Thysananthus amazonicus (Spruce) Schiffner (1893: 130)

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

2. Thysananthus amazonicus (Spruce) Schiffner (1893: 130) . Lejeunea (subg. Thysanolejeunea) amazonica Spruce (1884: 106) . Lectotype (designated by Fulford 1941): BRAZIL. Pará: Spruce s.n. (lectotype MANCH n.v.; isolectotypes G, MANCH, NY).

Plants autoicous and paroicous, with projecting growth, turning upwards and becoming ascending to erect or projecting downwards and becoming pendent, green to dull brown in the field, yellowish to reddish brown in herbarium specimens, up to 6 cm long × 2–3.5 mm wide. Stems rather rigid; ventral merophyte 7–9 cell rows wide; stem in cross section orbicular, 175–188 µm high × 161–167 µm wide, 12–14 cell layers high, composed of 29–32

22 • Phytotaxa 193 (1) © 2015 Magnolia Press

SUKKHARAK epidermal cells surrounding 63–73 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 oblong-falcate, 1.2–1.7 × 0.6–1 mm, apex apiculate, margin entire, dorsal base cordate, ventral margin incurved 1/2 × leaf length; cells elongate-hexagonal with acute ends, vitta absent, marginal cells 5–7.5 × 7.5–12.5 µm, median cells 15–27.5 × 7.5–10 µm, basal cells 50–70 × 10–22 µm, trigones cordate, often coalesced, intermediate thickenings 0–2(–3) per cell; oil bodies (3–)4–6 per cell. Lobules oblong-rectangular, 0.2–0.5 × 0.1–0.2 mm, 1/5–1/3 × 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 entire or with 1–2 triangular teeth, the first tooth consisting of 3–6 cells, being 2–3 cells wide at base and ending in a row of 1–2 cells, the second tooth consisting of 3–5 cells, being 1–2 cells wide at base and ending in a row of 1–2 cells, often absent. Underleaves imbricate, slightly squarrose, broadly spathulate, 0.7–1 × 0.5–0.9 mm, 3–5 × stem width, apex emarginate-lunulate, plane, margin entire or with up to 12 triangular teeth, the teeth consisting of 7–9 cells, being 2–3 cells wide at base, apex of one cell, 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 15–20 × 5–8 µm. Androecia below the gynoecium or terminal-intercalary on lateral branches, bracts and bracteoles in 2–8 pairs, bracts epistatic or hypostatic, 0.8–1 × 0.3–0.5 mm, apex acute, margins entire; antheridia 2 per bract ( Gradstein, 1994: 1–2 antheridia). Gynoecia with one lejeuneoid innovation forming a monochasial pattern; lobe ovate, 1.3–1.5 × 0.6–0.8 mm, apex apiculate, margins in upper 1/3 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; lobules broadly ovate, 1/2–2/3 × lobe length, apex apiculate to slightly bifid, margin 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; bracteoles spathulate, 1.1–1.4 × 0.5–0.7 mm, apex emarginate to short bifid, 1/3 × bracteole length with 12–14 triangular teeth, the teeth consisting of 3–7 cells, being 2–3 cells wide at base and ending in a row of 1–2 cells long at apex, margins recurved. Perianths oblong-cylindrical, 1.4–1.7 × 0.6–0.8 mm, keels in upper 1/3 with triangular teeth, the teeth consisting of 3–5 cells, being 2–3 cells wide at base and ending in a row of 1–3 cells; beak 30–60 µm (3–4 cells) in length. Fig. 10 .

Additional illustrations:— Fulford (1941, p. 38, Figs. 28 –40); Gradstein (1994, p. 85, Fig. 23 ).

Distribution and ecology:— Tropical America ( Costa Rica, Colombia, Amazonian part of Brazil, Venezuela, Trinidad, and Guianas; disjunct in eastern Cuba); from sea level to about 1000 m; on branches, twigs, trunks or lianas, occasionally on logs, in marsh forest, swamp forest, evergreen forests on white sand, and savannas; usually in the forest canopy. Fig. 5A .

