Curarea toxicofera (Wedd.) Barneby & Krukoff

Ortiz, Rosa del C., 2018, A taxonomic revision of Curarea Barneby & Krukoff (Menispermaceae), PhytoKeys 100, pp. 9-89 : 51-54

9. Curarea toxicofera (Wedd.) Barneby & Krukoff Figs 26A-H , 27A-F

Curarea toxicofera (Wedd.) Barneby & Krukoff, Mem. New York Bot. Gard. 22(2): 9. 1971. fig. 1.

Cocculus toxicoferus Wedd., in Castelnau, Expéd. Part. Cent. Amér. Sud. 5: 22. 1851. Type. Peru. "Vulg. Pani-base d’un poison pour les flèches utilisé chez les Indiens Pebas-Hte. Amazone", [1847], Castelnau s.n. (sterile) (lectotype designation effected by Krukoff and Moldenke 1939, pg. 338: P! [P00048602] which is annotated as Cocculus toxicoferus in the hand of Weddell and has the eight leaves mentioned by Krukoff and Moldenke. Moreover, this specimen was also annotated in 1939 by Moldenke as “Type”, photograph at NY!; presumed isolectotypes: F! [F-893667, frag.], P!). Peru, Florida, Río Putumayo, at mouth of Río Zubineta, forest, 200 m, Mar 1931-Apr 1931, (♂ fl), Klug 2042 (epitype, designated here: MO!; isoepitypes: BM!, F!, GH!, K!, NY!, US!). Note: Following Art. 9.8 of the Melbourne Code ( MacNeill et al. 2012), I am here designating an epitype to serve as an interpretative type of Cocculus toxicoferus , whose sterile condition makes it ambiguous for identification purposes.

Chondrodendron toxicoferum (Wedd.) Moldenke & Krukoff, Brittonia 3(2): 338. 1938. Type: Based on Cocculus toxicoferus Wedd., in Castelnau Expéd. Part. Cent. Amér. Sud. 5: 22. 1851.

Hyperbaena polyantha Diels, Verh. Bot. Ver. Brand. 50: 73. 1908. Type: Brazil. Amazonas: Juruá Miry, Lago de Esperança, Aug 1901, (♂ fl), Ule 5631 (lectotype designation effected by Krukoff and Moldenke 1938, pg. 23: B!, F neg. 4985]; isolectotypes: CORD! [image seen], F! [F-1014713, frag.], G!; K! [image seen], MG!, [image seen], NY! [NY00320584, frag.]).

Chondrodendron polyanthum (Diels) Diels, in Engler, Pflanzenr. 4(94): 78. 1910. Type: Based on Hyperbaena polyantha Diels.

Chondrodendron bioccai ? Chondrodendron bioccai G. Lusina, Revista Mus. Paul. Univ. São Paulo II. 8: 227, figs 1-2. 1954. Type: Brazil? Locality unknown, collector unknown.

Description.

