Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenberg

Sukkharak, Phiangphak, 2015, A systematic monograph of the genus Thysananthus (Lejeuneaceae, Marchantiophyta), Phytotaxa 193 (1), pp. 448-450 : 448-450

9. Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenberg in Gottsche et al. (1845: 287). Junermannia spathulistipa Reinwardt, Blume & Nees (1824: 212) . Jungermannia spathulistipa α, β Nees (1830: 38). Lejeunea spathulistipa (Reinwardt, Blume & Nees) Dumortier (1835: 12) . Frullania spathulistipa (Reinw., Blume & Nees) Nees (1838: 211) . Type: INDONESIA. Java [“Habitat in Javae montosis, v.c. in monte Leback Provinciae Bantam, freqaens”]: without date, Blume s.n. (holotype STR!; isotypes G 3 packets!, W!).

Thysananthus triquetrus (Mitt.) Stephani (1912a: 783) . Lejeunea triquetra Mitten (1864: 167) . Type: “Bagro river, W. Africa, 1861, Mann s.n. ” (holotype NY!; isotype BM!), syn. nov.

Thysananthus monoicus Stephani (1912a: 783) . Type : MADAGASCAR. without location, without date, Darbould s.n. (holotype: G!); Icon. Steph. nr. 10228.

Thysananthus sikkimensis Stephani (1912a: 798) . Type: “Himalayas”, without date, Levier s.n. (holotype G!); Icon. Steph. nr. 10212.

Thysananthus ( Lejeuneaceae , Marchantiophyta) Phytotaxa 193 (1) © 2015 Magnolia Press • 33

Thysanolejeunea amboinensis Schiffner ex Stephani (1912a: 799) , nom. inval.

Thysananthus hebridensis Stephani (1924: 565) . Lectotype (here designated): VANUATU. [“Novae Hebrides”], October 1911, Gunn s.n. (lectotype G!; isolectotypes G 2 packets!); Icon. Steph. nr. 10184.

Mastigolejeunea tarkwana Pearson (1931: 63) . (fide Wigginton & Grolle, 1996).

Thysananthus spathulistipus (Reinwardt, Blume & Nees) Lindenb. var. borneensis Herzog (1931: 199) . Type: MALAYSIA. Sarawak [West Borneo]: Bukit Mulu , 1000 m, 2 December 1924, H. Winkler 3333 (holotype JE!), syn. nov.

Thysananthus minor Verdoorn (1933b: 231) . Type: INDONESIA. Sumatra: Mt. Brastagi , “Gouv. S.O.K., in silvis pr. cataractas torrentis Petani, infra Brastagi, ca 1350 m ”, April 1930, Verdoorn s.n. (holotype FH!), syn. nov.

Thysananthus fuscobrunneus Horikawa (1934: 252) . Type : TAIWAN. Taito : Mt. Chipon, Kiriyama-Miyama, on bark, 31 December 1932, Y. Horikawa 10505a (holotype HIRO!), syn. nov.

Thysananthus grossidens Stephani ex Verdoorn (1934a: 168) , nom. inval.