Representative specimens:— Costa Rica. LIMÓN: Tortuguero, 29 November 1994, Cleef & Kapelle s.n. ( GOET) , Timme 11158 ( GOET) . Colombia. AMAZONAS: Río Caquetá , TRA 12 km, Mohr & Sosa 20 ( GOET, U) . Venezuela. BOLÍVAR: Cerro Guaiquinima, Sipman 26654, 26678 ( U) . Guyana. CUYUNI-MAZARUNI: Essequibo river , near Bartica, Richards 188 ( BM) , 510 (BM, GOET).— DEMERARA-MAHAICA: Timehri, Thomson’s farm, Gradstein 4709 (G, GOET) , Cornelissen & ter Steege C 017 ( U) .— UPPER DEMERARA-BERBICE: Mabura hill, Cornelissen & ter Steege C 111, C679, C680, C681, C682, C743, C744, C761, C762, C829, C860, C861 ( U) .— UPPER MAZARUNI: North slope of Mt. Roraima, Gradstein 5151 (G, U) ; Waruma river , Gradstein 4993 (G, U) ; Mt. Latipu , Gradstein 5560, 5650 ( U) ; Waramadan, trail to Mt. Pwipwi, Gradstein 5693 ( BR, G, NICH, U); Jawalla , at confluence of Kukui river and Mazaruni river , Gradstein 4835, 4911 ( U) , 4857 (G, U). Suriname. NICKERIE: Nickerie, area of Kabalebo dam project, Bekker 1086a, 1090, 1160b, 1164b, 1166, 1174, 1176, 1177, 1195, 1296, 1343, 1346b, 1361a, 1375, 1387, 1427b, 1459, 1471, 1475, 1483b, 1494b, 1514, 1522b, 1526b, 1530, 1554, 1555, 1749c, 1756a ( U) , Florschütz-de Waard & Zielman 5187A, 5189A, 5529A ( U) ; Jodensavanne-Mapane kreek area, Lindeman 3937 ( GOET, PC) .— PARAMARIBO: Paramaribo, Suringar 598 (L) .— SIPALIWINI: Blanche Marie valley , Muñoz 98–19 ( GOET) . French Guiana. CAYENNE: Trésor reservation, Hartmann & al. 04–052 ( GOET) ; Montagne de Kaw , Hartmann & al. 04–117, 04–118 ( GOET) , Cornelissen & ter Steege C 0265, C0311 ( U) , Gradstein 5900 ( U) ; forest between savanna one and “petit savanna”, Hartmann & al. 04–120, 04–121, 04–123 ( GOET) ; Emerald Jungle village , Holz FG 00–51A, FG00–55 ( GOET) ; la Trinité, lower Comté river, Cremers 5557 ( BR) ; bridge over Comté river, Gradstein 6658 ( U) ; Montsinery, along “Risque tout” forest track, Gradstein 5793 ( U) ; Saül, along La Fumée trail, Gradstein 6131 ( U) , Aptroot 15381, 15447 ( U) , Montfoort & Ek 1164, 1165, 1166, 1167, 1168, 1169 ( U) , Gehrig s.n. ( GOET) .— SAINT-LAURENT-DU-MARONI: Charvein, 20 January 1914, Benoist s.n. ( PC) ; along trail from St. Laurent to Apatoa, Cornelissen & ter Steege C 0270 ( U) . Brazil. AMAZONAS: Lages river, Vital & Yano 718 ( U) ; Manaus, Reserva Experimental do INPA, Prance & al .

Thysananthus ( Lejeuneaceae , Marchantiophyta) Phytotaxa 193 (1) © 2015 Magnolia Press • 23

18714 (U); Tarumã Fälle, Schäfer-Verwimp & Verwimp 9815 ( BR); along Manaus-Caracaraí road, 8 July 1974, Vital & al. s.n. ( U), 13 November 1973, Berg & al. P19514 ( U); Trombetas river , Campbell & al. P22268 ( U), Prance & al. 22208 ( U); Madeira river , Ule 585 (G); Juruá river , Ule 338 (G); Rio Negro, Ule 571 (G) .

Additionally reported from Trinidad ( Fulford, 1941; Gradstein, 1994).

Taxonomic notes:— Thysananthus amazonicus is morphologically most similar to the Asiatic T. comosus , from which it is readily distinguished by (1) slightly squarrose spathulate and emarginate-lunulate underleaves (flat obovate and truncate in T. comosus ), (2) monoicy (dioicy in T. comosus ), (3) epistatic or hypostatic male bract lobules (only hypostatic in T. comosus ), and (4) keels bluntly toothed-winged in the upper half to subentire, teeth 1–3 cells long (keels laciniate, with 3–6 cells long teeth in T. comosus ). Thysananthus amazonicus may also be confused with T. spathulistipus . However, T. spathulistipus differs by its toothed (sometimes entire) leaf margin, truncate underleaves, hypostatic male bract lobules and occurrence in the paleotropics.

Thysananthus amazonicus often grows more or less pendent, especially when occurring on small branches or twigs ( Gradstein 1994). The leaf lobules of T. amazonicus may have one or two teeth and sometimes there is no tooth at all. According to Gradstein (1994) the leaf lobule is usually 1/4 or less the length of the leaf but in some Cuban specimens the lobules may be slightly larger, up to 1/3 × leaf length.

Along the Río Caquetá, Colombia, Thysananthus amazonicus is called “lama” and is used as a painkiller against snake and scorpion poisoning (Mohr & Sosa in sched., fide Gradstein 1994).

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 10. Thysananthus amazonicus (Spruce) Schiffn. 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–J. Leaf apices. K. Margin cells of leaf lobe. L. Median cells of leaf lobe. M. Basal cells of leaf lobe. N–Q. Leaf lobule apices. R. Cross section of stem. (All from Gehrig s.n. (GOET)).

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 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)).