Bryophytes (hornworts, liverworts and mosses) of the British Indian Ocean Territory

Plants small, pale-greenish, in low turfs and clumps. Stems short, forked; rhizoids reddish brown. Leaves dimorphic, curled-contorted when dry, straight when wet, vegetative leaves oblong-lanceolate, gemmiferous leaves acuminate distally with often spatulate tips, 2.5–4.0 mm long, axillary hairs inconspicuous; costa percurrent, often long-extended beyond broad laminae and with narrow wings of laminae in gemmiferous leaves; cells of upper laminae isodiametric, obscure, smooth to unipapillose abaxially, mammillose-papillose adaxially; margins mostly thickened, sometimes unistratose, entire except sometimes irregularly serrate at shoulders; teniolae lacking; cancellinae usually broadly to narrowly acute distally. Gemmae common, borne on adaxial surface of costa, often in distinct saucer-shaped gemma receptacles on tips of modified leaves. Sporophytes not seen. less ↑

Plants small, pale, compact, in low turfs. Stems short, forked; rhizoids brown. Leaves only somewhat dimorphic, 2–3 mm long, not much contorted when dry, usually secund, straight and erect-spreading when wet, vegetative leaves oblong, gemmiferous leaves oblong or acuminate, axillary hairs inconspicuous; costa excurrent in gemmiferous leaves; cells of upper laminae isodiametric, smooth to unipapillose abaxially, mammillose-papillose adaxially; margins unistratose or slightly thickened, entire throughout; teniolae lacking; cancellinae narrow, typically truncate distally. Gemmae common, in pale spherical clusters all around on tip of excurrent costa. Sporophytes not seen. less ↑

Plants small, 5–10 mm high, grayish green, somewhat glossy, in loose or dense tufts and cushions. Stems short, sparsely branched; central strand absent. Leaves erect-patent to curved spreading, 4–6 mm long, flattened, strap-shaped to ligulate from an oblong-ovate base, apiculate at the apex; hyaline lamina cells small, appearing only on two sides of costa at the leaf base; margins entire, with minute serration at the leaf apex; costa broad and thick, filling almost the entire upper lamina, convex on abaxial side in cross section, costal leucocysts 2–5 layers on adaxial side, 1–4 layers on abaxial side, supporting a nearly centrally positioned chlorocyst layer, chlorocysts in cross section triangular in the upper parts of leaves, quadrangular at the leaf base. Autoicous. Setae ca. 4 mm long, yellow, smooth; capsules erect, cylindrical; peristome teeth 8, yellowish, lanceolate with a broad base, vertically striate; opercula long rostrate, with an oblique beak. Calyptrae cucullate, smooth, entire at the base. Spores not seen. less ↑

Plants small to medium-sized, pale glossy, whitish green, very compact, in dense turfs. Stems elongate, much-forked, rhizoids bright-red, sometimes conspicuous. Leaves straight and appressed wet and dry, 1.5–1.8 mm long, involute, lance-acuminate, composed mostly of cancellinae, with only small areas of green cells distally on both sides of costa, axillary hairs inconspicuous; green cells of upper laminae isodiametric, smooth abaxially, bulging to unipapillose adaxially; margins bordered all around with elongate hyaline cells, entire, more or less revolute distally along the green cells; cancellinae acute distally. Gemmae infrequent, inconspicuous, adaxial on tip of costa. Sporophytes frequent. Setae 5–7 mm long; capsules short-cylindric to somewhat urceolate, sulcate with age, 0.8–1.0 mm long, with few stomata at base; opercula long rostrate; peristome of low, blunt, imperfect teeth, to 60 µm tall, teeth narrowly segmented, irregularly patterned-papillose inside and outside in distal segments. Calyptrae 1.8 mm long, papillose-roughened distally. Spores green, 19–24 µm in diameter, finely papillose. less ↑

Taxiphyllum taxirameum

Taxiphyllum taxirameum (Mitt.) M. Fleisch., Musci Buitenzorg 4: 1435. 1923. Stereodon taxirameus Mitt., J. Linn. Soc., Bot., Suppl. 1: 105. 1859; Isopterygium taxirameum (Mitt.) A. Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1876–77: 439. 1878. Type: In[dia], Himalayae reg. temp., Simla et Kumaon, Thomson 1023 (lectotype: NY!, designated by Iwatsuki and Crosby (1979: 393); 1023b cited by Mitten in the type description is apparently an error; Ceylon, Gardner s.n. (syntype: NY!).

