Virola sebifera Aubl.

Santamaria-Aguilar, Daniel, Aguilar, Reinaldo & Lagomarsino, Laura P., 2019, A taxonomic synopsis of Virola (Myristicaceae) in Mesoamerica, including six new species, PhytoKeys 134, pp. 1-82 : 1

14. Virola sebifera Aubl. Fig. 28

Virola sebifera Aubl. Hist. Pl. Guiane. 2: 904. 1775. Type. French Guiana. Cayenne, 1775, [J.] Aublet s.n., (holotype: BM!*; isotypes: NY photograph of BM holotype). Fide Jaramillo et al. 2004.

Myristica panamensis Hemsl. Biol. Cent.-Amer., Bot. 3: 67. 1882. Virola panamensis (Hemsl.) Warb., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 185. 1897. Syntypes: Panama. shady forest near Cruces, n.d. (fls.), Seemann 545 (BM!*, K!*); [Canal Area], Lion Hill station, [28 Mar 1862, leaves, excl. fr], S. Hayes 618 (K!*).

Virola warburgii Pittier. Contr. U.S. Natl. Herb. 18: 143. 1916. Type. Panama. Panamá, collected in forests along the Chagres River above Alhajuela, 12 May 1911 [fls], H. Pittier 3505 (holotype: US; isotype: BM).

Distinctive characters.

Virola sebifera is recognised by its dense pubescence of pediculate and ferruginous trichomes ( Fig. 3N ) that cover almost all parts of the plant; leaf blades with few lateral veins (9-17 per side), well-spaced between them [3-4 (-5) veins per 5 cm, 1.3-2.7 cm apart] ( Fig. 8N ); the long staminate inflorescences that are very branched; the staminate flowers with the filament column short (0.1-0.4 mm long) and the anthers more than twice the length of the column of the filaments (0.6-0.8 mm long) and apiculate at the apex; its small fruits [1.1-1.7 (-2.3) × 0.8-1.3 (-1.6) cm] that are globose to subglobose ( Fig. 4K ), covered by a dense layer of trichomes that fall easily and a thin pericarp [0.3-0.6 (-0.9) mm]; and the thin aril that is laciniate in narrow bands. There are also usually many fruits per infructescence.

Distribution.

Virola sebifera is the most collected and widespread species in Mesoamerica. It is known from Honduras (Gracias a Dios), Nicaragua ( Atlántico Norte, Atlántico Sur, Jinotega, Matagalpa and Río San Juan), Costa Rica (Alajuela, Cartago, Guanacaste, Heredia, Limón, Puntarenas and San José) and Panamá (Bocas del Toro, Chiriquí, Coclé, Colón, Darién, Herrera, Panamá, San Blas and Veraguas) ( Fig. 27B ). It has been recorded between 0-1000 m elevation.

Common names.

Honduras: walus, banak. Nicaragua: banak, cebo, cebo macho, cebo sirsio, sangre drago. Costa Rica: coton, fruta dorada, miguelario. Panama: bogamani, copidijo, fruta dorada, gorgoran, malagueta de montaña, mancha, sangre, tabegua, velario colorado.

Phenology.

Herbarium specimens of flowering and fruiting individuals have been collected throughout the year. In the Osa Peninsula and Golfo Dulce, Costa Rica, flowering occurs in December to May.

Field characters.

Plants are shrubs or trees between 8-25 (-40) m tall and 7-40 cm DBH. Trunks are straight, sometimes with small buttresses and with long and horizontal branches. Bark is described as dark, minutely fissured vertically or flaking off in small pieces and aromatic. Exudate from different parts of the plant is red or reddish. The leaf blades are bright green above and whitish to greyish below in living material. Flowers have brown, cream, orange, yellow-brown or yellowish perianth and are sometimes fragrant, with a lemon or sweet aroma. The mature fruit is green and covered by a dense layer of trichomes, with a white seed with a red aril.

Notes.

The leaves of this species are very variable in shape and size. Sometimes, a single individual may have leaves of different sizes. Individuals with small leaf blades from Panama (e.g. G. McPherson 15341, MO; G. McPherson 20671, MO), that are sparsely pubescent and with sessile trichomes on the abaxial surface of the leaf, may be difficult to distinguish from V. elongata . However, specimens of V. sebifera tend to have thicker twigs, petioles and floriferous and fruiting peduncles.

Selected specimens seen.

