Bull. Natn. Sci. M us. , Tokyo, Ser. B, 23(3), pp. 81 - 102, September 23 , 1997

Oil Bodies and Oil Droplets of Some Hepatics from Sichuan, China
Tatsuwo Furuki 1 and Masanobu Higuchi 2
1

Depatment of Plant Sciences, Natural History Museum and Institute, Chiba,955- 2 Aoba-cho, Chuo-ku, Chiba, 260 Japan2
Department of Botany, National Science Museum,4-1 - 1 Amakubo, Tsukuba, Ibaraki, 305 Japan

Abstract The oil bodies and the oil droplets are described for 40 species in 24
genera of hepatics collected from Sichuan, China. Among them the oil bodies of 14
species are described for the first time. Radula gedena Steph. is new to China.
Key words:

oil body, oil droplet, hepatics, Sichuan, China.

The hepatic flora of Sichuan, China, was summarized by Piippo et al. (1997)
who reported 139 species of 55 genera. However, the oil bodies have been scarcely
described for hepatics in Sichuan as well as in China. The present paper deals withthe morphological observations of the oil bodies and the oil droplets of hepatics collected from Sichuan by the second author from August to September, 1996.
Materials and Methods
The materials and methods used for this study follow those of Furuki and
Higuchi ( 1996). The specimens examined are deposited in the herbarium of the National Science Museum, Tokyo (TNS), and duplicates in the herbaria of the Natural
History Museum and Institute, Chiba (CBM) and the Kunming Institute of Botany,
Academia Sinica (HKAS).
Results
The cells may include a greater number of oil bodies and oil droplets than thoserecognizable in the photographs because some oil bodies are out of the focal plane.
Four types of oil bodies were recognized: (l) homogeneous type - oil bodies homogeneous, (2) granulose type - oil bodies composed of minute granules, (3) botryoidal type - oil bodies segmented and composed of distinct granules and (4) absent
type - oil bodies absent. The number of oil droplets is too many to count in many
species, but few in others.
The arrangement offamilies and of genera in each family follows Grolle (1983).
Each species name is followed by the description of the oil bodies and the oil

82

Tatsuwo Furuki and Masanobu Higuchi

droplets, specimen(s) examined and notes.
Jungermanniales
Herbertaceae
Herbertus ad uncus (Dicks.) Gray, Nat. Arr. Brit. PI. I: 705 (1821 ). (PI. I: I, 2)
Oil bodies homogeneous type, 8- 15 per cell of vitta, 1- 5 per submarginal cell ofdisc, globose to oval, 3- 5 X 2--4 Jlm, often turbid, colorless. Oil droplets homogeneous
type, many per cell, spherical, less than 1 Jlm dia., colorless.
1.

Specimen examined. Kangding County, south of Jiagenba, 3250 m, 29°50'N, 10 I 0 30'E, on boulder, 8 Sept. 1996, Higuchi 30609.

Herbertus delavayi Steph., Hedwigia 34: 43 (1895) .
(PI. 1: 3, 4)
Oil bodies homogeneous type, 6- 15 per cell of vitta, 1- 5 (8) per submarginal
cell of disc, globose to oblong, 4- 8 X 3- 6 Jlm, colorless. Oil droplets homogeneous
type, many per cell, spherical, less than 1 Jlm dia., colorless.

2.

Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20 'N,
I 0 I 0 30 'E, on humus, I0 Sept. 1996, Higuchi 30829.

3.

Herbertus dicranus (Tay\.) Trev., Mem. R. Ist. Lombardo Sci . 4: 387 (1877).
(PI. I: 5, 6)
Oil bodies homogeneous type, (3) 5- 10 (15) per cell ofvitta, 1- 10 per submarginal cell of disc, oval to oblong, 2- 7 X 1--4 Jlm, colorless. Oil droplets homogeneous
type, many per cell, spherical, less than 1 Jlm dia., colorless.
Specimens examined. Luding County, Mt. Gonggashan, Hailuogou, 2500m, 29°30'N, 102°00'E,
on rotten log, 27 Aug. 1996, Higuchi 30206; Kangding County, Shad - Lugba, 3350 m, 29°30'N,
10! 0 30 ' E, on rock-cliff, 9 Sept. 1996, Higuchi 30768; Jiulong County, north of Jiulong, 3540m,
29° IO 'N, 10! 0 30'E, on trunk, 11 Sept. 1996, Higuchi 30976.

Herbertus kurzii (Steph.) Chopra, 1. Indian Bot. Soc. 22: 247 (1943).
(PI. II : I, 2)
Oil bodies homogeneous type, 18- 30 per cell of vitta, 2- 9 per submarginal cellof disc, oval to oblong, 2- 5 X2--4 Jlm, colorless. Oil droplets homogeneous type,many per cell, spherical, less than I Jlm dia., colorless.

4.

Specimen examined. Kangding County, south of Jiagenba, 3250 m, 29°50 'N, 10 I 0 30'E, on boulder, 8 Sept. 1996, Higuchi 30607.

