A taxonomic revision of Henckelia (Gesneriaceae) in South India with a new species, one new combination and seven lectotypifications

Abstract

The genus Henckelia Spreng. in South India is revisited based on field and herbarium data. Fifteen species are recognized. Henckelia wayanadensis is described new to science

Full text

48 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
with a new species, one new combination and seven
lectotypifications
Janeesha A.P.1 & S. Nampy2*
1Department of Botany, Providence Women’s College, Malaparamba P.O., Kozhikode, Kerala – 673 009, India.
2Department of Botany, University of Calicut, Calicut University P.O.,
Malappuram, Kerala – 673 635, India.
*E-mail: santhoshnampy2019@gmail.com
RHEEDEA
Journal of the Indian Association for Angiosperm Taxonomy
Vol. 30(1): 48–95 (2020)
ISSN: 0971-2313 (Print edition)
ISSN: 2582-2438 (Online edition)
https://dx.doi.org/10.22244/rheedea.2020.30.01.03
Abstract: The genus Henckelia Spreng. (Gesneriaceae)
in South India is revisited based on field and herbarium
data. Fifteen species are recognized. Henckelia
wayanadensis is described new to science while
Didymocarpus lyratus Wight var. protractus C.B.Clarke
and Henckelia sivagiriensis (Rajakumar, Selvak., S.Murug.
& Chellap.) E.S.S.Kumar are reduced to synonymy of
Henckelia lyrata (Wight) A.Weber & B.L.Burtt and
Henckelia wightii (C.B.Clarke) A.Weber & B.L.Burtt
respectively. Lectotypification of seven names are carried
out in this paper. Detailed descriptions, illustration/
photographic images and key for the identification of
South Indian species are provided.
Keywords: Conservation assessments, Endemic, H.
wayanadensis, Lectotypifications, New combination.
Introduction
The genus Henckelia Spreng. was first recognized
as Roettlera by Vahl in 1804. Since this name was
preoccupied with another genus, it was renamed
Henckelia by Sprengel (1817). Wallich (1819)
described Didymocarpus Wall. and transferred all
epithets under Henckelia to Didymocarpus. Later
taxonomists followed this treatment for almost 180
years. Weber and Burtt (1998[“1997”]) resurrected
the former name Henckelia from synonymy and
included about 180 species. In 2011, Weber et al.
(2011) remodelled Henckelia to include 56 species
formerly belonging to Henckelia sect. Henckelia,
Chirita Buch.-Ham. sect. Chirita (excluding species
Received: 10.12.2019; Revised & Accepted: 26.03.2020
Published Online: 11.05.2020
belonging to Damrongia Kerr ex Craib), the
monotypic genus Hemiboeopsis W.T.Wang and
excluded the species of Henckelia sect. Loxocarpus
(R.Br.) A.Weber & B.L.Burtt, sect. Didymanthus
(C.B.Clarke) A.Weber & B.L.Burtt, sect. Heteroboea
(Benth.) A.Weber & B.L.Burtt and sect. Glossadenia
A.Weber & B.L.Burtt. The genus under its current
definition is distributed in Sri Lanka, India,
Myanmar, Nepal, Bhutan, southern China, northern
Vietnam, northern Laos, and northern Thailand. A
few species were recently described bringing the total
number to c. 70 species worldwide (Manudev et al.,
2012; Middleton et al., 2013; Kumar, 2014; Janeesha
& Nampy, 2015; Ranasinghe et al., 2016; Krishna &
Lakshminarasimhan, 2018; Borah et al., 2019; Cai
et al., 2019; Sirimongkol et al., 2019; Yang et al.,
2019; Kanthraj et al., this issue). It is the eighth
largest genus in tribe Trichosporeae and tenth
largest in subfamily Didymocarpoideae (Möller &
Clark, 2013). Thirty five species and one variety
have so far been reported from India (Sinha &
Datta, 2016; Möller et al., 2017; Krishna &
Lakshminarasimhan, 2018; Borah et al., 2019;
Kanthraj et al., this issue). Lectotypification of some
species of this genus has been carried out by Vitek
et al. (2000), Janeesha and Nampy (2016) and
Ranasinghe et al. (2019). Henckelia is the largest
genus of Gesneriaceae in India with a high
proportion of endemism, particularly in South
India. However, since Clarke’s (1885[“1884”])
pioneering work in Hooker’s Flora of British India,
there has been no real revision of this genus in India.

49
Janeesha & Nampy
The purpose of the present study is to understand
the diversity, distribution and taxonomy of the
South Indian species of Henckelia, based on
extensive field work and consultation of specimens
and digital images deposited in major herbaria.
Fifteen species are recognized in South India while
two names are reduced to synonymy.
Materials and Methods
The materials for this study were obtained through
field trips in various parts of South India. Data sheets
were prepared for each population studied.
Herbarium voucher specimens were prepared and
deposited at Calicut University Herbarium (CALI).
The specimens consulted, including types, are held
at BLAT, BSD, BSI, CAL, CALI, DD, DEV, E,
FRC, JCB, KFRI, MH, RHT, SKU, TBGT and
online databases (https://plants.jstor.org; https://
www.ipni.org; https://www.tropicos.org; https://
www.biodiversitylibrary.org; https://www.
wcsp.science.kew.org; https://www.samlinger.
snm.ku.dk; https://www.linnean-online.org; http://
apps.kew.org/herbcat/navigator.do; https://
science.mnhn.fr; https://botanik.snm.ku.dk; https://
www.nhm.ac.uk). The acronyms of herbaria are
as per Thiers (continuously updated). Detailed
morphological descriptions were prepared after
studying fresh and herbarium specimens using the
terminology of Weber and Burtt (1998[“1997”])
and Simpson (2006). The genus description,
preceding those of the species from South India, is
applicable to the entire genus. Each species was
identified by consulting types, and protologues.
The nomenclature was updated with the latest
literature. Colour photographs of the dissected floral
parts were taken with a Leica EZ4HD stereo
microscope (Mannheim, Germany). The
distribution maps were prepared using the software
QGIS 3.12.1-Bucuresti (QGIS Development
Team, 2020) and conservation assessments were
done using IUCN Standards and Petitions
Subcommittee Criteria (2019). The information on
phenology, ecology, habitat and distribution of each
species was based on field studies and herbarium
specimens consulted. The chromosome numbers
and medicinal uses cited were gathered from the
literature and referenced in the text.
Taxonomic Treatment
Henckelia
Spreng., Anleit. Kenntn. Gew., ed. 2.
2(1): 402. 1817. Henckelia sect. Henckelia A.Weber
& B.L.Burtt, Beitr. Biol. Pflanzen 70: 334.
1998[“1997”]. Didymocarpus sect. Orthoboea Benth.
in Bentham & Hooker, Gen. Pl. 2(2): 1022. 1876.
Type: Henckelia incana (Vahl) Spreng.
Chirita Buch.-Ham. ex D.Don, Edinburgh Philos.
J. 7: 83. 1822. Chirita sect. Euchirita C.B.Clarke in
A.DC. & C.DC., Monogr. Phan. 5(1): 11. 1883,
nom. illeg. Roettlera sect. Euchirita (C.B.Clarke)
Fritsch in Engler & Prantl, Nat. Pflanzenfam. IV/
3b: 148. 1895. Didymocarpus sect. Euchirita
(C.B.Clarke) Chun, Sunyatsenia 6: 294. 1946.
Lectotype (designated by Burtt, 1954): Chirita
urticifolia Buch.-Ham. ex D.Don (=Henckelia
urticifolia (Buch.-Ham. ex D.Don.) A.Dietr.).
Perennial or annual herbs or subshrubs. Stem
reduced (rosette-forming) or erect, ascending,
rarely creeping. Leaves opposite, whorled or
alternate by reduction of one leaf at a node in
creeping species, sometimes one or two (rarely
three) in a plant, short to long petiolate; laminae
variable. Cymes axillary (or sometimes appearing
terminal by reduction of the vegetative shoot);
flowers (1)-10–38; bracts paired, ovate to linear,
narrowly ovate or narrowly triangular, sometimes
early caducous. Calyx of 5 sepals, free or fused into
a tube for part of the length; lobes linear, lanceolate,
ovate or triangular to narrowly triangular. Corolla
zygomorphic, variously shaped, infundibuliform to
campanulate, sometimes ventricose, 2-lipped, with
upper lip of 2-lobes and lower lip of 3-lobes,
colouration variable, often with yellow markings
in throat. Stamens 2, usually included; filaments
long, slightly bent (in South Indian species) to
geniculate; anthers cohering, glabrous or pubescent;
staminodes slightly bent, rarely straight or curved,
with or without a prominent antherode, glabrous
or pubescent. Disk cylindrical or cup shaped (absent

50 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
in South Indian species). Ovary conical or ovoid,
straight in relation to pedicel or with an angle
between the pedicel and the ovary (usually between
120–135° in South Indian Henckelia); stigma
chiritoid or obliquely subcapitate (in South Indian
species, except H. pradeepiana and H. repens).
Capsules linear, rarely globose, stipitate or sessile,
dehiscing loculicidally, splitting into 2 valves or
opening only along the dorsal side (mostly in
plagiocarpic ones), stipitate or sessile, orthocarpic
or plagiocarpic (plagiocarpic in South Indian
species). Seeds minute, numerous, ellipsoid.
Distribution: Sri Lanka, southern and northeastern
India, Nepal, Myanmar, Bhutan, southern China,
northern Vietnam, northern Laos and northern
Thailand (Weber et al., 2011; Sirimongkol et al.,
2019).
Key to the species of
Henckelia
in South
India
1. Plants with creeping stems or tuberous herbs
with 1–3 leaves ............................................... 2
1. Rosette-forming herbs with many leaves in
basal rosette ..................................................... 3
2. Tuberous herbs; leaves single (rarely up to 3);
laminae broadly ovate or elliptic; capsules ovoid
to sub-globose ............................. H. pradeepiana
2. Plants with creeping stems; leaves appearing
alternate by reduction of one leaf in a pair at a
node; laminae broadly ovate to sub-orbicular;
capsules linear ....................................... H. repens
3. Leaves distinctly petiolate; petioles not winged
.......................................................................... 4
3. Leaves sessile to ssub-sessile; if petioled, then
winged ............................................................ 5
4. Peduncles scaly towards apex; filaments
glabrous ........................................ H. missionis
4. Peduncles not as above; filaments puberulent
towards apex ................................ H. ovalifolia
5. Laminae obovate-spatulate or widely spatulate
to elliptic .........................................................6
5. Laminae not as above, but of variable shapes
......................................................................... 7
6. Peduncles 30–54 cm long, golden brown hairy
..................................................... H. macrostachya
6. Peduncles 10–18 cm long, glandular hairy
.............................................................. H. wightii
7. Leaf margins doubly crenate or doubly serrate;
ultimate segments irregular ........................... 8
7. Leaf margins crenate (occasionally doubly
crenate in H. meeboldii); ultimate segments
regular .......................................................... 11
8. Petioles lacerate; leaf bases lyrate; bracts pilose
................................................................ H. lyrata
8. Petioles not lacerate; leaf bases attenuate; bracts
not as pilose ................................................... 9
9. Leaves usually in whorls of three; calyx lobes
hispid on both sides .................. H. innominata
9. Leaves usually in whorls of four; calyx lobes
tomentose outside, glabrous inside ............ 10
10. Laminae ovate-widely ovate; bracts lanceolate
.............................................................. H. fischeri
10. Laminae narrowly elliptic to widely elliptic;
bracts ovate ............................ H. wayanadensis
11. Leaves usually in whorls of three ............... 12
11. Leaves usually in whorls of two, decussate
(occasionally whorls of three in H.
humboldtiana)................................................ 13
12. Capsules 13–15 mm long, tomentose ..............
...................................................... H. gambleana
12. Capsules 21–28 mm long, hispid ........ H. incana
13. Bracts 10–21 mm long, elliptic to obovate;
anthers bearded at the junction of the anther
lobes and filament ............................ H. bracteata
13. Bracts 3–5.1 mm long, linear; anthers glabrous
......................................................................... 14
14. Calyx lobes linear; corolla villous outside; ovary
8–9 mm long ........................ H. humboldtiana

51
Janeesha & Nampy
14. Calyx lobes ovate to lanceolate; corolla
glandular hairy outside; ovary 4–5.5 mm long
........................................................... H. meeboldii
Henckelia bracteata Janeesha & Nampy,
Willdenowia 45(1): 53. 2014. Type: INDIA, Kerala,
Kottayam district, Kurisumala, near to 10th Cross,
1100 m, 07.08.2014, A.P. Janeesha & Santhosh
Nampy 134270 (holo CALI!; iso CAL!, MH!) Fig. 1
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 8–14,
basal, decussate; petioles 2–9 cm long, pubescent,
winged; wings 2–3 mm broad; laminae elliptic-
ovate, 5.5–12.5 × 3–9 cm, acute or obtuse to
rounded at apex, narrowly cuneate to attenuate at
base, tomentulose on both surfaces, but more so
on the younger leaves and on lower veins,
becoming less so with age, margins shallowly
crenate, ultimate segments regular, surfaces usually
rugose; veins usually 5 on each side, much branched
and conspicuous beneath. Cymes axillary, dense,
1–11 per plant, 3–5(–6) times divided, 14–38-
flowered, pubescent; peduncles terete, 14–32 cm
long, reddish brown, pubescent; bracts two at each
branching, elliptic to obovate, 10–21 × 5–8 mm,
obtuse to rounded at apex, margins crenate,
tomentose; pedicels terete, 2–10 mm long,
pubescent. Flowers 10–20 × 4–9 mm. Calyx 5- or
rarely 6-partite, very deeply divided or basally
connate; lobes ovate, unequal, posterior lobe
smaller, 2.9–3.1 × 1.2–1.6 mm, others 3.3–4 × 1.6–
2 mm, slightly acute at apex, margins entire, reddish
brown, white tomentose. Corolla zygomorphic,
ventricose, two lipped, villous outside, glabrous
inside; tube 7–9 × 5.5–7.5 mm, lilac to pale lilac
with a yellow patch in the throat; lobes ovate to
rounded at apex, lilac to pale lilac, upper lip of two
lobes, 5.2–5.9 × 7.6–8.1 mm, lower lip of three
lobes, 5.6–7 × 7.4–8.3 mm. Stamens 2, adnate to
the base of corolla, included; filaments 4–6 mm
long, glabrous or sparsely pubescent towards the
region of the connective, green on the top and
colourless towards base; anthers reniform, 1.5–1.8
× 0.8–0.9 mm, milky white, bearded at the junction
of the anther lobes and filament. Staminodes 3
(sometimes 2); filaments 2.5–3 mm long, hyaline,
glabrous; antherodes not seen. Ovary ovoid to
oblong, 5–7 × 1.25–1.5 mm, green, pubescent; style
terete, 2–2.5 mm long, green, glabrous; stigma
obliquely subcapitate, 0.8–1.1 × 0.5–0.82 mm,
papillate, greenish white. Capsules cylindrical,
straight or randomly curved, 18–30 × 1.6–2 mm,
plagiocarpic, dehiscing loculicidally along the
dorsal side, reddish brown, tomentulose with
persistent stigma. Seeds numerous, elliptic, 0.4–0.6
× 0.2–0.25 mm, slightly acute at apex, truncate at
base; testa dark brown to black, reticulate.
Flowering & fruiting: Flowering fom July to
September and fruiting from August to October.
Habitat: In wet humus in granite rock crevices, at
altitudes 1100 m above sea level, along with
Mitracarpus hirtus DC. (Rubiaceae), Porella sp.
(Porellaceae) and species of moss, Selaginella sp.
(Selaginellaceae) and grasses (Poaceae).
Distribution: Endemic to the southern Western
Ghats, hitherto known only from the type locality
and adjoining areas (Fig. 6).
Specimens examined: INDIA, Kerala, Kottayam
district, Kurisumala, 17.08.2013, K.M. Manudev &
A.P. Janeesha 134206, 134209; on the way to 6th
cross, 07.08.2014, A.P. Janeesha & A.P. Roshan
134268; near to 6th cross, 07.08.2014, A.P. Janeesha
& A.P. Roshan 134269; near 7th cross, 26.06.2016,
P.S. Sreeja & A.P. Janeesha 137659; near 10th cross,
26.06.2016, P.S. Sreeja & A.P. Janeesha 137660; on
the way to Elaveezhapoonchira, 28.06.2016, A.P.
Janeesha & Santhosh Nampy 137665 (CALI).
Conservation status: The species can be found at two
localities in Kurisumala and Elaveezhapoonchira
about 20 km apart, with an estimated Extent of
Occurrence (EOO) of 34 km². Eighty two mature
plants were observed across these two localities and
the plants at Kurisumala and the series of crosses
represent a continuous population. A continuing
decline in population size has been observed over
three years because the localities are tourist
destinations. Therefore, this species is assessed here
as Endangered (EN), B1ab(ii,iii,iv,v), B2ab

52 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Fig. 1.
Henckelia bracteata
Janeesha & Nampy: a. Habit and habitat; b. A portion of the inflorescence; c. Flower; d. Calyx; e.e.
e.e.
e. Calyx with pistil;
f. Bract; g. Stamens; h. Staminode; i. Pistil; j. Capsule; k. Seeds (from
A.P. Janeesha & Santhosh Nampy
134270).

