Heteropsis flexuosa

5. HETEROPSIS FLEXUOSA (H. B. K.) G. S.

Bunting, Revista Fac. Agric. Univ. Centr. Venezuela 10: 201. 1979. Pothos? flexuosus H. B. K., Nov. Gen. Sp. 7: 151. (1825). Anthurium? flexuosum (H. B. K.) Kunth, Enumeratio Plantarum, vol. III, p. 82. (1841).

— TYPE: VENEZUELA. Montana de Javita, Humboldt & Bonpland 978 (holotype: P! ster).

Heteropsis jenmanii Oliv. , Hooker’s Icon. Pl. 20: t. 1949 (1890).

— TYPE: GUYANA. Mazaruni River, Kalacoon , May 1889, G. S. Jenman 5000 (holotype: K! fl).

Plant hemi-epiphytic, scandent; shoot densely leaved, internodes 2–5.5 cm long, 0.3–1 cm diam, subcylindric, one side flattened to slightly sulcate, pale green, dark brown, vertically striate when dry; lateral bud 3–6 mm long, generally extrorse, apex acute. Petiole 0.5–1 cm long, 2–2.6 mm diam, strongly canaliculate, usually twisted, margin smooth; geniculum 3–5 mm long; leaf blade 13–27 cm long, 3.5– 9.5 cm wide, lanceolate to obovate, subcoriaceous, subglossy, dark green adaxially, yellowish-green abaxially when living, pale to dark brown and opaque when dried, margin revolute, 0.24–0.60 mm thick, apex acuminate to attenuate, 1–2.4 cm long, base acute to cuneate, rarely obtuse, midrib flattened to etched or impressed adaxially, prominent abaxially, yellowish, primary lateral veins and interprimaries obscured on both surfaces, infra-marginal collective vein prominent, 1– 2.5 mm distant from the margin, external marginal vein 1. Inflorescence terminal or axillary, flowering shoot 2–14 (–18) cm long, internodes 0.5–4 cm long, 2–6 mm diam, dark brown when dried, slightly sulcate, axillary buds lacking or only present on the first internode; peduncle 2–5 mm long, 3– 4 mm diam, dark green; spathe 3.5–6 cm long, 1–1.5 cm diam, closed, ellipsoid, convolute when open, yellow, as long as the spadix, apex acute; spadix 3.5–6 cm long, 1–1.4 cm diam, ellipsoid, subclavate, apex obtuse, pale yellow, stipitate, stipe 3–5 + 3–5 mm, green, subcylindric. Stamens 1.5–2.5 mm long, 1–2.5 mm wide, anthers ovate; gynoecium ca. 3.5 mm long, 2.5 mm diam, slightly prismatic, apex 2–5 mm diam, ovary 1.5–2.5 mm long, 0.7–1 mm wide, oblong, locules 2 per ovary, ovules 1–2 per locule, 0.7–2 mm long, anatropous, subsessile, placentation sub-basal, stigma discoid-oblong. Infructescence when immature 4.6–6 cm long, when mature 8–13 cm long, 3–4 cm diam; berry 10–15 + 7–15 mm, green when immature, yellow when mature, with dark green apex, truncate; seed 8–14 + 0.5–1 mm 2 per berry, brown, glossy, oblong. Figures 1B View FIG , 4F, 4G View FIG , 6 View FIG , 7B, 7C View FIG , 8B View FIG , 15 View FIG .

Common Names— The following common names have been recorded for this species: cipó-titica ( Brazil), nibbi ou mibi ( Guiana), tamshi ( Peru).

Habitat and Distribution— Heteropsis flexuosa occurs in dense ombrophilous terra firme forest in all regions where it occurs, and in tall várzea forest in Amazonia, preferring in this case higher terrain. The species is widespread in South America being known from Bolivia, Brazil (Acre, Amapá, Amazonas, Bahia, Maranhão, Pará, Pernambuco, Rondônia, Roraima), Colombia, Ecuador, French Guiana, Guyana, Peru, Surinam, and Venezuela at altitudes between 50–1,000 m altitude.

Phenology— Flowering and fruiting specimens have been collected throughout the year.

Conservation Status— Using guidance from the Geocat (2012) online tool, the preliminary conservation assessment of H. flexuosa is least concern (LC), based on 39 localities and a 50 km cell width (auto value cell size option for area of occupancy).

