Thysananthus fruticosus (Lindenb. & Gottsche) Schiffner (1893: 130)

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

12. Thysananthus fruticosus (Lindenb. & Gottsche) Schiffner (1893: 130) . Bryopteris fruticosa Lindenberg & Gottsche (1847: 737) . Dendrolejeunea fruticosa (Lindenb. & Gottsche) Lacouture (1908: 104) . Caudalejeunea fruticosa (Lindenb. & Gottsche) Stephani (1912b: 14) . Lectotype (designated by Verdoorn 1934a): INDONESIA. Java: Mt. Sajira, without date, Blume s.n. (lectotype: STR!).

Bryopteris filicina (Sw.) Nees γ fruticosa Gottsche, Lindenberg & Nees (1845: 285) , nom. nud. syn. fide Verdoorn (1934a): 182.

Thysananthus manillanus Gottsche (1857: 342) . Type : PHILIPPINES. “ Manilla, Gaudichaud 115 ” (holotype PC; isotype PChb. Bescherelle!).

Bryopteris vittata Mitten (1871: 411) . Type : AUSTRALIA. Norfolk Island: without date, Robinson s.n. (holotype: G!).

Thysanolejeunea lanceolata Stephani (1896: 139) . Thysananthus lanceolatus (Steph.) Stephani (1912a: 796) . Type: PAPUA NEW GUINEA. Morobe: “Sattelburg”, Kärnbach 56 (holotype G!).

Thysananthus sinclairii (Mitt.) Stephani (1912a: 792) . Lejeunea sinclairii Mitten (1862: 19) . Type: FIJI. “Insulae Vitienses” “inter nr. 843”, Seemann s.n., “in Herb. Gourlie et Mitten” Sinclair s.n. cf. Geissler & Bischler (1987), syn. fide Verdoorn (1934a): 183.

Thysananthus abietinus Stephani (1912a: 794) . Type : VANUATU. “Insula Aneitum, 1882, Braithwaite s.n. ” (holotype G!); Icon. Steph. nr. 10165.

Thysananthus lauterbachii Stephani (1912a: 797) . Type: PAPUA NEW GUINEA. Madang: “Gogal”, Lauterbach 1060 (holotype G!); Icon. Steph. nr. 10193.

Thysananthus ovistipulus Stephani (1912a: 798) . Type : PAPUA NEW GUINEA. without location, without date, Musgrave s.n. (holotype G!); Icon. Steph. nr. 10202.

Thysananthus ( Lejeuneaceae , Marchantiophyta) Phytotaxa 193 (1) © 2015 Magnolia Press • 41

Thysananthus bowienus Stephani in Stephani & Watts (1914: 134). Type: VANUATU [“Novae Hebrides”]. Sanma: Santo , 1909, F.G. Bowie s.n. (holotype G!); Icon. Steph. nr. 10170.

Thysananthus densus Stephani (1924: 565) . Type: PAPUA NEW GUINEA. Milne Bay: 1911, Conway s.n. (holotype G!); Icon. Steph. nr. 10176.