Honduras. Gracias a Dios: Monte Alto de Anastasio, 90 m elev., 17 Mar 1994 (♂ fl), P. R. House 2019 (MO!); al sur de Krausirpe, 25 Mar 1995 (♀ fl), P. R. House 2351 (MO!); 30 minutos al sur de Krausirpe, 11 May 1994 (fr), P. R. House 2814 (MO!). Nicaragua. Atlántico Norte: 13 km above Kururia on road to San Jerónimo, 200 m elev., 02 Mar 1979 (♂ fl), J. J. Pipoly 3844 (MO!); Siuna, La Pimienta, [150 m elev.], 01 Jun 1984 (fr), F. Ortiz 1971 (MO!). Atlántico Sur: El Zapote, 40 km NE de Nueva Guinea, 130-150 m elev., 27 Feb 1984 (♂ fl), J. C. Sandino 4767 (MO!); Boca de Sábalo, 70 m elev., 24 Mar 1985 (♀ fl), P. P. Moreno 25639 (MO!). Jinotega: Wiwili, zona de amortiguamieno Bosawas, 170 m elev., 18 Jan 2006 (♀ fl), I. Coronado et al. 2949 (MO!); Wiwili, Reserva de Bosawas, 220 m elev., 09 Feb 2006 (♂ fl), I. Coronado et al. 3388 (MO!); Wiwili, Reserva de Bosawas, 170 m elev., 09 Jun 2007 (fr), I. Coronado et al. 3969 (MO!). Río San Juan: 2 km al NW del Río Santa Cruz, 70 m elev., 24 Feb 1984 (♀ fl), P. P. Moreno 23328 (MO!); El Castillo, Finca de Madrid, 100 m elev., 18 Jan 2005 (♂ fl), R. Loredo 2348 (MO!); entre el pueblo de San Juan del Norte Nuevo y la casa de Ramón Castillo, 0-100 m elev., 07 Jul 1994 (fr), R. Rueda et al. 1851 (MO!). Costa Rica. Alajuela: Reserva Biológica de Monteverde, río Caño Negro, 800 m elev., 19 Oct 1989 (fr), E. Bello 1402 (MO!); Upala, Estación San Ramón, 550 m elev., 16 Mar 1993 (♂ fl), R. Espinoza 803 (MO!); San Carlos, Fortuna, 255-400 m elev., 27 May 2004 (fr), A. Rodríguez 9046 (MO!). Cartago: Turrialba, El Zapote Nature Reserve, 700 m elev., 01 Mar 1987 (imm fr), W. Burger & J. Gómez-Laurito 12030 (CR!, MEXU!*). Guanacaste: Finca La Pasmompa, 400 m elev., 10 Dec 1990 (fl bud), C. Moraga 227 (MO!). Heredia: Tirimbina, Istarú farm, 220 m elev., 28 Jan 1972 (♂ fl), R. W. Lent 2329 (MO); Finca La Selva, [90 m elev.], 09 Jul 1984 (fr), D. Smith 1088 (MO!). Limón: Parque Nacional Barbilla, 300-400 m elev., 22 May 2000 (fl bud), E. Mora 1155 (MO); Cerro Coronel, 20-170 m elev., 16-23 Jan 1986 (♂ fl), W. D. Stevens 23823 (MO!). Puntarenas: Golfito, Mata Palo, 160-200 m elev., 09 Sep 1991 (fr), R. Aguilar 363 (MO!); Golfito, 1 km antes de llegar a la Palma, 8 m elev., 16 Jan 1993 (♂ fl), R. Aguilar 1586 (MO!); Punta Burica, 0-100 m elev., 22 Aug 1988 (fr), M. M. Chavarría et al. 292 (MO-2 sheets!). San José: Puriscal, San Martín, 800 m elev., 20 Nov 1993 (fr), J. F. Morales 2022 (MO!); vicinity of El General, 915 m elev., n.d. May 1937 (♂ fl), A. F. Skutch 3118 (MO!); Turrubares, San Rafael, 600-700 m elev., 08 Dec 2004 (fr), A. Soto & D. Santamaría 409 (MO!). Panama. Bocas del Toro: Isla Bastimentos, Bocatorito, 30 m elev., 15 Feb 1989 (♀ fl), P. M. Peterson & C. R. Annble 6872 (MO!). Chiriquí: 17 km NE of San Felix, 1000 m elev., 18-19 Mar 1974 (fr), M. Nee 10664 (MO!). Coclé: Caribbean side of divide at El Copé, 200-400 m elev., 04 Feb 1983 (fr), C. Hamilton & G. Davidse 2756 (MO!). Colón: Area near Guasimo, no elev., 22 Apr 1970 (♂ fl), T. B. Croat 9950 (MO!); Santa Rita ridge, 500 m elev., 13 Jan 1987 (fr), G. McPherson 10265 (MO!). Darién: Vicinity of Cana, 1750 ft [530 m] elev., 24 Jun 1959 (♂ fl), W. L. Stern 512 (MO!); serranía de Sapo, 300-800 m elev., 26 Nov 1990 (fr), H. Herrera & J. Polanco 793 (MO!). Herrera: Road between Las Minas and El Toro, 700-800 m elev., 23 Jan 1987 (fr), G. McPherson 10273 (MO!). San Blas: El Llano Cartí road, 350 m elev., 06 May 1985 (fr), G. de Nevers 5609 (MO!). Panamá: Barro Colorado Island, [10-100 m elev.], 13 Dec 1967 (fr), T. B. Croat 4275 (MO!); along Pipeline Road, 125 m elev., 10 Mar 1974 (♂ fl), M. Nee 10445 (MO!); Llano Cartí road, 350 m elev., 26 Jan 1986 (fr), G. McPherson & M. Merello 8148 (MO!). Veraguas: Cerro Hoya, subiendo por Cobachón, [750 m elev.], 21 Apr 1997 (fl), J. Deago 499 (PMA!*).

