Salpichlaena volubilis subsp. amazonica, 2019

b. subsp. amazonica G.G.Cárdenas & Tuomisto View in CoL , subsp. nov. — Fig. 3d View Fig , 4f, 9; Map 1 View Map 1

Salpichlaena volubilis subsp. amazonica can be recognised by its chartaceous to coriaceous lamina texture, clearly cartilaginous pinnule margins, long and wide pinnules and uniform abaxial lamina.

— Type: Tuomisto H, Moulatlet G, Ruokolainen K & Castro D 16904 ( holotype SP; GoogleMaps isotype INPA, GoogleMaps TUR (mounted on four sheets: 606298–606301)), Brazil, Amazonas, Carauari , c. 7 km south from the village of Vila Ramalho, Rio Juruá, S5°35' W67°29', 100 m, 14 June 2012. GoogleMaps

Etymology. The name refers to the geographical distribution of the subspecies in Amazonia.

Climbing and non-climbing fronds with chartaceous to coriaceous lamina texture, pinnae and pinnule margins clearly cartilaginous; scales on abaxial axes stick-like, formed by a long apical row of cells and usually two lateral smaller rows of cells, and sometimes also lanceolate scales with long appendices; stomata not (or very rarely) on white laminal protuberances. Non-climbing entire fronds up to 53 by 6.8 cm. Non-climbing 1-pinnate fronds 65–135 cm. Pinnae up to 3 pairs; apical one elliptic, lanceolate, 21.7–45.5 by 5.4–9.5 cm; lateral ones elliptic, oblanceolate, 18.5–30.8 by 5–7.7 cm; apex acuminate, cuspidate, with entire margin or with few small teeth on the cartilaginous margin; base obtuse, acute. Climbing fronds with pinnules subopposite to alternate. Fertile pinnules in 1–7 pairs; apical one lanceolate, elliptic (parallel-sided), 21–38.3 by 1.2–3.9 cm, apex attenuate, long-attenuate, acuminate, entire or with few small teeth (sometimes serration-like) on the cartilaginous border, base acute, strongly oblique; lateral ones elliptic-oblanceolate, lanceolate, parallel-sided, 14–25 by 1.6–2.2 cm, apex acuminate, attenuate, entire or with few small teeth (sometimes serration-like) on the cartilaginous border, base obtuse, equilateral. Sterile pinnules in 1–6 pairs; apical one elliptic, lanceolate or oblanceolate, 19.3–36.7 by 3.7–8.8 cm, apex acuminate, cuspidate, apicular, entire or with few small teeth (sometimes serration-like) on the cartilaginous border, base obtuse, acute, equilateral to strongly oblique; lateral ones elliptic, lanceolate, oblanceolate, 15.2–35.2 by 3.7–8.6 cm, apex cuspidate, acuminate (attenuate, apicular), entire or with few small teeth (sometimes serration-like) on the cartilaginous border, base obtuse, acute, equilateral to oblique; petiolulate, sessile (winged).

Distribution — Salpichlaena volubilis subsp. amazonica occurs mostly in Western Amazonia (eastern slopes of the Andes) with a few occurences in Central Amazonia and the Guyana Shield, in Bolivia, Brazil, Colombia, Ecuador, French Guiana and Peru.

Habitat & Ecology — Tropical rain forest, usually in terra firme, rarely in periodically inundated areas or swamps. On hills, mid-slope and flat terrain. On loamy soil. Altitude 50–1500 (–2378) m.

