Research Article |
Corresponding author: Roy Erkens ( roy.erkens@maastrichtuniversity.nl ) Academic editor: Thomas L.P. Couvreur
© 2017 Roy Erkens, Jessica Oosterhof, Lubbert Y.Th. Westra, Paul J.M. Maas.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Erkens RHJ, Oosterhof J, Westra LYT, Maas PJM (2017) Revisions of Ruizodendron and Pseudephedranthus (Annonaceae) including a new species and an overview of most up-to-date revisions of Neotropical Annonaceae genera. PhytoKeys 86: 75-96. https://doi.org/10.3897/phytokeys.86.13773
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We present revisions of the Neotropical genera Ruizodendron and Pseudephedranthus (Annonaceae). Ruizodendron includes a single species R. ovale. Pseudephedranthus now comprises two species, with the description of the new species P. enigmaticus sp. nov. extending the range of the genus beyond the Upper Rio Negro region of Brazil (Amazonas) and adjacent Venezuela (P. fragrans), to include Guyana, Suriname, and the Brazilian state of Pará. An overview is provided of current revisions of Neotropical Annonaceae genera that will aid in accessing proper species information for this frequently encountered tropical rain forest family.
Biodiversity, Neotropics, new species, systematics, taxonomy
Annonaceae are frequent components of tropical rain forests worldwide (
With respect to Neotropical Annonaceae, renewed revisionary efforts have been underway since the early eighties (
Two genera so far still awaiting revision were the monospecific genus Ruizodendron R.E.Fr. and the non-related genus Pseudephedranthus Aristeg.
34 Neotropical genera of Annonaceae (based on
Genus: | Most up-to-date revision: |
Anaxagorea A.St.-Hil. |
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Annona L. (including Rollinia A.St.-Hil.) | Ongoing work (for inclusion of Rollinia see |
Asimina Adanson (including Deeringothamnus Small) |
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Bocagea A.St.-Hil. |
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Bocageopsis R.E.Fr. |
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Cardiopetalum Schltdl. |
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Cremastosperma R.E.Fr. |
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Cymbopetalum Benth. |
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Desmopsis Saff. | Schatz et al. in prep. (pers. comm.) |
Diclinanona Diels |
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Duckeanthus R.E.Fr. |
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Duguetia A.St.-Hil. |
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Ephedranthus S.Moore | Carvalho Lopes in prep. (pers. comm.) |
Froesiodendron R.E.Fr. |
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Fusaea (Baill.) Saff. |
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Guatteria Ruiz & Pav. |
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Hornschuchia Nees |
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Klarobelia Chatrou |
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Malmea R.E.Fr. |
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Mosannona Chatrou |
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Onychopetalum R.E.Fr. |
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Oxandra A.Rich. |
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Porcelia Ruiz & Pav. |
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Pseudephedranthus Aristeg. |
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Pseudomalmea Chatrou |
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Pseudoxandra R.E.Fr. |
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Ruizodendron R.E.Fr. | This work |
Sapranthus Seem. | Schatz et al. in prep. (pers. comm.) |
Stenanona Standl. |
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Tetrameranthus R.E.Fr. |
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Tridimeris Baill. |
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Trigynaea Schltdl. |
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Unonopsis R.E.Fr. |
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Xylopia L. | Ongoing work |
The monospecific genus Ruizodendron occurs from Amazonian Brazil to Bolivia, Peru, Ecuador, and Colombia (Fig.
a Flowering branch of Ruizodendron ovale b The asymmetrical, transversely-ellipsoid monocarps of Ruizodendron ovale (van der Werff 22749, MO). Missouri Botanical Garden, used with permission. Photograph 1b downloaded from http://www.tropicos.org. Photograph 1a taken by Abel Monteagudo, 1b taken by Rodolfo Vásquez.
Ruizodendron was not originally described as a separate genus. During the field expedition of Ruiz and Pavón in South America in 1778–1788 they collected a specimen near Pozuzo in Peru that they described as Guatteria ovalis Ruiz & Pav. (
Drawing of Ruizodendron ovale. a Top of a branch b Flower c Base of flower with part of petals removed to show the interior d Fruit e Monocarp f Monocarp in longitudinal section showing laterally attached seed g Seed viewed clockwise starting from right: longitudinal view, hilum side, and in longitudinal section. Drawings by H. Rypkema.
