Journ. Hattori Bot. Lab. No. 62,' 191-200 (June 1987)

A PRELIMINARY STUDY OF THE GENUS RADULA FROM ,

KOHSAKU Y AMADA 1

Twenty-two species of the genus Radula have been recorded in the literature from Australia (induding Tasmania) by Hampe (1838), Stephani (1884, 1889, 1910), Castle (1963, 1964, 1967), Yamada (1979, 1982, 1984), and Windolf (1985). Eleven are en­ demic to Australia: Radula australiana, R . compacta, R. hicksiae, R. jovetiana, R. novae-hollandiae, R. ocellata, R. patens, R. pulchella, R. rathokowskiana, R. tasmanica, and R. wattsiana. However, most of them are known only in the southeastern part of Australia, such as New South Wales, and Tasmania. The Radula flora of Queensland has not been studied adequately, even though in recent years Yamada (1984) has re­ ported 9 species (induding 3 new species). Recently I had an opportunity to study many collections of Radula from Que­ ensland made by Dr. I. G. Stone, Dr. H. Streimann, Dr. B. M. Thiers, and Dr. M. 1. Hicks. In the present paper, I recognize 18 species, of which 2 species are new to science. I wish to express my sincere thanks to Dr. G. A. M. Se ott of Monash University (Aus­ tralia), Dr. M. L. Hicks of Appalachian State University (North Carolina), Dr. H. Streimann of Canberra Botanic Gardens (Canberra), and Dr. B. M. Thiers of The New York Botanical Garden (New York), for the loan of Radula specimens. Thanks are also due to Dr. S. Hattori of the Hattori Botanical Laboratory for his critical advice, and to Dr. A. J. Sharp of the University of Tennessee, for his kind help with the English of the manuscript.

Subgenus Odontoradula Yamada Key to the species 1. Leaf-lobes with vitta ...... 2. R. ocellata 1. Leaf-lobes without vitta ...... 2 2. Leaf-lobes with subacute apices, the margins of apical portion entire; walls of leaf- cells without trigones; female bracts of two pairs ...... 1. R. novae-hollandiae 2. Leaf-lobes with acute apices, the margins of apical portion with 1-3 acute teeth; walls of leaf-ceIls with medium-sized trigones; female bracts of one pair ...... 3. R. queenslandica

1. Radula novae-hollandiae Hampe In Lehmann, Nov. Min. Cogn. Strip. Pugillus 7: 24 (1838). Specim. exam.: Werwah, 80 m, J. Windolf813 (hb J. Windolf, hb K. Yamada); Mt. Bethongabel, 900 m, B. M. Thiers 1238 (NV); trail to Elebana Falls, 500 m, B. M. Thiers 1110 (NV); trail to Lyre Bird Lookout, 1000 m, B. M. Thiers 1194 (NV); track to Mt. Bethongabel, 900 m, B. M. Thiers & R. H.

1 941-47 Funaeyama, Seta-cho, Ise-shi 516, Mie-ken; also the Hattori Botanical Laboratory, Obi, Nichinan-shi, Miyazaki-ken 889-25. 192 Journ. Hattori Bot. Lab. No. 62 1 987

Hailing 2786 (AD, NY); Mt. Cool urn, 60-80 m, B. M. Thiers 2851 (AD, NY); Mt. Lewis SW of Mossman, M. L. Hicks 11134, 11394 (BOON, NICH); Mchugh Bridge along W of Innisfail, M. L. Hicks 10973 (BOON, NICH); Mt. Bartle Frere, M. L. Hicks 11031,11032,11228 (BOON, NICH); Devil's Thumb W of Mossman, 1150 m, M. L. Hicks 11274 (BOON, NICH); Mt. Spurgeon W of Mossman, M. L. Hicks 11101 (BOON, NI CH). Distr.: Endemic to Australia. This species has not known from Queensland until recently. Yamada (1984) and Windolf (1985) reported this species from Queensland where it seems to be one of the commonest species.