53
Janeesha & Nampy
(ii,iii,iv,v), D, according to IUCN’s guidelines
(IUCN, 2019).
Notes: This species is morphologically close to H.
humboldtiana in the shape of leaves, presence of
bracts and much longer scapes than the leaves
(Janeesha & Nampy, 2015; Janeesha, 2017).
However, it is distinguishable by its thick laminae
with narrowly cuneate to attenuate base, larger,
elliptic to obovate bracts, ovate calyx lobes that do
not reach the full length of the ovary and hairs at
the junction of the anther lobes and filaments.
Henckelia bracteata plants are robust and dull green
throughout while the peduncles, pedicels, calyx and
capsules are usually reddish brown. Although the
margins of the laminae are crenate, the crenation is
shallow when compared to H. humboldtiana. When
grown in the greenhouse at low altitudes, the colour
of the corolla changes from pale lilac to dark lilac
to violet.
Henckelia fischeri (Gamble) A.Weber &
B.L.Burtt, Beitr. Biol. Pflanzen 70: 344.
1998[“1997”]; T.S.Nayar et al., Fl. Pl. Western
Ghats 1: 527. 2014. Didymocarpus fischeri Gamble,
Bull. Misc. Inform. Kew 1923(93): 117. 1923, Fl.
Madras 2: 988. 1924; A.N.Henry et al., Fl. Tamil
Nadu Ind., Ser I: Analysis 2: 132. 1987;
M.Ahmedullah & M.P.Nayar, Endemic Pl. Indian
Region 1: 142. 1987; M.Mohanan & A.N.Henry,
Fl. Thiruvananthapuram 337. 1994; Sasidh. &
Sivar., Fl. Pl. Thrissur Forest 329. 1996; Sasidh.,
Biodivers. Doc. Kerala 6: 331. 2004; T.S.Nayar et
al., Fl. Pl. Kerala 342. 2006. Lectotype (first-step
designated by Vitek et al., 2000; second-step
designated here): INDIA, Tamil Nadu,
Coimbatore district, Anamalai hills, 500–600 m,
02.08.1915, C.E.C. Fischer s.n. (K [K000450837
digital image!]; isolecto K [K00122291 digital
image!]). Figs. 2 & 3
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 12–20,
basal, in whorls of four; petioles 0.7–5 cm long
(sometimes up to 7.5 cm), pubescent, winged;
wings 4–12 mm broad; laminae ovate-widely ovate,
8–23 × 6–12 cm, obtuse to slightly acute at apex,
attenuate at base, tomentose on both surfaces, with
small white patches on the upper surfaces, golden
brown on younger leaves and on lower veins,
becoming less so with age, margins doubly crenate,
ultimate segments irregular, surfaces usually rugose;
veins usually 6–8 on each side, much branched and
conspicuous beneath. Cymes axillary, 3–7 per plant,
4–7 times divided, 16–37-flowered; peduncles
terete, 17–38 cm long, green-pale reddish brown,
glandular hairy; bracts two at each branching point,
lanceolate, 2–4 × c. 1 mm, acute at apex, margins
entire, tomentose, with a tuft of hairs at the apex,
green to purple; pedicels terete, 6–24 mm long,
glandular hairy. Flowers 11–17 × 9–22 mm. Calyx
5-partite, very deeply divided or basally connate;
lobes linear to broadly lanceolate-ovate, unequal,
posterior lobe smaller, 2.5–3.8 × 0.8–1.6, others 3.6–
6 × 1.2–1.9 mm, slightly acute or obtuse to rounded
at apex, margins entire, reddish brown hairy.
Corolla zygomorphic, ventricose, two lipped,
glandular hairy outside, glabrous inside; tube 6.8–
12 × 5.7–7.9 mm, dark to pale lilac with a yellow
blotch at the throat; lobes rounded or very widely
ovate with undulate margins, dark to pale lilac,
upper lip of two lobes, 6.6–7.3 × 7.7–10 mm, lower
lip of three lobes, 6.9–7.8 × 8.6–10.9 mm. Stamens
2, adnate to the base of corolla, included; filaments
4–5.3 mm long, glabrous, yellow; anthers reniform,
1.5–2.75 × 0.75–1.16 mm, milky white to cream.
Staminodes 2 (the lateral ones), sometimes 3;
filaments 2.4–4 mm long, hyaline, glabrous;
antherodes sometimes present. Ovary ovoid to
oblong, 3.7–6 × 1–1.27 mm, green, glandular hairy;
style terete, 1–4.5 mm long, green, glabrous
towards the apex, basally glandular hairy; stigma
obliquely subcapitate, 0.65–0.87 × 0.9–1.1 mm,
slightly papillate, pale green. Capsules cylindrical,
straight or randomly curved, 14–37 × 1.8–2.4 mm,
plagiocarpic, dehiscing loculicidally along the
dorsal side, reddish brown, glandular hairy with
persistent stigma and calyx. Seeds numerous,
elliptic, 0.57–0.69 × 0.22–0.29 mm, slightly acute
at apex, truncate at base; testa dark brown, reticulate.
Flowering & fruiting: Flowering from May to
October and fruiting from July to December.

54 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Fig. 2.
Henckelia fischeri
(Gamble) A.Weber & B.L.Burtt: a. Habit and habitat; b. Inflorescence; c. Flower; d. Calyx; e. Calyx with pistil;
f. Stamens; g. Staminode; h. Pistil; i. Seeds; j. Immature capsule (a & b from
A.P. Janeesha, Habeeb Rahman & A. Kabeer
134216; c-j from
A.P. Janeesha & Santhosh Nampy
134224).

55
Janeesha & Nampy
Habitat: On moist slopes and in rock crevices,
usually at an altitude above 850 m, in association
with Adiantum philippense L., A. concinnum Willd.,
Hemionitis arifolia (Burm.) T.Moore (all
Pteridaceae), Spermacoce exilis (L.O.Williams)
C.D.Adams, Mitracarpus hirtus DC., Oldenlandia
umbellata L. (all Rubiaceae), Leucas aspera Link.
(Lamiaceae) and a few grass species.
Distribution: Endemic to southern Western Ghats
(Fig. 6).
Specimens examined: INDIA, Kerala, Idukki district,
Kottamala, 16.05.1995, Jomy Augustine 15309
(CALI); near Idukki dam, 24.08.1981, V.S. Raju
71173 (MH); Santhanpara, 2016, P. Soumya & A.P.
Janeesha 137678 (CALI); Palakkad district,
Karimala-grassland, 26.09.2013, A.P. Janeesha &
Santhosh Nampy 134227 (CALI); Nelliyampathy,
29.06.1994, A.E. Shanavas Khan & E.S. Santhosh
Kumar 20100 (TBGT); Ibid., s.d., V.P.K. Nambiar
1067 (KFRI digital image); Parambikulam-way to
Thunakkadavu dam view point, 24.09.2013, A.P.
Janeesha & Santhosh Nampy 134222; Seetharkund-
near the water fall, 17.09.2013, A.P. Janeesha, Habeeb
Rahman & A. Kabeer 134216; Vengoli hill top,
25.09.2013, A.P. Janeesha & Santhosh Nampy 134224
(CALI); Pathanamthitta district, Achankovil, s.d.,
A.G. Pandurangan 14432 (TBGT). Tamil Nadu,
Coimbatore district, Anamalai, 1857, s.coll. s.n.
(MH); Nilgiri district, Kodanad, 12.11.1970, E.
Vajravelu 36854 (MH); Tenkasi district, Courtallam,
s.d., K. Narayana Iyer 01874 (TBGT); Theni district,
Meghamalai, 25.08.1985, K. Ravikumar 2380 (MH);
Tirunelveli district, Kularatti estate, 21.12.1980, E.
Vajravelu 76416 (MH).
Conservation status: The species is widely distributed
in Kerala with a few localities in Tamil Nadu with
around twelve localities. The Extent of Occurrence
(EOO) is estimated to be c. 5600 km². The Area of
Occupancy (AOO) is likely less than 50 km², since
the plants occur in localized populations. For
instance, in Kerala five locations were surveyed and
a total of only 136 mature plants were found. The
locations in Kerala and Tamil Nadu are important
tourist destinations. Therefore, there is the potential
of disturbance due to tourist activities with
detrimental outcomes for the number of plants and
localities. Even though the species is widely
distributed, because of the likely small AOO the
species could be categorized at present as
Endangered (EN), B2ab(ii,iii,v), according to
IUCN (2019) criteria.
Notes: This is a fairly large plant with deeply crenate
leaves and many-flowered, long scapes. The flowers
are deeply pouched and horizontally oriented. Their
scapes are pale greenish-red under natural
conditions, but bright green when grown in the
greenhouse.
Typification: This species was described by Gamble
(1924) based on specimens collected by C.E.C.
Fischer from Anamalai hills. Vitek et al. (2000) cited
“South India, Anamalai hills, Coimbatore district,
500–600 m alt., 02.08.1915, C.E.C. Fischer (K)” as
the type. While searching for the type specimens,
Fig. 3. Lectotype of
Henckelia fischeri
(Gamble) A.Weber & B.L.Burtt.
© The Board of Trustees for The Royal Botanic Gardens, Kew.
Reproduced with permission.

56 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
two relevant sheets were found in K (K000450837,
K00122291). Both sheets were annotated,
“Didymocarpus Rottlerianus, Wall., Anamalai hills,
South India, 1700 ft., C. Fischer, 2.8.1915”. The
sheet K000450837 has a flowering plant, and two
fruiting scapes separately mounted on it, but
contains a note by Gamble “a letter from Mr.
Fischer dated 27.11.22 says that three specimens
were collected at a place called Pachchathani on
the lowest slopes of the anamalais ...... towards
mount Stuart, 13.1.23” while the other sheet has
two flowering plants. According to Art. 9.17 of
ICN (Turland et al., 2018), we take a second step
lectotypification by narrowing them to a single
specimen. We chose K000450837 that agrees with
the protologue and has the inscription by Gamble,
while the other specimen (K00122291) is designated
as isolectotype.
Henckelia gambleana (C.E.C.Fisch.) A.Weber &
B.L.Burtt, Beitr. Biol. Pflanzen 70: 345.
1998[“1997”]; T.S.Nayar et al., Fl. Pl. Western
Ghats 1: 528. 2014. Didymocarpus gambleanus
C.E.C.Fisch., Bull. Misc. Inform. Kew 1938: 36.
1938 (as ‘gambleana’); A.N.Henry et al., Fl. Tamil
Nadu Ind., Ser I: Analysis 2: 132. 1987;
M.Ahmedullah & M.P.Nayar, Endemic Pl. Indian
Region 1: 143. 1987. Didymocarpus rottlerianus Wall.
ex R.Br. var. lanuginosus C.B.Clarke in A.DC. &
C.DC., Monogr. Phan. 5 (1): 102. 1883.
Didymocarpus tomentosus Wight var. lanuginosus
(C.B.Clarke) C.B.Clarke in Hook.f., Fl. Brit. India
4: 353. (1885[“1884”]). Didymocarpus lanuginosus
(C.B.Clarke) Wight ex Gamble, Fl. Madras 988.
1924, nom illeg., non Wall. ex R.Br. 1839, non Wall.
ex A.DC. 1845. Lectotype (first-step designated by
Vitek et al., 2000; second-step designated here):
s.loc., s.d., Wight 561 (K [K000858182 digital
image!]; isolecto E [E00627530!], K [K000858183
digital image!]). Figs. 4 & 5
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 6–12,
basal, usually in whorl of three; petioles 0.7–1.8 cm
long, younger leaves subsessile, pubescent, winged;
wings 5–8 mm broad; laminae widely ovate, 4–7.7
× 3.2–6 cm, acute to obtuse at apex, narrowly
cuneate to attenuate at base, hispid on the upper
surface, golden brown hairy on lower surface, but
more so on younger leaves and lower veins,
becoming fewer with age, margins crenate,
ultimate segments regular, surfaces usually rugose;
veins usually 4 on each side, much branched and
conspicuous beneath. Cymes axillary, 2–4 per plant
(rarely up to 11), 3–5 times divided, 12–36-
flowered; peduncles terete, 8–19 cm long, green
to reddish brown, golden brown hairy; bracts two
at each branching point, lanceolate to triangular,
3–3.5 × 0.36–0.4 mm, obtuse to slightly acute at
apex, margins entire, white tomentose; pedicels
terete, 4–14 mm long, pubescent. Flowers 8–11 ×
5–6.5 mm. Calyx 5-partite, very deeply divided or
basally connate; lobes lanceolate to narrowly
oblong, unequal, posterior lobe smaller, 2.7–3 ×
0.42–0.53 mm, others 2.9–3.2 × 0.5–0.68 mm,
obtuse at apex, margins entire, reddish brown,
white hairy. Corolla zygomorphic, ventricose, two
lipped, villous outside, glabrous inside; tube 7–8 ×
5.8–6.4 mm, lilac to pale lilac with a yellow patch
in the throat; lobes widely ovate, rounded to
truncate at apex, lilac to pale lilac, upper lip of two
lobes, 3.8–4.3 × 3.5–4.1 mm, lower lip of three
lobes, 4.2–5.1 × 4–5 mm. Stamens 2, adnate to the
base of corolla, included; filaments 2.2–2.7 mm
long, glabrous, green on the top and colourless
towards base; anthers reniform, 1.3–1.5 × 0.6–0.7
mm, milky white, glabrous. Staminodes 2; filaments
1.6–1.8 mm long, hyaline, glabrous; antherodes not
seen. Ovary narrowly ovoid to oblong, 4–4.7 × 0.9–
1 mm, green, pubescent; style terete, 2.6–2.9 mm
long, green, glabrous; stigma obliquely subcapitate,
0.21 × 0.5 mm, papillate, pale green. Capsules
cylindrical, straight, 13–15 mm long, plagiocarpic,
dehiscing loculicidally along the dorsal side, reddish
brown, tomentose with persistent stigma and calyx.
Seeds numerous, elliptic, reticulate.
Flowering & fruiting: Flowering from July to late
September and fruiting from September to
November (sometimes up to March).

57
Janeesha & Nampy
Fig. 4.
Henckelia gambleana
(C.E.C.Fisch.) A.Weber & B.L.Burtt: a. Habit and habitat; b. A portion of the inflorescence; c. Flower; d. Calyx;
e. Calyx with pistil; f. Bract; g. Stamens; h. Staminode; i. Pistil (from
A.P. Janeesha & Santhosh Nampy
137677).

58 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Habitat: On moist hill slopes, along with Adiantum
raddianum C.Presl (Pteridaceae) and a few species
of moss.
Distribution: Endemic to the southern Western
Ghats (Fig. 6).
Specimens examined: INDIA, Tamil Nadu,
Coimbatore district, Bolampatti valley, 24.02.1917,
C.E.C. Fischer 4059 (FRC); Kanyakumari district,
Muthukuzhivayal grass land, 15.03.1979, A.N.
Henry 60703 (MH); Tenkasi district, Courtallam-
Thekkumalai, 11.11.1984, Jolly Jacob 4072; Ibid.,
11.11.1984, A.R. Sheela 3677; Ibid., 11.11.1984, K.
Leela 4344; Ibid., 11.11.1984, A.M. Rema 4878
(CALI); Tirunelveli district, Agasthyamalai peak,
01.07.1964, A.N. Henry & M. Chandrabose 19215;
Ibid., 24.08.1963, A.N. Henry 17317; Kannikathi,
13.06.1899, C.A. Barber 506; Neelikkattu,
11.10.1992, R. Gopalan 99487 (MH); on the way
to Kakkachi-near Manjolai estate board,
21.07.2016, A.P. Janeesha & Santhosh Nampy 137677
(CALI); Sevgaltai, 24.09.1915, s.coll. 12477 (MH);
Upper Kodayar, 02.09.1981, K.M. Matthew 17831
(RHT); way to Kannikkatti, 05.07.1964, A.N. Henry
& M. Chandrabose 19292 (MH). s.loc., s.d., T.
Surendran 5105 (CALI).
Conservation status: The species occurs in several
populations in three to four localities widely
disjunct in the SW and NW of Tamil Nadu.
Because of this disjunction, the Extent of
Occurrence (EOO) is with of c. 3500 km2 large.
The authors have observed only 27 mature plants
in one population in Kakkachi, in the Tirunelveli
district in Tamil Nadu. This might suggests that
the populations in general are rather small.
However, since the status of other localities is
unknown and potentially intervening localities may
exist, the species is best categorised as Data deficient
(DD).
Notes: This species can be easily differentiated from
the morphologically allied H. incana by its broadly
winged petioles, and much shorter (13–15 mm
long), tomentose capsules.
Typification: Henckelia gambleana was originally
described as a variety of Didymocarpus rottlerianus
Wall. ex R.Br. by Clarke (1883) based on Wight’s
manuscript name D. lanuginosus. Clarke
(1885[“1884”]), subsequently treated it as a variety
of D. tomentosus Wight (as D. tomentosus var.
lanuginosus). Gamble (1924), however, elevated it
to a distinct species (as D. lanuginosus). Gamble’s D.
lanuginosus, being a later homonym of D.
lanuginosus Wallich ex Candolle (1845), was
renamed as D. gambleanus (=H. gambleana) by
Fischer (1938). While searching for the type
specimens, three sheets were found, two in K
(K000858182, K000858183) and one in E
(E00627530). The sheet K000858182 with HRWP
label is annotated “D. lanuginosa, 561/1835” by
Wight. This sheet has two plants and two scapes
separately mounted on it. The other sheet
K000858183 has a field label, Courtallam 561.
There is one plant in vegetative stage and a scape
separately mounted on it. The specimen in E
(E00627530) has three plants, two in vegetative
stage and another flowering plant and a separate
Fig. 5.
Lectotype of
Henckelia gambleana
(C.E.C.Fisch.) A.Weber &
B.L.Burtt. © The Board of Trustees for The Royal Botanic Gardens,
Kew. Reproduced with permission.

