Ramalina caracasana Müller Argoviensis

Marcano, Vicente, Méndez, Antonio Morales & Prü, Ernesto Palacios, 2021, The genus Ramalina Acharius (Ascomycota, Lecanoromycetes, Ramalinaceae) in northern South America, Phytotaxa 504 (1), pp. 1-77 : 36-45

5. Ramalina caracasana Müller Argoviensis (Fig. 31)

Flora, Jena: 64: 86 (1881). Type :― VENEZUELA. Caracas: A . Ernst 230 (holotype G) .

Thallus corticolous, erect, coarser, robust, moderately to densely branched, up to 5 cm long. Branches pale yellow or greenish, solid, surface wrinkled-ridged. Pseudocyphellae punctiform, raised on tubercles. Soralia not seen. Pycnidia not seen. Apothecia lateral, subapical or apical, spurred. Ascospores fusiform, 16–19 x 4.5–5 µm.

Chemistry (TLC; HPTLC): Strain 1. Stictic, boninic, hypoprotocetraric, succinprotocetraric and protocetraric acids (Ortiz et al. 663). Strain 2. Protocetraric acid (Vareschi 4392, Vareschi 7759).

Ecology and distribution: This species grows over trees and shrubs in rainforests; 1000–1320 m. It is known only from northern Venezuela.

Remarks: Ramalina caracasana is a rare species which could be confused with R. africana , but the latter species cointains the sekikaic acid complex and has ellipsoid spores (11–14 x 3.5–4.0 µm). Although R. caracasana contains accessory stictic and boninic acids, it is characterized in the main by substances in the protocetraric acid complex. Krog & Swinscow (1975) reported the presence of protocetraric acid in this species.

Specimens examined: VENEZUELA: Miranda: La Guairita, Cave El Indio , 1000–1200 m, 15 August 1984, Ortiz et al. 663 ( VEN); near Tejerias, 1200 m, 12 July 1962, V . Vareschi 7759 ( VEN); Los Guayabitos, 1320 m, 6 November 1955, V . Vareschi 4392 (VEN).

. Ramalina celastri (Sprengel) Krog & Swinscow

Norw. J. Bot. 23: 159 (1976). – Parmelia celastri Sprengel, Syst. Veg. 4: 328 (1827). Type :― SOUTH AFRICA. Cap ., C. F. Ecklon (holotype S).

Thallus corticolous, conspicuous, rigid, subpendulous, very variable, up to 25 cm long, branching dichotomously anisotomic to irregular. Branches solid, flattened, lanceolate, surface smooth or furrowed, 0.9–6.0 mm wide, older branches longitudinally or reticulately ridged from strands of cartilaginous tissue, often with holes or cracks. Pseudocyphellae short linear, marginal and/or laminal, abundant, plane or concave. Chondroid tissue not cracked, continuous. Pycnidia not seen. Apothecia abundant, rounded, lateral or laminal, disc flat or convex, up to 1.7 mm diameter. Ascospores 1–septate, short fusiform, 13–17 x 3.5–5 µm.

Chemistry (TLC, HPTLC): Nil (Morales 92; Cleef 4190; Sipman & Velosa 32725; Rubiano 433, 434, 435; Sipman et al. 34117; Silvestone-Sopkin 3099; Aguirre & Sipman 6119; Vareschi 37, 2466–B, 2833, 3208–E, 3499, 3957, 5717, 6160, 6207–A, 7265, 7758, 8033, 8475, 8603, 8602, 8604, 8696, 9842; Pittier 11899; Morillo & Manara 390; Sipman et al. 34043), or usnic acid only (Allart 74; Vareschi 5721).

Ecology and distribution: This species is frequent in diverse habitats, where it is found growing on rocks and bark, in pastures, rainforests and paramo. In Colombia, it occurs on the bark of Salix sp. , Facara sp. ( Rutaceae ), Eucalyptus sp. , Fraxinus sp. , Xylosma sp. , Vallea sp. , Citrus sp. as well as other shrubs in pastures, and in forests with Anacardium sp. at 950–3500 m. It is known from Australia, Tasmania, East Africa, South Africa, and South America ( Chile, Brazil, Perú, Colombia, Venezuela, Uruguay and Argentina).

Remarks: Ramalina celastri is one of the most common Ramalina species in Colombia and Venezuela. Ramalina celastri resembles R. cumanensis but R. cumanensis has canaliculate branches and 1–septate, ellipsoid ascospores (8–12 x 4–5 µm). Both species lack medullary compounds.

