﻿New Species of Virola (Myristicaceae) from South America

﻿Abstract With about 70 species Virola, is the largest genus of Myristicaceae in the Neotropics, the genus ranked in the top ten genera of abundance across Amazonia. Ten new species are proposed in this striking genus, which are described based on morphology, and are illustrated. The new species were discovered thanks to herbarium specimens collected mainly in the 1980s and 1990s when field documentations were more active. The new species come from Colombia (V.calimensissp. nov., V.cogolloisp. nov., V.excisasp. nov., V.tuckeraesp. nov.), Ecuador (V.alvaropereziisp. nov., V.bombuscaroensissp. nov., V.calimensis, V.excisa, V.yasunianasp. nov.), Peru (V.aguarunanasp. nov., V.cumalasp. nov., V.excisa, V.parkerisp. nov.), and Brazil (V.excisa, V.yasuniana). Additionally, a lectotype is designated for V.macrocarpa, a name used to identify some specimens of the new species here described, and V.kwatae is reported for the first time for Brazil. We provide a comparation table between the new species and the species that is morphologically close to it, a preliminary list of species for the genus, and notes of how the new species were treated in floras, checklists, or collections that need more study and herbarium specimens.


Introduction
Virola Aubl. (Myristicaceae) is an ecologically and economically important genus of trees in the nutmeg family, Myristicaceae within the order Magnoliales (APG IV 2016). It is the fourth largest genus in Myristicaceae, a pantropical family with 21 genera and nearly 500 species; only three Asian genera, Horsfieldia Willd., Knema Lour., and Myristica Gronov., are larger. Virola occurs across the wet Neotropics, where it is distributed from Mexico to southern Brazil and in the West Indies [i.e. V. surinamensis (Rol. ex Rottb.) Warb.], though it is notably absent from El Salvador (Smith and Wodehouse 1938;Rodrigues 1980;Acevedo-Rodríguez and Strong 2012;Santamaría-Aguilar et al. 2019). Across Virola's ca. 70 species (including those described here), ca. 55 occur in South America (Fig. 1) and 15 in Central America, with just two widespread species [i.e., V. elongata (Benth.) Warb., and V. sebifera Aubl.] occurring in both (Table 1). The species richness of Virola is highest in Amazonia, where it is ranked in the top ten most abundant genera with 35 species (Cardoso et al. 2017). Like other genera that are most species-rich and abundant in Amazonian forests including Protium Burm. f. (Burseraceae) (Daly 2020;Daly et al. 2020), a smaller number of species are confined to montane forests, and an even smaller number of species occur in dry areas such as Brazilian (Rodrigues 1980). A few widespread species, including V. elongata and V. sebifera, can be found across different forest types (Aymard et al. 2020).
Virola is important to indigenous and rural communities for its several ethnobotanical attributes. It can be used as medicine to treat malaria, asthma, rheumatism, tumors of the joints, intestinal worms, skin diseases, erysipelas, hemorrhoids, bucal ulcerations, leishmaniasis, and halitosis; while its wood is used both for construction and paper pulp manufacturing, (Rodrigues 1972(Rodrigues , 1980Lourerio et al. 1989;Plotkin and Schultes 1990;Milliken 1997;Rodrigues et al. 2001). Resin from several species, including V. calophylla (Spruce) Warb. and V. calophylloidea Markgr., are potent hallucinogens (Schultes 1954). These hallucinogens, commonly ingested as a snuff known as yakee and yato in Colombia and paricá, kawabó, epena or nyakwana in Brazil (Schultes 1969(Schultes , 1978Aymard et al. 2020), are central to spiritual ceremonies of various indigenous communities in South America. Virola is likely also the source of the powerful hallucinogenic snuff hakudufha made by the Yekwana tribe in the upper Orinoco River and other tribes (Schultes 1954). For the elaboration of these snuffs, only the resin is used, or they mix it with the ashes of the leaves of Justicia pectoralis Jacq. (Acanthaceae), or ashes of the outer bark of Duguetia lepidota (Miq.) Pulle (Annonaceae), Elizabetha leiogyne Ducke (Fabaceae), or Eschweilera itayensis R. Knuth (Lecythidaceae); the preparation of snuff varies among different indigenous communities (Schultes 1976;Schultes and Raffauf 1994;Milliken et al. 1999).
