Virola koschnyi 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

7. Virola koschnyi Warb. Fig. 17

Virola koschnyi Warb. Repert. Spec. Nov. Regni Veg. 1: 71. 1905. Type. Costarica [Costa Rica]. [Alajuela] San Carlos, Th. Koschny s.n. (lectotype, here designated: F!*, fragment with B photo [649050, F0360191F]); Costa Rica. Heredia. [Sarapiqui] Parque Nacional Braulio Carrillo, bosque primario frente al Puesto La Ceiba, 450 m elev., 23 Dec 1988 (♂ fl), M. Ballestero 72 (epitype, designated here: CR! [16354]; isoepitypes: CR! [156734], MO! [5550256, MO-299037].

Virola merendonis Pittier., Contr. U.S. Natl. Herb. 20: 453. 1922. Type. [Honduras]. [ Copán:] Collected in the forests of Cuchillitas, between Arranca Barba Hills and Mohanes, in the Cordillera de Merendon, borders of Guatemala and Honduras, in fruit, [10-] 18 May 1919 [fr], H. Pittier 8530 (holotype: US!*; isotype: NY!*).

Virola costaricensis nomen nudum

Distinctive characters.

Virola koschnyi can be recognised by the densely tomentose leaf undersides with pediculate, dendritic trichomes ( Fig. 3G ) that are soft to the touch, the numerous lateral veins [(16-) 20-35] that form a clear submarginal vein and tertiary veins that are usually inconspicuous abaxially ( Figs 3G , 8G ). Adaxially, the new leaves are densely covered with dendritic sessile or subsessile trichomes that are deciduous at maturity, resulting in glabrous to glabrescent and smooth adxial surface on mature leaves. The staminate flowers have filament columns that are 0.7-0.9 (-1.4) mm long with anthers 0.5-0.7 (-1) mm long. Its fruits are 1.9-3.1 × 1.5-1.9 cm and ellipsoid or subglobose ( Fig. 4F ), with a pericarp 1.2-3.1 mm thick.

Distribution.

Virola koschnyi is known from Mexico (Chiapas), Guatemala (Izabal and Petén), Belize (Cayo, Toledo and Stann Creek), Honduras ( Atlántida, Cortés, Gracias a Dios and Olancho), Nicaragua ( Atlántico Norte, Atlántico Sur, Jinotega and Río San Juan), Costa Rica (Alajuela, Guanacaste, Heredia, San José and Limón), and Panamá (Bocas del Toro) ( Fig. 18A ). It is found mainly on the Caribbean slope, where it has been recorded between 10-1000 (-1700) m elevation.

We consider this species restricted to Mesoamerica. The collections identified as V. koschnyi from Ecuador in Catalogue of the Vascular Plants of Ecuador (i.e. C. H. Dodson et al. 6465 [RPSC, SEL]; Jørgensen and León-Yánez 1999) correspond to V. multinervia Ducke ( Jaramillo et al. 2004). Determinations of V. koschyni cited for Colombia by Cogollo et al. (2007) are doubtful and do not cite voucher specimens, while those of Gradstein (2016) could not be confirmed.

Common names.

Guatemala: Cedrillo, drago, sangre. Belize: banak, black banak. Honduras: bának, sebo, sangre de montaña, sangre real. Nicaragua: banak blanco, banak colorado, cebo, fruta dorada, sebo. Costa Rica: achiotillo, fruta dorada.

Phenology.

Flowering of Virola koschnyi has been recorded in January, March and May, July, September, October and December. Only three herbarium specimens with pistillate flowers were seen, two from Costa Rica and one from Nicaragua. Fruits are produced throughout the year, though most often collected in March, May and June.

Field characters.

Plants are trees (4-) 10-70 m tall and 12-80 cm DBH, with a trunk that is straight with well-developed buttresses. The bark is described as scaly and falling off in small plates or as smooth, red-brown or whitish in colour; clear, red or reddish exudate is released when damaged. Flowers usually have yellow perianth, though it can sometimes be white, brown, or orange. The mature fruit is yellow, orange or brown, with a red or pink-red aril. The testa of the seed is black, brown or white.

Typification.

Virola koschnyi was described by Otto Warburg based on a collection made by Theodor Koschny ( Warburg 1905). Warburg indicated in the protologue "Costarica: San Carlos, leg. Th. Koschny," but did not mention where the type was housed. The type is presumed to have been housed at Berlin (B) and subsequently destroyed during World War II.

