Lindsaea mesoamericana A.Rojas & Tejero, 2017

Rojas-Alvarado, Alexander Francisco & Tejero-Díez, José Daniel, 2017, Novelties and notes in Lindsaea (Lindsaeaceae) from Mexico and Central America, Phytotaxa 296 (2), pp. 147-160 : 152-157

Lindsaea mesoamericana A.Rojas & Tejero , sp. nov. ( Fig. 3 )

Lindsaea mesoamericana is similar to L. falcata Dryand. and L. leprieurii Hooker (1846: 208) but it differs in having 1 or 2-pinnate blade (vs. only 1-pinnate in L. falcata and L. leprieurii ), more segments per pinna (20–28 vs. 4–15 pairs per pinnae), blades (or distal pinnae in 2-pinnate specimens) that are narrowly triangular (vs. suboblong), with distal part of blade reduced up to 65–75% from baseline (vs. 10–15%), apical segments reduced and lobate (vs. the largest and hastate).

Type: — MEXICO. Oaxaca: Mpio. Santa María Chimalapa, cabecera del río Escolapa , ca. 15 km S de Santa María, 16°50 ʹ N, 94°41 ʹ W, 400 m, 22 October 1985, H. Hernández & C. González 1718 (holotype: MEXU!; isotypes: CHAPA!, IEB!, NY!) .

Perennial terrestrial herbs, rhizomes long creeping, 2–3 mm in diameter, with fronds every 5–8 mm; rhizome scales 1–3 × 0.3–0.5 mm, cordate-ovate with aristate apices on the stipe bases to lineate-deltate with short hair-like apices on the rhizome tips, orange-reddish to reddish-brown or dark reddish, lustrous, clathrate or occluded; fronds 40–50 cm long, ascending to arching; stipes 13–25 cm long, 1/3–1/2 the length of fronds, dark brown to grayish in the rachis, sharply angular, winged abaxially in distal part; blades 1 or 2-pinnate (then with 2–5 conform pinnae: 1–4 lateral ones plus terminal one); the 1-pinnate frond parts (entire blade or apical pinna in 2-pinnate plants), linear-deltate, 16–30 × 4–6 cm, the apical ones with presence of a petiolule 2–3 cm long, with (15–)20–28 pairs of segments, that are contiguous but not overlapping, gradually reduced towards apex (the distal reduced 60–75 % with respect to proximal ones); segments dimidiate, dark green, glabrescent with scattered 2–3 celled subcapitate hairs on the abaxial surfaces (0.15 mm); basal segments 2–3 × 0.8–1 cm, falcate-trapezoid, short-petiolulate (1–1.5 mm), decurrently on the rachis and form an adaxial wing; distal segments rectangular-trapezoidal with acroscopic side perpendicular to rachis, proximal segments short divergent and distally round-truncate, subsessile; apical segment subconform, stretch tringle, equal to or larger than adjacent lateral segment; lateral pinnae in 2-pinnate plants ascending 35–45º, narrowly lanceolate, anadromic, with a small basal acroscopical pinnule, subaxillar, flabellate (cuneate), 2–3 proximal segments reduced, quadrangular, subsequent segments rectangular-arched; pinnules with a thickened strand of elongate cells runs along the margin, highlighted veins, the primary forked sympodially 3–4 times; sori continuous, parallel and close to acroscopic margin; indusium 0.5 mm wide, entire, green.

Etymology:— The name of the new species refer to its known distribution.

Distribution:— Known from Mexico to Panama at (0–)300–600(–1050) m, in the understory of tropical rain forest.

