Virola laevigata Standl.

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

8. Virola laevigata Standl. Fig. 19

Virola laevigata Standl. Publ. Field Mus. Nat. Hist., Bot. Ser. 4(8): 209. 1929. Type. Panama. Province of Chiriquí, Progreso, [ July–Aug.] 1927 [♂ fl], G. P. Cooper & G. M. Slater 308 (holotype: F!*; isotypes: NY!*, WIS!*, US!*).

Distinctive characters.

Virola laevigata is distinguished by its glabrous or nearly glabrous vegetative parts (i.e. twigs, mature leaf blades on both surfaces [ Fig. 3H ], petioles); when trichomes are present, they are primarily on the leaf buds or very new leaves. Additionally, leaves have 12-20 lateral veins and margins and bases that are slightly revolute, staminate flowers have a straight filament column that is longer (0.8-1.3 mm) than the anthers (0.5-0.7 mm) and relatively small fruits (1.8-2.9 × 1.5-1.8 cm) ( Fig. 4M ) with pericarp that is 1.8-2.8 mm thick.

Distribution.

Virola laevigata is known from Costa Rica (Puntarenas and San José) and Panama ( Chiriquí) ( Fig. 18B ). It is found on the Pacific slope, where it has been recorded between 0-500 (1600?) m elevation. Jiménez (2007) suggested that the maximum elevation for this species in Costa Rica is 1600 m, potentially based on L. González 3089 from Cerro Turrubares (San José province) (B. Hammel pers. comm., Aug. 2019); however, we have not found any specimen that occurs this high.

Common names.

Costa Rica: fruta dorada. Panama: bogamani.

Phenology.

Flowering of Virola laevigata has been recorded in January, May, July and November. Fruits are produced from December to February. Pistillate flowers were not present on herbarium sheets studied.

Field characters.

Plants are trees 9-40 m tall and 35-60 cm DBH with a straight trunk and small (ca. 20 cm tall), triangular buttresses. The bark is described as finally grooved, smooth, flaking in vertical strips or scaly and is grey, blackish or reddish in colour, with exudate that is reddish or colourless and oxidising to reddish-cream. The leaves are bright green on both sides and have numerous pellucid dots that are most visible against the light. Flowers have yellow, yellow-brown or yellowish perianth, sometimes with a slight aroma in staminate flowers (N. Z amora & T. D. Pennington 1583, but the specimen label states pistillate flower). The mature fruit is yellow with a red aril (when immature, it is white). In the Osa Peninsula, where this species is frequent, it prefers riparian habitats.

Discussion.

Virola laevigata has traditionally been considered a synonym of V. guatemalensis (e.g. Smith and Wodehouse 1938; Standley and Steyermark 1946; Duke 1962), likely due to limited material. However, with new herbarium specimens, both species can be clearly distinguished vegetatively and with fruit characters. Vegetative material of V. laevigata can be distinguished by petioles and mature leaf blades that are abaxially glabrous (vs. diminutively pubescent with tiny stellate, sessile trichomes in V. guatemalensis ). Their fruits also differ in size, shape and thickness of the pericarp; in V. laevigata , they are smaller (1.8-2.9 × 1.5-1.8 cm), ovoid or subglobose and with thick pericarp (1.8-2.8 mm); while in V. guatemalensis , they are large (vs. 2.7-3.4 × 1.7-2.3 cm), ellipsoid and with a thin pericarp [0.4-1 (-2.5) mm].

In addition to Virola guatemalensis , herbarium specimens of V. laevigata have been determined as V. surinamensis (interpreted here as V. nobilis ). However, V. laevigata is distinguished by its glabrous or almost glabrous abaxial leaf surface ( Fig. 3H, O ) and mature fruits (vs. pubescent) and its tendency towards thinner pericarp [1.8-2.8 mm vs. 2.5-3.5 (-4.2) mm thick]. In the Osa Peninsula, where these two species grow together, they can be easily distinguished in the field: V. laevigata prefers riparian habitats, does not usually develop tall buttresses and the external bark has a greenish tone, while V. nobilis grows far from bodies of water, has tall buttresses and the external bark is reddish to brown. For a description and comparison of the bark of these two species, see Moya Roque et al. (2014), as Virola sp. A and V. surinamensis .

