Ramalina escorialensis V. Marcano & A. Morales, 1994

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 : 47-53

5. Ramalina escorialensis V. Marcano & A. Morales

Tropical Bryology 9: 194 (1994) . Type:― VENEZUELA. Mérida: Morro de San Francisco, Fila El Escorial , near Escaguey , 3500 m, 21 March, 1992, V. Marcano & Raul Vidal 4998 (holotype MERF!, isotype VEN!) .

Thallus corticolous, erect, delicate, pale yellowish, up to 6.5 cm height. Branches shiny, flat, 0.4–0.8 mm broad, having short and numerous divisions. Pseudocyphellae lateral, circular, sometimes becoming sorediate. Soralia helmet-shaped, abundant, subapical. Cortical tissue 18–40 μm thick. Peripheral chondroid tissue discontinuous, 50–70 μm thick. Medulla white, compact. Pycnidia and apothecia not seen.

Chemistry (TLC, HPTLC): Sekikaic acid and cryptochlorophaeic acid.

Ecology and distribution: Ramalina escorialensis grows on shrubs in paramo; 2500–3000 m. It is known only from the Venezuelan Andes.

Remarks: The morphological and anatomical features of this species have been described in detail by Marcano & Morales (1994c). Ramalina escorialensis is very similar to R. cochlearis as both species are characterized by helmetshaped soralia. However, they are readily distinguished by the anatomy and chemistry of the thalli: R. escorialensis has discontinuous chondroid tissue; flat branches with very short, numerous, divergent branchlets, and cryptochlorophaeic and sekikaic acids. Ramalina cochlearis , by contrast, has canaliculate branches, without branchlets, continuous chondroid tissue and sekikaic acid (and related substances), but never contains cryptochlorophaeic acid.

Additional specimens examined: VENEZUELA: Mérida: Raiz de Agua , Parque Sierra Nevada, near to La Mucuy Alta, 2500 m, 16 May 2020, V, Marcano 20–48 (herb. private V. Marcano) .

. Ramalina holstii Krog & Swinscow

Norw. J. Bot. 22: 275 (1975). Type:― KENYA. Eastern province: Machakos District, lava flow 5 km NW of Kibwezi, Krog & Swinscow 3 K 23 /145 (holotype O, isotype BM).

Thallus corticolous, densely and intrincately branched, pale gray or yellowish, up to 4.5 cm height. Branches solid, flat or broadly canaliculate, 0.3–1.5 mm broad, apically compressed, finely divided. Soredia not seen. Pseudocyphellae marginal, tuberculate. Pycnidia not seen. Apothecia frequently laciniate, with branchlets having soralia in the margins. Ascospores 1–septate, ellipsoid, 10–12 x 3–3.5 µm.

Chemistry (TLC, HPTLC): Strain 1. Salazinic, succinprocetraric, protocetraric and consalazinic acids, triterpenoids, (Dumont 6018). Strain 2. Salazinic acid (Vareschi 6162). Strain 3. Salazinic and protocetraric acids (Vareschi 6211). Strain 4. Salazinic, protocetraric and consalazinic (tr.) acids (Vareschi 3497).

Ecology and distribution: Ramalina holstii grows on trees and shrubs, in both primary rainforests and disturbed forests, at 400–1200 m. It is known from East Africa ( Kenya, Tanzania) and northern South America ( Venezuela).

Remarks: Ramalina holstii can be confused with R. africana , but the latter species has laminal pseudocyphellae, broader spores (4–5 µm wide) and the sekikaic acid complex as major substances. Ramalina crispata is also very similar as it also has laciniate apothecia, punctiform pseudocyphellae and salazinic acid as the major substance as well as similar-sized spores (11–12 x 3.5–4 µm); R. crispata , however, differs in possessing abundant pseudocyphellae on both its lower and upper surfaces. The thallus and branch sizes compare well with authentic material examined of this species, received from Dr H. Krog. Ramalina holstii exhibits four chemical strains, all of which contain salazinic acid as the major substance. Various β-orcinol depsidones occur as accessory substances. In East Africa ( Krog & Swinscow 1975, Swinscow & Krog 1988), this species has also salazinic acid and further contains cryptochlorophaeic acid. Other Ramalina species co-occurring in East Africa and northern South America are R. africana , R. aspera , R. asperula , R. celastri , R. calcarata , R. dendriscoides , R. peruviana , R. pusiola , R. reducta , and R. tenella ( Krog & Swinscow 1975, Swinscow & Krog 1988, Marcano & Morales 1993b, Marcano & Morales 1994b).

