Virola montana D.Santam., 2019

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

10. Virola montana D.Santam. sp. nov. Figs 21 , 22 A–D

Diagnosis.

Species most similar to Virola guatemalensis , from which it differs by the mostly caducous (vs. persistent) trichomes on the abaxial leaf surface that have 3-10 branches and are 0.2-0.6 mm long (vs. 3-6 branches 0.05-0.1 mm long), staminate flowers with a shorter filament column (0.6-0.9 mm vs. 1-1.2 mm long) and fruits with a thicker pericarp (3.2-5 mm vs. 0.4-1 [-2.5] mm thick).

Type.

Costa Rica. Cartago: Jiménez, Taus de Pejibaye, 900 m elev., 06 Apr 1994 (♂ fl), E. Lépiz & J. F. Morales 284 (holotype: CR! [201663]; isotypes: CR! [CR1578073], LSU! [0193696, LSU00199100], MO! [5551138, MO280083], USJ! [75968]).

Description.

Tree 6-35 m × (4-) 10-50 cm DBH; bark not described. Exudate described on one occasion as orange in bark, branches and fruits. Twigs 0.13-0.4 cm thick, angulate to lightly compressed, densely tomentose to puberulent, trichomes dendritic, yellowish, brownish to ferruginous. Leaves: petiole 0.5-1.2 × 0.16-0.3 cm, canaliculate, densely to sparsely pubescent, the trichomes dendritic to irregularly stellate; leaf blades (11.2-) 15-30.5 × (3.9-) 4.5-7.4 cm, oblong-elliptic; adaxial surface on mature leaf blades dark brown when dry, reddish-brown or greyish-blackish, glabrous or with trichomes very sparse and scattered, the surface smooth; abaxial surface pale brown, dark brown or whitish-greyish when dry, sparsely pubescent, but usually densely to sparsely pubescent along the lateral veins and midvein (in new leaves, blades with a dense layer of trichomes that fall readily when touched, covering the entire surface), trichomes dendritic or rarely irregularly stellate, sessile, yellowish to pale brown, with 3-10 branches 0.2-0.6 mm long, caducous; lateral veins (15-) 18-30 per side, (4-) 6-9 veins per 5 cm, 0.5-0.9 (-1.5) cm apart, the same colour as the adaxial surface or sometimes lighter, flat or very slightly sunken on adaxial surface, raised on abaxial surface, straight to slightly arcuate (especially towards the distal part), slightly anastomosing near the margin and without forming a very marked intramarginal vein; tertiary veins usually visible on both surfaces; midvein adaxially flat to slightly canaliculate, glabrous or with scattered trichomes, sometimes densely pubescent at the base, abaxially raised, rounded, densely tomentose (with trichomes that fall very easily to the touch) to glabrescent; base acute to rounded, not revolute, flat; margin flat; apex acute to acuminate. Staminate inflorescences 3-9 cm long, axillary either at the junction with a leaf or at leafless nodes, axes slightly flattened to irregularly angled, densely tomentose, with trichomes dendritic, brown to yellowish-brown; peduncle 1.7-3.5 × 0.05-0.12 cm; bracts 3-7 × 1.9-4 cm, pubescent on both sides, caducous; terminal fascicles dense, with 7-20 + flowers. Staminate flowers with the pedicel 1.7-3.4 mm long; receptacle 1.5-3 mm wide;; perianth (1.6-) 2-2.7 mm, subglobose, yellow, greenish-white or brown, possibly by the indumentum), connate by (0.2-) 0.5-0.8 mm long, abaxial surface pubescent, with brown to ferruginous trichomes, adaxial surface glabrous at the base, sparsely pubescent on the lobes; lobes 3, 1.5-2 × 1-1.4 (-1.8) mm; stamens 3 (-6), the filament column 0.6-0.9 mm long, straight or sometimes slightly thickened at the base and somewhat narrow at the apex, thin, not constricted at the apex; anthers 0.5-0.8 mm long; apiculus ca. 0.07-0.1 mm, inconspicuously apiculate. Pistillate inflorescences 3.5 cm long, at leafless nodes, with trichomes on the axes similar to those of the staminate inflorescences; peduncle 1.8-2.5 × 1-2 cm; bracts not seen; terminal fascicles of 5-6 flowers. Pistillate flowers with the pedicel 3.5-4 mm long; perianth 3.5-4.6 mm long, globose to subglobose, pale brown when fresh (possibly by the indument), connate by 1-1.5 mm long, abaxial surface pubescent, with brown trichomes, adaxial surface sparsely pubescent; lobes 3, 2.5-3.5 × 1.5-2.2 mm; gynoecium 2-2.7 × 1.5-1.6 mm, densely pubescent, subglobose, stipitate; stigmatic lobes ca. 0.3 mm, erect. Infructescence 3-6.2 cm long, with 1 (-2) fruits, peduncle 2-3.3 × 0.19-0.3 cm. Fruits (2.8-) 3-3.6 × 2-2.5 cm, ovoid-ellipsoid, sessile, densely tomentose to tomentulose, the trichomes dendritic to irregularly stellate, pale brown to ferruginous, the surface commonly rugulose or smooth when dry, the line of dehiscence carinate, the base obtuse, the apex acute, orange, golden or yellowish-brown when fresh; pericarp 3.2-5 mm thick; pedicel 0.7-1 cm long; seed 2.2-2.5 × ca. 1.5 cm, the testa dark brown, almost smooth; aril usually described as red when fresh, yellowish- or reddish-brown when dry, coriaceous, oily in texture, thick, laciniate in narrow bands distally.