Medium-sized understory lianas about 10-12 m tall; older stems more or less terete or less frequently flattened, then ca. 3 cm wide; bark greyish to dark brown, with shallow lengthwise fissures; branchlets brownish, greyish to silvery puberulent-strigillose. Leaves: blades 8-24 × 5-17 cm, narrowly to broadly ovate, chartaceous at all stages; surfaces discolorous, lustrous and glabrous adaxially, finely silvery tomentellous abaxially, sometimes creamish when older, indumentum mostly concealing the surface at all stages, base truncate, obtuse or shallowly cordate, apex acute or acuminate, long-acuminate when juvenile, 3-5(-7) palmati- or shortly plinerved, innermost pair of main veins acrodromous imperfect at all stages, midrib adaxially shallowly sunken at the base, becoming flat, conspicuously raised abaxially, secondary veins 2-3(-5) pairs, arising above the middle of the blade, raised on both surfaces, more conspicuous abaxially, veinlets slightly prominent adaxially, more conspicuous abaxially; petioles (2.7-)6-25 cm long, ridged, brownish to silvery strigillose-tomentellous or glabrate, distal pulvinus more conspicuous, rugulose, rounded, sometimes more or less flat adaxially. Staminate inflorescences solitary or fascicled, cauliflorous or axillary thyrsi, brownish, greyish or silvery strigillose-tomentellous, (trichomes spreading); axes, 10-42 cm long (less frequently simple dichasia arising in young terminal shoots, then the dichasia axes 2-4 cm long) (Fig. 26A ), densely brown; primary branches 1.4-9.4 cm long, with several (2-4) branching orders, these laxly arranged; bracts 0.5-1.2 mm long, narrow ovate to ovate, concave, moderately fleshy, indumentum as on the inflorescence. Pistillate inflorescences solitary or fascicled, cauliflorous, moderately stout, few-flowered thyrsi (Fig. 27A ), brownish strigillose- tomentellous; axes ca. 2.5 cm long; primary branches 0.7 cm long; bracts ca. 0.8 mm long, ovate, concave, moderately fleshy, glabrous adaxially, brown strigillose-tomentellous abaxially. Staminate flowers 1.6-2.4 mm long, greenish, green-yellowish or whitish; pedicels 1.6-6.0 mm long, mostly slender, ridged, indumentum as on staminate inflorescence; bracteoles 1-2, 0.2-0.6 × 0.1-0.4 mm, ovate, moderately fleshy, glabrous adaxially, greyish or silvery villous-tomentellous abaxially; sepals 6, glabrous adaxially, greyish or silvery villous-tomentellous abaxially; outer sepals 0.5-1.4 × 0.3-0.8 mm, narrowly ovate or elliptic, base truncate, apex obtuse; inner sepals 1.5-2.4 × 0.8-1.8 mm, elliptic, ovate, oblong or weakly obovate, base obtuse, cuneate or shortly clawed, apex acute or obtuse, tips mostly strongly reflexed past anthesis; petals 6, 0.7-1.6 × 0.6-1.1 mm, inner ones slightly smaller and narrower, obovate-trilobed or spatulate, weakly concave, membranous, glabrous adaxially, sparse silvery tomentellous abaxially, base cuneate or distinctly short- to long-clawed, lateral margins inflexed, partially clasping the filaments, apex obtuse or truncate; stamens (5)6, filaments 0.3-)0.7-1.4 mm long, clavate or clavate-sigmoid, free (shortly connate), glabrous adaxially, glabrous to mostly silvery tomentellous abaxially; anthers 0.3-0.6 mm long, erect or weakly incurved, especially when older, connective thicker adaxially and protruding as a hump (Fig. 26G ) or as a keel at the base of thecae (also apically); thecae not separating apically and, for the most part, not immersed in the connective. Pistillate flowers ca. 2.1 mm long, green; pedicels ca. 1.1 mm long, ridged, brown strigillose-tomentellous; bracteoles 3, ca. 0.4 × 0.3 mm, ovate, fleshy, weakly concave, glabrous adaxially, brownish villose-tomentellous abaxially; sepals 6-9, weakly concave and slightly fleshy to fleshy, glabrous adaxially, brownish, greyish to silvery villous-tomentellous abaxially; outer sepals ca. 0.7 × 0.6 mm, ovate, base truncate, apex acute; middle sepals ca. 1.6 × 1.8 mm, ovate to broadly ovate, base truncate, apex acute, inner sepals ca. 2.0 × 1.3 mm, broadly ovate, obovate, spatulate, elliptic or rhombic, base obtuse or truncate, apex acute or rounded, tips strongly reflexed past anthesis; petals 3, ca. 1.6 × 1.5 mm, spatulate, membranous, weakly concave, glabrous adaxially, moderate silvery tomentellous abaxially, clawed at base, apex obtuse or retuse; carpels 3, ca. 0.7 × 0.7 mm, brown villous tomentellous; style ca. 0.7 mm long. Infructescences axes (2-)8 × 0.3-0.7cm, indumentum as on pistillate inflorescences; fruiting pedicels 1.3-3.5 mm long, terete, slender to moderately stout; carpophores 3.8-11.3 mm long, free or basally connate up to 1/3 of their lengths, claviform or terete, spreading or incurved distally, brown velutinous-hispidulous. Drupelets 1.8-2.1 × 0.9-1.3 cm, dull orange to yellowish when ripe, oblongoid, ellipsoid or subglobose (Fig. 27F ), eccentrically attached, base attenuate or truncate, (shortly stipitate); stylar scar conspicuous; exocarp 0.6-0.8 mm thick, surface smooth, rugulose or weakly muriculate, silvery to greyish velutinous; mesocarp mucilaginous; endocarp 0.7-0.9 × 1.5-1.8 cm, papyraceous to chartaceous, smooth, with slightly prominent fibres or weakly rugulose throughout. Seeds with embryo 3.2-4 cm long, cotyledons equal.

Distribution and ecology.

