Taiwania, 57(4): 384–390, 2012 NOTE Studies on Oil-bodies in Some Liverworts from Sikkim, Eastern Himalaya, India Devendra Singh(1), Monalisa Dey(1) and D. K. Singh(2*) 1. Botanical Survey of India, Central National Herbarium, Howrah-711103, India. 2. Botanical Survey of India, CGO Complex, 3rd MSO Building, Block F (5th floor), Salt Lake Sector I, Kolkata-700 064, India. * Corresponding author. Telfax: +91 33 23214050; Email: [email protected] (Manuscript received 19 April 2012; accepted 10 May 2012) ABSTRACT: Oil-bodies in 17 species of leafy liverworts from Sikkim in Eastern Himalaya, belonging to order Jungermanniales, are described for the first time in Indian bryoflora. KEY WORDS: Eastern Himalaya, India, Liverworts, Oil-bodies, Sikkim. INTRODUCTION The plants were collected from different localities of North and West Sikkim during the months of November The studies on oil-bodies in Indian liverworts have 2009, July 2010 and May–June 2011 at altitudes ranging received considerable attention and so far 166 taxa between 1350–4000 m above mean sea level. spreading across 61 genera in 28 families of order Photomicrographs of the oil-bodies were taken within 10–15 Haplomitriales, Jungermanniales, Metzgeriales and days of their collection under Biological Research Marchantiales have been investigated for this feature Microscope (Olympus-CX 41) with the help of Olympus (Udar et al., 1970, 1978, 1982; Udar and Nath, 1971, Camedia C-7070 digital camera. All specimens have been 1976, 1977, 1978, 1979, 1981; D. Kumar and Udar, deposited in the Cryptogamic section of the Central National 1976; Udar and Shaheen, 1982; Udar and A.Kumar, Herbarium of Botanical Survey of India, Howrah (CAL). 1981, 1982a, b, c, d, 1983; Udar and Awasthi, 1981, 1982a, b, 1983, 1984a, b; Udar and Dh. Kumar, 1983; RESULTS Awasthi and Udar, 1984; Nath and Udar, 1984; A.Kumar and Udar, 1985; A.Kumar and Srivastava, Lepidoziaceae 1986; Awasthi and Srivastava, 1988; Srivastava and Sharma, 1990; Sharma and Srivastava, 1993; Srivastava 1. Bazzania himalayana (Mitt.) Schiffn. Fig. 1A and Srivastava, 1994; Asthana et al., 1995; Awasthi et al., 2000; Srivastava and Srivastava, 2002; Asthana and Oil-bodies grayish, 4–6 per leaf cell, elongated, Srivastava, 2003; D. Singh and Singh; 2006; Dey et al., ellipsoidal to more or less clavate, 6–12 × 4–5 μm, 2008; D. Singh et al., 2008a, b; Singh and D. Singh, smooth, sometimes digitately 2–3 (–4) segmented, with 2009; S.K. Singh and Singh, 2009; D. Singh et al., smooth segments. 2010; Dey and Singh, 2011; D. Singh, 2012; D. Singh Specimen examined: INDIA, Eastern Himalaya, South and Singh, 2012). The present paper deals with the Sikkim, Maenam Wildlife Sanctuary, ca. 3300 m, 04.06.2011, D. oil-bodies in yet another 17 species of jungermannioid Singh 51796. liverworts from Sikkim in Eastern Himalaya, India. Of 2. Bazzania imbricata (Mitt.) S.Hatt. Fig. 1B these, while oil-bodies in Anastrepta orcadensis (Hook.) Schiffn., Anastrophyllum joergensenii Schiffn., Oil-bodies grayish, 3–5 per leaf cell, A. minutum (Schreb.) R.M. Schust. and Mastigophora ovoid-ellipsoidal, 7.0–10.0 × 5.0–7.0 μm, rarely woodsii (Hook.) Nees have been studied earlier in extra spherical, 4.0–6.0μm in diameter, coarsely segmented, Indian