In 1925 the same species was collected by Steinbach in Santa Cruz in Bolivia, but also this specimen solely bore fruits. Only with the finding by Klug of a flowering specimen in the Upper (“oberen”) Río Huallaga region (San Martín; not far from the type locality of Guatteria ovalis) the placement of the genus within Annonaceae could be studied for the first time. From the flower it was clear that this indeed was not Guatteria.
More recently,
At this moment, based on molecular phylogenetic work (
Until now, Pseudephedranthus consisted of a single species: P. fragrans (R.E.Fr.) Aristeg. known only from Brazil in the state of Amazonas (Upper Rio Negro Region, Rio Cauaburí), and adjacent Venezuela (Piedra de Cucuy; Fig.
Pseudephedranthus (
Herbarium material was investigated from the following herbaria: A, AAU, CGE, E, F, G, GB, GH, INPA, K, L, LPB, MICH, MO, P, NY, RB, S, U, UC, US, WAG, and WIS. Dried herbarium material was used for measurements, colour indications and descriptions of surface structures. If measurements were done on material kept in spirit this is indicated by curly brackets { }. We have indicated the density of hair cover according to the following gradations: densely, rather densely and sparsely.
Ruizodendron R.E.Fr., Arkiv för Botanik Ser. 28B (4): 3. 1936.
Ruizodendron ovale (Ruiz & Pav.) R.E.Fr. (= Guatteria ovalis Ruiz & Pav.)
Trees, often with buttresses; young twigs lenticellate. Leaves: distichous, simple, entire, shortly petiolate, exstipulate; lamina medium-sized, mostly elliptic, venation brochidodromous to eucamptodromous, articulated at the base, primary vein slightly raised to flat above, glaucous below. Flowers actinomorphic, bisexual, perianth consisting of one whorl of sepals and two whorls of petals; pedicels with basal articulation and 2 soon falling bracts; sepals 3, imbricate, free, much shorter than the petals, soon falling; petals 6, imbricate, free, subequal; stamens ca. 40, extrorse, filament very short, connective apex discoid, glabrous; carpels ca. 25, spirally arranged, free, ovary 1-locular with 1 ovule attached below the middle, style absent, stigma ovoid, hairy. Fruit apocarpous, composed of few indehiscent monocarps, these transversely ellipsoid, the stipe laterally attached. Seed 1, filling the whole monocarp, orange-brown, rumination lamellate in 2 parts, raphe a fine, shallow groove.
Monotypic, occurring in the western part of South America (Fig.
Ruizodendron ovale (Ruiz & Pav.) R.E.Fr., Arkiv för Botanik Ser. 28B (4): 3. 1936. – Guatteria ovalis Ruiz & Pav., Systema vegetabilium florae peruvianae et chilensis 1: 146. 1798.
PERU. Huánuco: Pozuzo, anno 1782, Ruiz & Pavón s.n. (holotype MA [barcode MA811639], only photograph seen; isotype B).
Tree, 5–45 m tall, 10–70 cm diam., buttresses to ca. 1.5 m high, young twigs longitudinally fissured, sparsely covered with appressed, silvery hairs 0.1–0.4 mm long, soon glabrous. Leaves: petioles 5–10(–15) by 1–2 mm; lamina elliptic to ovate, rarely narrowly so, 6–25 by 4–12 cm (index 1.5–2(–3)), chartaceous, olive green to greenish brown and shiny above in sicco, glaucous to pale brown below, glabrous above, sparsely covered with appressed, silvery hairs to ca. 0.4 mm long mainly on larger veins below, base rounded with the extreme base slightly attenuate; apex obtuse, rarely acute; secondary veins 7–14 on either side of primary vein, slightly raised to flat above, shortest distance between secondary veins and margin 1–3 mm, tertiary veins strongly raised above, percurrent to reticulate. Inflorescences axillary, 1-flowered, rarely 2 together through auxiliary bud formation, often appearing on small, leafless lateral branches. Indument: pedicels, outer side of bracts and sepals densely covered with appressed, silvery hairs to ca. 0.4 mm long, petals sparsely so; pedicels 4–8 by 1 mm, fruiting pedicels to ca. 10 by 1–3 mm; bracts 2, soon falling, one bract placed just below the articulation, ovate, ca. 4 mm long, other bract placed at ca. 0.2 from the base of the pedicel, ovate-triangular, ca. 4 mm long; sepals ovate-triangular, 5–6 by 3–4 mm, soon falling; petals white to cream in vivo, black in sicco, narrowly elliptic, 20–50 by 3–5 mm, membranous, base widened enclosing the pollination chamber; torus slightly raised; stamens ca. 1.5 mm long; carpels ca. 1 mm long. Monocarps 1–10, green, maturing black in vivo, brownish black in sicco, transversely ellipsoid, 15–20{–25} by 10–16 mm, glabrous, apex excentrically apiculate (apiculum to ca. 1 mm long), wall 0.2–0.5 mm thick; stipes 8–25 by 1–1.5 mm, excentrically placed. Seed transversely ellipsoid, 14–22 by 9–13 mm, orange-brown.
