Colura valida Jovet-Ast, Rev. Bryol. Lichénol.

Sangrattanaprasert, Jiroat, Chantanaorrapint, Sahut & Zhu, Rui-Liang, 2019, The genus Colura section Gamolepis (Lejeuneaceae, Marchantiophyta) in Malesian region, with the description of Colura sigmoidea, Phytotaxa 387 (1), pp. 40-54 : 51

Colura valida Jovet-Ast, Rev. Bryol. Lichénol. 30: 6. 1961. ( Fig. 5 )

Type: — VIETNAM. Lam Dong: Blao, School of Agriculture, December 1959, P. Tixier s.n. (holotype: PC!).

Plants pale green, ca. 1 cm long; shoots 1.4–1.7 mm wide; irregularly pinnately branched, branches of Lejeunea - type. Stems 90–152 μm in diameter, in transverse section consisting of 7 cortical cells (36–60 × 14–50 μm) and 3 medullary cells (30–40 × 16–32 μm); ventral merophyte of stem 2 cells wide. Rhizoids numerous, fasciculate at base of underleaves. Leaves imbricate to remote, spreading from stem at 30°–80° angle. Lobes 1.04–1.4 mm long, 0.61–1.02 mm wide; dorsal margin ±plane, entire to crenulate. Lobe cells thin-walled, trigones large, intermediate thickenings very distinct; marginal cells isodiametric to subquadrate, 24–50 × 20–30 μm; median cells rectangular to slightly hexagonal, 34–68 × 20–30 μm; basal cells slightly rectangular to hexagonal, 40–80 × 26–30 μm. Cuticle smooth. Oil bodies numerous per cell, homogenous, ellipsoid, 4–8 × 1.6–2.4 μm. Lobules narrowly tubular, flaring toward sac, 1.42–2 mm long. Lobule sacs elliptic, strongly inflated, 0.7–1.1 mm long, 0.35–0.41 mm wide, sac surface mamillose, apex obtuse, without apical crest. Valves intermediate-type, ovate to elliptic, 116–130 × 70–80 μm, composed of one circle of 21–24 hyaline marginal and 22–32 median cells with one basal median cell, basal median cell adnate with a hinge cell not forming a sigmoid curve in outline. Hyaline papillae bulbous to spherical, 18–20 μm in diameter. Valve frames outline semicircular with cell uniseriate, 100–120 μm wide. Underleaves distant, deeply bilobed, sinus obtuse; lobes lanceolate, 360–496 × 104–168 μm, 12–16 cells long, 6–9 cells wide at base; margin entire. Asexual reproduction by discoid gemmae, occurring on lobule sacs; gemma cells subquadrate to isodiametric, consisting of 27–29 cells.