59
Janeesha & Nampy
scape. This sheet also has a field label 561,
Courtallam July 1835. Vitek et al. (2000) cited
“Wight 561 (K)” as the type. According to Art.9.17
of ICN (Turland et al., 2018), we take a second-
step lectotypification by narrowing them to a single
specimen at K (K000858182) that agrees with the
protologue and designate the rest of the specimens
as isolectotypes.
Henckelia humboldtiana (Gardner) A.Weber &
B.L.Burtt, Beitr. Biol. P flanzen 70: 346.
1998[“1997”]; T.S.Nayar et al., Fl. Pl. Western Ghats
1: 528. 2014. Didymocarpus humboldtianus Gardner,
Calcutta J. Nat. Hist. 6: 477. 1846; C.B.Clarke in
Hook.f., Fl. Brit. India 4: 353. 1885[“1884”];
Gamble, Fl. Madras 988. 1924; W.L.Theob. &
Grupe in Dassan. & Fosberg, Rev. Handb. Fl.
Ceylon 3: 84. 1981; A.N.Henry et al., Fl. Tamil
Nadu Ind., Ser I: Analysis 2: 132. 1987; Manil., Fl.
Silent Valley 200. 1988; Sasidh., Biodivers. Doc.
Kerala Part 6: Fl. Pl. 332. 2004; N.Anilkumar et al.,
Fl. Pathanamthitta Dist. 370. 2005; T.S.Nayar et
al., Fl. Pl. Kerala 342. 2006. Roettlera humboldtiana
(Gardner) Kuntze, Revis. Gen. Pl. 2: 476. 1891.
Lectotype (designated by Janeesha & Nampy, 2015):
SRI LANKA (CEYLON), Rambodde, Gardner 600
(K [K000858189 digital image!]; isolecto K
[K000858188 digital image!]).
Didymocarpus primulifolius Gardner, Calcutta J. Nat.
Hist. 6: 478. 1846, nom. illeg. Lectotype (designated
by Ranasinghe et al., 2019): SRI LANKA, Hantane
range, Near Candy, s.d., Gardner 601 (K
[K000858191 digital image!]; isolecto BM
[BM000617482 digital image!], K [K000858192
digital image!]). Fig. 7
Perennial, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 4–12,
basal, decussate, rarely in whorls of three; petioles
2–6 cm long, longer in older leaves, narrowly
winged, pubescent; wings 0.5–2 mm broad;
Fig. 6. Distribution of
Henckelia bracteata
Janeesha & Nampy,
H. fischeri
(Gamble) A.Weber & B.L.Burtt and
H. gambleana
(C.E.C.Fisch.)
A.Weber & B.L.Burtt in South India (drawn using QGIS v.3.12.1).

60 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Fig. 7.
Henckelia humboldtiana
(Gardner) A.Weber & B.L.Burtt: a. Habit; b. A portion of the inflorescence; c. Flower; d. Calyx; e. Calyx with
pistil; f. Bract; g. Stamens; h. Staminode; i. Pistil; j. Immature capsule; k. Seeds (a from
A.P. Janeesha & A.J. Robi
134201; b-k from
A.P.
Janeesha, Habeeb Rahman & S. Syam Radh
134221).

61
Janeesha & Nampy
laminae elliptic-ovate, 5–8 × 2.1–5.9 cm, slightly
acute at apex, attenuate at base, margins crenate,
ultimate segments regular, slightly undulate,
surfaces rugose, tomentulose on both surfaces, but
more so on the younger leaves and on lower veins,
becoming less so with age; veins 5–6 pairs on each
side, much branched and conspicuous beneath.
Cymes axillary, lax, 1–4 per plant, 2–3 times
divided, 6–10-flowered, pubescent; peduncles
terete, 8–12 cm long, pale green, pubescent; bracts
two at each branching point, linear to oblong, 4.2–
5.1 mm long, obtuse at apex, tomentose; pedicels
terete, 5–10 mm long, slender, pubescent. Flowers
10–12 × 6–7.5 mm. Calyx 5-partite, free or very
deeply divided; lobes linear, unequal, posterior lobe
smaller, 4.3–4.8 × 0.6–0.75 mm, others 5.1–6 ×
0.72–0.9 mm, slightly acute at apex, margins entire,
pale green with a pinch of reddish brown at the
base, tomentose. Corolla, zygomorphic, ventricose,
two lipped, villous outside, glabrous inside; tube
5–9 × 4.2–5.5 mm, pale lilac to white with a yellow
patch in the throat; lobes rounded, rounded at apex,
white to pale lilac, upper lip of two lobes, 3.1–3.4 ×
3.2–3.7 mm, lower lip of three lobes, 3.3–4.4 × 3.5–
5.6 mm. Stamens 2, adnate to the base of corolla,
included; filaments 5 mm long, slightly bearded,
pale green at the top, base hyaline; anthers reniform,
c. 1.3 × 0.8 mm, milky white, glabrous. Staminodes
2. Ovary ovoid to oblong, 8–9 × 1–1.1 mm, green,
pubescent; style terete, 3.5–4 mm long, green, base
slightly hairy; stigma obliquely subcapitate, 0.65–
0.8 × 0.34–0.42 mm, papillate, pale green. Capsules
linear, straight or slightly curved, 12–21 × 1.1–1.6
mm, plagiocarpic, dehiscing loculicidally along the
dorsal side, pale green, tomentulose with persistent
stigma. Seeds numerous, elliptic, 0.32–0.5 × 0.1–
0.19 mm, slightly acute at apex, truncate at base;
testa dark brown to black, reticulate.
Flowering & fruiting: Flowering from May to
September and fruiting from July to November.
Habitat: On moist slopes or on rocks, at medium
elevations of 350 m, along with Selaginella sp.
(Selaginellaceae), Pteris sp. (Pteridaceae), Impatiens
gardneriana Wight, I. acaulis Arn. (both
Balsaminaceae) and a few grass species.
Distribution: South India and Sri Lanka (Fig. 11).
Specimens examined: INDIA, Kerala, Ernakulam
district, Neriyamangalam, 20.08.1965, K.M.
Sebastine 25087 (MH); Idukki district, Munnar,
08.11.1985, P.C. Binoy 05047 (TBGT); Kollam
district, Thenmala, 01.10.1960, Joseph 2, 3 (BSI, E);
Kottayam district, Pambanar, 24.05.1964, K.
Vivekananthan 23998; Pullupara, 24.09.1964, K.
Vivekananthan 21314 (MH); Vagamon, 03.10.2015,
C. Pramod & A.P. Janeesha 137644, 137647 (CALI);
Kozhikode district, Kakkayam, 20.07.2013, A.P.
Janeesha & A.J. Robi 134201; Ibid., 26.05.2015, S.
Resmi & A.P. Janeesha 137618; Ibid., 26.05.2015,
A.P. Janeesha, P.K. Dilna & Santhosh Nampy 137620;
Kakkayam-Ghat road side, 03.10.2016, A.P.
Janeesha & K. Haseem 137687; way to Kakkayam
forest office, after 12th turn, 19.09.2013, A.P.
Janeesha, Habeeb Rahman & S. Syam Radh 134221
(CALI); Palakkad district, Koomankundu,
18.09.1982, K. Prasannan 10991; Pulippara,
18.09.1982, T. Sabu 10999 (CALI); Wayanad
district, Boys town-Kannur Mananthavadi route,
15.09.2016, Shimi Cheriyan, P.K. Dilna & A.P.
Janeesha 137687 (CALI); Travancore, s.d., C.C.
Calder & M.S. Ramaswami 1312 (CAL). Tamil
Nadu, Coimbatore district, Aliyar Dam,
24.08.1963, K.N. Subramanian 782 (FRC);
Anamalai, 04.11.1897, s.coll. s.n.; Bolamapatty hills,
s.d., s.coll. 14019 (MH); Dindigul district, Kodaikanal
hills, 14.10.1919, K.C. Jacob 16114 (MH);
Kodaikanal-Pulney, 20.10.1898, A.G. Bourne 193
(CAL); Perumal malai, 24.10.1977, M. Chandrabose
51666; Poombarai, 18.10.1977, M. Chandrabose
51365 (CAL, MH); way to Pannaikadu, 15.1968,
s.coll. 30849 (MH); Kanyakumari district,
Kurathividuthi estate, 11.11.1989, S.R. Srinivasan
86969 (MH); Madurai district, Anna-Manalur,
06.08.1985, K.M. Matthew 41821; Dolphin’s nose,
07.07.1959, K.M. Matthew 433; Ibid., 31.10.1985,
K.M. Matthew 42477 (RHT); Nilgiri district,
Coonoor, 27.07.1957, K.M. Sebastine 4060 (MH);
Ramanthapuram district, Deviar estate, 11.06.1979,
S.R. Srinivasan 63518 (CAL, MH); Virudhunagar
district, Mysore medu, 22.10.1988, S.R. Srinivasan

62 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
89452; s.loc., 11.11.1989, S.R. Srinivasan 86972
(MH). s.loc., 20.05.1913, s.coll. 691; s.loc., 23.08, s.coll.
229; s.loc., s.d., s.coll. 1784 (CAL).
Conservation status: This species is widely distributed
in Kerala and Tamil Nadu and in Sri Lanka and we
consider it as of Least Concern (LC) according to
IUCN (2019) criteria.
Notes: This species is close to H. bracteata and H.
meeboldii in leaf shape, but can be easily
distinguished by its linear to oblong bracts, linear
calyx lobes and hairy connectives.
Henckelia incana (Vahl) Spreng., Syst. Veg. 1:
38. 1824; Pallith., Fl. Sirumalai hills 180. 2001;
T.S.Nayar et al., Fl. Pl. Western Ghats 1: 528. 2014.
Roettlera incana Vahl, Enum. Pl. 1: 88. 1804.
Didymocarpus rottlerianus Wall., Numer. List 778.
1829. Type: INDIA, Madurai, s.d., Röttler (probably
lecto C [C10012757 digital image!]; a photograph
of the type from C is available at E [E00155176
photograph E!]).
Didymocarpus tomentosus Wight, Icon. Pl. Ind.
Orient. 4: t. 1349. 1848; C.B.Clarke in Hook.f., Fl.
Brit. India 4: 353. 1885[“1884”], excluding var.
lanuginosa; Rama Rao, Fl. Pl. Travancore 295. 1914;
Gamble, Fl. Madras 989. 1924; S.V.Ramaswamy &
Razi, Fl. Bangalore Dist. 545. 1973; Vajr., Fl.
Palghat Dist. 332. 1981; K.M.Matthew, Material
Fl. Tamil Nadu 287. 1981; A.N.Henry et al., Fl.
Tamil Nadu Ind., Ser I: Analysis 2: 133. 1987;
K.M.Matthew, Excursion Fl. Central Tamil
Nadu 359; M.Mohanan & A.N.Henry, Fl.
Thiruvananthapuram Dist. 338. 1994; Pull. & Ali
Moulali, Fl. Andhra Pradesh 2: 680. 1997; Sasidh.,
Biodivers. Doc. Kerala Part 6: Fl. Pl. 332. 2004;
T.S.Nayar et al., Fl. Pl. Kerala 343. 2006; Pull. et al.,
Fl. Eastern Ghats 4: 636. 2011. Didymocarpus
rottlerianus Wall. ex R.Br. var. tomentosus (Wight)
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5:
101. 1883. Lectotype (first-step designated by Vitek
et al., 2000; second-step designated here): INDIA,
Tamil Nadu, Jamalay near Coimbatore, 11.1847,
Wight 2352 (K [K000858196 digital image!];
isolecto K [K000858193, K000858195 digital
images!]). Fig. 8
Vernacular name: Elichuzhien (Malayalam).
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 6–18,
basal, usually in whorls of three, rarely two; petioles
0.8–3 cm long, pubescent, winged; wings 2.3–3.4
mm broad; laminae elliptic-widely ovate, 5.5–11 ×
3–9 cm, acute to obtuse at apex, attenuate at base,
margins shallowly crenate, ultimate segments
regular, surfaces usually rugose, pilose on the upper
surface, tomentose on lower surface, but more so
on the younger leaves and on lower veins,
becoming less so with age; veins usually 5 on each
side, much branched and conspicuous beneath.
Cymes axillary, dense, 2–6 per plant, 2–4 times
divided, 8–17-flowered, pubescent; peduncles
terete, 6–23 cm long, reddish brown, pubescent;
bracts two at each branching, ovate to lanceolate
2.2–2.8 × 0.5–0.6 mm, obtuse to narrowly acute at
apex, margins crenate, hispid along the margin;
pedicels 7–15 mm long, terete, narrow, glandular
hairy. Flowers 10–15 × 6–10 mm. Calyx 5-partite,
very deeply divided; lobes lanceolate, unequal,
posterior lobe small, 3.4–4.4 × 1–1.23 mm, others
4.1–5.2 × 1.2–1.3 mm, slightly acute to obtuse at
apex, margins entire, reddish brown, hispid. Corolla
zygomorphic, ventricose, two lipped, glandular
hairy outside, glabrous inside; tube 6.7–7.3 × 5.4–
7.1 mm, lilac to pale lilac with a yellow patch in
the throat; lobes oblate to widely, rounded to
slightly acute at apex, lilac to pale lilac, upper lip of
two lobes, 3.6–4.2 × 2.7–4.5 mm, lower lip of three
lobes, 4.3–4.8 × 3.9–5.2 mm. Stamens 2, adnate to
the base of the corolla, included; filaments 4–5.4
mm long, glabrous, rarely few (3–5) hairs towards
the top, green on the top and colourless towards
base; anthers reniform, 1.6–1.8 × 0.7–0.9 mm, milky
white, glabrous. Staminodes 2; filaments 2.4–3.1
mm long, hyaline, glabrous; antherodes globose or
not prominent, glabrous, if globose rarely with 1
or 2 hairs. Ovary ovoid to oblong, 3.8–4.3 × 1.16–
1.46 mm, green, glandular hairy; style terete, 3.23–
3.86 mm long, green, glabrous; stigma obliquely
subcapitate, 0.4–0.6 × 0.74–1 mm, papillate, pale
green. Capsules cylindrical, straight or slightly

63
Janeesha & Nampy
Fig. 8.
Henckelia incana
(Vahl) Spreng.: a. Habit; b. A portion of the inflorescence; c. Flower; d. Calyx; e. Calyx with pistil; f. Stamens;
g. Staminode; h..
..
. Pistil; i. Immature capsule; j. Seeds (from
A.P. Janeesha, P.M. Shahina & K.M. Manudev
134255).

64 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
curved, 21–28 × 2–2.4 mm, plagiocarpic, dehiscing
loculicidally along the dorsal side, reddish brown,
hispid, with persistent stigma and calyx. Seeds
numerous, elliptic, 0.5–0.56 × 0.22–0.26 mm,
slightly acute at apex, truncate at base; testa dark
brown to black, reticulate.
Chromosome numbers: n = 27 (Thathachar, 1942), n
= ± 45, 2n = ± 90 (Ratter & Prentice, 1967).
Flowering & fruiting: Flowering from May to
October and fruiting from July to December.
Habitat: On moist slopes and in rock crevices,
together with Begonia floccifera Bedd.
(Begoniaceae), Justicia japonica Thunb.
(Acanthaceae), species of moss and grass.
Distribution: Endemic to the southern Western
Ghats (Fig. 11).
Specimens examined: INDIA, Karnataka, Bangalore
district, Savandurga hilltop, 14.08.1979, K.P.
Sreenath 8954 (CAL); Mysore district, Devagiri
betta-Beduguli, 08.09.1961, Seshagiri Rao Rolla
73794; Ibid., 17.04.1962, A.S. Rao 80020 (E, CAL).
Kerala, Idukki district, Alampetty, Marayur,
30.07.1982, V.P.K. Nambiar & N. Sasidharan 2168
(KFRI digital image); Calvary mount, 13.11.1981,
V.S. Raju 71239 (CAL); Devikulam, 14.07.1993,
A. Nazarudeen 17834 (TBGT); Idukki dam,
24.08.1981, V.S. Raju 71172; Kattappana,
23.08.1977, K. Vivekananthan 50486; Ibid.,
23.08.1981, V.S. Raju 71162; Ibid., 27.09.1981, C.N.
Mohanan 72013 (MH); Kulamavu, 12.10.1982, C.N.
Mohanan 74583 (CAL, MH); Marakkanam,
23.08.1977, K. Vivekananthan 50486 (CAL);
Munnar forest division, Suryanelli, 19.10.1986, B.
Gurudev Singh & K.R. Sasidharan 12427 (FRC); on
the hill between Cheruthoni and Idukki dam,
24.08.1981, V.S. Raju 71172 (CAL); way to Calvary
mount, 13.11.1981, V.S. Raju s.n. (MH); Kollam
district, Aryankavu, 08.06.1977, E. Sarada Amma
21894 (CALI); Kazhuthuruthy-Thenmala,
20.12.1978, C.N. Mohanan 59572; Ottakkal hills
near Arnarkkad, 23.12.1969, M.V. Viswanathan
MVV307 (MH); Thenmala, 13.07.2010,
Geethakumary 69602 (TBGT); Kottayam district,
Kumarakam, 25.12.1980, A.S. Suletha 92205 (DEV);
Palakkad district, Nelliyampathy, 16.09.1998, K.
Radhakrishnan 29060 (TBGT); Ibid., 2016, P.
Soumya & A.P. Janeesha 137691; Parambikulam,
20.04.1976, Rugmini Devi 12129; Ibid.,
28.04.1976, M.P. Ramani s.n.; Ibid., 26.05.2014,
A. Kabeer & A.P. Janeesha 134262 (CALI);
Walayar forest, 10.07.1963, J. Joseph 17049 (E,
MH); Thiruvananthapuaram district, Arivikarai,
10.10.1928, V. Narayanaswami 934 (MH);
Ponmudi, 19.09.1968, K.M. Matthew 9190; Ibid.,
12.09.1970, K.M. Matthew 12411 (RHT); Ibid.,
16.08.1980, M. Mohanan 69225; Ibid., 17.11.1977,
M. Mohanan 52549; Ibid., 25.05.1979, M. Mohanan
63280 (MH); Ibid., s.d., M. Abdul Jabbar 13777; Ibid.,
29.10.1983, Satheesh Kumar 5921 (TBGT); Ibid.,
28.11.2012, A.P. Janeesha, P.M. Shahina & K.M.
Manudev 134255; Ibid., 31.07.2014, Santhosh Nampy
& A.P. Janeesha 134146, 134148; Ibid., 31.07.2014,
S. Syam Radh, Santhosh Nampy & A.P. Janeesha
134152, 134165, 134168 (CALI); Ponmudi-way to
upper sanatorium, 25.05.1979, M. Mohanan 63280;
Ibid., 16.08.1980, M. Mohanan 69225 (CAL);
Wayanad district, Edakkal cave-inside, 08.07.2016,
A.P. Janeesha & P.G. Arunkumar 137674 (CALI).
Tamil Nadu, Coimbatore district, Aliyar
submergible area, 28.07.1962, K.M. Sebastine 14686
(CAL, MH); Anamalai, 08.10.1901, C.A. Barber
3629; Ibid., 10.10.1901, C.A. Barber 3659 (MH);
Ibid., 26.06.1973, R.N. Kajal & G.N. Tribedi 567
(CAL); Andiparai shola R.F., 26.06.1973, E.
Vajravelu 44302 (MH); around Attakatti, s.d., J.
Joseph 12685 (CAL, MH); Bolampetty valley,
24.02.1917, C.E.C. Fischer 4059 (CAL); Hassanur,
25.08.1914, s.coll. 10634; Kurudimalai, 27.09.1956,
K. Subramanyam 865; Ibid., 17.11.1968, J.L. Ellis &
S. Karthikeyan 31325; Ibid., 20.01.1970, M.V.
Viswanathan MVV375 (MH); Kuridimalai-
northern slopes, 27.09.1956, K. Subramanyam
865 (CAL); Marudamalai, 24.06.1930, V.
Narayanaswami 3072 (MH); Ibid., 09.08.2014,
K. Thoiba & A.P. Janeesha 134280; Valparai-
Shivamalai, 13.12.2013, A.P. Janeesha & Santhosh
Nampy 134244; Velliangiri hills-way to temple,
29.05.2014, S. Syam Radh, R. Rajeevan & A.P.