Colombia (Amazonas, Cundinamarca, Meta), Ecuador (Morona-Santiago, Napo, Orellana, Pastaza, Sucumbios), Peru (Huánuco, Loreto, Madre de Dios, Pasco, San Martín, Ucayali), Bolivia (Beni, Cochabamba (?), La Paz, Pando, Santa Cruz), Brazil (Acre). Fig.
In tropical rain forest, often on alluvial soil. At elevations of up to 800 m. Flowering: July to September; fruiting: August to May.
Bolivia: Blanquillo (Vargas C. et al. 1133), Fariña seca de bajío (Dominguez & Gonzales 42), Ojojo blanco (DeWalt et al. 283, Neill & Quevedo 9386, Seidel et al. 5640), Ojoso (Vargas C. et al. 5366), Ojoso blanco (Del Aguila et al. 95), Ojoso negro (Cruz & Terceros 208, Meneces & Terceros 473, D.N. Smith et al. 14160, 14194A, 14215, 14346), Palo fino (Cruz & Medina 153), Periquina hoja menuda (Meneces & Hartshorn 777); Tumuqui (Tacana name) (DeWalt et al. 283). Brazil: Envira orelha de onça (Silveira et al. 1094), Piaçaba (Campbell et al. 9473a). Ecuador: Aparina cara caspi (Quichua name) (Zuleta 364), Shampipian (Ortega U. 233). Peru: Ochabaja (Begazo 61), Panjil ruro (Albán C. 5878, Schunke V. 1203), Paujil ruro (Tello 257), Paujil ruro amarillo (Kröll 217).
Bolivia. Beni: Prov. Moxos, 130 km S of San Ignacio, Del Aguila et al. 95 (U), 27 km from San Borja, along road to Trinidad, 260 m, D.N. Smith et al. 14160 (U), 14194A (U), 14215 (MO); Prov. Ballivian, 35 km on the road from Yucumo to Rurrenabaque, 230 m, D.N. Smith et al. 14346 (MO). La Paz: Prov. Iturralde, Comunidad de Santa Fé, 250 m, DeWalt et al. 510 (U), Upper Madidi ridge top, 7 km NE of camp, 370–380 m, Gentry et al. 70691 (MO); Prov. Iturralde, Ixiamas, 90 m, Couvreur & Vargas 213 (MO, RB); Prov. Sur Yungas, Basin of Río Bopi, Asunta, near Evenay, 690–750 m, Krukoff 10669 (A, F, G, K, MICH, MO, NY, S, U, UC, US); Prov. Sur Yungas, Upper Río Beni, concession of cooperativa Sapecho, 600 m, Seidel et al. 5640 (U). Pando: Prov. Manuripi, Batraja, 35 km E of Puerto Rico, 150 m, Chatrou et al. 453, 457 (U). Santa Cruz: San Rafael de Amboró, Centro de Estudios Ambientales, 400 m, Maas et al. 8762 (U); Prov. Ichilo, Parque Nacional Amboró, Campamento Molle, San Germán, road from Yapacaní to Río Ichilo, 400 m, Maas et al. 8773 (U); Prov. Ichilo, Reserve Forestal `El Choré’, 240 m, Meneces et al 473, 476 (U); Prov. Ichilo, Parque Nacional Amboró, 23 km S of Buena Vista, along Río Chonta, 420 m, Nee 36863 (LPB, MO, NY, U); Prov. Ichilo, Parque Nacional Amboró, 4 km SW of Buena Vista, S side of Río Surutu, 315 m, Nee 39391 (MO, NY, U, WIS); Prov. Ichilo, Reserva Forestal Choré, Río Ibabo, bosque experimental “Elias Meneces”, 180 m, Neill et al. 9386 (G, MO); Prov. Sara, 400 m, Steinbach 7248 (A, E, F, G, K, LPB, MO, NY, U); Prov. Ichilo, Parque Nacional Amboró, Río Yapojé and Saguayo, 8 km SW of El Carmén, 360 m, Vargas C. et al. 1129 (MO), 1133 (NY, MO); Prov. Ichilo, El Carmén, 8 km SSW of Buena Vista, on the way to Huayti, 400 m, Vargas