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

1. 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

2. 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.
ii

3. 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
iii

4. Sprengel 13
Buchanan-Hamilton 13
Don (D.) 14
Chamisso 14
Bentham 14
G. Don 16
Dalziel & Gibson 16
Hooker 16
Haines 17
Alston 17
Pennell 17
Borbás 18
Von Mueller 19
Urban 19
Von Wettstein 19
Merrill 20
Boldingh 20
Backer 21
3.4. Recent floristic accounts 21
Mukerjee 21
Sivarajan & Mathew 21
Banjeri 21
Philcox 21
Khan & Hassan 22
Diaz Miranda 22
iv

5. Cramer 22
Yamazaki 23
Quail Lewis 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
v

6. Testal pattern 46
5.1.1 SEM results and Yamazaki’s recent subgenera 59
5.2 Taxonomic Problems 63
5.2.1 Identity of the type species of the genus 63
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.
vi

7. 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
vii

8. 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
viii

9. 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.
1

10. 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 “Gratiolae 2,” 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.str 4. 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..
2

11. 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 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
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.
3

12. 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 1771 6. 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.) Philcox 7 was accepted as the type of the genus in all the accounts I
consulted during this revision.
6
It is possible, however, that Allioni made the combination in an as of yet unpublished work.
7
For L. procumbens (Krock) Philcox, the latter includes in synonymy, amongst others, Vandellia 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).
4

13. Fig. 1 a: Collections from Herb. Linn. 796.1; mixed
sheet of L. dubia L. on left; L. virginiana L. on the Fig. 1 b: Allioni’s illustration from Miscellanea
right. Wulfen reference in the illustration not found in Taurinensis 3, T5, F.1 (1766); open flower features
literature. 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 Fig. 1 d: Anagalloides procumbens
ovate, bivalved, 2-locular, valves elliptic. Krocker. 1790. Flora Silesiaca 2(1), t.
26.
5

14. Unfortunately, it has not been possible to verify the types of all the species listed in 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 Lindernia 8 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
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).
6

15. 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.
7

16. name G. cordifolia Colsm. even survived Pennell, who failed to synonymise it with his L.
anagallis (Burm.f.) Pennell. It has since been synonymized by Cramer (1981) in 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.
8

17. Figure 2 a: Roxburgh’s illustration of Gratiola Figure 2 b: Roxburgh’s illustration of G. parviflora Roxb.
parviflora Roxb. Roxburgh’s Icones 524, with detail; notice difference in
Pl. Corom. 3: 3, t. 203 (1819); with detail. venation compared with Fig. 2 a.
Maximovicz (1827-1891) treated several species in Decas 19 of Diagnoses 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
9

18. 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
10

19. 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,
11

20. 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
12

21. 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).
13

22. 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.
14

23. In Scrophulariae Indicae, Bentham used the genera Vandellia L., Bonnaya Link & Otto
(“filaments always abortive, simple, and club-shaped”) in the tribe Gratiolae to 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 erecta 12, Ilysanthes gratioloides, I.
capensis, and I. parviflora, all of which may indeed prove to form a tightly allied complex.
11
A cladogram of Bentham’s taxonomic concept as understood from DC X (1846) is attached in the
appendix 3.
12
See section 5. 2. 2
15

24. G. Don in his General System (1838) explained that the name Gratiola comes from gratia,
grace; on account of the supposed medicinal good qualities. His section on the Gratioleae
(Scrophulariaceae) included the genus Bonnaya, where he treated Bonnaya brachiata Link
& Otto, B. repens Spreng., B. veronicaefolia Spreng., and B. verbenaefolia Spreng. as 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
16

25. 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
17

26. 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.
18

27. Von Mueller, in Fragmenta Phytographiae Australiae (1882), listed Gratiola pedunculata
(Br. Pr. 435) 13 as dispersed throughout the whole of “Australiam extratropicam” but 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.
19

28. 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
20

29. 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
21

30. 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 characters 14 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.
22