Plants autoicous, with dendroid habit, having creeping stoloniform stems and upright, regularly pinnate branches, yellowish in herbarium specimens, up to 14.8 cm long × 1.8–2.1 mm wide. Stems rather rigid; ventral merophyte 6–11 cell rows wide; stem in cross section orbicular, 170–340 µm high × 135–300 µm wide, 9–20 cell layers high, composed of 30–62 epidermal cells, 25–50 subepidermal cells in 4–6 layers high surrounding 67–218 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 asymmetrically ovate-falcate, 1.1–2.7 × 0.5–1.3 mm, apex acute, dorsal base cordate, dorsal margin with 2–4 triangular teeth, the teeth consisting of 3–7 cells, being 3–4 cells wide at base, apex of one cell, ventral margin plane, with 2–6 triangular teeth, the teeth consisting of 3–7 cells, being 3–4 cells wide at base, apex of one cell; cells elongate-hexagonal with acute ends, vitta present in midportion of lobe, extending to base, 2/3 × lobe length, 8–10 cell rows wide, 17–26 cells long, marginal cells 7.5–15 × 7.5–12.5 µm, median cells 18–25 × 8–15 µm, vitta 38–60 × 13–23 µm, trigones cordate, often coalesced, intermediate thickenings 0–1 per cell; oil bodies unknown. Lobules oblong, 0.1–0.5 × 0.1–0.3 mm, ± 1/3 × lobe length, appendage on surface of lobule base not developed; keel with appendages on both sides of stem; lobule apex transverse, not or slightly continuing into the ventral lobe margin, entire or with one triangular tooth, the tooth consisting of 3 cells, being 2 cells wide at base, apex of one cell. Underleaves imbricate, slightly squarrose, ovate, 0.5–1 × 0.5–0.9 mm, 3 × stem width, apex truncate to emarginate, margin with 8–12 triangular teeth, the teeth consisting of 7–9 cells, being 2–3 cells wide at base and ending in a row of 6–7 cells, bases cuneate-slightly auriculate, underleaf bases not adnate with leaves; cells 23–43 × 8–10 µm. Androecia terminal-intercalary on lateral branches, bracts, and bracteoles in 5–21(–38) pairs, bracts hypostatic, 0.6–1.3 × 0.3–0.6 mm, apex acute, margin entire; antheridia 2 per bract. Gynoecia with 1–2 lejeuneoid innovations forming a monochasial or dichasial pattern; lobe lanceolate, 1.4–2 × 0.3–0.6 mm, apex apiculate, margins in upper 1/3 with triangular teeth, the teeth consisting of 3–10 cells, being 2–4 cells wide at base and ending in a row of 1–2 cells; lobule broadly ovate, 1/2 × lobe length, apex apiculate, margins with triangular teeth, the teeth consisting of 3–5 cells, being 2–3 cells wide at base and ending in a row of 1–2 cells; bracteole spathulate, 1.3–2 × 0.6–0.8 mm, apex truncate, 1/2 × bracteole length with triangular teeth, the teeth consisting of 3–7 cells, being 2–3 cells wide at base, apex of one cell, margins recurved. Perianths oblong-cylindrical, 1.3–2.3 × 0.7–0.9 mm, keels in upper 1/3 with laciniate teeth, the teeth consisting of 4–9 cells, being 2–3 cells wide at base and ending in a row of 2–5 cells; beak 33–75 µm (3–5 cells) in length. Figs. 23 , 24 .

Additional illustrations:— Thiers & Gradstein (1989, p. 72, Fig. 27 ); Gradstein et al. (2002, p. 21, Fig. 11 ).

Distribution and ecology:— Malesia, Australia (Queensland), and the Pacific region; 25–2000 m; on tree trunks, twigs, and rotten logs in lowland rain forests (primary and secondary), and montane forests. Fig. 5H .

Representative specimens:— Thailand. PHANG NGA: Sri Phang Nga National Park , Chantanaorrapint 2112 ( PSU) . Malaysia. JOHORE: Lubuk Tapa , Mohamed & Sabda 20, 42, 44, 59, 70, 110, 139, 142, 180 ( KLU) , Sabda 222 ( KLU) ; vicinity of Sungai Selor , Sabda 219, 223, 260 ( KLU) ; Gg. Sumalayang, Chin 744 ( KLU, SING) .— NEGERI SEMBILAN: Pasoh Forest Reserve , Kien-Thai 7425 ( KLU) .— PAHANG: Taman Negara , 19–23 June 1995, Mohamed s.n. ( KLU) ; Ulu Cheneras, Burkill 15669(b) ( SING) ; Kuala Tembeling , Holttum 20541 ( SING) .— SABAH: Danum valley conservation area, Ellis 38 ( BM) ; 2 miles north of Kinabatangan river at Bukit Garam, Wood 1437 ( BM) ; Sandakan, Sapagaya F.R., Kadir & Enggoh 10564 ( BM) ; Timbun Mata Island, Mapat river , Keith 7358 ( BM) ; Tawau , Kadir A 2023 ( BM) ; Mt. Rara Forest Reserve, Shea 2069 (KLU).— SARAWAK: Baram district, September 1892, Evereth s.n. (G 2 packets); Mt. Dulit , Richards 2277 ( BM, JE, SING) ; Buda National Park, Mohamed & Kien-Thai 5331 ( KLU) ; Mulu National Park, Mohamed & Kien-Thai 5434 ( KLU) .— SELANGOR: Hulu Langat, Congkat river, Ilkiu-Borges & Yong 3026 ( GOET) ; Gunung Nuang , 23 November 1998, Mohamed s.n. ( KLU) . Singapore. Chan Chu Kan , 1890, Ridley s.n. (G) . Indonesia. KALIMANTAN: peak of Balikpapan , Meijer B 1339e ( BM 2 packets, G, PC, SING), B1391 a ( SING) .— SULAWESI: Menado, without date, Vriese s.n. (F); without location, without date, Kiese s.n. (S 2 packets) .— SUMATRA: Engano Islands , “Modigliani” s.n., ex hb. Levier 152 ( BM, G 3 packets, PC) .— WESTERN NEW GUINEA: Bird’s Head peninsula, van Royen & Sleumer 7268 ( BR, U) , 16 February 1957, Bergman s.n. ( NICH) ; Mt. Arfak , 1872, Beccari s.n. ( JE) ; Andai, 1872, Beccari s.n. ( JE, U) ; “ Bogaljim ”, without date, Missionare s.n. (S 3 packets); Padang, 1924 ,