C. & Mendéz F. 5366 (U). Brazil. Acre: Upper Rio Moa near Fazenda Arizona, Campbell et al. 6196 (U); Fazenda Bom Sossego, between Igarapé Cujubim and Igarapé Jacamin, Campbell et al. 8633, 9473a (U); Mun. Cruzeiro do Sul, Igarapé Humaitá, afflente do Rio Juruá, Colocação Várzea Grande, Cid et el. 10453 U); Mun. Cruzeiro do Sul, Rio Juruá, entrada do Igarapé Viseu, Cid et al.10535 (U); Mun. Cruzeiro do Sul, Upper Rio Juruá, Fazenda Calila Sara, Cid et al. 10839 (MO, U), Mun. Sena Madureira, Basin of Rio Purus, Rio Iaco, right bank, Nova Olinda, between Igarapé Santo Antonio and Igarapé Boa Esperança, Daly et al. 7909 (U); Mun. Marechal Thaumaturgo, Basin of Rio Juruá, Rio Bagé, Daly et al. 10245 (U); Mun. Manoel Urbano, Rio Chandles (tributary of Rio Purus), downstream from Igarapé Canamarí, Daly et al. 11234 (U); Colonia Cinco Mil (“Seita do Cipó”), Nelson 818 (MICH, MO, NY, RB, U, US); Mun. Xapuri, Reserva Extrativista Chico Mendes, Serigal Floresta, Colocação Rio Branco, Figueiredo 270 (U); Tarauacá, Bacia do Alto Juruá, Rio Tarauacá, Reserva Indígena Praia do Carapanã, Colocação Nova Morada. Silveira et al. 1094 (U). Colombia. Linden s.n. (CGE); Cundinamarca: Mun. Yacopí, vereda El Morro, 405 m, Rangel et al. 13563 (U). Meta: Parque Nacional Natural Tinigua, Serranía Chamusa, Centro de Investigaciónes Primatológicas La Macarena, Stevenson 299 (COL). Ecuador. Napo: Parque Nacional Yasuní, Pozo petrolero Daimï, 200 m, Cerón et al. 4227 (MO); Cantón Tena, Estación Biológica Jatun Sacha, 8 km E of Misahuallí, 400 m, Cerón et al. 5976 (G, GB, MO, U), Cantón Aguarico, Reserva Etnica Huaorani, 260 m, Dik 1220 (U), Parque Nacional Yasuní, Pozo petrolero Daimï I, Conoco, 230 m, Hurtado et a1. 21 (MO, U); Cantón Tena, Estación Biológica Jatun Sacha, 8 km E of Misahuallí, 450 m, Maas et al. 8600 (U), Río Napo, 8 km down the river from Puerto Misahuallí, 400 m, Neill & Palacios 7130 (U); Reserva Biológica Jatun Sacha, Río Napo, 8 km Río E of Misahualli, 450 m, Neill & Manning 8014 (U); Estación Experimental INIAP Payamino, 5 km NE of Coca, 250 m, Palacios et al. 1036 (AAU, F, U); Reserva Biológica Jatun Sacha, Río Napo, 8 km down the river from Misahualli, on right side, 450 m, Palacios 1421 (U, MO); Cantón Loreto, Payamino-Loreto road, Comunidad Jumandi, 350 m, Palacios 10950 (F, U); 30 km NNW of Coca, Río Huashito, Site of Proyecto Huashito, 250 m, T.D. Pennington 10609 (U); Cantón Tena, Estación Biológica Jatun Sacha, 400 m, Revelo 65 (U); Cantón Orellana, Sector Huashito, 20 km N of Coca, Propriedad de Palmoriente, 250 m, Rubio 249 (G, GB, K, MO); Estación Biológica Jatun Sacha, confluence of Río Arajuno and Río Puní, 450 m, Zuleta 39 (U), Estación Biológica Jatun Sacha, 400 m, Zuleta 364 (U). Pastaza: Cantón Arajuno, Campamentos temporales 11–12, 785 m, Freire et al. 3468 (U). Sucumbios: Reserva de Producción Faunistica Cuyabeno, 1 km N of Laguna Grande, 265 m, Valencia et