lourn. HallOri BOI. Lab. No . 61: 281- 297 (Dec. 1986)

NOTES ON THE LEJEUNEACEAE. 12. MASTIGOLEJEUNEA HUMILIS AND ITS RELATED SPECIES FROM ASIA

MASAMI MIZUTANI1

1) Mastigolejeunea humilis (Gott.) Schiffn. In Engler & Prantl. Nat. Pft.-fam. 1, 3: 129 (1893).

Phragmicoma humilis Gatt. in Gatt. et aI., Synop. Hepat.: 299 (1845). Plychocoleus humilis (Gatt .) Trev., Mem. Istit. Lombardo 3. 4: 405 (1877). Lejeunea humilis (Gatt.) Spruce, Hepat. Amazon. et Andinae: 101 (1884). L. guahamensis Lindenb. in Gott. et aI., Synop. Hepat. : 333 (1845).

(Fig. 1)

Mastigolejeunea guahamensis (Lindenb.) Steph .. Denkschr. Math.-Nat. Kl. Acad . Wiss. Wien 85: 195 (1910). M. whightii Steph .. Spec. Hepat. 4: 773 (1912). Brachiolejeunea tylimanthoides Steph., Spec. Hepat. 5: 129 (1912). B. mioken.l'is Steph .. Spec. Hepat. 5: 132 (1912). Mastigolejeunea losbanosa Steph., Spec . Hepat. 6: 563 (1924). Thysananthus liukiuensis Horik., J . Sci. H iroshima Univ. ser. b. div. 2, 2: 250 (1934). Mastigolejeunea liukiuensis (Horik.) Hatt. . Bull. Tokyo Sci . Mus. 11: 139 (1944). M. liukiuensis (Horik.) Hatt. var. mayebarae Hatt ., J . Jap . Bot. 26: 299 (1951 J.

Plants small or medium in size, blackish brown in herbaria. Stems 10--20 mm long, 0. 18- 0.2 mm in diam. , with leaves 1.3- 1.8 mm wide, irregu larly branched; ventral merophyte of the stem 5- 7 cells wide; rhizoids few. Leaf-lobes closely imbricate, obliquely spread ing, ovate, 0.7- 1 mm long, 0.6--0.8 mm wide, apices obtuse to acute, ventral margins usually incurved , dorsal margins arched from auriculate bases; apical cells 8- 15 x 6- 8 j1, median cells 17- 25 x 12-18/1, walls thin, trigones large, cordate2

, intermediate thickenings few, small, basal cells abou t 38 x 25/1; cuticle smooth; leaf-lobule small, ovate, about 2/5 as long as the lobe, apex obliquely truncate, apical tooth small or obsolete, if present 1- 2 cells long and 1- 2 cells wide at base; keel nearly straight. Underleaves imbricate, obliquely spreading, about 3 timeS' as wide as the stem, ren iform, 0.4--0.52 mm long, 0.5- 0.75 mm wide, apices flat or recurved , lateral margins fl at or slightl y recurved, with minute basal auriculation , insertions sinuate.

Autoicous. Androecium terminal on stem or branch, often with apical innovation; bract­lobes 5-10 pairs, closely imbricate, obliquely to suberect-spreading, obliquely oblong, about 1/2 as long as leaf-lobes; bract-lobule ovate, about 3/4 as long as bract-lobe; bracteoles imbricate, nearly quadrate, slight ly smaller than underleaves. Gynoecium terminal on stem or branch, with 1-2 subfloral innovations, innovation usually again floriferous; bract-lobe ovate or orbicular, 0.65- 1 mm long and wide, apex usually obtuse, margin flat Or undulate; bract-lobule oblong, 1/2- 2/3 as long as bract-lobe, apex more or less crenate; keel 2/3-1 as long as bract-lobule, straight; bracteole oblong, 0.75- 0.9 mm long, 0.5- 0.55 mm wide, apex rounded, often recurved , lateral margins more or less recurved. Perianth obovate or oblong, \ - l.l mm long, 0.5- 0.7 mm

I The Hattori Botanical Laboratory, Obi, Nichinan-shi , Miyazaki-ken, 889-25.

2 "Cordate" is used when the trigones is bulging with two convex and one concave sides [after Gradstein (1975, p. 18)].

282 Journ . Hilttori Bot. Lab. No. 61 1 986

a q

q

FIG. 1. Mastigolejeunea humilis (Gott.) Schiffn . a- b. Portions of plants, ventral view, x 25. c- e. Leaves, x 25. f - i. Cells of leaf-lobes,f- g from apices, h- i from middle, x 400. j.

Apical portion of leaf-lobule, x 200. k- l. Underleaves, x 25. m- n. Female bracts, x 25. O .

Female bracteole, x 25. p-r. Perianths, p-q- ventral view, r--{:\orsal view, x 25. Nos. a, c,/, h,

j, k from isotype of Phragmicoma humilis from Java (herb. Lindenberg 6012) (G); others from specim. from Java, E. Nyman s.n. (NICH 369949).

wide, sharply 5-keeled, beak short. Specim. exam.: INDIA: Kabir, Amarkantak, 1134m, on trunk, J. Lal & N. S. Parihar 74.3 BURMA:

Thanbyuzayat , T. Nakajima S.n. ANDAMANS: E. H. Man S.n. - syntype of Mastigolejeunea whightii (G

12510); Port Blair, E. H. Man s.n. - syntype of M. whightii (G 12506). THAILAND: Payap, Fang, 600 m, on trunk, A. Touw 9410, 9423, O. Na-Thalang s.n.; Lampang, Khun Tan, 1000-1250 m, on trunks, O. Thaithong

3 All the cited specimens are in NICH, unless otherwise indicated.

M. MIZUTANI: Notes on the Lejeuneaceae. 12 283

251, 325, 326; Sakon Nakom, Kom Horn waterfall , on trunk , K. Sunlanachole 3; Nakhon Sawan, Doi Musae, 780-800 m, on fallen branch, A. Touw 8144; Uhtai Thani, Khoo Hin Dang, on climber, O. Thailhong 381, on trunk, J. Laohawanich 503; Prachinburi , Bang Bung Hills, on trunk, K. Larsen el al. 1161 ; Chan laburi, Soi Doa, S. Homchoen s.n.; akornsrithamarat , Ml. Luang, 250-620 m, on trunk , A. Touw 11381, on logs, A . Touw 11442, 11467, 11478; Phuket I. , 150-250 m, on trunk, A. Touw 11201. CAMBODIA: Angkor Wat , R. D. Svihla 53-1341. CHINA: Yunnan, Xishuangbanna, on decaying log, W. X. XU 6703; Hongkong, 50-200m, Z. Iwalsuki 34, 35, M. Mizulani 6031, 6032, 6034, 6040. JAPAN: Oki­nawa I. , T. Amallo 7107, 7200, 8189, 8213; Kumamoto-ken, l sshochi, 300m, on limestone, K. Mayeb~ra 1941 - holotype of Masligolejeunea liukiuensis var. mayebarae, 1940, 1943 - paratypes of M . liukiuensis var. mayebarae, 1952,2228, 2452,2785; Kumamoto-ken, Ono, 120 m, on rocks, K. Mayebara 2002 - paratype of M . liukiuensis var. mayebarae; Kochi -ken , Ml. Yokogura , on bark, S. Hallori 4166 - paralype of M. liukiuensis var. mayebarae . PHILIPPINES: Luzon, Semper S.n. - syntype of Brachiolejeunea Iylimanlhoides (G