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Blum s.n. (JE); Mt. Jaya , Edwards 4103A ( BM) . Philippines. BENGUET: Baguio and vicinity, Robinson 2–107, Bureau of science no. 14101 ( BM) .— LAGUNA: Siniloan national botanical garden, Alvarez & al. 0–771073 ( JE) .— NUEVA ECIJA: without location, Santos 212 ( JE) .— QUEZON: Botanical garden of University of the Philippines Los Banõs, Onraedt 85.P.11304 ( BR) ; Polillo, McGregor 2–39, Bureau of science no. 10499 ( BM, G), ibid., Bureau of science no. 10495 (G); Dimasingay, Baler, Santos 235 ( JE) .— SAMAR: Mt. Purog , without date, Edaño s.n. ( U) .— SURIGAO DEL SUR: Muni river , “Pahl” 85.P.11973 ( BR) .— TAYABAS: without location, Robinson 89604 ( BM, G). Papua New Guinea. CENTRAL: Moroka , July–August 1893, Loria s.n., ex hb. Levier 109 ( BM 2 packets, G 4 packets, PC); Koitaki plantation, Carr 12078, Hep. Sep. Crit. Verdoorn 449 ( BM 3 packets, BR, C, G, GOET, JE, PC, SING 2 packets, U 2 packets); 30 miles north of Port Moresby, Robbins 4132 ( JE) ; Angabanga river, Streimann & Naoni 16213 ( JE, LAE) ; Kuriva logging area, Streimann & Vinas 14399, 14408, 14426 ( JE, LAE) , 14434 (BR, JE, KLU, LAE); road to Muscrave river, Streimann & Naoni 15206 ( JE) , 15191, 15214, 15250 (JE, LAE).— EASTERN HIGHLANDS: Kassam pass, Streimann & Umba 11431 ( JE, LAE) .— EAST SEPIK: Hawain river range, Robbins 2081 ( JE) ; Wewak-Angoram, Robbins 2345 ( JE) ; Hunstein river, Hoogland & Craven 10.752 ( JE 2 packets, MEL); Kassam pass, Streimann & Umba 11451 ( JE, LAE) ; Prince Alexander Mts. , Robbins 2016 ( JE) .— MILNE BAY: Goodenough Island , Brass 22982 ( JE) ; north slope of Mt. Dayman , Brass 22185 ( JE) .— MADANG: Lower Ramu-Atitau , Robbins 1433, 1691 ( JE) .— MOROBE: Busu river logging area, 16 June 1968, Weber s.n. (F 3 packets, JE, U) ; Herzog Mts., Streimann & Umba 11038, 11055 ( KLU, LAE) , 11054 (JE, LAE), 11053 (BR, JE, KLU, LAE); Gurakor creek, Lae-Bulolo road, Bellamy 1248 ( JE, KLU, LAE) ; logging area, 15 km East of Bulolo, Streimann & Bellamy 13152 ( JE, KLU, LAE) , 13184 (BR, JE, KLU, LAE), 13762 (LAE); Gumi divide, head of Gumi creek , Streimann 25733 ( JE, LAE) ; Slate creek and Gumi creek divide, Streimann 13954 ( BR, KLU) ; heads Hump, Streimann 17437, 17440 ( JE) ; near Nauti village, Streimann & Elix 26229 ( JE) ; Nauti logging area, upper Watut river, Streimann & Kairo 17245, 17263, Streimann 17321, 17327 ( JE) , 17309 (GOET, JE, KLU, LAE); Sattelberg, Clemens 264/e ( JE) , ibid., June 1988, Nyman s.n. ( BM, C 2 packets), 21 July 1904, Pöch s.n., Museo Hist. Natur. Vindobonensi 4035 (G, S 2 packets, U) .— NEW IRELAND: Mussau Island, Bismarck Archipelago, KØie & Olsen 2051, 2061 (C).— ORO: Managalase area , Pullen 5685 ( JE) .— SOUTHERN HIGHLANDS: Batteri, Kagua-Erave road, Streimann 23433 ( JE, LAE) .— WEST NEW BRITAIN: Willaumez Peninsula , Kolema 28 ( JE, NICH) ; Mosa N.T.A., Lakemata reforest creek, Kolema 101 ( JE) ; Honde- Laulimi T.R.P., Kolema 121 ( JE) .— WESTERN SEPIK: headwaters of Kong Kong creek , Mundua 283 ( JE) . Solomon Islands. CENTRAL: Russel Islands , December 1935, Lever s.n. ( BM) . GUADALCANAL: without location, without date, van Zanten 68–2360, 68–2370 ( JE) .