al. 68558 (AAU). Peru. Huánuco: Tingo Maria, Asplund 13193 (G, NY, S); Prov. Puerto Inca, Distr. Yuyapichis, Dantas, 270 m, Kröll 217 (G, NY), 569 (G), 623 (G), 781 (G); Prov. Pachitea, Distr. Honoria, Schunke V. 1203 (F, G, GH, K, MO, NY); Distr. Yuyapichis, Prov. Puerto Inca, Unidad Modelo de Manejo y Producción Forestal Dantas, 270 m, Tello 257 (G, MO, ibidem, Timaná 143 (G); Shapajilla, 630 m, Woytkowski 32 (F). Loreto: Prov. Maynas, Yanamono, Explorama Tourist Camp, Río Amazonas halfway between Indiana and mouth of Río Napo, 130 m, Gentry et al. 42403 (MO). Madre de Dios: Prov. Tambopata, Río Heath, Pto. San Antonio, 210 m, Aguilar & Castro 916 (U), 1076 (U), Communidad Nativa, Sonene, ca 8 km above confluence with Río Madre de Dios, 180 m, Alexiades et al. 1210 (U); Prov. Manú, Parque Nacional de Manu, Estación Biológico Cocha Cashu, Chatrou et al. 157 (U), Parque Nacional “Bahuaja-Sonene”, Díaz et al. 9037 (U), Díaz & Ramírez 9548 (U); Cashucocha Camp, Río Manu, Parque Nacional de Manu, 380 m, Gentry et al. 26821 (U), 26952 (F); Cocha Cashu Biológical Station, Parque Nacional de Manu, 400 m, Gentry 43334 (U), Gentry 43537 (MO); Tambopata Nature Reserve, junction of Río La Torre and Río Tambopata, 250 m, Gentry et al. 57751 (MO); Prov. Tambopata, 15 km ENE of Puerto Maldonado, Cusco Amazónico, 200 m, Gentry et al. 68713 (MO); Prov. Manu, Puerto Maldonado, Los Amigos Biological Station, Madre de Dios River, Janovec et al. 2014 (U), ibidem, 270 m, Maceda 7 (U); Prov. Tambopata, Cusco Amazónico, 200 m, Núnez et al. 10607 (MO); Prov. Manu, Río Manu, Pakitsa - behind station, 350 m, Sobrevila et al. 1782 (F); Prov. Tambopata, Las Piedras, Cusco Amazónico, 200 m, Timaná et al. 1722, 2795, 3060 (MO). San Martín: Prov. Huallaga, Barranca, 380 m, Albán C. 5878 (F); Juanjuí, Alto Río Huallaga, 400 m, Klug 3798 (F, GH, K, MO, NY, S); Moyobamba - Tarapoto, km 20, 1000 m, T.D. Pennington & Daza 16708 (U; Prov. Mariscal Caceres, Distr. Tocache Nuevo, Quebrada de Saule Grande (margen derecho del Río Huallaga), Schunke V. 4387 (G, GH, K, MO, NY, US). Ucayali: Prov. Pucallpa, Estación Experimental Alexander von Humboldt, Begazo 61, 99 (U); Prov. Coronel Portillo, Humboldt station, near Campo Verde, ca. 34 km SW of Pucallpa, 200 m, Morawetz & Wallnöfer 11-19985 (U); Prov. Coronel Portillo, Bosque Nacional Alexander von Humboldt, km 68 of Pucallpa-Tingo Maria road, 270 m, Oliveira 15 (MO); Prov. Ucayali, Distr. Pampa Hermosa, Pampa Hermosa, 240 m, Reynel R. 832 (U); Prov. Coronel Portillo, Bosque Nacional Alexander von Humboldt, km 68 of Pucallpa-Tingo Maria road, 270 m, Salazar 32 (MO); Canchahuayo, Río Ucayali, 500 m, Vásquez et al. 6958 (AAU, F, NY, MO, U).
This species is easily recognizable by a combination of oval and on both sides rounded leaves with an olive green colour, long and narrow petals (often drying blackish), and stipes and apicules that are excentrically placed.