Specimens examined: COLOMBIA: Cauca: Popayan, Los Robles, campus of Fundación Universitaria, c. 15 km towards Timbío, 1750 m, 29 May 1986, H . Sipman & R . Velosa 32725 ( B). Valle: Municipio Zarzal, hacienda El Medio, Road Panamerican , between La Paila and zarzal, 950 m, 16 April 1987, F . A . Silverstone-Sopkin, N . Paz & J . Larrahondo 3099 ( B). Antioquia: Municipio Medellín, finca Yaromal, vereda Chuscal , c 10 km from San Antonio de Prado , 2300 m, 3 July 1986, H . Sipman, M . Escobar & J . Rubiano 34043 ( B); Municipio El Retiro, along road Medellín – La Ceja, between Las Palmas and El Retiro, Hacienda Fizebad, entrance of reconstructed farm, 2050 m, 6 July 1986, H . Sipman, M . Escobar & J Rubiano 34117 ( B). Huila: E-slope of Cordillera Central, La Plata, hacienda Limana , ca. 5 km from La Plata towards Paicol, 1000 m, 8 October 1984, J . Aguirre & H . J . M. Sipman 6119 ( B). Cundinamarca: Represa del Neusa, near rio Neusa , 3 km southern Represa Neusa, 2750 m, 26 May 1972, A . M . Cleef & R . Jaramillo 4190–c ( B); Tequendama, Archaeological Park , 2600 m, 15 September 1972, P . A . Florschütz 3714 ( B, COL, U); Zipaquirá , 2550 m, 16 june 1983, L . J . Rubiano 435, 436– A ( B) ; Zipaquirá, 2550 m, 15 April 1983, L . J . Rubiano 433, 434 ( B) . VENEZUELA: Dto. Federal, El Avila National Park , road Los Castillitos – El Rincón, 1200 m, 22 February 1971, G . Morillo & B . Manara 390 ( VEN); Dto. Federal, Caracas , Sabana Grande, 15 July 1950, V . Vareschi 37; El Avila National Park, between Cotiza and Los Venados, October 1924, Allart 74 ( VEN). Mérida: Los Topes, San Juanito , near Chiguará , 1200 m, 12 February 1984, A . Morales 92 ( B, MERF); Sierra Santo Domingo, Mucubaji lake , 3500 m, 19 May 1993, D. Martinez, V . Marcano & L . Galiz 190, 191, 195 (herb. lich. V. Marcano, VEN); Paramo Los Conejos road, 2200–2400 m, 6 April 1989, V . Marcano s/n (herb. lich. V. Marcano, VEN); Paramo La Negra , 2900–3100 m, 15 April 1954, V . Vareschi 3208– E ( VEN) ; Near Chachopo, above hacienda Schwarzkopf, 3150 m, 10 August 1969, V . Vareschi 8475 ( VEN); Chachopo, near Motatan river , 3100 m, 20 August 1972, V . Vareschi 8602, 8603, 8604 ( VEN); near Timotes , 3000 m, 5 september 1964, V . Vareschi 8033 ( VEN); Ejido , 1700 m, 10 November 1952, V . Vareschi 2466– B ( VEN) ; Chachopo, on rock, 3000 m, 11 November 1958, Vareschi 7265 ( VEN); near Chachopo , 3000 m, 5 September 1956, V . Vareschi 5214 ( VEN); Raiz de Agua, Parque Sierra Nevada de Merida, 2700 m, 28 July, 2020, V . Marcano & L . Castillo 20–92, 20–96 (herb. lich. V. Marcano). Miranda: Near Los Guayabitos , 1100 m, 1 August 1964, V . Vareschi 3499 ( VEN); Lomas de Sartaneja, near Los Guayabitos , 6 July 1955, V . Vareschi 3957 ( VEN); Sebucan , near Los Dos Caminos, 27 September 1925, H . Pittier 11899 ( VEN); Near Colonia Tovar , 2000 m, 5 November 1979, V ., Vareschi 9842 ( VEN); Tiara, near Tejerias , 1200 m, 12 July 1963, V . Vareschi 7758 ( VEN); Los Guayabitos, 1 January 1957, Vareschi 6160 ( VEN); Los Guayabitos , 8 January 1956, Vareschi 6207– A ( VEN) ; Los Guayabitos, 1200 m, 22 March 1953, V . Vareschi 2833 ( VEN). Táchira: Between Betania and Villa Paez , 2200–2400 m, 15 April 1992, V . Marcano s/n ( MER); Garcia lake, near Pregonero , 14 April 1992, V . Marcano A and B ( MER) ; Paramos, 3000 m, 3 October 1956, V . Vareschi 5717 ( VEN) .

. Ramalina chiguarensis V. Marcano & A. Morales

The Bryologist 97: 27 (1994) . Type:― VENEZUELA. Mérida: Chiguará, El Paramito, 1000 m, 12 February 1984, A. Morales 96 (holotype MERF!) .

Thallus corticolous, pendulous, soft, tufted from holdfast, densely and dichotomously branched, up to 12 cm long. Branches pale yellow, subshiny, complanate (1–1.5 mm broad). Pseudocyphellae lateral, laminal, linear, orbicular, ellipsoid, abundant on underside. Soralia abundant, lateral, laminal, producing coarse granules. Cortical tissue paraplectenchymatous, 6–8 μm thick, disrupted in lateral parts. Peripheral chondroid tissue sometimes forming continuous upper and lower layers, 30–45 μm thick. Pycnidia and apothecia not seen.

Chemistry (TLC, HPTLC): Strain 1. Sekikaic and homosekikaic acids (Lopez-Figueiras & Keogh 9224– B; Morales 123, 129). Strain 2. Sekikaic, homosekikaic and salazinic acids (Morales 97). Strain 3. Sekikaic acid (Marcano 174–92). Strain 4. Divaricatic acid (Morales 96; Aguirre & Sipman 5728–B). Strain 5. Divaricatic and cryptochlorophaeic acids and an unknown compound (Morales 96, 97).

Ecology and distribution: Ramalina chiguarensis occurs on the bark or decorticated wood of trunks and canopy branches; on dead wood in pastures ( Colombia); and on twigs in moist, secondary forests and paramo at 1000–3500 m. It is known only from northern South America ( Colombia and Venezuela).

Remarks: Ramalina chiguarensis closely resembles R. rectangularis as both species are characterized by dichotomous branches with lateral pseudocyphellae (Fig. 6) and complanate laciniae. However, the two species can be distinguished by both morphological and chemical character: R. rectangularis has isidiform soralia and contains salazinic acid, whereas R. chiguarensis lacks isidiform soralia and contains sekikaic acid (strains 1, 2 and 3) or divaricatic acid (strains 4 and 5) as major chemical substances. Ramalina chiguarensis is also very similar to R. andina (see above). Specimens of R. chiguarensis from a single population were found to contain different acids. For example, strains 1, 2 and 5 were collected in a single population on decorticated wood of trunks at Hacienda Los Topes, near Chiguará, Mérida. More surprisingly, different branches of a single thallus (Morales 96) were found to be of different strains (strains 2 and 5) including sekikaic, homosekikaic, divaricatic and cryptochlorophaeic acids.