65
Janeesha & Nampy
Janeesha 138187, 138189, 138191; Ibid., 30.09.2016,
Nikhil Krishna & A.P. Janeesha 137689 (CALI);
Dindigul district, Kodaikanal, 20.11.2012, K.M.
Manudev, A.P. Janeesha & Santhosh Nampy 134254
(CALI); Perumparai, 22.10.1977, M. Chandrabose
51612 (CAL); Ratter water fall, 25.02.2014, A.
Kabeer, K. Shinoj & A.P. Janeesha 134258 (CALI);
Sirumalai, 16.05.1958, J.M. Pallithanam 3356
(BLAT); Kanyakumari district, Kooladai,
08.09.1969, B.V. Shetty 32293; Maranthuvalmalai,
18.10.1985, R. Gopalan 83374; Sevgaltai,
24.09.1915, s.coll. 12447 (MH); Thekkumalai,
19.11.1956, K.M Sebastine 1368 (CAL, MH);
Madurai district, Amaravathi sagaram-lower camp,
22.06.1968, D.B. Deb 30121; Cumbum valley,
20.06.1959, K. Subramanyam 8048 (MH); High way
hut, 05.1917, E. Blatter & Hallberg 355 (BLAT,
CAL); Sirumalai, 23.08.1913, C.A. Barber 9058
(MH); Vengayapparai, 21.11.1985, K.M. Matthew,
A. Usha & N. Rajendran 42987 (RHT); way to
Perumparai, 22.10.1977, M. Chandrabose 51612
(MH); Namakkal district, Kolli hills, 01.08.1978,
N. Venugopal 15984 (CAL); Nilgiris district, Bimaka
shola, 26.08.1970, G.V. Subba Rao 36346 (MH); Kil
Kotagiri-on the way, 25.10.1956, K. Subramanyan
1171 (CAL); Kodanad view point, 24.07.1970, E.
Vajravelu 35084 (MH); Mudumalai, 03.06.1994,
S.K. Mandal 7214 (CAL); Naduvattam, 28.06.1970,
B.V. Shetty 34457; Ibid., 30.08.1970, B.D. Sharma
35926; way to Anakkatai from Ebanad, 27.07.1972,
G.V. Subba Rao 41563; way to Kilkotagiri,
25.10.1972, K. Subramanyam 1171 (MH); way to
Naduvattam from Devar shola, 21.07.1960, K.
Subramanyam 10517 (CAL, MH); Pykara falls,
08.1884, J.S. Gamble 15292 (CAL, BM digital
image); Ibid., 06.1884, J.S. Gamble 14241
(BM digital image); Pudukkottai district,
Ponnamaravathi, 22.07.1984, C. Arulappan s.n.
(MH); Ramnad district, Kendiparai-Ayyanar koil,
23.09.1971, E. Vajravelu 38704; Mudaliarattu,
21.09.1917, P.S. Jivanna Rao 15096 (MH); Salem
district, Kolli hills, 10.09.1992, A.A. Ansari 97066;
Retreat area-Yercaud, 24.07.1965, S. Karthikeyan
26853 (MH); Thiruchirapalli district, Trichy,
18.10.1985, P.C. Jose Kutty 1146 (DEV); Tenkasi
district, Courtallam, 04.07.1974, T.N. Reetha Bai
30815 (CALI); Tirunelveli district, Chunkankada
hills, near Sree Ayyappa College, 20.07.2016, A.P.
Janeesha, S. Resmi & P.G. Arunkumar 137676
(CALI); Karayar Dam, 30.07.1989, Shanavas Khan
5853 (TBGT); Kohimalai, 02.11.1914, s.coll. 11396;
Kulurathumottai grassland, 16.10.1992, R. Gopalan
9997; Mahendragiri, 17.09.1916, s.coll. 13139 (MH);
near Kodayar power house, 19.07.2016, A.P.
Janeesha, S. Resmi & P.G. Arunkumar 137675
(CALI); Neelikkathu R.F., 03.05.1989, R. Gopalan
90532 (MH); Papanasam project, 07.11.1959, K.M.
Sebastian 9560 (CAL); Thulukkam parai,
28.11.1969, B.V. Shetty 32947; Vasudevanallur R.F.,
03.10.1971, E. Vajravelu 38847 (MH); Vellore
district, slopes of Konamalai, 18.07.1961, C.P.
Sreemadhavan CPS741; s.loc., 27.09.1986, S.R. Raja
& Nagavath s.n. (MH). State unknown, s.loc.,
06.1883, J.S. Gamble 11780 (CAL). Peninsular India,
s.d., R. Wight 2352 (CAL).
Conservation status: This species is widely distributed
in Karnataka, Kerala and Tamil Nadu and the
thresholds for Extent of Occurrence (EOO) and
Area of Occupancy (AOO) are not met by this
species to fall into one of the threat categories. We
therefore consider it as of Least Concern (LC)
according to IUCN (2019) criteria.
Notes: Wallich (1829) in his catalogue listed
Didymocarpus rottlerianus and quoted Roettlera incana
as a synonym. D. rottlerianus Wall. is thus validated
by Roettlera incana and became nomenclaturally
superfluous (for more information, see Burtt, 1954)
and both names have Rottler’s specimen as type.
Typification: Vahl (1804) described Roettlera incana
(H. incana (Vahl) Spreng.) based on Röttler’s
specimen from mountains in Madurai, India. Vitek
et al. (2000) cited the sheet at C (C10012757 digital
image!) as the holotype. A further search revealed a
photograph of the above sheet from C at E with
the barcode E00155176 (photograph E!). The sheet
C10012757 contains two flowering specimens
mounted separately but do not bear any field label
or collector’s information to ascertain it as a type
material. A further search revealed one sheet with

66 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
two barcodes at K (K001129365, K001111886
digital images!) collected by Röttler. Except the one
specimen mounted on the upper side of the sheet,
all others agree with H. incana. However, the field
label attached at the top of the sheet indicated as
locality “ooty”. Hence, this specimen can also not
be considered for typification. According to Stafleu
and Cowan (1983), Röttler’s main collections are
kept at K and LIV and other materials are at C, ER,
FI, H, H-Aeh, JE, LE, M, MH, MO, MW and NY.
Even though a lectotypification is necessary, access
to some herbaria material has been impossible in
time, lectotypification is postponed here and will
be carried out in a separate paper.
Wight (1848) described Didymocarpus tomentosus
Wight and included an illustration of his collection
from Kaitie falls, on the Neilgherries. It was further
mentioned in the protologue “I have since received
many specimens from hills near Coimbatore, and
have often met with it in similar situations”. All of
this is original material and available for
lectotypification. Vitek et al. (2000) cited “Wight
(K)” as the type. We traced three relevant specimens
at K (K000858193, K000858195, K000858196
digital images!). One sheet K000858193 with an
HRWP label is annotated “Did. tomentosa?, Jamalay
Coimbatore” by Wight. The sheets K000858195
and K000858196 have the annotations “2352
Didymocarpus tomentosus Wt”, with field labels, one
with “Jamalay, June 1848” and the other “Jamalay
near Coimbatore, November 1847” respectively.
Specimens from Kaitie falls are, however, not
available in any herbaria. The sheet K000858196,
earliest among them with five mature plants and
scapes is chosen here as the second-step lectotype
while the others (K000858193, K000858195) are
consequently isolectotypes.
Henckelia innominata (B.L.Burtt) A.Weber &
B.L.Burtt, Beitr. Biol. Pflanzen 70: 347.
1998[“1997”]; T.S.Nayar et al., Fl. Pl. Western
Ghats 1: 528. 2014. Didymocarpus innominatus
B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 21:
201. 1954; B.D.Sharma et al., Fl. Karnataka Analysis
195. 1984; A.N.Henry et al., Fl. Tamil Nadu Ind.,
Ser I: Analysis 2: 132. 1987; M.Ahmedullah &
M.P.Nayar, Endemic Pl. Indian Region 1: 143.
1987; N.P.Singh, Fl. Eastern Karnataka 2: 475.
1988; Pull. & N.Yesoda, Fl. Anantapur Dist. 182.
1989; Sasidh. & Sivar., Fl. Pl. Thrissur Forest 329.
1996; Pull. & Ali Moulali, Fl. Andhra Pradesh 2:
679. 1997; Sasidh., Biodivers. Doc. Kerala Part 6:
Fl. Pl. 332. 2004; T.S.Nayar et al., Fl. Pl. Kerala
342. 2006; Pull. et al., Fl. Eastern Ghats 4: 361. 2011.
Lectotype (first-step designated by Vitek et al., 2000;
second-step designated here): s.loc., s.d., Wallich 778
(K [K001111885 (778B) digital image!]; isolecto K
[K00085184, K001111884 digital images!]).
Figs. 9 & 10
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 6–15,
basal, usually in whorls of three; petioles 2.5–9 cm
long, pubescent, winged; wings 1–3 mm broad;
laminae elliptic, 8.5–12 × 4.7–6.7 cm, acute at apex,
narrowly cuneate to attenuate at base, margins
doubly serrate, first serration deep and ultimate
segments irregular, surfaces usually rugose; hispid
on both surfaces, but more so on the younger leaves
and on lower veins, becoming less so with age, veins
usually 5 on each side, much branched and
conspicuous beneath. Cymes axillary, 2–7 per plant,
3–5 times divided, 12–22-flowered, pubescent;
peduncles terete, 18–24 cm long, green, pubescent;
bracts two at each branching point, linear, 3–5 ×
1.3–2.4 mm, obtuse at apex, margins entire, hispid;
pedicels terete, 4–14 mm long, narrow, pubescent.
Flowers 10–20 × 4–9 mm. Calyx 5-partite, very
deeply divided or basally connate; lobes elliptic to
oblong, unequal, posterior lobe small, 2–2.2 × 0.68–
0.77 mm, others 2.3–2.9 × 0.71–1 mm, obtuse at
apex, margins entire, reddish brown towards the
base and green towards the apex, hispid on both
sides. Corolla zygomorphic, ventricose, two lipped,
glandular hairy outside, glabrous inside; tube 6.8–
7.9 × 5.3–6.2 mm, pale lilac with a yellow patch in
the throat; lobes widely ovate to orbicular, rounded
at apex, slightly undulate, pale lilac, upper lip of
two lobes, 4.5–5.2 × 5.3–5.9 mm, lower lip of three
lobes, 5.4–5.8 × 5.3–6.2 mm. Stamens 2, adnate to
the base of the corolla, included; filaments 3.3–3.8
mm long, glabrous, green on the top and colourless

67
Janeesha & Nampy
Fig. 9.
Henckelia innominata
(B.L.Burtt) A.Weber & B.L.Burtt: a. Habit and habitat; b. A potion of the inflorescence; c. Flower; d. Calyx; e. Calyx
with pistil; f. Stamens; g. Staminode; h. Pistil; i. Capsule; j. Seeds (a from
A.P. Janeesha, Dani Francis, S. Resmi & K.S. Arathy
137682;
b-j from
A.P. Janeesha & Santhosh Nampy
137681)
.

68 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
towards base; anthers reniform, 1.45–1.6 × 0.62–
0.7 mm, milky white, glabrous. Staminodes 2;
filaments 1.8–2 mm long, hyaline, glabrous;
antherodes a small knob like structure. Ovary ovoid
to oblong, 3.1–3.5 × 1.15–1.3 mm, green, glandular
hairy; style terete, 2.4–2.6 mm long, green,
glabrous; stigma obliquely subcapitate, 0.53–0.58
× 0.3 mm, papillate, pale green. Capsules cylindrical,
straight, 12–19 × 1.4–1.6 mm, plagiocarpic,
dehiscing loculicidally along the dorsal side, reddish
brown, tomentulose with persistent stigma and
calyx. Seeds numerous, elliptic, 0.43–0.56 × 0.2 mm,
slightly acute at both ends; testa dark brown to
black, reticulate.
Chromosome numbers: n = 16, 2n = 32 (Ratter &
Prentice, 1967).
Flowering & fruiting: Flowering from (May) June
to September and fruiting from August to
December.
Habitat: On moist slopes and in rock crevices, along
with Selaginella sp. (Selaginellaceae), Spermacoce
alata Aubl. (Rubiaceae) and a few grass species.
Distribution: Endemic to the southern Western
Ghats (Fig. 11).
Specimens examined: INDIA, Karnataka, Coorg
district, s.loc., s.d., s.coll. s.n. (CAL). Kerala, Idukki
district, Chinnar, 19.01.1986, B. Gurudev Singh &
K.R. Sasidharan 12428 (FRC); Ibid., 07.10.1994, E.S.
Santhosh Kumar 17595 (TBGT); Kollam district,
Achencoil, 22.05.1979, C.N. Mohanan 63025 (MH);
Moonnamra-Thenmala, 17.08.2016, A.P. Janeesha
& Santhosh Nampy 137681; Moonnamra-near
constructing railway track, 17.08.2016, A.P.
Janeesha, S. Resmi, Dani Francis & K.S. Arathy
137682 (CALI); Palakkad district, Dhoni reserve
forest, 20.07.1963, J. Joseph 17220 (MH);
Pathanamthitta district, Achankovil to Mekkarai,
05.09.1913, M. Rama Rao 1652 (CAL). Tamil
Nadu, Coimbatore district, Attakatti, 04.07.1961,
J. Joseph 12685 (MH); Ibid., 21.11.1980, M.
Chandrabose & V. Chandrasekaran 69044 (MH,
CAL); Parappalar dam, 11.11.1986, K.M. Matthew
47293 (RHT); Siruvani, 14.08.1960, A.N. Henry 351
(MH); Siruvani-near Bungalow, 14.08.1960, A.N.
Henry 350 (BLAT). Dharmapuri district, Chitteri
hills, 12.08.1978, K.M. Matthew 16362; Ibid.,
09.01.1980, K.M. Matthew 25618 (RHT);
Kanyakumari district, s.loc., 29.10.2015, K. Thoiba
& A.P. Janeesha 137651, 137652 (CALI); Nilgiri
district, s.loc., s.d., G. Thomson 1351 (CAL); Ramnad
district, Ayyanarkoil, 23.09.1971, E. Vajravelu
38715; Cumbummedu, 14.03.1970, E. Vajravelu
33767; Yanaimutti rocks, 13.12.1972, E. Vajravelu
39412 (MH); Salem district, Krishnagiri,
24.09.1917, s.coll. 14935 (MH); Yercaud,
28.06.1980, V. Sainaba 28364 (CALI); Tenkasi
district, Courtallam, 18.07.1901, C.A. Barber 3374;
Ibid., 16.07.1907, C.A. Barber 8378; Ibid., 23.07.1957,
K. Subramanyam 3745 (MH); Ibid., 29.06.1980, K.
Indira Devi 5760; Ibid., 30.06.1980, K.S. Hema 8143;
Ibid., 30.06.1980, T. Usha 30144; Ibid., 30.06.1980,
M.S. Baby Usha Kiran 3783 (CALI); Tirunelveli
district, around Shembagadevi, 27.09.1975, K.K.N.
Fig. 10. Lectotype of
Henckelia innominata
(B.L.Burtt) A.Weber &
B.L.Burtt. © The Board of Trustees for The Royal Botanic Gardens,
Kew. Reproduced with permission.