Pseudolepicoleaceae(PI. II: 3)
Blepharostoma minus Horik., Hikobia I (2): I 03 (1951 ).
Oil bodies granulose type, 1- 3 (5) per cell of segment, globose, 2- 3 Jlm dia. ,composed of indistinct granules, colorless. Oil droplets homogeneous type, many per
5.

Oil bodies and oil droplets of some hepatics from Sichuan, China

83

cell, spherical, less than !pm dia. , colorless.
Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 3800 m, 29°20' N,
IOI 0 30'E, on soil, 11 Sept. 1996, Higuchi 30943.

6.

Pseudolepicolea trollii (Herz.) Grolle et An do, Hikobia 3: 177 ( 1963).

(PI. 11: 4)
Oil bodies not seen. Oil droplets homogeneous type, many per cell, spherical, 12 f.1m dia., colorless.
Specimen examined. Luding County, Mt. Gonggashan, Hai luogou, 3000 m, 29°30'N, I 02°00'E,
on rock-cliff, 9 Sept. 1996, Higuchi 30100.

Lepidoziaceae
Lepidozia reptans (L.) Dum., Recueil Observ. Jungerm.: 19 (1835). (PI. 11 : 5)
Oil bodies homogeneous type, 5- 15 per cell, globose to oval, 3- 8 X 3- 5 f..lm, colorless. Oil droplets homogeneous type, many per cell, spherical, less than 1 f..lm dia.,colorless.
7.

Specimen examined. Kangding County, Shad - Lugba, 3350 m, 29°30'N, I 0 I 0 30' E, on rotten
stump, 9 Sept. 1996, Higuchi 30728.

The above description of oil bodies are nearly the same as those reported fromJapan (Hattori 1953), except for shape (rice grain-like in Japanese plants).
Lepidozia robusta Steph., Mem. Soc. Sci. Nat. Cherbourg 29: 217 (1894).
(PI. II: 6)
Oil bodies homogeneous type, 5- 20 per cell, globose, 2-4 f.1m dia., colorless.
Oil droplets homogeneous type, many per cell, spherical, less than I f..lm dia., colorless.
8.

Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20'N,
I 0 I 0 30'E, on soil, I 0 Sept. 1996, Higuchi 30845b.

Lepidozia subtransversa Steph., Bull. Herb. Boiss. 5: 95 (1897).
(PI. Ill: I)
Oil bodies homogeneous type, 5- 15 per cell, globose, 2-4 f.1m dia., colorless.
Oil droplets homogeneous type, many per cell, spherical, less than 1 f..lm dia., colorless.
9.

Specimens examined. Luding County, Mt. Gonggashan, Hailuogou, 3050 m, 29°30'N, 102°00'E,
on trunk, 2 Sept. 1996, Higuchi 30308; Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m,
29°20'N, I 0 I 0 30'E, on humus, I 0 Sept. 1996, Higuchi 30848.

Cephaloziaceae
10.

Cephalozia bicuspidata (L.) Dum., Recueil Observ. Jungerm.: 18 (1835).
(PI. Ill: 2)
Oil bodies and oil droplets not seen.

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Tatsuwo Furuki and Masanobu Higuchi

Specimen examined. Jiu long County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20' N,
101 °30 'E, on humus, 10 Sept. 1996, Higu chi 30806.

Antheliaceae
11. Anthelia juratzkana (Limpr.) Trev. , Mem. R. 1st. Lombardo, Ser. 3. Cl. Sci. 4:416 (1877).
(PI. Ill: 3)
Oil bodies not seen. Oil droplets homogeneous type, many per cell, spherical,
less than 1 Jlm dia., coiorless.
Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20'N,
I 0 I 0 30'E, on soil, I 0 Sept. 1996, Higuchi 30842.

Jungermanniaceae
12. Anastrepta orcadensis (Hook.) Schiffn. in Engler & Prantl, Nat. Pfl.-fam. 1(3):85(1893).
(Pl.III:4)
Oil bodies homogeneous or botryoidal type, 3- 12 per cell, globose to oval,
2-6 X 2-4 Jlm, often turbid, sometimes composed of indistinct, large granules, colorless. Oil droplets homogeneous type, many per cell, spherical, less than 1 Jlm dia.,colorless.
Specimens examined. Kangding County, Shad - Lugba, 3350 m, 29°30'N, I0 I 0 30'E, on decaying
log, 9 Sept. 1996, Higuchi 30792; Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20' N,
IOI 0 30'E,onhumus, IOSept. 1996,Higuchi30808.

13.

Anastrophyllum joergensenii Schiffn., Hedwigia 49: 396 ( 191 0).
(PI. Ill : 5)
Oil bodies granulose type, 2- 6 per cell, globose to oval, 2- 6 X 2-4 Jlm, composed of minute granules, colorless. Oil droplets homogeneous type, many per cell,spherical, less than 1 Jlm dia., colorless.
Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20' N,
I 0 I 0 30'E, on humus, I 0 Sept. 1996, Higuchi 30840.

14. Barbilophozia lycopodioides (Wallr.) Loeske, Verh. Bot. Ver. Brandenburg 49 :
3 7 ( 1907).
(PI. Ill: 6)
Oil bodies granulose type, 1-5 per cell, nearly globose, 3- 7 Jlm dia., composedof minute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than 1 Jlm dia., colorless.
Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20'N,
IOI 0 30'E, on humus, 10 Sept. 1996, Higuchi 30846.