The only known original material of Virola koschnyi is a fragmentary specimen accompanied with a photograph at the Field Museum. The photo shows the original specimen was composed of three fertile branches with leaves (two with inflorescences and one with fruit), with a handwritten annotation of " V. costaricensis Warb", presumably by Warburg, though this name was never validly published. The fragment material consists of pieces of leaves, two inflorescence branches with a few immature flowers and a broken fruit comprising pericarp, testa and seed. Since they are mounted with the photo of the Berlin specimen, these fragments presumably originated from the holotype. Although it is not ideal type material due to its fragmentary nature, the specimen at F appears to be the only extant original material and is sufficient to confirm that it coincides with the protologue and concept of the species used and so is here designated as lectotype. In order to ensure the precise application of the name, given the fragmentary type material, an epitype was selected from the studied specimens.

Notes.

Most of the specimens from the Pacific slope of Costa Rica and Panama previously identified as V. koschnyi are here interpreted as V. chrysocarpa . The similarities and differences between V. koschnyi and V. chrysocarpa are discussed under the latter species. While most fruiting specimens of V. koschnyi have obtuse to rounded apices, G. Herrera 1092 (CR!, MO!) from Costa Rica and R. Rueda & H. Mendoza 17069 (MO!) from Nicaragua have apiculate apices. Aside from the fruit apex, these specimens conform to the species concept adopted here in all other characters (e.g. number of lateral veins, submarginal vein and tertiary veins that are usually inconspicuous abaxially and pediculate, dendritic trichomes).

Selected specimens seen.

