A taxonomic synopsis of Virola (Myristicaceae) in Mesoamerica, including six new species

Abstract A taxonomic synopsis of Virola (Myristicaceae) is presented for Mesoamerica. Fourteen species are recognised, amongst them six are described and published as new, based on morphology: V. allenii D.Santam. & Aguilar, sp. nov. from Costa Rica, V. otobifolia D.Santam., sp. nov. from Panama and V. amistadensis D.Santam., sp. nov., V. chrysocarpa D.Santam. & Aguilar, sp. nov., V. fosteri D.Santam., sp. nov. and V. montana D.Santam., sp. nov. from both Costa Rica and Panama. Additionally, a lectotype is designated for V. koschnyi, accompanied by an epitype in view of the fragmentary material. Finally, we recognise V. laevigata and V. nobilis as morphologically distinct species, though these are frequently considered synonymys of V. guatemalensis and V. surinamensis, respectively. Of the fourteen accepted species, twelve of them are endemic to Mesoamerica, while the remaining two species (V. elongata and V. sebifera) extend into South America. Illustrations, species diagnoses and distribution maps for each species are provided, as is an identification key to all species.

Virola is the largest genus of Myristicaceae in America and, with about 60 species, it is the fourth largest genus in the family. The plants have two particularly distinctive characteristics: their growth form and the mature fruit. The growth form, sometimes called myristicaceous branching (e.g. Gentry 1993), corresponds largely with Massart's Model (Hallé et al. 1978), which is characterised by a monopodial, orthotropic principal shoot with plagiotropic lateral branches that are developed in regular tiers by rhythmic branching of the main axis (Kühn and Kubitzki 1993). Fruits of Virola are distinctive and striking: they dehisce by two valves when ripe, displaying their single seed, which is covered by a brightly coloured laciniate aril that is rich in proteins, lipid and sugars. Birds and monkeys are primary dispersers of these seeds (Howe and Vande Kerckhove 1980;Howe 1981Howe , 1982Howe , 1993Sabatier 1997;Moreira et al. 2017). Not surprisingly, given the importance of their fruits as a food for wildlife and its abundance in many lowland forests, the ecological importance of Virola has been extensively documented (e.g. Frankie et al. 1974;Pitman et al. 2008;Steeves 2011;Riba Hernández 2017). Below, we provide a review of the morphology, biogeography, taxonomic history and ethnobotanical uses of Virola.

Morphology
As with many magnoliids, Virola has aromatic tissues. Unlike other magnoliids, though, when the bark is cut or a twig is broken, it yields an exudate that is initially watery and clear and quickly changes to red or pinkish. The leaves, which are exstipulate and simple with entire margins and occasionally have pellucid punctuation, are distichous. Most surfaces of the plant, including leaves, are usually covered with stellate or dendritic trichomes. Staminate inflorescences are paniculate, with few to many branches. Flowers, sometimes reported as fragrant, are unisexual, ebracteolate and inserted on a receptacle; they are produced in lax to dense clusters. Their yellow or brown perianth is connate to varying degrees and uniseriate, usually with 3-lobes. The androecium is compound, with filaments fused into a column, with 3-(4-6) anthers fused to the column. The pistillate inflorescences are shorter than staminate inflorescences. Pistillate flowers have a perianth that is more carnose than those of the staminate flowers and their gynoecia are pubescent and sessile or short-stipitate, topped by a stigma that is usually bilobed (Smith and Wodehouse 1938;de Wilde 1991;Kühn and Kubitzki 1993;Jaramillo et al. 2004).
In Mesoamerica, where five of the six native genera of American Myristicaceae occur ( Fig. 1), Virola can be identified using the following key:

Biogeography
Virola is found throughout tropical America. Species have been collected from Mexico to southern Brazil, though are notably absent in El Salvador. Occurrence in the Caribbean is limited: one species is found in the West Indies (V. surinamensis [Rol. ex Rottb.] Warb.), though a fossil Virola flower, recently described from Dominican Republic (Poinar and Steeves 2013), suggests that genus has persisted over geological time in the region. Species richness is highest in western Amazonia, with the greatest concentration of species in Brazil (~35 spp.), Colombia (~ 29 spp.) and Peru (~ 23 spp.), where they are found mainly in humid forests, though some species grow in dry formations (e.g. V. sessilis [A. DC.] Warb., V. subsessilis [Benth.] Warb.) (Rodrigues 1980(Rodrigues , 2015Gradstein 2016;Vásquez et al. 2018). Within Mesoamerica, the greatest concentration of Virola species occurs in Costa Rica and Panama. Together, these two countries contain 10 and 11, respectively, of the 14 Mesoamerican species (Fig. 2). Of the 14 species treated in this synopsis, 12 are endemic to Mesoamerica, while V. elongata (Spruce ex Benth.) Warb. occurs from Panama to Brazil and V. sebifera Aublet from Honduras to Brazil; both of these species are widespread in South America. The majority of Virola species occupy lowland wet forest below 800 m elevation, with a few species reaching montane forests up to 2000 m elevation (e.g. V. montana). A single montane species of Virola can occur on both slopes, though, within a country or region, there is a tendency to find them on only one, more often the Caribbean slope (e.g. V. fosteri, V. amistadensis, V. multiflora). A list of the current species of Virola in Mesoamerica with geographic distribution by country, elevations and slope is given in Table 1.