2. Radula ocellata Yamada J. Hattori Bot. Lab. 45 : 209 (1979). Specim. exam.: Mt. Lewis, 1. G. Stone (MUCV 6093); along N of Johnstone River, 750 m, B. M. Thiers & R. E. Halling 2180 (AD, NY); Mt. Bellenden Ker, 1500 m, B. M. Thiers & R. E. Halling 2452 (AD, NY); Mt. Lewis, 1000 m, B. M. Thiers & R . E. Halling 2620 (AD, NY); Mt. Windsor Tableland NW of Mossman, M. L. Hicks 11218, 11224, 11226 (BOON, NICH); Bartle Frere, M. L. Hicks 11236 (BOON, NICH); Mt. Spurgeon W of Mossman, M. L. Hicks 11103 (BOON, NICH). Distr.: Endemic to Australia. This species seems to be common in Queensland.

3. Radula queenslandica Yamada, sp. novo (Fig. 1) Planta minuta, olivacea; caulis 4-7 mm longus, irregulariter pinnatim parviramosus; lobi foliorum caulinorum laxe vel moderate imbricati, in piano ovati vel faicati-ovati, apice ± incufvo, sparsim 1- 3 dentato, dentibus acutis vel subacutis, parietibus cellularum medianae tenuibus, trigonis mediocribus; lobuli oblique patuli, subquadrati (vel raro ovati), apice obtuso, basi caulem haud tegente, carina leviter arcuata vel subrecta; dioica (androecia haud visa) ; gynoecia in caulibus terminalia; perianthia longa, plano-cylindrica, ore subtruncato. Plants small, olive-green in herb. Stem 4-7 mm long, ca. 0.05 mm in diam., with leaves 1.0-1.3 mm wide, irregularly pinnately branched, ordinary branches few, 2- 3 mm long, ca. 0.04 mm in diam., with leaves 0.7-0.9 mm wide; stem 4 cells thick, cortical cells as large as medullary cells, both cells thin-walled with medium-sized trigones, subhyaline. Leaf-lobes widely spreading, loosely to moderately imbricate, concave, ovate to falcate-ovate, 0.5- 0.65 mm long, 0.4-0.5 mm wide, apices with 1- 3 teeth, teeth acute to subacute, ± incurved, basal margins slightly arched, not auriculate, covering the stem 1/4-1/3 of the stem-width, insertions substraight; marginal cells 11-15 x 6-9 .urn, thin-walled with medium-sized trigones, median cells 15- 17(- 21) x 12-15 .urn, thin-walled with somewhat large trigones, basal cells 12- 15 x 6- 8 .urn; cuticle smooth; leaf-lobules obliquely spreading, subquadrate (or rarely ovate), ca. 1/2 (or less) the lobe-length, 0.18-0.2 mm long, 0.13-0.18 mm wide, apices obtuse, not elongate, abaxial margins slightly arched or substraight, not decurrent, adaxial margins substraight, bases not covering the stem, insertions substraight, carinal regions widely inflated; rhizoid-initial area convex, rhizoids rarely seen, pale brown; keel spreading at angles of 50° with the stem, 0.2- 0.3 mm long, slightly arched or substraight, not decurrent, sinuses wide. Dioecious (androecium not seen). Gynoecium terminal on stem, with two subfloral in­ novations; bracts in one pair; bract-lobe oblong-ovate with irregularly 4-8-toothed apex, bract-lobule oblong, with irregularly 2- 3-toothed apex and strongly sinuate keel; perianth long, flat-cylindric, ca. 2 mm long, 0.6 mm wide at mouth, mouth subtruncate. Type: Queensland: Mt. Bartle Frere, June 2,1982, I. G. Stone n.s. - holotype (MUCV 6004; dupl. in NICH). K. Y AMADA: The genus Radula from Queensland, Australia 193

FIG. 1. Radula queenslandica Yamada. a. Portion of stem with perianth, ventral view, x 26. b. Portion of stem, dorsal view, x 26. c. Cross-section of stem, x 533. d-f Cells of lobe of stem-leaf, d from margin, e from middle, f from base, aB x 533. g. Apical portion of leaf-lobule, x 533. h, i. Female bracts, x 26. Drawn from holotype.