Distinctive characters.

Virola montana is recognised by twigs, new leaf undersurfaces, petioles and inflorescences covered in indument of dendritic to irregularly stellate, caducous trichomes, with long branches; on the underside of the leaf, this indument is mainly found on the midvein and the lateral veins ( Fig. 3J ). Additional traits that distinguish this species include leaf blades with numerous lateral veins [(15-) 18-30 per side; Fig. 22C, D ]), the length of the filament column (0.6-0.9 mm long), nearly the same size as the anthers (0.5-0.8 mm long), fruits with thick pericarp that are densely tomentose ( Fig. 4H ) and carinate in the line of dehiscence (3.2-5 mm), as well as its montane habitat.

Etymology.

The specific epithet refers to the montane habitat where the species has been collected.

Distribution.

Virola montana is known from Costa Rica and Panama, where it has been collected on the Caribbean slope in the provinces of Alajuela, Cartago, Guanacaste, Heredia and Limón in Costa Rica and Bocas del Toro in Panama; it has only been collected on the Pacific slope in Puntarenas (Costa Rica) ( Fig. 18D ). It has been recorded between 700-2000 m elevation.

Preliminary conservation status.

Virola montana is of Least Concern following IUCN guidelines. It has both a large EOO (14,606 km2) and AOO (104 km2) and is known from nineteen localities.

Common names.

Costa Rica: fruta dorada.

Phenology.

Flowering of Virola montana has been recorded in January, March to May, November and December; only two herbarium specimens with pistillate flowers were seen. Fruits have been collected in March and June to December.

Field characters.

Leaf blades are lustrous, dark green above and whitish or silver below. Flowers have yellow, cream, greenish-white or brown perianth. Mature fruits are yellow, pale brown, brown yellow or orange. The seed is brown with a red aril.

Discussion.

As far as we know, the first specimen of this species was collected 115 years ago by Henri F. Pittier (1857-1950) in the mountains of El Rosario de Orosi, Cartago, Costa Rica (H. Pittier 16628, NY-2 sheets!*). Paul C. Standley treated this specimen as V. koschnyi in "Flora of Costa Rica" ( Standley 1937). Shortly after, Albert C. Smith and Roger P. Wodehouse (1938) included the Pittier specimen under V. guatemalensis and this remains the name that is most frequently misattributed to specimens of V. montana (e.g. Standley and Steyermark 1946; Jiménez 2007). However, V. guatemalensis , as interpreted here, is restricted to northern Mesoamerica (Mexico, Guatemala and Honduras). In addition to the differences given in the diagnosis, the V. montana can be distinguished morphologically from V. guatemalensis by the differences summarised in Table 6 . Although it is difficult to quantify, specimens of V. montana commonly have twigs, petioles and inflorescences covered with more trichomes with long branches (see Fig. 3F, J ).

In addition to Virola guatemalensis , herbarium specimens of this new species have been determined as three other species: V. koschnyi , V. sebifera and V. surinamensis (here treated as V. nobilis ). Vegetatively, V. montana can be distinguished from these species by its mature leaves that are abaxially sparsely pubescent (vs. covering the entire surface of the leaf blades abaxially). The first two species are distinguished by pediculate trichomes on abaxial surface of leaf blades (vs. sessile in V. montana , these primarily present in young leaves). Finally, V. nobilis has trichomes with short branches [0.05-0.1 mm ( Fig. 3L ) vs. 0.2-0.6 mm long ( Fig. 3J )], tends to have more lateral veins [25-34 vs. (15-) 18-30 per side] and has a preference for lower elevation habitats [0-500 (-1300) m vs. 700-2000 m elevation].