Etymology— The species epithet probably refers to the flexuose aerial roots which are widely used in Amazonia for fiber.

Notes— Heteropsis flexuosa has a broad distribution, exceeded only by H. oblongifolia to which it is somewhat similar morphologically (see commentary for the latter species). H. flexuosa has also often been confused in herbarium material with H. macrophylla , but the latter can be distinguished by the longer, non-twisted petioles (1–3 cm long), the narrowly oblong leaf blade, olive-green adaxially, paler green abaxially, with an obtuse to shortly cuspidate apex, numerous primary lateral veins which are prominent abaxially, infra-marginal collective vein close to the margin (ca. 0.25–5 mm distant) and usually lacking an external marginal vein, shortly stipitate to sessile spadix and prominently discoid stigma. H. flexuosa has an oblong to elliptic leaf blade which is yellowish-green abaxially, subcoriaceous, acuminate to attenuate apically, with the veins obscured on both surfaces, infra-marginal collective vein 1–2.5 mm distant from the margin with an external marginal vein, stipitate spadix and a discoid to oblong, but less prominent stigma.

In an ecological study of four species of Heteropsis , within an area of 2.4 ha. in the Reserva Florestal Adolpho Ducke ( Soares Morais 2008), H. flexuosa was found to be the most abundant species, comprising 37% of the recorded Heteropsis plants, and occurred on all types of terrain in different strata and on host trees of different diameters. However, it was most common in quadrats at higher elevations within the area (R 2 = 0.27; t = 3.92; p = 0.03). These results, together with label data gathered from specimens collected in other localities show that H. flexuosa occurs between 50 and 1,000 m alt. and is plastic ecologically.

Field observations at the Reserva Florestal Adopho Ducke confirmed that H. flexuosa produces spreading or somewhat pendent plagiotropic branches. Within each lateral shoot, the leaves (8 –14 in number) become progressively smaller toward the apex, thus forming an oval profile for the branch as a whole. This feature of the branch morphology is distinctive to H. flexuosa and differentiates it from the other species studied in the Reserva Ducke ( H. macrophylla , H. spruceana , H. steyermarkii , H. tenuispadix ).

Heteropsis flexuosa was originally described as Pothos flexuosus View in CoL by Kunth based on material collected by Humboldt and Bonpland. The origin of the collection was given as “Crescit inter Atures et Maypures (Missiones del Orinoco); item ad ripam Fluminis Nigri et Tuamini, prope Javitam” (“Occurs between Atures and Maypures (Orinoco Missions); also along the banks of the Rio Negro and Rio Tuani near Javita”). The type specimen in the Paris herbarium (P) is sterile with broad usually obovate leaves. The label reads “Herbier Humboldt & Bonpland (P), Pothos? flexuosus View in CoL –sub? N. 978 Montana de Javita ” and this information, together with the match of specimen and published description confirm it as the holotype. Kunth (1841) later made the new combination Anthurium flexuosum Kunth. Oliver View in CoL described H. jenmanii based on the collection Jenman 5000 from Guyana, which was later reduced to synonymy with H. flexuosa by Bunting (1979).

Additional Specimens Examined— BOLIVIA. Pando: Abuna, 9 50 ' S 65 40 ' W, 8 Jul 1992, Gentry & Alan Perry 77987 ( MO fl). GoogleMaps

BRAZIL. Acre: Marechal Thaumaturgo, 9 02 ' 52.92 '' S–9 02 ' 35.46 '' S 72 16 ' 24.48 '' W – 72 15 ' 59 '' W, 30 Nov 2000, Daly et al. 10286 ( MO fl). Amapá: Mazagão, 25 Feb 1988, Pires & Silva 2022 ( MG fl). Porto Grande, 1 Mar 2001, Pereira et al. 151 ( HAMAB, INPA fl) ; 1 Mar 2001, Pereira et al. 160 (HAMAB, INPA, MG fl); 1 Mar 2001 Pereira et al. 162 ( INPA ster).