The diagnostic characters of this species are (I) the stem 4 cells in thickness, (2) the widely spreading, loosely to moderately imbricate, ovate to falcate-ovate leaf-lobes with 1-3 teeth at the apices (teeth subacute to acute), (3) the thin-walled median cells of leaf-lobes with medium-sized trigones, (4) the subquadrate (or rarely ovate) leaf­ lobules with obtuse apices and slightly arched (or substraight), not decurrent keels, (5) the leaf-Iobules not covering the stem at the base, (6) the gynoecium terminal on the stem, with two subfloral innovations and one pair of bracts, and (7) the long, flat­ cylindric perianth with a subtruncate mouth. The present new species is closely related to Radula fissifolia Steph. known only 194 Journ. Hattori Bot. Lab. No. 62 1 987 from New Caledonia. However, the latter is d ifferent from R. queenslandica in (I) the thin-walled leaf-cells with large trigones, (2) the ovate leaf-lobes with usually incurved apices, (3) the female bract-Iobules with entire margins, and (4) the mouth of the perianth with long, sharp-pointed teeth. The present new species is also related to R. dentifolia Grolle known only from New Zealand, but the latter is different from the new species in having the two pairs of female bracts and the female bract-Iobes and bract-Iobules with entire margins except short pointed apices.

Subgenus Radula Key to the species 1. Plants with amentulose branches...... 2 1. Plants without amentulose branches...... 3 2. Leaf-10bules with wholly and strongly inflated carina! regions, keels strongly arched; amentulose branches usually strongly incurved at apices ...... 18. R. thiersiae 2. Leaf-lobules with slightly inflated carinal regions, keels substraight to slightly arched; amentulose branches not incurved at apices ...... 5. R. hicksiae 3. Gemmae present ...... 4 3. Gemmae absent...... 5 4. Plants usually growing on living leaves of trees or on fronds of ferns; gemmae oc­ curring on ventral margins of leaf-lobes ; apices of leaf-Iobules usually strongly turning away from the stem ...... 11 . R. nymanii 4. Pl ants growing on bark of trees; gemmae occurring on margins of leaf-lobes; apices of leaf-Iobu les not turning away from the stem ...... 4. R. acuti/oba 5. Apices of leaf-lobules usually strongly or weakly turning away from the stem ...... 6 5. Apices of leaf-Iobules not turning away from the stem...... 7 6. Stems densely branched; leaf-Iobules widely ovate (rarely subrhombic), apex always strongly turning away from the stem ...... 15a. R. retro/lexa var. /auciloba 6. Stems few-branched ; leaf- Iobules almost subrhombic, apex parallel with, or only slightly turning away from, the stem ...... 15. R. retro/lexa 7. Plants fragile, usually with caducous leaf-lobes...... 8 7. Plants not fragile, without caducous leaf-lobes ...... 9 8. Leaf-lobes falcate-oblong to ovate with obtuse apices and subacute sinuses; leaf­ lobules subquadrate, usually extending slightly beyond the farther edge of stem ...... 16. R. sharpii 8. Leaf-lobes widely ovate with rounded apices and obtuse si nuses; leaf-lobules sub- quadrate to quadrate, covering 1/2-1 /3 of the stem ...... 6. R. javallica 9. Leaf-Iobules not covering the stem ...... 10 9. Leaf-Iobules usually covering the stem ...... 11 10. Plants blackish brown; leaf-lobes ovate to ± obovate without widely rounded apices ; cells of leaf-lobes thin-walled without trigones; leaf-lobules subquadrate, keels sub- straight and spreading at angles of ca. 90° with the stem ...... 13. R. patells 10. Plants yellow-green ; leaf-lobes widely ovate with widely rounded apices; cells of leaf­ lobes thin-walled with large trigones; leaf-lobules ovate, keels usually arched and spreading at angles of ca. 50° with the stem ...... 17. R. tasmanica K. Y AMADA: The genus Radula from Queensland, Australia 195