Maas et al. 8773 from Bolivia: Inner layer of monocarps ca. 5 mm thick, orange-yellow, sweet and edible.
Pseudephedranthus Aristeg., Memoirs of the New York Botanical Garden 18(2): 43. f. 10. 1969.
Pseudephedranthus fragrans (R.E.Fr.) Aristeg. (= Ephedranthus fragrans R.E.Fr.).
Trees; young twigs glabrous. Leaves: distichous, simple, entire, shortly petiolate, exstipulate; lamina medium-sized, elliptic, venation brochidodromous, primary vein raised above. Inflorescences axillary, 1–4-flowered with 2nd order flowers originating from axils of lower bracts (or possibly also through accessory buds), often persisting on older leafless branchlets, pedicels with articulation in lower part and with 3–5 bracts, the uppermost bract above the articulation. Flowers actinomorphic, bisexual or staminate (androdioecious), 3-merous, perianth consisting of one whorl of sepals and two whorls of petals; sepals 3, valvate, basally connate, much shorter than the petals; petals 6, imbricate, elliptic, free, subequal; staminate flowers: torus conical, stamens numerous, extrorse, filament very short, apical prolongation of connective broadly ovoid in basal stamens to discoid in distal stamens; bisexual flowers: torus slightly raised, stamens numerous, but less so than in staminate flowers, apical prolongation of connective broadly ovoid; carpels numerous, spirally arranged, free, ovary 1-locular, with 1 basal ovule, style absent, stigma ovoid, papillate. Fruit apocarpous, composed of few, indehiscent monocarps, these ellipsoid, distinctly stipitate. Seed 1, pale brown, rumination lamellate in 2–4 parts, raphe a distinct groove.
Two species in the Amazon regions of Venezuela and Brazil, and in Guyana and Suriname.
1a | Petioles 3–5 mm long; leaf base acute to attenuate; outer side of outer petals (rather) densely covered with appressed hairs, inner side of outer petals and both sides of inner petals densely covered with white or greyish white curly hairs (Suriname, and the Brazilian state of Pará) P. enigmaticus | |
1b | Petioles 8–12 mm long; leaf base obtuse to less often acute, the extreme base shortly atttenuate or not; outer side of outer and inner petals densely covered with appressed, not curly hairs to ca. 0.1 mm, inner side of outer and inner petals sparsely covered with appressed hairs or glabrous (Upper Río Negro region of Brazil and Venezuela) | P. fragrans |
Differing from P. fragrans by shorter petioles. Moreover, petals in P. enigmaticus are for a large part covered by a very dense indument of curly hairs, those of P. fragrans covered by a less dense indument of appressed hairs. Also, seeds in P. enigmaticus are ellipsoid instead of ovoid and smaller than in P. fragrans.
SURINAME, Sipaliwini, Central Suriname Nature Reserve, ca. 4 km ENE of Kayserberg Airstrip, alt. 235 m, 4 June 2003, Evans et al. 3437 (holotype WAG! [barcode WAG.1584983]; isotype L! [barcode L.3724851]).
Tree, 3–15 m tall, 12–20 cm diam.; young twigs glabrous. Leaves: petioles 3–5 by 1–2 mm; lamina narrowly elliptic, 12–22(–26) by 4–6(–9) cm (index 2.8–4), chartaceous, pale gray to greenish gray above in sicco, somewhat bullate above in vivo, greenish brown to pale brown below in sicco, base acute, apex acuminate (acumen 5–10 mm long), primary vein raised above, secondary veins 6–10 on either side of primary vein, raised above, smallest distance between secondary veins and margin 4–7 mm, tertiary veins raised, rarely flat above, reticulate. Only staminate flowers seen, Inflorescence axillary, 1–2(–several)-flowered, pedicels 3–12 mm by 0.5–2 mm, rather densely to sparsely covered with erect to appressed, brown hairs to ca. 1 mm long, soon glabrous; bracts 4–5, depressed ovate, 1–2 mm long, outer side rather densely to sparsely covered with erect to appressed, brown hairs; flower buds ellipsoid; sepals shallowly ovate-triangular, ca. 2 by 2–3 mm, outer side rather densely to sparsely covered with erect to appressed, brown hairs; petals white, tinged with pale green in vivo, oblong-elliptic to narrowly so, 7–12 by 3–6 mm, outer side of outer petals densely to rather densely covered with appressed, brown hairs, inner side densely covered with whitish or greyish-white, curly hairs except for the glabrous base, outer side and apical part of inner petals densely covered with curly, white hairs; staminate torus conical, 2–2.5 mm long, ca. 1 mm diam. at base; stamens ca. 50, 2–2.5 mm long, apical prolongation of connective discoid, broadly elliptic. Monocarps 3–15, green in vivo, black in sicco, ellipsoid, 12–32 by 7–15 mm, glabrous or sparsely covered with appressed hairs, apex rounded, wall 0.2–0.5 mm thick, stipes 1–4 mm long, 1–1.5 mm diam. Seed ellipsoid, 12–19 by 7–10 mm, pale brown, transversely striate.
