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Accepted by Rafael Govaerts: 29 Mar. 2012; published online in PDF: 4 Apr. 2012 1
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright © 2012 Magnolia Press
Phytotaxa 50: 1–12 (2012)
www.mapress.com/phytotaxa/Article
Gaultheria stapfiana (Ericaceae), a species to be recognized: insights from
morphology, leaf anatomy and pollen morphology
SUBHASIS PANDA
Taxonomy & Biosystematics Laboratory, Post-Graduate Department of Botany, Darjeeling Government College, Darjeeling-734101,
India. E-mail: bgc.panda@gmail.com
Abstract
This study, based on an investigation of morphology (as assessed through field and herbarium studies), leaf anatomy
(stomata and vein islets), and pollen morphology (LM and SEM) reveals that Gaultheria stapfiana Airy Shaw is a
distinct species and not a variant within of the G. hookeri C.B.Clarke as proposed by Ruizheng & P.F. Stevens (2005).
Line drawings, photographs of living plants and specimens, photographs of leaf anatomical characters, and pollen of G.
stapfiana and its allies, show that this species is clearly delimited from both G. hookeri and G. fragrantissima Wall.
Key words: Gaultheria stapfiana, G. hookeri, Ericaceae, exomorphology, leaf anatomy, pollen morphology
Introduction
Gaultheria stapfiana Airy Shaw (Fig. 1 & 4A–B) was described by Airy Shaw (1952) based on a specimen
from Yunnan (China) collected by George Forrest in 1919 (No. 18021, K!). This species is similar to G.
fragrantissima Wall. var. racemosa Hook.f. ex C.B.Clarke, which was treated as a heterotypic synonym of G.
stapfiana by Airy Shaw (1952). Gaulthera frangrantissima var. racemosa was collected by J. D. Hooker
(1849) from Darjeeling and described by C. B. Clarke (1882). Clarke rightly distinguished Hooker’s specimen
(K!) from var. racemosa mainly due to the presence of long racemes (10–12.7 cm long), narrow lanceolate
leaves (110 × 12 mm), and median bracteoles. Hooker’s specimen was also distinguished from G. hookeri
C.B.Clarke by the presence of smaller bracts, sparsely pubescent branches, and a tall habit (not dwarf like G.
hookeri).
Airy Shaw (1952) accepted Gaultheria stapfiana as a new species and distinguished it from its two close
allies, G. fragrantissima (Fig. 3 & Fig. 4E–F) and G. hookeri (Fig. 2 & Fig. 4 C–D), on the basis of its sparsely
pubescent branches, oblanceolate, obovate to oblong-elliptic lamina, bract that are longer than those of G.
fragrantissima but shorter than those of G. hookeri, bracteoles median on the pedicel (as in G. hookeri) and
arborescent habit (like G. fragrantissima, but not shrubby as in G. hookeri). Airy Shaw (1952) described
several overlapping characters between G. stapfiana and G. hookeri, such as lamina size (4–14 × 1–4.5 cm),
raceme length (2–5 cm), bract length (3–5 mm), bracteole position (median), corolla length (3–4 mm), nature
of the stamens (1–2 mm long), ovary (globose, tomentose) and fruits (capsule enclosed in a fleshy bluish-
purple calyx, ovoid). Based on the above overlapping characters, Ruizheng & P.F. Stevens (2005) treated G.
stapfiana as a synonym of G. hookeri var. hookeri in Flora of China.
After Airy Shaw (1952), workers like Sealy (1973), Middleton (1991a; 1991b), Ruizheng & Stevens
(2005), Panda (2008) and Panda et al. (2009) worked on Gaulthera stapfiana. As a result of revisionary work
on Indian Ericaceae conducted as part of the Flora of India Project (1999–2004; which involved
morphological investigations, see Table 1) as well as laboratory-based studies of leaf anatomy (i.e., leaf-
stomata, Fig. 5, leaf areolar pattern, Fig. 6) and pollen morphology (Fig. 7) conducted from 2008–2010,
evidence shows that G. stapfiana Airy Shaw is a good species and not a synonym of G. hookeri.
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Materials and Methods
Exomorphology.—For this detailed study, flowers from living material and herbarium specimens (ASSAM,
BSIS, CAL, DD, MH) were dissected and examined. The floral parts were pasted on white paper cards (8" ×
5") using synthetic gum and measured. After considering the annotations of earlier workers and comparing
them with the type specimens, protologues and with live materials, complete and detailed amplified
descriptions of each of the three species discussed above were made (Table 1).