Amazonas: 19 Dec 1955, Coelho s.n. ( INPA 3140 fr). Manaus, Reserva Florestal Adolpho Ducke, 24 Nov 1993, Ribeiro et al. 1181 ( INPA fr); 15 Dec 1994, Ribeiro & Silva 1536 ( INPA fl); 31 Jan 1996, Ribeiro et al. 1799 ( INPA fr); 21 Feb 1995, Soares et al. 190 ( INPA fr), Soares et al. 497 ( INPA fl); 8 Sep 1994, Assunção s.n. ( INPA 176280 fr); 8 Sep 1994, Assunção et al. 47 ( INPA fl); Barcelos, 20 Apr 1955, Froes 28374 (IAN fr); Humaitá, 16 Jun 1980, Janssen 474 ( INPA fl) ; 3 07 ' S 58 59 ' W, 17 Aug 1986 Croat 62269 ( INPA, MO ster) GoogleMaps ; 16 Aug 1986, Croat 62208 (MO ster); Manaus-Itacoatiara, 12 Oct 1976, Adair s.n. ( INPA 72818 fl); 20 Nov 1968, Coelho & Lima s.n ( INPA 25953 fl); 24 Nov 2000, Souza et al. 418 ( INPA fl); 3 Nov 1973, Steward & Ramos P17656 ( INPA, U fl); Rio Javari, 8 Aug 1973, Lleras et al. P17214 (U fl); Santa Isabel do Rio Negro, 1 Sep 2003, Soares et al. 523 ( INPA fl, fr); Urucu, 18 Sep 2005, Soares et al. 669 ( INPA fr); 19 Sep 2005, Soares et al. 671 ( INPA fl, fr); 14 Dec 2005, Soares et al. 758 ( INPA fl); São Gabriel da Cachoeira, 10 Feb 2001, Hoffman & Farias 139 ( INPA fl). Bahia: Ilhéus, 29 Nov 2000, Gonc¸alves et al. 407 (CEPEC ster); 6 Mar 2001, Gonc¸alves et al. 791 (CEPEC ster); 11 Dec 1997, Gonc¸alves & Jardim 134 (CEPEC ster). Una, 7 Nov 2005, Soares et al. 753 (CEPEC ster); 16 Dec 1968, Santos 317 (CEPEC fl, fr); Goianesia, 8 Apr 2003, Bastos et al. 2956 ( MG fl); Uruc¸uca, 6 May 1996, Nadruz et al. 1180 (CEPEC ster). Maranhão: Rio Turiac¸u, 20 May 1979, Jangoux & Bahia 759 ( MG fl). Pará: 25 Sep 1948, Froes 23608 (RB ster); 11 Mar 1958, Froes 34197 (IAN fl); 19 Nov 1977, Prance et al. 25577 ( MG fl); 30 Mar 1974, Medeiros & Marinho s.n. (IAN 143489 fr); Jul 1952, Pires 7052 (IAN fl); 22 Nov 1996, Carlos et al. 1229 ( MG fr); 17 Aug 1969, Silva & Souza 2290 ( MG fr); 16 Nov 2001, Bastos et al. 2275 ( MG fl); 11 Sep 1908, Pessoal do Museu 9636 ( MG ster); 23 Mar 1986, Balée 2014 (MO ster); Belém, 26 Apr 1960, Oliveira 601 (IAN fr); 18 Sep 1963, Oliveira 2581 (IAN fr). Castanhal, 3 Oct 1942, Silva s.n. (IAN fr); Itaituba, 20 May 1983, Amaral et al. 1352 ( INPA fr); Mocambo, 1 25 ' S 48 25 ' W, 13 Aug 1986, Croat 62105 ( MO ster) GoogleMaps ; Tucuruí, 22 Oct 1983, Lima & Silva 90 ( INPA fl); 21 Oct 1966, Pires & Silva 10279 (IAN fr); Paragominas, 4 17 ' S, 47 32 ' W, 1 Mar 1980, Plowman et al. 9402 ( INPA, MO ster) GoogleMaps ; Santarém, Feb 1955, Froes 31473 (IAN fl). Pernambuco: Recife, 14 Feb 1990, Guedes 2267 ( UFBA fl). Rondônia: 28 Jun 1983, Silva 6506 ( INPA fl); Porto Velho, 26 Oct 1979, Vieira et al. 310 ( INPA fl); 27 Jul 1997, Lobato et al. 1830 ( MG fl). Roraima: 18 Feb 1971, Prance et al. 10598 ( INPA fl); 28 Feb 1971, Prance et al. 10761 ( INPA fl); 16 Nov 1977, Steward et al. 39 ( INPA fl).