materials (Hattori, 1953; Kitagawa, 1966; grape-cluster like. Schuster, 1969; Kis and Pócs 1997; Paton, 1999; Schill Specimen examined: INDIA, Eastern Himalaya, North and Long, 2003), the remaining have been investigated Sikkim, 12 km from Lachung towards Katau, ca. 3400 m, 24.05.2011, for the first time for their oil-bodies. D. Singh 52139. MATERIALS AND METHODS 3. Bazzania sikkimensis (Steph.) Herzog Fig. 1C 384 December, 2012 Singh et al.: Studies on oil-bodies in some liverworts from Sikkim Oil-bodies grayish, 5–9 per leaf cell, usually Oil-bodies grayish, 3–4per leaf cell, ovoid-ellipsoidal, ovoid-ellipsoidal, 6.0–11.0 × 4.0–5.0 μm, rarely 4.0–6.0 × 2.5–4.0 μm, coarsely segmented. spherical, 3.0–5.0μm in diameter, smooth, often Specimen examined: INDIA, Eastern Himalaya, North Sikkim, digitately 2–3 segmented with smooth segments like 12 km from Lachung towards Katau, ca. 3400 m, 24.05.2011, D. Singh those in B. himalayana. 52129. Schill and Long (2003) reported up to 7, coarsely Specimen examined: INDIA, Eastern Himalaya, West Sikkim, segmented oil-bodies per leaf cell in this species from Barsey Rhododendron Sanctuary (Barsey), ca. 2875 m, 16.07.2010, D. Singh 48999. Scotland. The Indian species of the genus can be categorized into three groups based on the characteristics of their 7. Anastrophyllum minutum (Schreb.) R.M.Schust. oil-bodies. The first group comprises B. himalayana and Fig. 1G B. sikkimensis, which have ellipsoid to more or less clavate oil-bodies with smooth surface, but often Oil-bodies grayish, 6–10 per leaf cell, digitately 2–3 (–4) segmented. The other group, ovoid-ellipsoidal, 6.0–13.0 × 5.0–8.0 μm, or spherical, represented by B. imbricata, B. sumbavensis and B. 5.0–9.0 μm in diameter, coarsely segmented. tridens, exhibit ovoid-broadly ellipsoidal, finely-coarsely Specimen examined: INDIA, Eastern Himalaya, North segmented grape-cluster like oil-bodies. Whereas, in B. Sikkim, 12 km from Lachung towards Katau, ca. 3400 m, 24.05.2011, tricrenata oil-bodies are spherical, subspherical to broadly D. Singh 52146. ovoid and smooth. While majority of the Indian species Schuster (1969) and Schill and Long (2003) have 3–9 oil-bodies in each leaf cell, B. sumbavensis and reported 2–6 oil-bodies in this species from Scotland, B. tricrenata show 7–15 and 2–4 oil-bodies per cell whereas Kis and Pócs (1997) reported only 1–4 respectively (Sharma and Srivastava, 1993; D. Singh et oil-bodies in the specimens from Africa. al., 2008a). 8. Scapania ligulata Steph. Fig. 1H Calypogeiaceae Oil-bodies grayish, (2–) 3–4 per leaf cell, ovoid, 4–8 4. Calypogeia aeruginosa Mitt. Fig. 1D × 3–6 μm, or spherical, 3–5μm in diameter, coarsely segmented. Oil-bodies bluish, 2–4 per leaf cell, Specimen examined: INDIA, Eastern Himalaya, West ovoid-ellipsoidal, 8.0–11.0 × 3.5–5.0 μm, or spherical, Sikkim, Barsey Rhododendron Sanctuary (4 km from Anden towards 4.0–6.0 μm in diameter, finely segmented. Okhrey), ca. 2260 m, 13.07.2010, D. Singh 48911. Specimen examined: INDIA, Eastern Himalaya, North The oil-bodies in the Indian species of the genus are Sikkim, 12 km from Lachung towards Katau, ca. 3400 m, 24.05.2011, usually similar in shape, but show little variation in D. Singh 52152. number per cell from 2–3 in S. contorta and S. All the Indian species of the genus investigated so ornithopodioides to 3–7 in S. griffithii (Srivastava and far for this feature, viz. C. aeruginosa, C. arguta, C. Srivastava, 1994; D. Singh et al., 2008a, b). azurea, C. lunata and C. marginella, show similar ovoid-ellipsoidal or spherical oil-bodies with Plagiochilaceae characteristic blue colour or the bluish tinge (Sharma and Srivastava, 1993; D. Singh et al., 2008a). 