69
Janeesha & Nampy
Fig. 11. Distribution of
Henckelia humboldtiana
(Gardner) A.Weber & B.L.Burtt,
H. incana
(Vahl) Spreng. and
H. innominata
(B.L.Burtt) A.Weber
& B.L.Burtt in South India (drawn using QGIS v.3.12.1).
Nair 1167 (CAL); forest near Papanasam project,
07.11.1959, K.M. Sebastine 9560; Kannakattii,
13.06.1899, C.A. Barber 505 (MH); Mekkarai-on
rock, 06.09.1913, C.C. Calder & M.S. Ramaswami
620 (CAL); Naterikal, 21.09.1914, s.coll. 10952
(MH); Sengalten, 16.02.1913, D. Hooper & M.S.
Ramaswami 39219 (CAL); Vasudevanallur,
13.11.1925, S.R. Raju 17801; Ibid., 03.10.1971, E.
Vajravelu 38845 (MH); s.loc., 01.09.1913, C.C.
Calder & M.S. Ramaswami 507. s.loc., 01.06.1905,
C.E.C. Fischer 22; s.loc., s.d., M.S. Ramaswami 607;
s.loc., s.d., s.coll. 67; s.loc., s.d., R. Wight 2678 (CAL,
E); s.loc., s.d., s.coll. 778 (CAL).
Conservation status: The species occurs widely in
13 districts in three states, Karnataka, Kerala and
Tamil Nadu. The Extent of Occurrence (EOO) is
estimated to be c. 34100 km². However, during field
surveys of two localities, only 106 mature plants
were observed, suggesting that the populations are
not large. Not all of the localities are in protected
areas and those outside face high risks of habitat
destruction, particularly one that is located very near
to a railway track construction site. However,
because of the number of populations and localities,
the species can be considered at present of Least
Concern (LC) according to IUCN (2019) criteria.
Typification: Wallich (1829) in his catalogue named
this species Didymocarpus rottlerianus Wall. but
quoted Roettlera incana Vahl as its synonym. The
name D. rottlerianus thus got validated and became
nomenclaturally superfluous (for more information,
see Burtt, 1954). Burtt (1954) named Wallich’s
specimen 778 as D. innominatus B.L.Burtt. Vitek et
al. (2000) cited “Wallich 778 [K]” as the type. We
have traced five Wallichian collections at K
(K001129365, K00085184, K001111885,
K001111884, K001111886 digital images!). In order
to narrow down the choice to a single specimen,
we carried out a second-step lectotypification here.
The sheets K001129365 (labelled 777C & D) and

70 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
K001111886 are mixed collections, and the
specimens mounted on the upper side alone belong
to H. innominata. K00085184, K001111885,
K001111884 are labelled 778, 778B & 778.1
respectively and all belong to H. innominata. Of
these, sheet K001111885 with a complete plant and
inflorescence is chosen here as the lectotype while
the other specimens (K00085184, K001111884) are
consequently isolectotypes.
Henckelia lyrata (Wight) A.Weber & B.L.Burtt,
Beitr. Biol. Pflanzen 70: 349. 1998[“1997”];
T.S.Nayar et al., Fl. Pl. Western Ghats 1: 528. 2014;
Janeesha & Nampy, Phytotaxa 268(1): 80. 2016;
M.P.Geethakumary et al., Phytotaxa 284(2): 147.
2016. Didymocarpus lyratus Wight, Icon. Pl. Ind.
Orient. 4: t. 1350. 1848; C.B.Clarke in Hook.f., Fl.
Brit. India 4: 353. 1885[“1884”]; Rama Rao, Fl.
Pl.Travancore 295. 1914; Gamble, Fl. Madras 2:
988. 1924; A.N.Henry et al., Fl. Tamil Nadu Ind.,
Ser I: Analysis 2: 132. 1987; M.Ahmedullah &
M.P.Nayar, Endemic Pl. Indian region 1: 143. 1987;
Pull. et al., Fl. Eastern Ghats 4: 361. 2011. Roettlera
lyrata (Wight) Kuntze, Revis. Gen. Pl. 2: 476. 1891.
Lectotype (designated by Janeesha & Nampy, 2016):
INDIA, Tamil Nadu, Courtallam, 09.1835, Wight
559 (E [E00155173 digital image!]; isolecto E
[E00155174, E00155173 digital images!], K
[K000858187 digital image!]).
Didymocarpus lyratus Wight var. protractus
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5:
102. 1883, syn. nov.; A.N.Henry et al., Fl. Tamil
Nadu Ind., Ser I: Analysis 2: 132. 1987. Lectotype
(designated by Vitek et al., 2000): INDIA, Pulney
hills, Beddome 8505 (BM). Fig. 12
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 6–9,
basal, usually in whorls of three; petioles 4.5–14 cm
long, pubescent, lacerate; laminae widely elliptic-
orbicular, 4.5–7.5 × 4.5–6.3 cm, rounded to obtuse
at apex, lyrate at base, margins doubly crenate to
serrate, sometimes the first crenation is very deep
and irregular, surfaces usually rugose, pilose on both
surfaces, but more so on the younger leaves and on
lower veins, becoming less so with age; veins usually
5 on each side, much branched and conspicuous
beneath. Cymes axillary, 3–9 per plant, 3–5 times
divided, 12–21-flowered, pubescent; peduncles
terete, 13–29 cm long, green, pubescent; bracts two
at each branching point, linear, 3–5 × c. 1 mm,
obtuse to rounded at apex, margins entire, pilose;
pedicels terete, 4–25 mm long, narrow, pubescent.
Flowers 10–14 × 5–7 mm. Calyx 5-partite, very
deeply divided; lobes broadly lanceolate, unequal,
posterior lobe small, 2–4.5 × 0.5–1.3 mm, others
4–11 × 0.7–3 mm, obtuse at apex, margins entire,
green, pilose. Corolla zygomorphic, ventricose, two
lipped, minutely villous outside, glabrous inside;
tube 8–10 × 5–6.7 mm, white to pale lilac with a
yellow patch in the throat; lobes widely ovate, ovate
to rounded at apex, white to pale lilac, upper lip of
two lobes, 3.7–4.1 × 4–4.8 mm, lower lip of three
lobes, 4–5 × 4.2–5.3 mm. Stamens 2, adnate to the
base of the corolla, included; filaments 3.5–4.2 mm
long, glabrous; anthers reniform, 1.5–1.6 × c. 0.62
mm, milky white. Staminodes 2; filaments 3.1–3.5
mm long, glabrous; antherodes highly reduced.
Ovary ovoid to oblong, 2.6–3 × 0.78–0.86 mm,
pubescent; style terete, 3–3.8 mm long, glabrous;
stigma obliquely subcapitate, c. 0.34 mm diam.,
papillate, white to pale green. Capsules cylindrical,
straight or slightly curved, 12–36 mm long,
plagiocarpic, dehiscing loculicidally along the
dorsal side, green, glandular hairy with persistent
stigma and calyx. Seeds numerous, oblong-ovoid
to elliptic, 0.4–0.51 × 0.17–0.21 mm, slightly acute
at apex; testa dark brown to black, reticulate.
Flowering & fruiting: Flowering from May to
October and fruiting from August to December.
Habitat: On moist rocks, in association with Porella
sp. (Porellaceae).
Distribution: Endemic to the southern Western
Ghats (Fig. 17).
Specimens examined: INDIA, Kerala, Idukki district,
Neymakkad gap, N 10°082 46.322', E 77°052
16.93', 1784 m, 22.10.2004, M.P. Geethakumary &
A.G. Pandurangan 55339 (TBGT); Kollam district,
Aryankavu, 02.12.1961, K.N. Subramanian 77355
(BSI); Palakkad district, Dhoni forest-Korakkallu

71
Janeesha & Nampy
Fig. 12.
Henckelia lyrata
(Wight) A.Weber & B.L.Burtt: a. Habit and habitat; b. A portion of the inflorescence; c. Flower; d. Calyx with pistil;
e. Stamens; f. Pistil; g. Immature capsule; h. Seeds (from
K.M. Prabhukumar
7559).

72 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
region, 27.09.2013, K.M. Prabhukumar 7559
(CALI). Tamil Nadu, Ramnad district, Kendiparai
slopes-Ayyanarkoil, 23.09.1971, E. Vajravelu 38715;
Kumbumedu, 14.03.1970, E. Vajravelu 33767;
Yanimutti Rock-Mudaliaruthu, 13.12.1972, E.
Vajravelu 39412 (MH); Virudhunagar district,
Srivilliputhur Wildlife sanctuary, 24.10.2013,
Kabeer & G. Gnanashekaran 130400 (MH).
Conservation status: The species occurs in Kerala and
Tamil Nadu with around five localities in five
districts, with one isolated disjunct locality in the
North. The estimated Extent of Occurrence (EOO)
is c. 7800 km2. The authors were able to study three
localities where they found a total of 26 mature
plants. Of the three populations, two populations
are located in unprotected areas and one in a wildlife
sanctuary. The situation at Aryankavu and
Mudaliaruthu are currently unknown. With view
to the location and small size of the populations a
further decline is likely. Thus, we assess the species
as Endangered (EN), B2ab(iii,iv,v) according to
IUCN (2019) criteria.
Notes: The lyrate, pilose leaves, shorter scapes and
small flowers identify this species in the field. Clarke
(1883) described another variety, Didymocarpus
lyratus var. protractus, differentiating from the typical
variety by its elliptic, narrowly winged lyrate, less
pilose, crenate leaves, 8–12 cm long petioles and
longer peduncles (up to 24 cm long). But most of
these characters are within the limit of H. lyrata (see
description above) and hence var. protractus is
treated here as conspecific.
Henckelia macrostachya (E.Barnes) A.Weber &
B.L.Burtt, Beitr. Biol. Pflanzen 70: 349.
1998[“1997”]; Shaju et al., Taprobanica 5(2): 138.
2013; T.S.Nayar et al., Fl. Pl. Western Ghats 1: 528.
2014. Didymocarpus macrostachya E.Barnes, Bull.
Misc. Inform. Kew 1938: 37. 1938; M.Ahmedullah
& M.P.Nayar, Endemic Pl. Indian Region 1: 143.
1987; Sasidh., Biodivers. Doc. Kerala Part 6: Fl. Pl.
332. 2004; T.S.Nayar et al., Fl. Pl. Kerala 342. 2006.
Lectotype (designated by Vitek et al., 2000): INDIA,
Travancore, high range, on Ottapparai ridge, 5500
ft., 17.09.1935, E. Barnes 1266 (K [K000858197
digital image!]; isolecto K [K000938534,
K000938535 digital images!]). Fig. 13
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 9–12,
basal, usually in whorls of three, sessile to sub-sessile;
laminae obovate-widely spatulate, 16–27 × 6–10
cm, rounded to slightly acute at apex, attenuate at
base, margins crenate, surfaces usually rugose, white
bullate hairy on the upper surface, with dense
golden brown woolly hairs on the younger leaves
and on lower veins, becoming less so with age; veins
7–8 on each side, much branched and conspicuous
beneath. Cymes axillary, 1–2 per plant, 3–6 times
divided, 16–28-flowered; peduncles terete, 30–54
cm long, stout, golden brown, covered with dense
golden brown woolly hairs; bracts two at each
branching point, broadly linear to elliptic, 1.5–4 ×
1–1.9 mm, rounded at apex, basally hairy, green;
pedicels terete, 3–11 mm long, slender, hairy.
Flowers 10–20 × 13–18 mm. Calyx 5-partite, very
deeply divided or basally connate; lobes ovate to
elliptic, unequal, posterior lobe small, 2–2.7 × 0.8–
1.3 mm, others 2.8–4.9 × 1–1.92 mm, slightly acute
to obtuse at apex, margins entire, reddish brown,
glandular hairy on the base, with golden brown
woolly hairs towards the apex. Corolla
zygomorphic, ventricose, two lipped, with white
woolly hairs outside, glabrous inside; tube 8–12 ×
4.3–5.5 mm, pale lilac to white with a yellow blotch
at the throat; lobes very widely obovate with
undulate margin, rounded at apex, white to pale
lilac, upper lip of two lobes, 5.5–6 × 7–7.8 mm,
lower lip of three lobes, 6.8–7.8 × 7.7–8.6 mm.
Stamens 2, adnate to the base of the corolla,
included; filaments 4.5–5 mm long, glabrous,
yellow; anthers reniform, 1.5–2.75 × 0.75–1.16 mm,
milky white to cream, bearded in the region of the
connective. Staminodes 2; filaments 2.4–3.3 mm
long, hyaline, glabrous; antherodes dome shaped,
0.4–0.62 × 0.5–0.7 mm, bearded in the region of
connective. Ovary ovoid, 4–6 × 2.1–2.64 mm, pale
green, with glandular and dense white woolly hairs
all over; style terete, 1.8–3 mm long, green, glabrous;
stigma obliquely subcapitate, 0.65–0.7 × 0.8–0.86
mm, papillate, pale green. Capsules cylindrical, stout,

73
Janeesha & Nampy
Fig. 13.
Henckelia macrostachya
(E.Barnes) A.Weber & B.L.Burtt: a. Habit and habitat; b. Flower; c. Calyx; d. Calyx with pistil; e. Bract;
f. Stamens; g. Staminode; h. Pistil; i. Immature capsule (from
A.P. Janeesha & Santhosh Nampy
137616).

74 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
13–28 × 2.2–2.7 mm, plagiocarpic, dehiscing
loculicidally along the dorsal side, reddish brown,
glandular hairy with persistent stigma and calyx.
Seeds numerous, elliptic, 0.57–0.69 × 0.22–0.29 mm,
slightly acute at apex, truncate at base; testa dark
brown, reticulate.
Flowering & fruiting: Flowering from July to early
September and fruiting from late August to
October.
Habitat: On moist slopes and in rock crevices, in
association with Cyanotis obtusa Trim.
(Commelinaceae), Christella sp. (Thelypteridaceae)
and some grass species.
Distribution: Endemic to the southern Western
Ghats (Fig. 17).
Specimens examined: INDIA, Kerala, Idukki district,
Devikulam, Pallivasal, 01.07.1944, N.R. Yull 236
(CAL); Eravikulam National Park, 22.08.2014, A.P.
Janeesha & Santhosh Nampy 137612; Ibid.,
22.08.2014, S. Syam Radh, Prashob & A.P. Janeesha
134192; Ibid., 19.05.2015, A.P. Janeesha, S. Resmi &
V. Veena 137615; Eravikulam National Park, after
view point, 19.05.2015, A.P. Janeesha & Santhosh
Nampy 137616 (CALI); near Idukki dam,
24.08.1981, V.S. Raju 71173 (MH); Pettimudi,
30.10.2012, Shaju & Shareef, 74028; Rajamalai,
16.09. 1998, S.D. Biju 35857; Ibid., 27.10.1998, S.D.
Biju 38393 (TBGT).
Conservation status: The species is currently known
only from four localities in one district outside
protected areas with an Extent of Occurrence
(EOO) of c. 74 km². The authors observed 58 plants
of different growth stages, of which 41 were mature
at two localities. The species is possibly surviving
as fragmented populations in a narrow range of
distribution, being subjected to habitat alteration/
destruction due to plantation activities and the
growing tourism in those areas. It is therefore
assessed here as Endangered (EN), B1ab(iii,iv,v),
D (IUCN, 2019).
Notes: This is a robust plant with spatulate leaves
and long scapes (about 50 cm long). The scapes,
calyx and capsules are completely covered with
golden brown woolly hairs.
Henckelia meeboldii (W.W.Sm. & Ramaswami)
A.Weber & B.L.Burtt, Beitr. Biol. Pflanzen 70: 349.
1998[“1997”]; T.S.Nayar et al., Fl Pl. Western Ghats
1: 528. 2014; Janeesha & Nampy, Phytotaxa 268(1):
80. 2016. Didymocarpus meeboldii W.W.Sm. &
Ramaswami, Rec. Bot. Surv. India 6: 43. 1914;
Gamble, Fl. Madras 2: 989. 1924; A.N.Henry et al.,
Fl. Tamil Nadu Ind., Ser I: Analysis 2: 132. 1987;
M.Ahmedullah & M.P.Nayar, Endemic Pl. Indian
Region 1: 143. 1987; Sasidh., Biodivers. Doc. Kerala
Part 6: Fl. Pl. 332. 2004; T.S.Nayar et al., Fl. Pl.
Kerala 342. 2006. Lectotype (designated by Janeesha
& Nampy, 2016): INDIA, Kerala, Peermade Ghat,
s.d., Meebold 12851 (CAL [CAL0000019179!];
isolecto CAL [CAL0000019180!, CAL00000
19181!, CAL0000019182!], E [E00155177!], K
[K000858198 digital image!). Fig. 14
Perennials, rosette-forming hairy herbs with
rootstocks. Roots adventitious, thin. Leaves 8–13,
basal, decussate; petioles 4–12 cm long, pubescent,
winged; wings 2–3 mm broad; laminae widely
elliptic-ovate, 6–16 × 5–11.5 cm, obtuse to acute
at apex, attenuate at base, margins crenate,
sometimes slightly doubly crenate, ultimate
segments regular, surfaces usually rugose, pubescent
on both surfaces, but more so on the younger leaves
and on lower veins, becoming less so with age; veins
5–7 on each side, much branched and conspicuous
beneath. Cymes axillary, 1–5 per plant, 6–9 times
divided, 10–18-flowered; peduncles terete, 10–26
cm long, reddish brown, glandular hairy; bracts two
at each branching point, linear, 3–4 × 0.8–1 mm,
acute at apex, margins entire, pilose; pedicels terete,
4–25 mm long, slender, glandular hairy. Flowers
13–15 × 11–13 mm. Calyx 5-partite, very deeply
divided or basally connate; lobes ovate to lanceolate,
unequal, posterior lobe small, 2–3.1 × 0.7–0.92 mm,
others 3.2–5 × 1–1.8 mm, slightly acute at apex,
margins entire, reddish brown, glandular hairy.
Corolla zygomorphic, ventricose, two lipped,
glandular hairy outside, glabrous inside; tube 9–11
× 3.7–4.2 mm, dark lilac with a yellow patch in the

75
Janeesha & Nampy
Fig. 14.
Henckelia meeboldii
(W.W.Sm. & Ramaswami) A.Weber & B.L.Burtt: a. Habit and habitat; b. A portion of the inflorescence; c. Flower; d. Calyx;
e. Calyx with pistil; f. Bract; g. Stamens; h. Staminode; i. Pistil; j. Immature capsule; k. Seeds (from
A.P. Janeesha & K.M. Manudev
134202).