COLOMBIA. 17 Sep 1991, Vester 569 ( MO ster).

ECUADOR. 19 Jun 1991, Palacios & Freire 7396 (MO fr). Napo: 12 Nov 1987, Ceron 2881 (MO fl) ; 00 52 ' S 76 05 ' W, 9 – 13 Jan 1988, Ceron & Coelho 3295 ( MO fr) GoogleMaps ; 00 57 ' S 76 13 ' W, 9 Jan 1988, Neill et al. 8353 ( MO fr) GoogleMaps ; 00 57 ' S 76 13 ' W, 9 Jan 1988, Neill et al. 8268 ( MO fl) GoogleMaps ; 00 52 ' S, 76 05 ' W, 11 – 17 Jan 1988, Coelho 14 ( MO fr) GoogleMaps ; 00 52 ' S, 76 05 ' W, 11–17 Jan 1988, Coelho 83 (MO fl fr); 0 59 ' S 76 12 ' W, 8 Jan 1995, Aulestia & Omehuat 3213 (MO fl); 1 Apr 1999, Leimbeck 123 (MO ster); 1 Jan – 8 Oct 1938, Gill 64 (K ster); Orellana Reserva Étnica Huaorani , 0 48 ' S 76 28 ' W, 6 Oct 1993, Dik 630 (MO fl) GoogleMaps ; 01 02 ' 33 '' S 76 53 ' 11 '' W, 30 Jan 2004, Freire & Narajo 478 (MO ster); 00 45 ' S 76 45 ' W, 21 Nov 1991, Neill & Rojas 10022 (MO ster); 0 38 ' S 76 9 ' W 14 Feb 2002, Koster et al. 926 (MO fl). Pastaza: 01 34 ' S 77 25 ' W, 25 Nov 1990, Gudiño 1132 (MO fl) GoogleMaps ; 18 Dec 1991, Gudiño & Gualinga 1619 (MO ster); 01 15 ' S 76 55 ' W, 1 Mar 1989, Vlastimil Zak 4051 (MO ster); 01 15 ' S, 76 55 ' W, 1 –6 Mar 1989, Vlastimil Zak 4143 (MO ster). Sucumbios, 00 00 ' S, 76 11 ' W, 15 Nov 1991, Palacios et al. 9258 (MO ster). San Martín: Provincia Mariscal Cáceres, 2 May 1981, Schunke-V. 12827 (MO ster) GoogleMaps .

FRENCH GUIANA. Reserve des Nouragues , 4 3 ' N 52 42 ' W, 6 Mar 2004, Poncy et al. 1807 (P, CAY, MO, U fl) GoogleMaps ; Montagne de Kaw , 1 Apr 1984, de Granville 6703 (P fr) ; Rivière Arataye 1 Feb 1969, Oldeman 2899 (P fl) ; 27 Mar 1981, Barrier 2830 (B, K, MO, NY, P, U, US fr) ; Marifasoula , 29 Aug 1961, Schnell 11543 (P ster) ; Sipomama 18 Jun 1984, Sauvain 139 (K, P ster) ; Wayãpi 4 Dec 1974, Grenand 560 (K ster) ; 28 Mar 1910, Anderson 432 (K ster); 29 Jun 1979, Prevost 670 (K ster), 03 37 ' N 53 12 ' W, 27 Mar 1983, Mori & Pipoly 15415 (MO, P fr); 15 May 1986, Mori & Pennington 18014 (MO fr).

