Cheilolejeunea trifaria (Reinwardt et al.) Mizutani (1964: 132) var. trifaria

Gil-Novoa, Jorge Enrique & Costa, Denise Pinheiro, 2023, Synopsis of the species of Cheilolejeunea (Marchantiophyta, Lejeuneaceae) in the Pacific dominion and Páramo province of tropical America, Phytotaxa 587 (2), pp. 73-120 : 110-111

29.1 Cheilolejeunea trifaria (Reinwardt et al.) Mizutani (1964: 132) var. trifaria . —

Jungermannia trifaria Reinwardt et al. (1824: 226) .

Type:— INDONESIA. Java, Blume s.n. (holotype L).— Fig. 17Y –HH.

Plants yellowish green to light brown, 0.8–1.4 mm wide. Stems in cross section 90–120 µm in diameter, (7–)8 epidermal cells in cross section, (14)21–35 × 10–22 µm; 10 medullary cells, 10–23 × 6–12 µm. Ventral merophytes 2 cells wide. Leaves imbricate, ovate-orbicular, flat, 500–980 × 350–700 µm, margin entire and apex rounded; dorsal and ventral margin curved; cells generally mammillose on the dorsal side, conspicuous trigones; basal cells 13–32 × 11–25 µm, median cells 14–34 × 12–24 µm, marginal cells 6–18 × 5–16 µm. Lobules trapezoid, 1/4–1/3 of leaf length, tooth acute 17 × 9 µm, free margin involute, tooth acute, keel arched. Underleaves imbricate, orbicular to broadly ovate, 300–650 × 350–780 µm, 4.5–6.5 × stem width, bifid to 1/3, with a V-shaped sinus, margin entire, base auriculate. Autoicous. Androecia on the main stem, with 3–5 pairs of bracts. Gynoecia with pycnolejeuneoid innovations, bracts obovate, 700 × 550 µm, apex acute. Perianth 5-keeled, keels weak to very strong, beak short. Vegetative reproduction unknown.

Distribution and habitat:—Pantropical, occurring from Mexico to Bolivia and Brazil, furthermore in the Antilles, tropical Africa, Southeast Asia and Oceania ( Bastos 2017, Gradstein 2021, Singh 2012). This species is very common and widely distributed in the study area, growing on tree bark, rocks, and decaying wood, in savannas, deciduous forests, and lowland and lower montane rainforests, from sea level up to 2400 m, in the Puntarenas-Chiriquí, GuatusoTalamanca, Guajira, Venezuelan, Magdalena, Cauca, Chocó-Darién, Galápagos Islands, and Trinidad provinces of the Pacific dominion. The records of C. trifaria var. trifaria occurring in the Páramo ( Fig. 19 ), actually belongs to the Cauca or Magdalena provinces.

Notes:—Characterized by autoicous sexuality, ovate-orbicular leaves with mammillose cells, ventral merophyte 2 cells wide, and imbricate, usually wider than long underleaves with auriculate bases.