Based on the number of herbarium specimens collected, Virola montana is the most common montane species of Virola in southern Mesoamerica and it is usually the only species where it occurs. However, three fruiting specimens of Virola sp., represented by R. Aguilar et al. 4327 (CR!, MO!), G. McPherson 8723 (MO!) and G. McPherson 10514 (MO!), have been collected in the general vicinity of V. montana in its preferred elevational range (1050-1500 m). These specimens clearly differ from V. montana in their small leaf blades [10.5-12.5 × 2.3-3.5 cm vs. (11.2-) 15-30.5 × (3.9-) 4.5-7.4 cm] with trichomes on the abaxial surface that have short branches (0.08-0.1 mm vs. 0.2-0.6 mm long) and cover the entire surface (vs. sparsely pubescent, but usually densely to sparsely pubescent along the lateral veins and midvein in V. montana ). These specimens share characteristics with two lowland species of Virola : V. nobilis and V. fosteri , including sessile, stellate trichomes on the abaxial surface of leaves. Additionally, these three specimens share their small leaves with V. fosteri , though not V. nobilis (9-17.2 [-27.5] × 2.5-5 [ –4.7– 7.1] cm vs. 10.5-12.5 × 2.3-3.5 cm). Further, the fruits of V. fosteri are smaller (1.5-2.3 × 1.2-1.8 cm vs. 2.8-3.2 × 2.2-2.7 cm). Additional collection and study is necessary to determine the identity of these specimens.

Specimens examined.