Ethnobotany
Virola has diverse ethnobotanical uses and there are many reports of local use of several species of Virola for many non-medicinal purposes. For example, the oils from the seeds of Virola (e.g. V. guatemalensis, V. sebifera) are used as lubricant in machinery and to make soap and candles that emit intense light, produce little smoke and smell pleasant; Pesce 2009). Instead of using the extracted oils, indigenous people of the Talamanca mountains of Costa Rica (Pittier 1978) and the Cuna tribe of Panama (Duke 1975) burn whole seeds as candles. Wood from many species, including Virola fosteri, V. koschnyi, V. sebifera and V. surinamensis, is used for construction and carpentry, to make matchsticks, as pulp for paper and to manufacture plywood (Rodrigues 1972;Flores-Vindas and Obando-Vargas 2014;Pérez and Condit 2018). The seeds and arils of Virola guatemalensis, a common shade tree in coffee plantations in Veracruz, Mexico (Sánchez , are used to flavour chocolate drinks and other beverages (Warburg 1897; Standley and Steyermark 1946;Pennington and Sarukhánn 2016).
Virola also produces many chemicals that are biologically active in humans. For example, Virola is perhaps best known for its hallucinogenic properties, which are often  incorporated into indigenous cultural practices of South America. The hallucinogen is usually obtained from the exudate of the inner bark of several species, including: Virola calophylla Warb., V. calophylloidea Markgr., V. duckei A. C. Sm., V. elongata, V. sebifera, V. surinamensis and V. theiodora Warb.; it has also been documented in Iryanthera and Osteophloeum platyspermum (Spruce ex A. DC.) Warb. (Schultes 1954(Schultes , 1969(Schultes , 1979Schultes and Holmstedt 1968;Prance 1970;Soares Maia and Rodrigues 1974;Bennett and Alarcón 1994). In addition to being psychoactive, the reddish exudate of various Virola species is thought to have medicinal properties. For example, the sap of V. melinonii (Benoist) A. C. Sm. and V. surinamensis is applied to treat skin conditions, alleviate tooth pain, soothe colic and as a styptic (i.e. a substance used to stop bleeding) to treat ulcerating sores and wounds, haemorrhoids and other sources of bleeding (Plotkin and Schultes 1990). The sap of an undetermined species of Virola has been reported to be used as a contraceptive by Indians on the Rio Negro of Brazil and is reputedly effective for a period of two or three years (Plotkin and Schultes 1990). Finally, the oil obtained from the seeds of Virola officinalis Warb, V. sebifera and V. surinamensis has many medical folk uses, including treatments for asthma, rheumatism, tumours of the joints, intestinal worms, skin diseases, erysipelas, haemorrhoids and bad breath (Rodrigues 1972(Rodrigues , 1980Plotkin and Schultes 1990).

Taxonomic history
Virola was first described by Aublet in "Histoire des plantes de la Guiane Françoise", based on V. sebifera Aublet (Aublet 1775). It is likely that the etymology of the generic name is derived from one of the common names mentioned in the protologue, which stated: "Nomen Caribaeum Oyapocenfibus VOIROUCHI; Sinemarienfibus VIROLA; Galliucum JEAJEAMADOU" (Aublet 1775). Subsequent authors have attributed this etymology to a Galibi or Sinemari vernacular name for V. sebifera in French Guiana (Rodrigues 1972(Rodrigues , 1980Cardona et al. 2010;Mari Mut 2018). However, others believe that the name is derived from the Latin viriola (bracelet), in reference to the aril that surrounds the seed (e.g. Flores 2010).
There is a long taxonomic history to the American species of Myristicaceae. This began with George Bentham (1853), who was the first to publish on these species in "Notes on the American Species of Myristica". In this treatment, he recognises 12 species of Myristica (nine of them new) and places these in three different groups, characterised by differences in the androecium (i.e. number of anthers and shape, thickness and length of the column) and leaf venation (i.e. number of veins and their degree of separation). Alphonse de Candolle published the second and broadest treatment of Myristicaceae, including species from both the New and Old World (de Candolle 1856). He recognised ca. 84 species, which he divided into 13 sections in a single genus, Myristica, which included two sections that contain species today recognised as Virola: Virola (Sect. III/ 8 spp.) and Sychoneura (Sect. IV/10 spp.). In this work, he described 12 new taxa that now belong to Virola. The British botanist William Bot-ting Hemsley was the earliest author to publish an account of the Mesoamerican species of Myristicaceae in "Biologia centrali-americana" (Hemsley 1882). In this work, three species are newly described: Myristica guatemalensis Hemsl., M. mexicana Hemsl.
Virola has been the subject of limited phylogenetic analysis. The first molecular phylogenetic study of Virola-and the only to date-demonstrated that the genus is divided into two large subclades, informally called "Multinervae" and "Sebiferae" (Steeves 2011). Morphologically, Multinervae is composed of tall emergent canopy trees that typically have more numerous secondary leaf veins, thicker pericaps and arils that are lacinate throughout their length, compared to the sub-canopy to canopy species of Sebiferae that have fewer secondary veins, thin pericarps and arils that are only lacinate for half of their length. This phylogeny, or future improved phylogenetic hypotheses, has the potential to provide information for future infrageneric reclassification of Virola.

Materials and methods
This work was undertaken as part of the "Flora Mesoamericana" project (http://www. tropicos.org/Project/FM) by the first author. However, our definition of "Mesoamerica" is different than that used in the Flora; here, it refers to the portion of Central America and southern Mexico from the Isthmus of Tehuantepec in Mexico in the north to the Panamanian-Colombian border in the south.
Species descriptions are based primarily on herbarium specimens, though observations during fieldwork by the first and second authors, especially in the Osa Peninsula of Costa Rica, were also important. If necessary and material permitted, flowers from herbarium specimens were rehydrated before measurement. A ruler was used to measure leaves and inflorescences; a digital Neiko caliper was used to measure fruits and seeds, as well as the thickness of the twigs, petioles and peduncles; and, finally, flowers, trichomes and thickness of the pericarp were measured with a micrometer calibration tool (1div = 1mm) under a dissecting stereoscope (Bausch & Lomb).
The preliminary conservation status of each new species was assessed using quantitative criteria recommended by the IUCN Red List (IUCN Standards and Petitions Subcommittee 2014). Georeferenced specimen data were used to determine the area of occupancy (AOO) and the extent of occurrence (EOO), which in turn were used to determine threat status. All analyses were performed in the R package conR (Dauby et al. 2017). When the recommendation differed between AOO and EOO assessments for a given species, we opted to conservatively recommend the more vulnerable status, following Knapp (2013).