9. Plagiochila denticulata Mitt. Fig. 1I Scapaniaceae Oil-bodies grayish, 4–6 per leaf cell, fusiform-ellipsoidal, 8.0–18.0 × 4.0–5.0 μm, usually 5. Anastrepta orcadensis (Hook.) Schiffn. Fig. 1E curved, finely segmented. Oil-bodies grayish, 4–8 per leaf cell, usually Specimen examined: INDIA, Eastern Himalaya, West spherical, 3.0–5.0 μm in diameter, or sometimes Sikkim, Barsey Rhododendron Sanctuary (Fedi), ca. 2900 m, 08.06.2011, D. Singh 51827. ovoid-ellipsoidal, 4.0–6.0 × 3.0–4.0 μm, nearly smooth. The characteristically curved, larger oil-bodies in Specimen examined: INDIA, Eastern Himalaya, North this species make it easily distinguishable from other Sikkim, 12 km from Lachung towards Katau, ca. 3400 m, 24.05.2011, Indian species of the genus, which usually show D. Singh 52126. Hattori (1953) and Kitagawa (1966) also reported ovoid-ellipsoidal or spherical oil-bodies (Udar et al., 7–9 and 6–10 oil-bodies per leaf cell respectively with 1970; Udar and Nath, 1976; Udar and Shaheen, 1982; nearly smooth texture in this species from Japan. D. Singh and Singh; 2006; D. Singh et al., 2008a). 6. Anastrophyllum joergensenii Schiffn. Fig. 1F 10. Plagiochila detecta M.L.So & Grolle Fig. 2A 385 Taiwania Vol. 57, No. 4 Oil-bodies grayish, 5–10 per leaf cell, spherical, 3–5 μm in diameter, or sometimes ovoid-ellipsoidal, 4.0–6.0 × 2.5–3.5μm, or sometimes ellipsoidal-fusiform, 5.0–8.0 × 3.0–4.5 μm, finely spherical, 2.5–3.5μm in diameter, coarsely segmented. segmented. Specimen examined: INDIA, Eastern Himalaya, West Specimen examined: INDIA, Eastern Himalaya, South Sikkim, Barsey Rhododendron Sanctuary (2 km from Barsey towards Sikkim, Maenam Wildlife Sanctuary, ca. 3300 m, 01.06.2011, D. Hee), ca. 2600 m, 16.07.2010, D. Singh 49043C. Singh 52173B. Oil-bodies in the Indian species of this genus are of 11. Plagiochila flexuosa Mitt. Fig. 2B two types, viz. smooth, usually spherical or rarely ellipsoidal-fusiform, as in the case of C. bhutanica, C. Oil-bodies yellowish, 4–8 per leaf cell, usually furcilobulata, C. udarii (Asthana and Srivastava, 2003; ovoid-ellipsoidal, 9–14 × 5–8 μm, or sometimes Dey and Singh, 2011), and finely segmented , usually spherical, 4–8 μm in diameter, smooth. ellipsoidal-fusiform or spherical as in C. latilobula, C. Specimen examined: INDIA, Eastern Himalaya, West longifolia, C. kashyapii, C. planissima and C. Sikkim, Barsey Rhododendron Sanctuary (Fedi), ca. 2900 m, minutissima (Asthana and Srivastava, 2003; D. Singh et 08.06.2011, D. Singh 51823. al., 2008a). However, C. dozyana is interesting in this In the presence of yellowish and smooth oil-bodies regard as it combines the features of both the groups in of this species, it is easily recognizable amongst other the presence of usually spherical yet finely segmented Indian species of the genus which have grayish, oil-bodies. Most of the species have 3–22 oil-bodies per finely-coarsely segmented oil-bodies (Udar et al., 1970; cell, except C. minutissima and C. bhutanica, which Udar and Nath, 1976; Udar and Shaheen, 1982; D. show 2–3 and 4–6 oil-bodies per cell respectively. Singh and Singh; 2006; D. Singh et al., 2008a). 