Guyana, Suriname, and the Brazilian state of Pará. Fig.
In periodically inundated or non-inundated forest, on sandy or loamy soil, alt. 100–600 m. Flowering: May, June; fruiting: June, July, September.
Brazil. Pará: Parque Indígena do Tumucumaque, Rio Parú de Oeste, Missão Tiriyo, Cavalcante 2579 (U); Rio Maicuru, Igarapé do Mutum, 31/2 hrs. por canoa de motor de poupa acima da pista de pouso do Lageiro, Jangoux & Ribeiro 1555 (L, RB); W bank of Rio Maicuru, ca. 23 km upstream from Lageira airstrip, N side of Mutum stream, Strudwick et al. 3808 (U). Guyana. Takutu-U Region, Rupununi River, between Kwattamang Landing and Rewa Village, 100 m, Clarke et al. 6750 (NY, US). Suriname. Suriname, Sipaliwini District, Sipaliwini River, Werehpai, 5 September 2010, Bánki et al. 1674 (L); Sipaliwini District, Sipaliwini River, Bánki et al. 1579 (L); Sipaliwini, vicinity of camp on W bank of Zuid River, across river (i.e. W and outside of) Central Suriname Nature Reserve, ca. 10 km straight-line distance SSE of Kayserberg Airstrip, 240 m, Evans et al. 3485 (L); Sipaliwini, Central Suriname Nature Reserve, on S slope of the first peak in Eilerts de Haan mountain range, ca. 7 km ENE of Kayserberg Airstrip, 400–600 m, Herrera C. et al. 9959 (L, WAG); Distr. Nickerie, area of Kabalebo Dam project, 30–130 m, Lindeman & de Roon 752 (U); Sipaliwini, Morro Grande camp forest island, 6 km W of Morro Grande dome, 360 m, Oldenburger et al. 416 (U); Sipaliwini, Central Suriname Nature Reserve, 2–5 km SE of E end of Kayserberg Airstrip, 235 m, Rosário 1796 (L); Sipaliwini, Central Suriname Nature Reserve, 2–5 km ENE of Kayserberg Airstrip, 235 m, Rosário et al. 1829 (MO); Sipaliwini, Central Suriname Nature Reserve, vicinity of camp at southern base of the first peak in Eilerts de Haan mountain range, 250–350 m, C.S. & D.O. Rosário 2176 (L).
Material of this species had previously been filed in herbaria under different generic names such as Cremastosperma, Guatteria, Klarobelia, Malmea, Oxandra, and Rollinia (which is quite aberrant!). The confusion is aptly expressed in the epithet “enigmaticus”. This new species fits quite well, however, within the genus Pseudephedranthus (segregated from Ephedranthus by Aristeguieta in
The specimens investigated here were either fruiting or flowering, the flowers all being staminate. Carpel bearing flowers are still needed to complete the description.
Drawing of Pseudephedranthus enigmaticus. a Flowering branch (Clarke 3420, U) b Flower in lateral view c Staminate flower in longitudinal section d Outer petal e Inner petal f Stamen (b–fEvans et al. 3437, WAG) g Fruit (Herrera C. 9959, WAG) h Seed, small part of seed coat removed to show lamellate ruminations i Cross section of monocarp and enclosed seed showing 4-parted rumination (h–iStrudwick et al. 3808, U). Drawings by Esmée Winkel.
Pseudephedranthus fragrans (R.E.Fr.) Aristeg., Memoirs of the New York Botanical Garden 18(2): 43. f. 10. 1969.
Ephedranthus fragrans R.E.Fr., Memoirs of the New York Botanical Garden 9(3): 327. 1957.