Specimens cited are listed first by country in a north to south sequence. Within country, specimens are listed alphabetically by major division and then alphabetically by province or department and, finally, in alphabetical order by the collector's surname. When the coordinates and/or elevation were not included on the herbarium label, but were present in the TROPICOS database, the values from TROPICOS are included. Dot-distribution maps were compiled from studied specimens and generated with SimpleMappr (Shorthouse 2010).
In the nomenclatural section for each new species, we cite both accession numbers and barcodes when present. Barcodes are included in square brackets and follow the format of a series of numbers preceded by a herbarium acronym (e.g. [GH00039891]); all other numbers correspond to accession numbers.
Knoweldge of the group was improved by numerous field trips since 1991 throughout Costa Rica, especially in the Osa Peninsula and La Selva Biological Station, by the second author and, more recently, by the first author in the same region.    Description. Tree 13-30 m × 10.4-50 cm DBH; bark sometimes described as smooth and reddish or dark brown. Exudate sometimes described as abundant and reddish or watery, but without specifying from where or red in the trunk. Twigs 0.16-0.22 cm thick, terete to slightly flattened laterally, puberulent, trichomes stellate to irregularly stellate, ferruginous. Leaves: petiole 0.5-1.4 × 0.13-0.24 cm, slightly canaliculate, tomentose, the trichomes stellate to irregularly stellate; leaf blades 16.2-29.2 × 3.2-7.3 cm, oblong-elliptic or rarely elliptic; adaxial surface of mature leaves olive or light brown (sometimes shining) when dry, glabrous or with scattered stellate trichomes, the surface smooth; abaxial surface pale brown to whitish when dry, densely but inconspicuously pubescent, trichomes stellate, sessile, the central part of the trichome reddish and contrasting in colour with the hyaline branches to reddishclear, with 4-10 branches, the branches ± 0.01-0.05 mm long, persistent; lateral veins 15-20 per side, 4-5 veins per 5 cm, 1.2-1.8 cm apart, the same colour as the adaxial surface or slightly transparent, on adaxial side flat to sunken, on abaxial side slightly elevated, arcuate-ascending, slightly anastomosing near the margin and without forming a very marked intramarginal vein; tertiary veins barely visible on both sides; midvein adaxially slightly elevated (sometimes flat, distally), abaxially raised, rounded to somewhat triangular, tomentose to glabrate; base acute or obtuse to rounded, not revolute, flat; margin flat; apex acuminate or rarely rounded. Staminate inflorescences 3.5-5.5 cm long, axillary, axes flattened, tomentose, with trichomes irregularly stellate, ferruginous; peduncle 1.2-1.9 × ca. 0.1 cm long; bracts not seen; terminal fascicles dense, with 15-40+ flowers. Staminate flowers with the pedicel 0.3-1.2 mm long; receptacle 1.2-2 mm wide; perianth 2-2.8 mm long, infundibuliform, yellow when fresh, connate by 1.1-1.7 (-2.3) mm long, external surface pubescent, with brown trichomes, internal surface glabrous or with few trichomes close to the margin of the lobes; lobes 3 (4), 0.8-1.5 × 0.6-0.9 (-1.2) mm; stamens 3, the filament column 0.5-0.6 mm long, glabrous, straight, thin, sometimes slightly thickened at the base, not constricted at the apex; anthers 0.6-0.9 mm long; apiculus 0.06-0.1 mm long, acute to apiculate, connate. Pistillate inflorescences and pistillate flowers not seen. Infructescence 2.5-7.5 cm long, with 2-13 fruits, peduncle 2-3.5 × ca. 0.47 cm. Fruits 2.7-3.5 × 1.5-2.5 cm, usually ellipsoid or rarely ovoid, stipitate, densely tomentose, Figure 6. Virola allenii A treetop B branching C lower trunk and buttress D branch with leaves, showing the adaxial surface E leaf blades on abaxial surface F exudate of the trunk G leaf blade surfaces, adaxial (above) and abaxial (below) H petiole and leaf base I mature fruit, inset left and arrow showing the galls on leaves and branches, respectively J fruits K fruit close-up. Photos by Reinaldo Aguilar. the trichomes dendritic, ferruginous and falling very easily to the touch (as dust), the surface rugose or smooth when dry, the line of dehiscence usually carinate, but not very conspicuous, the base obtuse, the apex acute to obtuse, green or golden and ferruginous by the pubescence when fresh; pericarp 3.2-3.8 mm thick; pedicel 0.4-0.7 cm long; seed ca. 2.5 × 1.3 cm, the testa when dry whitish-greyish, markedly grooved; aril usually described as red when fresh, pale brown when dry, membranaceous, the texture dry and thin, laciniate in narrow bands distally. Germination epigeal, seedling cryptocotylar, the first pair of leaves (sub)opposite (Ley López and Chacón Madrigal 2017; as V. macrocarpa).

Key to the species of Virola in Mesoamerica
Distinctive characters. Virola allenii is recognised by its narrow leaf blades with lateral veins that are well separated (Fig. 8A) with a whitish abaxial surface and covered with stellate, sessile trichomes with the central portion of the trichome reddish and contrasting in colour with the hyaline branches to reddish-clear (Figs 3A, 6F); the staminate flowers with the lobes of the perianth almost glabrous on the inner surface, the column of filaments straight and not constricted at the apex, anthers that are usually longer (0.6-0.9 mm long) than the column of the filaments (0.5-0.6 mm long) and an apiculus that is 0.06-0.1 mm long. It is also distinctive for its large, usually ellipsoid fruits (Figs 4A, 6I-K) with thick pericarp that are green when ripe and covered by ferruginous trichomes that fall very easily to the touch (Fig. 6K inset); and seeds with the testa markedly ribbed.