Additional revised specimens (paratypes):— MEXICO. Chiapas: Mpio. Ocosingo, 20 km arriba del arroyo Miranda (por Canoa ), entrando por el río Chajul , 150 m, 19 February 1985, G. Castillo et al. 3973 ( XAL!) ; Mpio. Ocosingo, Nvo. Veracruz, a 33 km al W del vértice del río Chixoy camino a Chanjul , en zona 1 Marqués de Comillas , 130 m, 10 January 1986, E. Martínez 15962 ( IEB!, MEXU!) ; ibídem, E. Martínez 16002 ( MEXU!), ibídem, E. Martínez 16038 ( MEXU!) ; Mpio. Ocosingo, en vértice del río Chixoy, sobre el camino Boca Lacantum Chajul, 130 m, 12 January 1986, E. Martínez 16159 ( MEXU!) ; Mpio. Ocosingo, en ejido Roberto Barrios, a 60 km al S de Boca Lacantum, camino a Chajul , 200 m, 18 April 1986, E. Martínez 18349 ( MEXU!) ; ejido Loma Bonita , 150 m, 24 January 1992, E. Martínez & R. Lombera 26076 ( IEB!, XAL!) ; Mpio. Benemérito de Las Américas, 2.8 km después de Loma Linda, rumbo a Benemérito de Las Américas, 16º04 ʹ 55 ʺ N, 90º34 ʹ 06 ʺ W, 179 m, 30 July 2011, A Mendoza et al. 1784 ( IEB!, UAMIZ!). Oaxaca: Mpio. Santa María Chimalapa , río Verde , 7 km en línea recta al NNE de Santa María Chimalapa, 360–430 m, 16°58 ʹ 17 ʺ N, 94°39 ʹ 45 ʺ W, 4 August 1985, P. Caletti et al. 75 ( CHAPA!) ; cañada del río Negro, en la embocadura del arroyo Huapontl , ca. 15 km al S de Santa María, 16°56 ʹ N, 94°39 ʹ 30 ʺ W, 300 m, 14 October 1985, H. Hernández & C. González 1628 ( CHAPA, MEXU!) ; El Callejón, vieja vereda a Lázaro Cárdenas, cerca del camino actual que sale a arroyo Chocolate , subiendo la falda NE de cerro Azul-Escolapa , ca. 8 km al SW de Santa María, 16°52 ʹ N, 94°44 ʹ W, 350 m, 27 February 1986, H. Hernández 2109 ( CHAPA!, MEXU!) ; ca. 10 km S de Santa María Chimalapa, por la vereda a la cabecera del Río Escolapa , 16°50 ʹ N, 94°41 ʹ W, 500 m, 24 June 1987, H. Hernández 2535 ( CHAPA!) ; filo del cerro entre el río Blanco (al E) y el río Los Milagros (al O), ca. 25 km al SE de Santa María Chimalapa, 16°51 ʹ N, 94°35 ʹ W, 600 m, 21 August 1987, H. Hernández 2674 ( CHAPA!) ; Mpio. San Juan Lalana, senda de Santiago Jalahui al río Cuñeta , 210 m, 23 June 1991, J. Calzada & M. Aranda 17039 ( MEXU!) ; Mpio. Ayotzintepec, Sierra de Juárez, entre Puerto Eligio y Comaltepec, km 150 entre Tuxtepec y Oaxaca, 29 October 1965, G. Martínez 440 ( UAMIZ!, MEXU!) ; 6 km antes de Puerto Eligio, rumbo a La Esperanza , km 56 de la carretera Tuxtepec-Ixtlán- Oaxaca, 17º43 ʹ 31 ʺ N, 96º19 ʹ 37 ʺ W, 600 m, 8 September 2006, A. Mendoza et al. 1029 ( UAMIZ!, XAL!) ; District of Villa Alta, valley of the Yelagago River , ca. 20 millas NE of Villa Alta, 17°25 ʹ N, 96°05 ʹ W, 762–1158 m [2500–3800 ft], 28 July 1962, J. Mickel 1048 ( MEXU!, NY) .

GUATEMALA. Chimaltenango: 1937–1941, J. Johnston s. n. ( EAP!). Izabal: El Estor , March 1972, E. Contreras 11396 ( MEXU!) ; vicinity Exmibal, Camp 2 ( La Gloria ), NW of Lake Isabal, 15º15–35 ʹ N, 89º0–25 ʹ W, 400–500 m, 9 May 1966, G. Jones & L. Facey 3338 ( EAP!) ; entre caseríos Cienega y Ceja , 15°40 ʹ N, 89°W, 13 March 1972, R. Tún 2383 ( EAP!) .