31. 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 Lewis 15 (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.
23

32. Species descriptions and keys were exclusively based on Nepalese material where possible,
and missing material was substituted from India or Malaya. A total of 105 herbarium 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).
24

33. 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.
Fig. 4.1 a: Roots of Lindernia a. Fig. 4.1 b: Roots of L. procumbens
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
25

34. of structures (esp. lamina and calyx), and which usually do not exceed a length of 0.1 mm
[Fig. 4.2 b], but seldom reach 0.2 mm (e.g., on young leaves of L. nummulariifolia). 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.
ciliate margin ▲
Fig. 4. 2 a: L. micrantha (D.Don) Wettst. Fig. 4. 2 b: L.antipoda.(L.) Alston
Lamina margin distantly, obscurely serrate, and Lamina margin serrate and ciliate. Not to be
smooth. confused with c., L. ciliata, which is not ciliate.
Also visible here are the sunken stomata.
Fig. 4. 2 c: L. ciliata (Colsm.) Pennell Fig. 4. 2 d: L. parviflora (Roxb.) Mukerjee
Lamina margin long-aristate serrate. Lamina margin entire to obscurely 2—3 dentate,
and smooth.
26

35. Fig. 4. 2 e: L. hookeri (C.B.Clarke) Wettst.
Lamina margin serrate and ciliate; usually Fig. 4. 2 f: L. hookeri (C.B.Clarke) Wettst.
bearing a longer protrusion on the projecting part Lamina base is often densely beset with long
hairs, whereas the apex has none [see 4. 1 e]. This
of the margin.
phenomenon is shared by a few Nepalese species.
Calyx (inset) also strongly hairy.
ciliate margin ▲
Fig. 4. 2 g: L. anagallis (Burm.f.) Pennell
Lamina margin serrate, but ciliate often only on
the projecting part of the margin. A further Fig. 4. 2 h: L. ruellioides (Colsm.) Pennell
character exhibited by many species of Lindernia Lamina margin deeply serrate, and ciliate.
(see also b + f) is the red colour of the margin.
Stems. All stems are quadrangular, and are winged to some extent. The most 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
27

36. slender plants; L. anagallis and L. indet-a are more stout. Features such as hairs and
colour are most obvious on the angles of the stem.
Fig. 4. 3 a: L. pusilla (Thunb.) Bold.
Stem strongly winged. Also visible are the Fig. 4. 3 b: L. anagallis (Burm.f.) Pennell
flattened hairs attached to the angles. Stem winged., not hairy. Also visible is the
reddening characteristic of this taxon.
Fig. 4. 3 d: L. parviflora (Roxb.) Mukerjee
Stem winged, not hairy.
Fig. 4. 3 c: L. hookeri (C.B.Clarke) Wettst.
Stem winged, sparsely hairy on angles.
Leaves. The leaves in Lindernia are highly variable in shape (base, margin, apex) as well 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.
28

37. 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.
29

38. Fig. 4. 4 b: L. indet-a.
Fig. 4. 4 a: L. micrantha (D.Don) Wettst. Three to five-veined from the base. Lamina sub-
Lamina obscurely serrate. Leaf length to leaf entire to undulate.
width ratio 7:1. Petiole not easy to define.
Fig. 4. 4 c: L. crustacea (L.) F.Muell. Fig. 4. 4 d: L. parviflora (Roxb.) Mukerjee
Lamina 3-veined from the base. Sunken stomata
visible. Margin obscurely dentate near apex.
Fig. 4. 4 e: L. ciliata (Colsm.) Pennell Fig. 4. 4 f: L. ruellioides (Colsm.) Pennell
Lamina uni-veined
30

39. 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.
Fig. 4. 5 a: Lamina base and node of Fig. 4. 5 c: Lamina base and node
L. muliflora (Roxb.) Mukerjee of L. anagallis (Burm.f.) Pennell
Fig. 4. 5 b: L. anagallis (Burm.f.) Pennell
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
31

