Gaulettia canomensis (Mart.) Sothers & Prance, 2014

Sothers, Cynthia, Prance, Ghillean T., Buerki, Sven, Kok, Rogier De & Chase, Mark W., 2014, Taxonomic novelties in Neotropical Chrysobalanaceae: towards a monophyletic Couepia, Phytotaxa 172 (2), pp. 176-200 : 184-192

Gaulettia canomensis (Mart.) Sothers & Prance , comb. nov.

Basionym:— Moquilea canomensis Martius (1827: 80) .— Hirtella canomensis (Mart.) Sprengel (1827: 341) .— Couepia canomensis (Mart.) Benth. Ex Hooker (1867: 42) . Type:— BRAZIL. Amazonas: Canuma, without date, Martius s.n. (holotype M!, isotypes BR!, G!, K!).

Parinari pilosa Standley (1937: 257) . Type :— BRAZIL. Amazonas : Mun. de Humaitá, plateau between Rios Livramento and Ipixuna, 7–18 November 1934, Krukoff 7109 (holotype F!, isotypes A!, BM!, BR!, G!, IAN!, K!, LE, MICH, MO!, NY!, RB!, U!).

Shrub or small tree to 10 m tall. Leaves ovate to oblong elliptic, 8.5–18.0 × 3.0–8.0 cm, prominently reticulate beneath with short grey-brown pubescence between reticulations. Inflorescences densely flowered axillary racemes. Receptacle subcampanulate-cylindrical, 5.5–10.0 mm long, dense ferrugineous-brown pubescent on exterior.Stamens 40–51, inserted in a nearly complete circle, with a short row of toothed staminodes opposite them. Fruit round to oblong/pyriform, 3.5–5.0 × 2.5–3.5 cm; epicarp short-velutinous; endocarp fragile, granular, sparsely-pubescent within. Fig. 6J .

Distribution and habitat: —Western and central Amazonia and Guyana, Colombia, Peru, Brazil (Amazonas, Pará, Rondônia) and Venezuela. Primary and secondary forest and clearings on non-flooded ground. Fig. 4 .

Conservation status: —The conservation status of this species is here evaluated as least concern, LC ( IUCN 2001). This is a widespread species occurring both in primary and secondary forests throughout the Amazon region. This species was previously assessed as LC for Colombia ( Calderón et al. 2002).

Selected specimens examined: — GUYANA. Rupununi District: Kanuku Mts., Two-Head Mt. , 28 January 1994, Jansen-Jacobs et al. 3434 ( K!, NY!, P!, U!) . VENEZUELA. Amazonas: Savannas of Santo Antonio, Río Orinoco , date unknown, Williams 15055 ( F!, US!, VEN!) . COLOMBIA. Vaupés: Mitú, lower Río Kubiyú, 17 April 1975, Zarucchi 1221 ( ECON!, NY!) . Caquetá: Araracuara , 28 January 1989, Gentry et al. 65297 ( K!, MO!) . BRAZIL. Amazonas: Mun. de São Paulo de Olivença, Estrada Bonfim , 23 November 1986, Daly et al. 4408 ( INPA!, K!, NY!); Manaus, Reserva Florestal Ducke , 03 December 1997, Souza 467 ( INPA!, K!, NY!) . Pará: Alto Rio Tapajós , São Raimundo, 23 May 1977, Rosa & Santos 1926 ( FHO!, MG!, NY!) . Rondônia: Mun. de Porto Velho , Samuel Dam area , 13 June 1986, Cid Ferreira 7427 ( INPA!, NY!) .

2d. Gaulettia cognata (Steud.) Sothers & Prance , comb. nov.

Basionym: Hirtella cognata Steudel (1843: 761) .— Couepia steudeliana Miquel (1851: 28) , nom. illeg. superfl.— Moquilea steudeliana (Miq.) Walpers (1852: 463) , nom. illeg. superfl.— Couepia cognata (Steud.) Fritsch (1889: 60) .— Parinari hostmannii Fritsch (1890: 13) , as ‘ Parinarium hostmanni’, nom. illeg. superfl. Type:— SURINAME. 1843, Hostmann 795 (holotype P!, isotypes BM!, CGE!, G!, GH!, K!, LE, NY!, OXF!, TCD!, U!, W!).