TABLE 1. Comparison of morphological characters among G. stapfiana Airy Shaw and the similar species G. hookeri
C.B. Clarke and G. fragrantissima Wall. (Figs. 1–4)
Stomatal slide preparation.—Small square pieces of leaf blades (ca. 1 sq cm) were excised from the base,
middle and apex (10 leaf samples from different natural habitats were taken). Slides were prepared using 10%
HNO3-boiling for 10 minutes and a 5% KOH overnight (12–24 hours) treatment with and without boiling.
Pieces were rinsed in sterilized water until clear. After clearing, the pieces were dehydrated in an ethanol
series followed by staining with 10% safranin and mounted onto a microscope slide in DPX (with pieces from
the basal, middle and apical regions on one slide). The slide was examined under an Olympus (Tokyo, Japan)
light microscope using 10× and 40× objectives and Camera Lucida drawings were made. The slides were
deposited in the laboratory of Taxonomy & Biosystematics, PG., Department of Botany, Barasat Govt.
College (Calcutta University). The descriptive terminology follows Metcalfe and Chalk (1950), Dilcher
(1974), Stace (1965, 1989), Fahn (1997), Carpenter (2005), and Judd et al. (2008).
Methodology of leaf clearing for venation study (areoles).—Entire mature leaves (10 leaf samples from
different natural habitats were taken) were immersed in 2.5% NaOH solution until clear (closed condition). In
the present study, most of the leaves were cleared after 11–14 days of NaOH treatment. After 14 days, these
NaOH-treated leaf samples were again immersed in 2.5% NaOH solution for 2–3 days followed by 1 drop
Vegetative and
reproductive
morphological
characters
G. stapfiana G. hookeri G. fragrantissima
1. Habit 0.3–2 m high. 0.3–1 m high. 0.5–2 (-3.5) m high.
2. Stem and branch
texture Indumentum Stem terete, blackish-
brown, glabrous. Branches
terete, light green, sparsely
hispid-setose.
Stem terete, blackish- brown,
sparsely setose. Branches
terete, light green to pink,
densely hispid-setose.
Stem terete, grey- brown to
light green, glabrous.
Branches winged, blood red,
deep pink to rarely light
green, glabrous, glaucous.
3. Abaxial leaf
indumentum Punctate. Setulose and punctate. Punctate.
4. Raceme length No.
of flowers 2.5–11 cm long. 10–24 (-
40)-flowered. 1.5–5 cm long. 10–15-
flowered. 2.5–11 cm long. 10–20 (-
40)-flowered.
5. length of flower 5–11 mm long. 7–15 mm long. 8–15 mm long.
6. Bract 3–5 × 1.5–2.5 mm. 5–7 × 3–4.5 mm. 2–4 × 1.5–2.5 mm.
6. Position of
bracteoles on pedicel Median. Subbasal to median. Apical to subapical.
7. Stamen length
Anther awns 1–2 mm long. Filaments
grey-white. Each lobe of
anther with 2 unequal awns
to rarely equal. Awn
minute.
2–3 mm long. Filaments
pinkish. Each lobe of anther
with 2 equal awns. Awn up
to 1 mm long.
3 mm long. Filaments grey-
white. Each lobe of anther
with 2 equal awns Awn 0.5–
1 mm long.
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GAULTHERIA STAPFIANA (ERICACEAE)
chloral hydrate treatment overnight. Leaf samples were then washed in distilled water. After clearing, an
entire leaf was dehydrated in an ethanol series followed by staining with 1% safranin and mounted onto
microscope slide in DPX (with entire leaf on a single slide). The slides were deposited in the laboratory of
Taxonomy & Biosystematics, PG. Department of Botany, Barasat Govt. College (Calcutta University). The
descriptive terminology follows Hickey (1973) and Dilcher (1974).
Preparation of pollen slides.—The method used in this study followed Erdtman (1952, 1969, 1986) and
Sarwar et al. (2006). The slides were deposited in the laboratory of Taxonomy & Biosystematics, PG
Department of Botany, Barasat Govt. College (Calcutta University). Pollen morphological data were based on
15 flower buds from a single population of each species. In the present study, three different populations
(Sikkim, Darjeeling and Arunachal Himalayas) were studied.
Slide preparation for SEM.—Acetolysed pollen grains were mounted on the metallic stub using double
stick tape (flower buds from single voucher specimen). Observations were made with Hitachi S530 (SEM,
Tokyo, Japan) in the high vacuum mode at an applied voltage of 15 KV (digital images).