15.

Chandonanthus birmensis Steph. , Bull. Soc. Roy. Bot. Belgique 38: 43 (1899).
(PI. IV: 1)
Oil bodies granulose type, 2- 5 per cell, nearly globose, 2- 6 Jlm dia. , composedof minute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,

Oil bodies and oil droplets of some hepatics from Sichuan, China

85

less than I Jlm dia., colorless.
Specimen examined.
Sept. 1996, Higuchi 30990.

Jiulong County, north of Jiulong, 3540 m, 29° I O' N, I 0 I 0 30'E, on soil, 11

16. Chandonanthus hirtellus (Web.) Mitt., J. Proc. Linn. Soc. London 22: 321( 1887).
(PI. IV: 2, 3)
Oil bodies granulose type, 2- 5 per cell, nearly globose, 2- 6 J..lm dia., composedof minute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than I J..lm dia., colorless.
Specimen examined. Jiulong County, north of Jiulong, 3540m, 29° IO 'N, IOI 0 30'E, on boulder,
11 Sept. 1996, Higuchi 30994.

Jamesoniella nipponica Hatt., J. Jpn. Bot. 19: 350 (1943).
(PI. IV: 4)
Oil bodies granulose type, 3- 6 per cell, globose to oval, 4-7X3- 5 J..lm, composed of minute granules, colorless. Oil droplets homogeneous type, many per cell,spherical, less than 1 J..lm dia., colorless.
17.

Specimens examined. Kangding County, south of Jiagenba, 3250 m, 29°50 'N, I 0 I 0 30' E, on baseof tree, 8 Sept. 1996, Hig uchi 30568; Kangding County, Shad - Lugba, 3350m, 29°30 ' N, IOI 0 30' E, on
soil , 9 Sept. 1996, Higuchi 30720; on rotten stump, Higuchi 30 756; on rotten log, Higuchi 30781.

The above description of oil bodies is nearly the same as those given for Japanese plants (Hattori 1951 as Jamesoniella autumnalis var. nipponica). The plants examined have strong verrucae on the surface of leaves, which suggests that thisspecies is distinct from Jamesoniella autumnalis with smooth leaf surface.
Jungramannia clavellata (Steph.) Amak., J. Hattori Bot. Lab. 22: 69, (1960).
(PI. IV: 5)
Oil bodies granulose type, 1- 3 per cell, oblong, 5- 12 X3-8j..lm, composed ofminute granules, colorless. Oil bodies in rhizoid cell homogeneous type, many, occupied cell lumen, oval, 2-4 X 1- 2j..lm, often turbid, colorless. Oil droplets homogeneous type, few per cell, spherical, less than 1 J..lm dia. , colorless.
18.

Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20 'N,
I 0 I 0 30 ' E, on soil, I 0 Sept. 1996, Higuchi 30845a.

19.

Lophozia excisa (Dicks.) Dum., Recueil Observ. Jungerm.: 17 (1835).
(PI. IV: 6)
Oil bodies granulose type, 5- 14 per cell, globose to oblong, 3- 8X3- 5 J..lm, composed of minute granules, colorless. Oil droplets homogeneous type, many per cell,spherical, less than 1 J..lm dia., colorless.
Specimen examined. Kangding County, south of Jiagenba, 3250m, 29°50'N, IOI 0 30'E, on rotten
log, 8 Sept. 1996, Higu chi 30536.

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Tatsuwo Furuki and Masanobu Higuchi

20. Lophozia longidens (Lindb.) Macoun, Geol. Survey Canada: Cat. Canad. PI. 7:
18 (1902).
(PI. V: I)
Oil bodies granulose type, 8- 20 per cell, nearly globose, 3- 7 f.1m dia. , composedof minute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than 1 f.1m dia., colorless.
Specimen examined. Kangding County, south of Jiagenba, 3250 m, 29°50'N, 101 °30' E, on boulder, 8 Sept. 1996, Higuchi 30544.

21. Lophozia morrisoncola Horik., J. Sci. Hiroshima Univ., Ser. B, Div. 2, Bot. 2:
150 (1934 ).
(PI. V: 2)
Oil bodies granulose type, 3- 6 per cell, nearly globose, 5- 12 f.1m dia., composedof minute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than 1 f.1m dia., colorless.
Specimen examined. Kangding County, Shad - Lugba, 3350 m, 29°30'N, I 0 I 0 30 ' E, on rock-cliff,
9 Sept. 1996, Higuchi 30738.

22. Tritomaria quinquedentata (Hud.) Buch, Memoranda Soc. fauna Fl. Fenn. 8:290 (" 1932" 1933).
(PI. V: 3)
Oil bodies granulose type, 5- 10 per cell, oval to oblong, 4- 10 X 3- 6 f.1m, composed of minute granules, colorless. Oil droplets homogeneous type, many per cell,spherical, less than 1 f.1m dia., colorless.
Specimens examined. Kangding County, Daxueshan Pass (between Kangding and Xindugiao),
4300 m, 30°00'N, I 0 I 0 50' E, on soil, 7 Sept. 1996, Higuchi 30494; Jiulong County, Jicoushan Pass (northof Jiulong), 4200 m, 29°20'N, I 0 I 0 30'E, on soil, I 0 Sept. 1996, Higuchi 30828.

