Virola guatemalensis (Hemsl.) Warb.

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

6. Virola guatemalensis (Hemsl.) Warb. Fig. 10 E–G

Virola guatemalensis (Hemsl.) Warb. Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 68: 220. 1897.

Myristica guatemalensis Hemsl. Biol. Cent.-Amer., Bot. 3. 66-67. 1882. Type. Guatemala. [no specific data in location], [no date], [seeds], [G. U.] Skinner s.n. (holotype: K!*).

Distinctive characters.

Virola guatemalensis is distinguished by many characters of its leaf, including overall size [12.3-17.5 (-24.1) × (2.4-) 3.8-5.5 (-8.9) cm], inconspicuous pubescence of tiny stellate and sessile trichomes on abaxial surfaces ( Fig. 3F ), 13-21 lateral veins on each side of the leaf that are barely distinctive abaxially ( Figs 8F , 10F ) and slightly revolute margins. Additionally, the staminate flowers have a filament column that is longer (1-1.2 mm long) than the anthers (0.5-0.8 mm long) and fruits that are 2.7-3.4 × 1.7-2.3 cm, ellipsoid ( Fig. 4G ), commonly glabrous or glabrous distally and with pubescence at the base with the line of dehiscence slightly carinate or smooth and a thin pericarp [0.4-1 (-2.5) mm thick].

Distribution.

Virola guatemalensis is known from Mexico (Chiapas, Oaxaca and Veracruz), Guatemala ( Sololá) and Honduras (Yoro) ( Fig. 9F ). It has been recorded between 150-1250 m elevation. Standley and Steyermark (1946) mention that it also occurs in Alto Verapaz, Suchitepéquez, San Marcos and Huehuetenango in Guatemala, while Standley (1931) indicates that it occurs in Lancentilla (Honduras), where it is noted to be one of the most common tree species. While Gentry (2001) postulated that V. guatemalensis is an expected species for Nicaragua, no Nicaraguan collections of this species are known.

Common names.

Mexico: cacao, cacao volador (Chiapas), cacaotillo, cedrillo (Veracruz), k’ik’ che’ [Lacandon name]. Guatemala: chucul, palo de sebo, cacao volador, cacao cimarrón. Honduras: sangre.

Phenology.

Flowering of Virola guatemalensis has been recorded in April and May and fruit production in January, March, August, October and December.

Field characters.

Plants are trees between 12-35 m high and 45-130 cm DBH with a straight trunk, sometimes with moderately sized buttresses. The bark is variously described as smooth or fissured and scaly and is brown to greyish-brown in colour and exudes watery reddish transparent sap when damaged. Flowers have yellow, green-yellowish or brown perianth. The mature fruit is yellow with a red aril.

Discussion.

While the name Virola guatemalensis has been applied to herbarium specimens from Costa Rica and Panama in the past, those are here interpreted as a distinct species, V. montana . Based on our interpretation, V. guatemalensis is restricted to Mexico, Guatemala and Honduras. It is distinguished from V. montana by a series of characters, described below. While V. laevigata was typified with material from the Pacific slope in Chiriquí, Panama and is frequently considered a synonym of V. guatemalensis (e.g. Smith and Wodehouse 1938; Standley and Steyermark 1946; Duke 1962), we treat these as morphologically distinct; differences between these two species are discussed under V. laevigata .

Notes.

The specimens, identified in Gómez-Laurito and Ortiz (2004), Haber (2014) and Monro et al. (2017) as V. guatemalensis or V. surinamensis , correspond to V. montana (R. Aguilar 1131, [E.] Bello 470, M. Chinchilla 3, J. Gómez-Laurito 11846, 11970, J. González 823) or V. amistadensis (A. Chacón 194). Additionally, a specimen cited as V. guatemalensis in Flora of Panama ( Duke 1962) corresponds with the type of V. laevigata (G. P. Cooper & G. M. Slater 308).