20362); Luzon , Ml. Bulusan, 450- 900 m, on trunk, Z. Iwalsuki el al. 15743b; Bataan, Olongapo, H. H. Bartlell 14094; Los Banos, Baker s. n. - syntypes of Masligolejeunea losbanosa (G 21822, 21823); Bohol I. , Bilar, 2000 ft , on trunk , M . Ramos 4114a; Mindanao, Abaga, 400 m, on trunk, A. L. Z wickey 597; Plawan I. , Mt. Balsahan, L. E. Ebalo 366. BORNEO: Sa bah, Mt. Kinabalu, 600-630 m, on decaying log, Z. Iwalsuki I 840a; Kanabatangan , Sungai Pin Timber, S. Kokawa & M. HOlla 1602; - , Batu Puteh, on trunk , G. Shea 3279. JAVA: Herb. Lindenberg 6012 - isotype of Phragmicoma humilis (G); Bandjar, 150m, on trunk, M .

Fleischer s.n. (Verdoorn, Hep. Se!. Crit. 6: 257b); Bu itenzorg, E. Nyman S.n. (NICH 369949); Bogor, 200 m, on trunk , W. Meijer 75a. CELEBES: Kendari , G. Kjellberg 25. SERAM: Seram Utara, Wahai - Wasa, 70- 160 m, on trunk , fI. Akiyama 2393 (KYO); Manusela Nat. Park , Kalva - Pasahari, 20-120 m, on root, fI. Akiyama 9465 (KYO); Taniwel , Taniwel - Buria, 1- 640 m, on fa llen tree, H. Akiyama 9932, 9934, 9938 (K YO); Bula, Gunung Ili , 20-100m, on trunk, H. Akiyama 1077 1 (KYO). NEW G UINEA: W Irian , Pulau Tsiof, Zane veld s.n .; NE New Guinea, Wstern Highlands, Miji , 1800m, on trunk, H. Slreimann 21395; - , Mt. Hagen, S. Saito 9; -, Morobe, Bulolo, 730- 1500 m, S. Shea 64R9, 6575, 6593, 8994, 9075, 9078a, H. Slreimann 13480, 14079; - , - , Aseki, 1500 m, H. Streimann & E. Tamba 12346, 12759. BISMARK ARCHIP .: Mioko, Micholitz S.n. - type of Brachiolejeunea miokensis (G 2020 1). MICRONESIA: Guam, Mertens s. n. - isotype of Lejeunea

guahamensis (G); Yap, 650 ft , D. R. Smith 2027. Range: India, Burma, Thailand, Andamans, Cambodia, S. China, Japan, Phil ippines, Borneo, Java,

Celebes, Seram, New Guinea, Bismark Archip ., Micronesia.

Masligolejeunea humilis is common, somewhat variable, and widely distributed in southeastern Asia. In M. humilis the followi ng extreme forms are recognized : in the first form the plants are small , and the leaf-lobes are obtuse at apices; in the second form the plants are medium in size, and the leaf-lobes are acute at apices. The type specimens of M . humilis, M . guahamensis, M . losbanosa, and most Asian specimens belong to the first form. The type specimens of M. whightii, M . miokensis, M. Iiukiuensis , its var. mayebarae, and Brachiolejeunea tylimanthoides belong to the second form .

2) Mastigolejeunea rep/eta (Tay!.) Evans Mem. Torrey Bot. Club 8: 131 (1902).

Lejeunea repleta Tay!., London J . Bot. 5: 392 (1846). Phragmicorna repleta (Tay!.) Tay!. in GOIl. et a!., Synop. Hepat. : 742 (1847). Lejeunea wardiana Milt. , Hepat. Indiae orient.: 109 (1861). Brachiv/ejeunea wardiana (Mitt.) Steph ., Spec. Hepat. 5: 129 (1912). Lejeunea alypos Schiffn. , Forsch. Gazelle 4: 22 (1890). Mastigolejeunea alypos (SchifTn . ) Steph., Spec. Hepat. 4: 768 (1912). M . speclabi/is Steph., Spec . Hepat. 4: 772 (1912).

(Fig. 2)

284 Journ, Hattori Bol. Lab. No. 61 I 986

b a

a h

FIG. 2. Masligolejeunea repleta (TayJ.) Evans. a. Portion of plant , ventral view, x 25. b. Leaf, x 25. ("- d. Cells of leaf-lobe, c from apex, d from middle, x 400. e-f Apical portions of leaf-Iobules, ex 250, Ix 400. g- h. Underleaves, x 25. i- j. Female bracts, x 25. k. Female bracteole, x 25. I. Perianth, ventral view, x 25. Drawn from specim. from Iridia, Foreau 128 (NY).

Plants large, blackish brown in herbaria. Stems 20-50 mm long, 0.2- 0.25 mm in diam., with leaves 2- 3 mm wide, irregularly branched; ventral merophyte of the stem 10-12 cells wide; rhizoids numerous, fasciculate. Leaf-lobes loosely to closely imbricate , widely spreading, oblong, 1.2- 1.6 mm long, 0.8- 1.2 mm wide, apices acute or apiculate, ventral margins usually incurved, often undulate, dorsal margins usually fiat, arched from auriculate bases; apical cells 7-12 x 5- 10 11, median cells 17- 25 x 12- 1511, walls thin, trigones large, cordate, intermediate thickenings few , small, basal cells about 45 x 3211; cuticle smooth; leaf-lobule small , ovate, 1/4-1 /3 as long as the lobe, apex obliquely truncate, apical tooth small, apiculate, 1- 2-celled, or

M , MIZUTANI: Notes on the Lejeuneaceae, 12 285

obsolete; keel nearly straight. Underleaves imbricate, obliquely to erect-spreading, orbicular or reniform, 0,6-0,75 mm long, 0,75- 1 mm wide, apices recurved or flat , lateral margins flat or recurved, insertions sinuate,

Autoicous, Androecium terminal on stem or branch, usually with an apical innovation , innovation often again ftor iferous; bract-Iobes 5- 10 pairs, closely imbricate, obliquely spreading, obovate, about 2/3 as long as leaf-lobes; bract-Iobule oblong, about 1/2 as long as bract-Iobe; bracteoles imbricate, nearly quadrate, slightly smaller than underleaves, Gynoecium usually terminal on the stem, with 1- 2 subftoral innovations, usually again ftoriferous; bract-Iobe oblong or ovate, 1.2- 1.6 mm long, 0,65-0 ,8 mm wide, apex acute or apiculate, margin usually undulate; bract-Iobule oblong or ovate, 2/3- 3/4 as long as bract-Iobe, apex crenate; keel slightly shorter than bract-Iobule, straight; bracteole oblong or obovate, 1-1 ,2 mm long, 0.8- 1 mm wide, apical and lateral margins usually recurved, Perianth oblong or obovate, 1.2- 1.5 mm long, 0,65--0,9 mm wide, sharply 3-keeled, beak short.

