flowering Plants of Nepal (Hara et al. 1981); the misidentified voucher

specimen TI 1488 is actually L. micrantha (D.Don) Wettstein.

Colsman worked on a thesis of the Gratiolae, entitled Species Gratiolae a D. König in

itinere 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]. The

9 König’s collections were described by both Colsman and Retzius, and are housed in Copenhagen and Lund, respectively.

8

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 the

Revised handbook for the Flora of Ceylon and by Press and Shresta (2000a), who list under

Scrophulariaceae Gratiola cordifolia Vahl, D. Don, Prodr. Fl. Nep.: 85 (1824). 'in Nepaliâ.

(Specimen not found). = Lindernia anagallis (Burm.f.) Pennell

Willdenow updated Linnaeus Sp. Pl. (1797) and created two names, Gratiola pusilla

[=Lindernia pusilla (Willd.) Bold.], and G. lucida [= L. crustacea (L.) F.Muell.]. He

omitted the reference to the König specimen made by Retzius when citing L. oppositifolia

Retz., which makes one wonder whether it was not clear at that time (1797) how important

type 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] in

Plants of the Coast of Coromandel II (1798) and III (1819). The type for G. parviflora is

often cited as an illustration from Pl. Corom. [sometimes as 203 (Fig. 2 a), or as 204, which

actually depicts G. rotundifolia). While 203 matches his description, his Icones 524 [Fig. 2

b] gives a much better depiction of the venation, which is 3—5-veined from the base. The

illustration of an open flower makes the plate very useful compared with the specimen

alone. 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 all

can readily be used in identification (in addition to the König specimens). His research is

summarized in Flora Indica of 1820 and Flora Indica ed. Carey of 1832. Roxburgh’s most

in-depth observations give extremely valuable references to the understanding of the Asian

Lindernia species.

Vahl described Gratiola cordifolia Colsman in Enumeratio Plantarum 1:95 (1804) without

reference to Colsman, but referring to his König type. Hence the species G. cordifolia Vahl

referred to in the literature is a synonym. Both names are now sunk into synonymy with

Lindernia anagallis (Burm.f.) Pennell.

9

Maximovicz (1827-1891) treated several species in Decas 19 of Diagnoses Plantarum

Novarum Japoniae et Mandshuriae. Here he stated that Linnaeus’ Capraria gratioloides

described in Systema Naturae 10 (2): 1117 of 1759 equalled Lindernia dubia L. from

Figure 2 a: Roxburgh’s illustration of Gratiola parviflora Roxb. Pl. Corom. 3: 3, t. 203 (1819); with detail.

Figure 2 b: Roxburgh’s illustration of G. parviflora Roxb. Roxburgh’s Icones 524, with detail; notice difference in venation compared with Fig. 2 a.

10

Species Plantarum 17 of 1753. His publication dealt with the errors that occurred in the

interpretation of this species identity; this extensive chapter on Lindernia All. promises

help to clarify the issues surrounding Allioni’s and Linnaeus’ descriptions. It cited

numerous works, which again cited numerous specimens with their European localities of

Lindernia 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. in

Asia. He stated that the latter, and Vandellia erecta Benth. are not conspecific, but a

transition exists between the two (as discussed in Fl. Ross). He further explained that a

cleistogamous form of L. procumbens probably exists, however, that its small, closed

corolla enticed plant collectors not to collect it, but rather to collect those plants with open

corollas instead. This work elaborated on the occurrence of Lindernia pyxidaria L. and

Vandellia 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 and

its affinities need to be elucidated, and because the new species from Nepal with a closed

corolla needs a name, Maximovicz’s work on the Asian Lindernia could prove important

during 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 Gratiola

Following the first century of descriptions and treatments, several authors attempted to

arrange the Lindernia complex into distinct genera. With increasing material, it had

become obvious that many of the species described as Gratiola were too distinct from the

other Gratiola spp. Genera had to be found to place these mainly tropical semi-aquatic

11

species. The genera created were to be used for the next century or so, until Pennell

successfully argued to have all of them merged into one genus, Lindernia (1935). The then

generic, now sub-generic distinctions were mainly based on androecium and length of fruit

(siliquosae versus brachycarpae), but dehiscence (Bentham 1835) and venation (Hooker

1884) were also suggested. For a while, Willdenow’s genus Hornemannia was used for

some short-fruited Lindernia [e.g., L. viscosa (Hornem.) Merr.] by Link and Otto (1820) I

recommend 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 he

made the important distinction “[...] stamina two fertile under the upper lip; anthers

unilocular; two sterile filaments under the lower lip. Ovary oblong; style compressed

above, 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 easily

amongst a pile of Lindernia specimens; these plants are decisively more slender than the

rest, 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 the

genus [L. ciliata (Colsm.) Pennell], emphasizing among other characters the unique long-

aristate margin which itself is framed in white, the narrow bracts subtending the – with

height of the plant shortening – pedicels, and interestingly, described the capsule as almost

quadrangular, 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 Selectarum

Horti Regii Botanici Berolinensis: 25, t. 11. Here they stated that the seed material was

brought from Manila by “Herr[n] Dr. von Chamisso.” The material did not withstand the

German climate but, planted in loose, sandy soil, seeded in the shelter of glasshouses. They

distinguished 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,

12

but the first description may possibly be of an earlier date (e.g., 1820; there is confusion in

the literature over abbreviations like Icon. Pl. Select. and Icon. Hort. Bot. Berol., which do

not refer to the same publication). According to Stearn (1972), this description of

Hornemannia ovata has been regarded as a combined description of a generic and specific

protologue by, e.g., Index Kewensis (as was the case in Bonnaya brachiata above), but

Link and Otto intended to only add another species to Willdenow’s Hornemannia genus.

Blume described the Scrophularinae of Buitenzorg/ Bataviam [Java] in Bijdragen tot de

Flora van Nederlandsch Indie(1825) and in it several species of Lindernia important to this

discussion. His treatment included Linnaeus’ Gratiola veronicaefolia, and Roxburgh’s G.

reptans and G. serrata. He distinguished these Gratiola L. from Diceros Lour., under

which 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 and

bilabiate (“labio superiore bilobo aut emarginato; inferiore trilobo”); the stamens having

two anthers with divergent locules, and two or three (! – this is the first time a botanist

observed the third, usually lost, anther) sterile anthers in Gratiola Linn.; in contrast, the

corolla being funnel-shaped, (“limbo subbilabiato, interdum subaequali”) and the stamens

didynamous, i.e. all 4 fertile in Diceros Lour. Blume, however, misunderstood Loureiro, as

Bentham (1846) in DC X points out, Diceros Bl., non Lour.)

He further distinguished the genus Mimulus L., in which he divided M. javanicus Bl. [a

synonym for L. crustacea (L.) F.Muell]. Distinguishing characters here were the tubular

calyx being 5-dentate (“quinquedentatis”) rather than 5-lobed (“quinquepartitus”), the

anther lobes divaricate, and the calyx completely covering the capsule (all these characters

making it more akin to Torenia, so Blume says). Lastly, he distinguished Torenia obtusa

Bl., which is a synonym for L. anagallis (Burm.f.) Pennell. This distinction was supported

based on the corolla character of “labio superiore obtuso aut retuso, ” and the anther

character of “antherarum lobis divaricatis, interdum effoetis. ” He admitted hereafter that

this genus is hardly distinguishable from Mazus.

Reichenbach made the combinations Tittmannia ovata (non Benth.)/ Tittmannia viscosa

from Hornemannia ovata Link & Otto and from Gratiola viscosa Hornemann in

Iconographia Botanica Exotica (1823/1824), herewith creating a new genus. According to

Bentham (1835), Reichenbach separated Tittmannia from all the other genera on the

13

grounds of ‘filaments bifid at base or appendaged, and from whence the fertile filaments

are arched over to the upper lip with connivent anthers’ by the short, not plicate calyx, and

a globular fruit [which were considered by Chamisso as the true Toreniae (and the ones

with a siliquose fruit and short calyx as the Vandelliae)]. The genus Tittmannia was

maintained by Bentham for several of his nomina nuda, but Bentham changed its status to

subgeneric 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 author

of one T. ovata, whereas the other is Bentham, whose Wallich type forms one basis of

Lindernia viscosa. It should also be noted that the basionym reference by Reichenbach to

G. viscosa Hornem. Hort. Hafn.: 19 is wrong, (and that to this day I have not seen the

protologue 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. veronicifolia

into Bonnaya.

Buchanan-Hamilton was the first botanist to collect in Nepal, i.e. 1802—1803. He was a

physician, who “spent 14 months in and around Kathmandu, recording information on all

forms of natural resources” (Press & Shresta 2000a). His botanical collection, which he

gave to Lambert (in London), formed the basis for Don’s Prodromus Flora Nepalensis (see

below). He published a few names in Lindernia, but none of them persist as basionyms.10

He moved Willdenow’s Gratiola lucida to Torenia lucida (which is L. crustacea), again a

move supported by both Blume, and Chamisso & Schlechtendahl, and also considered by

Bentham. He then created Torenia alba (already described as Capraria crustacea L.; hence

in one year (1831) he gave this taxon two different names, as can be traced in Wallich’s

Catalogue. An interesting specimen from Buchanan-Hamilton’s Nepal collection was

found 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 may

represent 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 that he gave a lot of his work to fellow contemporaries (e.g., Don, Wallich), who took over the publications without paying the tribute that we would expect today, and to the fact that a lot of his work remains unknown as of this day (Fraser-Jenskins, in preparation).

14

Don (D.) was the herbarium curator and librarian of Lambert. He was “comissioned [by

Lambert] with the approval of Hamilton” (Fraser-Jenkins, in preparation) to write an

account of the not yet studied Nepalese collections. Including Hamilton’s and Wallich’s

collections, but excluding Smith’s herbarium (to which Don did not have access) resulted

in 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 and

Vandellia in the sense of Linnaeus (Mant. 89) and Jussieu (Gen. 122). [Jussieu, in this

Genera 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 shortly

emarginate superior lip, an emarginate stigma and a bilocular capsule.] Unfortunately, Don

omitted most of the information that came with Buchanan-Hamilton’s detailed collection

notes; therefore, no specimen localities within Nepal can be traced through D. Don’s

Prodromus, nor through the Wallich Catalogue. While Don’s publication reads 1825, the

actual distribution of the material occurred before the close of the year 1824. This date

should 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 only

taxon within the Lindernia complex that has a tubular calyx with short lobes and with

seams (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 Botanical

Nomenclature, are nomina nuda; all specimens are in K-Wallich. They include Tittmannia

grandiflora Benth., (K-Wall. Cat. 3949!), now synonymized under L. anagallis (Burm.f.)

Pennell; Tittmannia angustifolia Benth., (K-Wall. Cat. 3951!), which had already been

described 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.

15

In Scrophulariae Indicae, Bentham used the genera Vandellia L., Bonnaya Link & Otto

(“filaments always abortive, simple, and club-shaped”) in the tribe Gratiolae to describe

most species of this treatment on Nepalese Lindernia, except Gratiola multiflora Roxb. and

Vandellia hookeri C. B. Clarke. In the beginning pages, elaborating the distinguishing

characters of all treated Scrophularineae, he commented (100 years ahead of Pennell) that

“perhaps it might be better to consider these three genera [Lindernia, Vandellia, and

Bonnaya] 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 tribe

Gratiolae (in 1835 solely encompassing Asian species). He called this subtribe the

Lindernieae (= at least posterior stamens fertile)11

, and herewith excluded them from the

genus Gratiola on the basis of the latter having a globose, 4-valved capsule. Gratiola was

placed 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 Bonnaya

Link & Otto, Vandellia L., Lindernia All. (which he knew only from Europe) together

with, 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 (however

included in Vandellia together with V. multiflora, V. hirsuta (= L. pusilla), V. scabra, and

V. glandulifera Bl. (both = viscosa), V. erecta (but L. pyxidaria was excluded into the only

Lindernia). He further listed V. nummulariifolia, V. pedunculata and V. angustifolia.

Treated as distinct from these were Ilysanthes gratioloides, I. parviflora, as well as

Bonnaya brachiata, B. reptans, B. veronici- and verbenaefolia, B. grandiflora, and B.

oppositifolia.