Shrub or small tree, 1–10 m tall (rarely large trees to 35 m). Leaves oblong to oblong-lanceolate, 2.5–6.0(–15.0) × 1.0–2.0(–4.5) cm, with grey- to brown-lanate pubescence beneath that usually obscures the prominently reticulate venation. Inflorescences terminal panicles, the axis and branches brown-villous. Receptacle cylindrical, 5–8 mm long, reddish-brown and often villous on exterior; petals white or pale pink. Stamens 12–25, unilateral, with a few teeth-like staminodes opposite them. Fruit globose; epicarp pubescent when young, verrucose and glabrous when mature.

Distribution and habitat: — Gaulettia cognata is distributed in Guyana, Suriname, Brazil (Amazonas) and Venezuela. It is found in savanna, savanna forest, scrubland, wallaba forest, primary and secondary forest, and tepuis (1100 m), as well as lower montane forests (200–300 m). Fig. 5 .

Conservation status: — Gaulettia cognata has a wide distribution in northern Brazil, Venezuela and the Guianas but often occurs in restricted habitat types, from ca. 50–1000 m. It is therefore here assessed as NT ( IUCN 2001 ) .

Selected specimens examined: — GUYANA. Upper Demerara-Berbice District: E of Linden, junction of road to Mabura Hill , 25 March 1987, Boom 7145 ( NY!, P!) . Demerara-Mahaica District: Linden Highway, 18 October 1997, Jansen-Jacobs & ter Welle 5616 ( K!, U!) . SURINAME. Jodensavanna , without date, BW 5229 ( A!, K!, MO!, NY!) . VENEZUELA. Bolívar: Mun. Raúl Leoni, 5 km from base of Cerro Camarón , 42.5 km SE of Entre Rios, 30 October–2 November 1988, Aymard & Fernández 7173 ( K!, MO!, PORT) . BRAZIL. Amazonas: Rio Içana, near Tunui , 28 March 1952, Fróes 28080 ( IAN!, NY!, UB!) .

Notes: —Three varieties of Gaulettia [ Couepia ] cognata have been recognized in the past ( Prance 1972a, 1974b, Prance & Sothers 2003b), but currently their taxonomy is uncertain and should be evaluated in light of more recently available material. The three varieties previously recognized are: var. cognata, var. major Prance (1972a: 249) and var. membranacea Prance (1972a: 249) .

2e. Gaulettia elata (Ducke) Sothers & Prance , comb. nov.

Basionym : Couepia elata Ducke (1935: 35) . Type :— BRAZIL. Amazonas : Manaus, 09 May 1932, Ducke RB 25001 (holotype RB!, isotypes K!, P!, U!) .

Tree to 40 m in height. Leaves ovate 4.0–7.0 × 2.5–3.0 cm, prominently reticulate beneath with silver-grey pubescence between reticulations. Inflorescences few-flowered lax racemes, with grey pubescence. Receptacle obconical-turbinate, 4–5 mm long, sparsely pubescent on exterior. Stamens 35–40, inserted in ½ a circle, with a row of short-toothed staminodes opposite them. Fruit globose, 5–6 cm in diameter, with a densely verrucose epicarp; endocarp fragile, granular. Fig. 6A–I .

Distribution and habitat: —French Guiana, Guyana, western and central Amazonian Brazil (Acre, Amazonas) and Colombia. Found in non-flooded forests. Fig. 7 .

Conservation status: —This species is primarily a terra firme forest canopy tree distributed over an extensive part of the Amazon region. Despite being infrequently collected, it is assessed here as least concern, LC, according to the IUCN criteria ( IUCN 2001). This species was assessed as NT (VU) for Colombia ( Calderón et al. 2002).

Selected specimens examined: — FRENCH GUIANA. Sinnamary R., above Petit Saut, 07 September 1993, Mori et al. 23651 ( K!, NY!). GUYANA. Cuyuni-Mazaruni Region: Pakaraima Mts. , Karowrieng R., 1 km SE of Maipuri Falls, 15 October 1992, Hoffman 3036 ( K!, NY!, US!) . COLOMBIA. Amazonas: Río Cahuina , 07 September 1988, Sánchez et al. 980 ( K!) . BRAZIL. Amazonas: Manaus, Reserva Florestal Ducke , 28 May 1997, Assunção 506 ( INPA!, K!, NY!, RB!); Rio Camanaú , Mun. de Novo Airão , 24 July 1989, Miller 173 ( INPA!, K!, NY!) . Acre: Cruzeiro do Sul, Estrada Alemanha , 15 April 1971, Prance et al. 11943 ( FHO!, INPA!, M!, NY!, RB!) .