Figures

Figure 3. Trichomes in Mesoamerican Virola A Virola allenii (R. Aguilar 2224, MO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (O. Valverde 970, MO) D V. elongata (G. McPherson 13674, MO) E V. fosteri (G. McPherson 20913, MO) F V. guatemalensis (R. Cedillo 3349, MO) G V. koschnyi (R. E. Gereau et al. 3455, MO) H V. laevigata (R. Aguilar 2004, MO) I V. megacarpa (J. A. Duke 15261, MO) J V. montana (E. Bello 855, MO) K V. multiflora (R. Rueda et al. 8196, MO; inset, J. C. Sandino 3302, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers & H. Herrera 7056, MO) N V. sebifera (F. Almeda et al. 5553, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO).

Figure 4. Fruits of Mesoameican Virola, organised in groups of morphologically similar species A V. allenii (P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9715, MO) C V. otobifolia (A. Gentry & S. Mori 14199, MO) D V. macrocarpa (A. E. Lawrance 675, MO) E V. chrysocarpa (B. Hammel et al. 16864, MO) F V. koschnyi (R. Robles 1587, MO) G V. guatemalensis (S. Sinaca 1542, MO) H V. montana (J. Utley & K. Utley 5054, MO) I V. fosteri (S. Mori & J. Kallunki 4891, MO) J V. multiflora (R. Rueda et a. l 9960, MO) K V. sebifera (T. B. Croat 5959, MO) L V. elongata (J. D. Dwyer & M. Correa 8420, MO) M V. laevigata (R. Aguilar 1585, MO) N V. nobilis from Barro Colorado (T. B. Croat 8090, MO) O V. nobilis from the Osa Peninsula (G. Herrera 4222, MO) P V. megacarpa (G. de Nevers 5184, MO). Scale bars: 1 cm.

Figure 8. Pattern of the lateral veins in Mesoamerican Virola A Virola allenii (K. Thomsen 1284, NO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (B. Hammel et al. 16864, MO) D V. elongata (M. Correa & R. L. Dressler 1078, MO) E V. fosteri (G. de Nevers 7226, MO) F V. guatemalensis (G. Ibarra 957, MO) G V. koschnyi (J. Miller & J. C. Sandino 1110, MO) H V. laevigata (N. Zamora & T. D. Pennington 1583, MO) I V. megacarpa (G. de Nevers 5184, MO) J V. montana (E. Lepiz & J. F. Morales 284, MO) K V. multiflora (J. Manzanares 3561, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers et al. 7530, MO) N V. sebifera (T. B. Croat 5959, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO).

Figure 27. Geographic distribution of Virola otobifolia (A) and V. sebifera (B).

Figure 28. Virola sebifera A leaf blade on adaxial surface B leaf blade on abaxial surface C, E leaf base on both sides D flowering branch G staminate flowers H infructescence I fruits and aril. Photos by Reinaldo Aguilar.