GUYANA. 5 30 ' N 58 22 ' W, 6 Jul 1982, Croat 53843 ( MO ster); GoogleMaps 03 37 ' N 53 12 ' W, 11 Feb 1993, Croat 74186 ( MO fl) GoogleMaps ; 11 Feb 1993, Croat 74186A (MO ster). Essequibo River, 15 –24 Dec 1937, Smith 2722 (P fl), 15 May 1997, Clark 4557 (U fr). Pomeroon – Supenaam 07 17 ' N 58 54 ' W, 11 Jul 1997, Hoffman & Ehringhaus 5104 ( MO ster) GoogleMaps ; Potaro – Siparuni 04 15 ' N 58 30 ' W, 17 Mar 1996, Hoffman & Allicock 5060 ( MO fr) GoogleMaps ; 04 12 ' N 58 30 ' W, 19 Mar 1996, Hoffman & Allicock 5052 (MO fr); 19 Mar 1996, Hoffman & Allicock 5061 (MO fr); 03 37 ' N 53 12 ' W, 19 Mar 1996, Hoffman & Allicock 5066 (MO ster); 04 12 ' N 58 30 ' W, 19 Mar 1996, Hoffman & Allicock 5067 (MO ster); 19 Mar 1996, Hoffman & Allicock 5070 (MO fr); 19 Mar 1996, Hoffman & Allicock 5071 (MO ster); 19 Mar 1996, Clark 1336 (U fl); 17 Dec 1997, Clark 3617 (U fr); 23 May 1997, Clark 4881 (U fl); Rupununi, 5 Feb 1991, Jansen-Jacobs et al. 2381 (MO, U fr); 02 00 ' N 59 15 ' W, 23 Oct 1992, Jansen-Jacobs et al. 3077 (K, MO, P, U fr).

PERU. 2 Jan 1973, Berlin 800 ( MO fr) ; 04 55 ' S 73 45 ' W, s.d., Baluarte s.n. (MO 5564058 ster). Reserva Tambopata, 15 Mar 1988, Bell et al. 88–178 (K fr); 14 Mar 1980, Huashikat 2262 ( MO fr). GoogleMaps Bagua : 04 55 ' S 78 19 ' W, 20 Oct 1995, Vásquez et al. 20362 ( MO fr) GoogleMaps ; 04 55 ' S 78 19 ' W, 14 Oct 1995, Vásquez & Jaramillo 20268 (MO fr); 04 55 ' S 78 19 ' W, 22 Oct 1995, Vásquez et al. 20393 (MO fr); 17 Nov 1997, Rojas et al. 583 ( MO fr). Loreto: Maynas , 6 Sep 1990, Ruiz et al. 1561 ( MO ster) ; 18 Nov. 1990, McCann 117 (MO ster); 18 Aug 1983, Rimachi 6935 (MO ster); 15 Aug 1990, Salaun & McCann 31 (MO ster); 22 Aug 1990, Salaun & McCann 40 (MO ster); 3 28 ' S, 72 50 ' W, 25 Jan 1990, Vásquez & Ayala 13411 (MO fl); 73 20 ' W, 94 10 ' S, 2 Jul 1988, Vásquez & Jaramillo 10829 (MO fr); 12 Apr 1977, Plowman et al. 6832 (MO ster); 3 May 1977, Plowman et al. 7204 (MO ster); Rio Ucayali, 75 10 ' W, 07 05 ' S, 27 Nov 1985, Vásquez et al. 6978 (MO fr). GoogleMaps Madre de Dios: 12 50 ' S 69 17 ' W, 1 Mar 1981, Gentry & Young 31848 (MO ster) GoogleMaps ; 12 49 ' S 69 18 ' W, 18 Feb 1984, Gentry et al. 45612 (MO fr); 12 49 ' S 69 43 ' W, 22 Jul 1985, Gentry et al. 51220 (MO fr); 12 49 ' S 89 18 ' W, 20 Aug 1990, Reynel et al. 5283 (MO fr); 71 52 ' W 11 40 ' S, 14 Oct 1986, Foster 11825 (MO fr); 12 49 ' S 69 17 ' W, 26 Jun 1980, Barbour 5804 (MO fr). GoogleMaps Pasco: 29 May 1984, Salick 7064 (MO ster) GoogleMaps ; 6 Apr 1985, Salick 7256 (MO ster); 10 09 ' S 15 12 ' W, 29 Aug 1986, Salick 7706 (MO ster); 10 09 ' S 15 12 ' W, 29 Aug 1986, Salick 7708 7 (MO ster); 29 Aug 1986, Salick 7708 (MO ster); Rio Santiago , 77 40 ' W, 3 50 ' S, 22 Dec 1979, Tunqui 416 (MO fr) GoogleMaps ; 17 Sep 1979, Huashikat 651 ( MO fr). GoogleMaps Puno: 13 21 ' S 69 40 ' W, 16 May 1992, Gentry et al. 76649 ( MO fr) GoogleMaps ; 30 Mar 1996, Aguilar & Castro 437 (MO fr); 12 50 ' S 69 17 ' W, 27 Sep 1994, Vásquez et al. 19286 (MO fr).