40. 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 Fig. 4. 6 b: Solitary, axillary inflorescence of L. pusilla
Pedicels slightly reflexed.
Fig. 4. 6 c: Solitary, axillary inflorescence of L. Fig. 4. 6 d: Solitary, axillary inflorescence of L.
anagallis parviflora
32

41. Calyx. The calyx consists of five lobes usually of slightly unequal length. 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. Fig. 4. 7 c: 5-partite calyx of
Fig. 4. 7 b: bilabiate calyx of L. L.antipoda
crustacea
hookeri
Fig. 4. 7 d: Glabrous calyx with Fig. 4. 7 e: Hairy calyx without
ciliate margin on apex of lobe (L. ciliate margin (L.pusilla)
anagallis)
33

42. Corolla. The corollae in Lindernia are bilaterally symmetric, semi-closed, and bilabiate.
The corolla of L. indet-a is closed, revealing cleistogamy. While accurate 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).
Fig. 4. 8 a: L. antipoda Fig. 4. 8 b: L. ruellioides
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
34

43. complex, (1) all 4 stamens bear fertile anthers (e.g., L. crustacea, L. pusilla, L. micrantha,
Fig. 4. 9 a, c, f ). In the next, (2) only the posterior pair bears fertile anthers, whereas 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.
35

44. Fig. 4. 8 c: L. anagallis (Burm.f.) Pennell Fig. 4. 8 d: L. parviflora (Roxb.) Mukerjee
DNEP 2B 191 DNEP 2B 125
Fig. 4. 8 e: L. antipoda (L.) Alston
Fig. 4. 8 f: L. nummulariifolia (D.Don) Wettst.
Fig. 4. 8 g: L. pusilla (Thunb.) Merrill Fig. 4. 8 h: L. micrantha D.Don
Herbarium Specimen EMAK soaked in OT2
36

45. Fig. 4. 9 a: Floral dissection of L. crustacea (L.) Fig. 4. 9 b: Floral dissection of L.
F. Muell.; WIL 326 nummulariifolia (D.Don) Wettst.; STA 4018.
Scale bar denotes millimeters.
Fig. 4. 9 d: Floral dissection of L. ruellioides
Fig. 4. 9 c: Floral dissection of L. pusilla (Willd.) (Colsm.) Pennell; SSW 9272. Anterior stamens
Bold.; as no flower for Nepal was available, a rudimentary.
specimen from area 6 was substituted.
ant. anther ►
post. anther ►
filament appendage ▲
Fig. 4. 9 f: Floral dissection of L. micrantha
Fig. 4. 9 e: Floral dissection of L. hookeri (D.Don) Wettst.; EMAK 174. Soaked in 6 (10 g
(C.B.Clarke) Wettst.; SSW 6767 aerosol OT2 / 100 g H2O) / 1 (acetone). Note
upper lip removed; anthers dorsifixed; appendage
club-shaped.
37

46. Gynoecium. The gynoecium is bi-carpellate, and the placentation axile. The style 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 Fig. 4. 10 b: Dehisced - and closed capsule of L. ciliata
(C.B.Clarke) Wettst. (Colsm.) Pennell
38

47. Fig. 4. 10 c: L. multiflora (Roxb.) Mukerjee Fig. 4. 10 d: L. nummulariifolia (D.Don) Wettst.
Fig. 4. 10 e: L. anagallis (Burm.f.) Pennell Fig. 4. 10 f: L. viscosa (Hornem.) Merrill
Fig. 4. 10 g: L. pusilla (Thunb.) Bold. Fig. 4. 10 h: L. crustacea (L.) F. Muell.
39

48. Fig. 4. 10 i: L. antipoda (L.) Alston Fig. 4. 10 j: L. parviflora (Roxb.) Mukerjee
Fig. 10 k: L. ciliata: Capsule with seeds
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
40

49. 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.
41

50. 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 /
42

51. 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
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52. 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
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