It becomes quite clear that Bentham worked mainly from herbarium material. Many

multiple entries treat the same taxon; had he had more material at hand, he might have

realized the synonymy between several of his listed species. Still, his comments, especially

those on the similarities between ‘species,’ are useful. For example, he treated Lindernia

pyxidaria, but explained that it looks like Vandellia erecta12

11 A cladogram of Bentham’s taxonomic concept as understood from DC X (1846) is attached in the appendix 3.

, Ilysanthes gratioloides, I.

capensis, and I. parviflora, all of which may indeed prove to form a tightly allied complex.

12 See section 5. 2. 2

16

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 part

of one complex on account of their long siliquose fruits, and (1) their racemose

inflorescence (most definitely erroneously including B. pusilla auctt.? in this list) – the

other part (2) consisting of B. grandiflora (Retz.) Spreng., B. peduncularis Benth., and B.

oppositifolia (Retz.) Spreng. with axillary flowers. The second complex was distinguished

by 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 Brachycarpae

on 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 the

calyx”, here listing V. nummulariifolia D. Don. He then treated the Siliquosae (with the

long pod), including V. diffusa, V. pedunculata, and V. angustifolia, ending with a few

poorly known species (V. cordifolia, V. roxburghii, and V. multiflora). Lastly he treated the

genus Lindernia, where he included L. pyxidara L.

Generally, his treatment closely followed Bentham’s Scrophularineae Indicae of 1835, but

included interesting facts about the eponomy of the genera. One nomenclatural question

begs 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 all

indigenous 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 he

pointed 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

17

taken in section 5. 2. 2, where the differences between B. parviflora and V. erecta are

explored.

Haines in The Botany of Bihar and Orissa (1922) reduced the four genera to two uniting

Bonnaya 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 Handbook

to the Flora of Ceylon (1931). This mistake in synonymy almost certainly must have had to

do with Alston not seeing Burman’s type from Java, as it could not have escaped Alston

that the Java specimen is synonymous not with R. antipoda, but with the Lindernia

cordifolia (Colsm.) Merr. that he described in the same account. This speculation was also

made by Philcox (1968). Alston’s fault was recalled and changed 50 years later by Cramer

in A revised handbook to the Flora of Ceylon 3 (1981). In the meantime, wrong

determinations of herbarium material were frequent.

Pennell in his Scrophulariaceae of eastern temperate North America merged all genera

described 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 anther

character [four anthers in Lindernia and two anthers in Ilysanthes] separates L. pyxidaria of

Europe 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 much

as an ultimate stage of evolutionary change, actual kinship being in other characters.” He

took 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 Brittonia

he argued that Torenia and Lindernia are separate, though a clear distinction is not easy to

18

draw. In Lindernia, the calyx is usually more or less deeply lobed, i.e. the sepals are

separate, whereas in Torenia, the calyx is united, with raised ridges above the five central

veins. However, there are exceptions to the rule, since L. crustacea has united sepals. A

second distinguishing character is Torenia’s larger corolla with an angular, open throat, and

Lindernia’s smaller corolla with a flattened throat. The capsule in Torenia is usually

completely covered by the calyx, whose apices are connivent, whereas in Lindernia the

capsule usually exceeds the calyx in length, and the calyx lobe apices point outwards. As

for similarities, Torenia also has a tendency to suppress the anterior stamens. It is that year

of 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 a

new species, Lindernia crenata Pennell from this expedition.

Pennell did not treat L. anagallis and L. cordifolia as conspecific – he said that L. cordifolia

has anterior filaments which are anther bearing, as opposed to L. anagallis without anterior

anthers (maybe that is the case in New Guinea). His description of L. cordifolia hints to a

mix 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 of

the Flowering Plants of Nepal, and by Mill in Flora of Bhutan). It could not be verified by

me that the Hungarian botanist made this new combination. Bekesvarmegye floraja [= The

flora of Bekes county] is cited for the location of the description, but in this to Scottish

botanists slightly obscure Journal [also cited as Magyar Tudomanyos Akademia.

Ertekesezek a Termeszettudomanyok Korebol. Kot. 11. szam 18 (1881)], Borbás wrote

Gratiola officinalis L. [...] Pyxidaria procumbens (Krock.) (Lindernia pyxidaria All.),

herewith showing the connection between the genera Pyxidaria, Anagalloides, and

Lindernia, but not officially combining Lindernia procumbens (Krock.) Borbás.

19

Von Mueller, in Fragmenta Phytographiae Australiae (1882), listed Gratiola pedunculata

(Br. Pr. 435)13

as dispersed throughout the whole of “Australiam extratropicam” but was

not 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 well

as G. serrata Roxb. (1793) into Lindernia, not aware that G. ciliata had already been

described by Colsman. I am fond of the name pedunculata because L. anagallis has the

longest 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 extent

of division of the upper lips, then the nature of the lower lips in great detail. He then went

on to describe the two forms of corollae, the closed kind and the opened kind, which

according to Urban can occur on the same plant, i.e., chasmogamy and cleistogamy seems

to depend on geographic location, but added that a closed corolla may have to do with

water-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 Asian

species of Lindernia pyxidaria are more adapted to pollination, while the European species

tend to inbreed. While Urban found anther characters good to distinguish the genera, he

later went on to hypothesize sliding scales rather than clear distinctions between the genera

listed 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 the

Himalaya [as well as L. sessiliflora (Benth.) Wettst.] as those with clayx 5 toothed, after

flowering 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] from

the whole of Southeast Asia, and L. scabra (Benth.) Wettst. [L. pusilla, in SE-Asia as well

as S. Africa and Madagascar] as those with a capsule not longer than calyx. Appendages on

13 I am fond of the name Gratiola pedunculata, because this species has the longest peduncles of all Nepalese Lindernia.

20

anterior filaments were described as “very small and bumpy [höckerig].” It is here that he

mentioned L. pyxidaria All. as flowering mostly cleistogamously in temperate Asia (= L.

pyxidaria All. et Aut.), and flowering only chasmogamously (= Vandellia erecta Benth.) in

East-India, and that both forms occur in localities between.

Sect. III Hornemannia Link et Otto Calyx 5-toothed. Capsule not longer than calyx

included 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 where

Hornemannia 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 had

been sunk into synonymy with L. antipoda: Vandellia grandiflora (Retz.) Merr., which

according to Philcox is conspecific with L. philippinensis. Merrill was the first author to

write about the ecology of Lindernia (1912). In these non-systematic Notes on the Flora of

Manila with special reference to the introduced element he spoke about the geographical

origin of species, stating that “in most treatments of tropical floras, whether of the eastern

or the western hemispere [...] species [such as Oxalis repens Thunb., Sida cordifolia L., and

Evolvus alsinoides L., i.e. various grasses and sedges] are usually listed as native ones. It is

far more probable that some have originated in one hemisphere [...] they have been

accidentally distributed by man within the past 400 years [...]and in most cases it is

probable that their original homes will never definitely be known.” He went on to explain

that “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 or

feathers of migratory wading or swimming birds.” A last quote, “while it is only reasonable

to suspect that most have been introduced by man (by sticking to clothing, to the hair of

animals) 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 lists

Vandellia crustacea Benth. as accidentally introduced, and of Oriental or European origin.

Boldingh in Zakflora voor de Landbouwstreken op Java (1916) made the combinations for

L. pusilla and L. viscosa. Since he omitted the basionyms, it is not clear whether Boldingh

referred to Thunberg’s Selago pusilla (1794) or to Willdenow’s Gratiola pusilla (1797), or

to another species altogether. Because the description of S. pusilla did not contradict the

21

species, I chose to favour Thunberg’s basionym as the earlier one. During this research, I

referred 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 were

illegitimate due to the fact that they were published in a descriptive key; the ICBN

however, appears to allow publications in descriptive keys pre-dating 1925, hence

Boldingh’s combinations can be accepted for the Flora of Nepal.

Backer in Onkruidflora der Javasche Suikerrietgronden (1931 – 34) continued Boldingh’s

work 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. van

Steenis). Here he included several illustrations, maintaining a narrow-leaved form of L.

anagallis var. angustifolia.

3.4 Recent floristic accounts

Mukerjee (1945) in his revision of the Indo-Burmese species of Lindernia listed 28 species

for this region. It is in this work that Mukerjee made the new combinations Lindernia

multiflora (Roxb.) Mukerjee and L. oppositifolia (L.) Mukerjee; the basionym authority of

the latter should be Retzius, not Linnaeus. In synonymy with L. pyxidaria he cited

Vandellia erecta Benth. and Gratiola integrifolia Roxb. This revision lacks species

descriptions and references to type material. Sivarajan & Mathew revised the genus

(1983) with 22 species for India, including species descriptions, which were lacking in

Mukerjee’s previous treatment.

Banjeri (1958) gave a detailed description of his expedition to Eastern Nepal; in his list of

collections 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 includes

a 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 Gratiola

ruellioides Colsm. (Type in Copenhagen) are distinct on account of the projection on the

teeth of the lamina. He further found that G. ruelliodes Colsm. and G. reptans Roxb. are

22

conspecific. In 1968, Philcox revised the Malesian species Lindernia, giving a reference for

cytologic 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 that

time. Philcox’s determination slips date among the latest found on herbarium sheets, and

several 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 experiments

concerning 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 is

remote, further concluding that capsules can form through self-pollination, and in some

cases can produce parthenocarpically.

Diaz Miranda (1977) included in his morphological account of Lindernia in South

America a phytochemical survey of leaf flavonoids research. The characters14

were found

to mirror the four sections Torenioides, Nummularia, Lindernia, and Brachycarpae. He

treated L. crustacea, placing it in Pennell’s section Torenioides, secondly L. diffusa (with

almost sessile calyces), thirdly Lindernia procumbens (Krock.) Philcox, synonymizing it

with 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, which

he synonymized with Capraria gratioloides L. and Ilysanthes riparia Rafin. Diaz Miranda

also 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 a

pantropic genus of about 80 species predominantly palaeotropic. He pointed out that until

Philcox traced Burman’s type of Ruellia anagallis, this taxon had been treated conspecific

with Ruellia antipoda L. With this revision of the Flora of Ceylon, he clarified Alston’s

combination 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 be extended beyond the five taxa treated here.

23

anagallis auctt. (non Burm.f. 1768). Further he commented that “the Lindernia-Ilysanthes

complex, 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 kept

Vandellia separate fron Lindernia in the Flora of Eastern Himalaya (1966). However, he

also sunk Lindernia ruellioides in synonymy with Vandellia antipoda (L.) Yamazaki. This

move meant a lot of confusion over the identity of these two taxa, which can be seen in

herbarium 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’s

synonymy of genera. His treatments of Lindernia spanned many years of consecutive

sections in New and noteworthy plants of Scrophulariaceae from Indo-China, all of which

were supplied with high-quality line drawings (Yamazaki 1978, 1980, 1983). He wrote the

Revision of the Indo-Chinese species of Lindernia All. (Scrophulariaceae) in 1981. A later

chapter will refer to his sub-generic classification of Lindernia.

Quail Lewis15

(2000) revised the New World Species of Lindernia, treating the L. dubia

complex as four distinct varieties, and treated an overall 12 distinct species. Although

dealing with New World taxa alone, the treatment is useful for Nepal in its overview of the

morphological characters, its chapter of the generic affinities, and its references to

glasshouse flowering experiments.

4. MATERIALS AND METHODS

The study used of herbarium specimens from Edinburgh (E, some of which included digital

images of live plants before collection), Kew (K), the Museum of Natural History (BM),

Nationaal Herbarium Nederland at Leiden (L). Specimens were also observed at the

Linnean Society (LINN). Digital images of type material were obtained from the

University 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 library

of 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 other works citing her as Lewis 2000. I am not sure whether some of her work appears elsewhere under Quail Lewis.

