Virola nobilis A. C. Sm.

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

12. Virola nobilis A. C. Sm. Figs 20 C–G , 24B

Virola nobilis A. C. Sm. Brittonia 2: 490. 1938. Type. Panama. Canal Zone, Barro Colorado Island, 07 Jan. 1932 [imm fr], [R. H.] Wetmore, [E. C.] Abbe & [O. E.] Shattuck 155 (holotype: GH!*; isotypes: A!*, F!*, MO!).

Distinctive characters.

Virola nobilis is recognised by its narrow, oblong leaf blades (9-17.2 [-27.5] × 2.5-5 [ –4.7– 7.1] cm) with numerous lateral veins (20-30 [25-32] per side) corresponding to 8-11 (5-7 [-9]) veins per 5 cm, as well the stellate, tertiary veins that are slightly sunken above ( Fig. 3L ) and sessile trichomes scattered on underside of the leaf ( Fig. 3L ); the staminate flowers with the filament column longer (0.8-1.1 [1.2-1.3] mm long) than the anthers (0.6-0.8 mm long); and usually ellipsoid or ovoid fruits (2.3-2.7 × 2.2-2.4 cm [2.1-3.1 × 1.6-2.5 cm]) ( Fig. 4N ) with thick pericarp (2.3-3.5 mm [2.5-4.2 mm]) and aril.

Distribution.

Virola nobilis is known from the Pacific slope of Costa Rica (Puntarenas, San José) and the Caribbean slope of Panama (Bocas del Toro, Colón, Panamá, San Blas). It is recorded from 0-400 (-1300) m elevation ( Fig. 18F ). Jiménez (2007) reported this species (as V. surinamensis ) from just one locality ( Chitaría) on the Caribbean slope in Costa Rica; however, we have not examined the specimen (Poveda 144, CR).

Common names.

Costa Rica: Fruta dorada. Panama: bogamani, coton cuinur gia, sabdurgia (Kuna name).

Phenology.

Flowers have been collected in January to April, July, August, November and December and fruits in almost all months except October.

Field characters.

Plants are large trees between (5-) 15-40 m tall and 20-70 cm DBH. The trunk is straight, usually with conspicuous buttresses and does not begin to branch until it reaches a great height. The bark is reddish and releases red exudate when damaged. The leaves are whitish with inconspicuous trichomes on the abaxial surface. Flowers have yellow perianth. The mature fruit is yellow to orange with a red aril.

Discussion.

This species has usually been treated, identified and included as a synonym of Virola surinamensis (e.g. Gentry 1975; Croat 1978; Correa et al. 2004; Jiménez 2007), which has similar leaf morphology (i.e. number of lateral veins, stellate and sessile trichomes scattered on the abaxial surface of the leave blades). However, Virola surinamensis ( Figs 23 G–J , 24A ) is characterised by having a shorter perianth that is also somewhat fleshy to submembranous and smaller fruits that are also ovoid to subglobose, glabrescent and with a thin pericarp ( Figs 23 I–J , 24A inset; also see illustration in Rodrigues 1972, 1980). Although both species may have similarly-sized leaves, they tend to be smaller in V. surinamensis , which also has inflorescence axes that are longer and with many more flowers ( Figs 23F , 24A ).

Notes.

Virola nobilis exhibits complex variation that requires more fieldwork, ideally in combination with molecular phylogenetic analysis, to clarify species boundaries. For example, specimens from the Pacific slope in Costa Rica (see specimens cited below and Figs 24C , 25 ) are included, with reservation, under V. nobilis . These specimens differ from other collections of V. nobilis , including the type and additional specimens from Barro Colorado Island (the type locality), in having wider leaf blades, fewer lateral veins, tertiary veins that are less inconspicuous below ( Fig. 3L, O ), a longer filament column and ovoid fruits with obtuse to rounded apices ( Fig. 4N, O ). These specimens also resemble V. reidii Little, a species from Colombia and Ecuador ( Jaramillo et al. 2004). The measurements presented in square brackets [] correspond with the Costa Rican material under question.

Other material that potentially belongs to V. nobilis includes a few collections from Panama, including both infertile and fruting specimens from El Llano Cartí and others from Cerro Jefe (e.g. G. de Nevers & H. Herrera 4333, imm fr, MO!; R. L. Liesner 657, fr, MO!; E. L. Tyson et al. 3353, fr, MO!). These differ from V. nobilis by having the abaxial surface of the leaf blade covered by a dense and a very inconspicuous layer of stellate, sessile and yellowish trichomes and fruits with a more or less smooth surface and that are not carinate in the line of dehiscence. Finally, R. Aguilar 3408 (CR-3 sheets!, MO!) from the Osa Peninsula of Costa Rica and G. McPherson 11474 (MO!; fr) from Panama differ from the species concept that we adopt in having trichomes on the abaxial leaf side with long branches (ca. 0.2 mm long) that are often shortly pediculate; and small fruits (ca. 2.1 × 1.6-1.7) that are ovoid to subglobose and apiculate at the apex.