Results
Gaultheria stapfiana is morphologically distinct from G. hookeri and G. fragrantissima in its sparsely hispid-
setose branches and glabrous stems, G. hookeri having densely hispid-setose branches and setose stems. The
branches of G. fragrantissima are glabrous, reddish, and winged. Additional differences are found in the
morphology (Table 1), leaf-anatomy (Table 2), and pollen morphology (Table 3) of the three species. Stems,
TABLE 2. Comparison of Leaf anatomical characters among G. stapfiana Airy Shaw and its two close allies i.e. G.
hookeri C.B. Clarke and G. fragrantissima Wall. (Figs. 5–6).
Leaf anatomical
characters G. stapfiana G. hookeri G. fragrantissima
Leaf-stomata Types: Para and
amphiparacytic. Average
dimension: 22.7(–32) ×
21.3(–28) µm.
Types: Para and pericytic.
Average dimension: 30 × 26
µm.
Type: only Paracytic.
Average dimension:
25.5(18–33) × 22(16–28)
µm.
Epidermal cells Epidermal cells: polygonal.
Epidermal walls in surface
view: arched
Epidermal cells: polygonal.
Epidermal walls in surface
view: straight
Epidermal cells: Hexa- to
polygonal. Epidermal
walls in surface view:
straight and sinuous.
Stomatal Index (SI) 15.05. average number of
stomata is 28 per 158
epidermal cells per
microscopic field.
26.2 average number of
stomata is 32 per 90 epidermal
cells per microscopic field.
15.0. average number of
stomata is 36 per 204
epidermal cells per
microscopic field.
Leaf venation Shape
of areole: Average
numbers of vein islets
(areoles) per sq. mm:
Vein endings: Vein
ends:
Quadrangular to pentagonal.
Larger areole (average): 998
× 353 µm. Smaller areole
(average): 294 × 176 µm. 6
per 1 sq. mm. 48 (average)
per 1 sq. mm, tapered to
bluntly acute, ends not
divided.
Quadrangular. Larger areole
(average): 823 × 306 µm.
Smaller areole (average): 270
× 317 µm. 5 per 1 sq. mm. 45
(average) per 1 sq. mm,
taperated, acute to rarely
bulbous, ends not divided.
Tri-, Qua- to pentagonal.
Larger areole (average):
1200 × 470 µm. Smaller
areole (average): 411 × 188
µm. 8 per 1 sq. mm. 102
(average) per 1 sq. mm,
tapered, acute to bulbous,
ends lobed or divided.
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TABLE 3. Comparison of pollen morphological characters among G. stapfiana Airy Shaw and its two close allies i.e. G.
hookeri C.B. Clarke and G. fragrantissima Wall. (Fig. 7).
bracts and bracteoles, corolla and stamens basically appear to be similar in most herbarium specimens of G.
stapfiana and G. hookeri due to the range of variation that exists in G. stapfiana, e.g., G. hookeri has densely
hispid-setose branches, 10–15-flowered short and congested racemes up to 5 cm long, and comparatively
large bracts (5–7 mm long) and stamens (2–3 mm long), while plants of G. stapfiana show sparsely hispid-
setose branches, 10–40-flowered long racemes up to 11 cm long, and comparatively shorter bracts (3–5 mm
long) and stamens (1–2 mm long). Pollen (tetrad and individual grain dimensions, colpi length and exine
surface), and leaf anatomy (stomatal dimension) also show consistent differences between these two species.
Discussion
Gaultheria stapfiana Airy Shaw is recognized here as a distinct species due to presence of distinctive
vegetative and reproductive morphological characters (i.e., the combination of sparsely hispid branches,
racemes up to 11 cm long, bracts 3–5 mm long, and minute anther lobes with 2 unequal or equal awns),
differences in leaf anatomy (smaller stomatal dimension, 22.7 × 21.3 µm, arched epidermal walls, 15.05
stomatal index, and bluntly acute vein ends), and the form of its pollen grains (smaller tetrads, 25–28 µm in
diameter, rugulate-psilate exine surface and comparatively long colpi, 10–14 µm long) (Figs. 1–7).
Key to three species (for easy identification in the field and herbarium)
1. Twig always glabrous, often winged; bracteoles apical on pedicel; leaf vein ends lobed or divided...........................