11. Plants regularly or irregularly dichotomously branched ...... 12 11. Plants regularly or irregularly pinnately branched ...... 13 12. Leaf-lobes loosely imbricate to contiguous, widely ovate with obtuse apices; leaf­ lobules subquadrate, covering 1/4-1/5 of the stem, adaxial margins usually depressed at middle, keels substraight to slightly arched ...... 7. R. jovefiana 12. Leaf-lobes moderately imbricate, narrowly ovate with rounded apices; leaf-Iobules quadrate, covering 1/2-2/3 of the stem, adaxial margins not depressed at middle, keels usually substraight ...... 9. R. loriana 13. Leaf-Iobules with narrowly recurved apices and abaxial margins usually decurrent to the ventral margins of leaf-lobes ...... 14. R. reflexa 13. Leaf-Iobules without recurved apices and with abaxial margins not decurrent to the ventral margins of leaf-lobes ...... 14 14. Leaf-Iobules usually with narrowly recurved bases; keels sinuate, slightly decurrent ...... 10. R. multiflora 14. Leaf-Iobules without recurved bases; keels substraight, not decurrent...... 15 15. Leaf-Iobules covering 3/4 (or more) of the stem; keel spreading at angles of ca. 40° with the stem ...... ••...... 8. R . kurzii 15. Leaf-Iobules covering 1/6- 1/5 of the stem; keel spreading at angles of ca. 60° with the stem ...... 12. R. parvitexta

4. Radula aculi/oha Steph. Hedwigia 28: 271 (1889). - Radula papulosa Steph., J. Linn. Soc. Bot. 29: 272 (1892), syn. novo Specim. exam.: Hurdle Gully, 14 km WSW of Moto, 300 m, H. Streimann 9873, 9880, 9883 (CBG, NICH). Distr. : Australia, New Guinea. Radula aculiloha was described by Stephani (1889) based on a specimen from Queensland (without definite locality). This species is closely related to R. constricta Steph. known from Asia. Recently I have examined the holotype specimen of R. papulosa Steph. from New Zealand preserved in G (22376), and confirmed that it is identical with R. aculi/oha Steph.

5. Radula hicksiae Yamada Cryptogamie Bryol. Lichenol. 5 : 191 (1984). Specim. exam.: Mt. Bartle Frere, on bark, Aug. 1982, M . L. Hicks 11047-holotype ( NICH; dupl. in hb M. L. Hicks); Mt. Hosie, Kirrma State Forest, I. G. Stone (MUCV 5882); Mt. Bellenden Ker south peak, on Alphitonia stem, H. Streimanl1 27486 (CBG, JE); Barron State Forest, Herberton Range, on trunks, H . Streimann 27304 (CBG). Distr. : Endemic to Queensland. This species is closely related to R. recurvifolia Yamada known only from New Caledonia.

6. Radula ja van ica Gott. In Gott. et aI., Syn. Hep.: 257 (1845). Specim. exam.: El Arish, 1. G. Stone (MUCV 6107); Dalymple track, I. G. Stone (MUCV 5927); Mt. Mackey State near Tully, I. G. Stone (MUCV 5890, 5892); Cedar Bay, I. G. Stone (MUCV 5845) ; "The Boulder", 7 km W of Babinda, 125 km, B. M . Thiers & R. E. Hailing 2400, 2424 (AD, NY); ca. 5 km W 196 Journ. Hattori Bot. Lab. No. 62 198 7

of Mossman, along Mossman River, 150 m, B. M. Thiers & R. E. Hailing 2562, 2576, 2589 (AD, NY); track to Tchupala Falls, 750 m, B. M. Thiers & R. E. Hailing 2362 (AD, NY); MiIlaa Millaa Falls National Park, 900 m, B. M. Thiers & R. E. Hailing 2239 (AD. NY); Mossman Gorge W of Mossman, M. L. Hicks 11182 (BOON, NICH); Rainforest W of Babinda, M. L. Hicks 10943 (BOON, NICH). Distr.: Widely distributed in the tropical and subtropical regions of SE Asia and the Pacific islands. The present species seems to be one of the commonest species in Queensland, as listed above. It is highly variable especially in forms of leaf-lobes and leaf-lobules.