24

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 sheets

was included for Nepal, of which at least three specimens per species (data permitting)

were measured to cover morphological variation, and hence to achieve representative

ranges for each Nepalese species. As mentioned by Davis and Heywood (1973),

“classification based primarily on herbarium material should not be pushed too far, unless

field 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 begun

to change with increased digital imaging of live plants. Colour images of live plants were

used in this section wherever possible. Sixty five characters found useful for identification

were scored in an Excel spreadsheet,16

a synthesis of which was used to create a

dichotomous, artificial 13 couplet-long key to the species. The descriptions were limited to

ca. 150 words, again focussing on characters aiding identification.

A Leo Supra 55VP Scanning electron microscope (SEM) was employed to investigate and

compare seed morphology of all thirteen taxa. Additionally, a pilot study to examine

differences in lamina structure was carried out. Since the herbarium specimens were

already dry, the untreated seeds were mounted on Agar Scientific Carbon tabs and

sputtercoated with platinum in a peltier cooled EMITECH K 575X. One advantage of not

hydrating and critical-point drying the seed material was that characteristcs actually change

when 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 accurate

measurement of seed dimensions.

The results were used to investigate the value of SEM study for species delimitation, as

well as their coherence with subgeneric delimitations established in previous literature on

this taxonomic group.

16 Appendix (6; CD-ROM).

25

4.1. MORPHOLOGICAL CHARACTERS

Duration and Habit. The Nepalese species of Lindernia are annual herbs. L. hookeri has

previously been described as a perennial herb in, e.g., the Flora of Bhutan, but no evidence

for this was found in Nepalese material. D.Don describes L. micrantha as a perennial (sign

interpreted in Stearn), but again the material available in this study did not exhibit this

character. The species of Lindernia are either erect, spreading, creeping, or prostrate. Their

height 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 to

short and slender (Fig. 4.1 b) as in most other species. Some decumbent species always

root at the nodes (e.g., L. ruellioides) while others do not exhibit this character at all.

Hairs and other structures. In this study, the use of this character was mainly based on

presence and absence. Only a few species of Lindernia are hairy. However, on glabrous

plants, particular parts exhibit small, short teeth, which ‘embroider’ the margins and apices

Fig. 4.1 a: Roots of Lindernia a.

Fig. 4.1 b: Roots of L. procumbens

26

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). Other

literature 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, the

stem 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 inversely

proportional to leaf size. Lastly, care should be taken when defining the state of the stem: It

may appear densely hairy at the nodes, but in fact perceived hairiness at the nodes is almost

always an increased hairiness of the decurrent lamina base margin.

Fig. 4. 2 b: L.antipoda.(L.) Alston Lamina margin serrate and ciliate. Not to be confused with c., L. ciliata, which is not ciliate. Also visible here are the sunken stomata.

ciliate margin ▲

Fig. 4. 2 a: L. micrantha (D.Don) Wettst. Lamina margin distantly, obscurely serrate, and smooth.

Fig. 4. 2 d: L. parviflora (Roxb.) Mukerjee Lamina margin entire to obscurely 2—3 dentate, and smooth.

Fig. 4. 2 c: L. ciliata (Colsm.) Pennell Lamina margin long-aristate serrate.

27

Stems. All stems are quadrangular, and are winged to some extent. The most extreme

examples of a winged stem can be found in L. hookeri (Fig. 4. 3 c), L. pusilla (Fig. 4. 3

a), and L. multiflora. Stem diameters were measured near the base of the plant as the

width of one of the four sides of the quadrangle; they ranged between 0.5—2 mm. The

taxa often referred to as of the subgenus Ilysanthes (i.e. L. parviflora) are the most

Fig. 4. 2 e: L. hookeri (C.B.Clarke) Wettst. Lamina margin serrate and ciliate; usually bearing a longer protrusion on the projecting part of the margin.

Fig. 4. 2 f: L. hookeri (C.B.Clarke) Wettst. Lamina base is often densely beset with long hairs, whereas the apex has none [see 4. 1 e]. This phenomenon is shared by a few Nepalese species. Calyx (inset) also strongly hairy.

Fig. 4. 2 g: L. anagallis (Burm.f.) Pennell Lamina margin serrate, but ciliate often only on the projecting part of the margin. A further character exhibited by many species of Lindernia (see also b + f) is the red colour of the margin.

Fig. 4. 2 h: L. ruellioides (Colsm.) Pennell Lamina margin deeply serrate, and ciliate.

ciliate margin ▲

28

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.

Leaves. The leaves in Lindernia are highly variable in shape (base, margin, apex) as well as

venation. They can be almost circular (L. nummulariifolia, L. pusilla) or linear and very

narrow (L. micrantha), almost rhombic (L. indet-a), or lanceolate (L. hookeri), and shape

can even vary within an individual plant. As Roxburgh (1832) pointed out, the shape of the

leaves in L. ruellioides varies between round on non-flowering and oval on flowering

shoots on the stoloniferous plants.

Fig. 4. 3 a: L. pusilla (Thunb.) Bold. Stem strongly winged. Also visible are the flattened hairs attached to the angles.

Fig. 4. 3 b: L. anagallis (Burm.f.) Pennell Stem winged., not hairy. Also visible is the reddening characteristic of this taxon.

Fig. 4. 3 c: L. hookeri (C.B.Clarke) Wettst. Stem winged, sparsely hairy on angles.

Fig. 4. 3 d: L. parviflora (Roxb.) Mukerjee Stem winged, not hairy.

29

Occasionally, the presence or absence of the petiole has been used as a character, but in

general 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 the

lamina 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. In

the 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) to

moderately (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 in

several cases to count the number of incisions per side of the lamina. The numbers are

surprisingly 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 the

lamina 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. 4

a + e), or pinnately veined (Fig. 4. 4 c, f + g). Just like the angles of the stem, veins tend to

be coloured reddish in a few species.

30

Fig. 4. 4 a: L. micrantha (D.Don) Wettst. Lamina obscurely serrate. Leaf length to leaf width ratio 7:1. Petiole not easy to define.

Fig. 4. 4 b: L. indet-a. Three to five-veined from the base. Lamina sub-entire to undulate.

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 f: L. ruellioides (Colsm.) Pennell

Fig. 4. 4 e: L. ciliata (Colsm.) Pennell Lamina uni-veined

31

Nodes. The nodes of the stem in Lindernia are very much influenced by the manner of

attenuation or petiolation of the opposite leaves. As mentioned above, the hairiness often

associated with the base margins of ciliate laminas tend to make the nodes appear hairy; the

origin of the hair, however, is the leaf and not the stem. Increased hairiness in these areas

seems to be associated with the fusion of tissue.

Inflorescence. Two main types of inflorescence were recorded: (A) About half the species

have solitary flowers in the axils of the leaves. These are frequently supported by pedicels

as 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 of

the plant. These taxa, namely Lindernia parviflora and L. procumbens may appear

subracemose 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 the

leaves 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 where

subtending the first pair of pedicels. Instead, they are small and shaped differently than the

leaves on the lower, vegetative part of the plant; the terminal raceme was defined as the

presence of a marked, abrupt change in leaf size from the vegetative to the flowering part

of the plant, neglecting the location of the first flowering internode, which often classified

Fig. 4. 5 b: L. anagallis (Burm.f.) Pennell

Fig. 4. 5 a: Lamina base and node of L. muliflora (Roxb.) Mukerjee

Fig. 4. 5 c: Lamina base and node of L. anagallis (Burm.f.) Pennell

32

as intermediate to, or same as vegetative part of the plant. The axillary, solitary type often

produces only one pedicel per node, and sometimes a shoot out of the opposite axil (4.6 c

and d).

Flowering time. Flowering times were estimated using the dates of material from Nepal. It

is possible that the flowering season may be slightly longer. The season spans roughly from

March to November.

Fig. 4. 6 a: Terminal raceme of L. multiflora Pedicels slightly reflexed.

Fig. 4. 6 b: Solitary, axillary inflorescence of L. pusilla

Fig. 4. 6 c: Solitary, axillary inflorescence of L. anagallis

Fig. 4. 6 d: Solitary, axillary inflorescence of L. parviflora

33

Calyx. The calyx consists of five lobes usually of slightly unequal length. Lindernia

crustacea 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 the

calyx, but glabrous taxa have glabrous calyces. If the calyx itself is glabrous, its calyx lobe

margins 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 may

appear hairy due to ciliate margins (Fig. 4.7 d).

Fig. 4. 7 a: 5-dentate calyx of L. crustacea

Fig. 4. 7 b: bilabiate calyx of L. hookeri

Fig. 4. 7 c: 5-partite calyx of L.antipoda

Fig. 4. 7 d: Glabrous calyx with ciliate margin on apex of lobe (L. anagallis)

Fig. 4. 7 e: Hairy calyx without ciliate margin (L.pusilla)

34

Corolla. The corollae in Lindernia are bilaterally symmetric, semi-closed, and bilabiate.

The corolla of L. indet-a is closed, revealing cleistogamy. While accurate measurements

can 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 upper

lip is shallowly 2- lobed, or not lobed at all, and ca. 2—4 mm. Corolla length ranges from 4

to 12 mm. The central lower lip lobe length can reach to 4 mm. Corolla colours range from

white to yellowish to blue to purple, but are never red because they are pollinated by small

insects, probably bees, not birds or bats. Comparison of the corollas indicates the close

relationship between L. ruellioides and L. antipoda. Their similarities can be acknowledged

by comparison of well-pressed specimens (Figs. 4. 8 a + b below).

Androecium.

All species of Lindernia are didynamous (Judd & Olmstead 2004), i.e. the length of the

anterior 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

Fig. 4. 8 a: L. antipoda

Fig. 4. 8 b: L. ruellioides

35

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 the

anterior pair bears sterile ones (e.g., L. hookeri Fig. 4. 9 e). The last case features (3) a

posterior 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 is

often 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 the

arrangement is just the reverse,” but this arrangement was not observed in Nepalese

material. Often both pairs are connivent, sometimes the anterior staminodes are not

connivent, but parallel.

36

Fig. 4. 8 c: L. anagallis (Burm.f.) Pennell DNEP 2B 191

Fig. 4. 8 d: L. parviflora (Roxb.) Mukerjee DNEP 2B 125

Fig. 4. 8 g: L. pusilla (Thunb.) Merrill

Fig. 4. 8 h: L. micrantha D.Don Herbarium Specimen EMAK soaked in OT2

Fig. 4. 8 f: L. nummulariifolia (D.Don) Wettst.

Fig. 4. 8 e: L. antipoda (L.) Alston

37

Fig. 4. 9 a: Floral dissection of L. crustacea (L.) F. Muell.; WIL 326

Fig. 4. 9 b: Floral dissection of L. nummulariifolia (D.Don) Wettst.; STA 4018. Scale bar denotes millimeters.

Fig. 4. 9 e: Floral dissection of L. hookeri (C.B.Clarke) Wettst.; SSW 6767

Fig. 4. 9 f: Floral dissection of L. micrantha (D.Don) Wettst.; EMAK 174. Soaked in 6 (10 g aerosol OT2 / 100 g H2O) / 1 (acetone). Note upper lip removed; anthers dorsifixed; appendage club-shaped.

Fig. 4. 9 d: Floral dissection of L. ruellioides (Colsm.) Pennell; SSW 9272. Anterior stamens rudimentary.

Fig. 4. 9 c: Floral dissection of L. pusilla (Willd.) Bold.; as no flower for Nepal was available, a specimen from area 6 was substituted.

post. anther ►

filament appendage ▲

ant. anther ►

38

Gynoecium. The gynoecium is bi-carpellate, and the placentation axile. The style is

penicilliform, simple, but often flattened into two stigmatic lobes at the apex. In some

species, the style is persistent until dehiscence of the capsule (Fig. 4. 10 e, f + j); in other

species, it leaves a ring on the capsule where it was attached, but falls off before the fruit

ripens. (In Pennell’s words (1943b) “[the capsule of Lindernia procumbens is] tipped by a

white 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 (C.B.Clarke) Wettst.

Fig. 4. 10 b: Dehisced - and closed capsule of L. ciliata (Colsm.) Pennell

39

Fig. 4. 10 c: L. multiflora (Roxb.) Mukerjee

Fig. 4. 10 d: L. nummulariifolia (D.Don) Wettst.

Fig. 4. 10 f: L. viscosa (Hornem.) Merrill

Fig. 4. 10 e: L. anagallis (Burm.f.) Pennell

Fig. 4. 10 g: L. pusilla (Thunb.) Bold.