Additional specimens examined: COLOMBIA: Huila: E-side of Cordillera Central , La Plata, vereda la Candelaria, Finca Merenberg, 2400 m, 30 September 1984, J . Aguirre & H. J. M . Sipman 5728– B ( B) . VENEZUELA. Mérida. Sierra la Culata , 3500 m, October 1993, V . Marcano 7168 ( MER); La Carbonera, between Mérida City and La Azulita, 1975, Lopez-Figueiras & Keogh 9224– B ( MERF); Hacienda Los Topes, near Chiguará, 1300–1400 m, 12 February 1984, A . Morales 96, 97 ( B); El Guamo, near Caña Brava, 1989, A . Morales 123 ( MERF); Chiguará, El Paramito, 1400 m, 1989, A . Morales 129 ( MERF) .

. Ramalina chilensis Bertero ex Nylander

Bull. Soc. Linn. Normand., sér 2, 4: 124 (1870) . Type :― CHILE. Quillota: 1829, C, G. Bertero 1213 (holotype H –NYL! 37513, isotype G) .

Thallus pendulous, rarely up to 30 cm long, contorted, flattened, whitish. Branches strongly flattened, apices subterete, reticulate, surface pale with prominent white striations, 1.8–5.2 mm broad, Soredia absent. Pseudocyphellae linear, striated, continuous. Pycnidia not seen. Apothecia lateral or laminal, common. Ascospores fusiform, 12–14(–16) x 2–2.5 µm.

Chemistry (TLC, HPTLC): Strain 1. Homosekikaic, sekikaic, boninic, succinprotocetraric (tr.) and protocetraric acids (Vareschi 2952, 2568; Morales 136). Strain 2. Sekikaic acid (Cleef 6043–b, 6055–a). Strain 3. Homosekikaic, sekikaic, boninic, and protocetraric (tr.) acids (Vareschi 2568).

Ecology and distribution: This species is found growing over trees, such as Quercus (in Colombia) in submontane and montane forests at 200–2780 m. It is known from South America ( Brazil, Chile, Colombia and Venezuela).

Remarks: This species is very similar to R. usnea , but the latter lacks pseudocyphellae and has long-fusiform ascospores (18–28 x 3–3.5 µm). In Colombia R. chilensis contains only sekikaic acid (strain 2), whereas in Venezuela, it contains sekikaic acid, homosekikaic and divaricatic acids as major components (strains 1 and 3), as well as accessory substances in the protocetraric acid complex. In Brazil, Kashiwadani & Kalb (1993) reported the presence of two strains, one containing sekikaic, boninic and 4ˊ– O –demethylsekikaic acids, and the other with norstictic acid. In Chile ( Rundel 1978a), R. chilensis contains salazinic, norstictic and sekikaic acids.

Ramalina chilensis could be confused with R. tenaensis and R. bogotensis . These species have contorted thalli. However, R. tenaensis can be distinguished by branches with a verruculose surface and containing divaricatic, boninic and 2ˊ– O –methylsekikaic acids whereas R. bogotensis has abundant granular soredia and long-fusiform spores (18–20 µm).

Specimens examined: COLOMBIA: Cundinamarca: San Cayetano, hacienda Portugal, Alto de Sendales , 2750– 2780 m, 8 November 1972, A . Cleef 6043–b, 6055–a ( B) ; VENEZUELA: Mérida: Near Chiguará, Los Topes, A ., 1200 m, Morales 136 ( MER) . Monagas: Near to Barrancas, in the margins of Orinoco river , 200–400 m, 25 August 1953, Vareschi 2952 ( VEN); Casacoima, margins Orinoco river , 7 January 1953, Vareschi 2568 ( VEN)

. Ramalina cochlearis Zahlbruckner

Bull. Herb. Boiss. ser. 2, 5: 542 (1905). Type :― BRAZIL. Cachoeira: Ad ramos arborum in sylvis, L . B . Damazio 1449 (holotype W) .

Thallus corticolous, fruticose, erect, up to 4.5 cm long. Branches dichotomous, solid, canaliculated ( Fig. 4 ), pale yellow or whitish, surface smooth, 500–925 μm broad. Pseudocyphellae not tuberculate, marginal, orbicular or ellipsoid (Fig. 7). Soralia helmet-shaped, present on the upper surface. Cortical tissue 18–22 um thick. Chondroid tissue continuous, 22–28 μm thick. Medulla white, loose. Pycnidia and apothecia not seen.