7. Radulajovetiana Yamada Cryptogamie Bryol. Lichenol. 5 : 193 (1984). Specim. exam.: Devil's Thumb W of Mossman, 1150 m, M. L. Hicks 11263-holotype (NICH; dupl. in BOON), 11264 (BOON, NICH); Mt. Bartle Frere, M. L. Hicks 11013,11045 (BOON, NICH); 33 km from Creek, Builyan, I. G. Stone (Mucv 5962); Mt. Haig, 1200 m, M. M. J. v. Balgooy 1584 (hb Lugd, CBG). Distr.: Endemic to Queensland.

8. Radula kurzii Steph. Hedwigia 23: 153 (1884). Specim.: Mt. Bartle Frere, M. L. Hicks 11020, 11025, 11229 (BOON, NICH); Mt. Lewis SW of Mossman, M. L. Hicks 11141 (BOON, NICH); , M. L. Hicks 10901, 10958 (BOON, NICH). Distr.: Ceylon, India, Andamann Is., Australia. Yamada (1984) first reported this species from Queensland.

9. Radula loriana Castle J. Hattori Bot. Lab. 21 : 6 (1959). Specim. exam.: Ca. 5 km W of Mossman, along Mossman River, 150 m, B. M. Thiers & R. E. Hailing 2565, 2596 (AD, NY); "The Boulder", 7 km W of Bobinda, 125 m, B. M . Thiers & R. E. Hailing 2419 (AD, NY); Mt. Mackay State Forest near Tully, I. G. Stone (MUCV 5895). Distr. : New Guinea, Australia. This species was previously known only from New Guinea.

10. Radula multiflora Gott. ex Schiffn. Forschungsr. "Gazelle" Bot. 4(4): 20 [1889] (1890). Specim. exam. : Mossman Gorge, M . L. Hicks 10860 (BOON, NICH), I. G. Stone 4653 (MUCV); Mt. Lewis SW of Mossman, M. L. Hicks 11173 (BOON, NICH). Distr.: Widely distributed in tropical Asia and the Pacific islands (for distribution see Yamada 1979). Yamada (1984) first reported this species from Queensland.

11 . Radula nymanii Steph. Spec. Hep. 4 : 229 (1910). Specim. exam.: Palmerston, I. G. Stone (MUCV 5851); track to Tchupala Falls, 750 m, B. M. Thiers & R. E. Hailing 2377 (AD, NY). Distr.: Widely distributed in tropical Asia and the Pacific islands; Australia. This species has not been reported previously from Australia. K. Y AMADA: The genus Radula from Queensland, Australia 197

12. Radula parvitexta Steph. Spec. Rep. 6: 513 (1924). Specim. exam.: Rerberton Range, 13 km S of Atherton, 1000 m, H. Sfreimann 27215 (CBG); Mt. Bellenden Ker, 23 km SEE of Gordonvale, 1550 m, H. Sfreimann 27331 (CBG); summit of central peak of Mt. Bellenden Ker, 1500 m, B. M. Thiers & R . E. Hailing 2459, 2473,2493,2495,2499,2521 (AD, NY); ca. 5 km W of Mossman along Mossman River, 150 m, B. M. Thiers & R. E. Hailing 2571 /a (AD, NY). Distr.: New Guinea, Guadalcanal 1. , Australia. This species seems to be rather common in Queensland, though it has not been reported previously from Australia.

13. Radula patens Yamada Cryptogamie Bryol. Lichenol. 5(1-2): 197 (1984). Specim. exam.: Salter creek W of of Mossman, Qct. 1982. M . L. Hicks 11328 - holotype (NICH ; dupl. in BOON); Mt. Lewis near Mossman, M. L. Hicks 11396, 11434, 11439, 11442 (BOON); Cape Tribulation N of Mossman, M. L. Hicks 11507 (BOON, NICH). Distr.: Endemic to Queensland. This species seems to be common on Mt. Lewis near Mossman.