Dioicous. Androecia terminal on lateral branches; male bracts in 3–5 pairs, hemispherical, apical keels crenulate with one projecting cell row, free margin entire; bract lobes oblong to semicircular, 264–296 × 136–176 μm; bract lobules semicircular to slightly oblong, 224–280 × 128–152 μm, antheridia 2 per bract; male bracteoles bilobed, 70–160 × 40–60 μm. Gynoecia terminal on lateral branches with 1 pycnolejeunoid innovation; female bracts elliptic to slightly falcate, apical keel entire, rarely crenulate with one projecting cell row; bract lobes elliptic to slightly falcate, 720–840 × 320–400 μm; bract lobules narrowly oblong, 640–720 × 104–144 μm; female bracteoles semicircular to slightly oblong, 60–130 × 80–100 μm, apex obtuse to emarginate. Perianths oblong, 1.26–1.5 mm long, 0.62–0.8 mm wide, with 3 keels; keels short, entire; beak 3 cells long. Sporophytes. Setae articulate, ca. 8 cells long; in transverse section with 4 inner and 12 outer cells. Capsules slightly spherical, 350–400 μm in diameter, splitting to ± 4/5 of its length into four erect valves after dehiscence; capsule valves 320–360 × 192–224 μm, wall 2–3 stratose, hyaline. Outer layer of capsule valve apex formed by 1 quadrate cell; cells of upper half of valve slightly rhombic, large, thin-walled; hinge butterfly-shaped, hinge cells rectangular to subquadrate, smaller than upper half cells, thin-walled; marginal cells rectangular, form 1 row. Inner layer of capsule valve cells in centre rectangular, walls with nodular thickenings; marginal cells of upper half quadrate to slightly rectangular, walls nodulose, formed by 2 rows, marginal cells of lower half rectangular, walls smooth; basal cell subquadrate, walls smooth. Elaters of 2 types: marginal and additional; marginal elaters 22 per capsule, each valve contains 4 or 5 upper marginal elaters and 2 or no lower marginal elaters; upper marginal elaters attached to upper half of valve margin by one end, 200–328 μm long, 14–18 μm wide, hyaline, with irregular thickenings, end attached to valve margin rounded, not expanded, free end dilated; lower marginal elaters attached to valve at both ends (near and parallel to basal margin), 200–250 μm long, 16–24 μm wide, hyaline, with irregular to annular thickenings, upper end narrowly tapered, lower end obtuse; additional elaters 4 per capsules (1 per each valve), 230–340 μm long, 6–10 μm wide, hyaline, with irregular thickenings, attached along their whole length to median part of valve, even upon dehiscence. Spores green, irregular-oblong or elongate-rectangular, 52–60 × 22–28 μm; spore surface covered with irregular lamellae; sporelings Lejeunea - type.

Habitat and Ecology: — Colura valida occurs on branches, bark and living leaves and was found in montane forest at elevation 850–1235 m.

Distribution: — Vietnam ( Jovet-Ast 1961, Shu et al. 2017), new to Malaysia and Thailand.

Selected specimens examined:— MALAYSIA. Pahang State: Genting Highlands, 1736–1745 m, 3 March 2013, Chantanaorrapint, Inuthai & Promma 1953 ( PSU). Pahang Stage: Cameron highland, Tanah Rata , rainforest near downtown, 04°28ʹ1ʺN, 101°22ʹ41ʺE, 1460 m, 16 February 2015, Sangrattanaprasert 276/15, 277/15D ( PSU) . THAILAND. Nakhon Si Thammarat: Ron Phiboon, Khao Ramrome, 08°14ʹ17.70ʺN, 99°48ʹ19.10ʺE, 930–1000 m, 25 September 2013, Sangrattanaprasert 13/13, 20/13, 21/13 B ( PSU) ; 29 July 2014, Sangrattanaprasert 188/14, 199/14 B, 200/14 B ( PSU) ; 25 February 2015, Sangrattanaprasert 326/15, 327/15 B ( PSU). Phatthalung: Kong Ra, Khao Loan , 850–1235 m, 27–28 May 2017, Chantanaorrapint & Suwanmala 953, 1139D ( PSU) .

Taxonomic notes: — Colura valida is characterised by large leaves (1.4–2.0 × 0.7–1.1 mm) without apical crest at lobule sac apex, lobule sac about 1/2 as long as lobule length, and large underleaf lobes with 12–16 cells long and 6–9 cells wide at base.

Figures

FIGURE 5. Colura valida Jovet-Ast: A. Portion of plant with gynoecium, dorsal view. B. Portion of plant, ventral view. C. Stem in transverse section. D–E. Lateral leaves, D. ventral view, E. dorsal view. F–G. Leaf lobe cells, F. median cells, G. marginal cells. H. Lobule sac apex. I. Valve (b = basal median cell, h = hinge cell, hy = hyaline papilla). J. Valve frame. K. Underleaf. L–M. Androecia, L. dorsal view, M. ventral view. N–O. Female bracts, N. dorsal view, O. ventral view. P–R. Female bracteoles. S. Perianth with young sporophyte.T. Spore.A, F–G, N–R. from Sangrattanaprasert 13/13 (PSU), B–E, H–M. from Sangrattanaprasert 326/15 (PSU), S–T from Sangrattanaprasert 21/13 (PSU).