CHEMISTRY (TLC, HPTLC). This species presents 12 strains which are summarized in the Table 5. Strain 1. Homosekikaic, sekikaic, ramalinolic, 4’– O –demethylsekikaic, 4ˊ– O –methylnorhomosekikaic and 4’– O – methylnorsekikaic acids. Strain 2. Boninic and 2ˊ– O –methylsekikaic acids. Strain 3. Sekikaic, 4ˊ– O –demethylsekikaic, boninic, 2ˊ– O –methylsekikaic and salazinic (tr.) acids (Vareschi 4333). Strain 4. Homosekikaic, sekikaic, 4ˊ– O – demethylsekikaic, boninic, 2ˊ– O –methylsekikaic and protocetraric (tr.) acids (Pittier 9283). Strain 5. Homosekikaic, sekikaic, 4ˊ– O –demethylsekikaic, boninic, hypoprotocetraric and protocetraric acids (Vareschi 5718). Strain 6. Homosekikaic, sekikaic and boninic acids (Sipman et al. 34116–A; Aguirre & Sipman 5744–B; Vareschi 768,1 Aguirre & Sipman 5558; Wolf 530). Strain 7. Homosekikaic and sekikaic acids (Rubiano 436–B). Strain 8. Homosekikaic, sekikaic, 4ˊ– O –demethylsekikaic and boninic acids (Sipman & Valencia 10296). Strain 9. Homosekikaic, sekikaic, ramalinolic (tr.), 4ˊ– O –demethylsekikaic, boninic, succinprotocetraric, protocetraric and fumarprotocetraric acids (Vareschi 3392, Sipman & Velosa 32727, Cleef & Jaramill 4190–e). Strain 10. Homosekikaic, sekikaic and protocetraric acids (Vareschi 5931). Strain 11. Homosekikaic, sekikaic, 4ˊ– O –demethylsekikaic, boninic and protocetraric acids (Sipman & Velosa 33711). Strain 12. Sekikaic and ramalinolic acids (Cleef & van Rens de Cleef 4849–B).

Ecology and distribution: This species is corticolous, growing in cloud forests and paramo (940–3750 m). In the Guayana Highlands, it grows associated with Usnea spp., Ramalina celastri , Teloschistes flavicans ( Swartz 1788: 147) Norman (1853: 229) , Heterodermia spp. and Parmotrema crinitum ( Acharius 1814: 196) M. Choisy (1952: 175) , whereas in the Andes it is associated with R. celastri , R. cumanensis , Teloschistes flavicans and Heterodermia sp. (Fig. 30). In Colombia, R. cochlearis grows in Quercus humboldtii Bonpland (1805: 155) forests, and on dead trunks in pastures; it is also found on bark of Weinmannia , Xylosma , Vallea , Salix , Cyatheaceae and Brunnellia. This lichen is known from the Caribbean, Central America ( México, Guatemala and Honduras) and South America ( Colombia, Venezuela, Perú and Brazil).

Remarks: Ramalina cochlearis is a very common species in northern South America. It closely resembles R. escorialensis , R. vareschii and R. xalapensis V. Marcano & A. Morales (1994c: 198) , but R. escorialensis can be distinguished by the presence of cryptochlorophaeic acid and R. vareschii by the tuberculate pseudocyphellae. Ramalina xalapensis is known only from México and is characterized by marginal isidial structures and a lack pseudocyphellae. The morphological and chemical characters of these species have been described in detail by Marcano & Morales (1994c).

Specimens of R. cochlearis from Colombia and Venezuela showed 12 chemical strains. All the specimens examined did not contain usnic acid in detectable amounts. The major chemical substances found were homosekikaic, sekikaic and boninic acids, whereas accessory substances included some sekikaic acid derivatives and substances in the protocetraric acid complex (Table 5). Kashiwadani (1988) and Kashiwadani & Kalb (1993) reported only two chemical strains for this species in Perú and Brazil, and Marcano & Morales (1994c) reported two strains from Honduras. The high chemical variability (chemosyndromic) of R. cochlearis in northern South America could be explained by an active and progressive chemical evolution where the chemosyndromes are ordinated by side-chain length of the constituent compounds ( Culberson & Culberson 1976). This chemical evolution could be hypothetically driven by differing evolutionary selection pressures within the Andean cordillera, such as those taking place in the steep altitudinal gradient viz. effects of exposing to ultraviolet radiation, and by other ecological constraints ( Rundel 1978a,b, Sheard 1978a,b, Culberson et al. 1990, Bjerke et al. 2002, Marcano et al. 2010, Marcano & Sipman 2021).