76 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
throat; lobes ovate to slightly elliptic, rounded at
apex, dark lilac, upper lip of two lobes, 3.7–4 × 5–
5.8 mm, lower lip of three lobes, 6–7 × 5–6 mm.
Stamens 2, adnate to the base of corolla, included;
filaments 4–6 mm long, puberulent, green on the
top and colourless towards base; anthers reniform,
1.5–1.7 × 0.9–0.95 mm, milky white, glabrous.
Staminodes 2; filaments 2.2–2.5 mm long, hyaline,
glabrous; antherodes 0.42 × 0.36 mm, hyaline,
glabrous. Ovary ovoid to oblong, 4.8–6.5 × 0.9–
1.2 mm, green, glandular hairy, sometimes slightly
curved; style terete, 4–5.5 mm long, green,
glabrous; stigma obliquely subcapitate, 0.3–0.35 ×
0.2–0.25 mm, papillate, pale green. Capsules
cylindrical, straight or slightly curved, 14–35 × 1.3–
1.5 mm, plagiocarpic, dehiscing loculicidally along
the dorsal side, reddish brown, tomentulose with
persistent stigma. Seeds numerous, elliptic, 0.38–
0.45 × 0.2–0.22 mm, obtuse to slightly acute at apex,
truncate at base; testa dark brown, reticulate.
Flowering & fruiting: Flowering from July to
October and fruiting from September to
November.
Habitat: On moist slopes, in association with
Adiantum philippenes L. (Pteridaceae), Christella sp.
(Thelypteridaceae), Selaginella sp. (Selaginellaceae)
and Cyanotis villosa Schult.f. (Commelinaceae).
Distribution: Endemic to the southern Western
Ghats (Fig. 17).
Specimens examined: INDIA, Kerala, Ernakulam
district, Shoolamudi, 11.07.2016, A.P. Janeesha,
Manu Philip & K. Shinoj 137692 (CALI); Idukki
district, Mannavan shola, 24.05.2015, K.M.
Manudev, S. Syam Radh & A.P. Janeesha 137619;
Munnar, 10.09.2013, A.P. Janeesha & Santhosh
Nampy 134214; near Kuttikanam-Kumali road,
30.06.2016, A.P. Janeesha & P.S. Sreeja 137662;
Neryamangalam to Adimali route, 25.07.2013, A.P.
Janeesha & K.M. Manudev 134202; Onnamthode-
Mathikettanshola National Park, 08.05.2015, S.
Syam Radh & A.P. Janeesha 137613 (CALI);
Peerumedu, 31.08.1992, A. Nazarudeen 14386; Ibid.,
12.09.2009, Geethakumary 58596 (TBGT);
Pothumade-Munnar, 25.07.2013, A.P. Janeesha &
K.M. Manudev 134203; Pynavu, 29.06.2016, P.S.
Sreeja & A.P. Janeesha 137667 (CALI); Upputhara,
30.09.1981, C.N. Mohanan 72086 (MH); Kottayam
district, near Ottayitti, 26.06.2016, P.S. Sreeja & A.P.
Janeesha 137661; Vagamon churam, 07.08.2014,
A.P. Janeesha & A.P. Roshan 134273, 134274,
134275; Ibid., 03.10.2015, C. Pramod & A.P. Janeesha
137645; Vagamon churam-200 mtr after view
point, 07.08.2014, A.P. Janeesha, A.P. Roshan &
Santhosh Nampy 134276 (CALI). Tamil Nadu,
Madura district, Palani-Kodaikanal ghat road,
27.11.1985, K.M. Matthew & N. Rajendran 43308;
Tiger shola-acroos to Korappar, 01.08.1986, K.M.
Matthew & M. Charles 46094 (RHT).
Conservation status: This species is widely distributed
in central areas of Kerala and some parts in western
Tamil Nadu, with an Extent of Occurrence (EOO)
of c. 3800 km². More than 240 mature plants were
observed from nine populations across Kerala. The
estimated Area of Occupancy (AOO) for all
localities is an estimated 48 km². The majority of
the localities are not in protected areas and are
subjected to human disturbance due to plantation
activities and tourism. Because if the small number
of mature plants we consider the species to be of
Endangered (EN) D, according to IUCN (2019)
criteria.
Notes: Ramaswamy (1914) described the corolla
tube and filaments as glabrous. But in the present
investigation, the outside of the corolla tube was
found to be glandular hairy and the filaments
bearded. The species is morphologically close to
H. humboldtiana, but can be distinguished by its
broadly elliptic to ovate leaves, glandular
indumentum on the pedicel, calyx lobes, corolla
tube and ovary.
Henckelia missionis (Wall. ex R.Br.) A.Weber &
B.L.Burtt, Beitr. Biol. Pflanzen 70: 350.
1998[“1997”], nom. cons. prop.; T.S.Nayar et al., Fl.
Pl. Western Ghats 1: 528. 2014. Didymocarpus
missionis Wall. ex R.Br., Pl. Jav. Rar. 119. 1839;
Rama Rao, Fl. Pl. Travancore 295. 1914; Gamble,
Fl. Madras 2: 988. 1924; A.N.Henry & Swamin., J.
Bombay Nat. Hist. Soc. 76: 373. 1977; A.N.Henry

77
Janeesha & Nampy
Flowering & fruiting: Flowering from May to July
and fruiting from July to September.
Habitat: On moist slopes and in rock crevices.
Distribution: Endemic to the southern Western
Ghats (Fig. 17).
Specimen examined: INDIA, Tamil Nadu,
Kanyakumari district, Kalikesam river side,
Balamore, 700 m, 31.08.1976, A.N. Henry 48151
(MH).
Conservation status: The population status at the type
locality (A.N. Henry 49421) is presently uncertain,
since no plants were found during recent fieldwork
in 2014. The situation of the nearby population at
Kalikesam river side at Balamore (A.N. Henry
48151) is presently unknown. Because of these
uncertainties, the species should be regarded as Data
Deficient (DD) according to IUCN guidelines
(IUCN, 2019).
Typification: Henckelia missionis (Wall. ex R.Br.)
A.Weber & B.L.Burtt (Gesneriaceae) was first
named (nom. nud.) as “Didymocarpus ? missionis” by
Wallich (1829). Subsequently Brown (1839)
validated the preceding name with a description
(“Caule brevissimo, pedunculis axillaribus
scapiformibus, folis cordato ovatis”) and as type, he
referred to “Wall. List. no. 639”. Kuntze (1891)
transferred this species to Roettlera as R. missionis
(Wall. ex R.Br.) Kuntze while Weber and Burtt
(1998[“1997”]) placed this species in Henckelia as
H. missionis (Wall. ex R.Br.) A.Weber & B.L.Burtt.
Meanwhile, Beddome (1874: 39, t. 176) described
another species, viz., Didymocarpus membranaceus
Bedd. from Pachemallai hills (Tamil Nadu) in South
India as “leaves all radical, ... membranaceous, from
obliquely cordate-ovate to cordate-lanceolate,
corol-tube subinfundibuliform, ......”. Clarke (1883)
synonymised Beddome’s species under D. missionis
and this treatment was followed by subsequent
authors (Clarke, 1885[“1884”]; Gamble, 1924;
Henry et al., 1987). This species is endemic and
very rare in South India (Nayar & Sastry, 1990;
Walter & Gillet, 1997), and has been re-collected
et al., Fl. Tamil Nadu Ind., Ser I: Analysis 2: 132.
1987; M.Ahmedullah & M.P.Nayar, Endemic Pl.
Indian Region 1: 143. 1987; M.P.Nayar & Sastry,
Red Data Book Indian Pl. 3: 157. 1990; K.S.Walter
& H.J.Gillet, IUCN Red List Threatened Pl. 291.
1997; Gopalan & A.N.Henry, Endemic Pl. India
114. 2000; T.S.Nayar et al., Fl. Pl. Kerala 342. 2006.
Roettlera missionis (Wall. ex R.Br.) Kuntze, Revis.
Gen. Pl. 2: 476. 1891. Type (cons. prop.): INDIA,
Tamil Nadu, Kanyakumari district, Kilaviarumalai-
Balamore, 480 m, 28.07.1977, A.N. Henry 49421
(MH!).
Henckelia membranacea (Bedd.) Janeesha & Nampy
comb. nov. Didymocarpus membranacea Bedd., Icon.
Pl. Ind. Or. 1: 39. 1874. Lectotype (designated here):
Beddome’s Icon Pl. Ind. Or. 1: t. 176. 1874. Figs. 15 & 16
Perennials, hairy herbs with rootstocks. Roots
adventitious, thin. Stems reduced. Leaves 6–10,
basal, decussate; petioles 11–16 cm long, pubescent,
not winged; laminae ovate to elliptic, 7.9–12.1 ×
5.2–7.3 cm, membranaceous, acute to acuminate
at apex, obliquely cordate at base, margins entire,
with white pellucid hairs, strigose on the upper
surface, surfaces rough; veins usually 6 on each side,
much branched and conspicuous beneath. Cymes
axillary, 1–5 per plant, 3–5 times divided, 6–10-
flowered; bracts two, linear, 4 mm long; peduncles
terete, 20–29 cm long, with scaly hairs towards
apex; pedicels terete, 6–9 mm long, slender,
pubescent. Flowers 20–25 mm long. Calyx 5-
partite, very deeply divided or basally connate; lobes
linear to lanceolate, 5–8 mm long, acute at apex,
margins entire, pilose. Corolla zygomorphic, sub-
infundibuliform, upper lip of two lobes, lower lip
of three lobes, glabrous; tube pale lilac with a yellow
patch in the throat; lobes pale lilac, ovate, rounded
at apex. Stamens 2, adnate to the base of corolla,
included, glabrous; anthers reniform, bearded.
Staminodes 2; antherodes present. Ovary ovoid;
style terete, glabrous; stigma obliquely subcapitate.
Capsules cylindrical, 27–33 mm long, plagiocarpic,
dehiscing loculicidally along the dorsal side, with
persistent stigma. Seeds not seen.

78 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Fig. 16.
Type (conserved) of
Henckelia missionis
(Wall. ex R.Br.)
A.Weber & B.L.Burtt. © Botanical Survey of India, Kolkata, Reproduced
with permission.
Fig. 15.
Henckelia missionis
(Wall. ex R.Br.) A.Weber & B.L.Burtt (from
A.N. Henry
49421, drawn by A.P. Janeesha).
only twice from Kanyakumari district in Tamil
Nadu (Madras Herbarium: Henry 48151, 49421).
While examining “Wallich List. No. 639” (as cited
by Brown l.c.), it was found that Brown’s (l.c.)
citation refers to Arenaria globiflora Wallich (1829:
19), belonging to Caryophyllaceae. It was found
that Wallich’s Didymocarpus missionis is more likely
referable to “Wall. List. No. 6396” (probably a
typographical error by Brown). According to
Stafleu and Cowan (1988), Wallich’s main
collections are kept at K (K-W). After a thorough
search in various herbaria, two sheets of “Wall. No.
6396” were found, one each at BM and K. The
lone sheet in BM (BM000617800 digital image!)
has two flower buds separately mounted and carries
two labels with annotations. The top one, probably
the original label reads “Didymocarpus missionis
Wall. Cat 6396 Herb Madras” and the second one,
made by curators at BM “Wallich 6396: TYPE
SPECIMEN Didymocarpus Wall. ex R.Br.,
Cyrtandreae: 119 (1839)” respectively. The flower
buds are not, however, unquestionably referable to
H. missionis. The sheet at K (K001123777 digital
image!) has a single specimen with three scapes and
a few leaves. There is an inscription on the sheet,
possibly by C.B. Clarke written in pencil:
‘Didymocarpus missionis Wall. C.B.Clarke (= D.
membranaceus Beddome Ic. t. 176)’. This sheet also
carries a Wallich label on which is written: “6396
Didymocarpus ? missionis Wall (caet. spec. 777-793).
Hb. Madras”. However, the ovate leaves with
cordate base, long pedicels, infundibuliform corolla,
and persistent calyx undoubtedly indicate that the
specimen belongs to Jerdonia indica Wight (Wight,
1848: 10, t. 1352) (Gesneriaceae). The original
description by Brown (1839) is applicable equally
to J. indica also. Since the type specimens of H.
missionis is demonstrably ambiguous and doesn’t

79
Janeesha & Nampy
Fig. 17. Distribution of
Henckelia lyrata
(Wight) A.Weber & B.L.Burtt,
H. macrostachya
(E.Barnes) A.Weber & B.L.Burtt,
H. meeboldii
(W.W.Sm.
& Ramaswami) A.Weber & B.L.Burtt and
H. missionis
(Wall. ex R.Br.) A.Weber & B.L.Burtt in South India (drawn using QGIS v.3.12.1).
permit a precise identification and application of
the name, the name is illegitimate (Art. 52.1) and a
proposal is under preparation to conserve the name
with a conserved type according to Articles 14.1
and 14.9 of ICN (Turland et al., 2018).
Beddome (1874) described Didymocarpus
membranacea Bedd. based on his specimens from
“Pachemallay (2000 feet elevation), in South
Travancore ghats”, India. According to TL-2
(Stafleu & Cowan, 1976), the herbarium and type
materials of R.H. Beddome are mainly at BM, some
perhaps at K and CAL. Even after extensive search
no relevant materials have been found in these
herbaria. Hence the illustration in the protologue
(Bedd., Icon Pl. Ind. Or. 1: t. 176. 1874) is selected
here as the lectotype, according to articles 9.3 and
9.4 of ICN (Turland et al., 2018).
Henckelia ovalifolia (Wight) A.Weber &
B.L.Burtt, Beitr. Biol. Pflanzen 70: 351.
1998[“1997”]; T.S.Nayar et al., Fl. Pl. Western
Ghats 1: 529. 2014; Janeesha & Nampy, Phytotaxa
268(1): 84. 2016. Didymocarpus ovalifolius Wight,
Icon. Pl. Ind. Orient. 4: 24. t. 1351. 1848, Ill. Ind.
Bot. 2: 131. t. 142, fig. 6. 1850; C.B.Clarke in
Hook.f., Fl. Brit. India 4: 354. 1885[“1884”]; Rama
Rao, Fl. Pl. Travancore 295. 1914; Gamble, Fl.
Madras 2: 988. 1924; M. Ahmedullah & M.P.Nayar,
Endemic Pl. Indian Region 1: 143. 1987;
A.N.Henry et al., Fl. Tamil Nadu Ind., Ser I:
Analysis 2: 132. 1987; Manilal, Fl. Silent Valley
200. 1988; M.Mohanan & A.N.Henry, Fl.
Thiruvananthapuram Dist. 338. 1994; Gopalan &
A.N.Henry, Endemic Pl. India 117. 2000;
M.Mohanan & Sivad., Fl. Agasthyamala 495. 2002;
Sasidh., Biodivers. Doc. Kerala part 6: Fl. Pl. 332.
2004; T.S. Nayar et al., Fl. Pl. Kerala 342. 2006.
Roettlera ovalifolia (Wight) Kuntze, Revis. Gen. Pl.
2: 476. 1891. Lectotype (designated by Janeesha &
Nampy, 2016): INDIA, Tamil Nadu, Courtallam,
08.1835, Wight 558 (E [E00155178 digital image!];