Plants autoicous, with projecting growth, turning upwards and becoming ascending to erect, green, brownish green to black in the older portion in the field, yellowish brown in herbarium specimens, up to 5 cm long × 1.7–3.0 mm wide. Stems rather rigid; ventral merophyte 7–9 cell rows wide; stem in cross section orbicular, 205–219 mm high × 163–175 µm wide, 14–18 cell layers high, composed of 32–46 epidermal cells surrounding 50–95 medullary cells, epidermal cells as large as medullary cells. Leaves imbricate, when dry suberect and convolute, when moist weakly convex, apical part plane, not recurved; dorsal lobe symmetrically ovate, 0.8–1.6 × 0.5–1.1 mm, apex acute, dorsal base cordate, dorsal margin entire or with 1–4 triangular teeth, the teeth consisting of 3–6 cells, being 2–3 cells wide at base and ending in a row of 1–2 cells, ventral margin recurved to inrolled 1/2 × leaf length, with 3–6 triangular teeth, the teeth consisting of 4–8 cells, being 2–3 cells wide at base, apex of one cell; cells elongate-hexagonal with acute ends, vitta absent, marginal cells 5–10 × 7–12 µm, median cells 17–27 × 5–12 µm, basal cells 25–62 × 15–25 µm, trigones cordate, often coalesced, intermediate thickenings 0–3 per cell; oil bodies 2–4 per cell ( Mizutani 1969: 4–6 oil bodies). Lobules oblong-rectangular, 0.3–0.5 × 0.1–0.2 mm, 1/4–1/3 × lobe length; appendage on surface of lobule base not developed; keel without appendage; lobule apex oblique, free margin continuing into the ventral lobe margin, apex with 1–2 triangular teeth, the first tooth consisting of 3–4 cells, being 2 cells wide at base and ending in a row of 1–2 cells, the second tooth of one cell only, often absent. Underleaves imbricate, slightly squarrose, broadly obovate-spathulate, 0.3–0.8 × 0.4–0.7 mm, 2–4 × stem width, apex truncate to slightly emarginate, plane, margin with 8–13 triangular teeth, the teeth consisting of 3–6 cells, being 2–3 cells wide at base and ending in a row of (1–)3–7 cells, bases cuneate, underleaf bases adnate with leaves on one side, on left-hand side for right branches and right-hand side for left branches; cells 7–17 × 5–12 µm. Androecia terminal-intercalary on lateral branches, bracts, and bracteoles in 2–8 pairs, bracts hypostatic, 0.6–0.8 × 0.2–0.6 mm, apex acute, margins with 1–2 teeth; antheridia 2 per bract. Gynoecia with 1–2 lejeuneoid innovation forming monochasial or dichasial pattern; lobe ovate, 1.3–1.9 × 0.6–1.1 mm, apex apiculate, margins in upper 1/3 with triangular teeth, the teeth consisting of 3–6 cells, being 2–3 cells wide at base and ending in a row of 1–2 cells; lobules broadly ovate, 1/3–1/2 × lobe length, apex apiculate, margins with triangular teeth, the teeth consisting of 5–11 cells, being 2–3 cells wide at base and ending in a row of 1–2 cells; bracteoles spathulate, 1.2–2.3 × 0.5–1 mm, apex emarginate to shortly bifid, 1/2 × bracteole length with triangular teeth, the teeth consisting of 3–8 cells, being 2–3 cells wide at base and ending in a row of 1–3 cells, margins recurved. Perianths oblong-cylindrical, 1.6–2.4 × 0.6–1.1 mm, keels in upper 1/3–1/2 with triangular teeth, the teeth consisting of 4–15 cells, being 2–4 cells wide at base and ending in a row of 1–2 cells; beak 50–55 µm (3–4 cells) in length. Figs.19 , 20 .

Additional illustrations:— Horikawa (1934, p. 251, Fig. 49 as Thysananthus fuscobrunneus ); Vanden Berghen (1950, p. 36, Fig. 1 ); Thiers & Gradstein (1989, p. 74, Fig. 28 ); Gradstein et al. (2002, p. 75, Fig. 47 as T. minor , p. 81, Fig. 51).

Distribution and ecology:— Africa, Indochina, Malesia, Australia (Queensland), and the Pacific region; from sea level to 2800 m; on tree trunks, roots, rotten branches and logs along road sides, in the swamp forest, mangrove forest, lowlnd rain forests (primary and secondary forests), and montane forests. Fig. 5B .

Representative specimens:—Ivory Coast. TAÏ: Taï National Park, Assi 12739/B (U). Ghana. WESTERN REGION: Ankasa river forest reservation, Jones 1578b (G), ibid., 14 February 1971, Jones s.n. (JE). Nigeria. BENIN: Okomu forest reserve, Jones 44, 174B (BM). Cameroon. SOUTHWEST REGION: Bibundi, September 1891, Dusén s.n. (S 4 packets). Gabon. without location, 24 June 1927, Testu s.n. (PC, S). Zaire. KIVU: Irangi forest station (Bryotrop Expedition loc. 120), Pócs 6601 (G, U), 6735 (G). Seychelles. MAHÉ ISLAND: Beau Vallon, forest above Le Niol, Norkett 16600 (BM).— PORT GLAUD: Morne Seychellois National Park, Onraedt 74.S.258 (JE), Norkett 16429, 16512, 16539, 16687 (JE), Pócs 00105, 00105/B, 9335/B (G); Congo Rouge Relict forest, Norkett