Etymology. The specific epithet honours the collector of the type specimen, Paul H. Allen (1911Allen ( -1963, who was probably the first person to collect this species 67 years ago (P. H. Allen 5763; 10 Jan 1951). During his five-year residency in Palmar Norte, Puntarenas, Costa Rica (Grayum et al. 2004), Allen made important collections and publications in this region (e.g. The Rain Forests of Golfo Dulce, Allen 1956). Distribution. Virola allenii is known only from Costa Rica (Puntarenas and San José) (Fig. 9A). It is found on the Pacific slope, at 0-350 (-1350) m elevation.
Preliminary conservation status. Virola allenii is Vulnerable following IUCN critera B1a and B2a. Justifying its status, it is known from seven localities and has an EOO of 3,424 km 2 and an AOO of 40 km 2 . Specimens have been collected regularly since the 1990s during botanical expeditions in the Osa Peninsula of Costa Rica, though only 22 specimens have been verified.
Common names. None recorded. Phenology. Flowering of Virola allenii has been recorded in January, March, April and December. Fruits have been observed in January, August to October. Pistillate flowers were not seen in the studied material.
Field characters. The bark is described as brown and smooth or as peeling in small pieces, sometimes with a strong, spicy scent. Twigs and leaves often have galls (e.g. Fig.  6I). Leaf blades are adaxially lustrous and abaxially whitish. Staminate flowers are yellow and fragrant. Fruits, which are ca. 5 × 3.2 cm when fresh, are green at maturity and covered with brown trichomes that fall very easily when touched and have a pericarp that is ca. 6 mm thick. The aril of mature fruits is red and white in immature fruits. Seeds have a white testa.  Discussion. Virola allenii is most similar to V. macrocarpa, a species from montane forests at 1100 m elevation in the Andes of Colombia (Boyacá) and this name has been previously applied to the species described here (e.g. Jiménez 2007;Cornejo et al. 2012;Aguilar et al. 2017 onwards). It is differentiated from V. macrocarpa by the characteristics presented in the diagnosis and in Table 2.
The comparison presented hereafter for Virola macrocarpa (Fig. 7C) is strictly based on the protologue (except for the pericarp thickness), from which we have been able to study two physical duplicates deposited at MO (A. Lawrance 675, MO-2 sheets!, fr [ Fig. 4D]) and the images at A! (st), F! (2-sheets, fr), G! (2-sheets, st.), K! (st), S (fr) and US! (2-sheets, fruits likely in the packet, but not seen). We confirmed all measurements from the protologue on Lawrance 675 (MO) and found them consistent with the exception of pericarp thickness; while 2-4 mm was stated in the protologue, our measurements ranged from 1.8-3 mm, which we present in Table 2 below. In our estimation, all other observed South American specimens, annotated with this name, represent an amalgamation of different identities (D. Santamaría-Aguilar, in prep.).   Based on a number of features listed in the diagnosis of V. alleni, including colour of the abaxial leaf blade, sessile trichomes, degree of separation of lateral veins and the length of the anther apiculus, V. allenii is similar to V. calophylla (Fig. 10A, B) and V. calophylloidea from South America; the latter was recently included as a synonym of V. calophylla (see notes below). Furthermore, Virola allenii also shares narrow oblong-elliptic leaf blades (3.2-8 cm broad) and short inflorescences with V. calophylloidea. However, it is distinguished from both species by the filament column that is constricted at or towards the apex (vs. not constricted in V. allenii) and the tendency towards short anthers (0.4-0.7 mm vs. 0.6-0.9 mm long). Additional differences amongst these three species are presented in the Table 2.
In Mesoamerica, Virola allenii can be confused with V. amistadensis and V. otobifolia (which are formally described as new below). All these species have lateral veins that are well spaced (Fig. 8A, B, M) and an abaxial leaf surface that is usually whitish with sessile stellate trichomes; for differences between these species see Table 3. Notes. The holotype, deposited at Field Museum (F), represents two sheets with hand written annotation ("Sheet 1 of 2," "Sheet 2 of 2"), which suggests that they represent a multi-sheet specimen of the same plant (ICN Art. 8.3) (Turland et al. 2018).
The specimen B. Hammel et al. 24041 (CR!; fr) from 1370 m elevation in the Tarrazú region of San José province (Costa Rica) differs from other members of this new species by its more rounded and pubescent fruits and its occurrence at a higher elevation than other specimens. It is included here with some reservation. This specimen is most similar to E. Alfaro 492 (CR!, LSU!, MO!; ♂ fl), which was also collected in montane forests on the Pacific slope of the Cordillera de Talamanca (1240 m elevation). E. Alfaro 492 differs from the rest of V. alleni in its larger staminate perianth (ca. 3.8-4 mm vs. 2-2.8 mm long) that is fleshy with dense pubescence on the entire adaxial surface (vs. glabrous or with sparse trichomes close to the margin of the lobes in V. allenii) and the column of the filaments (ca. 0.5-0.7 mm vs. 0.5-0.6 mm long) that is shorter than the anthers (ca. 1.2 mm vs. 0.6-0.9 mm long). This specimen was also discussed by Jiménez (2007) as V. macrocarpa.
Distinctive characters. Virola amistadensis is recognised by its elliptical to widely elliptical leaf blades with lateral veins that are well separated (Fig. 8B) and abaxially covered with stellate, sessile trichomes (Fig. 3B) and a dense layer of squamiform structures that are hyaline in colour; the small staminate flowers with a thickened filament column that is markedly constricted at the apex and shorter (0.2-0.4 mm long) than the anthers (0.6-0.7 mm), which are apiculate at the apex; and the ovoid to globose fruits ( Fig. 4B) with relatively thin pericarp (1-2 mm thick) (Figs 7B inset, 11D) that are covered by a dense, but inconspicuous layer of ferruginous trichomes. It is also distinctive for being a small tree (4-13 m tall) and its preference for premontane forests.
