Acta Botanica Hungarica 53(3–4), pp.

283–289, 2011
DOI: 10.1556/ABot.53.2011.3–4.10

THE LIVERWORT THYSANANTHUS SPATHULISTIPUS

(LEJEUNEACEAE) LONG-LOST IN INDIA
REDISCOVERED

A. E. D. DANIELS1, K. C. KARIYAPPA1 and P. DANIEL2

1
Bryology Laboratory, Botany Department and Research Centre, Scott Christian College
(Autonomous), Nagercoil–629 003, Tamil Nadu, India
2
36, Arulnagar, Nagercoil–629 001, Tamil Nadu, India

(Received 9 February, 2011; Accepted 30 May, 2011)

The liverwort Thysananthus spathulistipus, earlier known to be distributed in the Eastern

Himalaya, Northeast India and the Andamans in India, has been rediscovered in the Western
Ghats after more than a century. It was last collected by Decoly and Schaul in Kurseong in
1898. A detailed description of the species, an illustration and a distribution map are provided.

Key words: Peninsular India, rediscovered, Thysananthus spathulistipus, Western Ghats

INTRODUCTION

The genus Thysananthus Lindenb. belonging to the family Lejeuneaceae

has ca 10 species in the world (Gradstein 1992) and 2, viz., T. gottschei (J. B. Jack
et Steph.) Steph. and T. spathulistipus (Reinw., Blume et Nees) Lindenb. in India
(Bapna et Kachroo 2000, Parihar et al. 1994). Mitten (1861) who first reported T.
spathulistipus from India based on a Griffith material from the Khasia Hills,
once part of the colonial Assam, did so under the name Lejeunea spathulistipa
(Reinw. et al.) Dumort. Subsequently, Verdoorn (1934) who included T. spa-
thulistipus cited the Griffith material referred to above and added another ma-
terial collected by E. H. Man in 1890 in the Andaman (and Nicobar) Islands, an
archipelago in the Indian Ocean and cut off from the Indian mainland by a dis-
tance of about 1,200 km.
While exploring the erstwhile Tirunelveli-Travancore Hills, now a part of
the Agasthyamalai Biosphere Reserve in the southern Western Ghats for a
bryoflora, material of this species was collected at Pandimotta in Kollam Dis-

0236–6495/$ 20.00 © 2011 Akadémiai Kiadó, Budapest

284 DANIELS, A. E. D., KARIYAPPA, K. C. and DANIEL, P.

trict in Kerala, which is a rediscovery of the species in the Western Ghats in

Peninsular India. Though our primary objective is to bring out an inventory of
bryophytes of the southern Western Ghats eventually, one of the secondary
objectives is to identify/recollect the endemic and rare species particularly
those not collected after the British botanists during the colonial era. In an ear-
lier paper we reported the rediscovery of the liverwort Schistochila aligera
(Nees et Blume) J. B. Jack et Steph. (Daniels and Daniel 2008). Here we report
the rediscovery of Thysananthus spathulistipus made after more than a century.
With a view to facilitating its rediscovery elsewhere and fixing its status, this
species is described in detail, illustrated and its distribution in India mapped.

Thysananthus spathulistipus (Reinw., Blume et Nees) Lindenb.

