New Species of Virola (Myristicaceae) from South America Author Santamaria-Aguilar, Daniel https://orcid.org/0000-0002-6339-8250 Shirley C. Tucker Herbarium, Department of Biological Sciences, Louisiana State University, 103 Life Sciences Building, Baton Rouge, Louisiana 70803 - 1705, USA & Missouri Botanical Garden, St. Louis, Missouri, USA daniel.santamaria366@gmail.com Author Lagomarsino, Laura P. Shirley C. Tucker Herbarium, Department of Biological Sciences, Louisiana State University, 103 Life Sciences Building, Baton Rouge, Louisiana 70803 - 1705, USA text PhytoKeys 2022 2022-05-30 197 81 148 http://dx.doi.org/10.3897/phytokeys.197.81367 journal article http://dx.doi.org/10.3897/phytokeys.197.81367 1314-2003-197-81 37EA452A15155D1CB85CA350AF70A112 4. Virola calimensis D. Santam. sp. nov. Type . Colombia . Valle del Cauca : Bajo Calima , concesion Pulpapel / Buenaventura , 03°55'N , 077°00'W , 100 m , 03 Aug 1984 (fr), M. Monsalve 158 ( holotype : MO! [accession 3624779, barcode MO-2657915]; isotype: INPA [accession 147738; image!], JAUM [accession 007410]). Fig. 9 Diagnosis. Virola calimensis was previously confused with V. calophylla and V. macrocarpa . Perhaps because of stellate, sessile trichomes and lateral veins that are well separated. Morphologically it differs from V. calophylla in having flowers with longer staminate perianth (2.5-3.5 mm vs. 1-2.1 mm), the internal surface densely pubescent (vs. glabrescent), long anthers (0.6-1 [-1.2] mm vs. 0.4-0.5 mm), and large fruits (2.8-3.1 cm long vs. 1.6-1.5 cm). Differs from V. macrocarpa in leaf blades abaxially densely pubescent (vs. sparsely pubescent), fruits globose (vs. ellipsoid), shorter (2.8-3.1 cm long vs. [2.7-] 3.5-4.5 cm). Tree 5-20 or 30 m tall x 25-28 cm in diameter, inner bark described once as brown, cracked, rough. Exudate reddish, location of exudate on plant not stated, or described as being present in the inflorescence, but with no information about color. Twigs 0.3-0.33 cm thick, terete or slightly compressed, inconspicuously pubescent, trichomes dendritic, sessile, brown reddish, without lenticels. Leaves young terminal bud not seen; petiole 1.5-2.4 x 0.22-0.4 cm, slightly canaliculate, sometimes slightly winged, glabrescent, or inconspicuously tomentose to tomentulose, the trichomes dendritic; leaf blades 19-25 x 7.3-11 cm, widely oblong to elliptical; adaxial surface of mature leaves drying brown to olive, the surface smooth, sometimes shiny, glabrous; abaxial surface drying brown reddish to grayish brown, densely pubescent, the trichomes stellate, ca. 0.1 mm diameter, sessile, the central part of the trichome dark reddish, the branches pale reddish; lateral veins ca. 12-13 ( W. Devia 3086 ), 3-4 veins per 5 cm, spaced 1.3-2.3 cm apart, flat to slightly raised on adaxial surface, raised on abaxial surface and slightly darker than on adaxial surface, puberulent, densely tomentose to glabrescent above, densely pubescent to the sides, arcuate-ascending distally, slightly anastomosing near the margin and without forming a marked intramarginal vein; tertiary veins slightly visible on both sides; midvein adaxially slightly elevated, abaxially raised, rounded to rectangular, glabrescent to tomentose; base obtuse, not revolute, flat; margin flat; apex acuminate. Staminate inflorescence 6-8 cm long, axes flattened, tomentose, with trichomes dendritic, brown-reddish; peduncle 1.3-2.8 x 0.28-0.4 cm; main axes with 5-11 ramifications, the first pair opposite to subopposite, the others alternate; bracts not seen. Staminate flowers (in bud) in lax terminal fascicles of 3-5 flowers, on a receptacle 3-5 mm wide; perianth 2.5-3.5 mm long, elongate to rhomboid, fleshy, ferruginous when fresh (probably by the trichomes), connate to 1.1-1.7 mm in length, external surface densely pubescent with ferruginous and dendritic trichomes, internal surface densely pubescent (especially in the lobes) with ferruginous trichomes; lobes 3, 1.3-2 x 0.9-1.5 mm and 0.3-0.7 mm thick, without resinous punctuations when rehydrated; stamens 3, the filament column 0.4-0.6 (-0.8) mm