Notes.

The seedlings of V. laevigata are described by Ley López and Chacón Madrigal (2017) (though as " Virola sp. A"). Additionally, the species presented as " Virola sp. A" in Jiménez (2007) and as V. surinamensis in Jiménez Madrigal and Grayum (2002; RZ [= R. Zuñiga] 459) corresponds with V. laevigata . The illustration in Quesada Quesada et al. (1997) as V. guatemalensis is potentially also V. laevigata .

Selected specimens seen.

Costa Rica. Puntarenas: Golfito, 1 km antes de llegar a La Palma, 8 m elev., 16 Jan 1993 (fr), R. Aguilar 1585 (CR-2 sheets!, LSU!, MO!); fila Carbonera, cabo Matapalo, 300 m elev., 16 Jul 1993 (♂ fl), R. Aguilar 2004 (CR!, MO!); estación Sirena, 10 m elev., 12 Oct 1993 (fl bud, fr), R. Aguilar 2490 (CR-2 sheets!, LSU!, MO!); Golfito, orillas del camino a las torres de comunicación, 450-500 m elev., 12 Jan 1999 (fr), J. Gómez-Laurito & V. Mora 13191 (CR!); Playa Cacao, Fila entre quebrada Nazanero y el mar en Punta Voladora, 200 m elev., 26 May 1994 (♂ fl), G. Herrera & G. Rivera 7076 (CR!); Parque Nacional Corcovado, Sirena, 10 m elev., 28 Feb 1989 (fr), C. Kernan & P. Phillips 962 (CR-2 sheets!, MO!); Parque Nacional Corcovado, Sirena, Ollas Trail, 1-20 m elev., 19 Oct 1989 (fr), C. Kernan & G. Fonseca 1288 (CR-2 sheets!, INPA!*, MO!); Jiménez, Piro, camino a Laguna Silvestre, [0-100 m elev.], 01 Feb 2012 (fr), J. M. Ley-López 74 (USJ); Osa, fila Esquinas, 200 m elev., 26 May 1993 (♂ fl), M. Segura & F. Quesada 69 (CR!, LSU!, MO!); Rincón de Osa, in vicinity of airstrip, 40 m elev., 25 Jul 1974 (fl bud), J. Utley & K. Utley 1236 (CR!); Garabito, por Carara, cerca de la toma de agua, 97 m elev., 22 Nov 2006 (fl bud), L. D. Vargas & D. Castillo 1870 (CR!); Osa, Uvita, cerca de Dominical, 200 m elev., 28 Jan 1991 (♂ fl), N. Zamora & T. D. Pennington 1583 (CR-2 sheets!, INPA!*, MEXU!*, MO!); Puntarenas, Montezuma, camino a Cóbano por el río Montezuma, ca. 1.5 km oeste de la intersección con camino a Cabuya, 101 m elev., 11 Dec 2005 (fr), B. Hammel & I. Pérez 23944 (CR!); Montezuma, por el Canopy, 100 m elev., 08 Jul 2006 (fl bud), B. Hammel & I. Pérez 24149 (CR!). San José: Carara, sector Agrominas, sitio Carretera Costanera, 100 m elev., 20 Sep 1991 (fr), R. Zuñiga 459 (CR-sheets!, LSU!). Panama. Chiriquí: Río Platanal-Bugaba, 10 Dec 1975 (fr), M. M. Gutiérrez 21 (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 19. Virola laevigata A branch with staminate inflorescences B leaf blades on abaxial surface, inset showing the glabrous midvein C twig, petiole and leaf on adaxial surface D close up of staminate inflorescences E leaf base F leaf apex, inset showing leaf punctations G branch with immature fruits H close-up of immature fruit I open fruit, showing the aril. Photos by Reinaldo Aguilar.