Australia. NORFOLK ISLAND: 29°02' S 167°57' E, without date, Robinson s.n. ( MEL) .

Vanuatu. New Hebrides, 1909, Bowie s.n. ( BM). Fiji. REWA: Suva, 1898, Armstroy s.n. (G). Samoa. UPOLU: wihtout location, without date, Graeffe 628 (G), 1894, Reinecke s.n. (G). American Samoa. TUTUILA: without location, without date, Graeffe 664 (G). Niue. near Alofi, Sykes 289 (F). Cook Islands. Rarotonga, Gardner 72/e ( JE).

Reported from Australia (Queensland) by Thiers & Gradstein (1989).

Taxonomic notes:— Thysananthus fruticosus was originally described as a species of Bryopteris and was placed in a separate subgenus Dendrolejeunea (of Lejeunea s.l.) by Spruce (1884). Lacouture (1908) raised Dendrolejeunea to generic level but this has not been followed by later authors, who accepted Schiffner’s attribution (1893) of T. fruticosus to Thysananthus . The genus Dendrolejeunea was reinstated for T. fruticosus again by Gradstein (1992a). Thysananthus fruticosus differs from other members of Thysananthus by its dendroid habit, having creeping stoloniform stems and upright, regularly pinnate branches. The species shares the presence of a vitta with T. mollis , T. montanus , and T. retusus and was for this reason placed by Verdoorn (1934a) in Thysananthus sect. Vittatae . However, the molecular results showed that D. fruticosa is a member of Thysananthus ( Sukkharak et al. 2011b) .

Figures

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 11. Thysananthus appendiculatus Steph. A. Portion of shoot, ventral view. B. Leaves, ventral view: a = appendage. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. E. Female bracteole, ventral view. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apices. M. Cross section of stem. N. Portion of shoot, ventral view. O. Leaves, ventral view. P. Underleaves, ventral view. (A–C, H–M from the lectotype; D–G from Streimann 13859 (JE); N–P from Gradstein 3875 (GEOT)).

FIGURE 23. Thysananthus fruticosus (Lindenb. & Gottsche) Schiffn. A. Portion of shoot, ventral view. B. Leaves, ventral view: a = appendage. C. Underleaves, ventral view. D. Female bracts, ventral view: a = appendage. 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, showing the vitta. L–M. Leaf lobule apices. N. Cross section of stem. (A–C, H–N from the lectotype; D–G from Brass 22185 (JE)).

FIGURE 24. Thysananthus fruticosus (Lindenb. & Gottsche) Schiffn. A–B. Portion of shoot, ventral view: a = appendage. C–D. Leaves, ventral view: a = appendage. E. Underleaves, ventral view. (A, C, E from Beccari s.n. (JE); B, D from Streimann 25733 (JE)).

FIGURE 27. Thysananthus montanus Gradst., X.-L. He & Piippo. A. Portion of shoot, ventral view. B. Leaves, ventral view. C. Underleaves, ventral view. D. Female bracts, ventral view. E. Female bracteole. F. Perianth, dorsal view. G. Perianth, ventral view. H. Leaf apices. I. Margin cells of leaf lobe. J. Median cells of leaf lobe. K. Basal cells of leaf lobe. L. Leaf lobule apex. M. Cross section of stem. (All from the holotype).