Notes: — Gaulettia elata is a canopy terra firme forest tree and also occurs infrequently in sandy campinarana vegetation. It has less distinct reticulate venation than all other species of Gaulettia . The leaves and flowers also lack the dense pilosity of all other species,which may be due to differences in adaptation to forest versus open habitats.

2f. Gaulettia foveolata (Prance) Sothers & Prance , comb. nov.

Basionym: Couepia foveolata Prance (1972a: 221) . Type:— VENEZUELA. Bolívar: Río Caroní , below Urimán , 11 January 1955, Steyermark & Wurdack 62 (holotype NY!, isotypes BM!, F!, FHO!) .

Tree to 30 m tall, lamina oblong, 3.5–9.5 × 1.5–3.7 cm, prominently reticulate with hair-filled stomatal cavities beneath. Inflorescences of axillary racemes, few-flowered, densely ferrugineous-brown-pubescent. Bracts and bracteoles oblong, to 7 mm long, persistent. Receptacle subcylindrical, 8–10 mm long, hirtellous-pubescent on exterior. Stamens c. 20, unilateral, with tooth-like staminodes opposite. Fruit ovoid, 2.6 × 3.2 cm; epicarp verrucose and hard; endocarp granular, sparsely hirsute within.

Illustration:— Prance (1972a: 223).

Distribution and habitat: — Venezuela. Riverine forests of the Guayana Highland, 100– 600 m. This species occurs in the same region as G. steyermarkii but is ecologically distinct. Fig. 8 .

Conservation status: — Gaulettia foveolata occurs in a restricted habitat type in the Guayana Highland of Venezuela, and is thus here assessed as NT ( IUCN 2001 ) .

Selected specimens examined: — VENEZUELA. Bolívar: Salto Erepuchi, Alto Río Caroní , January 1949, Cardona 2555 ( NY!, VEN!); Mun. Raul Leoni, S of Urimán , March 1986, Fernández 2288 ( K!) ; Distr. Piar, Río Aparamán , rapids of Yuray-merú, SW base of Amaruay-tepui, 21 April 1986, Holst & Liesner 2669 ( MO!, NY!); Río Paragua between mouth and Quebrada del Trueno , 13 May–13 June 1987, Stergios 10186 ( MO!, NY!, PORT) .

Notes: —In Prance & Sothers (2003b) two specimens were erroneously cited as C. foveolata . Rosa & Cordeiro 1713 is now considered to be G. amaraliae , and the correct identification of Stevenson et al. 1011 is G. racemosa .

2g. Gaulettia parillo (DC.) Sothers & Prance , comb. nov.

Basionym: Couepia parillo Candolle (1825: 526) .— Moquilea parillo (DC.) Steudel (1841: 159) as ‘ parilla’. Type:— FRENCH GUIANA. Without date, Herb. Ventenat s.n. (holotype G!).

Couepia pauciflora Huber (1909: 372) . Type :— BRAZIL. Pará , Lago de Faro , Igarapé do Dedal, 4 September 1907, Ducke MG 8630 (holotype MG!, isotypes BM!, RB!).

Couepia villosa Fanshawe & Maguire (in Maguire et al. 1948: 378). Type :— SURINAME. Tafelberg, 17 September 1944, Maguire 24782 (holotype NY!, isotypes BR!, F!, K!, M!, MO!).

Tree to 20 m tall or shrubs. Leaves oblong or oblong-elliptic, 5.0–15.5 × 1.7–5.8 cm, prominently reticulate beneath, with silver-grey pubescence between the reticulations. Inflorescences racemose, 5–21 flowers, or rarely with solitary flowers, the axis with ferrugineous-brown pubescence. Receptacle tubular, 11–22 mm long, with dense hirtellous pubescence and some longer hairs on exterior, glabrous within or with hairs nearly to base on proximal side beneath ovary only; stamens 45–62, inserted almost in a complete circle, with a portion of toothed staminodes. Fruit globose, c. 2.5 cm diameter; epicarp yellow-brown-velutinous. Fig. 9 .

Illustration:— Prance (2002: 204).

Distribution and habitat: — Guyana, French Guiana, Suriname, Venezuela, Amazonian Brazil (Acre, Amazonas, Amapá, Roraima), Peru, Ecuador and Colombia. Savannas, primary and secondary forests. Fig. 10 .