Isopterygium planissimum Mitt., J. Linn. Soc., Bot. 12: 498. 1869; Taxiphyllum planissimum (Mitt.) Broth. in Engl., Nat. Pflanzenfam. ed. 2, 11: 463. 1925; Plagiothecium planissimum (Mitt.) E. B. Bartram, Bryologist 49(4): 122. 1946. Type. Jamaica, Wilson 675 (lectotype: NY!, here designated).

Isopterygium cavernicola Cardot, Rev. Bryol. 37: 56. 1910; Taxiphyllum cavernicola (Cardot) Thér., Rev. Bryol. Lichénol. 5: 109. 1932. Type: Mexico, Morelos, near Cuernavaca, Pringle, Plantae Mexicanae 15179 (isotypes: H!, NY!).

Plants in thin to dense mats, glossy, dark green to yellow-green. Stems to 5 cm long, 2–4 mm wide, complanate, simple or irregularly branched. Leaves 1–2 mm long, 0.3–0.6 mm wide, rigid, usually distant, wide-spreading, flat or somewhat concave, smooth or plicate, ovate- to oblong-lanceolate, symmetric or nearly so, acuminate or abruptly narrowed to an acute or rarely subobtuse apex; margins very narrowly recurved almost to apex, serrulate to serrate throughout; costa lacking or short and double; median cells 66–120 × 3–7 µm, smooth or prorulose at upper ends on dorsal leaf surface; alar cells poorly differentiated, long- to short-rectangular or rarely quadrate, in 1–3 rows with 1–5 cells in the marginal rows. Sex organs and sporophytes not seen on Latin American plants. Reported and observed to be dioicous; perigonia small, bracts ovate, acute; perichaetia small, bracts lanceolate to ovate-lanceolate, acuminate. Setae 0.7–1.2 cm long; capsules 1.0–1.5 mm long, reddish brown. Spores 11–13 µm in diameter.

Distribution and ecology: Common; West Indies, Mexico to Costa Rica, Colombia, Surinam, Ecuador, Brazil, and Bolivia (Figure 37); also known from eastern North America and eastern Asia; usually on moist, shaded limestone rocks and cliffs in woods, sometimes on soil banks beside streams and along roadsides, rarely on logs and bases of trees, at 40–2500 m.

Discussion: This is a distinctive species recognized by the dark green to yellow-green, rigid, complanate-foliate plants, the distant, wide-spreading, flat or somewhat concave, ovate- to oblong-lanceolate, acuminate or acute to rarely subobtuse, sometimes plicate leaves with very narrowly recurved margins, and the poorly differentiated alar regions of 1–3 rows of quadrate to rectangular cells with 1–5 cells in the marginal rows. It is the most common species of Taxiphyllum in Latin America, and it is especially prevalent in parts of Mexico and the West Indies.

Selected specimens examined: MEXICO. Chiapas: Rancho Monte Armenio, Ejido Chintul, Tecpatán, 300 m, Zamudio & Rodríguez 1499 (MEXU); Santa Elena, 650 m, 16°53'N, 91°35'S, Delgadillo M. 6324 (MO); above Paval N of Mapastepec, 1524 m, Sharp 4443 (MICH, TENN). Chihuahua: At bridge Puerto de Río Urique, 61 km S of Creel, 1707 m, Bowers et al. 5356 (CANM, MEXU, TENN), 5330-b, 5332-c, 5362b (TENN). Durango: Near Palos Colorados, W of Durango, 2499 m, Sharp 1499c (TENN). Hidalgo: 82.2 km SW of Tempoal on Molango Hwy., Spessard 696a (TENN); ˜26 km S of Tamazunchale, Bowers et al. 5049-a (TENN). Jalisco: Puente San Pedro, 8 km SW of Tecalitlán, ˜1200 m, 22 Sep 1958, McVaugh s.n. (MICH, NY); Arroyo de La Resolana, near Sapotillo, 305 m, Crum 643 (MICH, NY, TENN); near summit of La Cumbre, 16 km SW of Autlán, 1524 m, Crum 1345 (MICH, TENN, US); 27 km W of Ameca on road to Los Volcanes,