Specim, exam,: INDIA: Madras, Wight s,n, - isotype of Lejeunea repleta (G 1924); Perumalmalai - Palni

Hill , 6000 ft, Foreau 128 (NY), BHUTAN: Khegmupa, GriffiTh s,n" del. by Mitten as Lejeunea wardiana (G

19972), THAILAND: Payap, Mt. Chiengdao, A, Touw 8898; -, Fang, on trunk, A, Touw 9444; Uthai Thani,

ML Nang Rum, 400 m, on rock , J, Laohmranich 538; - , ML Hin Dang, on tr4.nk , R, Moennonsri 528b;

Prachinburi, Khao Yai , 750 m, on limestone, K. Larsen eT al. 135; Chantaburi , Pniu waterfall , K. Larsen eT ai, 1859; Rachaburi, Nam Tok, K. Larsen eT aI, 1402; Udawn, ML Luang, A, ToulV 10806, PHILIPPINES:

Luzon, Montalban, H , H. Bartlelf 14378; Dapitan, MicholiTz 33 - type of Mastigolejeunea spectahilis (G

21831), BORNEO: Sabah, Sandakan , W. Meijer 10878, G, Shea 1107; -, Madai cave, 20-100m, W. Meijer

4044a, Z , Iwatsuki 5447,5449,5450,5665, 5676, 5677,568 1,5697,5709, 5761a, NEW GUINEA: Micho{jTZ

11074, del. by Stephani as MasTigolejeunea aTYpos (G),

Range: India, Bhutan, Andamans, Thailand, Philippines, Borneo, New Guinea,

Mastigo/ejeunea rep/eta is closely related to M. humilis, The former can be separated from the latter by the much larger plants. Moreover, they are separated as shown in the following key, I, Stem with leaves about 2,5 mm wide; leaf-lobe usually oblong, apex usually acute; leaf­

lobule small , about 1/4 as long as the lobe; male bract-Iobe obovate, about 2/ 3 as long as leaf-lobe; male bract-Iobule oblong, about 1/2 as long as bract-Iobe; ventral merophyte of the stem 10- 12 cells wide; female bract-Iobe usually with acute or apiculate apex "" "" .

. """" ." ". " .. ""." ."."" ................ ...... ... .................... ...... ",, .......... M . repleta I, Stem with leaves about 1,7 mm wide; leaf-lobe usually ovate, apex usually obtuse; leaf­

lobule about 2/5 as long as the lobe; male bract-Iobe obliquely oblong, about 1/2 as long as leaf-lobe; male bract-Iobule ovate, about 3/4 as long as bract-Iobe; ventral merophyte of the stem 5- 7 cells wide; female bract-Iobe usually with obtuse apex ..... " .. "".,," M . humilis

Verdoorn (1934) reduced the following 4 species to synonyms of Mastigolejeunea rep/eta: M. whightii Steph., M. spiniloba Steph., M. indica Steph., and Brachiolejeunea tylimanthoides Steph. My recent study of the type specimens of these 4 species proved them to be different from M. repleta. M. whightii and B. tylimanthoides are better placed under synonymy of M . humilis. M . spiniloba is con specific with M. virens. M . indica is revived as a proper species.

3) Mastigolejeunea ligulata (Lehm. et Lindenb.) Schiffn. In Engler & Prantl, Nat. P/l ,·fam, 1,3: 129 (1893) ,

Jungermannia Iigulata Lehm, et Lindenb" Pugillus 6: 39 (1834),

(Fig. 3)

286 Journ. Hattori Bot. Lab. No. 61 I 986

q FIG. 3. Mastigolejeunea ligulata (Lehm. et Lindenb.) Steph. a. Portion of plant , ventral

view, x 25. b- c. Leaves, x 25. d-e . Cells of leaf-lobe, d from apex, e from middle, x 400. f­h. Apical portions of leaf-Iobules, fx 250, g- h x 400. i- j. Underleaves, x 25. k. Portion of androecium, ventral view, x 25. 1- 0. Female bracts, x 25. Jrq. Female bracteoles, x 25. r­s. Perianths, ventral view, x 25. Drawn from specim. from New Guinea, J. Klei 5 (NICH

378581 ).

Ptychanthus ligulatus (Lehm. et Lindenb.) Nees, Nat. Eur. Leberm. 3: 212 (1834). Phragmicoma ligulata (Lehm. et Lindenb.) Gott. et aI., Synop. Hepat.: 301 (1845). Lejeunea ligulata (Lehm. et Lindenb.) Mitt., Hepat. lndiae orient.: 110 (1861). L. novohibernica Schiffn., Forsch. Gazelle 4: 23 (1890). Mastigolejeunea novohibernica (Schiffn.) Steph., Spec. Hepat. 4: 770 (1912). M. obtusiloba Steph., Spec. Hepat. 4: 768 (1912).

M, M IZUTAN I: Notes on the Lejeuneaccae, 12 287

Plants large, brown in herbaria, Stems 20- 50 mm long, 0,17- 0,2 mm in diam" with leaves 2- 2,5 mm wide, irregularly branched; ventral merophyte of the stem 6- 7 cells wide; rhizoids few , Leaf-lobes loosely imbricate, widely or often obliquely spreading, oblong, 1- 1.3 mm long, 0,6-0,7 mm wide, apices rounded or often obtuse, margins nearly flat , dorsal margins somewhat arched from slightly auriculate bases; apical cells 12- 17 x 10- 13 jA, median cells 20--32 x 1 2-221~,

walls thin, trigones large, cordate, intermediate thic kenings few, small, basal cells abou t 32 x 30 jA ; cut icle smooth; leaf- lobule small, ovate, flat , 1/5- 1/4 as long as the lobe, apex truncate, apical tooth small, 1- 2-celled; keel nearly straight. Underleaves approxi mate or imbricate, reniform, 0.45-0,55 mm long, 0, 7-{),g mm wide, apica l and lateral margins flat or slightly recurved or often undulate, insert ions nearly transverse.