Gymnomitriaceae

23.

Marsupella commutata (Limpr.) H. Bernet, Cat. Hep. S.-0. Suisse: 29 (1888).
(PI. V: 4)
Oil bodies granulose type, 2- 3 per cell, globose to oblong, 4-8X4-6f.lill, composed of minute granules, colorless. Oil droplets homogeneous type, many per cell,spherical, less than 1 f.1m dia., colorless.
Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20'N,
101 °30'E, on rock-cliff, 10 Sept. 1996, Higuchi 30812.

24. Marsupella revoluta (Nees) Dum., Bull. Soc. Roy Bot. Belgique 13: 126(1874).
(PI. V: 5)
Oil bodies granulose type, 1- 3 per cell, nearly globose, 2--4f.1m dia., composedof exceedingly minute granules, colorless. Oil droplets homogeneous type, many percell, spherical, less than I f.1m dia., colorless.
Specimen examined. Luding County, Mt. Gonggashan, Hailuogou, 3000 m, 29°30'N, I 02°00'E,
on rock-cliff, 9 Sept. 1996, Higuchi 30117.

Oil bodies and oi l droplets of some hepatics from Sichuan, China

87

Scapaniaceae
25. Scapania ciliata Sande Lac. in Miquel, Ann. Mus. Bot. Lugduno-Batavum[Prol. Fl. Jap.] 3: 209 (1867).
(PI. V: 6)
Oil bodies granulose type, 20- 30 per cell, globose to oblong, 3- 8X3- 5 .urn,
composed of minute granules, colorless. Oil droplets homogeneous type, many percell, spherical, less than I .urn dia. , colorless.
Specimen examined. Luding County, Mt. Gonggashan, Hailuogou, 3000m, 29°30 'N, 102°00 ' E,on boulder, 9 Sept. 1996, Higu chi 30090.

Geocalycaceae
Lophocolea bidentata (L.) Dum. , Recueil Observ. Jungerm.: 17 (1835).
(PI. VI: 1)
Oil bodies botryoidal type, 2- 3 per cell, globose, 2-5 .urn, composed of large
granules, colorless. Oil droplets homogeneous type, few per cell, spherical, less thanI .urn dia., colorless.
26.

Specimen examined. Kangding County, Daxueshan Pass (between Kangd ing and Xindugiao),
4300 m, 30°00'N, I0 I 0 50'E, on humus, 7 Sept. 1996, Higuchi 30465.

Plagiochilaceae
Plagiochila elegans Mitt. , J. Proc. Linn. Soc. Bot. 5: 97 ("1860" 1861).
(PI. VI: 2)
Oil bodies homogeneous type, 4- 9 per cell, oval to oblong, 3- 7X3-4 .urn, often
turbid with septa, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than 1 .urn dia. , colorless.

27.

Specimens examined. Kangding County, south of Jiagenba, 3250 m, 29°50'N, I 0 I 0 30'E, on so il ,8 Sept. 1996, Higuchi 30532; Jiulong County, north of Jiulong, 3540m, 29° IO'N, IOI 0 30' E, on base oftree, 11 Sept. 1996, Higuchi 31008.

28. Plagiochila semidecurrens (Lehm. et Lindenb.) Lehm. et Lindenb. in Lindenb.
(PI. VI: 3-5)
et Gott., Spec. Hepat. (fasc. 5): 142 (1843).
Oil bodies granulose type, 3- 8 per cell, oval to oblong, 3- 6 X3- 5 .urn, 6- 10 per
cell of leaf-base , 5- 8 X 3-4 .urn, composed of minute granules, colorless. Oil droplets
homogeneous type, many per ce ll, spherical, less than I .urn dia. , colorless.
Specimen examined. Jiulong County, north of Jiulong, 3540m, 29° IO 'N, IOI 0 30'E, on boulder,
11 Sept. 1996, Higuchi 31021.

Radulaceae
29.

Radula gedena Steph., Hedwigia 23: 146 (1884).
(PI. VII: I)
Oil bodies granulose type, I (2) per cell, oval to oblong, like meniscus with con-

88

Tatsuwo Furuki and Masanobu Higuchi

vex-concave faces, 8- 15 X 6- 10 J.Lm, composed of minute granules, light brown. Oil
droplets not seen.
Specimen examined. Kangding County, south of Jiagenba, 3250 m, 29°50'N, l 0 l 0 30' E, on boulder, 8 Sept. 1996, Higu chi 30562.

Radula gedena is characterized by (I) the abundant discoid gemmae occurringon the margin of leaf-lobes, (2) the pronounced mammilliform rhizoid-initial area on
the inflated carina) region of leaf-lobes and (3) the uniformly thin-walled cells ofleaf-lobes which are very irregular in form and size (Yamada 1979). The distributionof this species extended northward from the known range, including Java, Borneo,
Thailand and Japan (Yamada 1979).
Porellaceae
Macvicaria ulophylla (Steph.) Hatt., J. Hattori Bot. Lab. 5: 81 (1951).
(PI. VII: 2)
Oil bodies homogeneous type, many per cell, oblong, 3- 5 X2- 3 J.Lm, colorless.
Oil droplets not seen.