1500 m, 20°34'N, 104°15'W, Whittemore 2107 (MO). México: Hacienda Lanzarote, município de Villa Nicolás Romero, 2350 m, Cárdenas 3820a (MEXU). Michoacán: Morelos Natl. Park, near Morelia, Norris & Taranto 15528 (CANM, TENN); Cascade de Caradacuan in Río Cupatzio, Patrick 615 (FH). Nayarit: ˜6 km from Compostela on road to San Pedro Jagunilla, Norris & Taranto 15079 (CANM, TENN); ˜3 km W of Compostela on road to Puerto Vallarta, Norris & Taranto 15161(CANM, TENN); 50 km E of Ruiz, 10.7 km E of San Pedro Ixcatán on road to Mesa de Nayar, 109 m, 22°02'N, 104°50'W, Whittemore 3227 (MO). Nuevo Léon: Cañon de San Francisco near Monterrey, Spessard 95 (CANM, TENN); Río Valle Alto, Spessard 436, 441 (TENN); Chipinque, Sierra Madre

Oriental, ˜14 km NW of Monterrey, 1158 m, Hermann 26039 (CANM), 26039b (US); Sierra Madre, near Monterrey, Pringle 15103 (NY). Oaxaca: Sierra Juárez above Río Valle Nacional, 20 Dec 1962, Robinson s.n. (MEXU, NY, US); below Tamazulapam Mixe, ˜70 km E of Oaxaca, Smith et al. 2994 (CANM, MEXU, TENN), 3001 (CANM, TENN). Puebla: Below Zacapoaxtla, near Xalacapa, 1524 m, Sharp 4240 (CANM, MICH, TENN); vicinity of Puebla, 2200 m, Arsène 584 (US). San Luis Potosí: W of Xilitia, 838 m, Sharp 5915 (MEXU, MICH, TENN, US); above Xilitla, 762 m, Sharp 5856 (MEXU, MICH, TENN, US); just E of San Francisco, 45 km E of San Luis Potosí, 2164 m, Magill 2609 (TENN); 49 km S of Antiguo Morelos, road to Huizache, Pursell 5367 (MO). Sinaloa: 45 km NE of Mazatlan, 152 m, Bowers et al. 5127 (CANM, F, MEXU, TENN). Tamaulipas: 14 km above San Antonio on Hwy. 101, 1773 m, Sharp et al. 1002-b (TENN); ˜16 km from Victoria on road to San Luis Potosí, 914 m, Smith & Sharp 112, 114 (CANM, TENN), 116 (TENN); ˜16 km N of Villa Aldama, Pursell 5458 (MO); South Mt., near Rancho del Cielo, Gómez Farias, 1037 m, 23°15'N, 99°12'W, Manuel 396 (MO); roadside of Hwy. 101, 37 km SW of Victoria, 1098 m, Bowers et al. 2776 (NY); Paso de las Calabazas, along Río Sabinas, near Gómez Farias, Sharp 8763 (CANM); just E of San Francisco, 45 km E of San Luis Potosí, 2164 m, Magill 2609 (TENN). Tlaxcala: Near falls of Río Zuahuapan, near Amaxac, Sharp 401 (MEXU, TENN). Veracruz: Sierra de Chiconquiaco, between Santa Rita and Bella Luz, ˜40 km NW of Xalapa, 1750 m, Hermann 28844 (CANM, TENN); above Poza Rica on Hwy. 130, 1000 m, Sharp et al. 1280 (CANM, TENN), 4858 (CANM, F, MEXU, TENN); El Esquilón, ˜1300 m, 19°37'N, 96°56'W, Buck 35518 (NY); La Concha, along Carretera La Concha a San Antonio, ˜1000 m, 19°36'N, 96°53'W, Buck 35406, 35416 (NY); just S of city of Xalapa on road to Coatepec, Instituto de Ecología, Rancho Guadeloupe, ˜1300 m, 19°31'N, 96°56'W, Buck 35380 (NY). Zacatacas: 7 km S of Laguna Grande, N of Monte Escobedo, 2030 m, Cárdenas 3057b (MEXU), 3028 (MEXU, NY); 8 km E of Monte Escobedo, 2030 m, Cárdenas 3070 (MEXU, NY). GUATEMALA. Alta Verapaz: Cerro Chinajá, between Finca Yalpemech and Chinajá, 150–700 m, Steyermark 45564 (F, FH); 3 km N of Coban, 1341 m, Sharp 2994 (FH). Baja Verapaz: Patal, 1615 