Dioicous. Androecium terminal on stem or branch, usually with an apical innovation , often again floriferous; bract-Iobes 10-25 pairs, loosely or some:.vhat imbricate, widely spreading, oblong, about 4/5 as long as leaf-lobes; bract-Iobules distant from each other, ovate with ligulate apices, about 2/5 as long as bract-Iobes; bracteoles imbricate, reniform, slightly smaller than underleaves. Gynoecium terminal on stem or branch, with 1- 2 subfloral innovations, innovation usually again floriferou s; bract-Iobe oblong or linear, 1.1- 1,3 mm long, 0.4-0.5 mm wide, apex rounded , obtuse to acute; bract-Iobu le oblong, linear or narrowly triangu.lar, 1/2-3/4 as long as bract-Iobe, apex truncate, rounded, obtuse to acute; keel about 1/2 as long as bract-Iobule, straight; bracteole oblong, 0.95- 1.1 mm long, 0.5- 0.55 mm wide, apex rounded or retu se, undulate or recurved, Perianth oblong, 1- 1.3 mm long, 0.5 0.6 mm wide, sharply 3-keeled, beak short,

Specim, exam,: THAILAND: Surat, Chumphon 50 m, K Larsen el ai, 1484; Yala , Kue Long waterfallS,

I 50- 250 m, C. Charoenphol el ai, 4177, MALAY PEN,: Pulo Penang, [herb, Hooker, herb, Kew,j without

collecter's name - isotype of Jungermannia ligulata (G 1707), BORNEO: Sabah, Gaya I., 1- 50 m, M, Mizutani 33 12; - , Sandakan, 400 ft , G, Shea 1081, 1085, S , Kokawa & M, Hotta 48; - , Madai cave,

10-30 m, Z . Iwatsuki 5409, 54 10, 5761b; - , Mt. Silam, 25- 250 m, Z. Iwatsuki 5622a, G. Shea 1903, 1984, AMBON: Air Besar, 10 m, on trunk , H. Akiyama 9252 (KYO), SERAM: W. v. Basse s,n" det. by Stephani as Mastigolejeunea novohibernica (G 11703); Manusela Nat. Park , Ka loa - Pasahari, 20- 120 m, H. Akiyama 9446 (KYO); -, Sawai , 1-130 m, H. Akiyama 9523 (KYO); Bula, 30 m, H. Akiyama 10749 (KYO);

Bula, Gung Sentoten, 30-100 m, on branches, H. Akiyama 10797 (KYO) , NEW Guinea: W. Micholit z s,n. - type of Mastigolejeunea obtusiloba (G 11690); Butaneng, L. Karubach 15 (G 1708); NE New Guinea,

Morobe, Gabensis Creek, 120m, H. Streimann & T. Umba 11269; - , - , Wau , S , Saito 5, 8; - , East S~pik ,

Kairurll I., 600 m, 0, W. Borrel! 81 - 9, 81- 10, J. Klei 5, MICRONESIA: Guam, D. R, Smith 1577. SAMOA:

Rechillger 3230 (G 1706), TAHITI: Nadeaud 15 (G 1705). Range: Thailand, Malay Pen " Borneo, Ambon, Seram, New Guinea, GlIa m, Samoa, Tahiti ,

Mastigolejeunea ligulata is readily separated from other Asian members of the genus Mastigolejeunea by the male bracts with ligulate lobules which are distant from each other. Moreover, this species has the following diagnostic characters: (I) the oblong leaf-lobe with a rounded or often an obtuse apex; (2) the male bract-lobe about same in fo rm and size as the leaf-lobe; and (3) the dioicous plant.

4) Mastigolejeunea virens (Aongstr.) Steph. Spec, Hepat. 4: 776 (1912),

Thysananthus virells Aongstr. , bfv, K . Vetcnsk , Akad. Forh, 5: 131 (1873) .

Lejeunea virens (Aongst r.) Steph , in Besch" J , de Bot. 12: 141 (1898), Mastigolejeunea laitica Steph" Hedwigia 35: 11 2 (1896),

Archilejeullea tahitensis Steph" Spec, Hepat. 4: 732 (191 1).

(Fig, 4)

288 Journ . Hattori Bot. Lab. No. 61 I 9 8 6

O e t

d

tfIl: pOr q ~g

FIG. 4. Mastigolejeunea virens (Aongstr.) Steph. a. Portion of plant, ventral view, x 25. b-e. Leaves, x25. I - i. Cells of leaf-lobes,f-g from apices, h- i from middle, x 400. j­

I. Apical portions of leaf-lobules, x 400. m- no Underleaves, x 25. o. Gynoecium and subfloral innovation, x 25. jrq. Female bracts, x 25. r. Female bracteole, x 25. s- t . Perianths, ventral view, x 25. Nos. a- c,/, h,j, m- a from isotype of Thysananlhus virens from Tahiti, N. J. Anderson S.n. (G 12495); others from Tahiti, H. Hurlimann 1123 (NICH 365550).

Mastigolejeunea appendiculifolia Steph. , Spec. Hepat. 4: 773 (\912). M. longispina Steph., Spec. Hepat. 4: 774 (1912). M. spiniloba Steph., Spec. Hepat. 4: 775 (\9 12). Phragmicoma taitica Gott. ex Steph. , Hedwigia 29: 8 (1890), nom. nud o

M. MIZUTANI: Notes on the Lejeuneaceae. 12 289

Plants small , olive brown in herbaria. Stems 10- 20 mm long, 0. 12- 0.17 mm in diam., with leaves 1.l-1.7mm wide, irregularly branched; ventral merophyte of the stem 5- 7 cells wide; rhizoids numerous, fasciculate. Leaf-lobes closely imbricate, obliquely spreading, oblong or ovate, 0.7-1 mm long, 0.5- 0.7 mm wide, apices obtuse, ventral margins incUfved, dorsal margins somewhat arched from somewhat auriculate bases; apical cells 7- 15 x 5- 8 j1, median cells 15- 30 x 12-15 j1, walls thin , trigones large, cordate, intermediate thicken ings few, basal cells about 45 x 30 j1; cuticle smooth; leaf-lobule small, 1/4-1 /3 as long as the lobe, oblong, apex obliquely truncate, apical tooth elongate, 4-6 cells long, 1- 2 cells wide; keels nearly straight. Underleaves closely imbricate, obliquely spread ing, about 3 times as wide as the stem, reniform, 0.3- 0.42 mm long, 0.45-0.55 mm wide, apical and lateral margins flat or slightly recUfved, insertions nearly transverse .