Additional specimens seen ( paratypes). BOLIVIA, La Paz, puente de Tora ( near Tipuani ), 800 m, Cárdenas M 1287 ( US); near Inglis-Inglis , 1524 m, 16 Aug. 1902, Williams RS 1295 ( US). – BRAZIL, Amazonas, Carauari , c. 4 km south from the village of Bom Jesus, Río Juruá , S5°24' W67°12', 100 m, 30 Mar. 2012, Tuomisto H et al. 16010 ( SP, TUR); GoogleMaps Itamarati, c. 5 km north-west from the town of Itamarati, Río Juruá , S6°24' W68°17', 120–150 m, 8 Apr. 2012, Tuomisto H et al. 16118 ( SP, TUR); GoogleMaps Eirunepe , c. 3 km south from the village of Vila União, Río Juruá, S6°47' W70°11', 140 m, 1 May 2012, Tuomisto H et al. 16550 ( SP, TUR). GoogleMaps – COLOMBIA, Amazonas, Río Caquetá , Araracuara , 24 Sept.1991, Vester H & Matapi A 653 ( Z); Caquetá, Valparaiso, vía Morelia-Valparaiso , finca Las Palmeras , N1°11'3.5" W75°39'21.6", 250 m, 19 Oct.2010, Rodríguez WD et al. 7001 ( NY). GoogleMaps – ECUADOR, Morona-Santiago, comunidad Shuar de Mutints, faldas orientales de la Coordillera del Cutucú , S2°11' W77°44', 600 m, Navarrete H 1257 ( AAU); GoogleMaps Napo, Parque Nacional Yasuní , S0°40' W76°23', 250–280 m, 30 Jan.1996, Navarrete H 1509 ( AAU); GoogleMaps Pastaza, Sarayacu, S01°52' W77°14', 1 Dec.2007, Sirén A et al.2023 ( QCA, TUR); GoogleMaps Sucumbios, Reserva Faunística Cuyabeno, one-hectare plot c. 1 km north of Laguna Grande and surroundings, S0°00' W76°12', 265 m, Poulsen A 80802 ( AAU). GoogleMaps – FRENCH GUIANA, D.Z. du Haut-Kourcibo , Bassin du Sinnamary , N4°28'23" W53°17'18", 80 m, 19Apr.1991, de Granville JJ et al. 11333 ( CAY, US). GoogleMaps – PERU, Amazonas, Bagua , along road from Chiriaco towards Bagua , S05°16'57" W78°23'10", 750 m, 21 Mar. 2001, Van der Werff H. et al. 16278 ( MO); GoogleMaps Cajamarca, San Ignacio, Huarango , S5°03'50" W78°43'19", 2378 m, 27 Aug. 2007, Perea J et al. 3902 ( AAU, US); GoogleMaps Loreto , Loreto, Upper Rio Pastaza, ± 20 km northwest of Nuevo Andoas community, S2°44' W76°37', 250 m, 26 Mar. 2005, Higgins M & Ruiz F 139 ( TUR); GoogleMaps Ramón Castilla, río Yavarí-Mirín , S4°25' W72°23', 100–200 m, 23 Mar. 2002, Cárdenas GG & Salovaara K 1290 ( AMAZ, TUR, USM); GoogleMaps Madre de Dios, Manu, Eastern bank of rio Colorado , 10 km from its confluence with rio Madre de Dios, S12°40' W70°28', 250–350 m, 2 Nov. 1998, Tuomisto H et al. 13601 ( TUR); GoogleMaps San Martín , Rioja , Yaracyacu , S5°58' W77°24', 1381 m, 10 June 2015, Suominen L et al. 219 ( TUR); GoogleMaps Pasco, Oxapampa , Palcazu Valley , Iscozacin , S10°12' W75°15', 400 m, 12 Jan. 1984, Foster R et al. 7894 ( F, USM). GoogleMaps

Notes — In general appearance, S. volubilis subsp. amazonica is similar to subsp. thalassica and non-climbing fronds can be confused with S. hybrida . However,usually subsp. amazonica can be distinguished from subsp. thalassica by the uniform appearance of the abaxial lamina in contrast to the typical white protuberances of the lamina associated with stomata in subsp. thalassica . The chartaceous-coriaceous lamina texture and cartilaginous pinnule margins distinguish subsp. amazonica from S. hybrida and also from subsp. volubilis , both of which have herbaceous texture and non-cartilaginous (or only slightly cartilaginous) margins. It is more difficult to separate subsp. amazonica from subsp. crenata because their characters sometimes overlap. In subsp. amazonica , the pinnule apices are entire or have small teeth that do not cut into the lamina, in contrast to subsp. crenata whose pinnule apices are usually crenate (sometimes slightly serrated or entire). Subsp. amazonica has wider pinnules, more chartaceous-herbaceous texture, and an obvious cartilaginous margin that is absent or not so obvious in subsp. crenata .

In the northern Andes, S. volubilis subsp. amazonica and subsp. thalassica are difficult to separate on morphological grounds. The distribution of our sequenced specimens suggests that S. volubilis subsp. amazonica is restricted to the eastern side of the Andes, whereas subsp. thalassica occurs only on the western side.