Specimens examined: COLOMBIA: Risaralda: Santa Rosa de Cabal , ca. 500 km E of Termales santa Rosa, 2325 m, 13 December 1985, J . Wolf & A . Wolf 2250 ( B, COL, U); Santa Rosa de Cabal, ca. 500 km E of Termales santa Rosa , 2460 m, 11 December 1985, J . Wolf 530 ( B, COL, U); W-slope of Cordillera Central, Santa Rosa de Cabal, along track from termales de Santa Rosa to Termales Antiguos, 2000 m, 19 September 1984, J . Aguirre & H . J . M. Sipman 5558 ( B, COL) . Huila: E-side of Cordillera central, La Plata, vereda La Candelaria, Finca Merenberg, 2400 m, 30 September 1984, J . Aguirre & H . J . M. Sipman 5744–b ( B) . Cundinamarca: ca. 5 km SW of La Calera, Valle Teusacá , 2850 m, 9 July 1972, A . Cleef & M . van Rens de Cleef 4849 ( B); between Bogota and Fusagasuga, ca. 5 km SW of roadtoll, 2700 m, 31 January 1979, H . Sipman & H . Valencia 10296 ( B, COL, U); Represa del Neusa, near Valle Rio Neusa , 3 km southern Represa, 2750 m, 26 May 1972, A . M . Cleef & R . Jaramillo 4190– E ( B) ; Zipaquirá, 2550 m, 16 June 1983, L . J . Rubiano 436– B ( B) . VENEZUELA. Aragua: Colonia Tovar, 1900 m, 20 March 1921, H . Pittier 9283 ( VEN) . Bolívar: Near Río Surukún, South Caroní river , 940 m, 30 March 1988, V . Marcano 1341 ( VEN) . Falcón. Sierra de San Luis, between Curimagua and La Tabla, 1400 m, López-Figueiras 19341 ( MERF); Sierra de Barbacoas, Paramo de los Népes , 2400–2500 m, 31 January 1980, López-Figueiras 22154 ( MERF) . Lara: Los Potreritos, near Paramo Los Nepes, 2100–2600 m, López-Figueiras 17158 ( MERF); between Fundo Buenos Aires and Humocaro Alto , 1700–1800 m, 29 March 1979, López-Figueiras & M . Hale 19643, 19653 ( MERF, US) ; Serranía de Ziruma or Empalado, around Cerro Azul, 1890 m, 17 August 1979, López-Figueiras & R . Wingfield 21622 ( MERF). Mérida: Laguna Coromoto, 3400 m, 13 January 1957, Vareschi 5931 ( VEN); Serranía La Culata , Monte Zerpa , 2500–3800 m, 28 February 1993, V . Marcano 4779, 4780, 4781 ( B, MERF); Las Playitas, near Bailadores , 2300 m, 14–April 1992, V . Marcano 4498, 4499, 4500 ( MERF); slopes next to El Morro, 2000 m, 18 February 1976, López-Figueiras & M . Keogh 12317 ( MERF); La Carbonera, Mérida-La Azulita road, 2200 m, López-Figueiras, T . Ahti & P . Jørgensen 17917 ( MERF); Cloud forest, Raiz de Agua, Parque Sierra Nevada, 2400 m, 18 May 2020, V . Marcano 20–56 (herb. V. Marcano, VEN). Miranda: Los Guayabitos , 1350 m, 20 June 1954, V . Vareschi 3392 ( VEN). Táchira: Near Villa Paez, Delicias , 1900–2100 m, 13 April 1992, V . Marcano, 6116– C ( MER) ; Paramos, 3000 m, 3 October 1956, V . Vareschi 5718 ( VEN); Near Laguna García , 1900 m, 25 February 1981, López-Figueiras & H . Rodriguez 25439 ( MERF); Valle del Tamá, between Betania and San Vicente, 2750 m, 5 January 1983, López-Figueiras 29924 ( MERF); Parque Nacional El Tamá, to South Betania , 2750 m, 15 April 1992, V . Marcano 4800 ( MERF); Pregonero , near La Grita, 1750–1800 m, 21 April 1993, V . Marcano 4861, 4862, 4863 ( B, MERF); Paramo La Negra , 9 September 1955, V . Vareschi 4333 ( VEN). Trujillo. Fila de San Isidro , SO Paramo de Cendé , 2300 m, 21 August 1981, Lopez-Figueiras & H . Rodriguez 26354 ( MERF); Las Palmas, road Carache-Agua de Obispo , 2300 m, 20 August 1981, Lopez-Figueiras & H . Rodriguez 26295 ( MERF), Lopez-Figueiras, J . Cuatrecasas & H . Rodriguez 23400 ( MERF) .

0. Ramalina complanata (Swartz) Acharius

Lich. Univ.: 599 (1810). – Lichen complanatus Swartz in Acharius, K . Vet. Nya. Handl. 18: 290 (1797). – Parmelia complanata (Swartz) Acharius, Meth. Lich. : 259 (1803). Type :― JAMAICA. India occidentalis: Swartz (holotype S; isotypes BM, H – ACH 1821 ) .

Ramalina denticulata (Eschweiler) Nylander, Acta Soc. Scient. Fenn. 7: 434 (1863).– Parmelia denticulata Eschweiler in Mart., Fl. Bras. 1 (1): 221 (1833).

Thallus caespitose, corticolous, more or less erect, pale yellow, up to 8 cm long, densely branched. Branches solid, more or less flattened, 0.9–3.2 mm wide, upper surface slightly sinuose, more or less rugose (Fig. 17). Soralia absent. Pseudocyphellae laminal, marginal, punctiform, tuberculate (Fig. 12). Chondroid tissue weakly to distinctly cracked, discontinuous. Pycnidia not seen. Apothecia subterminal to terminal, concave, not spurred. Ascospores 1–3 septate, long-ellipsoid, 10–14 x 3.5–4.5 µm.

Chemistry (TLC, HPTLC): This species presents 9 strains which are summarized in the Table 6. Strain 1. Stictic, salazinic, protocetraric and norstictic acids. Strain 2. Homosekikaic, 4ˊ– O –demethylsekikaic and salazinic acids (Vareschi 2834). Strain 3. Protocetraric acid (Vareschi 424–III, 2785, 4395, 7760). Strain 4. Salazinic, protocetraric and stictic (tr.) acids (Vareschi 5257, 6213). Strain 5. Salazinic acid (Morales 144–89, 145–89; Vareschi 3447, 5256). Strain 6. Salazinic and protocetraric (tr.) acids (Morales 2). Strain 7. Homosekikaic, sekikaic, salazinic, consalazinic and protocetraric acids (Marcano 146–92). Strain 8. Homosekikaic, sekikaic and salazinic acids (Morales 142–89, 153–89, 156–89; Sipman & Velosa 32765, 32728). Strain 9. Norstictic (tr.) and protocetraric acids (Vareschi 3208– D).

Substances (acids) Str.1 Str.2 Str.3 Str.4 Str.5 Str.6 Str.7 Str.8 Str.9 Total 4ˊ– O –demethylsekikaic X

Consalazinic X

Homosekikaic X X X

Norstictic X X(tr.) Protocetraric X X X X(tr.) X X

Salazinic X X X X X X X

Sekikaic X X

Stictic X X(tr.)

Usnic X X X X 4 Total 5 4

Ecology and distribution: This species is corticolous, growing on Citrus , Clusia and other trees and shrubs in rainforests and paramo from 1200–3800 m. It is known from the USA, Bermuda, the Caribbean ( Jamaica, Puerto Rico and Santo Domingo) and South America (the Galápagos Islands, Brazil, Paraguay, Colombia and Venezuela). In the USA, the Caribbean and the Galápagos Islands, R. complanata is a distinctly coastal species.

