Thysananthus appendiculatus Stephani (1912a: 794)

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

3. Thysananthus appendiculatus Stephani (1912a: 794) . Thysanolejeunea appendiculata Stephani (1896: 138) , nom. inval. Lectotype (designated by Verdoorn 1934a): PAPUA NEW GUINEA. Western: Fly River Branch, Bäuerlein 85 (lectotype G!); Icon. Steph. nr. 10166.

Plants dioicous, with projecting growth, turning upwards and becoming ascending to erect, yellowish brown in herbarium specimens, up to 10 cm long × 2–2.2 mm wide. Stems rather rigid; ventral merophyte 9–12 cell rows wide; stem in cross section orbicular, 228–260 µm high × 188–250 µm wide, 13–14 cell layers high, composed of 41–55 epidermal cells surrounding 95–118 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 oblong-falcate, 1.1–2.5 × 0.8–1.4 mm, apex apiculate, dorsal base auriculate, auricle 77–225 × 75–250 µm, dorsal margin with 3–5 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, ventral margin incurved 1/2 × leaf length, with 8–10 triangular teeth, the teeth consisting of 3–8 cells, being 2–3 cells wide at base and ending in a row of 1–2 cells; cells elongate-hexagonal with acute ends, vitta absent, marginal cells 7–13 × 5–8 µm, median cells 17–35 × 5–10 µm, basal cells 30–57 × 12–15 µm, trigones cordate, often coalesced, intermediate thickenings 0–1 per cell; oil bodies unknown. Lobules oblong-rectangular, 0.4–0.5 × 0.1–0.2 mm, 1/5–1/3 × lobe length; appendage on surface of lobule base not developed; keels with appendages on one side where leaves and underleaves are free and opposite to adnate ones; lobule apex oblique, free margin continuing into the ventral lobe margin, apex 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 of one cell only, often absent. Underleaves imbricate, slightly squarrose, broadly spathulate, 0.9–2.2 × 0.5–0.9 mm, 2 × stem width, apex truncate to emarginate, plane, margin with 9–11 triangular teeth, the teeth consisting of 3–6 cells, being 2–3 cells wide at base, apex of one cell, bases cuneate or auriculate, 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 × 5–7 µm. Androecia terminal-intercalary on lateral branches, bracts and bracteoles in 4–13 pairs, bracts hypostatic, 0.6–0.9 × 0.4–0.6 mm, apex acute, margins entire; antheridia 2 per bract. Gynoecia with 2 lejeuneoid innovations forming a dichasial pattern; lobe ovate, 2.2–2.4 × 1–1.1 mm, apex apiculate, margins in upper 1/3 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; lobules broadly ovate, 1/2 × lobe length, apex apiculate, 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, 1.8–1.9 × 0.9–1.1 mm, apex emarginate, 1/3 × bracteole length with 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, margins slightly recurved. Perianths oblong-cylindrical, 1.9–2.1 × 0.8–0.9 mm, keels in upper 1/ 3 with numerous laciniate teeth, the teeth consisting of 3–6 cells, being 2–3 cells wide at base and ending in a row of 2–4 cells; beak 62–65 µm (5 cells) in length. Fig. 11.

24 • Phytotaxa 193 (1) © 2015 Magnolia Press

SUKKHARAK

Distribution and ecology:— Endemic to Western Melanesia: Western New Guinea ( Indonesia) and Papua New Guinea; 400–2600 m; on trunks, branches, large vines, stems of treelets in old garden sites, grasslands with regrowth species, understory of lowland rain forests (secondary forests) and montane forests. Fig. 5J .

Representative specimens:— Indonesia. MOLUCCAS: Mt. Gunung Nona, Balázs & Pócs 95/6 ( NICH) .— WESTERN NEW GUINEA: Papua, Cycloop Mts., van Royen & Sleumer 6006 ( JE, L); West Papua, Arfak Mts. , October 1872, Beccari s.n. ( JE, L) . Papua New Guinea. CENTRAL: K. B. Sawmill, Streimann & Naoni 16567 ( JE, LAE) .— EASTERN HIGHLAND: Daulo Pass, Streimann 18124 ( JE) , Streimann & Bellamy 18093 ( JE) , Streimann & Kairo 18138 ( LAE) ; track to Mt. Michael , Streimann 18807 ( JE) .— GULF: Bema-Kaintiba road, Streimann 33647 ( JE) .— MOROBE: Wau, Mt. Kaindi , Gradstein 3875 ( U) , Kunai creek, Gradstein 3962 (G); Ogeramnang, Clemens 5414 ( JE, PC, S, W); Slate creek & Gumi creek divide, Streimann 13829 ( JE) , 13826, 13838 (LAE), 13834, 13859 (JE, KLU, LAE); Bulolo-Watut divide, Streimann 25022 ( JE) ; Kaisenik logging area, Shea 6336 ( JE) , 6364 (NICH); Upper Watut river , Streimann 23092 ( JE, LAE) ; Aseki-Bulolo road, Streimann 23191, 26124 ( JE, LAE) , 23195 (LAE); Pouyu village, Streimann & Tamba 12674 ( JE, LAE) ; Oomsis logging area, Streimann 25836 ( JE, LAE, W); Aiuwa-Bakia track, Streimann & Tamba 12358 ( JE, KLU, LAE) , 12276, 12362, 12377, 12413 (LAE).— SIMBU: Dirima mission, Toia 171 ( LAE) .— SOUTHERN HIGHLANDS: Margarima- Tari road, Streimann 24378, 24388 ( JE, LAE, W); Onim forestry station, Streimann 24628, 24740 ( JE) ; Komo-Tari road, Streimann 32618, 32631 ( JE) ; Kengaput, Streimann 26902 ( JE) , 23702 (LAE); Lama Sawmill logging area, Streimann 26422, 26562 ( JE) ; Tari gap, Streimann 32730 ( JE) .— WESTERN HIGHLAND: Kum Magei Mts. , Streimann 20679 ( JE, LAE) .