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17148 (BM, JE); La Misère road, Norkett 16512 ( BM) , 12 September 1973, Norkett s.n. ( BM) ; summit of Morne Blanc, Norkett 18603 ( BM) ; Sans Souci road, Norkett 16687 ( BM) ; old Capucin Mission , Trois Frêres road, Norkett 16429 ( BM, JE) ; Val Riche, Pócs 9333/L (F, G 4 packets, PC) ; NE escarpment of Trois Frêres, Pócs 9315/ W ( EGR, F, GOET); Morne Seychellois National Park , Copolia summit, Pócs 9335/B (G 2 packets), ibid., in a depression of Mare aux Cochons valley , 16 June 2000, Pócs s.n. (G 2 packets), ibid., submit of Montagne Palmiste, Pócs 9338/C ( EGR) ; Ridge of Brulée , Norkett 17933 ( BM) .— SILHOUETTE ISLAND: near Pota Eau ridge, Norkett 17741 ( BM) , without location, Alexejev 13 ( JE) . Madagascar. ALAOTRA-MANGORO: forest in Analamazoatra, Camboué 316, 334 ( PC) .— ANTSIRANANA: Diego-Suarez , Radama, Aptroot & Hensen 13198B ( U) ; Tamatave, Andasibe , Aptroot & Hensen 13433 ( U) .— ATSIMO-ANTSIRANANA: Renana stream, Humbert 3451 ( PC) .— HAUTE MATSIATRA: Fianarantsoa, Tixier 9009 (L). Réunion. SAINT-BENOÎT: between Grand Etang and road, “M” 17.855 ( BR 2 packages), Onraedt 73.R.1387 ( BR) .— SAINT-PAUL: Peak of the Furnace , Pócs 9611/CG (F). Mauritius. PLAINES WILHEMS: Le Patrin, “Guého” 48 ( PC) ; without location, without date, Bélanger s.n. ( BM) .

India. ANDAMAN AND NICOBAR ISLANDS: Andaman Islands, Port Blair, 14 November 1890, Man s.n., ex hb. Levier 773 (G), ibid., 1893, Man s.n., ex hb. Levier 741 (G), ibid., ex hb. Levier 774 (G 2 packets, JE, S 2 packets), ibid., October 1898, Man s.n., ex hb. Levier 1715 ( BM, G 2 packets, PC); Port Monate , without date, Kurz s.n. ( BM 3 packets, PC); without location, without date, Kurz 1622, 1631 (G). Sri Lanka. CENTRAL: Adam’s peak, Onraedt 87.L.8801 ( BR) .— SOUTHERN: Beraliya forest reserve, Onraedt 76.L.3284 ( BR, JE) ; Hiniduma, Onraedt 76.L.3508 ( JE) , 77.L.4304, 81.L.9291, 81.L.9292, 81.L9370 (BR), 76.L.3454, 76.L.3508 (BR, JE), 76.L.3684 (PC); Kammeliya forest reserve, Onraedt 77.L.4257 ( BR) . China. HAINAN: Kwangtung, Chen & al. 382, 435c, 633 (L) .