Remarks: Ramalina complanata is a very problematic species, the circumscription of which many authors have disagreed on ( Nylander 1870, Krempelhuber 1876c, Vainio 1890, Zahlbruckner 1902, Malme 1934, Jones 1973). For instance, R. denticulata ( Eschweiler 1833: 221) Nylander (1863: 434) , a species described from Brazil, was occasionally confused with R. complanata in the literature, because it agrees well morphologically and chemically with R. complanata . Hence, it is considered to be a synonym of R. complanata in this paper. Furthermore, the specimens examined of R. complanata from Venezuela and Colombia exhibited several morphotypes, which mainly varied in the width of the lobes with tubercles/papillae, and in the chemical composition. The spores of the Colombian and Venezuelan specimens are smaller than for specimens from Brazil ( Kashiwadani & Kalb 1993, Gumboski 2016), which are reported to be short-fusiform (12–16 x 4–5 µm). Kashiwadani & Kalb (1993) recognized two morphotypes of R. complanata in Brazil: one with narrow (up to 1 mm wide) and subcanaliculate branches, and a second with broad (1.5–3 mm wide) and more or less flat branches. Some Brazilian specimens, however, can be hard to place within these two types ( Kashiwadani & Kalb 1993).

In Colombia and Venezuela, the major substances found in R. complanata are salazinic and protocetraric acids, with sekikaic acid and related substances and other β-orcinol depsidones as accessory substances (Table 6). In the Galápagos Islands ( Aptroot & Bungartz 2007) this species has only one chemical strain characterized by salazinic acid, while in Brazil ( Kashiwadani & Kalb 1993) the species exhibits two strains: one with the divaricatic and salazinic acids, and another with salazinic acid.

Ramalina complanata is rare in Colombia ( Sipman et al. 2008) but common in Venezuela. In northern South America, R. complanata could be confused with R. africana and R. caracasana , but R. africana has spurred apothecia and contains the sekikaic acid complex whereas R. caracasana has fusiform spores (17–20 x 5–6 µm) and contains protocetraric acid. Most specimens here treated are reported from high-elevation rainforest and paramo whereas other are reported from coastline (e.g. Falcón, Vareschi 3447). In the USA, the Caribbean and Galápagos Islands, R. complanata appears as a coastal species occupying dry zones. Other Ramalina species likewise occupying wide altitudinal and ecological ranges are R. usnea and R. cochlearis . Therefore, ecological, morphological and chemical differences observed between specimens of R. complanata from Colombia, Venezuela, including Brazil, could suggests the occurrence of several varieties.

Specimens examined: COLOMBIA: Cauca: Popayan, Los Robles, campus of Fundacion Universitaria , c. 15 km towards Timbio , 1750 m, 29 May 1986, H . Sipman & R . Velosa 32728, 32765 ( B) . VENEZUELA: Aragua. Rancho Grande , 7 July 1956, V . Vareschi 5256, 5257 ( VEN) . Falcón: Near Tucacas , 10 m, 4 July 1954, V . Vareschi 3447 ( VEN) . Mérida: Near Mérida City, Arenal de Lourdes , 17 January 1992, V . Marcano & A . Morales 153 ( MER); Santo Domingo, Paramo de Mucubají , 3500–3800 m, March 1992, V . Marcano 146–92 ( MER); Paramo de la Culata, 3500 m, January 1992, V . Marcano 156–92 ( MER); near Chiguará, Los Topes, 1200 m, A . Morales 142, 144, 145 ( MER); San Juanito, Chiguará , A . Morales 2 ( MER); Paramo La Negra, 2900–3100 m, 15 Abril 1954, Vareschi 3208–d ( VEN) . Miranda. Gavilán, near Los Guayabitos, 23 February 1953, Vareschi 2785 ( VEN); Los Guayabitos, 6 November 1955, Vareschi 4395 ( VEN); Los Guayabitos, 1350 m, 9 September 1951, V . Vareschi 424–III ( VEN); Los Guayabitos, 2 March 1953, Vareschi 2834 ( VEN); Los Guayabitos, 8 January 1956, Vareschi 6213 ( VEN) .

. Ramalina crispata A. Morales & V. Marcano

The Bryologist 97: 28 (1994) . Type:― VENEZUELA. Mérida: Chiguará, Los Topes , 1200 m, 16 April 1988, A. Morales 200 (holotype MERF!) .

Thallus corticolous, more or less erect, rigid, fragile, intricately and irregularly branched, up to 1 cm high, pale yellow or gray-green. Branches solid, palmate to sublinear, 1–2(–3) mm broad, flat. Pseudocyphellae punctiform (Fig. 13), raised on tubercles, abundant on under and upper sides. Soralia absent. Cortical tissue paraplectenchymatous 14–18 μm thick, hyphae thick-walled, peripheral chondroid tissue forming discontinuous layers. Medulla very dense. Pycnidia not seen. Apothecia rare, lateral, subapical or apical, disc perforate, 2.6–4.0 mm diameter, margin pseudocyphellate, densely laciniate. Ascospores short-fusiform, 11–12 x 3.5–4 μm.

Chemistry (TLC, HPTLC): Salazinic acid.

Ecology and distribution: Ramalina crispata grows on shrubs and trees in a secondary forest at 1200–3200 m. It is known only from the Venezuelan Andes.