80 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
isolecto E [E00179262 digital image!], K
[K000858199 digital image!]). Fig. 18
Perennials, rosette-forming herbs with rootstock.
Roots adventitious. Stem reduced. Leaves 8–18,
decussate; petioles 4–14 cm long, hairy throughout,
not winged; laminae usually ovate-broadly ovate,
rarely orbicular, 3.3–12.8 × 2.5–7.7 cm, slightly
acute to obtuse at apex, unequal at base, margins
crenate to serrate, white strigose or tomentose above
and on the lower veins; veins 7–10 on each side,
much branched and conspicuous beneath. Cymes
axillary, 2–5 per plant, 2–3 times divided, 6–8-
flowered; bracts two, oblanceolate, 6.38 × 1.34 mm,
white tomentose hairy out; peduncles terete, 9–22.5
cm long, green-pale reddish brown, hairy
throughout; pedicels terete, 5–10 mm long, narrow,
hairy. Flowers 12–20 × 8–15 mm. Calyx 5-partite,
very deeply divided or basally connate; lobes linear
to lanceolate, unequal, posterior lobe small, 4.2–
5.1 × 0.8–1 mm, others 5.8–6.8 × 1.1–1.2 mm,
slightly acute at apex, margins entire, reddish
brown, white tomentose outside. Corolla
zygomorphic, ventricose, two lipped, white hirsute
outside, glabrous inside; tube 7.2–11.3 × 5–5.8 mm,
very pale violet to brilliant purple, deep purple at
the throat and a yellow blotch at the base of the
tube; lobes very widely ovate, slightly acute to
obtuse at apex, violet to brilliant purple, upper lip
of two lobes, 6.5–7.2 × 5.8–6.6 mm, lower lip of
three lobes, 8–8.4 × 7.1–7.25 mm. Stamens 2, adnate
to the base of the corolla, included; filaments 5.3–
5.7 mm long, puberulent towards the apex, light
yellowish green towards the top and milky white
towards the base; anthers reniform, 1.7–1.9 × 0.65–
0.8 mm, yellowish brown, glabrous. Staminodes 2;
filaments 3.5–3.7 mm long, hyaline, glabrous;
antherodes not prominent. Ovary narrowly ovoid
to cylindrical, tapering to a style, 4.9–5 × 1.1–1.21
mm, green towards the base and reddish brown
towards the apex, hairy; style terete, 4–4.1 × 0.35–
0.40 mm long, hyaline, hairy throughout up to the
middle; stigma obliquely subcapitate, 0.68 × 0.38
mm, papillate, milky white. Capsules linear,
straight, 36–44 × 2–2.2 mm, plagiocarpic, dehiscing
loculicidally along the dorsal side, reddish brown,
hairy with persistent stigma and calyx. Seeds
numerous, elliptic, 0.49–0.59 × 0.21–0.24 mm,
acute at apex, truncate at base; testa dark brown,
reticulate.
Flowering & fruiting: Flowering from June to
September and fruiting from August to October
(up to February).
Habitat: In moist shady areas of dense evergreen
forest, in humus rich soil or on fallen, moss covered
tree trunks. Grows in association with Impatiens
travancorica Bedd. (Balsaminaceae), Christella sp.
(Thelypteridaceae), Osbeckia sp. (Melastomataceae)
and some grass species.
Distribution: Endemic to the southern Western
Ghats (Fig. 17).
Specimens examined: INDIA, Kerala, Idukki district,
Kallar Reserve Forest, 16.07.1989, s.coll. 90687
(MH); Periyar–Koruthode, 25.09.1996, Jomy
Augustine 16811 (CALI); Kollam district,
Thenmala, 15.06.1984, N. Mohanan 165 (TBGT);
Palakkad district, Nillikkal South, 08.06.1982,
Prasannan 10392 (CALI); Thiruvananthapuram
district, Agasthyamala, 24.06.1993, N. Mohanan
11454 (CALI, TBGT); Ibid., 19.10.1993, G.
Gangaprasad & S. William Decruse 18419 (TBGT);
Ibid., 06.06.2014, K.M. Manudev & A.P. Janeesha
138919, 138925; Ibid., 21.10.2016, K. Shinoj & A.P.
Janeesha 137658 (CALI); Pongalapara, 25.08.1990,
N. Mohanan 10044 (TBGT); Ponmudi, 10.10.2014,
K.P. Vimal, M.G. Prasad & A.P. Janeesha 137602
(CALI); Western slopes of Agasthyamala,
06.10.1973, J. Joseph 44608 (MH); Wayanad district,
Gurukula botanical sanctuary (cultivated),
25.11.2014, A.P. Janeesha & S. Resmi 137609; Periya,
25.08.1984, R.T. Balakrishnan 40450; Thalapaya,
10.02.1986, R.T. Balakrishnan 41944 (CALI). Tamil
Nadu, Kanyakumari district, Muthukuzhivayal,
06.08.1977, A.N. Henry 49610 (MH); Ibid.,
01.09.1981, K.M. Matthew 17788 (RHT);
Tirunelveli district, Agasthyamala, 22.06.1901, C.A.
Barber 2918; s.loc., 1807, R.H. Beddome 164 (MH).

81
Janeesha & Nampy
Fig. 18.
Henckelia ovalifolia
(Wight) A.Weber & B.L.Burtt: a. Plant in natural habitat; b. Habit; c. A portion of the inflorescence; d. Flower;
e. Calyx; f. Calyx with pistil; g. Bract; h. Stamens; i. Staminode; j.j.
j.j.
j. Pistil; k. Immature capsule; l. Seeds (a from
K.P Vimal, M.G. Prasad & A.P.
Janeesha
137602; b-l from
K.M. Manudev & A.P. Janeesha
138919).

82 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Conservation status: This species is known from eight
locations in seven districts in Kerala and Tamil
Nadu with a narrow South-North distribution. The
estimated Extent of Occurrence (EOO) is c. 1100
km² and Area of Occupancy (AOO) is 32 km². The
authors surveyed three locations and found a total
of only 81 mature plants. Since these locations are
inside protected areas, the chances for a decline due
to anthropogenic activity is small. The situation of
just under half of the localities is known. However,
when extrapolating the findings on mature plants
to the entire distribution range, it may justify
categorisation of the species as Endangered (EN),
D (IUCN, 2019).
Notes: Henckelia ovalifolia and H. missionis are similar
in having simple ovate, long petioled leaves, large
flowers and long fruits. However, H. ovalifolia can
be identified by its leaves with serrate margins and
strigose hairs, scapes hairy throughout, hirsute
corolla, puberulent filaments and a hairy style.
Henckelia pradeepiana Nampy, Manudev &
A.Weber., Rheedea 22(2): 119. 2012; T.S.Nayar
et al., Fl. Pl. Western Ghats 1: 529. 2014.
Type: INDIA, Kerala, Kozhikode district,
Muthappanpuzha, N 11°26.6992', E 076°05.2882',
470 m, 19.08.2010, Santhosh Nampy & K.M. Manudev
3102 (holo CALI!; iso DEV!, W!). Fig. 19
Perennating tuberous herbs. Tubers 1–2.5 cm
diam., discoid. Leaves usually 1, rarely up to 3;
petioles 1.8–5.6 cm long, sparsely glandular hairy,
not winged; laminae broadly ovate or elliptic,
oblong-ovate, 9–16 × 5.5–9 cm, acute at apex,
slightly oblique or cordate at base, margins entire
to serrulate, glabrous above, glandular hairy below,
membranaceous, glandular hairy on the margins,
surfaces smooth; veins usually 7 on each side, much
branched and conspicuous beneath. Cymes axillary,
1–3 per plant, 1–6 times divided, 2–19-flowered,
glandular hairy; peduncles terete, 5–20 cm long,
green; bracts two at each branching point,
lanceolate to triangular, 4.8–6.1 × 1.2–1.7 mm,
acute at apex, margin entire, slightly hairy; pedicels
terete, 1–3.8 cm long, slender, glandular hairy.
Flowers c. 10 × 7 mm. Calyx 5-partite, basally
connate; lobes ovate-lanceolate, unequal, posterior
lobe small, 3–4.1 × 1–1.25 mm, others 4.3–7 × 1.3–
2.1 mm, keeled, blunt at apex, margins entire,
green, glabrous. Corolla campanulate, two lipped,
glabrous inside and outside (but with a few hairs at
the corolla mouth); tube 4–5.4 × 5.2–5.6 mm, lilac
to pale lilac with a yellow patch in the throat, with
7–10 purplish streaks; lobes ovate to rounded at
apex, lilac to pale lilac, upper lip of two lobes, 2–
3.1 × 3.4–4.51 mm, lower lip of three lobes, 3–3.8
× 3.6–4.3 mm. Stamens 2, adnate to the base of
corolla, included; filaments 2.4–3.3 mm long,
glabrous, greenish yellow; anthers reniform, 1.3–
1.5 × 0.53–0.73 mm, milky white, glabrous.
Staminodes 2; filaments 1–1.2 mm long, hyaline,
glabrous; antherodes ovate to oblate, c. 0.31 × 0.3
mm. Ovary ovoid, 2–2.4 × 1.35–1.75 mm, green,
glabrous; style terete, 4–5.22 mm long, green,
glabrous; stigma subpeltate, lower lip strongly
expanded and slightly emarginate at apex, 1.7–2.1
× 0.5–0.87 mm, papillate, yellow to yellowish green.
Capsules ovoid to subglobose, 5.6–6.3 × 5–6 mm,
plagiocarpic, dehiscing by a longitudinal slit on the
upper side, greenish brown, with persistent calyx
and a prominent beak. Seeds numerous, elliptic,
0.4–0.52 × 0.15–0.2 mm, acute at both ends; testa
dark brown, reticulate.
Flowering & fruiting: Flowering from July to August
and fruiting from August to September.
Habitat: On moist rocks and mountain slopes, found
in association with mosses and ferns.
Distribution: Endemic to the southern Western
Ghats (Fig. 20).
Specimens examined: INDIA, Kerala, Kozhikode
district, Muthappanpuzha, N 11°26.6992', E
76°05.2882', 470 m, 18.07.2010, K.M. Manudev
3022 (DEV); Ibid., 19.08.2010, A.K. Pradeep 90089
(CALI); Ibid., 19.08.2010, K.M. Manudev & Santhosh
Nampy 3105; Ibid., 17.09.2011, Santhosh Nampy &
K.M. Manudev 4547 (DEV); Ibid., 12.08.2014, A.P.
Janeesha & Santhosh Nampy 134287, 134288; Ibid.,
15.06.2015, A.P. Janeesha & S. Resmi 137624;
Muthappanpuzha-Kundanthode, 15.06.2015, A.P.

83
Janeesha & Nampy
Fig. 19.
Henckelia pradeepiana
Nampy, Manudev & A.Weber: a. Habit and habitat; b. A portion of the inflorescence; c. Flower; d. Calyx; e. Calyx
with pistil; f. Bract; g. Stamens; h. Staminode; i. Pistil; j. Dehiscing capsule; k. Seeds (from
A.P. Janeesha & Santhosh Nampy
134287).

84 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Janeesha & S. Resmi 137623; way to Vellarimala-
Olichuchattam, N 11°25.7842', E 76°05.1732',
1160 m, 19.09.1997, A.K. Pradeep 56009 (CALI).
Conservation status: The species is currently known
only from the type locality which is an unprotected
area, with two subpopulations separated by a
distance of 2 kilometres. The locality has a total of
79 mature plants, with the possibility of future
disturbances due to human activities since the area
is near a stream which is used by local people for
their daily needs. The Area of Occupancy (AOO)
is less than 10 km². The species may be categorized
as Critically endangered (CR), B2ab(ii,iii,iv,v), D
(IUCN, 2019).
Notes: This species has several unusual features not
found in any other Henckelia species in South India
or the genus as a whole. This is the only species of
Henckelia having tubers to enable the plants to
overcome unfavourable conditions or periods. The
plants usually have only one leaf, but sometimes
up to 3 arranged alternately. The stigma is unusual
for South Indian Henckelia in that they are chiritoid,
with the upper lobe reduced and the lower one
undivided and linguate (Weber et al., 2011;
Manudev et al., 2012). The capsules are globose in
contrast to the linear to cylindrical ones in other
South Indian Henckelia species or any other species
in the genus, but are plagiocarpic and dehisce
similar to other South Indian Henckelia. They are
reminiscent of those found in species of Loxocarpus,
another genus previously included in Henckelia
(Weber & Burtt, 1998[“1997”]) but recently
resurrected (Weber et al., 2011; Middleton et al.,
2013).
Henckelia repens (Bedd.) A.Weber & B.L.Burtt,
Beitr. Biol. Pflanzen 70: 354. 1998[“1997”];
T.S.Nayar et al., Fl. Pl. Western Ghats 1: 529. 2014.
Didymocarpus repens Bedd., Icon. Pl. Ind. Or. 1: 24.
t. 120. 1874; C.B.Clarke in Hook.f., Fl. Brit. India
4: 354. 1885[“1884”]; Rama Rao, Fl. Pl. Travancore
Fig. 20.
Distribution of
Henckelia ovalifolia
(Wight) A.Weber & B.L.Burtt and
H. pradeepiana
Nampy, Manudev & A.Weber in South India (drawn
using QGIS v.3.12.1).

85
Janeesha & Nampy
295. 1914; Gamble, Fl. Madras 2: 987. 1924;
M.Ahmedullah & M.P.Nayar, Endemic Pl. Indian
Region 1: 143. 1987; A.N.Henry et al., Fl. Tamil
Nadu Ind., Ser I: Analysis 2: 133. 1987; M.Mohanan
& A.N.Henry, Fl. Thiruvananthapuram Dist. 338.
1994; Gopalan & A.N.Henry, Endemic Pl. India
121. 2000; M.Mohanan & Sivad., Fl. Agasthyamala
496. 2002; Sasidh., Biodivers. Doc. Kerala Part 6:
Fl. Pl. 332. 2004; T.S.Nayar et al., Fl. Pl. Kerala
342. 2006. Rottlera repens (Bedd.) Kuntze, Revis.
Gen. Pl. 2: 477. 1891. Lectotype (designated by Vitek
et al., 2000): INDIA, Tinnevelly and Travancore
hills, 3000 feet elevation, 09.1868, Beddome 46 (K
[K000858200 digital image!]; isolecto MH [36684!,
36680!]). Fig. 21
Perennial creeping herbs, rooting at nodes and also
along the internodes, hairy throughout. Roots
adventitious. Stems slender, terete, hairy; internodes
6–12 cm long. Leaves two per node, one is highly
reduced to a scale; petioles 3.5–18 cm long, not
winged; laminae broadly ovate to sub-orbicular,
5.2–10 × 5–8.3 cm, acute at apex, cordate at base,
margins doubly crenate, hairy; veins usually 5–6
on each side, much branched and conspicuous
beneath. Cymes few-flowered, usually one from
each node, 1–3 times divided, 2–4-flowered;
peduncles 4–14.5 cm long, terete, green-pale
reddish brown, hairy throughout; pedicels 5–10
mm long, terete, narrow, glandular hairy. Flowers
12–20 × 8–15 mm. Calyx 5-partite, very deeply
divided or basally connate; lobes broadly lanceolate,
slightly acute at apex, reddish brown, margins
entire, hairy, unequal, posterior one small, 3.4–3.7
× 0.7–0.83 mm, others 3.8–4.2 × 0.83–1 mm.
Corolla zygomorphic, ventricose, two lipped,
glandular hairy outside, glabrous inside; tube 12–
13 × 4.5–5.4 mm, dark to pale lilac with a yellow
blotch at the throat; lobes elliptic to oblong,
rounded at apex, dark to pale lilac, upper lip of two
lobes, 4.1–4.9 × 4.7–5.2 mm, lower lip of three
lobes, 5–6.65 × 4.3–5.47 mm. Stamens 2, adnate to
the base of the corolla, included; filaments 7.5–8
mm long, broad and hairy at the middle, not
geniculate, yellow at the middle, other parts milky
white; anthers reniform, 2–2.1 × 0.5–0.7 mm,
purple coloured except the cohering region of the
lobes and also the region surrounding the
connective. Staminodes 2; filaments 3.6–4.1 mm
long, hyaline, hairy towards the middle; antherodes
trilobed, 0.42–0.44 × 0.36–0.37 mm, prominent.
Ovary narrowly ovoid, tapering to a style, c. 7.2 ×
1.21 mm, green towards the base and reddish green
towards the distal part, eglandular hairy; style terete,
4–4.4 × 0.5 mm long, hyaline, eglandular hairy
throughout; stigma oblique, subpeltate, 1.7 × 1.23
mm, glabrous. Capsules linear, straight, 20–22 ×
2–2.2 mm, plagiocarpic, dehiscing loculicidally
along the dorsal side, reddish brown, hairy, with
persistent stigma. Seeds numerous, elliptic, 0.49–
0.59 × 0.21–0.24 mm, slightly acute at apex,
truncate at base; testa dark brown, reticulate.
Flowering & fruiting: Flowering from (May) June
to October and fruiting from August to December.
Habitat: Henckelia repens is found as an undercover
in moist mixed evergreen and deciduous forests, in
association with Elatostema sp., Pouzolzia sp. (both
Urticaceae), Adiantum philippenes L. (Pteridaceae),
Christella sp. (Thelypteridaceae), Selaginella sp.
(Selaginellaceae) and some grass species.
Distribution: Endemic to the southern Western
Ghats (Fig. 26).
Specimens examined: INDIA, Kerala, Kollam
district, Kolatoor, s.d., s.coll. 1293 (MH);
Thiruvananthapuram district, Agasthyamala-
Mutththatti, 06.06.2014, K.M. Manudev, A.P.
Janeesha & Santhosh Nampy 138940 (CALI);
Athirumala, 12.08.1988, N. Mohanan 4227 (TBGT,
CALI); Ibid., 07.06.2014, K.M. Manudev, P.G. Arun
Kumar & A.P. Janeesha 138936; Attayar, 13.05.1988,
N. Mohanan 9614; Bonaccord, 27.04.1993, A.
Nazarudeen 17128 (CALI); Ibid., 21.12.1987, N.
Mohanan 9066 (TBGT, CALI); forest near
Bonaccord, 21.08.1975, J. Joseph 46470; Ibid.,
02.10.1973, J. Joseph 44493 (MH); Chemungi, s.d.,
Rajkumar 11775; Ibid., 18.08.1992, E.S. Santhosh
Kumar & M. Abdul Jabbar 14447; Ibid., 11.08.2011,
P.M. Sheeba 66275 (TBGT); Pandimottai,
19.05.1979, M. Mohanan 61853 (MH);
Pidichathamparamal, 31.05.2001, Non Wood Forest

86 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Fig. 21.
Henckelia repens
(Bedd.) A.Weber & B.L.Burtt: a. Habit and habitat; b. Flowering creeping shoot; c. Flower; d. Calyx; e. Calyx with
pistil; f. Stamens; g. Staminode; h. Pistil; i. Stigma; j. Capsule; k. Seeds (a
&
b from
K.M. Manudev, A.P. Janeesha & Santhosh Nampy
138940
;
K.M. Manudev, P.G. Arunkumar & A.P. Janeesha
138936).

