Ctenidium Moss

Ctenidium malacodes

Ctenidium malacodes Mitt., J. Linn. Soc., Bot. 12: 509. 1869. Type: Colombia. Andes Bogotensis, in sylvis supra Pacho ad arbores, 6000 ped., J. Weir 243 (lectotype: NY!, designated by Nishimura 1985); Ecuador. Guarapata, Spruce 1053 (syntype: NY!).

Cupressina anacamptopteris Müll. Hal., Bull. Herb. Boiss. 6: 123. 1898. Ctenidium anacamptopteris (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1048. 1908. Type: Brazil. Serra Itatiaia, 2100 m, E. Ule 1882 (isotypes: H, US!).

Meteorium terrestre Müll. Hal., Hedwigia 40: 97. 1901. Ctenidium terrestre (Müll. Hal.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 1048. 1908. Type: Brazil. Santa Catharina: Serra Geral, E. Ule 1206 (isotype: H-BR).

Ctenidium muenchii Broth. ex H. A. Crum & Sharp, Bol. Soc. Bot. México 15: 15. 1953, nom. nud. in syn. BASED ON: Mexico. Chiapas: distr. San Cristobal, “los Llanos,” 2500 m, 23 Dec 1907, Münch 7433 (H, NY!).

Plants medium-sized, often in extensive mats, lustrous, soft, yellow-green to golden. Stems to 6 cm long, creeping to ascending, subpinnately to pinnately branched, the branches often unequal, not complanate-foliate; stems in cross section with 3–5 rows of small thick-walled cells, the outermost row sometimes with ± thin outer walls, surrounding larger thinner-walled cells, central strand of small thin-walled cells; pseudoparaphyllia foliose, large, serrulate; axillary hairs consisting of a single short hyaline basal cell, 1–2 long-rectangular hyaline intercalary cells and a single elongate hyaline distal cell. Stem and branch leaves differentiated, stem leaves 0.8–1.1 mm long, erect-to wide-spreading with flexuose apices, broadly ovate-triangular, abruptly acuminate, the acumen flexuose, sometimes twisted, somewhat concave, usually somewhat plicate, broadly decurrent; margins serrate to sharply serrulate almost to base, plane or occasionally erect; costa short and double, the two forks not united at base; cells linear to oblong-linear, sparsely prorulose at upper ends on abaxial surface, firm-walled, becoming oblong to rectangular in 1–3 rows across the insertion; alar cells extending into the decurrencies, subquadrate to short-rectangular in a sizable group, surrounding 4–8 larger, oblong, ± inflated cells often in a somewhat excavated area. Branch leaves (0.5–) 0.7–1.0(–1.1) mm long, wide-spreading, sometimes ± secund, lanceolate to ovate-lanceolate, gradually acuminate, the apex curved to subflexuose and occasionally twisted, concave, subplicate, short-decurrent; margins serrate to serrulate above, subentire to serrulate below, plane above, usually erect below; costa short and double, or often none; cells linear to oblong-linear, sparsely to densely prorulose at upper and sometimes lower ends at back, firm-walled, becoming rectangular in 1–2 rows across the insertion; alar cells few, only scarcely extending above the short decurrencies, subquadrate to oblong. Asexual propagula none. Dioicous. Perichaetial leaves 2.0–2.5 mm long, erect, oblong, abruptly slenderly long-acuminate, concave, subplicate; margins serrate in the acumen, coarsely serrate at the shoulder, serrulate to entire below, plane or rarely erect; costa none; cells oblong-linear, smooth, becoming laxly rectangular toward the insertion; alar cells not differentiated. Setae 1.6–2.3 cm long, elongate, smooth, reddish; capsules 1.5–2.0 mm long, inclined to horizontal, arcuate, asymmetric, ovoid to cylindrical; exothecial cells subquadrate, the vertical walls thick, the cross walls thin, often ± sigmoid; annulus broad, of 2–3 rows of elongate firm-walled cells, deciduous; operculum short conic-rostrate; exostome teeth shouldered, bordered, on the front surface cross-striolate below, with fine, overlying papillae, the papillae becoming denser at midtooth and coarser above, trabeculate at back; endostome with a high, finely papillose basal membrane, segments finely papillose, keeled, perforate, sometimes gaping with age, about as long as the teeth, cilia finely papillose, in groups of 2–3, short to rudimentary. Spores 13–17 μm in diameter, spherical, finely papillose. Calyptrae cucullate, appearing naked or with a few scattered hairs, smooth.