Specimens examined:— BOLIVIA. Santa Cruz: Porongo , 17°54’22”S, 63°28’13”W, 1100 m, 16 January 2008, Linneo 1351 ( RB) . BRAZIL. Acre: Tarauacá , 15 November 1995, Costa 2535 ( RB) . Rio de Janeiro: Santa Maria Madalena , 21°57’54”S, 41°51’28”W, 125 m, 16 May 2007, Santos 859 ( RB) . São Paulo: Praia Grande , 24°0’57”S, 46°36’54”W, 20 m, 24 August 2016, Prudêncio 46 ( RB) . COLOMBIA. Antioquia: Cañas Gordas , 6°48’48.7”N, 76°5’17.6”W, 2000–2400 m, January 2015, Ospina 109 ( HUA) ; Cocorná , 6°2’41.8”N, 75°7’49.2”W, 975 m, 19 August 2016, Ordoñez 552 ( HUA) ; Ebéjico , El Palón, 6°9’54,34”N, 75°40’0,56”W, 1150 m, March 2017, Buitrago 47 ( UDBC) ; Guatapé , vereda de Santa Rita, 6°14’N, 75°10’W, 1850 m, 13 June 1997, Gutierrez 1210 ( HUA) ; Ituango , Corregimiento El Aro, 7°14’42”N, 75°30’44.2”W, 195 m, 26 August 2015, Trujillo 7306 ( HUA) ; Jardín , 1850 m, 12 May 1986, Escobar 799 ( HUA) ; San Roque, vereda Guacas Abajo , 17 October 2018, López 245 ( HUA) ; Támesis , 5°41’34.7”N, 75°42’29.5”W, 1376 m, Ordoñez 153 ( HUA) ; Urrao , vereda La Sexta, 6°23’36.6”N, 76°15’19”, 1139 m, 15 March 2019, Ordoñez 1123 ( HUA) . Boyacá: Santa María , La Cristalina, 4°51’33.1”N, 73°15’56.5”W, 825 m, 29 October 2013, Amortegui 8 ( UDBC) . Casanare: Sácama , camino a la vereda “Africa”, 1400 m, 16 August 1982, Aguirre et al. 2943 ( COL) . Chocó: Unguia, cerro Tacarcuna 1115 m, 2 April 2017, Gil-Novoa s.n. ( UPTC) . Cundinamarca: Guaduas , 1388 m, 1 September 2017, Malagón 851 ( COL) . Huila: Acevedo, Macizo colombiano, 1°36’59”N, 76°6’15”W, 2100 m, 26 November 2011, Castillo 2046p ( COL) . Meta: Acacías, vereda Frescovatele , 03°57’51.3’’ N, 73°49’22.4’’ W, 853 m, 3 December 2021, Gil-Novoa & Costa 3635, 3646 ( UPTC) ; Villavicencio, Reserva Forestal Buena Vista , 4°9’10.2”N, 73°39’19”N, W, 655 m, April 2017, Medina 1428 ( LLANOS) . Nariño: Mosquera , corregimiento La Vigia, 0 m, 10 November 1982, Linares 17 ( COL) . Putumayo: Orito , 300 m, 30 August 2015, Ruiz, 1246 ( COL) ; San Francisco , vereda La Esperanza, 01°12’21.2”N, 76°47’31.9”W, 1860 m, 28 September 2016, Solarte 441 ( COL) ; Valle del Guamuez , 0°23’41.4”N, 76°43’55”W, 290 m, 4 November 2016, Agudelo 16 ( UDBC) . Quindio: Circasia, finca San Agustin , 4°38’N, 75°39’W, 1700 m, Orrego 336 ( COL) . Risaralda: vereda La Colina, 4°56’N, 75°42’W, 1725 m, 19 August 1986, 1725 m, Wolf 1195 ( COL) . Santander: Hato , 6°35’31.3”N, 73°21’9.8”W, 2053 m, 9 August 2018, Lozano 236 ( UPTC) ; Tona , 7°10’29.2”N, 73°02’7.8”W, 1675–1715 m, 22 June 2004, Uribe 4275, 8284 ( COL) . COSTA RICA. San Domingo de Golfo Dulce , March 1896, Tonduz 15630 ( NY) . San José: Pacific slope, 2000 feet, 4 October 1947, Svilha 47-719 ( NY) . ECUADOR. Cotopaxi: Quevedo , 1°02’S, 79°16’W, 700 m, 27 October 1999, Andersson 3103 ( QCA) . Galápagos: Floreana , 550 m, 26 April 1976, Gradstein H177 ( COL, QCA) . Guayas: Camacho , 2°48,6’S, 79°38,9’W, 100 m, 15 March 2012, Schafer-Verwimp 32910 ( QCA) . Morona-Santiago: Macas , 2°07’S, 77°46,0’W, 650 m, 27 March 2012, Schafer-Verwimp 33212/ A ( QCA) . Orellana: Francisco de Orellana , 0°40’S, 76°24’W, 220–250 m, Schafer-Verwimp 32825 ( QCA) . FRENCH GUIANA. Saül . 150 m, 23 June 1986, Gradstein 6177 ( COL, HUA, PMA, RB) . PANAMA. Panama: Capira, Parque Nacional Altos de Campana , 850 m, Salazar-A. 9454p.p.3 ( PMA) . PUERTO RICO. Maricao: Maricao State Forest , 600–700 m, Gradstein 6530 ( PMA) . VENEZUELA. Amazonas: Rio Negro , 1500 m, 1°32’36”N, 66°50’W, 16 October 1987, Halling 5544 ( NY) . Lara: Cuenca del Río Gyamure , Griffin III, 239 ( PMA).

Figures

FIGURE 17. Cheilolejeunea rigidula A-B. Cladographs of fertile plants (solid ellipse = androecia, open ellipse = gynoecia with perianth, U= gynoecia without perianth); C. Shoot with androecia in ventral view; D. Perianth with bracts and bracteole; E. Shoot in ventral view; F. Ventral merophyte with underleaves; G. Leaf and lobule; H. Lobule; I. Underleaf; J. Leaf lobe cells with trigones. Cheilolejeunea schiavoneana K. Shoot in ventral view; L. Shoot in ventral view with caducous leaves; M. Ventral merophyte; N. Leaf and lobule; O. Lobule; P. Underleaf; Q. Leaf lobe cells with trigones. Cheilolejeunea tonduzana R. Cladograph of fertile plants (solid ellipse = androecia, open ellipse = gynoecia with perianth, U= gynoecia without perianth); S. Shoot with perianth in ventral view; T. Ventral merophyte; U. Leaf and lobule; V. Lobule; W. Underleaf; X. Leaf lobe cells with trigones. Cheilolejeunea trifaria var. trifaria Y-Z. Cladographs of fertile plants (solid ellipse = androecia, open ellipse = gynoecia with perianth, U= gynoecia without perianth); AA. Shoot with perianths and androecia in ventral view BB. Perianth in ventral view; CC. Shoot with androecium in ventral view; DD. Ventral merophyte; EE. Leaf and lobule; FF. Lobule; GG. Underleaf; HH. Leaf lobe cells with trigones.

FIGURE 19. Distribution of Cheilolejeunea tonduzana and C. trifaria var. trifaria in the Pacific dominion and the Páramo province.