30.

Specimen examined. Kangding County, Shad - Lugba, 3l50m, 29°30'N, l0l 0 20'E, on boulder, 9
Sept. 1996, Higuchi 30663.

31.

Porella gracillima Mitt., Trans. Linn. Soc. London, Ser. 2, 3: 202 (1891).
(PI. VII: 3)
Oil bodies homogeneous type, many per cell, oblong, 2- 5 X 2- 3 J.Lm, colorless.
Oil droplets not seen.
Specimens examined. Kangding County, south of Jiagenba, 3250 m, 29°50'N, l 0 l 0 30'E, on boulder, 8 Sept. 1996, Higuchi 30601; Kangding County, Shad - Lugba, 3350 m, 29°30'N, 101 °30'E, on rockcliff, 9 Sept. 1996, Higuchi 30739, 30770.

32. Porella revoluta (Lehm.) Trev. , Mem. Real. Ist. Lombardo, Ser. 3, 4: 407(1877).
(PI. VII: 4)
Oil bodies homogeneous type, many per cell , oblong, l- 2 X 1-4 J.Lm, colorless.
Oil droplets not seen.
Specimens examined. Kangding County, south of Jiagenba, 3250 m, 29°50'N, 101 °30'E, on boulder, 8 Sept. 1996, Higuchi 30600; Kangding County, Shad - Lugba, 3150 m, 29°30'N, 101 °20'E, on boulder, 9 Sept. 1996, Higuchi 30671.

Frullaniaceae
33. Frullania davurica Hampe fo. dorsoblastos (Hatt.) Hatt. et Lin, J. Hattori Bot.
Lab. 59: 132 ( 1985).
(PI. VII: 5).
Oil bodies granulose type, 4-10 per cell, oblong, 3-8X 3- 5 J.Lm, composed ofminute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,

Oil bodies and oil droplets of some hepatics from Sichuan, China

89

less than 1 J.lm dia., colorless.
Specimens examined. Kangding County, south of Jiagenba, 3250 m, 29°50'N, 101 °30'E, on boulder, 8 Sept. 1996, Higuchi 30587; Kangding County, Shad - Lugba, 3150m, 29°30 ' N, 101 °20'E, on
trunk, 9 Sept. 1996, Higuchi 30629; Kangding County, Shad - Lugba, 3350 m, 29°30'N, 101 °30'E, onroot, 9 Sept. 1996, Higu chi 30758; on trunk, Higuchi 30765a; Jiulong County, north of Jiulong, 3540 m,
29° 1O'N, 101 °30'E, on trunk , 11 Sept. 1996, Higuchi 30979.

The above description of oil bodies are nearly the same as that ofF davurica fo .
davurica reported by Hattori ( 1951) from Japan.
Frullania handelii Verd. in Handei-Mazzetti, Symb. Sinic. 5: 36 ( 1930).
(PI. VII: 6)
Oil bodies granulose type, 4-6 per cell, oblong, 2- 5X2- 3 J.lm, composed ofminute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than 1J.lm dia. , colorless.

34.

Specimen examined.
Sept. 1996, Higuchi 30790.

Kangding County, Shad - Lugba, 3350 m, 29°30'N, 101 °30'E, on trunk, 9

35. Frullania nepalensis (Spreng.) Lehm. et Lindenb., Nov. Minus Cogn. Strip.
Pugillus 4: 19 (1832) .
(PI. VIII: 1)
Oil bodies granulose type, 4- 7 per cell, oblong, globose to oval, 4-8X3- 5 J.lm,
composed of minute granules, colorless. Oil droplets homogeneous type, few per
cell, spherical, less than 1J.lm dia. , colorless.
Specimens examined. Kangding County, south of Jiagenba, 3250 m, 29°50 'N, 101 °30 'E, on boulder, 8 Sept. 1996, Higuchi 30521,30564,30570, 30604; on decaying log, Higuchi 30540.

36.

Frullania parvistipula Steph., Spec. Hepat. 4: 397 (191 0).
(PI. VIII: 2)
Oil bodies granulose type, 2- 5 per cell, nearly globose, 3- 5 J.lm dia., composedof minute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than 1 J.lm dia., colorless.
Specimens examined. Kangding County, south of Jiagenba, 3250 m, 29°50 'N, I 01 °30'E, on boulder, 8 Sept. 1996, Higuchi 30560; on trunk, Higuchi 30569; on trunk , Higu chi 30571.

37. Frullania tamarisci (L.) Dum. subsp. obscura (Verd.) Hatt., J. Hattori Bot.
Lab. 35:216 (1972).
(PI. VIII: 3)
Oil bodies granulose type, 3- 7 per cell, oblong, 2-5 X2- 3 J.lm, composed ofminute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than I J.lm dia., colorless.
Specimen examined. Jiulong County, Jicoushan Pass (north of Jiulong), 4200 m, 29°20'N,
101 °30'E, on base oftree, 10 Sept. 1996, Higuchi 30827.