Remarks: Ramalina crispata is morphologically similar to R. africana . Both species have punctiform pseudocyphellae raised on tubercles. However, R. crispata but differs in having an intricately branched, smaller thallus (up to 1 cm high); flat branches with abundant pseudocyphellae on both sides; apothecia with laciniate margins; and salazinic acid as a chemical constituent. Ramalina africana is characterized by linear, canaliculate, flat branches, that are 3–5 (–8) cm long, often with short secondary branchlets; abundant pseudocyphellae on the underside; and sekikaic acid as the principal chemical constituent.

Additional specimens examined. VENEZUELA. Táchira. Between Betania and Villa Páez, near Tamá National Park , 2800–3200 m, 1992, V . Marcano 5169 ( MERF) .

. Ramalina cumanensis Fée (Fig. 32)

Essai sur Les Cryptogames : 135 (1824) . Type:― VENEZUELA. Sucre: Cumana, Humboldt & Bonpland (holotype P).

Thallus corticolous, caespitose, dichotomously branched, up to 6 cm long. Branches canaliculate, smooth, pale yellow. Pseudocyphellae continuous, marginal and laminal. Soralia not seen. Pycnidia not seen. Apothecia abundant, disc concave or plane. Ascospores 1–septate, curved or ellipsoid, 8–12 x 4–5 µm.

Chemistry (TLC, HPTLC): Nil (López-Figueiras 12126; Vareschi 535–A, 4322, 7151, 8697; Marcano 7270, Marcano & Galiz 193; Sipman et al. 34115; Cleef 2290, 4849–D; Sipman & Valencia 10301; Flørschutz 3618–B), traces of usnic acid only (Martinez et al. 193; Sipman & Aguirre 27454; Sipman & Reyes 34456; Vareschi 7481, 8697, 9822), usnic acid and triterpenoid (Aguirre & Sipman 5563).

Ecology and distribution: This species is found growing on Podocarpus branches, Clusia , Pithecellobium dulce , Hypericum sp. and Fraxinus sp. ( Colombia) ; on shrubs, rocky banks; dead tree trunks in pastures; and on Weinmannia , Vallea stipularis and Miconia spp. in submontane, montane forests and paramo at 1000–3765 m (Fig. 30). It is known only from northern South America ( Colombia and Venezuela). Nevertheless, Krempelhuber (1876c) reported a specimen growing on bark and identified as R. cumanensis , collected by Glaziou (no. 1855) in the Brazilian Amazonas, near Rio Janeiro.

Remarks: Ramalina cumanensis is very common in Colombia and Venezuela ( Marcano et al. 1996, Sipman et al. 2008, Marcano & Castillo 2020). It could be confused with R. puiggarii because both species have continuous, marginal pseudocyphellae but lack soralia. However, R. puiggarii has multiseptate, longer fusiform spores (18–24 x 4– 4.5 µm). Krog & Swinscow (1976) considered R. cumanensis to be a growth form of R. celastri . Both species are rather common in northern South America. However, R. celastri has lanceolate, flat branches and short-fusiform ascospores (13–17 x 3.5–5 µm). Ramalina cumanensis has been confused in the literature with R. calicaris (L.) Fr., which occurs in northern Europe ( Krog & James 1977; see excluded species, below) and produces sekikaic and homosekikaic acids. Ramalina cumanensis , however (like R. puigarii and R. celastri ) always lacks medullary metabolites.

Specimens examined: COLOMBIA: Boyaca: Paramos to NW Belen, vereda San Jose de la Montaña , Altos de las Cruces , 3765 m, 6 March 1972, A . Cleef 2290 ( B); Municipio Cucaita, c. 10 km W of Tunja along road to Villa de Leiva , 2950 m, 12 July 1986, H . Sipman & O . Reyes 34456 ( B) . Huila: Municipio La Plata, E-side of Cordillera Central, vereda La Candelaria , Finca Merenberg , 2400 m, 30 September 1984, J . Aguirre & H . Sipman 5726 ( B, COL) . Cundinamarca: Valle Teusacá , c. 5 km SW La Calera, 2850 m, 9 July 1972, A . Cleef & M . van Rens de Cleef 4849–D ( B); between Bogota and Fusagasuga , c. 5 km SW of roadtoll, 2700 m, 31 January 1979, H . Sipman & H . Valencia 10301 ( B, COL, U); between Cogua and San Cayetano , near Laguna Seca, 3450 m, 13 September 1972, P . A . Florschütz 3618– B ( B) ; Municipio Supatá Alto El Tablazo, along track from radar station to Supatá , 3200 m, 20 October 1988, H . Sipman & J . Aguirre 27454 ( B) . Antioquia: Municipio El Retiro, along road Medellín-La Ceja, between Las Palmas and El retiro, Hacienda Fizebad , entrance of reconstructed farm, 2050 m, 6 July 1986, H . Sipman , M . Escobar & J . Rubiano 34115 ( B) . Risaralda: W-slope of Cordillera Central, Municipio Santa Rosa de Cabal, along track from Termales de Santa Rosa to Termales Antiguos , 2000 m, 19 September 1984, J . Aguirre & H . J . Sipman 5563 ( B). Valle: Cali, University of Valle , 1000 m, 1986, J . Rubiano s/n– B ( B) . VENEZUELA: Mérida: Mucumpís road, Paramo de Mifafí, La Culata , 2800–3200 m, October 1993, V . Marcano 7270 ( MER); Sierra de Santo Domingo, Mucubají lake , 3500 m, March 1992, V . Marcano s/n ( MER); Sierra de Santo Domingo, Mucubají lake , 3500 m, 19 May 1993, D. Martinez , V . Marcano & L . Galiz 193 ( MER); Sierra Nevada de Mérida, Las Escaleras, road Mucuchíes – El Carrizal , 2800–3100 m, 6 April 1979, M . Lopez-Figueiras & M . Hale 20526 ( MER); near Mucubaji Lake , 9 september 1958, V . Vareschi 7151 ( VEN); Near Chachopo, 3200 m, 31 November 1977, V . Vareschi 8697 ( VEN); Paramo La Negra , 9 September 1955, V . Vareschi 4322 ( VEN); near Timotes, 3200 m, 25 September , 1979, V . Vareschi 9822 ( VEN) . Miranda: Los Guayabitos , El Volcán, 1420 m, 28 December 1951, V . Vareschi 535– A ( VEN) . Trujillo: Near la Puerta , 2100 m, 28 July 1960, V . Vareschi 7481 ( VEN); Paramo de Tuñame, 3320 m, 4 February 1976, López Figueiras 12126 ( MERF, VEN) . Táchira: Garcia lake, near Pregonero , 14 April 1992, V . Marcano s/n ( MER): Paramo La China , bosque montano siempre verde y bajo seco, vegetación arbustiva, 2120 m, L . Castillo & V . Marcano 209 ( MER); Laguna La Verdosa , muscícola, subparamo, paramo, 3250–3300 m, L . Castillo & V . Marcano 042 ( MER) .

