Cheilolejeunea rigidula (Montagne) Schuster (1971: 105)

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 : 106-109

26. Cheilolejeunea rigidula (Montagne) Schuster (1971: 105) .—

Lejeunea rigidula Nees ex Montagne (1840: 336) .

Type:— FRENCH GUIANA. Mont Serpent, Leprieur 257 (PC-Mont. No. 3655 lectotype designated by Grolle in sched. 1979?).— Fig. 17A–J .

Cheilolejeunea polyantha Evans (1902: 141) , syn. fide Bastos & Gradstein (2020)

Cheilolejeunea serpentina (Mitten 1961: 112) Mizutani (1963: 171) , syn. fide Bastos (2012a)

Plants yellowish green to light green, 0.9–2.0 mm wide. Stems 60–80 µm in diameter, 7(–10) epidermal cells in cross section, 12–30 × 8–21 µm; 10–11 medullary cells, 5–17 × 3–12 µm. Ventral merophytes 2 cells wide. Leaves imbricate, ovate-orbicular, flat, 260–380 × 340–430 µm, margin entire, apex rounded; trigones inconspicuous to conspicuous; basal cells 10–30 × 11–19 µm, median cells 12–24 × 8–18 µm, marginal cells 7–18 × 5–15 µm. Lobules ovoid, inflated, 1/4–1/3 of leaf length, free margin involute, apical tooth oblong and obtuse. Underleaves distant to subimbricate, orbicular, 135–270 × 170–320 µm, 2–3.5 × stem width, bifid to 1/4–1/3, with a V-shaped sinus, base cuneate to rounded. Dioicous. Androecia on short-specialized branches or intercalary on long shoots, 2–6 pairs of bracts globose. Gynoecia with lejeuneoid innovations, bracts spathulate, 490 × 330 µm, apex rounded; bracteole 380 × 320. Perianth with 4 strong keels, beak short. Vegetative reproduction unknown.

Distribution and habitat:—Pantropical, from southern United States to northern Argentina, including the Caribbean islands ( Reiner-Drehwald 1998), and Africa ( Bastos 2017). This species is widely distributed in the study area, growing on tree trunks, humus, and rocks in the coastal desert and semi-desert areas, inter-Andean deserts and semi-deserts, savanna, deciduous forests, lowland rainforest, lower and upper montane rainforests, from sea level to 2800 m, in the Puntarenas-Chiriquí, Guatuso-Talamanca, Guajira, Venezuelan, Sabana, Magdalena, Cauca, Chocó-Darién, Galápagos Islands (Gradstein 2021), and Western Ecuador provinces of the Pacific dominion ( Fig. 18 ). Cheilolejeunea rigidula is the most common liverwort species in the Amazon basin ( Campos et al. 2019). The records of C. rigidula that apparently are from the Páramo ( Fig. 18 ), actually belong to the Cauca or Magdalena provinces.

Notes:—Characterized by having leaves orbicular, flat and apex rounded, underleaves 2–3.5 × the width of the stem, generally distant, dioicous sexuality, and with lejeuneoid innovations.

According to Bastos (2017) and Reiner-Drewald (1998) the lectotype was selected by R. Grolle “in sched.”, and there is an isosyntype, also designated by R. Grolle, in NY (NY-00920062).