Etymology. The specific epithet, amistadensis, refers to Parque Internacional La Amistad, a UNESCO World Heritage Site, shared between Costa Rica and Panama where the holotype and some of the paratypes of this species were collected. Distribution. Virola amistadensis is known from Costa Rica (Limón) and Panama (Bocas del Toro and Veraguas; Fig. 9B). It is found on the Caribbean slope at 650-1200 m elevation.
Preliminary conservation status. Virola amistadensis is Endangered following IUCN criteria B1a and B2a. This species is known from 4 localities and has an EOO of 2,573 km 2 and an AOO of only 28 km 2 . Only nine specimens were verified in this study. While it occurs in protected areas, its montane habitat is particularly prone to habitat disturbance.
Common names. None recorded. Phenology. Flowering of V. amistadensis has been recorded in April, June and July and fruiting in January to March, May, June and December.
Field characters. Plants are trees that are 4-13 m tall. Flowers have a yellowbrown perianth and brown fruits.
Discussion. Herbarium specimens of this new species usually have been identified as Virola calophylla (Figs 10A, B) or V. macrocarpa (Fig. 7C), probably because its leaves are abaxially whitish and covered with stellate, sessile trichomes with a reddish-clear to reddish centre that contrasts in colour with the hyaline branches and lateral veins that are well-spaced. Differences between V. amistadensis and V. calophylla are enumerated in the diagnosis, while those separating it from V. macrocarpa are listed in Table 4.
In Mesoamerica, Virola amistadensis is similar to V. allenii (Figs 6, 7A) and V. otobifolia (Figs 7D, 23A-E), two species from the lowland wet forest of Costa Rica and Panama. Their similarities include the characteristics of the leaf blades mentioned above. Their differences are summarised in Table 3.
Notes. Specimens from Veraguas Province (Panama), have smaller leaf blades and lateral veins that are more deeply sunken on the adaxial surface than the specimens from Limón and Bocas del Toro provinces.
Distinctive characters. Virola chrysocarpa is distinguishable for its leaf blades with pubescent adaxial surfaces that are rough to the touch in mature leaves (at least in herbarium specimens) and abaxial surfaces that are hirsute to hirsutulous with trichomes with long branches (0.2-0.6 mm long) (Fig. 3C), numerous lateral veins (28-32 per side), tertiary veins that are usually conspicuous on both surfaces (Figs 3C, 8C) and with a base that is usually markedly cordate; staminate inflorescences that are littlebranched ( Fig. 14D) with flowers with filament columns that are much longer (1.3-1.5 mm) than the anthers (0.6 mm); and fruits that are acute to apiculate at the apex (Fig. 14E, F, H). Additionally, as far as we are aware, this is the only Mesoamerica species that is completely deciduous (i.e. all leaves fall off the tree) (Fig. 13A).
Etymology. The specific epithet, chrysocarpa, is derived from the Greek chryso (gold) and carpo (fruit). This is in reference to its common name, "fruta dorada" (golden fruit), which is used by locals of the Osa Peninsula, Costa Rica, where this species is frequent.
Distribution. Virola chrysocarpa is known from Costa Rica (Puntarenas and San José) and Panama (Chiriquí) (Fig. 9C). It is found in the Pacific slope at 0-700 m in elevation.
Preliminary conservation status. Possible Near Threatened: This species has a small estimated AOO (60 km 2 ), though a relatively large estimated EOO of 5,334 km 2 . Its eighteen known specimens represent eleven localities. This limited number of specimens warrants a Possible NT status, though additional collection efforts may demonstrate the lack of conservation threat for this poorly known species.
Common names. Costa Rica: fruta dorada. Panama: bogamani. Phenology. Herbarium specimens of flowering Virola chrysocarpa have been collected in December to March and fruiting specimens from March to June. Herbarium specimens with pistillate flowers were not observed. In the Osa Peninsula, leaves fall completely during the dry season, which occurs in November to February (Allen 1956;Quesada Quesada et al. 1997;and R. Aguilar pers. obs., 2015and R. Aguilar pers. obs., , 2017and R. Aguilar pers. obs., , 2018and R. Aguilar pers. obs., , 2019. A study of vegetative, flowering and fruiting phenology has been published by Lobo et al. (2008;as V. koschnyi) in the Osa Peninsula and Golfo Dulce, Costa Rica. In this study, flowering was documented in January and February, when the canopy was deciduous. Fruiting occurred in the rainy season from June to August.
Field characters. Plants are large trees with boles that are straight and do not begin to branch until they reach a great height, with buttresses, up to 1.6-2.5 m tall. Bark is sometimes described as finely fissured. The new leaves are lime green in colour. Twigs, petioles and leaf blades on both surfaces (especially the youngest ones) are covered with golden, brown-reddish to rusty-red trichomes. Flowers have yellow or yellow-cream perianth and anthers. Mature fruits are yellow, orange or ferruginous (possibly due to their indumentum). Seeds are brown or blackish and covered with a red to scarlet aril.
Discussion. Virola chrysocarpa resembles a morphological group of species from South America that includes V. caducifolia W. A. Rodrigues, V. decorticans Ducke, V. guggenheimii W. A. Rodrigues, V. multicostata Ducke, V. multinervia Ducke, V. polyneura W. A. Rodrigues and V. rugulosa (Spruce) Warb. These species are characterised by having leaves that are evidently pubescent, some with dendritic to irregularly dendritic pediculate trichomes on the abaxial surface and leaf blades with numerous, conspicuous and comparatively dense lateral veins; staminate flowers with anthers that are subequal to or shorter than the filament column; and fruits with thick pericarp. Additionally, these species tend to be large, sometimes deciduous trees with cordate leaf bases and staminate flowers with the anthers that are obtuse at the apex. Table 5 presents the differences between these species and V. chrysocarpa.