Autoicous. Androecium terminal on stem or branch, usually with apical innovation; bract­lobes 10- 18 pairs, closely imbricate, obliquely spreading, obliquely oblong, about 2/3 as long as leaf-lobes; bract-lobule oblong with ligulate apex, about 3/5 as long as bract-lobe; bracteoles approximate or imbricate, nearly quadrate, slightly smaller than underleaves. Gynoecium terminal on stem or branch, with 1- 2 subfloral innovations, innovation usually again fioriferous; bract-Iobe oblong, about 1.1 mm long, 0.6 mm wide, apex obtuse or rounded; bract-lobule oblong, 2/5- 1/2 as long as bract-Iobe, apex irregularly crenate; keel about 1/2 as long as bract­lobule , straight; bracteole oblong, about 0.75 mm long, about 0.5 mm wide, apex rounded to retuse. Perianth oblong or obovate, 0.8-1.1 mm long, 0.6-0.75 mm wide, sharply 3-keeled, keels somewhat crenate in the upper half, beak short.

Specim. exam.: CEYLON: Paradenya, 500 m, V. Schiffner s.n. (Verdoorn, Hep. Se!. Crit. 6: 254 as M. humilis); Paradenia, 470 m, on trunk, N. S. Parihar s.n . THAILAND: Nakon Rachsima, Sakacrat, Wang

Chongarg, on trunk, R. Mernnonsee 616. MALAY PEN .: Penang, N. Kitagawa 14491. PHILIPPINES:

Luzon, Mt. Banahao, 800-1250 m, on decaying log, Z. Iwatsuki et al. 14415, on trunks, Z . Iwatsuki et al. 14421, 16023a, 16464; - , Baguio, on trunk, Alvarez & Saprid 78205, 78206. BORNEO: Sabah, Mt.

Kinabalu , 600-1500 m, M. Mizutani 2030, 2052, 2054, 2055, 3067, 3119; -, Ranau, 1500 ft, on trunk , G.

Shea 1206; - , Batn Puteh, on branches of bush, G. Shea 3281; Kalimantan , Taraken 1., low alt., on trunk, . W. Meijer 4838. JAVA: G. Megamendoeng, 1200 m, on trunk, V. Schiffner s.n. (Verdoorn, Hep . Se!. Crit. 6: 256 as M. humilis); Buitenzorg, 260 m, on trunk, V. Schiffner s.n. (Verdoorn, Hep. Se!. Crit. 6: 255a as M. humilis); Tjibodas, 1425 m, W. Meijer 3798; Gede, 1400-1700 m, van Leeuwen-Reijnvaan 11892, 12541 (NY); Bogor, M. Fleisher s.n. (NY); G . Pandansari , 1600m, F. Verdoorn s.n . (NY). SERAM; Manusela Nat. Park,

Uwelehu, 630-1000 m, H. Akiyama 9252 (KYO); -, Ruamah Sokat Batu, 1- 230m, on trunks, H. Akiyama 9893, 9898 (KYO); Taniwel , Batu Patih - Batu Soli , 400- 700 rn, H. Akiyama 10038 (KYo); -, Gunung Kakaela, 10-450 m, on cliff, H. Akiyama 10127, 10146 (KYO). NEW G UINEA: NE New Gu inea, Mt. Hagen,

1160 m, on trunk, H. Streimann 21045; - , Central, Narirogi creek, 440 rn , on rock , H. Streimann & E. K Naoni 15067; -, Bulolo, 800-1000 m, G. Shea 6574, 8970, 8972. MICRONESIA: Guam, D. R. Smith 1325. TAHITI: Visco s.n . - syntypes of Mastigolejeunea taitica (G 12174, 12179); Nadeaud 702 - type of Archilejeunea tahitensis (G 12316); Eimeo, N. J. Anderson s.n. - isotype of Thysananthus virens (G 12495); Papehue river, W. A . Setchell & H. E. Parks 5348; Papeete, 40 rn, on rock, H. Hurlimann 1123. NEW

CALEDONIA: Insula Pinarum, LeRat S.n. - type of Mastigolejeunea appendiculifolia (G 12395).

Range: Cey10n , Thailand, Malay Pen ., Philippines, Borneo, Java, Sera m, New Guinea, Guarn , Tahiti, New Caledonia.

Verdoorn (1934) reduced Mastigolejeunea virens to a synonym of M. humilis. However, my recent study of the type specimen, M . virens indicates that it should be revived as a proper species. Verdoorn (1934 ) reduced Mastigolejeunea taitica, M. appendiculifolia, M. /ongispina, Archi/ejeunea tahitensis and Phragmicoma taitica into the synonymy of M. humilis, and reduced M. spini/oba to a synonym of M. rep/eta.

290 Journ. Hattori Bot. Lab. No. 61 I 9 8 6

However, these 6 species and M . virens are conspecific . M. humilis is related to M . virens; however, they are separated as shown in the

following key. 1. Apical tooth of leaf-lobule elongate, 3- 6 cells long; leaf-lobe oblong, apex always obtuse:

leaf-lobule 1/4-1 /3 as long as leaf-lobe; keel of female bract about 1/2 as long as bract-lobule; plants olive-brown; stems and branches mostly creeping on substrata .... ... .... . .. .

... .. . .. .. . ..... .. ...... ... ..... ......... . .. .... .... .. . ...... . ..... .. .... .... ... .. . ....... . ... .. ... M . virens I. Apical tooth of leaf-lobule small or obsoletc, 0- 2 cells long; leaf-lobe ovate, apex rounded,

obtuse or acute; leaf-lobule about 2/5 as long as leaf-lobe; keel of female bract 2/ 3- 1 as long as bract-Iobule; plants blackish brown; stems and branches mostly ascending from substrata ... ... .. .... .. .. .... .. .... . ... . .. ... .......... .. . .. ......... ..... ............ ... ..... .. M. humilis

5) Mastigolejeunea undulata Gradst. et Grolle (Fig. 5) In Hiepko & Schultze-Motel, Mensch, Kultur u. Umwelt im zentr. Bergland von W-Neuguinea 7: 13 , f. 7 (1981).

Lopholejeunea robinsonii Steph., Icones Hepaticarum inedit. : 8485; Verd ., Ann . Bryol. suppl. 4: 104 (1934), nom. nudo

Plants medium in size or large, olive-green to reddish brown in herbaria . Stems 10- 50 mm long, 0.13-0.15 mm in diam., with leaves 1.5- 2 mm wide, irregularly branched; ventral mero­phyte of the stem 6- 8 cells wide; rhizoids few . Leaf-lobes densely imbricate, obliquely to widely spreading, ovate, 0.85-1.1 mm long, 0.5- 0.75 mm wide, apices acute or apiculate, ventral margins incurved or somewhat undulate , more or less auriculate at bases, dorsal margins flat or sometimes narrowly recurved , arched from slightly auriculalte bases; apical cells 5- 12 x 5- 10 IJ, median cells 20-30 x 12- 181J, walls thin, trigones large , cordate, intermediate thickenings few but large, basal cells about 35 x 251J; cuticle smooth; leaf-lobule small, ovate, about 1/4 as long as the lobe, apex truncate, apical tooth elongate, 3- 4 cells long, 1- 2 cells wide at base; keel nearly straight. Underleaves loosely to densely imbricate, obliquely to erect-spreading, reniform, 3- 4 times as wide as the stem, 0.35- 0.5 mm long, 0.45- 0.7 mm wide, apices truncate or retuse, usually recurved, lateral margins flat or slightly recurved, auriculate at bases, insertions sinuate.