Specimens examined. Venezuela: Sucre: Vicinity of Los Pocitos, near Manacal , NW Irapa, 400 m, 11 July 1972, Dumont et al. 6018 ( NY, TRTC, VEN) . Miranda: La Mata, near Los Guayabitos, 1 August 1954, V . Vareschi 3497 ( VEN); Los Guayabitos, 8 January 1956, Vareschi 6211 ( VEN); Los Guayabitos , 1200 m, 1 January 1957, Vareschi 6162 ( VEN)

. Ramalina lopezii A. Morales & V. Marcano

The Bryologist 97: 29 (1994) . Type:― VENEZUELA. Trujillo: East Paramo del Jabón, near Carache , 2900–3100 m, 22 August 1981, Lopez-Figueiras & H. Rodriguez 26552 (holotype MERF!) .

Thallus corticolous, erect, pale yellow or pale green, up to 5 cm high, with distinct branches arising from a common holdfast. Branches simple or sparingly furcate, mostly terete towards the base, more or less flattened above (Fig. 1), 1.3–1.6 mm broad, surface foveolate, undulate, occasionally with furrows. Pseudocyphellae not seen. Peripheral chondroidal tissue discontinuous. Medula compact. Pycnidia frequent, apical, blackened. Pycnospores rod-shaped, 3–3.5 x 0.8–1 µm. Apothecia lateral, subapical, occasionally terminal, sessile, disc convex, up to 4.2 mm diameter (Fig. 1). Ascospores 1–septate, straight or slightly curved, 10–11 x 3.5–4 µm.

Chemistry (TLC, HPTLC): Stictic and norstictic acids.

Ecology and distribution: Ramalina lopezii is known only from the type locality where it grows on shrubs in the paramo. It is known only from the Venezuelan Andes.

Remarks: Ramalina lopezii is a very rare species known only from the type locality in Venezuela. Morphologically, R. lopezii is very similar to R. cuspidata ( Acharius 1810: 605) Nylander (1870: 158) and R. siliquosa ( Hudson 1762: 460) A. L. Smith (1918: 172), two widespread European lichens ( Krog & James 1977, Sheard 1978a,b, LaGreca et al. 2020). Ramalina lopezii differs from R. cuspidata in having a foveolate surface; however, both species share blackened pycnidia, a pale yellow or pale green thallus, a shiny surface, and norstictic and/or stictic acids. Ramalina siliquosa , on the other hand, has a foveolate surface much like R. lopezii , but is distinguished by its glaucous, matte surface and the presence of hypoprotocetraric, protocetraric and/or salazinic acids. There are vast ecological differences between these species as well: R. lopezii is corticolous at high elevations, while R. cuspidata and R. siliquosa both inhabit rocky shores. Recent molecular evidence reveals that these two taxa are not phylogenetic sister species ( LaGreca et al. 2020).

. Ramalina maegdefraui V. Marcano & A. Morales sp. nov.; Mycobank #838605 (Fig. 34)

Thallus corticolus, erectus. Laciniae intricate vel irregulariter ramulosae, complanatae. Pseudocyphelliis punctiformis. Soralia lateralia vel laminalia. Pycnidia non visa. Apothecia lateralia. Acida usnicum, salazinicum et protocetraricum continens. Type:― VENEZUELA. Lara: 30 km to the West of Barquisimeto , 28 March 1958, K . Mägdefrau 548–a (holotype VEN) .

Thallus corticolous, rigid, erect or partly decumbent, fragile, up to 2.5 cm high, intricately and irregularly branched, with several laciniae arising from a common holdfast. Branches solid, flattened, with a rugose or wrinkled surface, pale yellow, 0.5–2 mm broad. Pseudocyphellae punctiform, flattened, raised on tubercles, abundant on lower and upper sides, soredia present. Soralia abundant, lateral and laminal, producing coarse granules, abundant. Cortical tissue paraplectenchymatous, 150–180 µm thick, peripheral tissue prosoplectenchymatous, 250–300 µm thick, medulla dense. Pycnidia not seen. Apothecia subapical to apical, marginal, abundantly spurred, with rugose or wrinkled amphithecium, disc flat or concave, 0.3–1 mm diameter; thalline exciple ridged. Ascospores 1–septate, colorless, short ellipsoid, 10–12 x 3–4 µm.