m, Sharp 2827 (FH, TENN). Chimaltenango: Quisaché, 1800 m, Standley 62310 (F, FH). Quezaltenango: Above Santa María de Jesús, ˜1650 m, Standley 67279 (F, FH). Retalhuleu: Near Chivolandia (Dept. Quezaltenango) along road to San Felipe, ˜650 m, Standley 87209a (F), 87213 (DUKE, F, FH, MICH, NY). Sacatepéquez: Above Pastores, ˜1680 m, Standley 60768 (F, FH, MICH, NY, US). Zacapa: Trail between Río Hondo and waterfall, 250–400 m, Steyermark 29413 (F, FH, MICH). BELIZE. Cayo: S side of Rapaculo Branch, 16°49'N, 88°56'W, Allen 18458, 18474 (MO). Toledo: Solomon Camp, vicinity of junction of Richardson creek and Bladen Branch, foothills of the Maya Mts., 80–420 m, Davidse & Brant 32073B (MO); Upper Bladen Branch, Bladen Nature Reserve, 16°54'N, 88°55'W, Whittemore 5800 (MO, NY); Gabriel Camp-Edwards Central Camp, Boutin & Schlosser 5155 (MO); Columbia River Forest Reserve, 16°22'N, 89°07'W, Allen 18678 (MO). HONDURAS. Comayagua: Vicinity of Siguateqeque, ˜1050 m, Standley & Chacón 6599 (F); Quebrada Agua Helado, above San José del los Planes, ˜15 km E of Lago Yojoa, cerro Meámbar, 14°47'N, 87°51'W, Allen 13717 (MO). Cortés: Along Río Lindo, N of Lake Yojoa, 500–600 m, Morton 7875, 7876 (US); near Jaral, 600 m, Standley 7021 (F); Nacimiento del Río Lindo, near Jaral, ˜600 m, Standley 7075 (F). El Paraíso: Vicinity of Danlí, 700–800 m, Standley 16394 (F). Olancho: Slopes of Montana Pena Blanca, 11.5 km S of Gualaco on road from San Francisco de La Paz to Gualaco, 980–1000 m, 14°57'N, 86°08'W, Allen 12405, 12426 (MO). EL SALVADOR. Ahuachapán, near Ataco, 1200–1400 m, Standley & Padilla 2689, 2694, 2701 (F). NICARAGUA. Jinotega: Between Las Camelias and La Salvadora, along tributary of Río Jiguina, ˜13°05'–06'N, 85°53'–54'W, Stevens & Grijalva 15359 (MO). COSTA RICA. Cartago: Río Macho, 1100 m, Morales et al. 963 (USJ). Puntarenas: ˜2 km SE of Monteverde, 1500–1550 m, Gentry & Burger 2755D (MO). Aserri, 1400 m, Alfaro 70b (FH). BERMUDA. Tucker’s Town, Church Cave, Britton 418 (NY). CUBA. Guantánamo: Loma Lagunato, ESE of El Ramón, 300–400 m, Reyes 9193B (NY). Oriente: Loma del Gato, Hioram 12044 (F). Pinar del Río: Sierra la Buira, Leon 4710 (NY). Santo Domingo: Barrabos, 400 m, 1887, von Eggers s.n. (H). Santiago de Cuba: La Gran Piedra, 1000 m, Buck 7609, Shaw 5098, 5153 (NY). Seibo: Azui, Taylor 278b (NY). Sierra Maestra: “El Peru,” in the valley of the Río Yara, Pócs 9073c (NY). Trinidad Mtns., Santa Clara, León 14149 (NY). JAMAICA. Manchester: Marshall’s Pen, 5 km WNN of Mandeville, 701 m, Crosby 13825 (NY); 4 km N of Plowden Hill, 500 m, Crosby 13848 (MO, NY). St. Andrew: Newcastle road, between 16 & 17 mile posts, 762–1069 m, Farr 104 (CANM). St. Catherine: Byndloss Gully, 1.6 km N of Linstead, 91 m, Farr 106 (CANM). St. Thomas: Corn Puss Gap and vicinity, ˜14 km N of Bath, 550 m, Buck 5561 (NY); Bath, along Sulphur River, 91 m, Crosby 13631 (MO, NY). Trelawny: Vicinity of Windsor Cave, 140 m, Buck 5932 (NY); Cockpit Country, just N of Wilson’s Run, 9 km N of bridge at Troy, 500 m, Crosby 