Dioicous. Androecium terminal on stem or branch, often with an apical innovation; bract­lobes 5- 12 pairs, imbricate, obliquely spreading, ovate, slightly shorter than leaf-lobes, apices apiculate; bract-Iobule ovate , about 2/5 as long as bract-Iobe, apex apiculate; bracteoles imbricate or approximate, obliquely or erect-spreading, reniform, about as large as underleaves, apices recurved . Gynoecium terminal on stem or branch, with 1-2 subfloral innovations, innovation usually again floriferous; bract-Iobe oblong, 1- 1.4mm long, 0.42-0.6 mm wide, apex acute or apiculate or obtuse; bract-Iobule oblong, 1/2- 3/5 as long as bract-Iobe, apex rounded to acute; keel straight, 1/2- 2/3 as long as bract-lobule; bracteole oblong, 0.85-1 mm long, 0.55- 0.7 mm wide, apex rounded to slightly retuse, recurved or undulate. Perianth oblong. 1.3- 1.8 mm long, 0.55-0.75 mm wide, usually lO-keeled (5 ventral, 2 lateral, 3 dorsal), lateral keels sharp, often with scale-like appendages, ventral and dorsal keels rounded and smooth, beak short.

Specim. exam.: PHILIPPINES: Luzon, Nueva, Vizcaya, McGregor 2021 7, det. by Stephani as Lopholejeunea robinsonii (G). SERAM: Taniwel, Buria, 300-400 m, on trunks, H. Akiyama 9965, 9967, 9972, 9985 (KYO); Manusela Nat. Park , Sawai, 1- 130m, on trunk, H. Akiyama 9501 (KYO); - , Wae Ili - Sawai, 80{}-1230m, on trunk, H. Akiyama 9890 (KYO); - , Hatu - Piliana, 1-400 m, on trunk, H. Akiyama 10247 (KYO); - , Hatumete - Hoale pass, 20{}-640m, on trunk, H. Akiyama 10467 (KYO). NEW GUINEA: NE New Guinea, Western Highlands, Nebilyer river, 2760 m, on trunk, H. Streimann 20530; - , Eastern Highlands,

M. MIZUTANI: Notes on the Lejeuneaceae. 12

-- p FIG. 5. Mastigolejeunea undulata Gradst. et Grolle. a. Portion of plant, ventral view,

x 25. b. Leaf, x 25. c- e. Cells of leaf-lobes, c-d from apices, e from middle, x 400. I - h. Apical portions of leaf-Iobules, x 400. i- j. Underleaves, x 25. k- n. Female bracts, x 25. 0 -

p. Female bracteoles, x 25. q- s. Perianths, q- ventral view, r-s-<lorsal view, x 25. Drawn from specim. from Seram I., H. Akiyama 9967 (KYO).

291

Goraka - Kundiawa, 2650 m, on base of tree, van Zanten 68- 447/a; - , Morobe, Smanzing, 1800m, Clemens 94 I Obis/Hf - syntype of Mastigolejeunea undulata (NICH 360161); - , - , Kasanombe - Namba, 1600-1700 m, H. Inoue 32 137 (TNS); - , Central , Woitape, 1500m, H. Inoue 30428 (TNS); -, -, Mt. Albert Edward, 2000--2300m, H. Inoue 31262, 31381., 31568, 31581 (TNS).

Range: Philippines, Seram, New Guinea.

The distinguishing characters of Mastigolejeunea undulata are: (1) the leaf-lobe with an acute orapiculate apex; (2) the leaf-lobule with an elongate tooth; (3) the leaf-

292 Journ. Hattori Bot. Lab. No. 61 I 986

lobe with more or less undulate and arched ventral margin; (4) the small, auriculate bases of lateral margins of underleaf; (5) the dioicous plants; (6) the perianth with 10 keels; and (7) the female subfloral innovation with a conspicuously small first lateral leaf which is about 1/2 as long as the neighboring leaves. The characters 6 and 7, mentioned above, are quite different from the other Asiatic species of M. humilis group, which has a 3-keeled perianth and a large first lateral leaf of the female subfloral innovation, which is slightly smaller than, or about as large as, the neighboring leaves.

In appearance M. undulata resembles Thysananthus planus Lac., particularly in the outline of leaves and underleaves. However, they are separated as shown in the following key. 1. Leaf-lobe with long vitta and flat dorsal margin; leaf-cells with thick walls and triangular

trigones; female bract-Iohe and bracteole with dentate margins; perianth 3-keeled ........ . .... ......... ..... . ... . .... . ..... . .. ......... ..................... . ... . . ...... . . ....... Thysananthus planus

1. Leaf-lobe with recurved dorsal margin and without vitta; leaf-cells with thin walls and cordate trigones; female bract-Iobe and bracteole with entire margins; perianth 10-keeled

.... ........ . . .. .. .... . . . .. . . . . . . . . . . .. ..... . . . . ... . . . ......... ... ..... . .. . ...... Masligolejeunea undulata

6) Mastigolejeunea truncata Mizut. , sp. novo (Fig. 6) Planta minor, fusco-olivacea, fiaccida. Caulis brevis , irregulariter ramosus. Folia imbricata, ovata ve l

triangu lata, apice obtuso, margine undulato, recurvo, lobulus parvus, rectangulatus . Amphigastria reni­formia , transverse inserta. Autoica. Androecia in ramis term in alia, bracteis 5- 10 jugis. Gynoecia in caulis vel ramis terminalia. perianthia oblonga, alte triplicata, apice late truncato, margine integerrimo. recurvo.

Plants small, flaccid, olive-brown in herbaria. Stems 7- 15 mm long, 0.12-0.15 mm in diam., with leaves 1.2-1.7 mm wide, irregularly branched; ventral merophyte of the stem 6-8 cells wide; rhizoids few. Leaf-lobes imbricate, widely spreading, ovate or triangular, 0.85- 1 mm long, 0.55-0.68 mm wide, apices obtuse, more or less incurved, ventral and dorsal margins undulate and narrowly recurved; apical cells 12- 17 x 7- 10 fl, median cells 25- 35 x 10-13 11, walls thin , trigones large, cordate, intermediate thickenings absent, basal cells about 45 x 32/1; cuticle smooth; leaf-lobule very small , rectangular, 1/5- 1/4 as long as the lobe, apex obliquely truncate, apical tooth absent, merely angular; keel st raight-. Underleaves distant, about 3 times as wide as the stem, reniform, 0.32- 0.37 mm long, 0.45- 0.55 mm wide, margins flat or slightly recurved , insertions nearly transverse .