15. Drepanolejeunea longii Grolle & R.L.Zhu Mastigophoraceae Figs. 2F & G Oil-bodies grayish, 3–5 per leaf cell in the middle of 12. Mastigophora woodsii (Hook.) Nees Fig. 2C leaf lobe, spherical, 4–6 μm in diameter; up to 6 in basal leaf cells, spherical 4–6 μm in diameter or ellipsoidal, Oil-bodies grayish, 3–8 per leaf cell, 5–8 × 4–5μm, coarsely segmented. ovoid-ellipsoidal, 5.0–9.0 × 2.5–5.0 μm, or spherical, Specimen examined: INDIA, Eastern Himalaya, South 3.0–4.5 μm in diameter, very finely segmented. Sikkim, Maenam Wildlife Sanctuary, ca. 3200 m, 31.05.2011, D. Specimen examined: INDIA, Eastern Himalaya, North Singh 51718A. Sikkim, 12 km from Lachung towards Katau, ca. 3400 m, 24.05.2011, So far, only D. erecta and D. yunnanensis were D. Singh 52156. studied in India for their oil-bodies (Udar and Awasthi, Paton (1999) also reported up to 8, finely segmented 1984; D. Singh et al., 2008a), and both are more or less oil-bodies in this species from British Isles. similar to D. longii in this feature. Lejeuneaceae 16. Lejeunea cocoes Mitt. Fig. 2H 13. Cheilolejeunea subopaca (Mitt.) Mizut. Fig. 2D Oil-bodies grayish, (1–) 2–3 per leaf cell, ovoid-ellipsoidal, 4.0–8.0 × 3.0–4.5 μm, or spherical, 3–4 Oil-bodies grayish, one per leaf cell, ellipsoidal, 12– μm in diameter, coarsely segmented. 20 × 8–11 μm, coarsely segmented. Specimen examined: INDIA, Eastern Himalaya, South Sikkim, Specimen examined: INDIA, Eastern Himalaya, North Maenam Wildlife Sanctuary, ca. 3200 m, 01.06.2011, D. Singh 51729. Sikkim, Dobang valley, ca. 2984 m, 21.05.2011, D. Singh 52009. Unlike C. birmensis, C. laeviscula and C. 17. Lejeunea eifrigii Mizut. Fig. 2I trapezia with 1–2 oil-bodies per cell; C. serpentina with 1–4 oil-bodies and C. trifaria with 2–3 oil-bodies Oil-bodies grayish, (1–) 2–3 per leaf cell, spherical, (Asthana et al., 1995; D. Singh et al., 2008a; D. Singh, 4–5 μm in diameter, or ovoid-ellipsoidal, rarely 2012), C. subopaca possesses a single oil-body only in fusiform, 5–8 × 3–5 μm, coarsely segmented. each cell. Specimen examined: INDIA, Eastern Himalaya, North Sikkim, Chungthang (along the Lachung Chhu), ca. 2050 m, 14. Cololejeunea dozyana (Sande Lac.) Schiffn. 23.11.2009, D. Singh 46680. Fig. 2E Except L. kashyapii and L. obscura, which show 18–35 and 40–55, smooth oil-bodies per cell Oil-bodies grayish, 4–6 per leaf cell, usually respectively, the other Indian species of the genus have segmented oil-bodies which vary in number from 2–3 386 December, 2012 Singh et al.: Studies on oil-bodies in some liverworts from Sikkim per cell in L. cocoes, L. eifrigii and L. punctiformis to Lophoziaceae) in the Himalayan region and Western 2–4 in L. wallichiana, 2–6 in L. indica, 4–6 in L. flava, China. J. Hattori Bot. Lab. 94: 115–157. 3–8 in L. tuberculosa and 3–12 in L. nepalensis (Udar Schuster, R.M. 1969. The Hepaticae and Anthocerotae of North America (East of the Hundredth Meridian) II. and Awasthi, 1981; Udar and Shaheen, 1982; D. Singh Columbia University Press, New York. pp. 1–1062. and Singh, 2006; Dey et al., 2008; D. Singh et al., Srivastava, S. C. and D. Sharma. 