Heteropsis oblongifolia, Kunth, Enum. Pl.

Soares, M. Lourdes, Mayo, Simon J. & Gribel, Rogério, 2013, A Preliminary Taxonomic Revision of Heteropsis (Araceae), Systematic Botany (Basel, Switzerland) 38 (4), pp. 925-974 : 953-955

9. HETEROPSIS OBLONGIFOLIA Kunth ,

Enum. Pl. 3: 60. 1841.

— TYPE: BRAZIL. F. Sellow (as “ Sello ”) 207 (lectotype, here designated: K! fl).

Dracontium integerrimum Vell. , [Fl. Flumin. 9: t. 119 (1831), nom. nud.] Arquivos do Museu Nacional 5: 389 (1881). Heteropsis integerrima (Vell.) Stellfeld , Arq. Mus. Paranaense 8: 179 (1950).— TYPE: BRAZIL. Published plate t.119, original citation: “Dicitur etiam Timbôpeba... Habitat silvis maritimis.“ No specimen is known to exist.

Plant a scandent hemi-epiphyte; shoot densely leaved, internodes 1.9–5 cm long, 1.0– 6 mm diam, dark green, subquadrangular, one side flattened to slightly sulcate, pale brown when dried, longitudinally striate; axillary bud 2.7– 5 mm long, extrorse to straight, apex pointed. Petiole 4–9 + 1.5–2.4 mm, strongly canaliculate, margin pale, sinuate towards the apex; geniculum 3–4 mm long; leaf blade 10– 23 cm long, 3.2–7 cm wide, ovate, lanceolate, oblong to oblanceolate, coriaceous, subglossy, dark green adaxially, paler green abaxially when living, opaque pale brown when dried, 0.14–0.58 mm thick, margin revolute, apex abruptly cuspidate to acuminate, base cuneate, acute, subacute to obtuse, midrib sulcate adaxially, prominent, yellowish abaxially, primary lateral veins and interprimary veins prominent abaxially when dried, infra-marginal collective vein prominent, 0.6–3 mm distant from margin, external marginal veins 1–2. Inflorescence terminal, axillary, flowering shoot 1–14 cm long, internodes 0.2–3 + 0.12– 0.46 cm diam, pale brown when dried, sulcate, normally with lateral bud on the apical internode; peduncle 0.5– 3 mm long, 2–4 mm diam, dark green; spathe as long as the spadix, 3.5–5 cm long, 0.9–1.3 mm diam when closed, ca. 5 cm wide when expanded, oblong to ovate, inflated, convolute, yellow, with a white stripe ca. 1.5 cm wide at the margin, apex acuminate to abruptly cuspidate, sometimes somewhat twisted; spadix 1.5–3.5 + 0.5–0.8 cm diam, ellipsoid, apex rounded-acuminate, pale yellow, stipitate, stipe 2.3–4 + 2–4 mm, green, subcylindric. Stamens 3–3.5 mm long, 1–1.25 mm wide, anthers ovate to elliptic; gynoecium 1.5–3.5 + 1.5–3 mm diam, prismatic, with tannin cells in the stylar region, apex 2 – 3.5 mm diam, ovary 2-locular, 1– 1.5 mm long, obpyramidal, locules with sticky, translucid mucilaginous contents, ovules 2 per locule, ca. 0.5 mm long, anatropous, sessile, placentation sub-basal, stigma punctate-discoid to oblong. Infructescence 6–10 cm long, 2–3.3 cm diam; berry 10–17 + 8–17 mm, green when immature, brown when pre-mature, red with dark brown apex (or orange-yellow to red in Costa Rica) when mature, truncate to prismatic; seed 7–10 + 4–8 mm, 2–4 per berry, brown (black in Costa Rica), oblong to elliptic. Figures 2A , 7D, 7E , 9D , 19 .

Common Names— The following common names have been recorded for this species: cipó-titica ( Brazil), bejuco de hombre ( Peru).

Habitat and Distribution— Heteropsis oblongifolia occurs in dense, ombrophilous forest, seasonal montane forest, secondary forest or in disturbed areas with rock outcrops in Bolivia, Brazil (Acre, Amapá, Amazonas, Bahia, Espírito Santo, Maranhão, Mato Grosso, Minas Gerais, Pará, Paraná, Pernambuco, Rio de Janeiro, Roraima), Colombia, Costa Rica, Ecuador, Nicaragua, Peru, and Venezuela, at between 50 and 2,000 m alt.

Phenology— Flowering and fruiting specimens have been collected almost throughout the year, but specimens have not been seen in flower in June and September or in fruit in August.