Autoicous. Androecium terminal on branch, apical innovation absent; bract-Iobes 5- 10 pairs, densely to loosely imbricate, obliquely spreading, oblong, 1/2-2/3 as long as leaf-lobes, apices obtuse; bract-Iobule oblong, about 2/3 as long as bract-Iobe; bracteoles distant or imbricate, obliquely spreading, nearly quadrate, slightly smaller than underleaves. Gynoecium terminal on stem or branch, usually with I subfloral innovation, innovation usually again floriferous; bract-Iobe oblong or lanceolate, 1- 1.\ mm long, 0.5- 0.55 mm wide, apex obtuse or acute; bract-Iobule oblong, about 1/2 as long as bract-Iobe , apex rounded or slightly crenate; keel 1/3- 1/2 as long as bract-Iobule, straight; bracteole oblong or obovate, 0.7- 0.8 mm long, 0.65-0.7 mm wide, margin irregu larly undulate and recurved . Perianth oblong, 1-1.3 mm long, 0.55 mm wide, sharply 3-keeled, mouth wide, narrowly recurved .

Specim. exam.: BORNEO: Sabah, Madai cave, 30-70 m, on limestone, Z. lwatsuki 5426 ~ holotype (NICH

255426). SERAM: Seram Utara, Wasa - Roho, 60 m, H. Akiyama 2429 (K YO); Manusela Nat. Park, Elemata Makulaina, lOOm, on boulder, H. Akiyama 9321 (KYO).

Range: Borneo, Seram.

M. MI ZUTANI: Notes on the Lejeuneaceae. 12

g

v p

FIG. 6. Mastigolejeunea truncata Mizut. a. Portion of plant, ventral view, x 25. b- d. Leaves, x 25. e- g. Cells of leaf-lobes, e-f from apices, g from middle, x 400. h- i. Apical portions of leaf-Iobules, x 400. j-k. Underleaves, x 25. 1-0. Female bracts, x 25. p-q. Female bracteoles, x 25. r- t. Perianths, r, t-ventral view, s-dorsal view, t- young perianth, x 25. Drawn from holotype from Borneo, Z. Iwatsuki 5426 (NICH 255426).

293

This new species has a very unique character for the Lejeuneaceae, i.e. the perianth has a wide mouth even in every growing stage; the mouth is about as wide as the perianth. Moreover, this species has the following diagnostic characters: (1) the leaf-lobe with an undulate and narrowly recurved margin; (2) the leaf-lobe gradually narrower toward the suddenly narrowed obtuse apex; (3) the very small leaf-lobule about 10 cells long; (4) the small underleaves usually distant from each other; (5) the

294 Journ. Hattori Bot. Lab. No. 61 198 6

flaccid, olive-brown, not secondarily pigmented plants. However, this species belongs to Mastigolejeunea because of (1) the leaf with a small, folded lobule, (2) the reniform underleaves, (3) the gynoecium with 2 bracts and 1 bracteole, (4) the Radula-Lejeunea type of branching of female subflora l innovation, (5) the female and male bracts and bracteoles with entire margins, (6) the leaf-cells with thin walls and cordate trigones, (7) the sharply 3-keeled perianth, (8) the Lejeunea type branching of the vegetative branches, and (9) the small flagella on the basal portion of the stem.

7) Mastigolejeunea recurvifolia Mizut., sp. novo (Fig. 7) Planta minor, fusco-olivacea, fragillima . Cauli~ brevis , irregulariter ramosus. Folia imbricata, ovata,

apice obtuso, margine recurvo, lobulus parvus, ovatus. Amphigastria reniformia, transverse inserta. Autoica. Androecia in caulis vel ramis terminalia, bracteis 5'- 10 jugis. Gynoecia in caulis vel ramis terminalia, perianthia oblonga, alte triplicata, rostro brevissimo.

Plants small, pale olive-brown in herbaria, somewhat fragile. Stems 10-20 mm long, 0.15- 0.2 mm in diam., with leaves 1.2- 1.7 mm wide, irregularly branched; ventral merophyte of the stem 6-7 cells wide; rhizoids few. Leaf-lobes imbricate, widely spreading, ovate, 0.65- 0.8 mm long, 0.48- 0.53 mm wide, apices obtuse, ventral and dorsal margins recurved; apical cells 10-15 x 8-10 11, median cells 20-30 x 12- 1811, walls thin , trigones large, cordate, intermediate thickenings absent, basal cells 42 x 3711; cuticle smooth; leaf-lobule very small , ovate, 1/5- 1/4 as long as the lobe, apex truncate, apical tooth apiculate, 1- 2-celled, or merely angular; keel straight. Underleaves imbricate, obliquely spreading, reniform, 3- 4 times as wide as the stem, 0.3- 0.42 mm long, 0.45- 0.55 mm wide, apical and lateral margins flat or slightly recurved , insertions transverse.

Autoicous. Androecium terminal on stem or branch, usually with an apical innovation, innovation often again floriferous; bract-Iobes 5- 10 pairs, densely imbricate, obliquely to suberect-spreading, oblong, about 2/3 as long as leaf-lobes, apex obtuse; bract-Iobule oblong, 2/3- 4/5 as long as bract-Iobe, apex rounded; bracteoles distant to imbricate, obliquely to suberecl-spreading, nearly quadrate, slightly smaller than underleaves. Gynoecium terminal on stem or branch, with I subfloral innovation, innovation usually again flor iferous; bract-Iobe ovate or oblong, 0.9- 1.2 mm long, 0.5-0.6 mm wide, apex obtuse; bract-Iobule ovate or oblong, 1/2- 2/ 3 as long as bract-Iobe, apex irregularly crenate; keel 1/2- 2/3 as long as bract-Iobule; bracteole obovate or oblong, 0.8- 0.95 mm long, 0.6- 0.7 mm wide, margin more or less crenate and recurved. Perianth oblong, 1- 1.3 mm long, 0.5- 0.6 mm wide, sharply 3-keeled, keels smooth, beak short.

Specim. exam. : BORNEO: Sabah, Madai Cave, 30-100 m, on limestones, Z. Iwatsuki 5427, 5685; - , Batu Puteh, on limestones, G. Shea 3278 - holotype (NICH 334833), 3293.

Range: Borneo.