BELIZE. N boundary of the BFREE reserve, in the NE Bladen watershed, 16°34 ʹ 33 ʺ N, 88°42 ʹ 11 ʺ W, 65 m, 7 June 2008, S. Brewer 4367 ( MEXU!) ; upper Mare ʹ s Nest Branch in the Cockscomb Basin Wildlife Sanctuary , 16°44 ʹ 4 ʺ N, 88°41 ʹ 49 ʺ W, 250 m, 21 May 2008, S. Brewer & Z. Goodwin 4262 ( MEXU!). Cayo District: Maya Mountains , Smokey Branch, 16.33°N, 89.02°W, 450 m, 17 March 1994, A. Monro & T. Helgason 219 ( MEXU). Stann Creek District: 28 mi section, Humming Bird Hwy, 28 January 1957, P. Gentle 9305 ( MEXU!) ; helicopter landing site 500 (HLS 500), 3.5 km E of Union Camp, 16°24 ʹ 02 ʺ N, 89°06 ʹ 40 ʺ W, 940 m, 19 February 1997, T. Hawkins 1474 ( MEXU!, MO!, SEL!). Toledo: Columbia River Forest Reserve , ca. 2 km SW of SW end of Little Quartz Ridge, 16°22 ʹ 58 ʺ N, 89°07 ʹ 10 ʺ W, 700 m, 11 February 1997, B. Holst 5675 ( MEXU!, MO!, SEL!) .

HONDURAS. Atlántida: Mpio. Esparta, 41.5 km E of Tela on the Tela-Ceiba Hwy , then 2.7 km N alog old timber road, 15°39 ʹ N, 87º16 ʹ W, 100 m, 17 April 1994, A. Brant & D. Hazlett 2859 ( EAP!, MO!) ; campamento Quebrada Grande, ca. 10 km SW of Pa Ceiba, at base of N slope of Pico Bonito , from camp to 2 km E of camp, 15°42,N, 86°51 ʹ W, 80–180 m, 10 May 1993, R. Liesner 26151 ( EAP!, TEFH!) ; montaña La Manga, 30 km SE de La Ceiba, 1050 m, 13–18 April 1976, C. Nelson et al. 3279 ( EAP!). Gracias a Dios: cuenca del río Plátano, crique Sulawala , 12–14 May 1977, M. Erazo 338 ( TEFH!) ; NE Honduras, near the río Patuca , under 175 m, 30 August–3 September 1973, A. Clewell 4576 ( EAP, TEFH) ; orilla del río Dursuna , 70 m, 14–21 February 1979, C. Nelson & E. Vargas 4946 ( TEFH!) ; ibídem, C. Nelson & E. Vargas 4947 ( EAP!) ; near road crossing of río Dursuna , SW of Dursuna forest camp, 150 m, 5 February 1981, G. Proctor 38898 ( TEFH!) .

NICARAGUA. San Juan del Norte: en las lomas El Encanto, a 30 km al W de San Juan del Norte sobre el río Indio, 11°05 ʹ 30 ʺ N, 83°53 ʹ W, 7 September 1982, E. Martínez & R. Riviere 2015 (MEXU!).

COSTA RICA. Cartago: carretera a Paraíso en junta con la de Buenos Aires, November 1979, L. Gómez 7209 ( CR!) ; Turrialba, Reserva Forestal Río Pacuare , cuenca del Matina , sendero principal hasta río Barbilla , 9°58 ʹ 24 ʺ N, 83°27 ʹ 02 ʺ W, 500 m, 18 September 1999, E. Mora & A. Rojas 482 ( CR). Heredia: Sarapiquí, Puerto Viejo, estación biológica La Selva, lindero sur, ca. 10º26 ʹ N, 84º01 ʹ W, 110 m, 10 September 2000, M. Blanco & J. Horn 1575 ( USJ!) ; Sarapiquí, La Selva Biological Station , 10º26 ʹ N, 84º01W, 50–150 m, 20 June 2002, M. Jones & P. Olivas 492 ( CR!, USJ!) ; cerros Sardinal, ca. 2–2.5 km N of Chilamate de Sarapiquí (finca La Martita), 10°28´N, 84°04´W, ca. 80–160 m, 21 January 1986, A. Smith et al. 1805 ( CR!). Limón: 7 km SW of Bribrí , 100–250 m, 4 May 1983, L. Gómez et al. 20420 ( CR!, MEXU!) ; Talamanca, Sixaola, Gandoca, El Lano, Bosque de Orey, 9°37 ʹ 10 ʺ N, 82°38 ʹ 00 ʺ W, 10 m, 3 April 1995, G. Herrera 7686 ( CR) ; Talamanca, Cahuita, Gandoca, El Llano, humedales de Manzanillo, 9°37 ʹ 00 ʺ N, 82°38 ʹ 00 ʺ W, 0–5 m, 17 October 2010, A. Rojas 9055 ( CR!) . PANAMA. Coclé: along trail from caño Sucio to cerro Tife at base of waterfall, caño Sucio is a 5 hr walk from the sawmill at El Cope on the Atlantic slope, 366–427 m [1200–1400 ft], 3 February 1980, T. Antonio 3699 ( CR) .