Chemistry (TLC, HPTLC): Salazinic and protocetraric acids (Mägdefrau 548–A).

Ecology and distribution: Ramalina maegdefraui is known only from the type locality, where it grows on shrubs ( Peireskia sp. ) in moist forests at 1000 m.

Remarks: Ramalina maegdefraui is distinguished by the complanate, sorediate thallus, flattened, tuberculate pseudocyphellae (Fig. 10) and a smooth upper surface. This species is morphologically and chemically very similar to R. mirandensis but differs in having soralia and a flattened thallus. Both species are characterized by punctiform pseudocyphellae, short ellipsoid spores and salazinic and protocetraric acids as chemical compounds. Ramalina maegdefraui could initially be confused with R. africana , R. crispata and R. holstii , but all these species lack soralia.

The name of the species is in honor of Dr Karl Mägdefrau, who collected the type material.

. Ramalina microphylla A. Morales & V. Marcano

The Bryologist 97: 30 (1994) . Type:― VENEZUELA. Falcón: Península de Paraguaná, Fila Tausabana , between El Rodeo and Machuruca, 90 m, 12 October 1984, Lopez-Figueiras & R. Wingfield 31022– A (holotype MERF!, isotype VEN!) .

Thallus saxicolous, erect or sunpendulous, rigid, 0.8–1 cm long, unbranched or dichotomously to irregularly branched, with a number of short laciniae from a broad holdfast. Branches palmate or sublinear, 1.5–2.7 μm broad, with short conspicuous laminal striae along their entire length. Surface puckered or wrinkled. Pseudocyphellae lateral, linear to ellipsoid. Soralia subapical to apical, producing coarse granules, capitate or labriform. Cortical tissue paraplectenchymatous, 10–16 µm thick, peripheral chondroid tissue forming discontinuous layers. Medulla dense with imbedded chondroid strands. Pycnidia marginal or laminal, with pale, honey-coloured ostioles. Pycnoconidia uniformly rodshaped, 4–4.5 μm x 0.6–0.9 μm. Apothecia not seen.

Chemistry (TLC, HPTLC): Salazinic acid only (Lopez Figueiras et al. 32498).

Ecology and distribution: Ramalina microphylla is a very rare species known only from the type locality where it grew abundantly on exposed igneous rocks in a dry forest.

Remarks: Ramalina microphylla somewhat resembles R. lopezii . Both are characterized by rigid, erect, branched thalli and medullary β-orcinol depsidones. Ramalina lopezii , however, is corticolous; lacks soralia and pseudocyphellae; bears blackened apical pycnidia; and produces stictic and norstictic acids. Ramalina dicipiens Montagne from the Canary Islands (Krog & Østhagen 1980). However, it exhibits smaller thalli with lateral soralia and lacks pseudocyphellae, whereas R. decipiens has longer thalli [5–12(–18) cm], longitudinal pseudocyphellae and lacks soralia.

Additional specimens examined: VENEZUELA: Falcón: Península de Paraguaná, Fila Tausabana, 1984, Lopez- Figueiras & R. Wingfield 31022–B and C (IUTC, MERF); Peninsula de Paraguaná, Fila Tausabana, El Rodeo – Muchuruca road, 90–100 m, M. Lopez-Figueiras, R. Wingfield & A. Morales 32498 (MER).

0. Ramalina mirandensis V. Marcano & A. Morales sp. nov.; Mycobank #838606

Thallus corticolus, erectus. Laciniae irregulariter ramulosae, complanatae. Pseudocyphelliis punctiformis, laminalis. Soralia non visa.

Pycnidia non visa. Apothecia laminalia. Sporae ellipsoideae. Acida salazinicum et protocetraricum continens. Type:― VENEZUELA. Miranda: Near La Mata , 1100 m, 1 August 1954, V . Vareschi 3484 (holotype VEN) .

Thallus corticolous, flattened or subterete, up to 0.9 cm long, dichotomously or irregularly branched. Upper branches terete. Soralia absent. Pseudocyphellae tuberculate, elliptical, laminal. Chondroid tissue prosoplectenchymatous, continuous, never cracked. Medulla dense. Pycnidia not seen. Apothecia abundant, terminal, margiinal, disc plane or concave. Ascospores 1–septate, short-ellipsoid, 8–9 x 3.5–4 µm.