Frullania tubercularis Hatt. et Lin., J. Jpn. Bot. 60: 107 (1985).
(PI. VIII: 4)
Oil bodies granulose type, 3- 8 per cell, oblong, 3- 7 X 2- 3 J.lm, composed ofminute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,

38.

90

Tatsuwo Furuki and Masanobu Higuchi

less than I J.lm dia., colorless.
Specimens examined. Kangding County, Shad - Lugba, 3350 m, 29°30' N, 101 °30 ' E, on rotten
stump, 9 Sept. 1996, Higuchi 3072 7; on trunk, Higuchi 30765b.

Lejeuneaceae
39.

Lejeunea parva (Hart.) Mizut., Mise. Bryol. Lichenol. 5: 178 ( 1971 ).
(PI. VIII: 5)
Oil bodies granulose type, 2- 6 per cell, oblong, 3- 12 X3- 5 J.lm, composed ofminute granules, colorless. Oil droplets homogeneous type, few per cell, spherical,
less than I J.lm dia., colorless.

Specimen examined. Luding County, Mt. Gonggashan, Hai1uogou, 2500 m, 29°30 ' N, 102°00 ' E,
on trunk, 28 Aug. 1996, Higuchi 29845.

Metzgeriales
Metzgeriaceae
40. Apometzgeria pubescens (Schrank.) Kuwah ., Rev. Bryol. Lichenol. 34: 214( 1966).
(PI. VIII: 6)
Oil bodies and oil droplets not seen.
Specimen examined. Kangding County, Shad - Lugba, 3350 m, 29°30'N, 101 °30 ' E, on rock-c1i ff,
9 Sept. 1996, Higuchi 30746.

Discussion
The oil bodies of 14 species were described for the first time (Table I). Among
the remained 26 species most of species have the oil bodies which agree with the descriptions of oil bodies in the previous reports. However, in Lophocolea bidentata thetype of oil bodies of Chinese plants differ from that of North American plants which
Schuster ( 1980) described as "finely granular-segmented type ." Also in Lepidozia
reptans their shape is different from that of North American plants which Schuster(1969) described as "homogeneous and 2- 3 segmented ones," although it agrees withthat of Japanese plants reported by Hattori ( 1953). In these species the careful comparative study of oil bodies will be needed.
This is the first report of the oil droplets in Chinese hepatics. The oil droplets
were recognized in the cells of most of species examined in this study, except for

Cephalozia bicuspidata, Radula gedena, Macvicaria ulophylla, Pore/la gracillima
and Pore/la revoluta. Oil droplets were almost uniformly similar in all the taxa examined, that is, they are spherical, homogeneous and colorless. In Chandonanthus
hirtellus oil droplets were recognized in Chinese plants, although Furuki & Higuchi( 1996) reported the absence of oi l droplets in New Caledonian plants. As noted by
Furuki & Higuchi ( 1996) the taxonomic significance of oil droplets in hepatics may

Oil bodies and oil droplets of some hepatics from Sichuan, China
Table I.
Species
Herbertus aduncus
Herbertus delavayi
Herbertus dicranus
Herbertus kurzii
Blepharostoma minus
Pseudolepicolea trollii
Ledidozia retans

Lepidozia robusta
Lepidozia subtransversa
Cephalozia bicuspidata
An thelia juratzkana
Anastrepta orcadensis
Anastrophyllun joergensenii
Barbi lophozia lycopodioides
Chandonanthus birmensis
Chandonanthus hirtellus

Jamesoniella nipponica
Jungermannia clavellata
Lophozia excisa
Lophozia longidens
Lophozia morrisoncola
Tritomaria quinquedentata
Marsupella commutata
Marsupella revoluta

91

Types of oil bodies in 40 species of hepatics examined.
Type of oi l bodies
homogeneous
homogeneous
homogeneous
homogeneous
homogeneous
granulose
granuloseabsenthomogeneous
homogeneous
homogeneous
homogeneous
homogeneous
homogeneous
absentabsentabsentabsenthomogeneous
homogeneous, botryoidal
granu lose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
grnaulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose
granulose

Locality
Japan
China
China
China
China
Japan
China
China
Japan
N. America
China
China
Japan
China
N. America
China
N. America
China
Japan
China
China
N. America
China
Japan
China
Japan
Japan
New Caledonia
China
Japan
China
China
N. America
China
N. America
China
Japan
China
Japan
China
Japan
China
N. America
China

References
Hattori ( 1951 , 1952)*
present study
present study
present study
present study
Hattor (1951, 1953)
present study
present study
Hattori (1953)
Schuster ( 1969)
present study
present study
Hattori ( 1951)
present study
Schuster ( 1974)
present study
Schuster ( 1974)
present study
Kitagawa ( 1966)
present study
present study
Schuster (1974)
present study
Kitagawa ( 1966)
present study
Hattori ( 1951)
Kitagawa ( 1965)
Furuki & Higuchi ( 1996)
present study
Hattori ( 195 1)
present study
present study
Schuster ( 1969)
present study
Schuster ( 1969)
present study
Kitagawa ( 1966)
present study
Kitagawa ( 1965)
present study
Hattori (1953)
present study
Schuster ( 1974)
present study