The specimen J. A. Steyermark 47624 (MO-3 sheets!) seems to represent two different individuals - two of the sheets have staminate flowers, while the third has immature fruits. The duplicate of this collection at the Field Museum (digital image) has two sheets, both from a pistillate individual: one sheet carries flowers and the other immature fruits.

Virola guatemalensis , along with another Mesoamerican species, V. koschnyi (see below), produce the largest grains of pollen in Virola ( Walker and Walker 1979).

Selected specimens seen.

Mexico. Chiapas: Tila, Chewupaj, 1000 m elev., 10 Dec 1982 (fr), A. Méndez 5223 (MEXU!*, MO!); Peltalcingo, slope of Ahk’ulbal Nab above Peltalcingo, 1700 m elev., 27 Feb 1981 (fr), D. Breedlove 49868 (MEXU!*); La Trinitaria, 10 km east northeast of Dos Lagos above Santa Elena, 1170 m elev., 19 Jan 1982 (fr), D. Breedlove & F. Almeda 57553 (MO!); La Independencia, ridge, 45-50 km E of Lagos de Montebello National Park on road to Ixcán from Santa Elena, 760 m elev., 22 Jan 1982 (fr), D. Breedlove & F. Almeda 57735 (MO!). Oaxaca: Matías Romero Avendaño, ± 11 Km. al S de Aserradero La Floresta, ± 24 Km. al S de Esmeralda, lomas al S del Río Verde, 300 m elev., 24 Apr 1981 (fl), T. Wendt et al. 3235 (MEXU!*). Veracruz: Cima del cerro Vigia, 450 m elev., 19 Apr 2005 (♂ fl), E. Velasco-Sinaca 678 (MEXU!*, MO!); La Escondida, 3 km NO de Estación de Biología Tropical Los Tuxtlas, 200 m elev., 05 Apr 1983 (♂ fl), G. Ibarra 604 (MEXU!*, MO!); Estación de Biología Tropical Los Tuxtlas, 200 m elev., 29 Oct 1983 (fr), G. Ibarra 957 (MO!); camino Laguna Escondida, 2 km NW de la Estación de Biológia Tropical Los Tuxtlas, 200 m elev., 08 May 1984 (♂ fl), G. Ibarra & G. Gómez 1607 (MO!); Ocotal Grande, 5 km N de Mecayapan, 1000 m elev., 14 Mar 1985 (fr), G. Ibarra et al. 2357 (MEXU!*, MO!, NO!); 2 km al NW del rancho Rubén Sánchez, 250-350 m elev., 28 May 1985 (♂ fl), G. Ibarra 2450 (MO!); Santiago Tuxtla, Alta Luz, 540 m elev., 11 May 1968 (♂ fl), M. Sousa 3680 (MO!); Catemaco, Tebanca, no elev., 20 Oct 1971 (fr), J. I. Calzada 615 (MO!); Cerro El Vigia de Santiago Tuxtla, 800 m elev., 18 Feb 1967 (fr), R. Cedillo 9 (MO!); Catemaco, El Chinchero, 14.2 km al SE de Tebanca camino al río Huacinapan, no elev., 19 Dec 1984 (fr), R. Cedillo & G. Pérez 2958 (MEXU!*, MO!, NO!); Hidalgotitlán, lomitas al SE de Poblado 6, 150 m elev., 27 Apr 1982 (♂ fl), T. Wendt et al. 3898 (LSU!, MO!). Guatemala. Sololá: Bordering barranco on Finca Olas de Mocá, just west of Finca Mocá, south-facing slopes of Volcán Atitlán, 1000-1100 m elev., 15 Jun 1942 (♂ fl, imm fr), J. A. Steyermark 47624 (F!*, MO-3 sheets!). Honduras. Yoro: Cascada de Río Guán Guán, 400-440 m elev., 20 Apr 1995 (♂ fl), T. Hawkins & M. Merello 768B (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 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 9. Geographic distribution of Virola allenii (A), V. amistadensis (B), V. chrysocarpa (C), V. elongata (D), V. fosteri (E) and V. guatemalensis (F).

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).