long and 0.2-0.4 mm wide, glabrous, straight, not constricted at the apex; anthers 0.6-1 (-1.2) mm long and 0.2-0.3 mm wide; apiculus 0.1-0.2 mm long, acuminate, slightly separated distally. Pistillate inflorescence and flowers unknown. Infructescence ca. 4.5 cm long, with 2 fruits, peduncle ca. 1.7 x 0.5 cm. Fruits ca. 3.3-4.2 x 2.8-3.1 cm, color not described when fresh, globose, shortly stipitate, densely tomentose, the trichomes dendritic, sessile, ferruginous, the trichomes not falling easily like dust, the surface slightly rugose, the line of dehiscence slightly carinate, the base obtuse, the apex acute; pericarp ca. 3.8-4 mm thick; pedicel ca. 0.6 cm long. Seed ca. 2.5 x 2.1 cm, the testa when dry brown reddish, slightly ribbed from the base to the apex; aril color when fresh not described, blackish to brown reddish when dry, the texture dry and thin, laciniate almost to the base, in narrow bands distally. Distinctive characters. Virola calimensis can be recognized by its broad staminate inflorescence, few-flowered inflorescences, elongate to rhomboid, fleshy, densely pubescent perianth, very fleshy filament column and shorter (0.4-0.6 [-0.8] mm long) than the anthers (0.6-1 [-1.2] mm long), and globose, tomentose fruit with trichomes that do not fall easily and with a thick (3.8-4 mm) pericarp (Fig. 6D ). Like other species described here, the abaxial side of V. calimensis ' leaves are covered with stellate, sessile trichomes whose central portion is darker than the branches on their abaxial surfaces. It further has lateral veins that are well separated (1.3-2.3 cm spaced) (Fig. 4D ) and a relatively long and thick petiole. Etymology. The specific epithet of the new species refers to the Bajo Calima region (Valle del Cauca department, Colombia), the region where most specimens of this new species come from. Distribution. Virola calimensis occurs in the Pacific coast of Colombia (Valle del Cauca department) and Ecuador (Esmeraldas province) at elevations ranging from 5 to 260 m (Fig. 18C ). According to the field notes of the specimens collected in Colombia (e.g., M. Monsalve 158 ), the species grows in nutrient poor soils with a high concentration of aluminum. See Gentry (1986) , Faber-Langendoen and Gentry (1991) and Marcano-Berti and Aymard C. (2021) for more information about this region. Phenology. Staminate flowers of Virola calimensis have been recorded in March, May, June, July, and September. Fruits have been observed in June and August. Pistillate flowers were not seen in the studied material. Common name and uses. Cuangare (Colombia; M. Monsalve 158 ). Preliminary conservation status. Virola calimensis is Endangered following IUCN criteria B1a and B2a. It is known from three localities, has an EOO of 313 km2, and an AOO of 12 km2. While its small range justifies this preliminary status, V. calimensis benefits from growing in regions with relatively low rates of deforestation compared to the rest of the region ( Antonelli 2022 ), which is in part due to collective land titling by Afro-Colombian communities in Valle del Cauca, Colombia ( Velez et al. 2020 ) and proximity to the Awa Reserve ( Oviedo 2006 ). Discussion. Herbarium specimens of Virola calimensis were previously identified as V. calophylla and/or V. macrocarpa . This misidentification was probably due to characteristics shared with V. calophylla : stellate, sessile trichomes and lateral veins that are well separated, as well as their lax, few-flowered staminate inflorescences that are relatively wide (these traits are unknown in V. macrocarpa ). However, V. calimensis differs from V. calophylla in having fleshier, longer staminate perianth with indument on the internal surface and a filament column that is shorter than the anthers. The new species also differs in the size and shape of its fruit as compared to V. calophylla and V. macrocarpa ). A summary of the characteristics that differentiate these three species is presented in Table 4 , and 5 . Table 4. Comparison of Virola calimensis , with V. macrocarpa . †From Smith and Wodehouse (1938) .