92

Tatsuwo Furuki and Masanobu Higuchi
Table I
Species

Scapania ciliata
Lophocolea bidentata
Plagiochila elegans
Plagiochila semidecurrens
Radula gedena
Macvicaria ulophylla
Porella gracillima
Porella revoluta
Frullania davurica fo. dorsoblastos
Frullania handelii
Frullania nepalensis
Frullania parvistipula
Frullania tamarisci subsp. obsucra
Frullania tubercularis
Lejeunea parva
Apometzgeria pubescens(continued)

Type of oil bodies
granulose
granulose
granulose
botryoidal
homogeneous
granulose
granulose
granulose
homogeneous
homogeneous
homogeneous
homogeneous
homogeneous
granulose
granulosgranulose
granulose
granulose
granulose
granulose
granulose
granulose
granuloseabsentabsentLocality
Japan
China
N. America
China
China
Japan
China
China
Japan
China
Pakistan
China
China
China
China
Japan
China
China
Japan
China
China
Japan
China
Japan
China

References
Hattor ( 1951)
present study
Schuster ( 1980)
present study
present study
Hattori ( 1951)
present study
present study
Hattori (1951)
present study
Furuki (1993)
present study
present study
present study
present study
Hattori ( 1953)
present study
present study
Hattori ( 1951)
present study
present study
Inoue ( 1976)
present study
Hattori ( 1951)
present study*Previous reports are also included, but voucher specimens are not examined.

be much less than that of oil bodies.
Acknowledgments

We wish to express our sincere thanks to Dr. Hiroshige Koyama of the National
Science Museum, Tokyo, for giving us the opportunity to study Chinese hepatics. Junior author is also indebted to Mr. Liu Peigui and Mr. Li-Song Wang of the Kunming
Institute of Botany, Academia Sinica for their kind help in the field research. This
study was partly supported by a Grant-in-Aid for Overseas Scientific Survey (no.
07041151) from the Ministry of Education, Science, Sports and Culture, Japan.

Oil bodies and oil droplets of some hepatics from Sichuan, China

93

References
Furuki, T., 1993. Studies on oil bodies of some hepatics collected from Pakistan. In T. Nakaike and S.
Malik (eds.), Cryptogamic Flora of Pakistan, Vol. 2: 231 - 238 .
Furuki, T. & M. Higuchi, 1996. Studies of oil bodies and oil droplets of some hepatics (Jungermanniales) from New Caledonia. Bull. Natn. Sci. Mus ., Tokyo , Ser. B, 22(2): 59- 75.
Hattori, S. , 1951. Oil bodies of Japanese Hepaticae (1). J Hattori Bot. Lab. 5: 69- 97, pis. I-VII. (InJapanese.)
Hattori, S., 1953. Oil bodies of Japanese Hepaticae (2). J Hattori Bot. Lab. 10: 63- 78, pis. 1- III. (InJapanese.)Inoue, H., 1976. Illustrations of Japanese Hapeticae 2. 193 pp. Tsukiji Shokan, Tokyo. (In Japanese. )Kitagawa, N., 1965. A revision of the family Lophoziaceae of Japan and its adjacent regions. I. J Hatfori Bot. Lab. 28: 239- 291.
Kitagawa, N., 1966. A revision of the family Lophoziaceae of Japan and its adjacent regions. II. J Hatfori Bot. Lab. 29: 101 - 149.
Piippo, S., X.-L. He & T. Koponen, 1997. Hepatics from northwestern Sichuan, China, with a checklist
ofSichuan hepatics. Ann. Bot. Fennici 34: 51 - 63 .
Schuster, R. M., 1969. The Hepaticae and Anthocerotae of North America. Vol. 2. xii+ 1062 pp. Columbia Univ. Press, New York.
Schuster, R. M. , 1974. The Hepaticae and Anthocerotae of North America. Vol. 3. xvi + 880 pp. Columbia Univ. Press, New York.
Schuster, R. M. , 1980. The Hepaticae and Anthocerotae of North America. Vol. 4. xviii + 1334 pp. Columbia Univ. Press, New York.
Yamada, K. , 1979. A revision of Asian taxa of Radula , Hepaticae. J Hattori Bot. Lab. 45: 201 - 322.