Thailand. CHANTHABURI: Khao Sa-Bab , 24 December 1972, Sapanuchart s.n. ( BCU) .— CHIANG MAI: Mt. Inthanon , Touw 9613 ( BKF) .— LOEI: Mt. Phu Luang , Thaithong 201 ( BCU) , Touw 10530 ( BKF) , ibid., north ridge, Tagawa & Kitagawa T 1366, T1377 (G), ibid., eastern slope, Tagawa & Kitagawa T 1876 (G), ibid., north-eastern ridge, Tagawa & Kitagawa T 1686, T1873 (G); Phu Kradung, Touw 10959 ( BKF) , Tagawa & Kitagawa T 1019 (G), ibid., Pen-Pop waterfall, Thaithong 972 ( BCU) .— N AKHON R ATCHASIMA: Khao Khiao, Tixier 319 ( PC) .— NAKHON SI THAMMARAT: Mt. Khao Luang , Smitinand 1027 ( BCU, BKF) , Touw 10667, 11354, 11360, 1176 8, 11875, 11883, 12015, 12023 ( BKF) , 11686, 11738 (BKF, BR), Tagawa & Kitagawa T 5067, T5072, T5154 (G); Mt. Khao Nan Yai , Sukkharak 50, 130, 170, 180, 183, 189 ( BCU) , ibid., San Yen, Sukkharak 739, 746, 751, 752, 762, 766, 768, 784, 788, 793 ( BKF, GOET) .— PHANG NGA: on the way to Roe-Si-Sawan cave, Boonkerd 14 ( BCU) ; Sri Phang Nga National Park , Chantanaorrapint 2089, 2096, 2099, 2100 ( PSU) . — PRACHINBURI: Khao Yai National Park , Touw 12104, 12360 ( BKF) , 12103 (BKF, BR).— RANONG: La-Oon mangrove forest, Thaithong 981 ( BCU) ; hot spring, Tixier 3895 ( PC) . — SATUN: Taritao Island , Chantanaorrapint 2072 ( PSU) . — TRAT: Chang Klau , Hui Range, Smitinand 1453 ( BCU, BKF) ; Koh Chang, 10 February 1900, Schmidt s.n. (C). — TRANG: Mt. Khao Chong, Tagawa & Kitagawa T 6877, T6965, T6970, T6974 (G). Cambodia. Tonle Sap, Tixier 3864 ( PC) ; Kampot, Mt. Bokor, Tixier 2669, 3006, 3027, 3196, 3225, 3313, 3603, 3710 ( PC) ; Tonle Sap , Tixier 3835 ( PC) . Vietnam. LÂM ĐỒNG: Da Lat, 21 September 1957, Tixier s.n. ( PC) . Malaysia. JOHORE: Mt. Ophir , Verdoorn 58, 66, 100, 135, 170 ( SING) .— KEDAH: Gunung Raya , Haniff & Nur 7120 ( SING) ; Gunung Raya Forest Reserve, Mohamed & Kien-Thai 5089 ( KLU) ; Kedah peak, Tixier 6058 ( PC) , Holttum 14889 ( SING) ; Ulu Muda Forest Reserve , Kien-Thai 3326 ( KLU) .— KELANTAN: Gunung Stong Forest Reserve , Kien-Thai 3935, 4013 ( KLU) .— MALACCA: Gg. Tunduk, Ridley 701 ( SING) .— NEGRI SEMBILAN: Bukit Putus, Ridley 751 ( SING) .— PAHANG: Cameron Highlands , Wee 942 ( NICH) ; trail to Gunung Jasar, Gradstein 10400/1, 10400/2, 10404/1, 10404/2 ( GOET) ; 2.5 miles on Genting Highland road, 21 April 1972, Een s.n. ( JE, S); Fraser’s Hill, Tixier 4753 ( PC) , ibid., 21 May 1997, Schäfer-Verwimp & Verwimp s.n. (G); the road to Genting Hightland, Sukkharak 716, 719, 722 ( BKF, GOET) .— PENANG: Penang hill, Allen 754 (L), Ridley 525 ( SING; between Penang hill and Tiger hill, Kitagawa 14565 ( KLU) .— PERAK: Taiping, Ilkiu-Borges & al. 2981 ( GOET) , Yamaguchi 25770 ( HIRO) ; Gunong Batu Putih, Wray 1180 ( BM, G, SING, W); Buyong Malacca, Ridley 723 ( SING) ; Taiping, Quah 24 ( KLU) .— SABAH: Mt. Kinabalu, Kamborangah , November 1931, Holttum s.n. ( SING) ; Silau-Silau trail, Sukkharak 562 ( BCU) , Sukkharak 549, 550, 593 ( BKF, GOET) ; Kiaw View trail, Shea 3006 ( KLU); 5°31'40'' N 116°32'20'' E, 4 miles up Mt. Trus Madi from Kpg, Shea 1461 ( KLU, NICH) .— SARAWAK: Mt. Dulit, August 1932, Richards s.n., Hep. Sel. Crit. Verdoorn 400 ( BR, C, G, GOET, PC, SING, U, W), ibid., 1932, Richards 2276, Hep. Sel. Crit. Verdoorn 450 ( BR, C, G, GOET, JE, PC, S, SING, U, W), ibid., October 1932, Richards s.n., Hep. Sel. Crit. Verdoorn 397 (C, GOET) , ibid., November 1932, Richards s.n., Hep. Sel. Crit. Verdoorn 398 (C), Richards 1208, 2339 ( SING) , Synge 2083 ( BM) , ibid., Dulit trail, Richards 1182 ( JE) ; Sibu, Loba Kabang protected forest, Wood 1420 ( BM, L) ;