The genus has also been well-studied by botanists and ecologists as it is a tractable system for studying seed dispersal (e.g., Howe and Vande Kerckhove 1980;Howe 1981Howe , 1993Queenborough et al. 2007;Ratiarison and Forget 2013;Moreira et al. 2017).
The last complete taxonomic revision of Virola was published over 80 years ago by Smith and Wodehouse (1938). This treatment recognized 38 species [including V. oleifera (Schott) A. C. Smith, now placed in its own genus Bicuiba W. J. de Wilde]. Smith and Wodehouse (1938) divided the genus into six groups: Mollissimae (I), Sebiferae (II), Calophyllae (III), Rugulosae (IV), Surinamenses (V), and Subsessilis (VI). The characters used to delimit these groups included trichome type, degree of Table 1. List of species of Virola in Central and South America, and Antilles. Region: C = Central America; S = South America; A = Antilles; M = Mexico. For the number of species per country in South America see Fig. 1. References next to a country correspond to species for which we have not seen a herbarium specimen for that country.

Species
Country of original material
During a review of herbarium material from South America as part of the Myristicaceae treatment for "Flora Mesoamericana" (D. Santamaría-Aguilar, in rev.), we found several specimens that we consider fall outside of the morphological species concepts under which they were identified or treated. Most of this material was collected in the 1940s, 1980s, and 1900s, and was identified as V. calophylla, V. macrocarpa A. C. Sm., V. obovata Ducke, V. peruviana (A. DC.) Warb., and V. sebifera, or V. multin- ervia Ducke. We describe these as new species here. Morphologically, the new species appear similar to the species with which they were previously identified; however, detailed observation of complete specimens (i.e., those in which both flowers and mature fruits are present) reveals clear morphological differences in leaves, trichomes, inflorescences, staminate flowers, and, especially, fruits. It is likely that pistillate flowers are also distinct, although these are infrequent in herbarium specimens. With the exception of V. cumala, whose leaves have numerous and close lateral veins and dendritic trichomes, the other new species have leaves with well-separated lateral veins and stellate, sessile trichomes (usually with the central portion darkened) on the abaxial surface of leaf blades (Fig. 4). These characteristics are also shared with V. allenii D. Santam. & Aguilar, V. amistadensis D. Santam., and V. otobifolia D. Santam from Central America.  Below, we describe 10 new species of Virola from Colombia, Ecuador, Peru, and Brazil. This increases the total number of species in Virola to ca. 70,with 33,20,27 and 38 species, respectively, now known from each of these countries. Continued exploration and collection in South America likely will continue to reveal an abundance of new species and new distribution records of Virola.

Materials and methods
Approximately 3000 physical herbarium specimens from Meso and South American Virola were examined for this study from the following herbaria: COL, CR (including ex INB), JAUM, LSCR, LSU, MO, NO, NY and USJ (acronyms follow Thiers 2021 [continuously updated]), although specimens from MO and NY represent the majority of the material studied. All type specimens, as well as general collections, hosted by virtual herbaria, were consulted, including those maintained by: Species descriptions are based primarily on herbarium specimens. 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 = 1 mm) under a dissecting stereoscope (Bausch & Lomb).
Specimens cited are listed first by country. Within a country, specimens are listed alphabetically by major division and then alphabetically by department, province or state 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 TROPI-COS database, the values from TROPICOS are included. Dot-distribution maps were compiled from studied specimens and generated with SimpleMappr (Shorthouse 2010).
Distribution, habitat, phenological data, common name and uses, flower and fruit colors, habit, bark, and exudate data were obtained from herbarium labels.
In the nomenclatural section for each new species, we cite both accession numbers and barcodes when present.
Distinctive characters.. Virola aguarunana is recognized by its very short staminate inflorescences (2.2-2.5 cm long) with dense fascicles of flowers, relatively thin perianth that is externally densely pubescent and internally moderately pubescent, and a filament column that is shorter (0.4-0.5 mm long) than the anthers (0.6-0.9 mm long). It is further distinguished by its pyriform to subglobose fruits covered with ferruginous trichomes (Fig. 6A) that fall easy to the touch and leaf blades usually that are narrow with well-separated later veins and scattered pubescence of stellate and sessile trichomes, on the abaxial surface (Fig. 4A).