Fig. 4. 10 h: L. crustacea (L.) F. Muell.

40

Seeds. 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 can

be green during different stages of maturity (Fig. 10 k above). Most seeds of Nepalese

Lindernia are alveolated, i.e., the seed coat is depressed between protruding ridges.

According to Rahmanzadeh, three goups can be distinguished within Linderniaceae based

Fig. 4. 10 i: L. antipoda (L.) Alston

Fig. 4. 10 j: L. parviflora (Roxb.) Mukerjee

Fig. 10 k: L. ciliata: Capsule with seeds

41

on seed characters. The first group is characterized by alveoli, exhibited also by

Craterostigma Hochst., parts of Artanema and parts of Torenia. The second group is

characterized by those taxa not exhibiting alveolated seeds, part of which are the Lindernia

s.str., represented by, e.g., L. parviflora. The third group of the Linderniaceae,

characterized by aulacospermous seeds (Rahmanzadeh et al. 2005) was not represented by

Nepalese Lindernia. Alveolation is strongly associated with appendages, or hook-like

thickenings 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), sometimes

long 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 to

appression to the other seeds in the often tightly-packed capsule. The testal patterns show

different densities of tuberculation.

42

5. RESULTS AND DISCUSSION

The account for the genus in Nepal is attached in the appendix (1). This chapter deals first

with results from the SEM seed study (5.1), followed by a comparison of Yamazaki’s

subgeneric limits with those subgeneric distinctions apparent in seed characteristics (5.1.2).

Secondly, this chapter discusses specific taxonomic problems and the research needed to

solve 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 or

ribbed. The seed coat is alveolate or pitted (e.g., L. nummulariifolia), coarsely or irregularly

ribbed (L. hookeri), or areolate (L. procumbens). In this study, four sections could be

distinguished.

The seeds of the Nepalese species of Lindernia show considerable variation and a rough

grouping of species was possible by comparison of features such as their alveolate ridges,

appendages, and tuberculate seed coats. The comparisons in this chapter, summarized in

Table 5 below, are made under the assumptions that like was compared with like, but in

some cases the results can but guide further research. Further, care must be taken when

generalizing from a small study like this one: the seeds used most certainly were collected

at different stages in the life cycle, and were most certainly exposed to different drying

methods. 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 this

particular sample. It is known, however, that scrophulariaceous seeds are often influenced

by tight packing within the capsule (Elisens & Tomb 1983; Lewis 2000). This study

confirmed 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 now

clear that the taxon initially determined as L. procumbens is actually L. parviflora, and

therefore 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 /

43

L. procumbens?’ throughout. A separate, post-SEM study was conducted, comparing seeds

of L. parviflora with what is now hypothesized to represent L. procumbens. While this

study needs further investigation, and L. procumbens, once identified, should be included

in 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 been

treated 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 Lindernia

parviflora and L. multiflora (0.22 x 0.16 mm) to the more than twice as large (0.52 x 0.4

mm) seeds of L. ruellioides (note reduced magnification in micrograph). Although seed

size within one accession did not vary much, samples in which the seeds were convoluted

showed 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 strongly

convoluted (L. anagallis, L. antipoda, L. hookeri, L. crustacea, and L. multiflora). The

dimensions given in Table 5 and in the captions of Figs. 5. 2 a—p are taken from the

micrographs; it is obvious that the width of the strongly flattened L. anagallis seed, for

example, is smaller than the width measured in the micrograph. These convoluted seeds

would benefit from the soaking and critical point drying method applied to material before

mounting 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 wider

than 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 of

the 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 that

frame six relatively flat sides. As to the taxon represented by Norkett 8037 (Fig. 5.2 i), the

44

densely rugose appendiculation leaves the shape underneath open for speculation. Here, a

latitudinal dissection of the seed would be of great value.

Colour. The colours of the seeds ranged from yellowish-orange to red to bright green. The

colour of the seed was not used to distinguish taxa; it is likely to vary with maturity of the

seed. More important are differences in angularity and size (Mill pers.comm. Aug. 2005).

Longitudinal and latitudinal alveolate rows. Longitudinal alveolate rows are known from

other 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. Taxa

could be grouped into those whose seeds exhibit alveolate rows (Lindernia s.l.), and those

which do not (Lindernia s. str., Rahmanzadeh et al. 2005). Where they do, the longitudinal

and latitudinal rows form a structural network between which the seed coat is depressed by

approximately 20 μm. Yamazaki (1981) used the term ‘scrobiculate’ to describe this

character 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 are

shallower than in those seeds mentioned above. The same phenomenon arose in the

interpretation of L. ruellioides, for example, which seems to have but obscure longitudinal

rows and depressions (hence the entry in Tab. 5: n/a). This difficulty in discerning the

number of rows on a seed may be due to the absence of convolution. Therefore, counting

the 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 and

that of mere ‘angulation.’

A row was defined as housing depressions of the seed coat. Angulation was defined as a

simple structural line not associated with a depression in the seed coat. This distinction was

also made by Lewis (2000), who described the seeds as ‘faintly to strongly angled or

ribbed). Lindernia antipoda and L. ciliata had the highest estimated number (10—12) of

longitudinal 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

45

rectangular, and framed by crisp latitudinal ridges. Lindernia indet-a is distinct in that its

seed coat is mildly convolute, however it does not have depressions associated with the

alveolate 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 (see

Tab. 5; depression number, “DEP#”). In most cases, this number is a function of the

number 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, and

circumstances during the sample’s drying. Apparently distinct from the majority of taxa

examined are Lindernia indet-a, which does not show alveoli, and L. parviflora, whose

depressions are elongated rather than circular. Lindernia viscosa also stands out due to its

angulation and the lack of appendages (see following paragraph).

Appendiculation of testa. All but two taxa show some form of appendaged tissue attached

to 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 can

be seen that L. indet-a has an indistinct reticulate pattern, much reduced in comparison with

the strong reticulation exhibited by L. pusilla (Fig. 5.1 n and 5.2 N). Lindernia ruellioides

exhibits 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 belong

to the same (i.e., L. parviflora) taxon, and that G – > O – > I are a series of increasingly

mature seeds. When comparing Figs. 5.2 α + β, the differences in appendage dimension

between 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. anagallis

rather than L. ruellioides.

Lindernia crustacea and L. multiflora (Figs. 5.2 J + K) have filiform appendages. While

several seeds were sampled of each taxon, and all show this form of appendiculation, it is

not clear whether the origin of this tissue derives from the inside of the capsule, or whether

it is an actual outgrowth from the seed coat. Lindernia pusilla is the most distinct species of

46

all Lindernia; its reticulate wall thickening (Barthlott 1981; Canne 1979) is not found in

any other seeds in this study, except indistictly in L. indet-a. Distinct from the majority

here 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, whose

testal pattern is granular (Fig. 5.2 m + α), reminiscent of a stucco wall. L. anagallis has

tuberculate 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 the

appendage. 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 the

species, but an even greater magnification would have to be applied to properly analyze

these differences. However, when comparing the general appearance of tuberculation in the

Lindernia 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 group

are Lindernia ruellioides (granular), L. viscosa (densest), and L. indet-a (partially lacking

obvious tuberculation; Fig. 5.2.H).

47

Lindernia COLL # LTH WTH SHP ALV LO-R LA-R APP. C. DEP DEP # TESTA

anag. SSW 569 0.28 0.22 R

Y 7--9

8—10 W+S Y M 56--90 TUB+SM

antip. PSW 5664 0.28 0.24 R

Y

10--12

8—10 W+S Y M

80--100

TUB- WK

hook. SSW 6767 0.4 0.33 R

Y 6--8 5—6 W+S Y D 30--48 TUB

cili. SSW 7581 0.28 0.24 A

Y

10--12 7—8 W+S N D 70--96 TUB

micr. PSW 5856 0.36 0.22 A--L

Y 7--9 6—8 W+S N D 42--72 TUB

num. SSW 4018 0.45 0.25 R

Y 8--10 8 ca W+S N M 64--80 TUB

parvi. TI 1492 0.24 0.12 A--C

N 6 ca 6—7 R N S n/a TUB

indet-a. PSW 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

crust. PSW 5624 0.38 0.29 R

Y 8--10 6 F Y D 48--52 TUB

multi. BUC 1387 0.22 0.16 R

Y 7--9? 6—7 F+W Y D 42--60 TUB

visco. SSW 7056a 0.35 0.26 A

N 7--9 6—7 0 N S n/a

TUB-DEN

ruell. TI 1484 0.52 0.4 R

n/a n/a n/a W+L N S 32--36 GRAN

pusi. TI 1478 0.38 0.39 R--A

n/a 7--8 5—6 0 N S 32--36 TUB

parvi SSW 8696 0.24 0.14 A--C

N 6 ca 6—7 R N S n/a TUB

Mazus delavayi

DNEP 2B67 1.6 0.67 R

Y

14--16 n/a W+S N M n/a WK

Tab. 5: Seed and seed-surface characters from SEM study. Columns show taxa; COLL#= collectors’ codes [see below for 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 of depressions 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 of collectors 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

48

Fig. 5.1 a. L. anagallis (Burm.f.) Pennell SSW 569 Dimensions: 0.28 x 0.22 mm. Longitudinal alveolate rows: 7—9. Latitudinal ridges: 8—10. Surface scrobiculate/ depressed in pits between ridges, ca. 56—90 pits/ seed. Tendency to convolution. Appendages present. Fig. 5.1 b. L. antipoda (L.) Alston PSW 5664 Dimensions: 0.28 x 0.24 mm. Longitudinal alveolate rows: 10—12. Latitudinal ridges: 8—10. Surface scrobiculate/ depressed in pits between ridges, ca. 80—100 pits/ seed. Tendency to convolution. Appendages present. Fig. 5.1 c. L. hookeri (C.B.Clarke) Wettst. SSW 6767 Dimensions: 0.4 x 0.33 mm. Longitudinal alveolate rows: 6—8. Latitudinal ridges: 5—6. Surface scrobiculate/ depressed in pits between ridges, ca. 30—48 pits/ seed. Tendency to convolution. Appendages present.

49

Fig. 5.1 d. L. ciliata (Colsm.) Pennell SSW 7581 Dimensions: 0.28 x 0.24 mm. Longitudinal alveolate rows: 10—12. Latitudinal ridges: 7—8. Surface scrobiculate/ depressed in pits between ridges, ca. 70—96 pits/ seed. Raphe visible. Appendages present. Fig. 5.1 e. L. micrantha D.Don PSW 5856 Dimensions: 0.36 x 0.22 mm. Longitudinal alveolate rows: 7—9. Latitudinal ridges: 6—8. Surface scrobiculate/ depressed in pits between ridges, ca. 42—72 pits/ seed. Raphe well visible. Appendages present. Fig. 5.2 f. L. nummulariifolia (D.Don) Wettst. SSW 4018 Dimensions: 0.45 x 0.25 mm. Longitudinal alveolate rows: 8—10. Latitudinal ridges: ca. 8. Surface scrobiculate/ depressed in pits between ridges, ca. 64—80 pits/ seed. Appendages present.

50

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. 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.

XX

51

Fig. 5.1 j. L. crustacea (L.) F. Muell. PSW 5624 Dimensions: 0.38 x 0.29 mm. Longitudinal alveolate rows: 8—10. Latitudinal ridges: ca. 6; not strong. Tendency to convolution/ scrobiculation. Estimated number of depressions 48—52. Surface depressions well visible. Long hair-like appendages. Fig. 5.1 k. L. multiflora (Roxb.) Mukerjee BUC 1387 Dimensions: 0.22 x 0.16 mm. Longitudinal alveolate rows: 7—9? Latitudinal ridges: 6—7; not strong. Estimated number of depressions 32. Surface deeply depressed in pits between ridges, ca. 42—63 pits/ seed; strongly convolute. Fig. 5.1 l. L. viscosa (Hornem.) Merrill SSW 7056a Dimensions: 0.35 x 0.26 mm Longitudinal ridges: 7—9? Latitudinal ridge: 6—7; Surface depressions discernible, but shallow.