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Banting , Beccari 1867 ( U) ; Bukit Setiam , Mohamed & Bakar 3383, 3436 ( KLU) . Singapore. Chua Chu Kang , Ridley 441 ( SING 2 packets); Gardens, Ridley 600 ( SING) . Brunei. TEMBURONG: 1.5 km NE of Kampong Semabat, Seah 534 ( SING) . Indonesia. JAVA: East Java , Tretes, on N-slope of Mt. Ardjuno, Sipman 6992 ( U) ; Dorowatti , May 1929, Clason s.n. (L); West Java , Mt. Salak , Fleischer 76 (S); Mt. Gede , Artja , April 1894, Schiffner s.n., Hep. Sel. Crit. Verdoorn 289 ( BM, BR, C, G 2 packets, GOET, JE, PC, S, SING, W), ibid., Perbawati , July 1930, Verdoorn s.n., Hep. Sel. Crit. Verdoorn 290 ( BM 2 packets, BR, C, G 2 packets, JE, PC, S, SING, U 2 packets, W); Cibodas, April 1894, Schiffner s.n., Hep. Sel. Crit. Verdoorn 291 ( BM 2 packets, BR, C, G 2 packets, GOET, JE, PC, S, SING, U, W), ibid., February 1931, Renner s.n. ( JE 2 packets), 12 February 1913, Fleischer s.n. (C); Tjibodas, 12 February 1913, Fleischer s.n. (F); Tjibureum, November 1927, Burgeff 8103 ( JE, S), “Noeste” 308 ( JE) ; Mt. Pangerango , without date, Kurz s.n. ( BM 2 packets).— KALIMANTAN: East Kalimantan, peak of Balikpapan, Meijer B 1332c, B1416 (S), Yamaguchi 28690 ( HIRO) ; Nunukan , Meijer B 4921 ( BM) ; Kutai , Meijer B 1454 (L).— LESSER SUNDA ISLANDS: Bali, Bali Handara country club, Schäfer-Verwimp & Verwimp 20790 ( GOET) .— SULAWESI: Central Sulawesi , Lore Lindu National Park , Mt. Nokilalaki, Gradstein & Ariyanti 11000, 11041, 11052 ( GOET) , Central Sulawesi , November 1888, Warburg s.n. ( JE) .— SUMATRA: Bangka- Belitung , Balai , 1869–1870, Teysmann s.n., Hep. Sel. Crit. Verdoorn 399 ( BR, C, G, GOET, JE, PC, S, SING, U, W); Mt. Menoembing, Teysmann 30 ( GOET) , 30/A, 30/B (L); North Sumatra , Bandar Baroe , Staal S –249 (L); West Sumatra, Singalang, July 1894, Schiffner s.n., Hep. Sel. Crit. Verdoorn 288 ( BR, C, PC, S, SING, U); Lubuk Sikaping, 1931, Renner s.n. ( JE, S); Padang, Schild 107 ( JE, W); Singalang, Schiffner 2924, 4009 ( U) ; Agam , 29 August 1922, Bleeker s.n. (L).— WESTERN NEW GUINEA: Papua, Cycloop Mts., van Royen & Sleumer 6256 ( BR, JE, U) ; Kadubaka , Swart valley, Bergman H. 50, H.51 (S); West Papua, Vogelkop Peninsula, Royen & Sleumer 7966 ( BR, JE, U) ; Bian Katem , van Zanten 229 ( U) ; Tanah Merah , van Zanten 182 e ( U) . Philippines. AGUSAN DEL NORTE: Butuan, Weber 1343 ( BM) ; Mt. Urdaneta , October 1912, collector unknown ( BM, G, PC, U, W).— BENGUET: Baguio, February 1928, Burgeff s.n. (S).— ORIENTAL MINDORO: Mt. Malasimbo, Bartlett 13865 ( JE) ; Mt. Banahao , Alvarez 0–77512 ( JE) .— QUEZON: Botanical Garden of University of the Philippines Los Banõs, Onraedt 85.P.11342 ( BR) .— SAMAR: Silanga , Merrill 9005 ( BM) .— ZAMBALES: Malaat creek , Gruezo 5396 ( NICH) . Papua New Guinea. CENTRAL: Varirata National Park, Sloover 42.564 ( BR) ; Boridi , Carr 13537 ( JE) , 13528 ( BM, L 2 packets); near Dabamura on Owers Corner road, Streimann & Naoni 14879, 14950 ( JE, LAE) ; Kuriva logging area, Streimann & Vinas 14421 ( LAE) , 14448 (JE, LAE); Bereina-Angabanga river road, Streimann & Naoni 16303 ( JE, LAE) ; K. B. sawmill, Ehu creek , Streimann & Naoni 16499, 16633 ( LAE) , 16668 (JE, LAE); road to Musgrave river , Streimann & Naoni 15239, 15953 ( JE, LAE) .— EAST SEPIK: Ambunti subdistrict, along Yapa, Hoogland & Craven 10121, 10738 ( JE) .— MOROBE: Wau , Mt. Kaindi , Kunai creek, Gradstein 3964 (G 2 packets, U 3 pakcets), 3965 ( U) ; Kaisenik logging area, Shea 6404 ( JE) , 6416 (NICH); Mt. Missim track, Streimann 22911 ( GOET) , 22919 (JE, LAE); Molukkerne , 27 November 1898, Nyman s.n. (S); Yinimba, Streimann 19032, 19095 ( JE, W); Windowi village, Streimann & Kumei 22648 ( JE, LAE) ; logging area, 7°10' S 146°45' E, Streimann & Bellamy 13311 ( JE, LAE) ; Slate-Gumi creeks divide, Streimann 13831 ( JE, LAE) , 13973 (LAE); Gumi divide, Streimann 22743 ( LAE) , 22733 (JE, LAE); Aiuwa-Bakia track, Streimann & Tamba 12274 ( LAE) , 12399 (JE, LAE); Herzog Mts. , Streimann & Umba 10838, 10854, 10869, 10916, 10964, 11012 ( JE, LAE) ; Araulu logging area, Streimann 13565 ( JE, LAE) , 13553, 13571 (LAE); upper Watut river, Streimann 23100 ( BR, JE, LAE) ; Kaisenik logging area, Shea 6404, 9904, 9005 ( NICH) ; Wau-Salamaua track, Streimann 25392 ( JE) ; upper Nawata Banda , Streimann 19519, 24868, 24872, 33962 ( JE) ; Bulolo-Watut divide, Streimann 17480, 24968, 24995 ( JE) , ibid., Manki Trig , Streimann & Bellamy 13066 ( LAE) ; 3 miles south-east to Bulolo , Shea 6488, 9096 ( NICH) ; Angabena ridge, Aseki-Bulolo road, Streimann 25918 ( JE) ; Gumi divide, head of Gumi creek, Streimann 25066, 25136 ( JE) ; Heads Hump , Streimann 17374, 17402 ( JE) ; Kaisenik logging area, Shea 6404 ( JE) ; Oomsis logging area, Streimann 25839 ( JE) ; Mt. Kaindi , Blue Point , Gradstein 3793 (G, U 3 packets); Pouyu village, Streimann & Tamba 12662 ( LAE) ; Nauti logging area, Streimann & Kairo 17299 ( LAE) .— NEW BRITAIN: slopes of Mt. Ulawun , Streimann 41386 ( NICH) .— NEW IRELAND: Mussau Island , Bismarck Archipelago, KØie & Olsen 2070 (C, JE) .— SOUTHERN HIGHLANDS: Tari-komo road, Streimann 32712 ( BR, JE, LAE, W); Piribu sawmill, Streimann 32473 ( JE) ; Lama sawmill, Streimann 26605, 26611 ( JE) .— WEST NEW BRITAIN: slopes of Mt. Ulawun , Streimann 41386 ( JE) ; Nakanai Mts. , Streimann 40478 ( JE) , ibid., Geleo-Lasilai logging area at Mt. Laliti, Streimann 41086 ( JE) .— WEST PROVINCE: Star Mts , Sibil-valley , van Zanten 559b (L), ibid., Antares Mts , van Zanten 559/B (L).— WESTERN HIGHLANDS: Baiyer river-Ruti road, Jimi valley, Streimann 22061 ( JE, LAE) , 22242 (BR, JE, LAE).— WEST SEPIK: Antingin , Eliptamin, Macrosimnok 4 ( JE) .