Chemistry (TLC, HPTLC): Salazinic, stictic (tr.) and protocetraric acids (Vareschi 3484).

Ecology and distribution: This species is known only from the type locality where it was collected growing on shrubs in disturbed forests at 1000–1200 m.

Remarks: Ramalina mirandensis seems to be a very rare species in northern South America ( Marcano 2020). Chemically it produces β-orcinol depsidones only. This species could be confused with R. complanata (strain 11, salazinic, stictic, and protocetraric acids), but the latter species has multiseptate, long-ellipsoid ascospores (10–14 x 3.5–4.5 µm). Furthermore, R. complanata has flat (more or less canaliculate) branches, whereas R. mirandensis has flattened to subterete thalli with terete upper branches. In northern South America, R. mirandensis could be confused with R. africana and R. caracasana , but R. africana has short, spurred apothecia and contains the sekikaic acid complex whereas R. caracasana has fusiform spores (17–20 x 5–6 µm) and contains protocetraric acid ( Marcano and Morales 1994b).

The name of the species refers to the type locality.

. Ramalina morrocoyensis A. Morales & V. Marcano (Fig. 35).

Mem. Soc. Ci. Nat. La Salle 55: 35 (1995) . Type:― VENEZUELA. Falcón: Parque Nacional Morrocoy, Playa Azul , sea level, 27 March 1991, A. Morales 316 (holotype MERF!, isotype VEN!) .

Thallus corticolous, erect, more or less decumbent, up to 2 cm high, 0.5–1 mm wide, with a number of short branches originating from a broad holdfast. Branches solid, complanate, irregularly branched, 0.5–1 mm wide, 220–240 μm thick, apices rod-shaped. Pseudocyphellae absent. Soralia absent. Chondroid layer prosoplectenchymatous, 50–70 μm thick, forming a continuous cylinder. Medulla dense. Pycnidia not seen. Apothecia numerous, marginal, subterminal, shortly pedicellate, disc convex, 0.5–1.5 mm in diameter. Ascospores 1–septate (occasionally multiseptate), fusiform, 17–19 x 3.5–4 µm.

Chemistry (TLC, HPTLC): Strain 1. Sekikaic and boninic acids (Steyermark & Manara 110846). Strain 2. Divaricatic acid (Steyermark & Manara 110308).

Ecology and distribution: This species grows on the bark of shrubs ( Suriana maritima L., Surianaceae ) in dry forests in exposed, coastal areas at 0– 2 m. It is known only from northeast Venezuela (Fig. 30).

Remarks: Ramalina morrocoyensis is characterized by a corticolous, more or less decumbent thallus, a smooth surface, a lack of pseudocyphellae and soralia, complanate branches and fusiform spores ( Morales & Marcano 1995). Ramalina morrocoyensis can be confused with R. tenaensis , but the latter has contorted branches a verruculose surface, and grows in Colombian submontane forests. Ramalina morrocoyensis is found associated with R. paradisensis in similar ecological niches, along the west coast Venezuela. Chemically and anatomically R. morrocoyensis is very similar to R. paradisensis , but the latter has branches with furcate, terete apices as well as pseudocyphellae and soralia.

Additional specimens examined. Venezuela: Falcón: Cayo Borracho, NE Chichiriviche, 1–2 m, 28 August 1974, J . Steyermark & B . Manara 110308 ( MO, VEN); Cayo Boca Seca, Northern Boca Grande , 1 m, 29 August 1974, J . Steyermark & B . J . Manara 110846 ( VEN) .

. Ramalina paradisensis A. Morales & V. Marcano (Fig. 36)

Mem. Soc. Ci. Nat. La Salle 55: 38 (1995) . Type:― VENEZUELA. Falcón: Parque Nacional Morrocoy, Playa Azul , sea level, 27 March , 1991, A. Morales 315 (holotype MERF, isotype VEN) .

Thallus corticolous, erect, dichotomously and intricately branched, 3–6 cm long, 1–2 mm wide, pale yellow. Branches solid, fragile, complanate, 0.5–0.6 mm wide, 240 μm thick; apices furcate, terete. Pseudocyphellae marginal, linear, punctiform, elliptic. Soralia laminal, forming coarse granules. Chondroid layer prosoplectenchymatous, 40–78 μm thick, forming a continuous cylinder, algal layer 9–14 μm thick. Pycnidia and apothecia not seen.