...................................................................................................................................... 3. Gaultheria fragrantissima
1. Twig always hispid-setose, never winged; bracteoles subbasal to median on the pedicel; leaf vein ends never
divided.......................................................................................................................................................................... 2
2. Twig sparsely hispid-setose; racemes up to 11 cm long; bract 3-5 mm long; anther lobes with minute awns; average
dimension of leaf-stomata 22.7 × 21.3 µm, stomatal Index (SI) 15.05; exine surface of pollen tetrads rugulate-psilate
................................................................................................................................................................1. G. stapfiana
Pollen morphological
characters G. stapfiana G. hookeri G. fragrantissima
Tetrads Tetrahedral Tetrahedral Tetrahedral
Tetrad size (D) 25–28 µm diameter 32–34 µm diameter 32–36 µm diameter
P/E × 100 85.7 (sub-oblate) 79.4 (sub-oblate) 87.4 (sub-oblate)
Individual grain (d) 19.2–21.5 µm diameter 22–24 µm diameter 12.6–20.8 µm diameter
‘d’ with/without furrows ‘d’ possesses distinct
transverse furrows. ‘d’ possesses distinct
transverse furrows. ‘d’ does not possess
distinct furrows.
Exine 1.9–5.2 µm thick surface:
rugulate-psilate 1.9–3.2 µm thick surface:
faintly striate 2–2.8 µm thick surface:
faintly striate
D/d 1.30 1.43 2.03
Colpi 10–14 µm long 5.8–8.8 µm long 8.9–10.2 µm long
2f/D (Ratio of colpus
length (2f) to tetrad
diameter (D)
0.37–0.52 µm 0.17–0.26 µm 0.26–0.3 µm
Septum thickness 1.6–3.8 µm 1.6–3.8 µm 1.2–2.8 µm
Phytotaxa 50 © 2012 Magnolia Press • 5
GAULTHERIA STAPFIANA (ERICACEAE)
2. Twigs densely hispid-setose; racemes up to 5 cm long; bract 5-7 mm long; anther lobes with ca. 1 mm long awns;
average dimension of leaf-stomata 30×26 µm, stomatal Index (SI) 26.2; exine surface of pollen tetrads faintly striate
.................................................................................................................................................................. 2. G. hookeri
1. Gaultheria stapfiana Airy Shaw (1952: 171). TYPE: CHINA. YUNNAN: Western flank of the Shweli-
Salwin divide, 25°40'N, 9000 ft, June 1919, G. For rest 18021 (HOLOTYPE: K, photo!).
Gaultheria fragrantissima Wall. var. racemosa Hook.f. ex C.B.Clarke (1882: 458). TYPE: INDIA: Darjeeling, 1849, J.D.
Hooker s.n. (HOLOTYPE: K, photo!). Figs. 1 & 4A–B.
FIGURE 1. Gaultheria stapfiana Airy Shaw. A. habit; B. flower; C. ovary (t.s.); D–E. bracts; F–G. bracteoles; H. corolla (inside); I–J.
calyx lobes; K. corolla lobe; L–M. pistils; N–S. stamens. — Scale bars: A = 1 cm; B = 2 mm; C, D, E, F, G, H, I, J, L, M = 1 mm; K =
0.5 mm; N–S = 1 mm (A–S: drawn from S. Panda 29906). Drawn by S. Panda.
Shrubs, stout, erect, dwarf, 0.3–2 m high, often hanging down from rock crevices. Stem terete, blackish-
brown, profusely branched, glabrous; branchlets terete, light green, sparsely hispid-setose non-glandular
multicellular hairs. Leaves alternate; petioles stout, 3–8 mm long, sparsely setulose non-glandular
multicellular hairs up to 3 mm long; lamina oblong-obovate, oblanceolate, narrowly elliptic to obovate, 3–10
× 1–2.5 cm, serrulate at margin, narrowly cuneate at base, mucronate at apex, dark green, glabrous above,
light green, punctate beneath; venation conspicuous brochidodromous with 4–5 pairs lateral veins. Racemes
axillary, rarely pseudoterminal, perulate; rachis light green, 2.5–7(–11) cm long, 10–24-flowered, puberulous.