1990. A new species of 2008a). Jubula from Milam in Kumaon (Western Himalaya). Proc. Indian Acad. Sci. Pl. Sci. 100: 85–89. ACKNOWLEDGEMENTS Sharma, D. and S. C. Srivastava. 1993. Indian Lepidoziineae (A taxonomic revision). Bryoph. Biblioth. The authors are thankful to the Chief Wildlife Warden, 47:1–353. Sikkim Forest Department for facilitating the exploration work Singh, D. 2012. A note on Cheilolejeunea trifaria (Reinw. et in Sikkim. One of us (MD) is also thankful to the Ministry of al.) Mizut. from Indian Himalayan region. Lindbergia 35: Environment and Forests, New Delhi for financial assistance 18–21. under the All India Coordinated Project on Taxonomy Singh, D. and D. K. Singh. 2006. Studies on oil-bodies in (AICOPTAX). some foliose liverworts from East Sikkim. Indian J. Forest. 29: 457–461. LITERATURE CITED Singh, D. and D. K. Singh. 2012. A new species of the genus Lepidozia (Marchantiophyta, Lepidoziaceae) from Sikkim, India. Nova Hedwigia 94: 221–225. Asthana, G. and S. C. Srivastava. 2003. Indian Singh, D., M. Dey and D. K. Singh. 2008a. Studies on Cololejeunea - A taxonomic study. Bryoph. Biblioth. 60: oil-bodies in some foliose liverworts from East Sikkim II. 1–155. Indian J. Forest. 31: 315–320. Asthana, G., S. C. Srivastava and A. K. Asthana. 1995. The Singh, D., D. K. Singh and M. P. Singh. 2008b. Scapania genus Cheilolejeunea in India. Lindbergia 20: 125–143. contorta Mitt. (Subgen.: Scapania, Sect., Awasthi, U. S. and S. C. Srivastava. 1988. Status of Plicaticalyx) with a note on diversity and distribution of Brachiolejeunea (Spruce) Schiffn. in India. Proc. Indian the genus in India. Indian J. Forest. 31: 151–158. Acad. Sci. Pl. Sci. 98: 1–12. Singh, D., M. Dey and D. K. Singh. 2010. Leptolejeunea Awasthi, U. S., S. C. Srivastava and D. Sharma. 2000. apiculata (Horik.) 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Botanical Survey of India, Central National Herbarium, Howrah-711103, India. 2. Botanical Survey of India, CGO Complex, 3rd MSO Building, Block F (5th floor), Salt Lake Sector I, Kolkata-700 064, India. * 通信作者。Telefax: +91 33 23214050; Email: [email protected] （收稿日期：2012年4月19日；接受日期：2012年5月10日） 摘要：本文研究17種在錫金（東西馬拉雅山脈）的葉蘚目物種之油滴體，為印度苔蘚植物 誌之首次描述。 關鍵詞：東喜馬拉雅山脈、印度、蘚類、油滴體、錫金。 388 December, 2012 Singh et al.: Studies on oil-bodies in some liverworts from Sikkim Fig.1. Photomicrographs of oil bodies. A: Bazzania himalayana (Mitt.) Schiffn. B: Bazzania imbricata (Mitt.) S.Hatt. C: Bazzania sikkimensis (Steph.) Herzog. D: Calypogeia aeruginosa Mitt. E: Anastrepta orcadensis (Hook.) Schiffn. F: Anastrophyllum joergensenii Schiffn. G: Anastrophyllum minutum (Schreb.) R.M.Schust. H: Scapania ligulata Steph. I: Plagiochila denticulata Mitt. 389 Taiwania Vol. 57, No. 4 Fig. 2. Photomicrographs of oil bodies. A: Plagiochila detecta M.L.So & Grolle. B: Plagiochila flexuosa Mitt. C: Mastigophora woodsii (Hook.) Nees. D: Cheilolejeunea subopaca (Mitt.) Mizut. E: Cololejeunea dozyana (Sande Lac.) Schiffn. F: Drepanolejeunea longii Grolle & R.L.Zhu (from middle). G: The same from base. H: Lejeunea cocoes Mitt. I: Lejeunea eifrigii Mizut. 390