Mexico. Chiapas: Ocosingo, En ejido Chaju, 150 m elev., 17 Mar 1993 (fl), E. Martínez & C. H. Ramos 26336 (MEXU-4 sheets!*); Boca de Chajul, 800 m al SE del poblado, 350 m elev., 21 May 1992 (fr), G. Domínguez 438 (MEXU!*). Belize. Cayo: At mile 28.5 m on Hummingbird highway, 200-300 ft [60-90 m] elev., 14-21 Jun 1973 (fr), T. B. Croat 24562 (MO!). Stann Creek: Mountain Cow, [200 m elev.], 25 Mar 1940 (imm fr), P. H. Gentle 3277 (MO!); Big Eddy Ridge, [50-200 m elev.], 21 May 1940 (fr), P. H. Gentle 3347 (MO!); 22 miles Stann Creek, 200 ft [60 m] elev., 27 Jun 1927 (fr), W. A. Schipp 949 (MO-2 sheets!). Toledo: Bladen Nature Reserve, 20 m from the Bladen River, 45 m elev., 27 Feb 1997 (♂ fl), S. W. Brewer 178 (MO!); Southern Maya mountains, Bladen Nature Reserve, 250 m elev., 23 May 1996 (fr), G. Davidse 36211 (MO!). Guatemala. Izabal: Between Seja and Fronteras, no elev. 08 May 1971 (fl bud), E. Contreras 10751 (MO!); Livingston, Creek Jute, Biotopo Chocón Machacas, [0-5 m elev.], 28 Jul 1988 (fr), P. Tenorio et al. 14953 (MEXU!*, MO!). Petén: Chinchila, Sebol road, [20 m elev.], 16 May 1967 (fr), E. Contreras 6924 (MO!); La Cumbre, [300 m elev.], 30 Jul 1969 (fr), E. Contreras 8790 (MEXU!*, MO!). Honduras. Atlántida: Lancetilla S of Tela, 20-50 m elev., 25 Jun 1970 (fr), G. Davidse & R. Pohl 2178 (MO!). Cortés: [Santa Cruz de Yojoa], 660 m elev., 13 Feb 1952 (fl), P. H. Allen 6456 (GH!*, EAP-2 sheets!*). Gracias a Dios: La Mosquitia, Ahuas Bila, 100 m elev., 5-13 May 1985 (fr), C. Nelson & G. Cruz 9484 (MO!). Olancho: San Esteban, montaña El Carbón, 460 m elev., 02 Feb 1994 (fl bud), A. Meras 02 (MO); a 3 km comunidad El Carbón, 430 m elev., 12 Mar 1997 (fl), I. Rivas 08-C1 (EAP!*). Nicaragua. Atlántico Norte: Bonanza, Musawas, 50-150 m elev., 17 Oct 2002 (♂ fl), C. Aker et al. 624 (MO!); Siuna, Waspado, 100-120 m elev., 06 Oct 1982 (imm fr), F. Ortíz 252 (MO!); sur de río Wawa, 40 m elev., 16 Mar 1971 (imm fr), E. L. Little Jr. 25164 (MO!). Atlántico Sur: Kurinwac[s]ito, 80-100 m elev., 18-22 Mar 1984 (♀ fl, fr), P. P. Moreno 23682 (MO), P. P. Moreno 23701 (MO!); El Zapote, 40 km NE de Nueva Guinea, 130-150 m elev., 29 Feb 1984 (♂ fl), J. C. Sandino 4808 (MO!). Estelí: El Zacatón, entre Mesas Plan Helado y la laguna de Miraflor, 1300-1400 m elev., 29 Mar 1983 (imm fr), P. P. Moreno 23788 (MO!). Jinotega: Cua Bocay, Reserva de Bosawas, comunidad de San Andrés, 180 m elev., 30 Jun 2005 (fr), I. Coronado et al. 1951 (MO!). Río San Juan: El Castillo, Reserva Indio Maíz, 100-200 m elev., 15 Mar 1999 (♂ fl), R. Rueda el al. 10319 (MO!); San Juan del Norte, Reserva Indio Maíz, 30 m elev., 18 May 2002 (fr), R. Rueda & H. Mendoza 17069 (MO!); río Sábalos, 2 km de Santa Eduviges, 80 m elev., 18 Feb 1984 (♂ fl), P. P. Moreno 23042 (MO!). Costa Rica. Alajuela: Parque Nacional Rincón de La Vieja, 900-1000 m elev., 07 Mar 1988 (♀ fl), G. Herrera 1607 (CR!, MO!); Upala, Dos Ríos, 500 m elev., 02 Nov 1987 (fr), G. Herrera 1092 (CR!, MO!); Upala, Estación San Ramón, 550 m elev., 27 Jan 1995 (fr), F. Quesada 243 (CR!, LSU!, MO!); Upala, Finca San Gerardo, 600 m elev., 20 Feb 1998 (fl bud), F. Quesada et al. 577 (CR-2 sheets!, MO!, USJ!). Cartago: Turrialba, CATIE, 650 m elev., 08 Feb 1975 (fr), C. Vaughan 93 (USJ!); Bosque de Florencia de Turrialba, 650 m elev., 31 Jul 1969 (fr), R. Ramalho 7990 (USJ!). Guanacaste: Santa Cruz, road from Santa Cecilia to La Esperanza, 340 m elev., 29 Dec 1989 (fr), R. E. Gereau et al. 3455 (CR-2!, MO!); Parque Nacional Guanacaste, Estación Pitilla, 700 m elev., 02 Mar 1991 (♂ fl), P. Ríos 309 (CR-2 sheets!, MO!). Heredia: Parque Nacional Braulio Carrilo, Estación El Ceibo, 500-600 m elev., 01 Oct 1989 (fl bud), R. Aguilar 9 (MO!, LSU!, USJ!); Los Arbolito, al N de Puerto Viejo, 20 m elev., 09 Mar 1993 (♂ fl), F. Araya 197 (CR-2 sheets!, LSU!, MO!); between La Selva entrance and bridge before Puerto Viejo, [100 m elev.], 18 Apr 1981 (fr), J. Folsom 9791 (MO!). Limón: Refugio de Vida Silvestre Gandoca Manzanillo, sendero Cerillo, 1 m elev., 03 Mar 1999 (fr), U. Chavarría 1918 (CR-2 sheets!, MO!); Cordillera de Talamanca, ridge separating Río Madre de Dios from Quebrada Cañabral, 440 m elev., 02 Sep 1988 (♀ fl), M. Grayum et al. 8692 (CR-2 sheets!, MO-2 sheets!, USJ!); Sixaola, Quebrada Mata de Limón, 20-40 m elev., 27 Jan 1987 (fr), M. Grayum et al. 8004 (CR!, MO!); Río Peje, no elev., 14 Jun 1979 (fr), R. Ocampo 2582 (CR!); Parque Nacional Tortuguero, Estación Agua Fría, 40 m elev., 02 Feb 1988 (fr), R. Robles 1587 (CR!, MO!); Cerro Coronel, 20-170 m elev., 16-23 Jan 1986 (♂ fl), W. D. Stevens 23768 (CR, MO). San José: Z. P. [Zona Protectora] La Cangreja, Santa Rosa de Puriscal, 350 m elev., 01 Oct 1992 (fr), J. F. Morales 764 (CR). Panama. Bocas del Toro: Isla Colon, camino central, unos 4 km de Boca de Drago, 2-25 m elev., 16 Mar 1993 (fr), R. Foster et al. 14549 (F!*).

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 17. Virola koschnyi A lower trunk and buttresses B trunk C branching D leaf blades on abaxial surface E leaf blades on adaxial surface F twig, petiole and leaf base G leaf venation on adaxial surface (left) and abaxial surface (right) H branch with fruits, showing the interior part of the fruit I seed J mature fruit. All photos by Reinaldo Aguilar from https: // sura. ots. ac. cr / florula 4 / index. php, except G by D. Santamaria-Aguilar.

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