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Flowers 6–11 mm long; pedicels 2.5–5 mm long, white puberulous to rarely glabrous; bract 1, basal, light
green, ovate-elliptic to elliptic, 3–5 × 1.5–2.5 mm, ciliolate at margin, acute at apex, puberulous; bracteoles 2,
opposite, light green with pinkish stripes, median on pedicel, ovate-elliptic to elliptic, 2–3 × 1–1.5 mm,
ciliolate at margin, acute at apex, puberulous; calyx lobes light pink to white, ovate-triangular to elliptic, ca. 2
× 1.5 mm, ciliolate at margin, acute at apex, glabrous outside, puberulous inside; corolla globose to tubulo-
urceolate, grayish-white, 4–5 × 2–3 mm, glabrous outside, pilose inside, minutely lobed; stamens 10, 1–2 mm
long, loosely epipetalous with grayish-white filaments up to 1.5 mm long, slender, pilose or glabrous, dilated
at base; anthers brown, oblong, up to 1 mm long, glabrous, each lobe with 2 unequal or equal warty apical
awns; pistil 2.5–3.5 mm long; ovary globose, light green, ca. 1 mm long, ca. 1.5 mm diameter, sparsely to
densely white tomentose, the placentation axile, the ovules numerous in each locule; nectar disc minutely 10-
dentate; style light green, ca. 2.5 mm long, slender, glabrous; stigma truncate. Capsule loculicidally 5-valved,
enclosed in a fleshy, bluish-purple accrescent calyx, ovoid, ca. 7 × 5 mm, with 2–4 mm long glabrous pedicel.
Seeds numerous, minute, obconical, scariose.
Distribution.—India: Eastern Himalaya (Sikkim, West Bengal and Arunachal Pradesh); SC China and N
Myanmar.
Habitat.—This is a rare species grown in discontinuous patches in moist humus-covered rocky slopes,
rarely in loose humus-covered boulders, often hanging down from rock crevices in association with
Gaultheria hookeri, G. tetramera and G. semi-infera at altitudes ranging from (1600–) 2200–3500 m.
Flowering.—May–June.
Fruiting.—July–September.
Vernacular name.—Sikkim: Pochu (Lepchas of Chhaten).
Specimens examined in India.—Arunachal Pradesh: Simbi Hotspring, Lohit district, 14 Sep 1994,
Haridasan 7050 (APFH). Sikkim: East district: Zuluk, 28 Jul 1985, D.C.S. Raju 3999 (BSHC); North district:
Zemu Valley, 9500 ft, 9 Jul 1909, Smith & Cave 1026 (CAL); Yumthang, 1901, Dr. Prain’s Collector 365
(CAL); Lachen to Chhaten, 8000 ft, 8 May 2001, S. Panda 29911 (CAL); Yangdin, Lachen to Thangu, 10000
ft, 31 May 2002, S. Panda 29975 (CAL); between Chhaten & Lachen, 2550 m, 7 Jun 1999, D. Maity 21304
(BSHC); without precise locality, 7000 ft, J.D. Hooker s.n., acc. no. 265849 (CAL). West Bengal
(Darjeeling): near Sonatek, 5000 ft, Jul 1906, G.H. Cave s.n., acc. no. 6630 (Lloyd Bot. Garden Herbarium,
Darjeeling); Tonglu, 9000 ft, 6 Dec 1924, G.H. Cave s.n. (CAL); Jalapahar, 7500 ft, 16 Dec 1910, G.H. Cave
4865 (CAL).
2. Gaultheria hookeri C.B.Clarke (1882: 458). TYPE: INDIA. SIKKIM: Lachen, 10000 ft, 3.6.1849, J. D.
Hooker s.n. (top “in flowers”)(HOLOTYPE: K, photo!).
G. hookeri var. angustifolia C.B.Clarke (1882: 458). TYPE: INDIA. SIKKIM: YakLa, 9000 ft, 10.05.1876, C. B. Clarke
27837A (HOLOTYPE: K!; ISOTYPE: CAL!).
G. veitchiana Craib (1912: 188). TYPE: W CHINA. 6000–10000 ft, May, 1904, E. H. Wilson 3916 (HOLOTYPE: K,
photo!). Figs. 2 & 4C–D.
Shrubs, stout, erect, bushy dwarf, 0.3–1 m high, often hanging down from rock crevices. Stem terete,
blackish-brown, profusely branched, sparsely setose; branchlets terete, light green to pink, densely hispid-
setose. Leaves alternate; petioles stout, 3–6 mm long, glabrous or punctuate; lamina oblong-elliptic, oblong to
rarely oblanceolate or obovate, 2.5–5.5 (– 7.5) × 1.5–3 cm, serrulate-ciliate at margin, cilia deciduous and up
to 2 mm long, broadly cuneate at base, mucronate at apex, dark green above, light green, setulose and punctate
beneath, venation conspicuous brochidodromous with 4–5 pairs lateral veins. Racemes axillary to rarely
pseudoterminal, perulate; rachis light green, (1.5 –) 2–5 cm long, 10–15-flowered, densely white puberulous.