Notes: —In the herbarium, most specimens of this species have been determinated previously as Lindsaea lancea . Linnaeus (1753) described this latter species, based on a specimen from Suriname, as: “ pinnate fronds: pinnae opposite, lanceolate-oblong, with triangular hastate termination.... Pinnulae recurved, oblong, obtuse: the terminals [apical segment] cordate or hastate-triangular shaped, oblong, acute ”. This description does not correspond to L. mesoamericana here described, because it is primarily one-pinnate or 2-pinnate with one (two) pair of pinnae at the base, the blade is gradually reduced, and the apical segment varies between lanceolate to flabellate but not hastate and large. While medial segments of Lindsaea mesoamericana are recurved, the two or three basal pairs are distinctively arched. The Linnaean description is also not consistent with the type of L. falcata (BM-000937670!: N. Funck & L.J. Schlim 592, Venezuela), a name used in infraspecific combination of L. lancea , since it is 1-pinnate, the blade is suboblong with few leaflets, and the basal ones are strongly falcate (the apical segment are similar to that described by Linnaeus). Lindsaea mesoamericana differs from L. lancea by its 1 or 2-pinnate (vs. full 2-pinnate) blades, pinnules that are up to three times longer than wide (vs. ca. 2–2.5 times longer than wide), the basal and medial segments that are commonly falcate and curved down at apex (vs. pinnae with basiscopic side perpendicular to ascending with respect to rachis), the distal segments that are slightly or not reduced (vs. distal ones about 1/2 the size of the lower ones) and strongly asymmetric (vs. symmetric) terminal segment. For differences with L. leprieurii see the notes under that species.

Lindsaea mesoamericana is the most common species in Mesoamerica and is easily distinguished from other species in this area by the following combination of characters: blades (or terminal pinna in 2-pinnate specimens) with larger basal segments strongly recurved (falcate), towards the apex reduced in size and less recurved, ending in a subconform apical segment; lamina olive (dark green); stipe and rachis brown. The green wings along the rachis facilitate the recognition.

Lindsaea subalata (Kramer) A.Rojas & Tejero , stat. nov. Basionym: Lindsaea quadrangularis subsp. subalata Kramer (1957a: 190) .Type:— COSTA RICA. San José: from the vicinity of El General, Skutch 2241 (holotype US; isotypes K, MO, NY, S-PA)

Distribution:— Southern Mexico to Panama.

Note:— According to Kramer (1957a), this taxon is easily distinguishable from the other sub-species of Lindsaea quadrangularis by blades being gradually reduced (vs. slightly reduced in L. quadrangularis subsp. terminalis Kramer (1957a: 192)) , spores trilete (vs. monolete in L. quadrangularis subsp. quadrangularis ) and stipe reddish brown to dark (vs. stramineous in L. quadrangularis subsp. antillensis Kramer [1957a: 194] . Also, the status change considered here is based on the geographical distribution, as this taxon is typically Mesoamerican (vs. West Indies L. quadrangularis subsp. antillensis , and southeast Brazil and Paraguay in L. quadrangularis subsp. terminalis and L. quadrangularis subsp. quadrangularis ).

The other subspecies need to be raised to species status.