Basionym: Jungermannia spathulistipa Reinw., Blume et Nees (1824), Nova Acta

Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 212; Sprengel (1827), Syst. Veg. 4(1): 222,
No. 67; Nees (1830), Hepat. Jav.: 38, No. 48. ≡ Lejeunea spathulistipa (Reinw., Blume et Nees)
Dumort. (1835), Recueil Observ. Jungerm. 1: 12; Mitten (1861), J. Proc. Linn. Soc., Bot. 5: 109;
Spruce (1885), Trans. Proc. Bot. Soc. Edinburgh 15: 106. ≡ Frullania spathulistipa (Reinw.,
Blume et Nees) Nees (1838), Naturgesch. Eur. Leberm. 3: 211. ≡ Phragmicoma spathulistipa
(Reinw., Blume et Nees) Mitten (1871), Seemann: Fl. Vit., p. 412. ≡ Thysananthus spathulistipus
(Reinw., Blume et Nees) Lindenb. in Gottsche et al. (1845), Syn. Hepat., p. 287; Jack et
Stephani (1894), Bot. Centralbl. 60(4): 104; Schiffner (1898), Consp. Hepat. Arch. Ind., p. 307,
No. 386; Verdoorn (1934), Ann. Bryol. (Suppl.) 4: 167; Mizutani (1969), J. Hattori Bot. Lab.
32: 138; Campbell (1971), J. Roy. Soc., New Zealand 1: 13; Kitagawa (1973), J. Hattori Bot.
Lab. 37: 271; Onraedt (1985), Cryptog. Bryol. Lichénol. 6: 173; Thiers and Gradstein (1989),
Mem. New York Bot. Gard. 52: 71; Parihar et al. (1994), Hepat. Anthocerot. India, p. 38;
Grolle (1995), Bryophyt. Biblioth. 48: 139; Srivastava (1998), Topics Bryol., p. 72; Bapna and
Kachroo (2000), Hepatic. India 2, p. 292; Gradstein et al. (2002), Acta Bot. Fenn. 174: 80;
Piippo et al. (2002), Ann. Bot. Fenn. 39: 120; Wigginton (2002), Trop. Bryol. Res. Rep. 3: 73. –
Type: Indonesia, Java, Bantam, Leback. Coll.: Blume, s.n. (Holotype: STR; isotypes: G, W).
(Figs 1–18).
= Lejeunea ungulata Mitten (1861), J. Proc. Linn. Soc., Bot. 5: 110. – Type: Burma, Rangoon.
Coll.: McClelland, s.n. (BM).
= Thysano-lejeunea amboinensis Schiffner (“1889” 1890), Forschungsreise S. M. S. “Gazelle” 4:
22. – Type: Indonesia, Amboina (FH).
= Thysananthus sikkimensis Steph. ex [Schiffner (1899), Österr. Bot. Zeitschr. 49: 205, No. 30,
nom. nud.] Stephani (1912), Sp. Hepat. 4: 798. – Types: India, (Sikkim-) Himalaya,
prope Kurseong, 6,000 ft. Coll.: Rev. P. Decoly and Schaul, s.n., 28 March 1889
(Schiffner No. 516), Bryoth. E. Levier (NY); (Bhutan), “zwischen Maria Basti & Labar,
5,000–6,000 ft”. Coll.: Rev. L. Durel (159), 12 April 1898, Herb. Levier. (G).
= Thysananthus aculeatus Herzog (1931), Ann. Bryol. 4: 89; Mizutani (1969), J. Hattori Bot.
Lab. 32: 137. – Type: Philippines, Mt Banahaom C. Luzon. Coll.: Baker (7079/7083), 12
Dec. 1913 (BM).
= Thysananthus fuscobrunneus Horik. (1934), J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2: 251.
– Type: Taiwan, Mt Chipon, Prov. Taitô. Coll.: Horikawa (10505), Dec. 1932 (HIRO).

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 THYSANANTHUS SPATHULISTIPUS LONG-LOST IN INDIA REDISCOVERED 285

Figs 1–18. Thysananthus spathulistipus (Reinw. et al.) Lindenb. – 1 = plant, 2 = cross section of
stem, 3–4 = leaves, 5 = leaf apical cells, 6 = leaf median cells, 7 = leaf basal cells, 8 = cells with
oilbodies, 9 = leaf lobule teeth, 10–12 = underleaves, 13 = male inflorescence, 14–15 = female
bracts, 16 = female bracteole, 17 = perianth, 18 = cross section of perianth (1–18 from K. C.
Kariyappa 15)

Acta Bot. Hung. 53, 2011

286 DANIELS, A. E. D., KARIYAPPA, K. C. and DANIEL, P.

Plants monoicous, corticolous, pendant, 3–5 cm long, olive green. Stems

with branching of Frullania-type, pinnate or bipinnate, 0.14–0.18 µm in diam.,
with 12–15 cells across; cortical cells thick-walled, non-trigonous, quadrate-
hexagonal, 4–20 × 4–8 µm; medullary cells with walls thinner than that of corti-
cal ones, non-trigonous, quadrate-hexagonal, 8–20 × 8–12 µm. Leaves incubous,
wide-spreading, 1.2–1.5 × 0.56–0.8 mm, oblong-ovate, asymmetric, cordate at
base, acute and toothed at apex; margin widely arched dorsally, incurved ven-
trally; keel arched; cells thick-walled, with bulging trigones and intermediate
nodular thickenings; apical and marginal cells 6–8 × 4–6 µm; median ones
20–40 × 12–20 µm; basal ones 28–48 × 16–20 µm; oil bodies 3–8 per cell, seg-
mented, 6–12 × 2–4 µm, ovoid to spindle-shaped; lobules ca 1/3 as long as
leaves, rectangular, 2-toothed at apex; teeth distinct, 3- or 4-celled. Under-
leaves imbricate, transversely inserted, not appendiculate, roughly cuneate,
irregularly denticulate at margin above, sometimes faintly 2-lobed, recurved
at lateral margin, 3–4 times as broad as stem, 0.64–1.12 × 0.56–0.96 mm, smaller
towards base and larger towards female inflorescences. Male inflorescences
terminal on lateral branches, 1.5–3 mm long; bracts 8–12, smaller than leaves,
0.35–0.4 × 0.25–0.32 mm, toothed at apex; lobules swollen, many-toothed;
bracteoles imbricate, smaller than underleaves, 0.55–0.7 × 0.15–0.25 mm,
toothed at margin above. Female inflorescences terminal on main stem or
branches, with subfloral innovations; bracts larger than leaves, 1.5–2 × 0.9–1.1
mm, oblong-ovate, toothed at margin above; lobules ovate-oblong, 0.7–0.9 ×
0.4–0.6 mm, toothed; bracteoles twice as long as underleaves, 1.5–1.8 × 0.7–0.9
mm, lingulate, inflexed and undulate at margin above, slightly bifid at apex.
Perianth 1.5–1.7 × 0.7–0.9 mm, oblong, inflated, 3-keeled; keels toothed.
Sporogonium seen immature.
Habitat: Corticolous, in evergreen forests, ca 200 m.
Distribution: Tropics. India: Andamans, Eastern Himalaya (W Bengal),
NE India (Meghalaya) and Western Ghats (Fig. 19).
Specimens examined: Kerala, Kollam District, Westren Ghats, Shankili
Forest, Pandimotta, ca 200 m. Coll.: K. C. Kariyappa (15), 02.06.2009. Herb.
Scott Christian College, Nagercoil (SCCN).