Flowers 7–14 mm long; pedicels greyish-white, (2 –) 4–7 mm long, white puberulous to rarely glabrous;
bract 1, basal, light green with pinkish stripes, broadly ovate or ovate-elliptic to rarely lanceolate, 5–7 × 3–4.5
mm, ciliolate at margin, acute to rarely acuminate at apex, sparsely puberulous or glabrous; bracteoles 2,
opposite, subbasal or median on pedicel, greyish-white with pinkish stripes, broadly ovate or elliptic to
Phytotaxa 50 © 2012 Magnolia Press • 7
GAULTHERIA STAPFIANA (ERICACEAE)
ovate-triangular, 2–3.5 × 1–1.5 mm, ciliolate at margin, acuminate to acute at apex, glabrous; calyx lobes
greyish-white to light pink, ovate-triangular or elliptic to rarely ovate-oblong, c. 2 × 1.5 mm, ciliolate at
margin, acuminate at apex, glabrous to pilose inside; corolla ovoid-urceolate, globose-urceolate to tubular-
urceolate, greyish-white to light pink, 5–7 × 2–4.5 mm, glabrous outside, pilose inside, lobes minute; stamens
10, 2–3 mm long, loosely epipetalous; filaments pinkish, 1–2 mm long, slender, papillose, pilose to glabrous,
dilated at base; anthers orange brown, oblong, c. 1 mm long, glabrous, each lobe with 2 equal minute warty
apical awns of 0.5–1 mm long; pistil 2.5–4.5 mm long; ovary globose to subglobose, light green, (0.5 –) 1–2 ×
1–2 mm, tomentose, ovules numerous on axile placenta in each locule; disc dark green, minutely 10-oblong-
dentate; style pinkish, 2–3 mm long, slender, glabrous; stigma truncate.axillary, rarely pseudoterminal,
perulate; rachis light green, 2.5–7(–11) cm long, 10–24-flowered, puberulous. Capsule loculicidally 5-valved,
enclosed in a fleshy accrescent calyx, globose to subglobose, sky blue, 9–11 × 9 mm including c. 2 mm long
persistent style with c. 5 mm long glabrous pedicel, tomentose. Seeds numerous, minute, obconical, scariose.
FIGURE 2. Gaultheria hookeri C.B. Clarke. A. habit; B. flower; C–D. bracts; E–G. bracteoles; H–I. calyx lobes; J. corolla lobe; K–L.
stamens; M. pistil; N. ovary (t. s.). — Scale bars: A = 1 cm; B = 2 mm; C–I, K–N = 1 mm; J = 0.5 mm (A–N: drawn from S. Panda
30872, CAL). Drawn by S. Panda.
Distribution.—India: Eastern Himalaya (Sikkim, West Bengal and Arunachal Pradesh) and NE India
(Nagaland), E Nepal, Bhutan, SW China and N Myanmar.
Habitat.—This is a rare species grown in discontinuous patches in moist humus-covered rocky slopes and
in loose humus-covered boulders, often hanging down from rock crevices in association with other ericaceous
species viz. Gaultheria nummularioides, G. tetramera, G. stapfiana, G. semi-infera, Pieris formosa, Vaccinium
nummularia, Rhododendron barbatum and R. cinnabarinum at altitudes ranging from 2400–3600 m.
Flowering.—April–June.
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Fruiting.—June–August.
Vernacular name.—Sikkim: Kalum, Kalumba (Bhutias of Lachen).
Specimens examined in India.—Nagaland: Saramati ridge, Tuensang district, 2800 m, 2.4.2003, S. Panda
30872 (CAL); Kohima district, Japhu Hill, 9900 ft, 25.10.1885, C. B. Clarke 41339 (ASSAM). Sikkim: East
district, Yak La, 10000 ft, 10.05.1876, C. B. Clarke 27850 (CAL); North district, Lachen, 9000 ft, May, 1885,
Dr. King’s Collector s.n., acc. no. 265793 (CAL); Tangkar Mt., 11500 ft, 01.08.1892, G. A. Gamm ie 480
(CAL); On the bank of Lachung Chhu River, 8000 ft, 01.05.2001, S. Panda 30057 (CAL); Lachung to
Yumthang, 10000 ft, 02.05.2001, S. Panda 30061 (CAL); Yumthang to Lachung, 11000 ft, 02.05.2001, S.
Panda 30063 (CAL); Lachen to Thangu, 9000 m, 07.05.2001, S. Panda 29903 (CAL); Yumthang to Lachung,
26.05.1986, S. K. Rai & S. Pradhan 5571 (BSHC); West district, Tsoka to Dzongri, 11000 ft, 14.05.2002, S.
Panda 29941 (CAL). West Bengal (Darjeeling): Above Dali, near Darjeeling town, 2150 m, 16.11.1980, A.