This new species is related to Mastigolejeunea truncata . Both species have in common: (1) the small leaf-lobule; (2) the leaf-lobe with a recurved margin and an obtuse apex; (3) the olive-brown, not secondarily pigmented plants; and (4) the gynoecium usually with I subfloral innovation which is usually rapidly again floriferous . However, M . lruncala can be separated from M. recurvifolia by the following key. 1. Mouth of perianth wide; margin of leaf-lobe undulate; trigones of cells of leaf-lobule small

or absent; underleaves distant; plants flaccid ..... . .... . .......... .............. . .. . M . lruncata

M. MIZUTANI: Notes on the Lejeuneaceae. 12

o k

FIG. 7. Masligo/ejeunea recurvijolia Mizut. a. Portion of plant, ventral view, x 25. b­d. Leaves, x 25. e-f. Cells of leaf-lobe, e from apex, f from middle, x 400. g- i. Apical portions of leaf-Iobules, x 400. j- k. Underleaves, x 25. /-0. Female bracts, x 25. p- q. Female bracteoles, x 25. r- s. Perianths, ventral view, x 25. Drawn from holotype from Borneo, G. Shea 3278 (NI CH 334833).

295

1. Mouth of perianth beaked; margin of leaf-lobe not undulate; trigones of cells of leaf-lobule large; underleaves imbricate; plants fragile .............. .......... ....... ........ M. recurvifolia

8) Mastigolejeunea indica Steph . (Fig. 8) Spec. Hepat. 4: 776 (1912).

Plants medium in size, dark brown in herbaria. Stems 10- 30 mm long, 0.17- 0.2 mm in diam., with leaves 1.5- 2 mm wide, irregularly branched; ventral merophyte of the stem 8-10 cells

296 Journ. Hattori Bot. Lab. No. 61 I 9 8 6

d

o· J G

f

e

o k

n Q

FIG. 8. Mastigalejeunea indica Steph. a. Portion of plant, ventral view, x 25. b-f Leaves, x 25. g- h. Cells of leaf-lobe, g from apex, h from middle, x 400. i. Teeth of leaf­lobule, x 250. j - k. Underleaves, x 25. I-a . Female bracts, x 25. p-q. Female bracteoles, x 25. r. Perianth, ventral view, x 25. Drawn from specim. from Thailand, O. Thaithang 371

(NICH 359994).

wide; rhizoids numerous, brown, fasciculate. Leaf-lobes densely imbricate, obliquely spreading, ovate, 0.8-1 mm long, 0.6-0.75 mm wide, apices rounded or obtuse, often incurved, ventral margins flat or incurved, often slight ly undu late, dorsal margins arched from more or less auriculate bases; apical cells 5- 8 x 5- 10 /l, median cells 20- 25 x 15- 18/1, walls thin, trigones large, cordate, intermediate thickenings few but large, basal cells about 37 x 30 /l; cuticle smooth; leaf-lobule large, ovate or triangular, about 1/2 as long as the lobe, apex acuminate, free margin with 2- 4 large teeth, teeth triangular, 3- 8 cells long, 4-8 cells wide at base, apices acute or obtuse, margins of teeth more or less recurved and often with a small secondary tooth; keel

M . MIZUTANI: Notes on the Lejeuneaceae. 12 297

straight or slightly arched . Underleaves approximate or imbricate, 3-4 times as wide as the stem, reniform, 0.25- 0.4 mm long, 0.5- 0.7 mm wide, margins usually narrowly recurved, insertions sinuate.

Autoicous. Androecium terminal on stem or branch, apical innovation rare; bract-Iobes 5- 8 pairs, densely imbricate, obliquely to erect-spreading, oblong, 1/ 2- 2/3 as long as leaf-lobes; bract-Iobule slightly shorter than bract-Iobe, oblong; bracteoles imbricate, obliquely spreading, nearly quadrate, slightly smaller than underleaves. Gynoecium terminal on stem or branch, with 1- 2 subfloral innovations, innovation often again floriferous; bract-Iobe ovate or oblong, 0.8- 0.9 mm long, 0.45- 0.6 mm wide, apex usually obtuse, margin somewhat undulate; bract­lobule oblong or obovate, 1/2- 3/4 as long as bract-Iobe, apex rounded or irregularly crenate; keel 2/3- 3/4 as long as bract-Iobule, nearly straight; bracteole oblong or obovate, 0.7- 0.8 mm long, 0.5-0.55 mm wide, apical and lateral margins flat or irregularly recurved. Perianth oblong or obovate, 0.9- 1.2 mm long, 0.6-0.75 mm wide, 3-keeled, ventral keel usually with 1- 2 weak additional keels, beak short.

Specim. exam.: THAILAND: Udawn, Ban Na Luang, 370 m, on trunk , A. Tou>!' 10364; U-Thai Thani, Kao Hin Dang, on fallen branch. O. ThailllOng 371; Nakon Rachsima, Sakaerat, on trunk . R. Mernnonsee 624;

Rachaburi, Huai Yang. K. Larsen el al. 1343. NICOBARS: E. H. Man s.n . - type of Masligolejeunea indica (G

14988). CHINA: Yunnan: Xishuangbanna , on trunk . W. X. XU 11027. Range: Thailand. Nicobars. China.

Verdoorn (1934) reduced Masligo/ejeunea indica to a synonym of M . rep/eta. However, my recent study of the type specimen of M. indica revealed it to be a proper species. In sterile condition M . indica resembles a species of Acro/ejeunea because of the large leaf-lobule with large wave-like teeth. However, M. indica belongs to the Masligolejeunea because of (I) the 3-keeled perianth, (2) the entire leaf-lobes, underleaves, male and female bracts and bracteoles, (3) the cordate trigones of leaf­cells, and (4) the gynoecium with 1- 2 subfloral innovations of Radu/a-Lejeunea type branching.

ACKNOWLEDGMENTS . r thank the directors and curators of the following herbaria , who sent me on loan many important specimens, including types: Conservatoire et Jardin bota­niques, Geneve (G), Kyoto University. Kyoto (KYO), The New York Botanical Garden , New York (NY), and National Science Museum, Tokyo (TNS). Thanks are also due to Dr. S. Hallori of the Hattori Botanical Laboratory for his critical advice, and to Dr. A. J. Sharp of the University of Tennessee for his suggestions concerning the English text.

REFERENCES

Gradstein , S. R. 1975. A taxonomic monograph of the genus Acrolejeunea (Hepaticae). Bryo­phytorum Bibliotheca 4: 1- 162. pI. 1- 24.

Verdoorn, F . 1934. Die Lejeuneaceac Holostipae der Indomalaya unter Beriicksichtigung siimtlicher aus Asien, Australien, Neu Seeland und Ozeanien angefiihrten Arten . Ann . Bryol. suppl. 4: 40-192, f. 4-7.