Distribution and ecology: Mexico to northern South America; controversially in eastern North America; growing typically on rocks but also soil and logs, in humid and often open forests, above 1500 m.

Illustrations in publications: Fig. 180G–J in Bartram (1949: 408); Pl. 130 in Buck (1998: 330); Fig. 776 in Nishimura and Ando (1994: 1047).

Discussion: Ctenidium malacodes is distinguished by its broadly ovate-triangular stem leaves that have large decurrencies of differentiated alar cells. The stem leaves are also abruptly acuminate and sharply serrate. The branch leaves are only shortly decurrent and gradually acuminate but usually more serrate than the stem leaves. The cells are prorulose on the abaxial surface of the leaves.

Ctenidium malacodes has been reported by Nishimura (1985) as occurring in eastern North America, where collections of the genus had previously been called C. molluscum (Hedw.) Mitt. Pedano (1996) suggested that this material is neither C. molluscum nor C. malacodes but rather a taxon intermediate between the two and worthy of species rank. Buck and Allen (2004) made the appropriate combination as C. subrectifolium (Brid.) W. R. Buck & B. H. Allen.

Selected specimens examined: MEXICO. Chiapas: San Cristoval, 2500 m, Münch 7433 (H, NY). México: Mpio. Ocuilán, Barranca de Mexicapa, 2200–2350 m, Buck 28148 (NY). Michoacán: 4.1 km W of Mil Cumbres, ˜2800 m, McGregor 16574 (NY). Oaxaca: 34.5 km above Valle Nacional in Sierra Juárez on Hwy. 175, Sharp et al. 4750-b (TENN). Veracruz: Paß an der Straße Orizaba–Tehuacan, 23 Mar 1979, Frahm s.n. (NY). GUATEMALA. Huehuetenango: Sierra de los Cuchumatanes, 2400 m, Steyermark 50030 (NY). San Marcos: Barranco Eminencia, 2500–2700 m, Standley 86365 (NY). Sololá: Volcán Santa Clara, 2100–3000 m, Steyermark 46954 (NY). Totonicapán: Above Totonicapán, 2800–3100 m, Standley 84472 (NY). COSTA RICA. Alajuela: La Palma de San Ramón, Brenes 113 (NY). PANAMA. Chiriquí: Boquete, Helion 812 (H, NY). JAMAICA. Blue Mountains, St. Helen’s to Morce’s Gap, Britton 1075 (NY). DOMINICAN REPUBLIC. La Estrelleta: Sierra de Neiba, ˜1700 m, Buck 4790 (NY). La Vega: Vicinity of La Lagunita, ˜2600 m, Norris et al. B5738 (H). Peravia: San José de Ocoa, La Nevera, ˜2100 m, Mejía & Zanoni 8721 (JBSD, NY). COLOMBIA. Boyacá: Mpio. Duitama, Páramo “La Rusia,” 3550 m, 06°N, 73°W, Ireland 23626 (CANM, US). Caldas: Cordillera Central, Magana, 3000–3300 m, Killip & Hazen 12098 (NY); Mpio. Río Sucio, on the Río Sucio, 1900–2080 m, ˜05°20'N, 75°15'S, Churchill & Arbeláez A. 15658 (US). Nariño: Mpio. Pasto, between km 13 and 15, Cocha Lagoon, 3100 m, 01°12'N, 77°12'W, Churchill & Arbeláez A. 15960-b (US). Norte de Santander: Cordillera Oriental, 2800–3000 m, Killip & Smith 19868 (NY). VENEZUELA. Trujillo: Distr. Urdaneta, S and above La Mesa de Esnujaque, 3100 m, Griffin III et al. 1377 (FLAS, NY, TNS). GUYANA. Mt. Roraima, Quelch & McConnell 340 (H). ECUADOR. Andes Quitenses, ˜1800 m, Spruce 1051 (H-SOL); Timgiragia. 2700−3050 m, Spruce s.n. (NY). PERU. Convención: Dept. Cuzco, 3200 m, Bües 1468 (NY). BOLIVIA. Santa Cruz: Vallegrande, Calaculo, ˜26 km S of Vallegrande, road to Khasamonte, 2450 m, 18°38'S, 64°02'W, Churchill & Arroyo P. 21200 (US); Tablas, ˜3400 m, Herzog 2817 (JE); Río Tocorani, ˜2200 m, Herzog 4082 (H, JE). BRAZIL. Santa Catarina: Serra Geral, Ule 1157 (H).