52

Fig. 5.1 m. L. ruellioides (Thunb.) Bold. TI 1484 Dimensions: 0.52 x 0.4 mm. Longitudinal and latitudinal ridges not protruding. Conspicuous, branched trichome-like appendages covering shallow depressions in coat. Estimated number of depressions 32—36. Note magnification reduced to 500 x. Fig. 5.1 n. L. pusilla (Thunb.) Bold. TI 1478 Dimensions: 0.375 x 0.385 mm. Longitudinal ridges: 7—8. Latitudinal ridges: 5—6. Strongly reticulate seed wall thickenings. Estimated number of depressions 32—36. Note magnification reduced to 500 x. Fig. 5.1 o. L. parviflora (Roxb.) Haines SSW 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 in seed coat 36—42; not pit-like, but rather laterally elongated.

53

100 μm

100 μm

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. Fig. 5.2 C. L. hookeri (C.B.Clarke) Wettst. SSW 6767 General lack of trichome-like appendages. Density of tuberculation not variable.

54

100 μm

100 μm

100 μm

Fig. 5. 2 D. L. ciliata (Colsm.) Pennell SSW 7581 Trichome-like appendages. Density of tuberculation not variable. Fig. 5. 2 E. L. micrantha D.Don PSW 5856 Trichome-like appendages. Density of tuberculation variable. Fig. 5.2 F. L. nummulariifolia (D.Don) Wettst. SSW 4018 Trichome-like appendages. Density of tuberculation not variable.

100 μm

100 μm

100

μm

55

100 μm

Fig. 5. 2 G. L. parviflora (Roxb.) Haines TI 1492 Trichome-like appendages rugose/ rounded. Density of tuberculation not variable. Fig. 5.2 H. L. indet-a PSW 4476. No appendages. Seed coat relatively smooth; occasionally minutely tuberculate. 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.

100 μm

56

Fig. 5.2 J. L. crustacea (L.) F. Muell. PSW 5624 Filiform cuticular projections. Density of tuberculation variable. Fig. 5.2 K.. L. multiflora (Roxb.) Mukerjee BUC 1387 Filiform cuticular projections. Density of tuberculation mildly variable. Fig. 5.2 L. L. viscosa (Hornem.) Merr. SSW 7056a General lack of cuticular projections. Density of tuberculation mildly variable.

100 μm

100 μm

100 μm 100 μm 100 μm

100 μm

57

Fig. 5.2 M. L. ruellioides (Colsm.) Pennell TI 1484. Trichome-like appendages conspicuously large. Granular tuberculation evenly dense; i.e. ‘stucco – effect. Fig. 5.2 N. L. pusilla (Thunb.) Merr. TI 1478 Reticulate appendages. [This shape is reminiscent of some Pseudolysimachion seeds described by Martinez-Ortega & Rico (2001).] Tuberculation evenly dense. Fig. 5.2 O. L. parviflora (Roxb.) Haines SSW 8696 Compare with same taxon (Fig. G.). Trichome-like appendages rugose/ rounded, appearing reticulate at times, comparable but much reduced from 5.3 I. [The shape is reminiscent of some Veronica seeds (Martinez-Ortega & Rico 2001.] Tuberculation consists of evenly distributed bumps.

58

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. However, trichome-like appendages prove to be not genus-dependent.

100 μm

Fig. 5. 2 α. Attachment of appendage in L. ruellioides (Colsm.) Pennell. Magnification: 4500 x Appendage stretching over depression in seed coat. Seed coat coarsely granular/ strongly pitted throughout (like stucko).

Fig. 5. 2 β. Attachment of appendage in L. anagallis (Burm.f.) Pennell. Magnification: 4500 x Projections much smaller than in α. Seed coat smooth away from the appendages, with surface features more dense closer to the attachments.

59

5.1.1 SEM results and Yamazaki’s recent subgenera

Yamazaki included seed characters in his subgeneric classification of Lindernia (1981). As

to 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

Lindernia

Subgenera: TittmanniaLinderniaEuilysanthes

Bonnaya

L. ciliataL. ruellioidesL. antipoda

L. parvifloraL. procumbens

GENUS:

Section:Lindernia Tittmannia

L. pusillaL. viscosaL. hookeri

Angustifolia

L. anagallisL. micrantha

Nummularia

L. nummulariifolia

Torenioides

L.crustacea

Bonnaya

Figure 5.3: Classification of Lindernia based on Yamazaki (1981).

60

I (1): Seeds of L. procumbens were not examined due to lack of material. They were

described 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” differs

from the above in shape and size. This distinction between the above conforms to my

preliminary results.

II (1): His description of Lindernia pusilla did not mention the reticulate pattern observed

in our sample. However, under 50x microscope, this reticulation is not obvious. He

described the seeds as “shortly cylindric, ca. 0.3 x 0.2 mm, scrobiculate.” Following in this

same section is L. viscosa with exactly the same description. The last species belonging to

Yamazaki’s group is L. hookeri, but here only the variety L. hookeri var. cochinchinensis

Bonati is described as “globose or broadly ellipsoidal” (followed by the same dimensions

and testal pattern).

II (2): Yamazaki’s grouping of Lindernia anagallis with L. micrantha is sound, but for

seed 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 0.15

mm, 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 filiform

appendages. I am not aware of any studies showing whether these appendages are typical

of 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, a

grouping based on its similar fruits, but which cannot be upheld based on seed characters

alone.

Assuming that seed characteristics delimit groups within Lindernia, and further assuming

that the characters observed here are independent from each other (an assumption that may

well be rejected when tested), I propose the following key to groups:

61

1A Granular testal pattern with long hooked appendages.......................... L. ruellioides.

1B Tuberculate testal pattern with short, or no appendages............................................2

2A Alveolate seed coat....................................................................................................3

2B Non-alveolate seed coat.............................................................................................5

3A Alveolae with regularly dispersed appendages.........................................................4

3B 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................................6

6A 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 by

Yamazaki. A solid key to the species based on seed characters would require a large

sample of, e.g., all Linderniaceae s.l. I agree that Lindernia parviflora belongs to a distinct

group. Further, Yamazaki’s section Bonnaya corresponds in its capsule with section

Angustifolia, being distinct in their inflorescence types. Their shared alveolate seed

characteristic suggests that inflorescence type is a weak character, whereas capsule shape

may be a stronger indicator of species affinity. Yamazaki’s series Tittmannia included three

species that have little in common as far as the seed coat is concerned. In fact, the

subgeneric affinities of Tittmannia and Bonnaya shown in the SEM strongly support

Pennell’s choice to unite the two groups into Lindernia.

From this small study I interpret two major natural groups of alveolate, and non-alveolate

taxa, as has been found by Rahmanzadeh et al. (2005; he described a third group, that does

not occur here). These are further divided based on the nature of the appendages. It is

62

conceivable, however, that a larger study would reveal a strong connection between alveoli

and appendages. Lindernia ruellioides was found distinct based on its granular testal

pattern and its large, broad appendages. I further argue for the distinctness of Lindernia

parviflora based on its strongly angled, columnar shape also found for L. microcalyx Stehlé

by Rahmanzadeh et al. (2005). Lindernia pusilla is distinct because of its strongly

reticulate seed coat pattern; L. indet-a’s seed coat lacks projections, is mildly reticulate, but

appears smooth in those places where the seed has been exposed to pressure from

neighbouring seeds. Wile a large SEM study may prove to differ, the results from this small

study suggest that seed characters alone may not resolve the relationships of taxa within

Lindernia. It would, however, be useful to extend the study to include neighbouring taxa in

the Linderniaceae.

Regarding the role of morphology in phylogeny reconstruction, it has been argued that it is

better to include few rigorously studied anatomical characters rather than a large data

matrix with ambiguous characters (Scotland et al. 2003). SEM-based seed characters have

proved informative in other genera17

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 appendage

characters are useful in predicting affinities within this taxonomic group, and further, to test

whether any of the characters explored above have evolved in other clades also. I suggest

to include seed, capsule, and venation characters amongst others in a large morphological

data matrix. It would be very useful also to include phytochemical characters in a combined

morphological 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. This

pilot 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).

63

5.2 Taxonomic problems

This section discusses major and minor problems and open questions concerning the

completeness 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 genus

The state of identity of the type of the genus is problematic, because Allioni did not specify

an 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 Lindernia

pyxidaria L., using Allioni’s legitimate genus description indirectly referring to the type

specimen Allioni had in mind for the genus Lindernia. No other Lindernia had been

legitimately described since 1766. However, because Linnaeus cited in synonymy

Capraria gratioloides L., which again is thought to be based on Gratiola dubia L., a taxon

distinct from L. pyxidaria, the fact that he also cited von Lindern’s illustration of Allioni’s

type did not save him from having made an illegitimate description before the International

Code of Botanical Nomenclature, and therefore, in 1965, Philcox published a correct (Mc

Neill, pers. comm. Aug. 2005) lecto-typification [in Taxon (1): 30] when chosing

Krocker’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 new

type of the genus in their database, which was created much later than 1965. It seems

unfortunate that this modern lecto-typification, with all the material available today, was

not connected with an actual herbarium specimen, but in contrast, is based on a copper

etching on which not even lamina venation is discernable (recall Fig. 1 d). I am not against

using illustrations as types, but I believe that a single specimen seen and personally

annotated by Linnaeus could be found superior to Krocker’s illustration (recall Fig. 1 b).

64

The type for Linnaeus’ Capraria gratioloides will have to be determined. The scans

requested from the Linnean Society for this project included a specimen entitled

gratioloides (Fig. 5.3 a + b), but it is not certain whether this is the type specimen. Once

determined, 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 Lindernia

specimen 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 specimen

from 1763 should be found. Last, I suggest that natural variation between all specimens

claimed to represent Allioni’s first description (these are spanning the globe) should be

carefully studied, and compared with the type Capraria gratioloides L. I hypothesize that

the results will either reject synonymy between Vandellia erecta Benth. and Anagalloides

procumbens Krock., or include Capraria gratioloides L. in it.

Fig. 5. 4 a: Herb. Linn. 796.3 bears resemblance with L. procumbens (Krock.) Philcox. Type candidate for Capraria gratioloides L. ?

Fig. 5. 4 b: Same as a; cut, pasted & enlarged.

65

Maximovicz’s work would represent a good starting point to find European specimens. For

example, his reference to Döll’s Flora von Baden (1859) suggested two putative type

specimens for L. pyxidaria All.: Reichenbach 1698 and Schultz 70. This description cited

10 further localities with specimens from Germany [also revealing that the leaves are

blue—green, ovate-elongate, but obovate at the base of the plant; the seeds germinate even

after several years of drought, and describing in detail the shape and structure18

of the seeds

and their manner of attachment to the septum].

Krocker’s protologue19 indicates that he described a cleistogamous plant, whose flowers

are 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 of

the plant (= laying down); hünerdarm translated literally means chicken’s intestine. Further

clues to the identity of the plant are given: The peduncles are longer than the leaves, the

four anthers have no filaments, the style is subulate, the stigma is penicilliform. As to its

location, the next time I am near the river Oder at the border of Poland and Germany, I will

certainly be on the lookout to re-collect it (although it would not be surprising were it at the

brink of extinction by now20

).

5.2.1.1 Identity of the Asian form of the type species

The problem of typification we face is compounded by the difficulty of having a

cleistogamous European species synonymized with a chasmogamous Asian species. Unless

a lecto-type specimen can be chosen for our as of yet elusive Lindernia pyxidaria L. in

Asia, as may be possible for Allioni’s Lindernia (pyxidaria) through careful study of the

specimens 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, Wallich

3943; Figs. 5. 4 a – d) can (emphasis added) be synonymized with Krocker’s Anagalloides

18 Döll’s description of the seeds: numerous, very small, long-threadlike, usually with 5 longitudinal edges, 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.

66

procumbens or with Allioni’s Lindernia pyxidaria, the question of illegitimacy set aside for

a moment21

.