14. Radula reftexa Nees et Mont. In Mont., Ann. Sci. Nat. Bot. ser. 2, 19 : 255 (1843). Specim. exam. : Cape Tribulation N of Mossman, M. L. Hicks 11506 (BOON, NICH). Distr.: Cera m I., Borneo, New Guinea, Australia. This species has not been reported from Australia previously.

15. Radula retroftexa Tayl. London J. Bot. 5: 378 (1846). Specim. exam.: Mt. Windsor Tableland NW of Mossman, M. L. Hicks 11191, 11200 (BOON, NICH); Nat. Park, 900 m, B. M. Thiers & R. E. Hailing 2221 , 2248 (AD, NY). Distr.: Widely distributed in tropical Asia and the Pacific islands. Yamada (1984) made the first report of this species from Queensland.

15a. Radula retroftexa Tayl. var.fauciloha (Steph.) Yamada J. Hattori Bot. Lab. 45 : 282 (1979). Specim. exam.: Mt. Coolum, J. Windolf 1142 (NICH; hb Windolf). Distr.: Widely distributed in tropical Asia and the Pacific is lands. Windolf (1985) reported this variety from Sunshine coast in Queensland.

16. Radula sharpii Yamada J. Jap. Bot. 60: 360 (1985). Specim. exam. : Mossman Gorge, T. G. Srone (MUCV 6101). Distr.: New Guinea, Australia. This species has not been reported previously from Australia.

17. Radula tasmanica Steph. Spec. Rep. 4: 212 (1910). Specim. exam.: Barron State Forest, Rerberton Range, 1000 m, H. Sfreimann 27215 (CBG, JE, NICH, H); Mt. Bethongabel, 900 m, B. M. Thiers 1220 (NY); Ballenden Ker Nat. Park, 125 m, B. M. 198 Journ. Hattori Bot. Lab. No. 62 1 9 8 7

Thiers & R. E. Hailing 2394, 2404, 2795, 2801 (AD, NY). Distr.: Endemic to Australia (including Tasmania). This species has not been reported from Australia previously, but does not seem to be rare in Queensland.

18. Radula thiersiae Yamada, sp. novo (Fig. 2) Planta mediocris, flavobrunnea ; caulis irregulariter pinnatim ramosus, ramis amentulosis (cum foliis minutis), valde hamatis ; lobi foliorum caulinoruro moderate imbricati, concavi, in piano late ovati, basi parum auriculata; lobuli remoti, omnino inflati, ovati (in piano subquadrati), area carinali valde inflata, margine abaxiali subrecto, decurrente, basi caulem ca. 1/5 (vel minus) tegente, carina va lde arcuata; dioica? (gynoecia haud visa); androecia in caulibus lateralia. Plants medium-sized, creeping on bark, yellow-brown in herb. Stem 25- 35 mm long, ca. 0.15 mm in diam., with leaves 2.4-2.6 mm wide, irregularly pinnately branched, branches 3- 7 mm long, ca. 0.1 mm in diam., with leaves 1.6-2.0 mm wide; amentulose branches numerous, 1.8- 2.5 mm long, with 5- 9 pairs of small, hook-shaped leaves usually with strongly incurved