Conservation Status— Using guidance from the Geocat (2012) online tool, the preliminary conservation assessment of H. oblongifolia is least concern (LC), based on 34 localities and a 50 km cell width (auto value cell size option for area of occupancy).

Etymology— The species epithet refers to the more oblong leaf shape, in contrast to the lanceolate leaf shape of H. salicifolia , the latter being the only other species described by Kunth (1841).

Notes— Heteropsis oblongifolia is the most widely distributed species, occurring from Central America ( Nicaragua and Costa Rica) to tropical South America although it has not been recorded from the Guianas and Suriname. This wide distribution is accompanied by great morphological and ecological variation, with habitats ranging from sea level to 2,000 m. Grayum (2003: 118) notes that in view of the wide disjunction between Costa Rica and eastern Brazil, there is doubt that the Central American taxon is really conspecific with the Atlantic Forest taxon. A more detailed comparison of living plants (beyond the scope of the present study) from the two regions is needed to resolve this question.

The species has some similarities to H. flexuosa , especially in the morphology of the leaf blade and this has led to many dubious identifications in sterile herbarium specimens. However, H. oblongifolia differs from H. flexuosa by its straight lateral buds, the apical sinuations of the petiole margin, less coriaceous leaf blade, infra-marginal collective vein up to 3 mm distant from the margin, and presence of 1–2 external marginal veins. When fertile, the main characters that differentiate the two species are to be found in the spathe, which in H. oblongifolia has a white stripe along the margin, an acuminate to abruptly cuspidate apex and red mature berries, dark brown at the apex and yellowish towards the base in Costa Rican material. In H. flexuosa the petiole has a smooth margin and is usually twisted, the leaf blade is coriaceous, the flowering shoot has at most a lateral bud on the internode below the spathe or none at all, the spathe is yellow, and the berries are yellow when mature.

In the original description of Heteropsis oblongifolia Kunth cited only “Brasilia meridionalis (Sellow legit.)”, i.e. a collection by Friedrich Sellow (also known as Sello) from southern Brazil. Kunth’s herbarium was deposited at Berlin (B) and this type specimen must have been included in it. However, this specimen did not survive the damage to B during the Second World War. The microfiche of Schott’s collection of drawings at Vienna ( Nicolson and Riedl 1984; Schott 1984) includes a drawing (Icon no. 3577) which is the only one determined by Schott as Heteropsis oblongifolia and which bears the annotation “Her. G. Berol. 207 ''. Since Schott attempted whenever possible to have drawings made of type specimens, it is probable that this specimen represents the collection Sellow 207. This is confirmed by the presence of a duplicate specimen at K, in the herbarium of W. J. Hooker with more detailed label information: “207. Ex Herb. Reg. Berolinense 1859, Heteropsis oblongifolia Kunth. Doubletten Brasilia. Sello. ” We have therefore selected this specimen, doubtless originally an isotype, as the lectotype of H. oblongifolia Kunth.

Vellozo’s name Dracontium integerrimum (Flora Fluminensis, t. 119) was invalid when first published ( Vellozo 1831) because there was no description or any analysis in the figure (i.e. a separate figure or element showing details). It remained a nomen nudum until the first publication of Vellozo’s complete text of the Flora Fluminensis ( Vellozo 1881), by which time Kunth’s epithet had priority in the genus Heteropsis ; the combination Heteropsis integerrimum was later made by Stellfeld (1950) in his study of Vellozo’s work on Araceae . A comparison of the habit, leaves and post-floral inflorescences leaves little doubt that H. oblongifolia and H. integerrima denote the same species.

Additional Specimens Examined— BOLIVIA. 30 Jul 1982, Sperling & King 6409 ( K, MO fl, fr).