In north-western Amazonia, including the eastern slopes of the Andes in Peru (Fig. 25 ). The species commonly grows in periodically flooded forests, but also in non-flooded lowland forests and up to 350 (500) m in elevation. Flowering and fruiting material were collected all year round.

Common names and uses

(sterile specimens are indicated as st). Brazil: fruits edible ( Fróes 26364). Colombia: “taufe-lleida” (Diaz 36, st); “Ñamitá” (en Karijona) ( García-Barriga 14578, st); cure for fevers, antimalarial (Grassl 10076, st); said to be "formerly used by Karijona Indians in arrow-poison" ( Krukoff and Barneby 1970) (Schultes 5526, st). Ecuador: to hunt wild animals, “ambi-huasca” (Quichua) ( Cerón et al. 39668, st). Peru: “pani’ (Castelnau s.n.,1851); "abuta amarilla, “isaveño” (Huitoto) (Martin & Lau-Cam 1266, st); “ampihuasca” (Mathias & Taylor 3900, imm fr); "abuta amarilla" (Rimachi 10636, mat fr; Tina & Tello 2066, ♂ fl); “sarívana” (Weiss 132, st); “ampihuasca”, medicinal (Woytkowski 108, st); “ampihuasca” (Woytkowski 5108, st); "abuta negra" (Rimachi 10503, st).

Note.

Sterile collections such as Fox 12 (K), from Peru and Schultes 3522 from Colombia, had previously been identified as Chondrodendron toxicoferum (= Curarea toxicofera ) ( Krukoff and Barneby 1970), the first being reported as "poison used for blow pipe" and the second as a source of curare with the common name of "sa pe pa" ( Kofán) ( Krukoff and Barneby 1970). Both specimens cannot be determined with certainty and are here tentatively identified as Curarea sp.

Etymology.

Presumably in reference to the toxic effects attributed to the plant, as the type specimen was collected in the Pevas region, amongst the Yaguas who use the plant in the preparation of arrow poison ( de Castelnau 1851: 22).

Conservation status.

The Extent of Occurrence (EOO), based on 34 collections corresponding to 28 localities of C. toxicofera , is calculated as 419,469 km2 and an Area of Occupancy (AOO) of 108 km2. The 28 localities correspond to 27 subpopulations, of which two are within protected areas in Ecuador and four subpopulations are found in private reserves near Iquitos, Peru. In addition the species is widespread across its distribution. Therefore, C. toxicofera is assigned a preliminary category of "Least Concern" (LC).

Discussion.

The typical form of C. toxicofera as circumscribed here, ranges from the Amazonian lowlands of Ecuador, Colombia, Peru and Brazil (including the sterile type specimen of Cocculus toxicoferus Wedd., Castelnau s.n., Peru and the type of Hyperbaena polyantha Diels, Ule 5631, Brazil).

Individuals are frequently found in periodically flooded forests, but can also be found in non-flooded forests up to 500 m in elevation. Most have ovate leaves with 5 main veins and staminate and pistillate inflorescences with very short, brownish to greyish, appressed indumentum, but slightly longer (up to 0.3 mm long) and silvery or cream indumentum is also observed in specimens from the lower Amazon basin in Brazil (e.g. Prance 11272, Fróes 29639 and Ducke 2134).

In the staminate condition, C. toxicofera is readily distinguished from C. tomentocarpa by its brown strigillose-tomentellous indumentum (vs. a rufescent to silvery hispidulous) and its larger flowers -they are the largest in the genus- with greyish to silvery villous-tomentellous, adpressed indumentum (vs. rufescent or silvery spreading indumentum). However, separation from C. iquitana is more challenging and at present is distinguished by its slender pedicels in staminate flowers (vs. thicker in C. iquitana ). However, more field and taxonomic work remains to be done in order to be able to confidently match the staminate and pistillate specimens of the taxa involved.

Chondrodendron bioccai is here included in the synonymy with hesitation. As discussed by Krukoff and Barneby (1970), the species appears to have been described from two collections (from the region of Rio Tiquiê and the region of Rio Uaupés by Biocca and Giacone, respectively). A holotype was not stated, two photographs (listed as Figs 15 , 16 , but published as Figs 1 , 2 ) presumably of the original material were published in the protologue ( Lusina 1954). Krukoff and Barneby (1970) hesitantly placed Chondrodendron bioccai as a synonym of Chondrodendron toxicoferum and later as a synonym of Curarea toxicofera (Barneby & Krukoff, 1971). The repository of the type collection remains uncertain at the present time and hence the taxonomic identity cannot be established firmly.