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Australia. QUEENSLAND: Cardwell, Stoney creek , 21 May 1982, Stone s.n. ( U) ; El Arish, 11 July 1982, Stone s.n. ( U) ; Cook district, Bellenden Ker National Park, Thiers & Halling 2397 ( U) ; Main coast range, Daintree River National Park, Pócs & Streimann 9987/L ( GOET) ; Cape Tribulation area, Gap creek, Pócs & Streimann 99105/K ( GOET) ; Boonjie state forest, Streimann 27553 ( JE) ; Cooroo logging area, Streimann 29943, 29970 ( JE) ; Dunn creek, Kirrima road, Streimann 31234, 31241, 31255 ( JE) , 31257 ( JE, NICH); Cardwell range, Streimann 28591 ( JE) ; Koombooloomba dam road, Streimann 28910, 28923 ( JE) ; Culpha creek catchment, Streimann 28982 ( JE) ; Mossman Gorge , 26 May 1975, Stone s.n. ( JE) ; Main coast range, Streimann & Gray 27172 ( JE) ; Walter Hill range, Streimann 30477 ( JE) ; Mt. Lewis NW of Cairns, Hicks 11576 ( U) , ibid., 29 June 1982, Stone s.n. ( U) , ibid., 29 June 1984, Stone s.n. ( U) ; The Boulders Rain Forest Reserve along Babinda creek, Pócs & Streimann 9983/L (G); North Kennedy , Renner 1679 ( NSW) , ibid., Babinda creek , Renner 1842c ( NSW) , ibid., Little Birthday creek , 29 June 2005, Brown 2005/86 & Renner s.n. ( NSW) .