As discussed in section 2.1 of the Taxonomic History, and in section 5.2.1 of the results

above Lindernia procumbens (Krock.) Philcox was chosen as the new type species of the

genus Lindernia by Philcox (1965). During this study I was not able to develop a clear

understanding of this species because of the lack of material from Nepal. The relationship

between L. procumbens and L. parviflora (Roxb.) Haines is unclear. Theoretically, the

species are distinct on the basis of the number of anthers [4 fertile anthers in L. procumbens

and 2 fertile anthers in L. parviflora] and the lamina margin [entire in L. procumbens and

minutely 1—2-dentate in L. parviflora (Yamazaki 1981). Both of these have been reported

from 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 third

problem I have is whether the cleistogamous plants from Europe (i.e. L. procumbens) are

the same species as chasmogamous Nepalese plants (i.e. L. parviflora).

The differences between the inbreeding European and outcrossing Asian plants may be

due to the environment. Taking this thought a step further, would it be possible for a

species to have 4 fertile anthers when flowering early in the year, and 2 fertile anthers

later? To find out whether this might be possible, one would have to study the

pollination syndrome and foraging behaviours of the insects in the localities and

climates 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 intraspecific

adaptations to cleistogamy and chasmogamy. Recently, it has been reported by Lewis

(2000) that in typically chasmogamous flowers of L. procumbens and L. dubia, early

flowers are often cleistogamous, and set fruit by self-pollination. In greenhouse conditions

21 And to take the question of illegitimacy back up immediately, would not Bentham’s creation of Vandellia erecta, in which he cited a wrong synonymy (i.e., Tittmannia trichotoma = L. multiflora), be illegitimate?

67

on the other hand, e.g., L. grandiflora Nutt. will not produce capsules, and must

obligatorily outcross. While this evidence makes it more credible that L. procumbens

(Krock.) Philcox from Europe may be conspecific with Nepalese material, the geographic

variability and pollinator availability must be considered when searching for the factors

responsible for a variation in the number of fertile stamens.

According to ontogenetic studies (Walker-Larsen & Harder 2000), zygomorphy in the

Lamiales is often associated with partial loss of the stamen whorl. While generally in the

asterids this reduction occurred only seldomly, Lindernia represents one case where such a

reduction has led to the formation of a didynamous androecium. Walker-Larsen & Harder

claim that the persistence of staminodes after reduction of the stamen whorl occurs in the

asterids I clade only in some Solanaceae, and bilabiate-flowered Scrophulariales. Of the

antesepalous whorl, the three stamens opposite the lower calyx have moved to the top of

the upper lip, losing one stamen (probably in order to support the function of the two

remaining stamens). For the two stamens associated with the posterior two calyx lobes, the

insertion remains antesepalous, but the filaments are much extended to allow the anthers to

be connivent below the posterior lip. If these two anterior stamens have lost their primary

function of pollen production, it might be to prevent self-pollination. Studies on pollination

syndromes suggest that such transformations aid pollinator attraction, easily verified by a

look at L. antipoda’s corolla with bright yellow staminodes. It is interesting to note that the

partial loss of a stamen whorl is reversible. It is further interesting to note that Blume

reported Linderniae with two fertile stamens and two, or sometimes three sterile stamens in

Java.

It would be very interesting indeed to explore the relationship between L. parviflora with

two, and L. procumbens with 4 fertile anthers, and to see whether they are actually

conspecific, and have adaptated differently due to flowering times of the year in different

geographic areas, and/ or pollinator- availability in temperate zones during separate

seasons.

68

5.2.2 Vandellia erecta and Lindernia parviflora

In this section, first or early historic taxonomic treatments are compared to find evidence

for potential conspecificity of the material that has been treated as distinct since its first

description. 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 identities

of the above are dubious at best, respective treatments and protologues were compared.

It is now clear that Krocker’s Anagalloides procumbens is cleistogamous. Philcox’s

Lindernia procumbens, however, includes both cleistogamous and chasmogamous forms of

L. 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 also

included L. parviflora in synonymy of his [broad] species concept. In search of clues

whether it would be possible that one species has been treated as two distinct species for

185 years based on one character alone (although a strong one), it was thought interesting

to compare the descriptions of the two according to Bentham, and to see whether other

characters 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’s

Scrophularineae Indicae with Vandellia erecta (left column), and Bonnaya parviflora

(right column), and below moving on to the respective entries in De Candolle’s Prodromus

X [DC X] of 1846. Pertaining to V. erecta, Bentham omitted the synonymy of Tittmannia

trichotoma Benth. in his later treatment. He was then able to change “pedunculis axillaribus

racemosisve” to “pedunculis omnibus axillaribus.” No characters openly oppose each

other, and throughout the treatments (1835 & 1846), he points out the similarity between V.

erecta and Bonnaya / Ilysanthes parviflora, and his incapacity to distinguish them from

each other by any character other than the stamen number. Especially his later description

of I. parviflora points to the typical manner of inflorescence, which does tend to appear

subracemose toward the apex of the plant, with relatively shorter latitudinal branches.

69

In DC X, Bentham continues his treatment 11 years later:

“V. erecta suberecta, glabra, foliis subsessilibus ovatis oblongisve basi angustatis, pedunculis axillaribus racemosisve calyce 2-5 plo longioribus, calycibus ante anthesin 5-partitis vel profunde 5-fidis, laciniis lanceolatis membranaceis capsulas ovatas superantibus.—Tittmannia trichotoma, Benth. in Wall. Cat. n. 3943.—Tittmannia erecta, Benth. in Wall. Cat. n. 3947 (ubi cum Bonnaya parviflora miseitur) [Fig.].—Valde affinis V. crustaceae, sed satis distincta videtur. Hab. in Peninsula, Deyra Dhoon, Munghyr, Napalia, Sillet, Assufghur, Regno Birmannico, &c., Wallich, Royle, &c.”

FROM SCROPH. IND. OF 1835

“B. parviflora (Bentham in Wall. Cat. n 3867), humilis, ramosa, foliis oblongis lanceolatisve, floribus axillaribus subracemosisve, corollis calyce duplo longioribus.—Gratiola parviflora Roxb. Pl. Corom. 3. 3. t. 203” [see Fig. 2. a) above]. “Hab. in Hindustania et Sillet, Wallich, China, Nelson.”

FROM SCROPH. IND. OF 1835

“V. erecta; erecta, ramosissima vel subdiffusa, glabra, foliis infimis ovatis in petiolum brevem angustatis, floralibus sessilibus vel semiamplexicaulibus ovatis oblongis lanceolatisve, pedunculis omnibus axillaribus calyce pluries longioribus, calycis segmentis lineari-lanceolatis, capsulis ovato-globosis. [..] in India orientali frequens a Peninsula ad Nepal et ripas Irawaddy. Tittmannia erecta Benth.! in Wall. cat. n. 3947. Ilysanthi parviflorae ita similis ut vix nisi staminibus distingueres. Filamentorum anticorum appendicula longa linearis. (v.s.)”

FROM DC X OF 1846

“I. parviflora ; ramosissima, gracilis, foliis oblongis lanceolatisve, inferioribus in petiolum angustatis, superioribus sessilibus vel semiamplexicaulibus, floribus superioribus subracemosis, corolla calyce subduplo longiore, filamentis anticis leviter glandulosis infra apicem lobulo minutissimo glabro auctis. [...] in India orientali frequens a peninsula (Wight! herb. n. 2200) ad Napaliam (Wall.! h. DC.); Gratiola parviflora Roxb.! Pl. Corom. 3. p. 3. t. 204. Scroph. Ind. p. 34. A simillima I. capensi distinguitur habitu gracili, floribus minoribus, filamentorum anticorum ramo glabro, latitudine rami majoris subbreviore. (v. s.)

FROM DC X OF 1846

70

At species level, he writes:

Fig. 5. 4 c: Wallich 3947C; Bentham quoted Wallich 3947 as voucher for V. erecta Benth. in 1835 & 1846.

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 e: Entry of Tittmannia erecta Benth. in the Wallich Catalogue 3943

Fig. 5. 4 f: Wallich 3947B; another L. procumbens from Nepal.

Fig. 5. 4 g: Wallich s.n., Nepal; determined as Ilysanthes parviflora

71

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 it

were 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 that

Bentham may have had trouble to reach a conclusion about Ilysanthes parviflora and Vandellia

erecta. 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 of

British 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.”

72

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 integrifolia22

. 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. It

becomes clear that Hooker would rather move Vandellia erecta Benth. to the genus

Ilysanthes, [i.e. Ilysanthes erecta (Benth.) Hooker]. I agree with Bentham that the

staminal 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 be

viewed as a hypothesis still in need of testing. Corolla size in L. parviflora is known to

vary (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 in

22 A specimen determined Gratiola integrifolia Roxb. was recovered at E. It indeed resembles (and might be synonymous with) Vandellia erecta Benth. It is from India.

73

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 as

conspecific with L. hyssopioides (L.) Haines by Paul (1978) for Bihar and Orissa, a

taxon that appears very distinct from L. parviflora in Indian collections. As discussed

above, it has been impossible to find strong distinguishing characters between L.

procumbens and L. parviflora with the material available for this study. It is however

well-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 (van

der Vlugt 1985). The best solution to the question lies in collecting fresh material, and

specifically, to dissect, e.g., fifty, corollas from several populations to see whether

intraspecific variability occurs.

L. procumbens has been included in the key, but the material from which the key was

constructed 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 be

retrieved and dissected all proved to be a typical Ilysanthes sensu Rafinesque with two

fertile stamens. The desription included in the key was made from the Wallich specimen at

E, 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 Lindern

had its fruits more or less developed, but lacked flowers altogether.

5.2.3 The identity of L. indet-a

PSW 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 at

2500 m, the highest altitude recorded for collections of this genus in Nepal. This

Nepalese species was found to most probably be cleistogamous, since its corolla is

74

almost completely closed. It could either be a cleistogamous form23

of another species

of Lindernia, its affinities not having been identified, or it could be a separate species. I

have preliminarily referred to it here as Lindernia indet-a. While it is quite possible that

this species has been described already, and lack of time during this research is to be

blamed for it not having been properly identified, it is as feasable that we are dealing

with a new species. However, a similar individual was collected in Japan (Miyoshi

Furuse 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 individual

plants.

If monophyly of Lindernia in Asia was ever tested, it would be interesting to investigate

the question whether Lindernia indet-a shares a most recent common ancestry distinct

from the remaining Nepalese taxa. As to morphological characters, Lindernia indet-a

appears to have strong affinities with the L. dubia complex known from both America

and Europe. The phenotype of L. indet- a looks similar to, e.g., that of the Clayton

specimen 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 be

affected by migrating aquatic birds, and (3), known disjunctions between Asia and the

Eastern seaboard of the United States exist (Xiang et al. 1998), the origin of this species

can only be speculative. Furthermore, several specimens from Europe were found to

resemble, but not match Lindernia indet-a. The following species names bear

resemblance with L.indet-a: Gratiola inaequalis Walter – America; Lindernia dubia

(L.) Pennell sensu lato (1753)—based on Clayton specimen from Virginia; Gratiola

anagallidea Michaux (1803); Lindernia dilatata var. anagallidea H.L. Muehl.; L. dubia

var. 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. For

this 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.

75

which they reported a broad range of morphological variation within the complex, and

found the 4 varieties treated by Lewis (2000) should be treated as one species (Abstract

detail 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 be

ascertained: 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 (ranging

between 0.4 and 0.8). Further, measurements were taken to describe the manner of

inflorescence. The obviously stronger decrease in leaf length toward the apex of the

plant 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

0

5

10

15

20

25

1 2 3 4 5 6 7 8 9 10

Node number from Base to Apex

Leaf

leng

th [m

m]

LL av (p)LL av (a)

Figure 5.5: Depiction of average leaf length (measured from three flowering specimens of the same species, respectively) from the base of the plant (left) to the apex (right). LL av (p) denotes leaf length average of L. parviflora/ L. procumbens? (Krock.) Philcox; LL av (a) denotes leaf length average of Lindernia indet-a. Nodes are numbered from 1 (base) to 9 (apex).

76

Fig. 5.6: Lindernia indet-a – BM collection. The geographic locality is 29º 14’ N, 82 º 07’ E, altitude is ca. 2500 m.