Selected specimens examined.

BRAZIL. Acre. Proje. RADAM-Sub-base de Cruzeiro do Sul-Ponto 7-Sb-18-ZD, 23 Feb 1976, (imm fr), Marinho 291 (NY!) . Amazonas: Paraná do Careiro ( Boca do Solimoes ), (varzea, igapó do Lago Capitari ), 8 Jun 1948 (♂ fl), Ducke 2134 (COL!, GH!, NY!, R!); Mun. Eirunepe, Lago Dois Unidus, Ituxy, restinga, 30 Nov 1946, (♂ fl), Fróes 21802 A (NY!); Beira do lago de Badajós, igapó, 24 Ago 1950, (imm fr), Fróes 26364 (IAN!); Vicinity of Manaus, Igarapé Ipixuna, Lower Rio Negro opposite Manaus, Igapó, 1 Apr 1971, (♂ fl), Prance, et al. 11272 (F!, MG!, NY!, US!) . Rondônia: Esperança (ad ostium fluminis Javari), silva loco alto, high forest, 27 Oct 1945, (♂ fl bud), Ducke 1968 (NY!, R!); Rio Machado, curso inferior, igapó, Jan 1981, (♂ fl), Goulding 1187 (MG!) .

COLOMBIA. Amazonas: Quebrada El Mochilero, afluente del Yarí, márgenes y zona de rebalse aluvial, 150 m, 24 Apr 1986, (imm fr), Galeano et al. 1135 (COL!); Río Caquetá, La Pedrera, 1-4 Oct 1952, (st.), García-Barriga 14578 (COL!, MO!); Puerto Nariño, Trocha de Panduro a San Martin, bosque no inundable, 03°50'S; 070°20'W, 100 m, 24 Feb 1993, (♂ fl), Madriñán 702 (MO!); Leticia, Parque Nacional Natural Amacayacu, Centro Administrativo Mata-matá (Inderena), a orillas de la quebrada Mata-matá en zona de várzea, 03°47'S; 070°15'W, 100 m, 11 Mar 1991, (imm fr), Rudas et al. 1528 (MO!); Corregimiento de Tarapacá; Caño Pupuña (afluente del Rio Cotuhe ), a las orillas del caño, 02°59'S; 070°02'W, 100 m, 25 Jun 1991, (♂ fl), Rudas et al. 2512 (MO!) . Vaupés: Mayaca River, vicinity of Cachivera del Diablo and mouth of river, 300 m, 1 May 1943, (mat fr), Schultes 5526 (AMES!) .

ECUADOR. Napo: Rio Yasuni, periodically inundated forest, ca. 80 km upriver from Nuevo Rocafuerte, 225 m, 17 Sep 1977, (♂ fl bud & fl), Foster 3722 (F!) . Orellana: Cantón Nuevo Rocafuerte Isla aproximadamente 10 Ha. entre la Laguna de Jatun Cocha, cerca del Rio Yasuni , Parque Nacional Yasuni , Igapó, 00°01'S; 075°28'W, 280 m, 17 Sep 1999, (st), Cerón et al. 39668 (MO!) . Pastaza: Curaray , SE of the airstrip, rain forest and Mauritia várzea, understorey dominated of Melastomataceae , Rubiaceae and palms, 01°22'S; 076°57'W, 250 m, 20 Mar 1980, (mat fr), Holm-Nielsen et al. 22169 (AAU n.v., NY!) . Sucumbíos: Lago Agrio, Reserva Cuyabeno, Rio Cuyabeno , 2-3 km arriba de Laguna Grande , área inundada estacionalmente por aguas negras, 00°00'S; 076°14'W, 230 m, 16 Nov 1991, (♂ fl), Palacios et al. 9032 (MO!); Rio Zabalo , bosque húmedo tropical, bosque afectado por inundaciones de aguas negras, 00°22'S; 075°45'W, 230 m, 22 Nov 1991, (imm fr), Palacios et al. 9493 (MO!, NY!) .