Morphological character	V. calimensis	V. macrocarpa
Leaf blades size, and pubescence abaxially	19-25 x 7.3-11 cm; densely pubescent (Fig. 4D )	20-40 x 7-11 cm†; sparsely pubescent (Fig. 4J )
Spaced lateral veins	1.3-2.3 cm apart	1.7-3 cm apart
Fruit shape, and long	Globose, 2.8-3.1 cm (Fig. 6D )	Ellipsoid, (2.7-) 3.5-4.5 cm (Fig. 6H )
Pericarp thickness	ca. 3.8-4 mm	(2.7-) 3-4.7 mm
Seed size	ca. 2.5 x 2.1 cm	2.2-2.5 x 1.5-1.7 cm†
Habitat	Lowland rain forest, Colombia (Valle del Cauca), and Ecuador (Esmeraldas), at 20 to 260 m elevation	Montane forests, Andes of Colombia ( Boyaca ), at 1100 m elevation†

Notes. The first collection of Virola calimensis was made 76 years ago by Jose Cuatrecasas ( J. Cuatrecasas 17540 ; 5-15 May 1944) in Rio Cajambre, Valle del Cauca, Colombia. Six years later, Smith (1950) attributed this Cuatrecasas collection to V. macrocarpa , which he used to describe staminate inflorescences; however, he expressed doubt in this and stressed the need for more material, stating that this collection " has leaves considerably smaller , although similar in texture, shape, and indument ." Smith (1950) refers to two additional collections from Colombia with very young inflorescences, which we could not study, as V. macrocarpa ; these are J. Cuatrecasas 15596, 16613 (A, F [as Ch]) from the Western Cordillera (1250-1400 m elevations) and Choco region (5-50 m elevations) respectively. Table 5. Comparison of Virola calimensis , with V. calophylla .

Morphological character	V. calimensis	V. calophylla
Leaf blade size, and base	19-25 x 7.3-11 cm, obtuse	(15-) 20-60 x 10-16 cm; (usually) deeply cordate to truncate (obtuse)
# of lateral veins	ca. 12-13	11-28
Length of staminate inflorescences	6-8 cm	6-30 cm
Staminate perianth long, and internal surface	2.5-3.5 mm; densely pubescent	1-2.1 mm; glabrescent
Long filament column	0.4-0.6 (-0.8) mm	0.2-0.6 mm
Long anthers	0.6-1 (-1.2) mm	0.4-0.5 mm
Fruit size	ca. 3.3-4.2 x 2.8-3.1 cm (Fig. 6D )	2.5-3 x 1.6-1.5 cm (Fig. 6E )

Walker and Walker (1979 ; fig. 44) illustrated the pollen of the first collection of this species (again, J. Cuatrecasas 17540 ; 5-15 May 1944), then attributed to V. macrocarpa . It was assigned to Pollen Type I (more information can be found in their publication). The specimens C. Jativa & C. Epling 1143 and M. Monsalve 158 , mentioned under V. macrocarpa in Flora of Ecuador ( Jaramillo et al. 2004 ) and La Familia de Arboles Tropicales Myristicaceae en el Departamento del Valle del Cauca, Colombia ( Taylor and Devia Alvarez 2000 ) correspond to this new species. The digital images of W. Devia et al. 5091 (st), 5461 (st), all at TULV, appear to correspond with V. calimensis as well. Specimens examined. Colombia . Valle del Cauca : Bajo Calima , 15-20 m , 28 Jun 1961 (fr), I. Cabrera 556 (COL) ; Costa del Pacifico , Rio Cajambre , Silva , 5-80 m , 5-15 May 1944 ( ♂ fl), J. Cuatrecasas 17540 (A!, F [image!]); Buenaventura , Carton de Colombia , Via a Malaga Km. 22, Frente Hans , [ 03°59'47"N , 076°58'28"W ], 20 m , 01 Mar 1990 (fl bud), W. Devia 3086 (MO-2 sheets!, TULV [image!], UPCB [n.v]); B/ventura [Buenaventura], Canalete, km 28 via Malaga , 50 m , [s.d.] May 1991 (fr), W. Devia 3253 (NY!); Bajo Calima, concesion Pulpapel/ Buenaventura, 03°55'N , 077°00'W , 100 m , 11 Mar 1986 ( ♂ bud fl), M. Monsalve 968 (INPA [image!], MO!); ibid., 22 Sep 1987 ( ♂ bud fl), M. Monsalve 1769 (F [image!], INPA [image!], MO!). Ecuador . Esmeraldas : At Tobar Donoso, junct. of Rio San Juan and Rio Camumbi , [ 01°11'24"N , 078°30'30"W ], 260 m , 25 Jul 1966 ( ♂ fl), C. Jativa & C. Epling 1129 (NY!); ibid., 27 Jul 1966 ( ♂ fl), C. Jativa & C. Epling 1143 (MO!, NY!, US [image!]) .