P. D as 309 (North Bengal University Herbarium, Siliguri).
3. Gaultheria fragrantissima Wall. (1820: 397). TYPE: NEPAL. 1818, Wallich s.n. (HOLOTYPE: K,
photo!).
Arbutus laurifolia Buch.-Ham. ex D.Don (1825: 151). TYPE: NEPAL. NARAINHETTY: March 10, 1803, Buchanan-
Hamilton s.n. (HOLOTYPE: BM, n.v.).
Gaultheria ovalifolia Wall. (1829: 1523). TYPE: INDIA. TAMILNADU: Nilgiri hills (“Neilgherry”), October 14, 1825,
Noton in Wall. list no. 1523 (HOLOTYPE: CAL!).
G. leschenaultii DC. (1839: 593). TYPE: INDIA. TAMILNADU: Nilgiri hills, Leschenault s.n. (HOLOTYPE: P, n.v.).
Leucothoe katagherensis DC. (1839: 606). Andromeda katagherensis (DC.) Hook. (1840: 246). TYPE: INDIA.
TAMILNADU: Nilgiri hills, Kotagiri, L. B. E. Schmid s.n. (HOLOTYPE: W, n.v.).
Gaultheria forrestii Diels (1912: 210). TYPE: CHINA. SE XIZANG: Tali range, 25° 40´N, 10000–12000 ft, July, 1906,
G. Forrest 4183 (HOLOTYPE: K, photo!). Figs. 3 & 4E–F.
Shrubs to treelets, stout, erect, 0.5–3.5 m high, often hanging down from rock crevices. Stem terete, grey-
brown, light green to light brown, profusely branched, glabrous; branchlets winged to triangular, blood red to
dark pink to rarely light green, glabrous, glaucous. Leaves alternate; petioles stout, 4–12 mm long, glabrous or
punctuate beneath; lamina ovate-elliptic, oblong-elliptic, ovate-lanceolate to rarely ovate or obovate, 4–10 (–
14) × 1.8–4 (– 8) cm, serrate to serrulate at margin, broadly cuneate at base, mucronate at apex, deep green,
glabrous above, light green, punctate beneath; venation conspicuous brochidodromous with 3–5 pairs lateral
veins. Racemes axillary, rachis 2.5–8 (– 11) cm long, 10–20-flowered, puberulous. Flowers 8–14 mm long;
pedicels greyish-white, 2–8 mm long, puberulous; bract 1, basal, light green, broadly ovate, 2–4 × 1.5–2.5
mm long, ciliate at margin, acute or subacute at apex, glabrous; bracteoles 2, opposite, apical or subapical on
pedicel, pink with greenish stripes, broadly ovate, 1.5–2.5 × 1–2 mm, ciliolate at margin, acute at apex,
puberulous; calyx lobes light pink, ovate-triangular, 2–2.5 × 1–1.5 mm, ciliolate at margin, acute at apex,
puberulous inside; corolla ovoid-urceolate, greyish-white to light greenish-white, c. 5 mm long, 3–4 mm
across, glabrous outside, pilose inside, lobes minute, c. 0.5 × 1 mm; stamens 10, c. 3 mm long, loosely
epipetalous; filaments greyish-white, c. 1 mm long, papillose, pilose, basally dilated; anther lobes blackish-
brown, oblong, c. 1 mm long, glabrous, each lobe with 2 apical awns of c. 1 mm long or minute. Pistil 2–3.5
mm long; ovary globose to subglobose, light green, 1–1.5 × 1.5–2 mm, tomentose, ovules numerous on axile
placenta in each locule; disc minutely 10-dentate; style light green, 1–2 mm long, slender, glabrous; stigma
truncate. Capsule loculicidally 5-valved, enclosed in a fleshy accrescent calyx, light green (immature) to deep
blue or sky blue (mature), globose to subglobose, 5–8 × 3–6 mm including 2–4 mm long pedicel, pubescent.
Seeds numerous, minute, obconical, scariose.
Distribution.—India: Eastern Himalaya (Sikkim, West Bengal and Arunachal Pradesh), NE States
(Meghalaya, Nagaland and Manipur) and Hill-tops of SW Ghats (Tamil Nadu, Kerala and Karnataka); E
Nepal, Bhutan, SC China, N Myanmar, Sri Lanka, N Vietnam and Malesia.