Notes. The illustration presented in Manual de Plantas de Costa Rica (Jiménez 2007) as V. koschnyi is a mix of these two species. The branch with leaves and inflorescences (and, most likely, the trichomes) are based on material that represents V.    Distinctive characters. Virola elongata it is recognized by its relatively small leaf blades [10.8-18.5 × 2.7-3.7 (-5.1) cm] with a sparsely pubescent abaxial surface with stellate to dendritic-stellate trichomes that are usually sessile (Fig. 3D) and few lateral veins (9-14 per side) that are well separated [(0.8-) 1-1.7 cm apart] (Fig. 8D); staminate inflorescences with thin axes and flowers with short filament columns (0.3-0.4 mm long), anthers that are apiculate at the apex (apicula 0.1-0.2 mm long) and more than twice the length of the column (0.6-0.9 mm long); and its small fruits (1.6-1.9 × 0.9-1.1 cm) (Fig. 4L) that are green when ripe (drying light brown in herbarium specimens with the surface blistering to rough), inconspicuously to sparsely pubescent (densely pubescent when young) with trichomes that are dendritic and brown to ferruginous in colour (when present), pericarp that is 0.5-0.7 mm thick and an aril that is thin and laciniate in narrow bands. Distribution. In Mesoamerica, Virola elongata is only known in Panama (Colón, Panamá and San Blas) (Fig. 9D). It has been recorded from between 50-450 m elevation.
Distinctive characters. Virola fosteri is recognised by its small leaf blades (7.8-12 × 1.4-2.7 cm) and fruits (1.5-2.3 × 1.2-1.8 cm) (Figs 4I and 16F-H), as well the stellate, sessile trichomes on the abaxial surface of the leaf (Fig. 3E). It is also distinguished by its leaf blades that have numerous lateral veins (16-24 per side) that are prominent on adaxial surface, the revolute leaf margin and the base (Fig. 16C), the midvein that is laterally compressed adaxially and sometimes resembling a cutting edge; staminate flowers with a filament column (0.9-1.3 mm long) that is longer than the anthers (0.6-0.9 mm long); and its thick pericarp (1.5-2.5 mm).
Etymology. The specific epithet honours one of its collectors, Robin B. Foster (1945-), ecologist and botanist at Field Museum in Chicago (F) who pioneered the cataloguing of the flora of Barro Colorado Island (BCI) in Panama, where V. fosteri occurs. Robin noted on one of his collections (R. B. Foster 2931) that it could represent a new species. In addition, on the same herbarium sheet, he observed one of the taxonomic characters that we here use to distinguish this as a new species: "Leaves are consistently small throughout the tree and on juvenile plants." Distribution. Virola fosteri is known from Costa Rica (Limón) and Panama (Bocas del Toro, Colón, Panamá, San Blas and Veraguas) (Fig. 9E). It is found on the Caribbean slope from 0-350 (-800) m elevation.
Preliminary conservation status. Virola fosteri is Vulnerable following IUCN criterion B2a. While the EOO for this species is large (25,645 km 2 ), the small AOO (40 km 2 ) with only eight known localities warrants its conservative status.
Common names. Panama: bogamani, fruta dorada. Phenology. Flowering of Virola fosteri has been recorded in January to April, June and October and production of fruits in January to April.
Field characters. Plants are large trees with tall buttresses. Bark exudes reddish watery exudate when damaged. Their small leaves are white or grey below. Flowers have pale orange or yellow perianth. The mature fruit is yellowish or golden brown with a red aril and brown seed.
It is also comparable to V. micrantha A. C. Sm. from Colombia due to the similar size of the leaf blades, which also have sessile stellate trichomes on the abaxial surface and short staminate inflorescences. Virola micrantha is a name apparently

R. L. Liesner 6778 [MO!] and J. Velazco 851 [MO!]).
Virola fosteri differs from this species by its attenuate leaf base and acute to acuminate apex (vs. obtuse on both sides, also mucronulate at the apex), longer perianth of staminate flowers (2-2.5 [-3] mm vs. ca. 1 mm long) and longer anthers (0.6-0.9 mm vs. ca. 0.3 mm long) (Smith 1953 Ducke (Brazil) are other species with similarly-sized leaves with revolute margins, traits shared with V. fosteri. Additionally, with V. coelhoi, which it is more likely to be confused, V. fosteri shares overall leaf shape and the type of trichomes on the abaxial surface (i.e. stellate, sessile and yellowish). The new species is distinguished from V. coelhoi by the abaxial surface that is pale brown to reddish-brown when dry and puberulent (vs. abaxial surface yellowish, very densely pubescent), staminate inflorescences with small bracts (2-5 × ca. 2.5 mm vs. 2.5-9 × 4-6 mm), and staminate flowers with longer filament columns [0.9-1.3 mm vs. (0.3-) 0.6-0.7 long] and anthers (0.6-0.9 mm vs. 0.4-0.5 mm long). It can be differentiated from V. parvifolia by its leaf blades and inflorescences that are glabrous or nearly so (vs. pubescent in V. fosteri) (Ducke 1936); additionally, Ducke (1936) mentions that V. parvifolia has numerous small granules or tubercles on the branches, leaf blades and peduncles that are lacking in V. fosteri. Jiménez (2007; as Virola sp. B) mentions that the new species is similar to Virola pavonis (A. DC.) A. C. Sm. from South America. This is probably because both species have sessile and stellate trichomes on the abaxial surface of the leaf, sometimes a similar leaf size, a similar number of lateral veins (though there is a tendency towards higher numbers of veins in V. pavonis) and the length of the filament column and anthers. However, the new species differs in its shorter staminate inflorescences [2.5-5.3 cm vs.