13879 (MO, NY). HAITI. Vicinity of St. Louis du Nord, Morne Colombot, Leonard & Leonard 14249 (NY, US), 14319 (F, FH, NY, US). Vicinity of Port au Prince, Pétionville, Leonard & Leonard 15792 (FH, NY, US). Nord: vicinity of Marmelade, 800 m, Leonard 8139 (CANM, FH, NY, US). Sud’est: Massif de la Selle, 19 km S of Kenscoff, 1600 m, Buck 9015 (NY). Vicinity of Bombardopolis, 610 m, Leonard & Leonard 13560 (FH, US). Les Trois Sources, between Chambellan and Cayaba, 442 m, Bartlett 17792 (DUKE, MICH). DOMINICAN REPUBLIC. La Altagracia: Between Higüey and Miches, ˜20 km N of Otra Banda, Jones & Norris 1403A, 1405A (NY). Dajabón: Cañada Tirolis, 1 km S of Villa Anacaona, 396 m, Reese 15386, 15390 (NY). La Estrelleta: Sierra de Neiba, from 8 to 15 km N of Angel Féliz, 1402–1829 m, Buck 4605 (NY). La Vega: Vicinity of Piedra Blanca, 200–500 m, Allard 16886, 16941d, 16962 (NY, US). Pedernales: 4 km E of Mencia, 457 m, Reese 15073 (NY). Samaná: Parque Nacional Los Haitises, S of Naranjo, Smith 10416 (NY); Hacienda Nydia, 3.5 km E of Las Terrenas, 122 m, Steere 23193 (NY). San Cristóbal: Mano Matuey, 12 km from Cambito Garabito, 549 m, Buck 5159, 5197, Reese 15608, 15622, 15628 (NY). Sánchez Ramírez: Loma La Guacara, N of Comedor Arriba, ˜5 km S of Fantino, 100–240 m, Zanoni & Peláez 16185 (MO, NY, U). PUERTO RICO. Arecibo, between Manati and Morovis, ˜300 m, Buck 16149 (DUKE, MO, NY). ˜6 km SSW of Florida, just S of junction with Río Yúnes, Reese 14434 (NY). Cordillera Central, 13 km S of Utuado, 6 km N of Adjuntas, Reese 14511, Buck 3759, 3763 (NY). Cordillera Central, above Villalba, Dona Juana Recreation Area, 800-900 m, Allen 6390 (MO). Between Arecibo and Utuado, Britton 2077 (F, FH, MICH, NY). LEEWARD ISLANDS. Nevis: St. George Gingerland Parish, Golden Rock Estate, Golden Rock Nature Trail, ˜300 m, 17°08'N, 62°34'W, Buck 29538 (NY). WINDWARD ISLANDS. Guadeloupe: Rivière Beaugendre, LeGallo 651 (CANM), 705 (MICH). COLOMBIA. Antioquia: Mpio. Liborina, N of Liborina en la carretera a Sabanalarga, 910–1030 m, Churchill et al. 14653 (NY); Mpio. Dabeiba, 2–3 km NE of Dabeiba, 460 m, Churchill et al. 17048 (CANM, NY). Caldas: Mpio. of Río Sucio, on the Río Sucio, Vereda el Tabor, 1900–2080 m, Churchill & Arbeláez 15646 (NY). Huila: Transecto Neiva, San Vicente del Caguán, Río Las Ceibas, 680 m, Churchill & Betancur 16767 (CANM, NY). Nariño: Mpio. Alban, Campo Bello, 01°28'N, 77°04'W, Ramírez P. 5868 (MO); Mpio. de Barbacoas, km 1 El Mirador–Junín, 1500 m, 01°08'N, 78°09'W Ramírez P. 3631 (NY); Mpio. de Sandoná, 1900 m, Ramírez 5678 (NY). SURINAM. Without locality, Korthals s.n. (H). ECUADOR. Azuay: Gualaquiza, Jun 1909, Allioni 8343 (H). BRAZIL. Paraná: Mpio. Foz do Iguaçu, Parque Nacional do Iguaçu, forest along Rio Iguaçu, 100–200 m, ˜25°40'S, 54°25'W, Vital & Buck 12074 (NY). São Paulo: Serra de Paranapiacaba, Eldorado, Caverna do Diabo, 500 m, Schäfer-Verwimp & Verwimp 11082 (CANM). BOLIVIA. Santa Cruz: Aneres Ibanes, Espejillas, 20 km W of San José, 17°56'S, 63°26'W, Fuentes 140A (MO). less ↑