Caroline Island. Mt. Nanlaut , 28 June 1949, Glassman s.n. (F). New Caledonia. NORTH: Diahot, l’Ignambi, Hürlimann 2858a (G, GOET, PC) , 2858 ( U); Mt. Panie , Morat 6909 ( PC) .— SOUTH: l’Est , Hürlimann 2215 (G, GOET, PC, U) ; Mé Amméri , Guillaumin & Baumann-Bodenheim 9097 ( GOET) ; south of Kouanémoa Mts. , Hürlimann 2334 (G, GOET, PC, U) ; Rivière Bleue , Guillaumin & Baumann-Bodenheim 10894 ( GOET, PC, U) ; Mt. Ouénarou , Guillaumin & Baumann-Bodenheim 11875A ( GOET) ; Mois de Mai , Yaté, Baumann-Bodenheim 14028 (G, GOET, PC) ; Ni , Hürlimann 2709 (G, PC, U) ; Tao , January 1910, Franc s.n. (G, PC, S, W); Yaté, Kee 42654 ( PC) . Fiji. NAITASIRI: Mt. Tomanivi , Hürlimann T 1096e ( GOET) . Samoa. UPOLU: montane forest west of Tiavi , Schutze-Motel 3601 ( JE) .

Taxonomic notes:— The difference among Thysananthus minor , T. spathulistipus , T. spathulistipus var. borneensis , and T. triquetrus is only the number of tooth of the leaf margins ( Figs. 19A, B ; 20A, B, D–F, H ). Variation in leaf dentation is common and is also seen in T. comosus and T. discretus . I, therefore, reduce T. minor , T. spathulistipus var. borneensis , and T. triquetrus to synonyms of T. spathulistipus . Thysananthus spathulistipus is morphologically most similar to T. appendiculatus . Differences are discussed under the latter species.

The original material of the species in the Nees herbarium in STR contains 10 specimens, mostly from " Bantam " but only one annotated as originating from "Leback". The latter specimen is considered the holotype .