87
Janeesha & Nampy
Product’s (NWFPs) Team 46274; Pinarummodu,
25.09.2008, C.G. Vishnu 64727 (TBGT); Ponmudi,
28.11.2012, A.P. Janeesha, P.M. Shahina & K.M.
Manudev 134240; Ibid., 10.10.2014, M.G. Vimal,
Prasad & A.P. Janeesha 137603; s.loc., 21.12.1987,
N. Mohanan 9066 (CALI). Tamil Nadu,
Tirunelveli district, Ambasamudram to
Kannikkatti, 31.12.1977, K.M. Matthew 15627; Ibid.,
01.01.1978, K.M. Matthew 15796 (RHT); Chittar
river-way to Agasthyamalai, 22.08.1963, A.N.
Henry 16971; Kannikkatti, 12.06.1899, C.A. Barber
396; Kannikkatti to Kalivayalpil, 31.05.1901, C.A.
Barber 3002; Papanasam hills, 1867, R.H. Beddome
s.n.; Valayar forest, 11.07.1976, P. Bhargavan 47445;
way to Ainthuthalai, 14.10.1989, R. Gopalan 90662;
way to Pothgai malai, 16.10.1989, R. Gopalan
91605; way to Kalivarapil, 23.01.1991, R. Gopalan
94658; s.loc., 1877, R.H. Beddome s.n. (MH) ); s.loc.,
s.d., R.H. Beddome 5008, 5805 (BM digital image).
Conservation status: The species is distributed in 4
districts in southern Kerala and southern Tami
Nadu and where seven locations are known. Only
five of these fall in protected areas. The Extent of
Occurrence (EOO) is calculated to be c. 600 km2
and the Area of Occupancy (AOO) is 28 km2.
During field studies, the authors observed a total of
only 52 mature plants at four locations. This might
indicate that the species occurs with very few
individuals in total and should belong in the
Endangered category (EN), D (IUCN, 2019).
Notes: Henckelia repens is the only creeping species
in South India while the other species, H. burttii
D.J.Middleton & Mich.Möller, with a very similar
habit is found in Northeast India (Burtt & Panigrahi,
1965; Wood, 1974). Henckelia repens has a longer
corolla tube (up to 1.5) compared to other South
Indian species. Beddome’s illustration (Icon. Pl. Ind.
Or. 1: 24. t. 120. 1874) and drawings on the type
sheet at K differ with the protologue in two points.
The anthers are cohering in the specimens but they
are drawn in the icon and on the illustration on the
type sheet as non-coherent. Similarly, the filament
is broad except at the apex, but it is drawn as narrow
along its entire length in the type sheet and broad
at the basal region in the icon.
Henckelia wayanadensis Janeesha & Nampy, sp.
nov. Figs. 22 & 23a1-i1
Henckelia wayanadensis is morphologically similar to H.
fischeri, but can be readily identified by its elliptic to widely
elliptic leaves (vs. ovate), ovate hispid bracts (vs. lanceolate
tomentose bracts), small flowers 12–14 × 6.7–11 mm (vs.
large flowers 11–17 × 9–22 mm), and purple glands on the
ovary (vs. hyaline glandular hairs on the ovary).
Type: INDIA, Kerala, Wayanad district,
Kaduvakuzhi hill top, 1200 m, 08.07.2016, A.P.
Janeesha & Santhosh Nampy 137672 (holo CALI!; iso
MH!)
Perennials, rosette-forming, herbs with rootstocks.
Roots adventitious, thin. Leaves 8–20, basal, usually
in whorls of four; petioles 4–11.5 cm long,
pubescent, winged; wings 3–8 mm broad; laminae
elliptic to widely elliptic, 8–19.2 × 6–12 cm, obtuse
to slightly acute at apex, attenuate at base, margins
doubly crenate, ultimate segments irregular, first
crenation very deep, surfaces usually rugose, white
tomentose on both surfaces, becoming less so with
age; veins 6–8 on each side, much branched and
conspicuous beneath. Cymes axillary, 4–10 per
plant, 3–5 times divided, 35–63-flowered;
peduncles terete, 23–30 cm long, green-pale
reddish brown, tomentose; bracts two at each
branching point, ovate, 5.5–6.6 × 1–2 mm, slightly
acute at apex, margin entire, hispid, green; pedicels
terete, 4–26 mm long, narrow, tomentose. Flowers
12–14 × 6.7–11 mm, held horizontal. Calyx 5-
partite, very deeply divided or basally connate; lobes
linear to ovate-elliptic, unequal, posterior lobe
small, 2.7–3.4 × 0.5–0.9 mm, others 3.55–4.35 ×
0.8–1.4 mm, green, obtuse at apex, margins entire,
glandular hairy. Corolla zygomorphic, ventricose,
two lipped, glandular hairy outside, glabrous inside;
tube 5–9 × 3.4–4.5 mm, dark to pale lilac with a
yellow blotch at the throat, swollen towards the
mouth; lobes widely ovate, rounded at apex dark
to pale lilac, upper lip of two lobes, 3.7–4 × 4–5.1
mm, lower lip of three lobes, 3.9–4.4 × 4.2–5.8 mm.
Stamens 2, adnate to the base of corolla, included;
filaments 3.3–4.3 mm long, glabrous, yellow;
anthers reniform, 1.2–1.7 × 0.4–0.76 mm, milky

88 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Fig. 22.
Henckelia wayanadensis
Janeesha & Nampy:
a. Plants in natural habitat; b. Habit; c. Bract; d. Flower; e. Calyx; f. Calyx with pistil;
g. Hairs on the pedicel; h. Stamens; i. Staminode; j. Pistil; k. Purplish glandular hairs on the ovary; l.l.
l.l.
l. Immature capsule; m.m.
m.m.
m. Seeds (a-l from
A.P.
Janeesha
&
P.G. Arunkumar
137671; m from
A.P. Janeesha
&
K. Haseem
137690).

89
Janeesha & Nampy
Fig. 23.
Comparison of
Henckelia wayanadensis
Janeesha & Nampy and
H. fischeri
(Gamble) A.Weber & B.L.Burtt: a1-i1
H. wayanadensis
;
a2-i2 & a3-i3
H. fischeri
: a. Habit; b. Flowers; c. Calyx; d. Calyx with pistil; e. Stamens; f. Staminodes; g. Pistil; h. Immature capsules; i. Seeds.

90 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
white to cream. Staminodes 2; filaments 1.3–3 mm
long, hyaline, glabrous; antherodes a small knob-
like structure. Ovary ovoid to oblong, 2.7–3.5 ×
0.9–1.3 mm, green, glandular hairy, gland tip
purplish; style terete, 2.2–3.6 mm long, green,
glabrous towards apex, glandular hairy towards
base; stigma obliquely subcapitate, 0.4–0.5 × 0.22–
0.24 mm, papillate, pale green. Capsules cylindrical,
straight or slightly curved, 20–22 × 1.6–2 mm,
plagiocarpic, dehiscing loculicidally long the dorsal
side, reddish brown, glandular, hispid, with
persistent stigma and calyx. Seeds numerous,
elliptic, 0.65–0.72 × 0.20–0.24 mm, truncate at both;
testa dark brown, reticulate.
Flowering & fruiting: Flowering from July to August
and fruiting from August to September.
Etymology: The species epithet ‘wayanadensis’ refers
to the locality ‘Wayanad’, the floristically rich
district of Kerala in which the new taxon was
discovered.
Habitat: In rock crevices, usually at an elevation
above 900 m, in association with Parahemionitis sp.
(Pteridaceae), Cyanotis sp. (Commelinaceae) and
some moss and grass species.
Distribution: Hitherto known only from the type
locality (Fig. 26).
Specimens examined: INDIA, Kerala, Wayanad
district, Kaduvakuzhi hill top, 08.07.2016, A.P.
Janeesha & P.G. Arunkumar 137671; Ibid.,
15.09.2016, A.P. Janeesha & K. Haseem 137690; on
the way to Kaduvakuzhi, 500 m away from
Adivaram, 08.07.2016, A.P. Janeesha & P.G.
Arunkumar 137670 (CALI).
Conservation status: The new species is currently
known only from one locality with three
subpopulations in an unprotected area, separated
by a distance of 50–60 meters, with a total of 61
mature plants. There is the possibility of
disturbances in the future due to human activities,
where local peoples are exploring the area for wood,
forage grasses etc. The Area of Occupancy (AOO)
is likely to be less than 10 km2. Therefore the species
is categorized as Critically endangered (CR),
B2ab(iii,iv,v) according to IUCN criteria (IUCN,
2019).
Henckelia wightii (C.B.Clarke) A.Weber &
B.L.Burtt, Beitr. Biol. Pflanzen 70: 359.
1998[“1997”]. Didymocarpus rottleriana Wall.,
Wight, Icon. Pl. Ind. Orient. 4: 24. t. 1348. 1848.
Didymocarpus rottlerianus Wall. ex R.Br. var. wightii
C.B.Clarke in A.DC. & C.DC., Monogr. Phan. 5:
101. 1883, C.B.Clarke in Hook.f., Fl. Brit. India 4:
354. 1885[“1884”]. Didymocarpus wightii
(C.B.Clarke) Gamble, Fl. Madras 2: 989. 1924;
A.N.Henry et al., Fl. Tamil Nadu Ind., Ser I:
Analysis 2: 133. 1987; M.Ahmedullah & M.P.Nayar,
Endemic Pl. Indian Region 1: 143. 1987; Sasidh.,
Biodivers. Doc. Kerala Part 6: Fl. Pl. 332. 2004;
T.S.Nayar et al., Fl. Pl. Kerala 343. 2006, Fl. Pl.
Western Ghats 1: 529. 2014. Lectotype (designated
here): INDIA, Tamil Nadu, Tirunelveli district,
Shevghery, Courtallam, s.d., Wight 2323 (K
[K000858201 digital image!]; isolecto: E
[E00155179 digital image!]).
Didymocarpus sivagiriensis Rajakumar, Selvak.,
S.Murug. & Chellap., Indian J. Forest. 32(3): 481.
2009; T.S.Nayar et al., Fl. Pl. Western Ghats 1: 527.
2014, syn. nov. Henckelia sivagiriensis (Rajakumar,
Selvak., S.Murug. & Chellap.) E.S.S. Kumar, Polish
Bot. J. 59(1): 149. 2014. Type: INDIA, Tamil
Nadu, Tenkasi district, Thalaiyanai reserve forest,
s.d., Rajakumar, Selvakumari, Murugesan &
Chellapperumal 829 (holo JCH). Figs. 24 & 25
Perennials, rosette-forming, hairy herbs with
rootstocks. Roots adventitious, thick. Leaves 12–
20, basal, in whorls of 4–7, sessile to sub-sessile;
laminae spatulate-obovate to elliptic, 3.8–10.7 ×
1.6–3.6 cm, obtuse to rounded at apex, attenuate at
base, margins slightly crenate and undulate, surfaces
smooth, velvety, with white woolly hairs on both
surfaces, but more so on the younger leaves and on
lower veins, becoming less so with age; veins 4–6
on each side, much branched and conspicuous
beneath. Cymes axillary, dense; 2–6 per plant, 3–4
times divided, 5–13-flowered; peduncles terete, 10–
18 cm long, greenish, glandular hairy; bracts two
at each branching point, minute, completely

91
Janeesha & Nampy
woolly, c. 2 mm long; pedicels terete, 9–18 mm
long, narrow, glandular hairy. Flowers 11–15 × 10–
14 mm. Calyx 5-partite, very deeply divided; lobes
oblong-lanceolate, unequal, posterior lobe small,
1.8–2.1 × 0.45–0.6 mm, other lobes, 2–3 × 0.6–0.8
mm, slightly acute at apex, margins entire, with
white woolly hairs on the upper side. Corolla
zygomorphic, ventricose, two lipped, glandular
hairy outside, glabrous inside; tube lilac to pale lilac
with a yellow patch in the throat; lobes orbicular,
ovate to rounded at apex, lilac to pale lilac, upper
lip of two lobes, lower lip of three lobes. Stamens
2, adnate to the base of corolla, included; filaments
c. 2 mm long, glabrous; anthers reniform, glabrous.
Staminodes 3 (sometimes 2); filaments c. 1.2 mm
long, glabrous; antherodes not seen. Ovary ovoid
to cylindrical, c. 2.5 mm long, covered with dense
white woolly hairs, completely enclosed within
calyx; style terete, c. 1.3 mm long, glabrous; stigma
obliquely subcapitate, papillate, pale green. Capsules
linear-cylindrical, straight, 11–17 mm long,
plagiocarpic, dehiscing loculicidally along the
Fig. 24
..
..
. Henckelia wightii
(C.B.Clarke) A.Weber & B.L.Burtt: a. Habit showing whorls of five leaves; b. Flower; c. Immature capsules
(from plants cultivated at Gurukula Botanical Sanctuary, Periya).
Fig. 25. Lectotype of
Henckelia wightii
(C.B.Clarke) A.Weber &
B.L.Burtt. © The Board of Trustees for The Royal Botanic Gardens,
Kew. Reproduced with permission.

92 A taxonomic revision of
Henckelia
(Gesneriaceae) in South India
Fig. 26. Distribution of
Henckelia repens
(Bedd.) A.Weber & B.L.Burtt,
H. wayanadensis
Janeesha & Nampy and
H. wightii
(C.B.Clarke)
A.Weber & B.L.Burtt in South India (drawn using QGIS v.3.12.1).
dorsal side, green, tomentulose with persistent
stigma and calyx. Seeds numerous, reticulate.
Flowering & fruiting: Flowering from July to
October and fruiting from August to November.
Habitat: On moist rocks.
Distribution: Endemic to the southern Western
Ghats (Fig. 26).
Specimens examined: INDIA, Kerala, Idukki
district, Karadippara-Munnar, 25.07.2013, A.P.
Janeesha & K.M. Manudev 134203 (CALI). Tamil
Nadu, Erode district, Attamalai hills, 24.07.1988,
N. Venkatasubramanian 1424 (FRC); Tenkasi
district, New Courtallam, 16.07.2015, A.P.
Janeesha, S. Resmi & P.G. Arunkumar 137628
(CALI); Virudhunagar district, Kendiparai to
Cumbumedu-Ayyanarkovil, 23.09.1971, E.
Vajravelu 38720; Mudaliarattu, 11.12.1971, E.
Vajravelu 39339 (MH).
Conservation status: The species is known from four
districts in Kerala and Tamil Nadu where a total of
five localities were known. The Extent of
Occurrence (EOO) is estimated to be c. 2100 km²
and Area of Occupancy (AOO) is 20 km². During
a period between 2013 and 2017, one of the authors
(JAP) visited four localities except Ayyanarkovil and
was unable to observe more than 5 mature plants
in two localities, i.e., Munnar and Courtallam. Thus,
a decline in number of localities from five to two
over five years has been observed, in (1) EOO (ii)
AOO; (iv) number of locations or subpopulations
and (v) number of mature individuals. Because of
the very low number of plants found, we consider
this species to be Critically endangered (CR), D
according to IUCN guidelines, with a high risk of
extinction in the near future (IUCN, 2019).
Notes: Rajakumar et al. (2009) described Henckelia
sivagiriensis (as Didymocarpus sivagiriensis Rajkumar,

93
Janeesha & Nampy
Semvak., S.Murug. & Chellap.) and differentiated
it from H. wightii by its longer leaves, presence of
glandular hairs, longer dichotomously branched
scapes, bluish corolla, puberulous style and longer
capsules. Critical examination of specimens and
protologues of both these taxa revealed that the
characters outlined for separating the former from
the latter are intergrading (see description above)
and hence H. sivagiriensis is reduced to synonymy
of H. wightii.
Typification: Wight described this species based on
material collected from the Shevghery hills,
Courtallam in Tamil Nadu. Since Vitek et al. (2000)
cited only syntypes, it cannot be considered a
lectotypification. Two sheets of Wight’s materials
were located, while searching for the type
specimens, one each in E (E00155179) and K
(K000858201). The sheet K000858201 has an
HRWP label and bears a field label, “Shevghery
hills August 1836” and an annotation by Clarke
“the specimen from which Wight Ic. 1348 was
taken”. The sheet itself has the no. 2323/1837 in
the bottom left hand corner. The sheet in E
(E00155179) has an HWP label annotated
“Didymocarpus rottlerianus var. wightii” and “Herb.
Wight. propr. n. 2679” in the top right hand corner
and another in the lower left hand corner “1837,
No 2323”, presumably referring to the year 1837.
The specimen at K (K000858201) used for
illustrating this species by Wight is selected here as
the lectotype and the other at E (E00155179) as the
isolectotype.
Acknowledgements
The authors are grateful to Curators of BM, CAL,
E and MH for permitting consultation of the
specimens; Dr. J.V. Sudhakar (MH), Dr. Mark
Carine (BM), Dr. K. Karthigeyan (CAL) and Dr.
Lesley Walsingham (K) for comments on the
annotations on the herbarium materials; Dr. Kanchi
N. Gandhi (Nomenclature Registrar, Harward
University) for Nomenclatural suggestions; Dr.
Anton Weber (Vienna) for literature; Dr. A.K.
Pradeep (Calicut University), Dr. K.M. Manudev
(St. Joseph’s College (Autonomous), Kozhikode;
Gurukula Botanical Sanctuary, Periya, Wayanad
and Dr. (Sr.) Jaseena Joseph (Principal, Providence
Women’s College, Kozhikode) for various helps;
Dr. Shahina P.M., Mr. Arunkumar P.G., Ms. Resmi
S, Mr. Syam Radh S., Dr. Shinoj K., Dr. Thoiba
K., Dr. Smitha K., (Research Scholars, Calicut
University), Mr. Habeeb Rahman, Mr. Rajeevan
and Mr. Ahmadul Kabeer for helping collections.
We are thankful to Dr. Michael Möller (E) and Dr.
David Middleton (Singapore Botanic Gardens) for
their valuable comments for improving the
manuscript. We thank Dr. K.M. Prabhukumar
(CMPR, Kottakkal) and Dr. M.G. Prasad (NBRI,
Lucknow) for photographic images of some species
of Henckelia. SN is thankful to SERB New Delhi
(EMR/2016/007346 dated 01/11/2018) for funding
and JAP to University Grant Commission for
Fellowship.
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