77

5.2.4 Lindernia indet-b

The 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 the

leaf shape does not correspond to that known from other Nepalese collections. It is to be

found in the L. nummulariifolia folder in BM. A digital image, cropped and pasted, can be

viewed in Figure 5. 7 below. Unfortunately, the original handwriting on the bottom was cut

off 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.

78

5.2.5 The type of L. micrantha D.Don

The 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 with

this 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 Nepalese

material, and that the type may just be temporarily lost. The last time it was cited from the

BM as “Wallich s.n. in type folder” was by Press and Shresta in their Catalogue of Type

specimens from Nepal (2000b).

Fig. 5.8 a: Lectotype of V. angustifolia Benth.

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5.2.6 The type of Lindernia oppositifolia (Retz.) Mukherjee

Lindernia oppositifolia (Retz.) Mukerjee is cited by Hara in the Enumeration of the

Flowering Plants of Nepal [EFPN], but no specimen was recovered from Nepal during

this 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 in

the EFPN, or whether other specimens have been collected in Nepal and been deposited

elsewhere 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 within

scrub jungles” (Fig. 5. 8 c).

Fig. 5.8 b: Type of Gratiola oppositifolia Retz. (LD) Digital image of photograph.

Fig. 5.8 c: Lindernia oppositifolia Retz. K.M.Matthew P282 (1981); India.

80

6. SUMMARY

This 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 not

be underestimated, despite missing data. The revision demonstrated the importance of

alpha taxonomy and its rewards to science: The use of correctly typified nomenclature is

fundamental for plant science on all levels.

To understand Lindernia in its full geographical extent, a combined morphological and

molecular phylogenetic study would be very useful. The morphological characters

evaluated in this study should be coded rigorously. This analysis should include

morphometric coding of the corolla shape, a character for which the current availability of

herbarium material will not suffice. A research study should be designed to test the

variability of corolla and androecium particularly to resolve the questions pertaining to the

Lindernia parviflora/ L. procumbens complex. This would involve sampling of several

populations across the whole of several flowering seasons across national boundaries.

Further, the knowledge about phytochemical compounds in this group should be extended

through, e.g., further chromatography of flavonoids, to include these characters in the

phylogenetic analysis, because its application has proven useful in delimiting taxa (Diaz

Miranda 1977). While it would be favourable to include all geographic regions in the

phylogenetic study, a recollection of specimens known from Nepalese herbarium material

with the simultaneous production of detailled line-drawings may be treated as a priority to

directly benefit completion of the Lindernia section in the Flora of Nepal.

81

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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.

85

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87

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88

APPENDIX (1): ACCOUNT OF THE GENUS FOR THE FLORA OF NEPAL

Lindernia 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 inflorescence

intermediate. 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 unequally

5-lobed and bilabiate; lobes lanceolate, imbricate in bud,; calyx lobes 1—5-veined, margins

smooth 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 one

of the three anterior lobes. Androecium didynamous, anterior filaments longer, arching to

posterior side, with club-shaped geniculum, posterior filaments straight, without

geniculum; anthers connivent or free, either all fertile, or posterior fertile and anterior

sterile 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 long

as 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 of

anthers; each pair always consists of two, adding up to four anthers. If all four are fertile, I

write “all stamens fertile.” All here refers to the number “4.” If only the posterior pair is

fertile, I write “anterior stamens sterile” or “anterior staminodes,” and “posterior stamens

fertile,” herewith saving to repeat the superfluous number “2.” The species distribution

listed below may not be complete. Additional voucher specimens recovered in the future

should be added.

89

KEY TO THE SPECIES OF LINDERNIA

1A Calyx with lobes at least 1/3 of total calyx length or bilabiate .......................….2

1B Calyx cup-like with teeth up to 1/4 of total calyx length…….….…2. L. crustacea

2A Inflorescence a terminal raceme, sometimes very condensed and appearing

umbellate; upper bracts differently shaped than leaves below, marking an abrupt transition

to the flowering part of the plant..........................................................................................3

2B Flowers solitary, axillary; upper leaves subtending pedicels similarly shaped to

leaves below and, if smaller, decreasing in size gradually toward the apex of the plant

........................................................................................................….………...…..............8

3A Capsules ≥ 10 mm, cylindrical, much longer than the calyx.................................4

3B Capsules ≤ 5 mm, ovoid, about the length of the calyx...……......…....................6

4A Lamina serrations usually exceeding 14 per side, except on round leaves, where it

could be as few as 10 ...........................................................................................................5

4B Lamina serrations usually not more than 6--8 per side, except on very long, linear

leaves, where it could be as many as 10.............................................................1. L. antipoda

5A Leaves sessile or subsessile, marginal serrations extended into a long bristle; plant

not rooting at the nodes...........................................................…………...………5. L. ciliata

5B Leaves distinctly petiolate, marginal serrations not extended into a long bristle;

plant rooting at the nodes ................................................................................6. L. ruellioides

6A Lamina hairy above and at least on veins below; calyx hairy.............................7

6B Lamina and calyx glabrous...............................................................11. L. multiflora

7A Plant erect, not rooting at the nodes; length: width ratio of lamina > 2.2, bracts ≥ 3

mm, calyx > 4 mm, 2 stamens fertile, 2 sterile................................................13. L. hookeri

7B 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

90

[inflorescence solitary, axillary]

8A Stem densely hairy...............................................................................................9

8B Stem glabrous ....................................................................................................10

9A Lamina hairy, entire or with 3—4 serrations per side; leaf size markedly decreasing

toward apex of plant; capsule shorter than calyx...............................7.L. pusilla

9B Lamina glabrous, 6—10 distinct serrations per side; leaf-size not markedly

decreasing toward apex of plant; capsule longer than calyx................12. L. nummulariifolia

10A Lamina venation pinnate or uni-veined.............................................................11

10B Lamina three to five –veined from the base......................................................12

11A Lamina uni-veined, length: width ratio > 5, all 4 stamens fertile.....10.L. micrantha

11B Lamina venation pinnate, ratio <5, anterior stamens sterile................3. L. anagallis

12A Leaves subtending pedicels markedly decreasing in size toward the apex of the

plant, pedicels 1.5—2 times the length of the leaf .........................................................13

12B Leaves subtending pedicels not decreasing in size much toward the apex of the

plant, pedicels more or less the length of the leaves.......................................14. L. indet-a

13A Stamens 4, all fertile.......................................................................4. L. procumbens

13B Stamens 2 posterior fertile, 2 anterior sterile....................................9. L. parviflora

SPECIES DESCRIPTIONS

1. 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.: Habitat

in 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.

91

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 2

mm 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 than

leaves. 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 staminodes

geniculate, 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 X

1—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)

DNEP2 A55 (E)

DNEP2 B40 (E)

DNEP2 B44 (E)

Gardner, P.C. 156 (BM)

Gardner, P.C. 157 (BM)

Hara et al. 1486 (BM)

Kanai et al. 1456 (BM)

D.H.Nicolson 2824 (BM)

Nicolson, D.H. 3060 p. p. (mixed) (BM)

92

Norkett 8066 (BM)

Norkett 9165 (BM)

Noshiro et al. 9760068 (BM)

B.D. Pande 23 (BM)

PSW 3664 (BM)

PSW 5664 (E)

J.D.A. Stainton 1538 (E)

SSW 37 (BM)

SSW 2778 (BM)

SSW 5664 (E; BM)

SSW 5831 (BM)

SSW 7045 (BM)

SSW 7181 (BM)

Wallich 3857F (K-W)

Wallich 3858C (K-W)

K. Wesche s.n. (E)

Wraber 34396 (BM)

2. Lindernia crustacea (L.) F. Muell., Syst. Cens. Austral. Pl. 1: 97. 1882. Lectotype: LINN

785.3!

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.

93

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. Lamina

pinnately nerved, petiolate, ovate, ovate--oblong, or broadly ovate, to 15 X 9 mm, upper

surface glabrous, lower surface with sparse short hairs on protruding midrib, base

subtruncate or rounded, apex obtuse, margin sparsely ciliate. Inflorescence solitary and

axillary, 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 mm

in flower, to 2 mm in fruit. Corolla blue or purple, ca. 6 mm, pubescent inside. Anterior

filaments geniculate, stamens all fertile. Capsule ovoid, ca. 3 X 2 mm. Seeds ca. 0.4 X 0.25

mm, alveolate, filiform appendage.

Flowering Time: June—December.

Fruiting Time: June—October.

Habitats: On grassy or stony, dry or moist places, terraced banks, in sun

and 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 a

calyx so shortly lobed, it would better fit in with Torenia, was it not for its much smaller

corolla. According to “Checklist of medicinal plants in Southeast Asia”, used “against

boils, and itchiness. In Indonesia, cures scable: kudis.”

Chromosome number: 2n = 42 (Darlington & Wylie 1955)

Specimens seen:

Buch-Ham 1390 (Wall.Cat. 3961)

Norkett 8082 (BM)

Noshiro et al. 9760521 (BM)

Noshiro et al. 9760067 (BM)

PSW 5624 (BM)

SSW 4103 (E; BM)

SSW 6454 (E; BM)

SSW 7055 (E; BM)

94

SSW 7578 (BM)

Wallich 3945C (K-W)

Wallich 3962 (E)

Wesche No. 7/ 92 (BM)

L.H.J.Williams 326 (BM)

T. Wraber 36770 (BM)

T. Wraber 36785 (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 graminosis

humidusculis Zeylon[a]e’ König s.n. (C!)

Gratiola cordifolia Vahl, Enum.1: 97. 1804.—Holotype: ‘in graminosis humidusculis

Zeylon[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 Provinciarum

occidentalium 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 to

subpetiolate, ovate to broadly deltoid, 10--25 X 6--12 mm, glabrous base truncate to

attenuate, 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. Calyx

5—6.5 mm in flower and fruit, glabrous, calyx lobes 4—5 mm in flower and fruit, margin

and apex ciliate [or apex a white tip]. Corolla purplish pink and white, to 13 mm, central

95

anterior lip lobe to 3.5 X 6 mm wide, posterior lip 2 mm, throat internally, tube externally

hairy; ovary 2 mm, style ca. 5 mm. Anterior filaments ca. 3 mm, geniculate, posterior

subsessile in throat; anterior stamens sterile, subconnivent, anthers ca. 1 mm; posterior

stamens fertile. Capsule golden brown, elliptic to elongate, 7—10 X 1.5 mm . Seeds

alveolate, appendaged.

Flowering Time: March—late November.

Fruiting Time: April—November.

Habitats: Base of path, mixed sedge and herb vegetation. Moist terraces

near 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 exists

throughout the species.

Specimens seen:

DNEP 2B191 (E)

PSW 1833 (BM)

PSW 3648 (BM)

J.D.A.Stainton 65 (E; BM)

SSW 569 (E; BM)

SSW 2777 (E; BM)

SSW 5216 (E; BM)

SSW 7180 (BM)

Wallich 3949F (E; K-W)

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).

96

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 clasping

the stem, apex acute—acuminate, margin subentire to distantly serrate. Inflorescence

solitary, axillary, or subracemose; pedicels winged, 5—14 mm, often 1.5—2 times longer

than lamina; leaves subtending pedicels 3—10 mm long, 1—4 mm wide. Calyx to 3 mm in

fruit, glabrous; calyx lobes ca. 1.5 in fruit. Corolla not seen. Stamens all fertile. Capsule

striate, 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 did

not include any geographic information. It looks similar to L. parviflora, but has 4 fertile

anthers. Corolla not seen; it may fall off shortly after collection (I would advise to make a

drawing 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.

97

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. 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 10 cm, stem simple or diffusely branching, not rooting at nodes, to 1 mm

diameter, glabrous or shortly hairy on angles. Lamina sessile, elliptic to lanceolate, to 3—4

X 1 cm, glabrous to shortly hairy, base attenuate, apex acute—acuminate, margin with

usually 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 flower

and fruit, calyx lobes ca. 1.5—2 mm in flower, ca. 2.5—3 mm in fruit. Corolla white or

pink, 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 or

sandy 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 midrib

and the long-aristate margin.