In Mesoamerica, other species with leaf blades that are covered with stellate and sessile trichomes on the abaxial surface (Fig. 3E, F, L) and a filament column that is longer than the anthers are V. guatemalensis and V. nobilis. However, these two species have larger leaf blades (12.3-17.2 [-27.5] cm vs. 7.8-12 cm long) and fruits ([2.1-] 2.3-2.7 [-3.1] cm vs. 1.5-2.3 cm long). Similarities with V. nobilis, specifically, include their distribution pattern (at least in Panama), leaf blades with more lateral veins per side (20-30 [25-32] vs. 16-24) that are markedly elevated abaxially (vs. flat to slightly elevated), the leaf margin and base that are usually not revolute (vs. revolute) and fruits that generally have thick pericarp (2.5-3.5 mm vs. 1.5-2.5 mm).
Notes. The species referred to as Virola sp. B in the Manual de Plantas de Costa Rica (Jiménez 2007) and as V. multiflora (G. de Nevers 7608, MO!) in the Catálogo de las plantas vasculares de Panamá (Correa et al. 2004)   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.
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, greenyellowish 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) 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.
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.  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).  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.

Virola laevigata Standl.
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. Zamora & 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)  Distinctive characters. Virola megacarpa can be recognised by its large and oblong leaf blades (20.3-37 × 7-13 cm) with numerous [(32-) 40-50 per side], dense lateral veins and a densely pubescent abaxial surface with dark brown to ferruginous dendritic trichomes (Fig. 3I). It is also the species with the largest fruits (4-5.7 × 2-2.9 cm) in the region and likely the genus; these are also densely pubescent with an acuminate to rostrate apex (Fig. 4P) and thick pericarp (3-6 mm). Distribution. Virola megacarpa is only known from Panama (Colón, Panamá, San Blas and Veraguas) (Fig. 18C) from 50-550 m elevation. This species is attributed to Colombia in Cogollo (2011), based on the specimen J. Brand 1252 (JAUM!*; fr). It is also mentioned as occurring in Colombia in Gradstein (2016) and Ulloa Ulloa et al. (2017), though these references do not mention a voucher specimen (likely they refer back to the same specimen cited by Cogollo [2011]). The first author has seen a digital image of this specimen (J. Brand 1252, JAUM) and it appears that the leaf undersurface and fruits are scarcely pubescent, as well as smaller. This specimen clearly corresponds to a species of the group Surinamenses sensu Smith and Wodehouse (1938) and not V. megacarpa. For that reason, V. megacarpa is considered restricted to Mesoamerica.
Common names. None recorded.
Phenology. The only observed herbarium specimen with flowers (these staminate) was collected in August. Fruits were collected in February and March and August to November.
Field characters. Plants are trees 12-30 m tall and 21.5-53 cm DBH. Damaged bark releases exudate that is red or that oxidises reddish-brown. Flowers have pale yellow perianth. Fruits are densely pubescent with brown trichomes and a red aril.
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 km 2 ) and AOO (104 km 2 ) 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 (1857Pittier ( -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)  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  ] 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. 11. Virola multiflora (Standl.) A. C. Sm. Fig. 22E Distinctive characters. Virola multiflora is recognised by its usually small and narrow leaf blades [5.5-15.5 × 1.5-3.6 (-4.8) cm] with an inconspicuously pubescent abaxial surface with stellate and sessile trichomes (Fig. 3K), lateral veins that are not very prominent (Fig. 8K) and long and thin petioles [0.5-0.9 (-1.1) cm long]; staminate flowers with the filament column longer (0.7-1 mm long) than the anthers (0.3-0.6 mm long); and for its small fruits [1.3-1.9 × 0.9-1.2 (-1.4) cm] (Fig. 4J) with the pericarp 0.7-1 mm thick. Distribution. Virola multiflora is known from Belize (Cayo, Stann Creek and Toledo), Honduras (Gracias a Dios), Nicaragua (Atlántico Norte, Atlántico Sur, Jinotega, Matagalpa and Río San Juan) and Costa Rica (Alajuela, Cartago, Heredia and Limón) (Fig. 18E). Throughout its range in Mesoamerica, it is only known from the Caribbean slope. It has been recorded from between 0-650 (-1400) m elevation.
While V. multiflora is not documented from Guatemala in herbaria, Standley and Steyermark (1946) postulated that the species is to be expected to occur in Izabal (Guatemala) and Jiménez (2007) attributes it to that country. Conversely, while it is mentioned as occurring in Peru by Jiménez (2007), specimens could not be located. The Peruvian specimen attributed to V. multiflora (R. Vásquez & C. Grández 17507, MO!;fr) in Vásquez M. et al. (2018) corresponds to a species related to V. multinervia. Finally, the report presented in the Nuevo Catálogo de la Flora Vascular de Venezuela (Rodrigues 2008) is errorenous. Based on this evidence, V. multiflora is considered as restricted to Mesoamerica. Phenology. Flowering of Virola multiflora has been recorded in March to August, with a noted peak in July; just one herbarium specimen with pistillate flowers was seen and it is from Nicaragua. Fruits were collected in December through April.
Field characters. Plants are trees between 6-30 m tall and 17-35 cm DBH. Bark exudes latex red. The leaf blades are sometimes whitish abaxially. The flowers usually have yellow, golden or orange perianth. The mature fruit is yellow or orange with a red aril. Notes. We were not able to locate any Panamanian specimens of V. multiflora: all fertile Panamanian specimens identified as V. multiflora that the first author has studied are actually V. fosteri. However, V. multiflora is to be expected in Panama because it occurs in physiognomically similar forests in Costa Rica near the border. The similarities and differences between V. multiflora and V. fosteri (which is formally described above) are discussed under the latter species.   4N) with thick pericarp (2.3-3.5 mm [2.5-4.2 mm]) and aril. Distribution. Virola nobilis is known from the Pacific slope of Costa Rica (Puntarenas, San José) and the Caribbean slope of Panama (Bocas del Toro, Colón, Panamá, San Blas). It is recorded from 0-400 (-1300) m elevation (Fig. 18F). Jiménez (2007) reported this species (as V. surinamensis) from just one locality (Chitaría) on the Caribbean slope in Costa Rica; however, we have not examined the specimen (Poveda 144, CR).