Lindsaea horizontalis Hooker (1844: 214) . Type:— BRAZIL. Rio de Janeiro: Organ Mountains , 1838, G. Gardner 157 (lectotype BM!; isolectotypes E!, K!)

Distribution:— Costa Rica, Panama, Colombia, Venezuela and Brazil.

Notes:— Kramer (1957a) treated this taxon as a synonym of Lindsaea arcuata . However, analysis of the types and other specimens demonstrates that it is distinct in having triangular pinnules, commonly 1-pinnate (vs. 2-pinnate) blades, when L. horizontalis is 2-pinnate have an irregular 1–2 couples of pinnae (vs. 3–8 pairs).

Lindsaea moritziana Klotzsch (1844 [1845]: 548). Lindsaea stricta var. stricta fo. moritziana (Klotzsch) Kramer (1957a: 228) . Type:— Venezuela. Monagas: “Caripe, ad rivul. i. pasc. mont.”, J. Moritz 164 (lectotype B!, lectotype designed by Kramer 1957a: 228; isolectotype BM!). The duplicate in MPU! is from Colonia Tovar and belongs to Lindsaea feei C. Chr.

Lindsaea quadrangularis Raddi subsp. mexiae Mickel (in Mickel & Smith, 2004: 367).— Type: Mexico. Jalisco: West of San Sebastian, Hacienda del Ototal , Arroyo de Los Hornos , Y. Mexia 1837 (holotype: NY!; isotypes: F, G, GH, IEB!, MICH, MO, UC, US).

Distribution:— Mexico, Guatemala, Honduras, Greater Antilles and South America.

Notes:— Kramer (1957a) and Moran (1995) considered this taxon as a form of Lindsaea stricta ( L. stricta var. stricta fo. moritziana . Mickel (in Mickel & Smith 2004) indicates that it is closest to L. quadrangularis (and form a new subsp.= L. quadrangularis subsp. mexiae ). However, it is obvious that the characters of L. moritziana are sufficiently divergent from both L. stricta and L. subalata (= L. quadrangularis subsp. subalata ) (see key). Lindsaea moritziana has fewer pinnae pairs (2–3 pairs vs. (2–)4–7 pairs), and fewer pinnules (12–18 pairs vs. 20–30(–40) pairs), that are shorter (5–6 mm long vs. 8–17 mm long), proportionally (1.2–1.5 times longer than wide vs. 2–3 times), and ending in an ovate to deltate segment (vs. pinnatifid). This species is likely more closely related to L. brasiliensis Desvaux (1811: 330) (= L. stricta var. parvula Kramer, 1957a: 230 ) based on its stramineous axes and small number of pinnae pairs, but it differs in having fewer (12–18 vs. (15–)25–50) pinnule pairs with an ovate to deltate terminal segment (vs. pinnatifid).

The presence in Jalisco, Mexico of this species ( Mexia 1837 IEB!, NY!, etc.) is the northernmost distribution of the genus.

New distributional records:

Lindsaea cubensis Underwood & Maxon (1907: 336) . Type:— CUBA. Villa Clara: Cuba Occ., Loma Pelado, 1859–1860, C. Wright 3947 (holotype: NY-00127167 !; isotypes: B-20 0141631!, GH-00021502!, GH-00359569!,

K-000644030!, NY-00127168!, P-00539202!, P-00539203!, P-00539204!, S-R3260!, U-0007333!, US-00066449!, US-01100897!)

Distribution: — Cuba, Nicaragua (first record).

Specimen examined:— NICARAGUA. Zelaya: Puerto Cabezas, 0–100 m, 12 March 1971, J. Atwood 4516 (MEXU!).

Notes:— Lindsaea cubensis is close to L. stricta in having strongly vertical fronds and similar habitat preferences (open areas near seasonal water bodies), but it differs in having subdimorphic fronds (vs. monomorphic), 1-pinnate (vs. 1–2-pinnate, but more commonly 2-pinnate) blades, fewer pinnate pairs ((5–) 12–22 pairs vs. 25–70 pairs), and pinnae with ascending basiscopic sides in respect to the costae (vs. perpendicular to reflexed).