The report of Thysananthus appendiculatus from India by Daniels & Raja (2011 a, 2011b) is erroneous according to Sukkharak & Gradstein (2011).

Taxonomic notes:— The most outstanding character of Thysananthus appendiculatus is the foliar appendage, which is developed on the keels of leaves on one side of the stem only, on the side where leaves and underleaves are free and opposite to adnate ones. This character is otherwise found in T. anguiformis , T. convolutus var. laceratus , T. discretus , T. fruticosus , and T. gottschei var. continuus . Differences are discussed under T. discretus . Thysananthus appendiculatus is similar to T. spathulistipus , the latter species differing essentially by the absence of the appendage and by the triangular, 1–2 cells long perianth teeth.

The epithet “ appendiculatus ” is derived from the foliar appendage which is found on the keels of leaves as described by Stephani in the original description. The original drawing of Thysananthus appendiculatus in Stephani’s “Icones Hepaticarum Ineditae” is erroneous in showing a foliar appendage on both sides of the stems. The illustrated left-hand side appendages are probably parts of the dissected underleaves. Note that the illustration of the species in Gradstein et al. (2002: Fig. 44) does not show the foliar appendage.

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 12. Thysananthus ciliaris (Sande Lac.) Sukkharak. 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. Leaf lobule apices. M. Cross section of stem. (All from Gradstein 10366 (GOET)).

FIGURE 13. Thysananthus comosus Lindenb. A. Portion of shoot, ventral view. B–C. Leaves, ventral view. D–E. Underleaves, ventral view. F. Female bracts, ventral view. G. Female bracteole, ventral view. H. Perianth, dorsal view. I. Perianth, ventral view. J. Leaf apex. K. Margin cells of leaf lobe. L. Median cells of leaf lobe. M. Basal cells of leaf lobe. N.–P. Leaf lobule apices. Q. Cross section of stem. R. Oil bodies (A, C, E–Q from the lectotype, B, D from Schiffner s.n. (GOET), R from Sukkharak 730 (BKF, GOET)).

FIGURE 14. Thysananthus convolutus 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 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. Oil bodies. (A–M from the lectotype; N from Sukkharak 717 (BKF, GOET)).

FIGURE 15. Thysananthus convolutus var. laceratus (Steph.) Sukkharak. A. Portion of shoot, ventral view. B. Leaves, ventral view. 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. L. Leaf lobule apices. M. Leaf lobule base. N. Cross section of stem. (All from the lectotype).

FIGURE 16. Thysananthus discretus Sukkharak & Gradst. A–B. Portion of shoot, ventral view. C–D. Leaves, ventral view. E–F. Underleaves, ventral view. G. Female bracts, ventral view: a appendage. H. Female bracteole, ventral view. I. Perianth, dorsal view. J. Perianth, ventral view. K. Leaf apex. L–M. Margin cells of leaf lobe. N. Median cells of leaf lobe. O. Basal cells of leaf lobe. P. Leaf lobule apices. Q. Cross section of stem. (A, C, E, G–K, M–Q from the holotype; B, D, F, L from Streimann 13622 (JE)).

FIGURE 17. Thysananthus gottschei (J.B.Jack & Steph.) Steph. A. Portion of shoot, ventral view. B. Leaves, ventral view. 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. L. Leaf lobule apex. M. Cross section of stem. (A–C, H–M from the lectotype; D–G from Gradstein 10324 (GOET)).

FIGURE 18. Thysananthus gottschei var. continuus Sukkharak. 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. Margin cells of leaf lobe. I–J. Leaf lobule apices. K. Leaf lobule base. (All from the lectotype).

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 21. Thysananthus anguiformis (Hook.f. & Taylor) Taylor ex Gottsche, Lindenb. & Nees. 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–J. Leaf apices. K. Margin cells of leaf lobe. L. Median cells of leaf lobe. M. Basal cells of leaf lobe. N–R. Leaf lobule apices: h = hyaline papilla. S. Cross section of stem. (All from the lectotype).

FIGURE 22. Thysananthus pancheri (Steph.) Hürl. A. Portion of shoot, ventral view. B. Leaves, ventral view. 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–I. Leaf apices. J. Margin cells of leaf lobe. K. Median cells of leaf lobe. L. Basal cells of leaf lobe. M–N. Leaf lobule apices: h = hyaline papilla. O. Cross section of stem. (All from the lectotype).

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)).