. Ramalina dendriscoides Nylander

Flora, Jena 59: 412 (1876) . Type :― CUBA. C. Wright , Lich. Cub. Ser. 2: 738 (lectotype H –NYL 37025, fide Krog & Swinscow 1976; isotypes FH, G) .

Thallus corticolous, shrubby to intrincately branched, up to 10 cm high, yellowish. Branches solid, flat or more or less terete, surface smooth sorediate, apical on short lateral branchlets, 0.6–1.2 mm wide. Pseudocyphellae elliptic. Soralia scattered, terminal in secondary branches. Soralia isidiform not seen. Chondroid tissue cracked, discontinuous. Medulla white, compact. Pycnidia not seen. Apothecia marginal, laminal, abundant.Ascospores elliptic, 10–12 x 3–3.5 µm.

Chemistry (TLC, HPTLC): This species presents 7 strains which are summarized in the Table 7. Strain 1. Salazinic and consalazinic (tr.) acids (Vareschi 5255, 6019). Strain 2. Salazinic and fumarprotocetraric acids (Dennis 1538). Strain 3. Salazinic acid and UV + rose pigments (Vareschi 6204–B, Vareschi 6205, Mägdefrau 673). Strain 4. Salazinic acid (Vareschi 5260, 6204–A, 6214; Dennis 1513–B). Strain 5. 4ˊ– O –demethylsekikaic, stictic and salazinic acids (Vareschi 3208). Strain 6. Salazinic, succinprotocetraric and protocetraric acids (Vareschi 6199). Strain 7. Sekikaic, ramalinolic and 2ˊ– O –methylsekikaic acids (Sipman et Escobar 34050).

Ecology and distribution: Growing on the tree branches and shrubs in submontane forest, montane forest and paramo; in Colombia epiphytic on Quercus ; 400–3100 m. Known from East Africa ( Kenya, Tanzania), México, Florida and Texas ( USA), Cuba, Puerto Rico and South America ( Brazil, Colombia and Venezuela).

Remarks: Ramalina peruviana and R. tenella can be confused with R. dendriscoides , but R. peruviana lacks apical and lateral pseudocyphellae, has thinner branches (0.1–0.4 mm wide) and contains the sekikaic acid complex as major medullary substances, while R. tenella has a shorter and more delicate thallus (up to 4 cm high), with more or less flattened branches, predominantly dichotomous branching and produces salazinic acid only. The latter species also lacks accessory substances ( Krog & Swinscow 1976, Stevens 1987). Ramalina dendriscoides contains salazinic acid as the major substance together with sekikaic, ramalinolic, 4ˊ– O –demethylsekikaic and 2ˊ– O –methylsekikaic acids and substances in the protocetraric acid complex as accessory compounds (Table 7). In Brazil ( Kashiwadani & Kalb 1993, Gumboski 2016) this species has only one chemical strain characterized by salazinic acid and atranorin (traces). In East Africa ( Krog & Swinscow 1976) it contains salazinic acid only.

Specimens examined: COLOMBIA: Antioquia: Municipio Santa Rosa de Osos, vereda Santa Rosa, Vallecitos , 2550 m, 5 July 1986, H . Sipman & M . Escobar 34050 ( B) . VENEZUELA: Aragua: Rancho Grande National park , 7 July 1956, V . Vareschi 5255, 5260 ( VEN) . Carabobo: Near Puerto Cabello , 6 April 1958, K . Mägdefrau 673 ( VEN). Dto. Federal: near to Chichiriviche, 6 J 1958 , Dennis 1538 ( VEN) . Miranda: Cortada del Guayabo, 1200 m, 9 July 1958, Dennis 1513– B ( VEN); Los Guayabitos, El Volcán, 1400 m, 5 January 1957, V . Vareschi 6199, 6204, 6205 ( VEN); Los Guayabitos, 1400 m, 8 January 1956, V . Vareschi 6214 ( VEN) . Mérida: Paramo La Negra , 2900–3100 m, 15 April 1954, V . Vareschi 3208– A ( VEN) . Sucre: Vecinity Los Pocitos, above Manacal, Northwest Irapa , 400 m, 11 July 1972, Dumont 6019 ( NY, TRTC, VEN) .

Figures

FIGURES 4– 5. Upper surface of R. venezuelensis showing pseudocyphellae as observed by SEM. 14. Lateral pseudocyphellae and small pore-like structures, scale = 15 mm. 15. Lateral pseudocyphellae, scale = 30.5 µm.