BRAZIL. Acre: Brasiléia, 1 Nov 1980, Lowrie 670 ( INPA, MO fl). Amapá: Laranjal do Jarí, 13 Nov 2004, Pereira & Cardoso 748 ( HAMAB, INPA fr). Amazonas: Humaitá, 11 Oct 1934, Krukoff 6493 ( U fl). Rio Negro, 2 Ma 1973, Loureiro et al. s.n. ( INPA 37938 fr); Santa Isabel do Rio Negro, 11 Jun 1976, Marinho 516 ( IAN ster). Bahia: Ilhéus, 19 Mar 1998, Queiroz & Leoni 4984 ( CEPEC, K fl); Mata da Esperanc¸a, 8 Nov 2005, Soares et al. 751 ( CEPEC, INPA fl); 8 Nov 2005, Soares et al. 752 ( CEPEC, INPA fl); 16 Apr 1991, Mayo & Santos 751 ( K, MO ster); 18 Apr 1991, Mayo et al. 755 ( CEPEC, K, MO ster); 6 Oct 1970, Santos 1276 ( CEPEC fr, K fl); 14 Dec 1988, Santos 4454 ( CEPEC fr); Serra Bonita, 9 Nov 2005, Soares et al. 755 ( CEPEC, INPA fl); Itacaré, km 15, Estrada Taboquinhas 24 Apr 1991, Mayo et al. 770 ( K, MO ster); 24 Apr 1991, Mayo et al. 792 ( CEPEC ster, K); 26 Apr 1991, Mayo et al. 801 ( CEPEC, K, MO ster). Juc¸ari, 11 May 1991, Mayo et al. 813 ( CEPEC, K, MO ster). Maraú, 25 Feb 1980, Carvalho et al. 159 ( CEPEC fl); Porto Seguro, 11 Jan 1977, Harley et al. 17851 ( CEPEC, K ster); Prado, 22 Oct 1993, Thomas et al. 10156 ( CEPEC fr); Una, 28 Jul 1994, Jardim et al. 519 ( CEPEC ster); 7 Nov 2005, Soares et al. 754 ( CEPEC, INPA ster); 11 Jan 1977, Harley et al. 18232 ( CEPEC, K ster); 14 May 1991, Mayo et al. 821 ( K, MO fr); Wenceslau Guimarães, 31 May 1991, Mayo & de Carvalho 866 ( CEPEC, K, MO ster). Espírito Santo: Domingos Martins, 30 Jul 1986, Croat 61816 ( MO ster); Linhares, 15 Oct 1992, Folli 1664 ( MO ster); 2 Oct 1971, Santos 2046 ( CEPEC, K fl); Santa Teresa, Aparecidinha, 9 Nov 1998, Kollmann et al. 914 ( INPA, MBML, RB fl); 27 Oct 1998, Kollmann et al. 802 ( MBML fl); 24 Nov 2004, Kollmann et al. 1082 ( MBML fl); 27 Oct 1999, Demuner et al. 189 ( INPA, MBML fl); Estação Biológica de Santa Lúcia, 17 Nov 1994, Soares 600 ( INPA, MBML fl). Maranhão: Monção, 12 Apr 1985, Balée & Gely 859 ( MO ster). Mato Grosso: 4 Oct 1975, Lisboa et al. 659 ( INPA fl). Minas Gerais: Carangola, 30 Jan 2001, Gonc¸alves et al. 706 ( UFMG fr). Pará: 14 Dec 1979, Maciel et al. 528 ( MG fl); 6 May 1968, Pires & Silva 11645 ( IAN fl); 13 Mar 1968, Silva 2731 ( IAN fr); Porto Trombetas, s.d., Evandro 390 ( INPA fl); Santarém, 15 Nov 1977, Prance et al. 25429 ( MG fl); 17 Aug 1969, M. Silva 2291 ( MG fl); Tucuruí, 5 Jun 1980, M. G. Silva & Rosário 5380 ( MG fr). Paraná: Antonina, Bairro Alto, 4 Feb 1983, Hatschbach et al. 46085 ( K, MBM, UFMG ster). Pernambuco: Cabo, 13 Jan 1993, Nadruz & Mayo 887 ( K ster); Curado, Lins & Silva s.n. ( RB 321894 ster); Recife, 26 Jan 1996, Lins & Cleidson 137 ( RB ster). Rio de Janeiro: 18 Feb 1981, Lima et al. 559 ( MO fl); Aug 1965, Strang 680 ( GUA fl); Campos dos Goytacazes, 2 Feb 1984, Kuhlmann 6507 ( RB ster); 1 Oct 1987, Ribeiro et al. 1146 ( GUA ster); Magé Paraíso, Farney et al. 521 ( RB fl). Roraima: 21 Feb 1971, Prance et al. 10660 ( INPA, U ster).

COLOMBIA. 24 Apr 1990, Londono et al. 1626 ( MO ster) ; 24 Jul 1990, Alvarez et al. 882 ( MO ster). Caquetá: 8 Nov 1991, Duivenvoorden et al. 848 ( MO ster) .

COSTA RICA. Lankester 153 ( K fr) ; 9 May 1960, Lankester 1762 ( K fr) ; 4 May 1983, Gomez et al. 20424 ( K ster). Aguabuena, 3 km W of Rincón 15 Apr 1993, Thomsen 336 ( K fl). Heredia, Parque Nacional de Bráulio , 5 Jul 1990, Acevedo 96 ( K fr) .