Phenology. Flowers have been collected in January to April, July, August, November and December and fruits in almost all months except October.
Field characters. Plants are large trees between (5-) 15-40 m tall and 20-70 cm DBH. The trunk is straight, usually with conspicuous buttresses and does not begin to branch until it reaches a great height. The bark is reddish and releases red exudate when damaged. The leaves are whitish with inconspicuous trichomes on the abaxial surface. Flowers have yellow perianth. The mature fruit is yellow to orange with a red aril.
Distinctive characters. Virola otobifolia is recognised by its large leaf blades with well-spaced lateral veins (Fig. 8M) and a whitish abaxial surface that is covered with stellate, sessile trichomes (Fig. 3M); staminate flowers that have a filament column that is straight and very thickened throughout its length, except where it is constricted at the apex and anthers that are almost the same length as the filament column; and its large fruits with thick pericarp that are densely tomentose and with a conspicuously carinate line of dehiscence (Fig. 4C). Etymology. The specific epithet refers to the similarity of the leaf blades with Otoba, another member of Myristicaceae. This epithet was, in part, inspired by the fact that some of the first collections of this new species were initially confused with this genus (as Dialyanthera Warb.; e.g. A. Gentry & S. Mori 14199, MO). Distribution. Virola otobifolia is only known in Panama (Colón, Panamá and San Blas), where it is found on the Caribbean slope (Fig. 27A). It has been recorded between 50-850 m elevation.  Preliminary conservation status. Virola obtobifolia is Endangered following IUCN criteria B1a and B2a. Justifying this status, it has both a small EOO (3,269 km 2 ) and AOO (36 km 2 ) and is known from only five localities.
Common names. Panama: velario, miguelario; cuinur burwi, putmas (Kuna). Phenology. Virola otobifolia has been recorded with flowers and fruits in February to April and one collection with fruits was made in October.
Field characters. Exudate is slow to appear and is watery and red-black. Leaf blades are whitish abaxially. Flowers have yellow perianth. Fruits are green, but appearing brown (possibly due to pubescence).
Discussion. Specimens of Virola otobifolia have been confused with and identified as V. macrocarpa (e.g. Correa et al. 2004;Condit et al. 2011) (Figs 4D, 7C), probably due to the similar size of the fruits and the leaf morphology (i.e. large discolorous leaf blades, with stellate, sessile trichomes with a reddish centre, and lateral veins that are well-separated). However, it is easily distinguished by the leaf width, pubescence on abaxial surface of the leaves, the number of lateral veins, fruit morphology and habitat; a comparative table of these two species is presented in Table 7.
Two similar species from Mesoamerica that resemble Virola otobifolia are: V. allenii (Figs 6, 7A), which ocurrs in the lowland wet forest on the Pacific slope of Costa Rica and V. amistadensis (Fig. 7B) from montane forests of Costa Rica and Panama on the Caribbean slope. Morphological comparison between these species is presented in Table 3.
Notes. The holotype, deposited at Missouri Botanical Garden (MO), represents a single collection mounted on two sheets that are clearly labelled ("Sheet 1 of 2," "Sheet 2 of 2) as being parts of the same specimen (ICN Art.  Distinctive characters. Virola sebifera is recognised by its dense pubescence of pediculate and ferruginous trichomes (Fig. 3N) that cover almost all parts of the plant; leaf blades with few lateral veins (9-17 per side), well-spaced between them [3-4 (-5) veins per 5 cm, 1.3-2.7 cm apart] (Fig. 8N); the long staminate inflorescences that Table 7. Morphological comparison between Virola otobifolia and V. macrocarpa.
2.7-3.3 × 2-2.3 cm * ; tomentellous, with cauducous trichomes that are dust-like when they fall; the surface smooth when dry, the line of dehiscence slightly carinate. Pericarp (2.7-) 3-4.7 mm thick 1.8-3 mm thick Habitat Lowland or premontane rain forest, Central Panama (Panamá, San Blas) at 50-850 m elevation  the highest species diversity of Virola, with 10 and 11 species, respectively. We clarified the application of some names, the resurrection of others and the distribution of some species previously considered to occur in Mesoamerica (e.g. South American V. macrocarpa) or South America (e.g. the Mesoamerican V. multiflora). This has resulted in a much modified taxonomy for Virola, summarised here: Mesoamerican collections previously identified as V. calophylla and/or V. macrocarpa are here interpreted as three new species: V. alleni, V. amistadensis and V. otobifolia, while Mesoamerican specimens previously determined as V. surinamensis are treated as V. nobilis. As a result of this, V. calophylla and V. macrocarpa are now restricted to South America and V. surinamensis to South America and the Antilles. Additionally, two new species are identified within what was previously V. guatemalensis: montane collections from the southern range now belong to V. montana and we have resurrected V. laevigata, previously considered a synonym. Under our concept, Virola guatemalensis has a restricted range, found only in northern Mesoamerica. Finally, the majority of V. koschnyi specimens from the Pacific coast of Costa Rica and Panama correspond to V. chrysocarpa, while most of Panama's collections, identified as V. multiflora, are now considered V. fosteri. These contributions came to light via detailed comparisons across the complete range of these species thanks to hundreds of collections made by numerous botanists and herbaria who care, maintain and provide access to the specimens, including making them available online.
H. van der Werff for their help provided during herbarium visits at MO; also, to MO for providing substantial material as loans and donations. He is also grateful to Jennifer Kluse (LSU) for her help with with loans and herbarium supplies and to the Shirley C. Tucker Herbarium at Louisiana State University for providing space and access to facilities. Last, but no less important, we appreciate the work of all the botanists and institutions -past and present -who contributed the many herbarium specimens studied and ensure their longterm care.