Conservation status: —This species meets the criteria of least concern, LC ( IUCN 2001). It is widespread, occurring throughout the Amazon region, in a number of habitats including secondary forests. Gaulettia parillo (as C. parillo ) was also assessed as least concern, LC for Colombia ( Calderón et al. 2002).

Selected specimens examined: — GUYANA. Potaro-Siparuni Region: Iwokrama Reserve , 26 September 1996, Clarke 2598 ( K!, NY!, US!) . SURINAME. Sipaliwini: Blanche Marie Falls , Nickerie R., 25 March 1997, Evans et al. 2671 ( K!, MO!) . VENEZUELA. Amazonas: Dept. Río Negro, Río Mawarinuma, Neblina Massif , 09–14 July 1984, Davidse & Miller 27217 ( MO!, NY!, US!, VEN!) . COLOMBIA . Vaupés: near Mitú , 02 October 1991, Betancur et al. 2865 ( COL!, K!, NY!, US!) . ECUADOR. Napo: Aguarico Cantón, Res. Faunística Cuyabeno, Laguna Zancudo Cocha , 28 September 1991, Palacios 7751 ( K!, MO!) . PERU. Loreto: Prov. Loreto, Nauta, Río Marañón , 10 November 1982, Vásquez & Jaramillo 3463 ( MO!, NY!) . BRAZIL. Acre: Mun. de Cruzeiro do Sul , 44 km E of Cruzeiro do Sul, 18 October 1984, Cid Ferreira et al. 5336 ( INPA!, K!, NY!, RB!) . Amazonas: Mun. de Barcelos , 3 km S of Central Serra of Serra Aracá , 04 March 1984, Amaral 1698 ( INPA!, NY!) . Amapá: Mun. de Oiapoque , 109 km SSE of Oiapoque road to Calçoene , 05 December 1984, Mori et al. 17186 ( K!, MG!, NY!) . Roraima: Rio Catrimani , Igarapé Camogi , 20 April 1974, Pires et al. 14059 ( IAN!, NY!) .

Notes: — Gaulettia parillo and G. canomensis have similar leaves and are sympatric, widely distributed species. They are more easily distinguished by the size and pilosity of their flowers and inflorescences. The two are also the only species of Gaulettia with villous mature fruit.

2h. Gaulettia racemosa (Benth. ex Hook.f) Sothers & Prance, comb. nov.

Basionym: Couepia racemosa Benth. ex Hooker (1867: 43) . Lectotype (selected by Prance 1972a):— BRAZIL. Amazonas: Falls of São Gabriel, March 1951, Spruce 1776 (K!, isolectotypes OXF!, P!).

Couepia racemosa var. reticulata Pilger (1906: 150) . Lectotype (designated here):— BRAZIL. Amazonas : Manaus, 17 May 1902, Ule 6149 (K!; isolectotypes G!, HBG, MG!; holotype B destroyed).

Couepia calophlebia Standley (1937: 248) . Type:— BRAZIL. Amazonas: Rio Embira , 19 June 1933, Krukoff 4913 (holotype F!, isotypes A!, BM!, K!, M!, MO!, NY!, RB!, U!, US!) .

Tree to 30 m tall. Leaves elliptic to oblong, 5.0–18.5 × 2.0– 7.5 cm, prominently reticulate beneath, or with stomatal cavities, with appressed grey pubescence. Inflorescences little branched, densely flowered panicles, the axis and branches with brown tomentellous pubescence. Receptacle cylindrical-turbinate, 5.0– 7.5 mm long, with brown pubescence on exterior. Stamens 24–30, unilaterally placed, with a smooth, glabrous, untoothed portion of the circle opposite them. Fruit elliptic, pointed at apex, 3.0–5.0 × 2.0– 3.5 cm; epicarp verrucose and pericarp undifferentiated, fragile, sparsely pubescent within.

Distribution and habitat: — Amazon region of Venezuela, Colombia, Peru and Brazil (Amazonas, Pará). Occurs in non-flooded forest or on periodically flooded ground, riverine forests, riverbanks and secondary forests up to 100 m. Fig. 11 .

Conservation status: — The conservation status of this species is here evaluated as least concern, LC ( IUCN 2001). As with G. parillo and G. canomensis , this is a widespread species occurring throughout the Amazon region , in a number of habitat types, including secondary forests. This species was also assessed as LC for Colombia ( Calderón et al. 2002) .