PERU. Loreto: Yanamono, campamento Explorama Lodge, terreno de altura, 03°25'S; 072°45'W, 120 m, 31 May 1979, (mat fr), Díaz et al. 1187 (MO!); Requena , Cano Yarina , a 300 m de la Base Yarina en la Zona Reservada del Rio Pacaya , margen izquierda del Río Ucayali, bosque inundable, 6 Apr 1977, (♂ fl), Encarnación 1094 (G!, MO!, NY!, US!); Rio Itaya , inundated tahuampa forest, [03°46'S; 073°15'W], 120 m, 20 Mar 1977, (♂ fl), Gentry et al. 18505 (F!, MO!, NY!); Maynas, tahuampa near Río Amazonas between Punchana and Santa Clara de Nanay, outskirts of Iquitos, 120 m, 3 Feb 1978, (♀ fl & imm fr), Gentry et al. 21624 (F!, MO!); Alto Amazonas, Balsapuerto ( lower Rio Huallaga basin); dense forest, [05°49'S; 76°33'W], 150-350 m, 28-30 Aug 1929, (♂ fl), Killip & Smith 28665 (F!, NY!); Alto Amazonas, Fortaleza, near Yurimaguas, [05°54'S; 76°07'W], 140 m, Nov 1932, (♂ fl), Klug 2782 (BM!, F!, G!, GH!, K!, MO, NY!, US!); Distrito Las Amazonas, Quebrada Sucusari, bosque maduro en tierra firme, camino de ACEER hacia el Napo, 03°15'S; 072°55'W, 140 m, 4 Feb 1996, (♂ fl), Ortiz et al. 157 (MO!); Distrito Iquitos, Rio Itaya , San Juan de Muniches , borde de purma, 16 Oct 1976, (mat fr), Revilla & Carillo 1501 (F!, MO!, NY!); Indiana, Yanamono, Explorama Lodge, Bosque inundable estacional (aguas blancas), 106 m, 03°28'S; 072°50'W, 19 Feb 1989, (♂ fl), Vásquez & Jaramillo 11714 (MO!); Explor Napo Camp at Rio Sucusari , primary forest, flooded forest, collected from canoe along Rio Sucusari , 03°20'S; 072°55'W, 120 m, 22 Mar 1996, (♂ fl buds & fl), van der Werff & Vásquez 13990 (MO!) . Ucayali: Prov. Coronel Portillo, Distrito Calleria, Quebrada Pumayacu, margen izquierda del Rio Utiquinia , al. borde de la quebrada, en bosque alto, 150-175 m, 08°09'S; 074°15'W, 13 Ma 2003, (♂ fl bud & fl), Schunke 15323 (F n.v., MO!) .

VENEZUELA. Amazonas: Embanchure du Guaviare, Plantes du Haut Orenoque, 1887, (♂ fl), Gaillard 184 (P!); Forest immediately behind "El Tobogan de La Selva" camping area; 35 km south of Puerto Ayacucho , 85 m, 21 Feb 1979, (st), Plowman 7712 (F!) .

Figures

Figure 1. Schematic representation of affinities and drupelet: A phylogenetic affinities of Curarea and related taxa, modified from Ortiz et al. 2016, three asterisks (***) indicate 100 % support in all analyses in the referred study B schematic illustration of a Curarea drupelet showing the length and width planes measured, ca. / abbrev> = carpophore, frpe = fruiting pedicel, rs = remnant of style.

Figure 2. Transversal sections of stem of Curarea: A relatively round stem of C. aff. tomentocarpa (Ortiz 197), showing wide rays B flattened stem of C. tecunarum (Ortiz & Vasquez 214) showing successive partial rings. Scale bar: 0.5 cm (A); 1.7 cm (B).

Figure 15. Curarea crassa staminate plant: A flowering branch B detail of inflorescence C flower bud D inner sepal, adaxial surface E-F outer and inner petals, abaxial views G-H outer stamens, abaxial-lateral surfaces (based on Thomas et al. 10900).

Figure 16. Curarea crassa pistillate plant: A fruiting branch B innermost sepal, adaxial surface C petal, abaxial surface D carpels (based on Jardim et al. 609).

Figure 25. Geographic distribution of Curarea tomentocarpa and C. toxicofera.

Figure 26. Curarea toxicofera staminate plant: A flowering branch B detail of inflorescence C flower bud D inner sepal, adaxial surface E-F outer and inner petals, abaxial and latero-adaxial surfaces G-H outer stamens, lateral and abaxial surfaces (A-B based on Ule 5631 C-H based on Encarnacion 1094).

Figure 27. Curarea toxicofera pistillate plant: A flowering branch B flower bud C innermost sepal, adaxial surface D petals and carpels E petal abaxial surface F infructescence (A-E based on Gentry et al. 21624 F based on Revilla et al. 2566).