Lindsaea divaricata Klotzsch (1844: 547) . Type:— GUYANA. “Guiana angl.”, M.R. Schomburgk 368 (holotype B-200079709!)

Distribution:— Costa Rica (first record), Panama, Guadeloupe, Venezuela, Guyana, French Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia and Paraguay.

Specimens examined: — COSTA RICA. Limón: Talamanca, Sixaola, Gandoca , El Llano , bosque de Orey, 9°37 ʹ 10 ʺ N, 82°32 ʹ 04 ʺ W, 10 m, 3 April 1995, G. Herrera 7694 ( CR!) ; Talamanca, Sixaola, Gandoca , El Llano, entre filas Manzanillo y río Mile Creek , 9°37 ʹ 00 ʺ N, 82°41 ʹ 00 ʺ W, 50–100 m, 27 March 1995, G. Herrera & E. Sandoval 7597 ( CR!, MO!) ; Parque Nacional Tortuguero, 600 m SW de Tortuguero sobre el río Tortuguero , se toma al W en zona cenagosa, 10°31 ʹ N, 83°30 ʹ W, 1 December 1987, R. Robles 1408 ( CR!, MO!) ; Talamanca, Cahuita, Gandoca , El Llano , humedales de Manzanillo , 9°37 ʹ 00 ʺ N, 82°38 ʹ 00 ʺ W, 0–5 m, 17 October 2010, A. Rojas 9054 ( CR!). Heredia: Sarapiquí , La Selva Biological Station , 10º26 ʹ N, 84º01 ʹ W, 50–150 m, 29 May 2007, M. Jones & M. Putkonen 616 ( USJ!) ; Sarapiquí, Estación Biológica La Selva , 10º25 ʹ 49 ʺ N, 84º00 ʹ 25 ʺ W, 18 January 2008, M. Sundue et al. 1514 ( USJ!) .

Notes: —The mentioned specimens of Lindsaea divaricata from Costa Rica were previously determinated as L. quadrangularis based on their narrow terminal segment, but they differ in having brown to reddish-brown (vs. stramineous) stipes and rachises that are not angulate (vs. angulate) abaxially. Lindsaea divaricata is also similar to Lindsaea quadrangularis subsp. terminalis because both have strongly ascending pinnae and pinnules, but the later species has blackish stipes and rachises, fewer pinnae (commonly 2–3 vs. commonly 4–6 pairs), fewer pinnules ((5–)10–15 vs. (13–)20–25) pairs, and a deltate to hastate terminal segment (vs. narrowly lanceolate).

Lindsaea horizontalis Hooker (1844: 214) . Type:— BRAZIL. Rio de Janeiro: Organ Mountains , 1838, G. Gardner 157 (holotype BM!; isotypes E!, K!)

Distribution:— Costa Rica (first record), Panama (first record), Colombia, Venezuela and Brazil.