Specimens seen:

SSW 7581 (BM)

EMAK 9 (E)

K. Wesche No. 7/91 (1995)(BM)

Norkett 6788 (BM)

T. Wraber 187 (BM)

K. N. Sharma E192 (BM)

O. Polunin 1928A (BM)

O. Polunin 1961B (BM)

P.C. Gardner 1550 (BM)

A.H. Norkett 8080 (BM)

PSW 5813 (BM)

SSW 6850 (BM)

98

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 et

India 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); Hara

et al. in Enum. Fl. Pl. Nep. 3: 117 (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 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, base

attenuate, 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 lobes

narrow, 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 externally

orange, 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 argillacious

earth, 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 it

lacks the long-aristate teeth on the lamina margin. Also, it roots profusely at the nodes; the

99

leaves on the non-flowering shoots are roundish, on the flowering shoots elongate, and

wider than in L. ciliata.

Specimens seen:

SSW 7054 (E)

SSW 7045a (BM)

W&S 8251 (BM)

Wraber 243 (BM)

SSW 9272 (E; BM) Wraber 79 (BM)

Wraber 79 (BM)

TI 1484 (BM)

SSW 6798 (BM)

Wesche 5/ 178 (BM)

Wallich 3856 D (K-W)

7. 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. in

Enum. Fl. Pl. Nep. 3: 117 (1982); Deyuan et. al. in Fl. Chin. 18: 33 (1998); Press et al. in

Ann. 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, calyx

lobes 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 middle

of lower lip, ca. 5—6 mm wide, central lower lip lobe ca. 3 mm. Stamens all fertile or

posterior fertile. Capsule green, ovoid—globose, ca. 2.5—3.5 X 3 mm. Style not persistent

in fruit. Seeds ca. 0.4 x 0.4 mm, shortly octagonally cylindric, reticulation protruding ca. 5

μm .

100

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 and

globose capsule with non-persistent style make this Lindernia easy to spot. We need more

collections.

Specimens seen:

TI 1478 (1963, BM)

Buch-Ham. 1381 (E)

Buch-Ham. 1395 (E)

Wall. Cat.3945F (K-W)

DNEP 2B—192 (E)

101

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. in

Enum. Fl. Pl. Nep. 3: 117 (1982); Deyuan et. al. in Fl. Chin. 18: 33 (1998); Press et al. in

Ann. 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. Leaves

sessile to 5 mm petiolate. Lamina pinnately veined, ovate to lanceolate, 15—40 X 12—22

mm, sparsely, coarsely hairy on both sides, increasingly so on veins below, hairs adpressed,

lamina margins dentate or undulate, shortly ciliate, longer on younger leaves. Inflorescence

racemose, bracts to 1.5 mm, pedicels shorter than leaves. Calyx hairy, to 3 mm in flower, to

4 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. Capsule

ovoid, almost as long as calyx. Seeds ca. 0.3 X 0.4 mm, ellipsoid—oblong, slightly

tapering 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].

102

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 al.

in Ann. Check. Fl. Pl. Nep. 294 (2000).

Annual, to 15 cm, stem winged, 0.5--1 mm diameter, not rooting at nodes, glabrous. Leaves

sessile, lamina 3—5-veined from base, lanceolate, 8—16 X 3—7 mm, glabrous; base

rounded 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 3

mm 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 stamens

sterile, 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. Seeds

cylindrical 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 occasionally

obscurely toothed lamina margin and its anterior pair of anthers being sterile. A third

character might be the callous style base that remains on the non-dehisced capsule of L.

procumbens, but not on L. parviflora.

103

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 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).

Annual to 20 cm, stem simple or branching, erect, not rooting at nodes, 0.5—1.3 mm

diameter, 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, apex

rounded, margin entire to obscurely incised, glabrous or ciliate. Inflorescence solitary,

axillary, leaves subtending pedicels same size as leaves below, smaller when subtending

pedicels of immature capsules; pedicels 4—15 mm in flower, 10—28 mm in fruit. Calyx

3.5—4.5 in flower, 3.5—5 mm in fruit, glabrous; calyx lobes 2.7—3.6 mm in flower, ca. 4

mm in fruit, margins and/ or apex ciliate. Corolla white—pink, 6—9 mm, lower lip ca. 8

mm wide, central anterior lip lobe ca. 3 mm wide; posterior lip ca. 5 mm wide; anterior 2

filaments ca. 5 mm, appendaged; posterior 2 filaments ca. 1 mm, stamens all fertile, anthers

0.8—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.

104

Distribution: C Nepal; [W. Himalaya; E. Asia; SE. Asia].

Notes: This species is easily recognized by its narrow, linear leaves; as to

this character, however, it is posible that we are dealing with L. oppositifolia, whose

existence in Nepal has been reported (but without voucher specimens). The inflorescence

of 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, glabrous

or 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

105

mm in fruit, glabrous, margins ciliate. Corolla purplish—white, 3.5—4 mm, central lobe of

lower lip ca. 0.6 mm. Stamens all fertile, ca. 0.3 mm. Capsule ovoid, ca. 3 x 2 mm, apex

acute. 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. The

plant resembles L. viscosa, but is glabrous. The slightly reflexed pedicels in fruit and the

relatively 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 at

nodes, hairy to 0.2 mm. Leaves sessile or subsessile, lamina pinnately 3—5-veined (but

seemingly palmate), ovate to suborbicular, 10—15 X 10—15 mm, glabrous and glossy

above, sparsely ciliate on veins below; base rounded, apex rounded or obtuse, margin to

10-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 usually

about equal the lamina length, glabrous, but shortly hairy toward the apex; bracts 0.7—1.3

mm. Calyx 2—3 mm in flower, 2.5—3.5 mm in fruit, glabrous; calyx lobe margins ciliate

on apex. Corolla pink and white, 5—7 mm, central lower lobe ca. 1--4 mm, upper lip ca. 3

106

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, posterior

anthers fertile. Capsule linear—cylindric, acuminate, 5—6 X 2 mm wide, 1.5—2 times

length 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 other

Lindernia.

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).

107

Annual to 15 cm. Stem erect, to 1.5 mm diameter, sparsely pubescent at angles, hairs to 1

mm long. Leaves shortly to 6 mm petiolate, lamina pinnately veined, ovate or lanceolate, to

25 X 10 mm, lamina base attenuate to cuneate, apex acuminate, margin serrate, with 8 to

10 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 terminal

cyme and/ or solitary, axillary. Pedicels approximately as long as leaves. Subtending bracts

approximate leaf size at base of inflorescence, much smaller, bract-like at apex. Calyx

bilabiate, 5-lobed, lobes subequal, divided to middle in flower, +/- to base in fruit and fully

covering 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. Capsule

light 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 its

fat capsules.

Specimens seen:

SSW 6767 (E; BM)

Banjeri, Shresta & Upadhyay 2528 (BM)

SSW 7056 (BM)

14. Lindernia indet-a

Annual to 15 cm, stem erect or occasionally procumbent and rooting at nodes, to 2 mm

diameter, glabrous. Lamina 3—5-veined, sessile, broadly elliptic to rhombic, to 25 X 12

mm, glabrous, base decurrent, apex obtuse, lamina margin subentire—undulate.

Inflorescence solitary, axillary, leaves subtending pedicels 14—22 X 5—12 mm; pedicels

to 10 mm, shorter than leaves. Calyx to 4 mm in fruit, glabrous; calyx lobes to 3 mm in

108

fruit, calyx lobe margin smooth. Corolla white, closed. Filaments absent, anterior

staminodes, posterior stamens fertile. Capsule ca. 3 x 2 mm, ovoid, striate, glabrous, only

base of style (to 1 mm) persitent on capsule. Seeds ca. 0.35 x 0.2 mm, not alveolate, not

appendaged, seed coat smooth.

Flowering Time: at least July

Fruiting Time: at least July

Habitats: 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 Williams

4476); the 10 plants collected varied little. A good character may be that during

inflorescence, one side develops a peduncle, while the opposing side develops a further

shoot that carries two leaves. Next node up, the reverse.

Specimens seen: PSW 4476 (E; BM).

APPENDIX (2)

Malesia Bhutan Thailand Indo-China Nepal

Indo-Burma

W. Himal China

Sri Lanka Java

L. anagallis (Burm.f.) Pennell 1 1 1 1 1 1 1 1 L. antipoda (L.) Alston 1 1 1 1 1 1 1 1 L. blancoi (Merr.) Philcox 1 L. brevipedunculata Migo 1 L. cambodgiana (Bonati) Philcox 1 L. celebica Philcox 1 L. cephalantha T. Yamaz. 1 1 L. cerastioides T. Yamaz. 1 L. ciliata (Colsm.) Pennell 1 1 1 1 1 1 1 L. crenata Pennell 1 L. crustacea (L.) F. Muell. 1 1 1 1 1 1 1 1 L. cyrtotricha P. C. Tsoong 1 L. delicatula P. C. Tsoong 1 L. dictyophora Tsoong 1 1 1 L. dubia (L.)Pennell 1 L. elata (Benth.) Wettst. 1 1 L. glabra Philcox 1 L. hookeri (C.B.Clarke) Wettst. 1 1 1 2 L. hyssopioides (L.) Haines 1 1 1 L. jiuhuanica X. H. Guo & X. L. Liu 1 L. junciformis (Bonati) T. Yamaz. 1 1 L. kiangsiensis P. C. Tsoong 1 L. kerrii T. Yamaz. 1 1 L. khaoyaiensis T. Yamaz. 1 L. laotica Bonati 1 L. latifolia ? 1 1 L. maxwellii T. Yamaz. 1 1 L. macrobotrys P. C. Tsoong 1 L. megaphylla P. C. Tsoong 1 L. micrantha D. Don 1 1 1 1 1 1 L. mollis (Benth.) Wettst. 1 1 1 1 L. molluginoides Wettst. 1

110

L. Montana ? 1 L. multiflora (Roxb.) Mukerjee 1 1 1 L. nummulariifolia (D.Don) Wettst. 1 1 1 1 1 1 L. oblonga (Benth.) Merr. & Chun 1 L. parviflora (Roxb.) Haines 1 1 1 L. philippinensis Philcox 1 L. pierreanoides T. Yamaz. 1 1 L. procumbens (Krock.) Philcox 1 1 1 (1?) 1 1 1 L. pusilla (Willd.) Bold. 1 1 1 1 1 1 L. rivularis Kerr ex Barnett 1 L. rotundifolia ? 1 L. ruellioides (Colsm.) Pennell 1 1 1 1 1 L. satakei T. Yamaz. 1 L. scapigera R. Br. 1 L. scutellariiformis T. Yamazaki 1 L. sessiliflora (Benth.) Wettst. 1 L. setulosa (Maxim.) Tuyama ex. Hara 1 L. srilankana Cramer & Philcox 1 L. stemodioides (Miq.) Kerr 1 L. stolonifera T. Yamaz. 1 L. stricta P. C. Tsoong & T. C. Ku 1 L. subconnivens Philcox 1 L. succosa (Kerr. Ex Barnett) Philcox 1 L. subulata R. Br. 1 L. taishanensis F.Z.Li 1 L. tenuifolia (Colsm.) Alston 1 1 1 L. thorelii Bonati 1 L. udawnensis T. Yamaz. 1 1 L. umbellata T. Yamaz. 1 L. urticifolia (Hance)Bonati 1 L. viatica (Kerr. Ex Barnett) Philcox L. viscosa (Hornem.) Bold. 1 1 1 1 1 1 1 L. yaoshanensis P. C. Tsoong 1 L. indet-a 1 L. indet-b (1?) Philcox Mill Yamazaki Yamazaki (Baden) Mukerjee Pennell Deyuan et al. Haines Backer

111

BENTHAM’S GENERA (1846)

Tribe VIII: Gratiolae

Lindernieae

Genus: Vandellia Lindernia Ilysanthes Bonnaya

B. brachiataB. reptansB. veronicaefolia

I. parvifloraL. pyxidaria

V. scabraV. hirsutaV. multiflora

Subtribe:

EllobumSection

V. angustifoliaV. pedunculata

NummulariaTittmannia

IlyogetonTorenioides

V. nummulariifolia

V. crustacea

V. erecta

APPENDIX (3)

112

Appendix (4) Krocker’s

protologue of Anagalloides procumbens

nostra

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___

___________________________ __________________________

114

115

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

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116