SURINAME. Kamueatta , 2 Jul 1918, Hohenkerk s.n. ( K fr). GoogleMaps Kariako, Barama River 07 22 ' N 59 42 ' W, 4 Sep 1996, van Andel et al. 1039 ( U ster) GoogleMaps ; 15 Mar 1949, Jonker et al. 2715 (IAN fr); 4 Mar 1918, Wallnofer 3803 (K fr); 12 Feb 1919, B.W. 4255 (P fl); 30 Jun 1924, B.W. 6555 (MO ster); 6 Feb 1942, Stahel s.n. (K, MO fl); 16 Mar 1942, Stahel s.n. (MO fr); 15 Mar 1949, Lanjouw & Lindeman 2715 (K fr); 2 58 ' 18 '' N 54 33 ' 14 '' W, 11 Apr 1998, Evans & Peckham 2857 (MO fr); Coppename River , 10 Mar 2004, Clarke & Rhodes 11360 (U fr) ; Mapane , 25 Apr 1967, Vreden 11650 (K ster) .

VENEZUELA. 6 Feb 1990, Stergios & Velazco 14706 ( MO ster), 5 37 ' N 67 22 ' W, 29 Jan 1991, Romero et al. 2270 (MO fl); GoogleMaps 5 37 ' N 67 22 ' W, 6 Feb 1992, Romero et al. 2317 (MO ster) GoogleMaps ; 5 37 ' N 67 22 ' W, 9 Feb 1992, Romero et al. 2343 (MO fl); 18 Apr 1990, Perez 962 (MO ster); 6 21 ' N 64 59 ' W, 25 Jan 1994, Salas s.n. (MO 05033605 ster). Amazonas: 5 37 ' N 67 22 ' W, 1 Feb 1991, Romero et al. 2287 (MO fl); 23 Mar 1983, Clark 83–1 (MO ster). Atabapo: 02 24 ' N 64 24 ' W, Oct 1991, Marin 1684 (MO fr); 14 Feb 1990, Aymard & Delgado 7852 (MO ster); 3 39 ' N 65 42 ' W, 6 Mar 1985, Liesner 18387 (MO fl, fr); 0 50 ' N 66 10 ' W, 25 Nov 1984, Croat 59314 (MO ster). Atures: Nov 1989, Sanoja et al. 3178 (MO fr); 9 May 1980, Steyermark 122135A (MO ster); 0 50 ' N 66 10 ' W, 10 May 1980, Steyermark et al. 122221 (MO fl); 5 35 ' N 67 15 ' W, 2 Nov 1988, Liesner 25679 (MO fr). Bolivar: 04 55 ' N 62 49 ' W, 1 Sep 1986, Fernandez 3322 ( INPA, MO fl); 04 23 ' N 61 38 ' W, 21 Oct 1985, Holst & Liesner 2385 (MO ster); Nov 1994, Knab-Vispo 224 (MO ster); Feb 1986, Elio Sanoja 199 (MO ster); 5 43 ' N 64 07 ' W, 30 Oct–2 Nov 1988, Aymard & Angel Fernandez 7248 (MO ster). Rio Negro: 22 Jan 1992, Aymard et al. 9728 (MO ster); 30 Mar 1953, Steyermark 74792 (MO ster); 14 May 1953, Steyermark 73349 (MO fl); 22 May 1980, Steyermark et al. 117194 (MO ster); 4 59 ' N 61 10 ' W, 9 May 1988, Liesner 24394 (MO ster); 4 23 ' N 61 38 ' W, 22 Oct 1985, Liesner 18962 (MO ster); 4 30 ' N 61 35 ' W, 7 Nov 1985, Liesner 19635 (MO fr); 20 Apr 1986, Liesner & Host 20076 (MO ster); Cassiquiare, Rio Javita, 7 Apr 1981, Clark & Ribeiro s n. (MO ster).