ECUADOR: Esmeraldas, 3 Nov 1994, Pennington 14959 ( K ster). Morona Santiago, 26 Aug 1985, Anananch 129 ( MO ster). Napo: Apr 1988, Paz & Mino 81010 ( MO ster); 13 Mar 1968, Pitman & Romero 287 ( MO ster); 0 40.853 ' S 76 23.697 ' W, 20 Jun 1995, Acevedo-Rodriguez & Cedeno 7423 ( MO fl, fr); Sucumbios, Apr – Oct 1988, Paz & Mino 81010 ( MO ster). Pastaza: 26– 31 Jan 1989, Neill & Hurtado 8822 ( MO ster).

NICARAGUA. Rosita, 14 08 ' 58 '' N 84 16 ' 00 '' W, 21 May 1994, Rueda, et al. 1459 ( K ster).

PERU. Bagua: 04 55 ' S, 78 19 ' W, 3 Feb 1996, Jaramillo et al. 1060 ( MO fr); 12 50 ' S, 69 20 ' W, 6 Oct 1985, Smith et al. 589 ( MO fl). Loreto: Maynas, 04 29 ' S, 73 35 ' W, 24 Nov 1990, Grandez & Ruiz 2143 ( MO fr); 18 Aug 1978, Haxaire 5157 ( P ster); 13 Jul 1982, Rimachi 6203 ( MO fr); 6 Apr 1989, Vásquez et al. 11999 ( MO fr); 9 Nov 1982, Vásquez & Jaramillo 3424 ( MO fr); 12 Apr 1977, Plowman et al. 6832 ( MO ster). Madre de Dios: 30 Nov 1991, Timana & Jaramillo 3620 ( K ster); 28 Feb 1981, Gentry & Young 31789 ( MO fr); 12 15 ' S, 69 17 ' W, 5 Nov 1984, Young & Stratton 186 ( MO fl); 27 Mar 1981, Young 205 ( MO fr); 2 Jun 1980, Barbour 5487 ( MO fr); Apr 1988, Foster et al. 81010 ( MO ster); Monte Virgem, 6 Sep 1979, Huashikat 410 ( MO fr); Rio Santiago, 21 Feb 1980, Huashikat 2168 ( MO fr); Tambopata, 3 Mar 1981, Gentry & Young 31969 ( K, MO fr).

VENEZUELA. Amazonas: 5 37 ' N 67 22 ' W, 1 Feb 1991, Romero et al. 2286 ( MO ster). Bolivar: 04 58 ' N, 63 24 ' W, Apr 1988, Fernandez 4506 ( MO fr) ; Caño Mosquito , 30 Jan 1976, Colchester 2126 ( K ster) ; 21 Feb 1976, Lister 259 ( K ster) ; 23 Feb 1976, Lister 267 ( K ster) .

Figures

FIG. 2. Geographical distribution of species of Heteropsis. A. H. oblongifolia and H. peruviana. B. H. rigidifolia, H. robusta, H. salicifolia, and H. spruceana. Drawn by Francisca D. A. Matos.

FIG. 7. A–M. Inflorescences in bud and at anthesis. A. Heteropsis duckeana. B–C. H. flexuosa. D–E. H. oblongifolia. F–G. Bud and open inflorescence of H. salicifolia, respectively. H. H. spruceana. I–J. Bud and open inflorescence of H. steyermarkii, respectively. L–M. H. tenuispadix. (photos A–C, H–J and M by M. L. Soares; D–G by Hélio; L by Projeto Flora Reserva Florestal Adolfo Ducke).

FIG. 9. A–F. Mature and immature infructescences of Heteropsis. A. H. duckeana. B–C. Immature and mature infructescences of H. spruceana, respectively. D. H. oblongifolia. E–F. Immature and mature infructescences of H. tenuispadix, respectively. (photos: A, C, E, F by M. L. Soares, B by Projeto Flora Reserva Florestal Adolfo Ducke, D by André Amorim).

FIG. 19. Heteropsis oblongifolia. A. Habit. B. Venation of right abaxial surface of leaf blade. C. Inflorescence. D. Frontal view of stamen. E. Lateral view of gynoecium. F. Transverse section of ovary. G. Longitudinal section of gynoecium. H. Lateral view of ovule. (A–H prepared from Soares et al. 600 (INPA)). Drawn by Felipe França.