FIG. 2. Radula thiersiae Yamada. a. Portion of stem with amentulose branches, ventral view, x 16. b. Portion of cross-section of stem, x 480. c, d. Stem-leaves, x 16. e-g. Cells of lobe of stem-leaf, e from margin, f from middle, g from base, all x 480. h. Androecium and leaf-Iobules on stem, x 23. Drawn from holotype. K. Y AMADA: The genus Radula from Queensland, Australia 199 apices; stem 11 - 12 cells thick, cortical cells as large as medullary cells, both cells thick-walled with large trigones, pale yellow. Leaf-lobes moderately imbricate, widely spreading, concave, widely ovate, 1.1-1.4 mm long, 1.5- 1.6 mm wide, apices rounded, not incurved, basal margins arched, slightly auriculate at bases, dorsally extending more than stem-width beyond the farther edge of stem, insertions inverted J-shaped; marginal cells 7-9 x 11 -13 .urn; median cells (16-)20--23 x (12-)16-19 .urn, thin-walled with medium-sized trigones (intermediate thick­ enings not seen), basal cells 23-26 x 10--13(-16) .urn; cuticle smooth; leaf-Iobules remote, wide­ ly spreading, usually wholly inflated, ovate (when flat subquadrate), ca. 2/3 the lobe-length, 0.85-0.9 mm long, 0.5--0.65 mm wide, apex (when flat) bluntly angular, abaxial margins sub­ straight, usually decurrent and ± incurved along the ventral margins of leaf-lobes, adaxial margins ± narrowly recurved along margins, substrajght (rarely slightly arched) toward bases, the bases arched, covering only ca. 1/5 (or less) of the stem-width, insertions substraight, carinal regions usually strongly inflated; rhizoid-initial areas convex, rhizoids red-brown, in a bundle; keels spreading at angles of 50- 70° with the stem, 0.85- 0.92 mm long, strongly arched, not decurrent, sinuses very wide. Dioecious? (gynoecium not seen). Androecium lateral on stem, shortly spicate with 3-4 pairs of bracts. Type : Queensland : Cool district, Bellenden Kef National Park, summit of central peak of Mt. Bellenden Ker, vicinity of Telecom Transmitting Station, 1500 m alt., July 10, 1984. B. M. Thiers & R. E. Hailing 2500 - ho!otype (NY; dupl. in AD, NICH). The present species is characterized by (I) the medium-sized (25-35 mm long) plants, (2) the stem 11-12 cells thick, stem-cells thick-walled (with large trigones), (3) the numerous, amentulose branches usually with small, hook-shaped leaves, (4) the moderately imbricate, widely spreading, widely ovate leaf-lobes with rounded apices and arched basal margins, (5) the remote, wholly inflated, ovate (when flat subquadrate) leaf-lobules with strongly inflated carinal regions and strongly arched keels, (6) the bases of the leaf-lobules covering only ca. 1/5 (or less) of the stem, and (7) the abaxial margins of leaf-lobules usually decurrent to the ventral margins of leaf-lobes. This species is very closely related to R . scariosa Mitt., known from Fiji and New Caledonia, but the latter is different from the present species in (1) the amentulose branches with 3-4 pairs of leaves not hook-shaped and not so much reduced, (2) the narrowly ovate leaf-Iobules which do not cover the stem, and have 3- 4 hyaline marginal cell-rows along the adaxial margins, and (3) the stem which is only 8-9 cells thick. The amentulose branches of the present species are unique, and no close relative is known in the section Amentulosae.

LITERATURE CITED Castle, H. 1963. A revision of the genus Radula. Part. H. Subgenus Acroradula. Section 6. Saccarae. Rev. Bryol. Lichenol. 32 : ]-48. --.1964. --. Section 8. Acutilobulae. Rev. BfYOI. Lichenol. 33: 185-210. --.1967. --. Section 11. Complanatae. Rev. Bryol. Lichenol. 34 : 1- 94. Hampe, F. 1838. In J. G. G. Lehmann, Novarument minus congnitarium stirpium. Pugillus 7 : 1-41. Stephani, F. 1884. Die Gattung Radula. Hedwigia 23 : 145- 159. - - . 1889. Hepaticae Australiae. Hedwigia 28: 257-278. 200 Journ. Hattori Bot. Lab. No. 62 1 987

--. 1910. Radula. In Species Hepaticarum 4: 151-234. Windolf, J. 1985. Survey of the Hepaticae and Anthocerotae of the Sunshine Coast regions, Que- ensland. J. Hattori Bot. Lab. 59: 171 - 176. Yamada, K. 1979. A revision of Asian taxa of RadII/a, Hepaticae. J. Hattori Bot. Lab. 45: 201 - 322. --. 1982. Some new species of Radula (Hepaticae). J. Hattori Bot. Lab. 51 : 323-328. --. 1984. A Radllla collection made by Dr. M. L. Hicks in Queensland, Australia. Crypt. Bryol. Lichenol. 5(1- 2): 191 - 199.