Selected specimens examined: — VENEZUELA. Amazonas: Dept. Atures, Río Cataniapo, 2 km from San Pedro de Cataniapo , 11 November 1980, Guanchez 396 ( MYF) . COLOMBIA. Amazonas: Resguardo Aduche, Correg. de Pto. Santander, 01 December 1993, Cárdenas & Andoque 4282 ( COAH, K!) . PERU. Loreto: Maynas, Llachapa, Río Napo , 19 January 1983, Vásquez & Jaramillo 3762 ( MO!, NY!) . BRAZIL. Amazonas: Mun. de Tefé, Lago Tefé , Vila Nogueira , 13 October 1982, Amaral et al. 56 ( INPA!, NY!, US!) ; Manaus, Estrada Mauá , 26 March 1971, Prance et al. 11631 ( FHO!, INPA!, K!, NY!, RB!) ; Rio Negro at confluence with Rio Uaupés , Serra Camaleão , 22 November 1987, Stevenson et al. 1011 ( INPA!, K!, NY!) . Pará: 25 km up Rio Mapuera , 07 June 1974, Campbell et al. P 22563 ( INPA!, NY!, US!) .

2i. Gaulettia steyermarkii (Maguire) Sothers & Prance , comb. nov.

Basionym: Couepia steyermarkii Maguire (1952: 252) . Type:— VENEZUELA. Bolívar: SE slopes of Ptari-tepui , 10–11 November 1944, Steyermark 60025 (holotype F!, isotypes K image!, NY!, VEN!) .

Tree or shrub with elliptic to oblong-ovate leaves, 4.0–6.0 × 2.7–4.0 cm, prominently reticulate beneath, grey-lanate between the reticulations. Inflorescences few-flowered axillary racemes. Receptacle cylindrical, 8–10 mm long, with hirtellous pubescence on the exterior. Stamens inserted around 2/3 of the circle; number of stamens unknown. Young fruit to 21 × 10 mm, epicarp appressed velutinous (lanate), with long yellow hirtellous hairs, apex pointed.

Distribution and habitat: — Venezuela (highlands of Bolívar state). Forests from 1000–1600 m and on tepuis (rarely at 500 m); one specimen from secondary forest (1112 m). Fig. 12 .

Conservation status: — Gaulettia steyermarkii has a restricted distribution and is known from only a few collections. It is here assessed as endangered, EN D, according to the IUCN criteria ( IUCN 2001).

Specimens examined: — VENEZUELA. Bolívar: Sifontes Dept., Mun. de Urdaneta , 30 January 1985, Hernández & Dezzeo 112 ( MYF, NY!); Distr. Roscio, 15 km ESE of San Ignacio de Yuruaní, 01 March 1984, Huber et al. 9152 ( MYF) .

Notes: —When Maguire(1952) described this species, he noted the similarity of the pubescence of the inflorescences with C. pauciflora , a synomym of G. parillo , and that it was “suggestive” of G. canomensis . He referred to the venation of G. steyermarkii as “reticulate veinwork”. This species has a deeply reticulate venation and distinct stomatal cavities on the abaxial surface of the leaves, and the fruits are tomentose on the exterior; however, only immature fruits have been collected.

Figures

FIGURE 4. Distribution of G. canomensis.

FIGURE 5. Distribution of G. cognata.

FIGURE 6. Gaulettia elata (Ducke) Sothers & Prance.A. Flowering habit × 1(2/3); B. Underside of leaf × 1.5(1); C. Detail of underside of leaf venation × 9(6); D. Bud × 4½(3); E. Flower × 6(4); F. Flower L.S. × 6(4); G. Stamen, two views × 40(30); H. Fruit × 1(2/3); I. Detail of fruit surface × 9(6); J. Fruit of Gaulettia canomensis × 1(2/3).A–G: Freitas et al. 760; H, I: Souza et al. 449; J: Nascimento & Silva 710. Scale bar: A, H, J = 3 cm; B = 2cm; C, I = 3 mm; D = 7 mm; E, F = 5 mm; G = 0.8 mm (drawn by Lucy T. Smith).

FIGURE 7. Distribution of G. elata.

FIGURE 8. Distribution of G. foveolata.

FIGURE 9. Fruit of Gaulettia parillo (DC.) Sothers & Prance (Souza et al. 211), from Brazil. © Projeto Flora da Reserva Ducke.

FIGURE 10. Distribution of G. parillo.

FIGURE 11. Distribution of G. racemosa.

FIGURE 12. Distribution of G. steyermarkii.