Thysananthus spathulistipus is a rather polymorphic species with respect to the dentation of leaves, which varies from edentate to strongly dentate as seen in T. comosus , T. discretus , and T. mollis . In montane forests of Sulawesi, Gradstein (pers. comm.) noted that phenotypes from below 2000 m were green in color and rather small whereas those from the upper montane cloud forest at 2300–2600 m were brown-green when fresh and more robust. Molecular analysis has resolved T. spathulistipus as polyphyletic ( Sukkharak et al. 2011b).

Figures

FIGURE 1. Method of measurement in Thysananthus. A. Leaf, ventral view; a = appendage, d = dorsal leaf margin, f = lobule free margin, k = keel, l = leaf length, lb = lobule length, t1,2 = lobule teeth, u = auricle, v = ventral leaf margin, w = leaf width, wb = lobule width. B. Underleaf, ventral view; u = auricle, l = underleaf length, w = underleaf width. C. Perianth, ventral view; b = beak, l = perianth length, w = perianth width. D. Portion of shoot with Lejeunea-type branching, ventral view; c = collar. E. Portion of shoot with androecia, gynoecium, and lejeuneoid innovation (dichasial pattern), ventral view. F. Flagellum at stem base, ventral view. G. Dorsal leaf bases are distinctly extending across the dorsal midline of the stem, indicating that lateral merophytes are dorsally interlocking; h= hyaline papillae. H. Underleaf base in longitudinal section. (A from the lectotype of T. appendiculatus; B–C from the holotype of T. discretus; D from T. spathulistipus, Sukkharak 766 (BKF, GOET); E from T. spathulistipus, Sukkharak 746 (BKF, GOET); F from isolectotype of T. comosus; G from T. spathulistipus, Sukkharak 716 (BKF, GOET); H from holotype of T. discretus).

FIGURE 5. Distribution of Thysananthus species. A. Thysananthus amazonicus (Spruce) Schiffn. B. Thysananthus spathulistipus (Reinwardt, Blume & Nees) Lindenb. C. Thysananthus convolutus Lindenb. D. Thysananthus mollis Steph., E. Thysananthus pancheri (Steph.) Hürl. F. Thysananthus anguiformis (Hook.f. & Taylor) Taylor ex Gottsche, Lindenb. & Nees G. Thysananthus comosus Lindenb. H. Thysananthus fruticosus (Lindenb. & Gottsche) Schiffn. I. Thysananthus aculeatus Herzog. J. Thysananthus appendiculatus Steph. K. Thysananthus montanus Gradst., X.-L. He & Piippo. L. Thysananthus retusus (Reinw., Blume & Nees) B.M.Thiers & Gradst. M. Thysananthus gottschei (Jack & Steph.) Steph. N. Thysananthus ciliaris comb. nov. O. Thysananthus discretus Sukkharak & Gradst.

FIGURE 19. Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenb. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral view. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apex. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L–M. Leaf lobule apices. N. Cross section of stem. O. Oil bodies. (A–N from the holotype; L from Pócs 9358/B (EGR); O from Sukkharak 716 (BKF, GOET)).

FIGURE 20. Thysananthus spathulistipus (Reinw., Blume & Nees) Lindenb. A–D. Portion of shoot, ventral view. E–H. Leaves, ventral view. (A, E from the holotype of T. spathulistipus var. borneensis; B, F from the holotype of T. minor; C, G from the holotype of T. fuscobrunneus; D, H from the holotype of T. triquetrus).

FIGURE 28. Thysananthus retusus (Reinw., Blume & Nees) B.M.Thiers & Gradst. A–C. Portions of shoot, ventral view. D. Leaves, ventral view. E. Underleaves, ventral view. F. Female bracts, ventral view. G. Female bracteole, ventral view. H. Perianth, dorsal view. I. Perianth, ventral view. J–L. Leaf apices. M. Margin cells of leaf lobe. N. Median cells of leaf lobe. O. Basal cells of leaf lobe, showing the vitta. P–Q. Leaf lobule apices. R. Cross section of stem. S. Oil bodies, non-vittate cells. T. Oil bodies, vittate cells. (A–R from the holotype; S–T from Sukkharak 733 (BKF, GOET)).