94

Tatsuwo Furuki and Masanobu Higuchi

Explanation of Plates
Plate I. Oil bodies, oil droplets and ch loroplasts in ce lls at leaf-vitta ( I, 3, 5) and at submargin of leafdisc (2, 4, 6). I, 2. Herbertus aduncus (D icks.) Gray (Higuchi 30609). 3, 4. Herbertus de/avayi
Step h. (Higuch i 30829). 5, 6. Herbertus dicranus (Tay!.) Trev. (Higuchi 30206). Scales all I 0 pm.
Plate 11. Oil bodies, oil droplets and chloroplasts in cells at leaf-vitta ( I), at submargin of leaf-disc (2),at leaf-segment (3), at margin of disc (4) and at middle of leaf-disc (5, 6). I, 2. Herbertus kurzii(Steph.) Chopra (Higuchi 30607). 3. Blepharostoma minus Horik. (Higuchi 30943). 4. Pseudo/epico/ea trol/ii (Herz.) Grolle et Ando (Higuchi 30100). 5. Lepidozia reptans (L.) Dum. (Higuchi
30728). 6. Lepidozia robusta Steph. (Higuchi 30845b). Scales all 10 pm .
Plate Ill. Oil bodies, oil droplets and chloroplasts in cells at middle of leaf-disc (1 - 3) and at leaf-midd le (4-6). I. Lepidozia subtransversa Steph. (Higuchi 30848). 2. Cepha/ozia bicuspidata (L.) Dum.
(Higuchi 30806). 3. Anthe/ia juratzkana (Limpr.) Trev. (Higuchi 30842). 4. Anastrepta orcadensis(Hook.) Schiffn. (Higuchi 30808). 5. Anastrophyl/um joergensenii Schiffn. (Higuchi 30840). 6. Barbilophozia lycopodioides (Wallr.) Loeske (Higuchi 30846). Scales all 10 pm.
Plate IV Oil bodies, oil droplets and ch loroplasts in cells at middl e of leaf-disc (I, 2), at apex of leaf
(3) and at leaf-middle (4-6). I. Chandonanthus birmensis Steph. (Higuchi 30990). 2, 3. Chandonanthus hirtellus (Web.) Mitt. (Higuchi 30994). 4. Jam esoniel/a nipponica Hatt. (Higuchi 30720).
5. Jungermannia clavellata (Steph. ) Amak. (Higuchi 30845a) . Arrow shows a rhizo id ce ll.
6. Lophozia excisa (Dicks.) Dum. (Higuchi 30536). Scales all 10 pm .
Plate V Oil bodies, oil drop lets and chloroplasts in middle cells at leaf ( 1- 5) and at ventral leaf- lobe
(6). I. Lophozia /ongidens (Lindb.) Macoun. (Higuchi 30544). 2. Lophozia morrisonco/a Horik.
(Higuchi 30738). 3. Tritomaria quinquedentata (Hud.) Buch. (Higuchi 30828). 4. Marsupel/a commutata (Limpr.) H. Bern. (Higuchi 30812). 5. Marsupe/la revo/uta (Nees) Dum. (Higuchi 30117).
6. Scapania ciliata San de Lac. (H iguchi 30090). Scales a ll I 0 pm .
Plate VI. Oi l bodies, oi l droplets and chlorop lasts in cells at leaf-middle ( 1-4) and at leaf-base (5).
I. Lophoco/ea bidentata (L.) Dum. (Higuchi 30465) . 2. Plagiochila elegans Mitt. (Higuchi 31008).
3- 5. P/agiochila semidecurrens (Lehm. et Lindenb.) Lehm. et Lindenb. (Higuchi 31021 ). 4. Anotherfocus of Fig. 3. Arrows show oil droplets. Scales all I 0 pm.
Plate VII. Oil bodies, oil droplets and chloroplasts in middle ce lls at leaf-lobe (1 - 6). I. Radu/a gedena
Steph. (Higuchi 30562). 2. Macvicaria ulophyl/a (Steph .) Hatt. (Higuchi 30663). 3. Pore/la gracil/ima Mitt. (Higuchi 30770). 4. Pore/la revoluta (Lehm.) Trev. (Higuchi 30600). 5. Frul/ania davurica
Hampe fo. dorsoblastos (Han.) Hatt. et Lin. (Higuchi 30765a). 6. Frul/ania handelii Verd. (Higuchi
30790). Scales all 10 pm.
Plate VIII. Oil bodies, oil droplets and chloroplasts in middle cells at leaf-lobe (1 - 5) and at wing ofthallus. I. Frullania nepalensis (S preng.) Lehm. et Lindenb. (Higuchi 30604). 2. Frul/ania parvistipula Step h. (Higuchi 3057 1). 3. Frul/an ia tamarisci (L.) Dum. subsp. obsucra (Verd.) Hatt. (H iguchi
30827). 4. Frullania tubercularis Hatt. et Lin. (Higuchi 30765b). 5. Lejeunea parva (Hatt.) Mizut.
(Higuchi 29845). 6. Apometzgeria pubescens (Schrank.) Kuwah. (Higuchi 30746). Scales a ll 10 pm.

Oil bodies and oil droplets of some hepatics from Sichuan, China

Plate I

95

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Tatsuwo Furuki and Masanobu Higuchi

Plate 11

Oil bodies and oil droplets of some hepatics from Sichuan, China

Plate Ill97

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Tatsuwo Furuki and Masanobu Higuchi

Plate IV

Oil bodies and oil droplets of some hepatics from Sichuan, Chin a

Plate V99

100

Tatsuwo Furuki and Masanobu Higuchi

Plate VI

Oil bodies and oil droplets of some hepatics from Sichuan, Ch ina

Plate Vll

101

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Tatsuwo Furuki and Masanobu Higuchi

Plate VIII