Specimens examined:— BRAZIL. Amazonas: S „ o Gabriel da Cachoeira , 0°24’S, 66°23’W, 130 m, 5 July 1979, Yano 1713 ( RB) . Rio de Janeiro: PARNA Tijuca, 22°56’51”S, 43°17’30”W, 607 m, 19 April 2006, Santos 443 ( RB); Arboreto do Jardim Botânico do Rio de Janeiro, 15 December 2000, Costa 186 ( RB); Santa Maria Madalena, Muribeca do Imbé , 21°53’73”S, 41°54’67”W, 582 m, 16 May 2017, Costa 1790 ( RB) . COLOMBIA. Amazonas: Comunidad “Peña Roja”, 0°34’S, 72°08’W, 200 m, 8 July 2000, Idárraga 2338, 2349, 2353 ( HUA); La Pedrera , 1°18’S, 69°36’N, 325 m, 9 March 1990, Churchill 16090-b ( HUA) . Antioquia: Anori, El Retiro, 1500 m, 30 June 2016, Farfán C1423 ( COL); Buriticá , vereda Higragra , 6°41’59.5”N, 75°53’30.6”O, 1067 m, Carmona 454 ( HUA); Caracolí, 700 m, July 2017, Soto, s.n, ( CUVC); Corcorna, 6°2’23.43”N, 75°7’1574”W, 830 m, 19 August 2016, Ordoñez 241 ( HUA); Guatapé, vereda Santa Rita, 6°14’N, 75°10’W, 1850 m, 14 June 1997, Gutiérrez 1228 ( HUA); Ituango, vereda Organí Medio, 7°15’3.540”N, 75°27’26.447”W, 197 m, 28 September 2015, Trujillo 7330 ( HUA); Medellín, corregimiento Santa Elena, 6°16’27”N, 75°30’23”W, 2446 m, Londoño 133 ( HUA); Urrao, vereda La Clara, 894 m, 19 July 2016, Morales 1065 ( HUA). Boyacá: Paez, vereda Yamonta, 1345 m, 2017, Bravo 980, 981 ( JBB); Santa María , vereda Caño Negro , 4°50’58”N, 73°16’53”W, 850 m, 7 October 2000, Aguirre 12898 ( COL). Cauca: Morales, vereda Santa Barbara, 2°48’8.4”N, 76°43’10.5”W, August 2015, Gómez 556 ( HUA). Chocó: Unguía, Cerro Tacarcuna, 1113 m, 4 April 2017, Gil-Novoa 3431 ( UPTC); Nuquí , Amargal , 30 m, 6 August 1992, Gradstein, 8864 ( COL). Córdoba: Montelíbano, May 2015, López 183 ( HUA); Puerto Libertador, vereda Las Claras , 7°50’26.1”N, 75°41’53.5”O, 84 m, 5 April 2017, Pérez 519 ( HUA). Guaviare: San José del Guaviare, 2°34’13.9’’ N, 72°53’05.9’’ W, 301 m, 4 December 2021, Gil-Novoa & Costa 3657, 3658 ( UPTC). Huila: Acevedo , Macizo colombiano, 1°36’59” N, 76°6’15” W, 2100 m, 26 November 2011, Castillo 2358 I ( COL). Magdalena: Santa Marta , cuchilla de San Lorenzo, 1800 m, 17 January 1967, Winkler C173 ( COL). Meta: Villavicencio, Reserva Forestal Buena Vista, 4°9’09.1”N, 73°39’11.3”W, 645 m, April 2017, Medina 1422 ( LLANOS). Nariño: Mosquera , 0 m, 9 November 1992, Linares 3b ( COL). Quindío: Circacia, 4°35’N, 75°42’W, 1400 m, 15 April 2000, Orrego 337 ( COL). Risaralda: west side of cordillera Occidental , 800 m, 29 July 1992, Gradstein 8639 ( COL). Santander: Floridablanca, 7°70’14.8”N, 73°04’7.6”W, 1620–1871 m, 23 June 2014, Uribe 4315, 4322 ( COL); Puerto Parra , 126 m, 2014, Castaño T3A2C1M2 ( UPTC) . Tolima: Santa Isabel, 2960 m, 28 July 1980, Aguirre 1583 ( COL); Venadillo, vereda Planadas , 1150 m, 11 August 1980, van Reen 2482 ( COL) . COSTA RICA. San José: Pacific slope, 2000 feet, 4 October 1947, Svilha 47-736 ( NY) . ECUADOR. Esmeraldas: Nr. Lita, 500 m, 20 August 2014, Wilson 04-20 ( QCA) . PANAMÁ. Darien: Pinogana, 50 m, Salazar-A. 9200 (PMA) . Panamá: Barro Colorado, N . Salazar-A. 6079 ( PMA); Panamá City , 22 April 1991, Gradstein 7886 ( HUA, RB) . SURINAM. Nickerie: Bekker 1295ab ( PMA) . VENEZUELA. Amazonas: near mouth of Cañon Grande, 140 m, 00°50’N, 66°10’W, January-February 1985, Halling s.n. ( NY) . Merida: Distr. Campo Elias , 28 January 1976, Schuster 76-798 ( F) .

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 18. Distribution of Cheilolejeunea rigidula and C. schiavoneana in Pacific dominion.