DISCUSSION

Emilio Levier (1839–1911) bought specimens from many collectors and

afterwards sold them to museums and other collectors under “Bryotheca E.
Levier” with about 2,400 specimens (Winter, in litt.). Schiffner (1899) based on
material collected by Rev. R. Durel on 12 April 1898 in “British-Bhutan” be-
tween Maria Basti and Labar, which he had received from his Italian friend the

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 THYSANANTHUS SPATHULISTIPUS LONG-LOST IN INDIA REDISCOVERED 287

said Levier, listed 35 liverworts including T. sikkimensis without a description,

but with remarks on its affinity with allied species and attributed the name to
Stephani. He also mentioned in the paper that F. Stephani had a nice collection
from Kurseong (in Sikkim). Schiffner had found that some plants of the Durel
collection and those from Stephani were the same. As he anticipated Stephani
to publish these, Schiffner mentioned such plants only by their manuscript
names. Long (1979) discussed the issue of Durel not having collected in the
so-called British-Bhutan and pointed out that Schiffner (1899) wrongly as-
sumed so as may be evident from the title of his paper and that these areas are
near Kalimpong (Darjeeling District) in the State of West Bengal in India.
Stephani (1912) validly published the name T. sikkimensis, which is con-
sidered a synonym of T. spathulistipus by recent workers. In the protologue
Stephani (1912) mentioned only “Hab. Himalaya” concerning a type. How-
ever, the microfiche of the unpublished icones of Stephani (image 10212) men-
tions Herb. Levier. and the Decoly and Schaul material at NY, which was re-
ceived from G has information such as India (Sikkim-) Himalaya, prope Kur-

Fig. 19. Distribution of Thysananthus spathulistipus in India (l earlier localities; ª present lo-
cality)

Acta Bot. Hung. 53, 2011

288 DANIELS, A. E. D., KARIYAPPA, K. C. and DANIEL, P.

seong and Bryotheca Levier to boot. Since the material at G and NY came from
the same source “Bryotheca E. Levier” it is reasonable to assume that they are
original material.
Granting that Durel was the last to collect material of T. spathulistipus (= T.
sikkimensis) in 1898, the present collection, made in the Western Ghats far away
from the earlier known region at the other end of the country and a totally dif-
ferent phytogeographical region, is indeed an interesting rediscovery of this
species after more than a century, 112 years to be exact. Though this species is
said to be distributed in the tropics and somewhat common in other regions
(vide Specimens examined in Onraedt 1985, Piippo et al. 2002, Thiers and
Gradstein 1989), it appears to be rare in India.

*
Acknowledgements – The senior author is thankful to the Ministry of Environment and
Forests, Govt. of India, New Delhi, for funding a project under the AICOPTAX and to Dr M.
Sanjappa, former Director, Botanical Survey of India, Kolkata, for encouragement and a
copy of D. G. Long’s paper. The authors are thankful to the Kerala State Forest Department
for permission to explore the area in question and help in the field, Dr R. L. Zhu (HSNU), for
confirming the identity of the material, Mr M. J. Wigginton (Peterborough, England) and Dr
X.-L. He (Botanical Museum, University of Helsinki, Finland), for help with literature, to Dr
G. Winter (Senckenberg Natural History Museum, Germany), for help with literature, in-
formation on image 10212 in the microfiche of Stephani’s unpublished icones, on the labels
of the material of T. sikkimensis at G and NY and unravelling the E. Levier connection and Dr
M. Price (G), for information on the label of T. sikkimensis material at G. The senior author is
thankful to Dr S. C. Rose (Principal, Scott Christian College), for encouragement.

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