Selected specimens.

Costa Rica. Puntarenas: Golfito, camino a Piro, finca de Adrian, 50 m elev., 16 Apr 2005 (fr), R. Aguilar & X. Cornejo 9739 (MO!); Osa, Punta Pargos, 0 m elev., 17 Apr 2008 (fr), R. Aguilar 11186 (MO!, USJ!); Osa, Rancho Quemado, siguiendo el nuevo camino a Drake, 400 m elev., 20 Jun 1990 (fr), G. Herrera 4222 (MO!); Trocha de La Tarde rd. 10 km SW of La Palma, 150-200 m elev., 28 Apr 1988 (fr), B. Hammel & R. Robles 16722 (MO!); Osa, Fila Ganado, 350 m elev., 15 Dec 1990 (♂ fl), B. Hammel et al. 18017 (CR-2 sheets!, MO!); Osa, Fila Ganado hasta Guerra, 1-300 m elev., 28 Mar 1991 (fr), B. Hammel et al. 18170 (MO!). San José: Tarrazú, 900-1300 m elev., 19 Aug 1997 (fr), O. Valverde & A. Estrada 135 (CR!). Panama. Bocas del Toro: North of Fortuna Dam, on road to Chiriquí Grande, 500 m elev., 18 Jan 1986 (fr), G. McPherson 8098 (MO!). Colón: Carretera Gatún-Piñas, 0-50 m elev., 26 Jul 1994 (fr), C. Galdames et al. 1415 (CR!, MO!). Panamá: Barro Colorado Island, [10- 100 m elev.], 23 Jan 1969 (fl bud), T. B. Croat 7488 (MO!); ibid, 22 Feb 1969 (fr), T. B. Croat 8090 (MO!); El Llano Cartí, 1100-1200 m [335-365 m] elev., 28 Dec 1974 (fr), S. Mori & J. Kallunki 4151 (CR!). San Blas: El Llano-Cartí, 100-350 m elev., 03 Nov 1985 (imm fr), G. de Nevers et al. 6172 (MO!).

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 18. Geographic distribution of Virola koschnyi (A), V. laevigata (B), V. megacarpa (C), V. montana (D), V. multiflora (E) and V. nobilis (F).

Figure 20. A. Virola megacarpa A branch with fruits, both leaf blade surfaces shown B fruit. Virola nobilis from Barro Colorado C lower trunk and buttress D branch with fruits, leaf blades on both surfaces E pistillate flowers F staminate flower and bud G fruits. Photos by Carmen Galdames (A, B), Rolando Perez (C, D, G), Steven Paton (E, F); all photos from https: // stricollections. org / portal / index. php.

Figure 23. Virola otobifolia A branch with leaf blades showing both surfaces and inflorescences B leaf blade, venation and base on abaxial surface C infructescence D fruits E detail of fruit, showing an aril of an immature fruit. Virola surinamensis. F Branch with inflorescences G, H leaf blade on adaxial (G) and abaxial surface (H). I Infructescence J detail of fruits, including a longitudinal section of an immature fruit. Photos by Rolando Perez (A, B from https: // stricollections. org / portal / index. php); Jerry Harrison (C), Alwyn H. Gentry (D, E from http: // www. tropicos. org) and John P. Janovec (F-J from http: // atrium. andesamazon. org /).

Figure 24. Comparisons of Virola surinamensis with collections of V. nobilis A V. surinamensis (S. Mori & R. Cardoso 17467, NY; inset showing fruits from W. E. Broadway 5264 a, MO) B V. nobilis (T. B. Croat 7488, MO) C V. nobilis from Costa Rica (B. Hammel 18017, MO). A, image courtesy of New York Botanical Garden.

Figure 25. Virola cf. nobilis from the Osa Peninsula, Costa Rica A trunk B branch with fruits C leaf blades on adaxial surface D twig, petiole and leaf base on adaxial surface E exudate on a twig F, G leaf blades on abaxial surface and trichomes (G). H Leaf margin I leaf apex J staminate inflorescences K detail of staminate inflorescences L detail of staminate flower M branch with fruits N fruits O seed. Photos by Reinaldo Aguilar.