Costa Rica. Alajuela: Reserva Biológica Monteverde, Río Peñas Blancas, parcela de Badilla, 800 m elev., 23 Oct 1988 (fr), E. Bello 470 (CR-2 sheets!, INPA!*, LSU!, MO!); Reserva Biológica Monteverde, Parcela de Jesús Rojas, 850-900 m elev., 05 May 1989 (♂ fl), E. Bello 855 (CR-2 sheets!, INPA!*, MEXU!*, MO!); Reserva Biológica Monteverde, Ref. Aleman, 900 m elev., 10 Nov 1993 (♂ fl), E. Bello 5404 (CR!, LSU!, MO!); Reserva Forestal de San Ramón, 850-1150 m elev., 12-14 Mar 1987 (♂ fl), W. Burger et al. 12152 (CR!, F!*); San Ramón, Reserva Biológica Alberto Brenes, Estación San Lorencito, 900 m elev., 03 May 2000 (♂ fl), K. Caballero 2 (CR-2 sheets!, MO!); Reserva Forestal de San Ramón, 900-1200 m elev., 12-15 Mar 1987 (♂ fl), J. Gómez-Laurito 11446 (CR!, USJ!); Reserva Forestal de San Ramón, 900-1200 m elev., 16-19 Apr 1987 (♂ fl), J. Gómez-Laurito 11484 (CR!, F-2 sheets!*); Reserva Forestal de San Ramón, Estación Río San Lorencito, 850-900 m elev., 18 Oct 1989 (fr), J. Gómez-Laurito et al. 11846 (CR!, MEXU!*, MO!, USJ!); La Balsa, 1100 m elev., 02 Jul 1990 (st), J. Gómez-Laurito & J. A. López 11970 (USJ!); Reserva Biológica Monteverde, Río Peñas Blancas, 850 m elev., 31 Mar 1987 (♂ fl), W. Haber & E. Bello 6978 (CR!, F!*, INPA!*, MEXU!*, MO!); Reserva Biológica Monteverde, finca Wilson Salazar, 800 m elev., 07 Nov 1987 (fr), W. Haber & E. Cruz 7698 (CR!, INPA!*, MO!, USJ!); Río San Lorenzito, 800-1000 m elev., 30 Mar 1987 (♂ fl), G. Herrera et al. 514 (CR!, INPA!*, LSU!, MO!); La Fortuna, Finca El Jilguero, 1160 m elev., 20 Nov 1992 (fr), G. Herrera 5544 (CR!); 2 km N.E of La Balsa de San Ramón, 900 m elev., 26 Sep 1976 (fr), R. Lent 3899 (CR-3 sheets!, INPA!*, MARY!*, MEXU!*, MO!, PMA!*, U!*); Reserva Forestal San Ramón, Estación Río San Lorenzo, sendero al Volcán Muerto, 1100 m elev., 26 Apr 1993 (♀ fl), F. Quesada 20 (CR!, LSU!, MO!); Lago Coter [Cote], Hotel Eco-Lodge, 700 m elev., 09 Apr 1997 (♂ fl), J. Rivera & B. Petruzzi 2933 (CR!). Cartago: Taus, faldas del Cerro Alto El Humo, 900-1200 m elev., 06 Apr 1994 (♂ fl), J. F. Morales & E. Lépiz 2654 (CR!, MO!); El Rosario de Orosi, no date, Jan 1903 (fr), 1120 m elev., H. Pittier 16628 (NY-2 sheets!*); Turrialba, Monumento Nacional Guayabo, Santa Teresita, sobre los ríos Guayabo, Lajas y Torito, 700-1800 m elev., 08 May 1992 (♂ fl), G. Rivera 1659 (CR!, MO!, USJ!); límite Sur del Monumento Nacional Guayabo, sector Las Ventanas, 1100 m elev., 07 Jul 1992 (fr), G. Rivera 1910 (CR!, F!*, MO!, USJ!); [ Paraíso] 5 km N.W of Río Grande de Orosi at Tapanti, 1300 m elev., 27 May 1976 (♂ fl), J. Utley & K. Utley 5054 (CR-2 sheets!, MEXU-2 sheets!*, MO-2 sheets!). Guanacaste: San Gerardo, 1000 m elev., 21 Nov 1998 (fr), E. Bello 553 (CR!, INPA!*, MO!). Heredia: Sarapiqui, Parque Nacional Braulio Carrillo, puesto El Ceibo, 750 m elev., 05 Mar 1994 (st), B. Boyle et al. 2938 (MO!); Cariblanco, 800 m elev., n.d. Mar 1950 (fl bud), J. León 2368 (USJ!); Cariblanco, 800 m elev., 10 Aug 1950 (fr), LRH [L. R. Holdridge] 2618 (USJ!). Limón: Talamanca, Bratsi, Alto Lari, Kivut, 1300-1500 m elev., 15 Mar 1992 (fr), R. Aguilar & H. Schmidt 1131 (CR!, MO!). Puntarenas: Coto Brus, Zona Protectora Las Tablas, sitios Las Juntas, 1500 m elev., 03 Jul 1999 (fr), E. Alfaro 2357 (CR-2 sheets!, MO!); [Coto Brus], Las Tablas, río Cotoncito, [1300-1500 m elev.], 10 Dec 1983 (fr), I. Chacón et al. 1819 (INPA!*, LSU-2 sheets!, MO!); Parque Internacional La Amistad, Estación Pittier, 1650 m elev., 28 Jan 1995 (♀ fl), M. Chinchilla & IV curso de paratoxónomos 3 (CR-2 sheets!, LSU!, MO!); Zona Protectora Las Tablas, 2000 m elev., 01 Sep 1992 (fr), A. Fernández 354 (CR!); Fila Tigre, SE of Las Alturas, 1350-1450 m elev., 29 Aug 1983 (fr), G. Davidse 24178 (CR!, INPA!*, MEXU!*, MO!); Parque Internacional La Amistad, Santa María de Pittier, 1700 m elev., 13 Jun 1995 (fr), J. González 823 (CR-2 sheets!); Coto Brus, Zona Protectora Las Tablas, Hacienda La Amistad, 1600-2000 m elev., 28 Dec 2003 (♂ fl), R. Kriebel et al. 4174 (CR-2!, MO!). Panama. Bocas del Toro: Bocas del Toro-Chiriquí border above Fortuna Dam, 1200 m elev., 04 Dec 1985 (imm fr), G. McPherson 7756 (INPA!*, MO!, 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 10. Virola calophylla A branch with leaves, showing both sides B branch with inflorescences. Virola elongata C leafy branch with inflorescences, inset fruit D inflorescences. Virola guatemalensis E, F leaf blades showing adaxial (E) and abaxial (F) surface. G young twigs and petioles. Photos by Robin Foster (A, B), from https: // plantidtools. fieldmuseum. org / en / nlp; Steven Paton (C, D), Rolando Perez (C inset), from https: // stricollections. org / portal / index. php and Angela Rojas (E-G).

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 21. Virola montana A branch with leaves and inflorescences B fruit C flowers. Drawn by Pedro Juarez based on G. Herrera et al. 514 (A-C), and R. Lent 3899 (B).

Figure 22. Virola montana A juvenile tree showing branching pattern B leaf blades on adaxial side C leaf blades on abaxial side D venation. Virola multiflora. E Branch with staminate inflorescences, inset showing twig and flowers F twig, leaf blade on abaxial surface and inflorescences, inset showing immature fruit G lateral view of a perianth H close-up of staminate flowers. Photos by J. Esteban Jimenez (A-D); B. Hammel (E-H), Indiana Coronado (F, inset).