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FIGURE 3. Gaultheria fragrantissima Wall. A. habit; B. flower; C–D. bracts; E–F. bracteoles; G–H. calyx lobes; I. corolla lobe; J–K.
stamens; L. pistil; M. ovule; N–O. ovary (t. s.); P–Q. fruits; R. seeds. — Scale bars: A = 1 cm; B, P = 2 mm; C–H, J–L, N, O = 1 mm;
Q = 5 mm (A–R: drawn from S. Panda 30701).
Habitat.—This species grows gregariously in moist rocky soil, often hanging down from rock crevices,
rarely in loose humus-covered boulders or in landslide areas in association with other ericaceous species viz.
Gaultheria nummularioides, Lyonia ovalifolia, Leucothoe griffithiana, V. dunalianum, Rhododendron
formosum and R. virgatum at altitudes ranging from (1350–) 1600–2300 (2700) m.
Flowering.—March–May; December–January.
Fruiting.—May–October.
Vernacular name.—Sikkim: Kaloma (Tibetans of East district), Jathyroid (Sherpas), Machino,
Dhasingare (Nepalese of Yuksum), Goenhli (Nepalese of Hilley), Kalomba (Lepchas of Lachung). West
Bengal: Chanchhewaa (Tamang Nepalese of Darjeeling). Arunachal Pradesh: Shep-Sheng (Moplas of
Tawang). Meghalaya: Jirhapkynthai, Jirhap, Soh-lyngthrait, Dieng-la-sirhap, Jirhapiong (Khasis of
Shillong), Lathynrait (Khasis of Jongsha Village). Tamil Nadu: Kolakkaai, Moolai (Tamilese of Nilgiri hills).
Specimens examined in India.—Arunachal Pradesh: Dibang Valley district, Myodia to Tewarigaon,
17.11.2000, D. K. Singh & Party 97502 (BSD); Lohit district, Walong to Helmet top, 1450 m, 28.04.2003, S.
PANDA10 • Phytotaxa 50 © 2012 Magnolia Press
Panda 30886 (CAL); Lower Subansiri district, Pange to Talle Valley, 2500–2800 m, 31.12.2002, S. Panda
3084 (CAL); Tawang district, Moukto, 7000 ft, August, 2001, B. Balodi 10238 (ARUN); West Kameng
district, Rupa to Jabrang, 23.05.1958, G. Panigrahi 16046 (ASSAM). Kerala: Sispara to Palghat, Palakkad
district, 2000 m, 28.03.1983, N. C. Nair 77213 (CAL); Devicolam, Idduki district, 6000 ft, December, 1909,
A. Meebold 13337 (CAL). Meghalaya: East Khasi Hill district, Shillong Peak, 5000 ft, 13.03.1970, R. P. Patil
21011 (ASSAM); Shillong Peak, 5500 ft, 08.03.1885, C. B. Clarke 37496 (CAL); Shillong Peak, 5000 ft,
17.03.2002, S. Panda 30703 (CAL). Manipur: Mao, Senapati district, 11.02.1958, D. B. Deb 1536 (CAL).
Nagaland: Pegwima to Japhu Hill, 6000 ft, 19.05.1882, G. Watt 6873 (CAL). Sikkim: Yuksum to Bakhim,
West district, 6000 ft, 12.05.2002, S. Panda 29928 (CAL); Phamtam, South district, 10.05.1991, R. C.
Srivastava 13162 (BSHC). Tamil Nadu: Dindigul district, Bearshola, Kodaikanal, 2233 m, 10.03.1958, K.
Subramanyam 5555 (CAL); Nilgiris district, Bison Camp, 7000 ft, 03.02.1911, C. E. C. Fischer 253 (CAL).
West Bengal (Darjeeling): Sonada, 7000 ft, 19.05.1913, G. H. Cave s.n., acc. no. 265787 (CAL).
Acknowledgements
The author is grateful to Dr. M. Sanjappa, former Director, Botanical Survey of India for awarding a research
fellowship. Thanks are also due to Curators in CAL, BSIS, ASSAM, DD, MH for providing all facilities and
permission to consult herbarium specimens. The author is also grateful to James L. Reveal (BH) for kindly
reviewed the manuscript.
FIGURE 4. A–B. Gaultheria stapfiana Airy Shaw (A. live,
B. cibachrome type photo); C–D. G. hookeri C.B. Clarke (live
and cibachrome type photo); E–F. G. fragrantissima Wall (live
and cibachrome type photo.
FIGURE 5. A–B. Stomatal complex of Gaultheria stapfiana
(40X); C–D. Stomatal complex of G. hookeri (40X); E–F. Sto-
matal complex of G. fragrantissima (40X).
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GAULTHERIA STAPFIANA (ERICACEAE)
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