VENEZUELA. Amazonas: Río Negro, occasional on lower slopes of Piedra Nunca, just N of Piedra de Cucuy, 100–150 m, 10 April 1953, Maguire & Wurdack 34954 (holotype S [barcode S.85211]; isotypes F, GH, NY, P, S).
Tree, 10–20 m tall, 15–30 cm diam.; young twigs glabrous, fissured, covered with a white, waxy surface. Leaves: petioles 8–12 by 1–2 mm; lamina lamina elliptic to narrowly elliptic, (5–)10–25 by (2.5–)4–7(–9) cm (index 2–3), chartaceous to coriaceous, dark greenish-brown above, pale green below, shiny and glabrous on both sides, base obtuse to less often acute, the extreme base shortly attenuate or not, apex shortly acuminate (acumen to ca. 15 mm long), secondary veins 5–9 on either side of primary vein, raised above, smallest distance between secondary veins and margin 2–6 mm, tertiary veins raised above, reticulate. Inflorescence axillary, 1–2(–more)-flowered, usually on older parts of branchlets; pedicels 4–8 by ca. 1 mm, fruiting pedicels to ca. 10 by 2–4 mm, densely covered with appressed hairs to ca. 0.1 mm long; bracts 3–5, triangular, 0.7–1 mm long, outer side densely covered with appressed hairs to ca. 0.1 mm long; flower buds broadly ovoid; sepals shallowly ovate-triangular, 1.5–2 by 3 mm, outer side densely covered with appressed hairs to ca. 0.1 mm long; petals white to cream in vivo, narrowly elliptic, 9–15 by 3–5 mm, outer side densely covered with appressed hairs to ca. 0.1 mm long; staminate flowers: torus concave, ca. 2.5 mm long, stamens ca. 50, 1.5–2 mm long, basal stamens with a broadly ovoid prolongation of connective, the distal ones with a discoid prolongation of connective, carpels absent; bisexual flowers: torus slightly raised, ca. 1 mm long, stamens ca. 15, 1.5–2.5 mm long, apical prolongation of connective broadly ovoid, carpels ca. 25, ca. 3 mm long, densely covered with appressed hairs to ca. 0.1 mm long, stigma ca. 1 mm long. Monocarps 3–10, green, maturing orange in vivo, pale brown in sicco, ellipsoid, 25–32 by 12–15 mm, apex rounded, sparsely covered with appressed hairs to ca. 0.4 mm long to glabrous, wall 0.2{–1} mm thick; stipes 2–4 by 1.5 mm. Seed ovoid, 25–30 by 13–15 mm, pale brown, transversely striate.
Restricted to the Upper Rio Negro region of Brazil and adjacent Venezuela. Fig.
In lowland rain forest (in forested hills at the base of Piedra de Cucuy, Maas, pers. comm.). At elevations of 50–600 m. Flowering: April, December; fruiting: October, December.
Venezuela. Amazonas: Río Negro, forested base of Piedra de Cucuy, 100–200 m, Maas et al. 6878 (INPA, MO, NY, RB, S, U, WU). Brazil. Amazonas: Rio Cauaburí, Caño Tucano, 125 m, Maguire et al. 60189 (F, GH, MO, NY, S), 60191 (MO, NY, S, US); Rio Negro, Rio Cauaburí, Rio Tucano, Camp Tucano (Camp No. 2), 850 ft., Maguire et al. 60332 (MO); between Palmito Camp and Camp Tatu, 400–600 m, N.T. Silva & Brazão 60683 (NY, US); Rio Negro, Rio Cauaburí, Rio Maturacá, between Camp III and Maloca, N.T. Silva & Brazão 60738 (NY).
We thank Esmée Winkel (Naturalis Biodiversity Center) who prepared the drawing of Pseudephedranthus enigmaticus in this manuscript. We also are grateful that Rodolfo Vásquez (MBG) makes his photographs of Ruizodendron ovale available via the Tropicos website (www.tropicos.org) and that Abel Monteagudo allowed publication of his photograph as well. Furthermore, we thank the curators of the following herbaria for allowing us to borrow and/or study material: A, AAU, CGE, E, F, G, GB, GH, INPA, K, L, LPB, MICH, MO, P, NY, S, U, UC, US, WAG, and WIS. Finally we wish to thank Richard Saunders, Adriana Lobão, and an anonymous reviewer for reviewing the document and providing us with useful feedback to improve our manuscript.