Chemistry (TLC, HPTLC): Sekikaic acid.

Ecology and distribution: Ramalina paradisensis is a very rare species known only from the type locality where it occurs abundantly on the bark of Suriana maritima L. ( Surianaceae ) in mangrove forests near the shore (Fig. 30). It is known only from Venezuela.

Remarks: Ramalina paradisensis is characterized by a corticolous, erect, intricately branched thallus; complanate branches, terminating in furcate terete apices; marginal pseudocyphellae; and laminal soralia ( Morales & Marcano 1995). Ramalina paradisensis could be confused with R. peruviana , as both species have mainly terete branches, soralia and contain sekikaic acid and/or aggregates. However, R. peruviana has a thinner thallus and bears marginal or lateral soralia. Ramalina paradisensis also resembles R. farinacea (Linnaeous 1753: 1146) Acharius (1810: 606) , a common and widespread species in western Europe and North America ( Krog & James 1977, Krog & Østhagen 1980, Brodo et al. 2001). However, R. paradisensis bears marginal pseudocyphellae and contains sekikaic acid, while R. farinacea lacks pseudocyphellae and contains β-orcinol depsidones.

. Ramalina peruviana Acharius

Lich. Univ.: 1599 (1810). Type :― PERÚ. Lagasta: (holotype H – ACH) .

Thallus corticolous, densely branched, up to 7 cm high, pale yellow, growing from a common holdfast. Branches solid, thin, terete, striate, surface smooth, 0.2–1.0 mm in wide, rarely with cylindrical branchlets. Pseudocyphellae concave, scattered, ellipsoid or elongate. Soralia lateral, laminal, punctiform, covering the tips of branches, producing soredia. Chondroid tissue not cracked, irregular, continuous. Medulla white, loose or compact. Pycnidia not seen. Apothecia marginal, laminal. Ascospores fusiform, 15–16 x 3–4.5 µm.

Chemistry (TLC, HPTLC): Strain 1. Homosekikaic, sekikaic, ramalinolic, 4ˊ– O –demethylsekikaic, boninic, succinprotocetraric (tr.) and protocetraric acids (Vareschi 3356). Strain 2. Protocetraric acid and triterpenes (Marcano 172–92). Strain 3. Hypoprotocetraric acid (tr.) (Aguirre & Sipman 5562). Strain 4. Homosekikaic, sekikaic, ramalinolic and boninic acids (Vareschi 4334).

Ecology and distribution: This species occurs on bark and rocks in rainforests, disturbed forests and paramo at 1400–3500 m. It is known from East Africa ( Kenya, Tanzania and Uganda), Australia, New Zealand, the Pacific Islands, Texas ( USA) and South America (the Galápagos Islands, Perú, Brazil, Paraguay, Colombia, Venezuela).

Remarks: Ramalina peruviana could be confused with R. dendriscoides and R. tenella , but these species have capitate soralia, ellipsoid spores (10–12 x 3–3.5 µm) and contain salazinic acid. Four distinct chemical strains are found in specimens of R. peruviana from Colombia and Venezuela. One strain (strain 1) contains sekikaic acid as major medullary substance together with associated meta-depsides and β-orcinol depsidones (protocetraric acid complex); strain 4 contains only sekikaic acid and its aggregates and strains 2 and 3 only contain the protocetraric acid complex. In North Atlantic Islands ( Aptroot & Schumm 2008), East Africa ( Krog & Swinscow 1976, Swinscow & Krog 1988), Australia ( Stevens 1987), New Zealand ( Galloway 2007), Perú and Brazil ( Kashiwadani 1987, Kashiwadani & Kalb 1993, Gumboski 2016) R. peruviana only contains sekikaic acid and its aggregates, and lacks substances in the protocetraric acid complex. The strains with substances in the protocetraric acid complex may suggest the occurrence of a South American evolutionary line that originated after the uplift of the Andean chains and separation of the Gondwanaland.

Specimens examined: COLOMBIA: Risaralda: Municipio Santa Rosa de Cabal , W-slope of Cordillera Central, 2000 m, 19 September 1984, J . Aguirre & H . Sipman 5562 ( B, COL) . VENEZUELA: Mérida: Miranda: El Volcan, 1400 m, 2 May 1954, Vareschi 3356 ( VEN) . Táchira: Southern Betania , near Villa Paez, 3500 m, 13 April 1992, V . Marcano 172–92 ( MER) .