Specimens examined:— COSTA RICA. Cartago: Paraíso, Parque Nacional Tapantí, sendero Dos Quebradas , 9 April 1986, J. Berrocal & J. Sánchez 140 ( CR!) ; cerro entre cerro Chimú y cerro Matama , 1200 m, 30 April 1985, L. Gómez 23569 ( CR!) ; Paraízo, Parque Nacional Tapantí-Macizo de La Muerte, cuenca del reventazón, sendero Árboles Caídos , 9°45 ʹ 00 ʺ N, 83°47 ʹ 00 ʺ W, 1200–1300 m, 9 July 1997, R. Guzmán 113 ( CR, MO) ; Turrialba, Moravia de Chirripó , 1400 m, 1 January 1976, R. Ocampo 1184 ( CR!) ; Turrialba, valle del Reventazón, Jicotea , finca La Pradera , Jicotea , 9.7875°N, 83.5472°W, 1000 m, 22 June 1995, G. Rivera & A. Rojas 2472 ( CR!) ; Paraíso, Cordillera de Talamanca, Estación Biológica Río Macho , 300 m SW del embalse, 9°45 ʹ 11 ʺ N, 83°51 ʹ 48 ʺ W, 1700 m, 30 May 1994, A. Rojas 1047 ( CR) ; Turrialba, Cordillera de Talamanca, Tayutic , Jicotea , 9°47 ʹ 15 ʺ N, 83°32 ʹ 50 ʺ W, 1100–1600 m, 22 June 1995, A. Rojas et al. 2006 ( CR!) ; Paraíso, Parque Nacional Tapantí, cuenca del río Reventazón , sendero Árboles Caídos y Oropéndula , 9°45 ʹ 20 ʺ N, 83°47 ʹ 00 ʺ W, 1300–1500 m, 7 April 1999, A. Rojas & A. Soto 5020 ( CR!, MO) ; ibídem, 19 November 1997, A. Rojas et al. 3971 ( CR!, MO, USJ) ; Cartago, Agua Caliente, Muñeco , 9°46 ʹ N, 83°54 ʹ W, 1700 m, 2 April 2004, A. Rojas et al. 5570 ( CR!, MO) ; Paraíso, Parque Nacional Tapantí-Macizo de La Muerte, sendero Árboles Caídos , cerca de 200 m antes de la salida del sendero, 9º45 ʹ 05 ʺ N, 83º46 ʹ 55 ʺ W, 1300–1350 m, 22 April 2012, A. Rojas et al. 9730 ( CR!, MO, USJ) ; Orosi, Parque Nacional Tapantí-Macizo de La Muerte, sector Tapantí , sendero Árboles Caídos , en la última parte del circuito, 9º46 ʹ 00 ʺ N, 83º 47 ʹ 20 ʺ W, 1220–1300 m, 5 June 2012, A. Rojas et al. 9830 ( CR!, MO, USJ) ; 5.6 km of San José, side of road leading to Palmichal, steep new dirt road up hill from Pan American Hwy, 1700–1800 m, K. Walter 79490 ( CR). Limón: Talamanca , trail ca. 5 km up Alto Lari , 9.4042°N, 83.0847°W, 1300 m, 27 February 1992, J. Bittner 1335 ( CR!) ; Talamanca, camp by Kivut, trail between Kivut and Alto Lari , 1 km up to Alto Lari , 9.4042°N, 83.0847°W, 1200–1300 m, 6 March 1992, J. Bittner 1450 ( CR!) ; Talamanca, Alto Urén, cerro Laúbeta , entre río Lorni y quebrada Chacho , 9.3861°N, 83.0069°W, 1190 m, 27 July 1989, G. Herrera 3381 ( CR!) ; Limón, Almirante, cuenca superior del río Xichiari , 9.7639°N, 83.3292°W, 1300 m, 14 August 1995, G. Herrera 8480 ( CR!) ; near Moravia de Chirripó, 1300 m, 7 April 1949, L. Williams 16186 ( EAP!). Puntarenas: Las Cruces , ridge 6 km S of San Vito de Java, 4500 ft, February 1974, McAlpin 2421 ( CR) ; Buenos Aires, Parque Internacional La Amistad, Cuenca Térraba-Sierpe , Potrero Grande , La Lucha , cabeceras del río Guineal , 9°06 ʹ 38.36 ʺ N, 83°05 ʹ 21.84 ʺ W, 1600– 1700 m, 22 February 2008, A. Rodríguez et al. 11850 ( CR, PMA). San José: Tarrazú, camino entre cerro Pito y cerro Toro, 9°34 ʹ 50 ʺ N, 84°04 ʹ 10 ʺ W, 1200–1400 m, 26 January 1998, A. Estrada 1430 ( CR) ; El Guarco, Cerros de Escazú-La Carpintera, La Cangreja , 9°48 ʹ 00 ʺ N, 83°57 ʹ 50 ʺ W, 1900 m, 26 June 1993, A. Rojas 303 ( CR) .

PANAMA. Chiriquí: along road from Gualaca to Fortuna dam site, 5.9 mi NW of Los Planes de Hornito, 8.7167°N, 82.2833°W, 1370 m, 9 April 1980, T. Croat 49868 (CR!, MO!).

Notes:— Lindsaea horizontalis has been included as synonym of L. arcuata , for differences see up in the validation of this species.

Figures

FIGURE 3. Lindsaea mesoamericana (H.Hernández & C. González 1718, MEXU).A. General aspect of type specimen. B. Abaxial blade detail.