Etymology. The specific epithet honors the Aguaruna people, who live in the area where this species was collected.
Phenology. Staminate buds and flowers of Virola aguarunana have been recorded in January, October, and November; pistillate flowers were not seen in the studied material. Fruits have been collected in August and September.
Common name and uses. No common names or uses are mentioned among the herbarium specimens observed.
Preliminary conservation status. Virola aguarunana is Endangered following IUCN criteria B1a and B2a. It is known from two localities, has an EOO of 98 km 2 , and an AOO of 12 km 2 . Further justifying this status, this very small distribution is combined with occurrence in areas known to be impacted by forest declines driven by shifting agriculture demands (Antonelli 2022). Of the few species we were able to verify, the most recent was collected in 1997.
Discussion. Most of the specimens of Virola aguarunana were previously determined as V. calophylla or V. sebifera, both widely distributed in South America. The new species shares some characteristics with V. calophylla, including a mixture of stellate and sessile trichomes and well-separated lateral veins (a feature shared with V. sebifera). However, V. aguarunana can be distinguished by its sparsely pubescent abaxial leaf surface (vs. densely pubescent; Fig. 4A, E), longer staminate perianth ([2-] 2.5-2.7 mm vs. 1-2.1 mm long), a filament column that is shorter than the anthers (vs. longer than anthers), and fruits that are covered by a dense layer of trichomes (vs. covered with an inconspicuous layer of trichomes; Fig. 6A, E). Virola aguarunana shares the following traits with V. sebifera: a filament column that is shorter than the anthers, fruits covered by a dense layer of trichomes, and the same leaf traits as shared with V. calophylla. The new species differs from V. sebifera, by the sessile trichomes on the abaxial leaf surface (vs. usually pediculate; compare with Fig. 4M) and larger fruits (2.9-3.5 × 1.8-2.2 cm vs. 1-1.9 [-2.1] × 0.7-1.4 [-1.7] cm). Additionally, V. aguarunana can be differentiated in both species for its short and narrow staminate inflorescences with flowers borne in dense terminal fascicles. Further, V. calophylla and V. sebifera tend to have larger leaf blades.
Notes. The only seed that could be measured is from one of the two fruits of the specimen C. Díaz et al. 7195 (MO), which is not well preserved. The typical seed size for V. aguarunana is likely larger than presented here.  Fig. 7 Diagnosis. Virola alvaroperezii is more similar to V. macrocarpa and V. otobifolia from Colombia, Panama, respectively. All these species have relatively large leaf blades and fruits, with lateral veins that are well separated and the abaxial leaf surface covered with stellate, sessile trichomes. Morphologically, it differs from V. macrocarpa and V. otobifolia by the abaxial leaf surface densely pubescent (vs. sparsely pubescent). Additionally, it differs from V. macrocarpa in having lateral veins that are more separated ([2-] 2.4-2.7 cm apart vs. 0.8-1.5 cm apart), and large fruits (4.3-4.5 × 3-3.6 cm vs. 2.7-3.3 × 2-2.3 cm). It differs from V. otobifolia by its wide fruit (3-3.6 cm vs. [1.9-] 2.3-2.9) and thin pericarp (2.8 mm vs. [2.7-] 3-4.7 mm).
Distinctive characters.Virola alvaroperezii is a distinctive species characterized by generally large, wide leaf blades with well-separated lateral veins that are abaxially covered with dense pubescence of stellate, sessile trichomes ( Fig. 4B) and large fruits (4.3-4.5 × 3-3.6 cm) with thick pericarp (2.8 mm) (Fig. 6B). It is also distinctive for being a large tree (30-40 m tall).
Etymology. It is a great pleasure to dedicate this new species to the Ecuadoran botanist Álvaro Javier Pérez Castañeda. He is an excellent botanist, collector and expert of the flora of Ecuador, especially the flora of Yasuní. Among other contributions, he has described at least 26 from different angiosperm families (e.g., Pérez et al. 2013;Torke and Pérez 2013;Pérez 2015, 2016) and is coauthor of Árboles emblemáticos de Yasuní, Ecuador (Pérez et al. 2014) and the treatment of Myrtaceae for the Flora of Ecuador (Kawasaki et al. 2019). Pérez Castañeda collected specimens of some of the Virola species described here.
Distribution. Virola alvaroperezii is know from Carchi and Esmeraldas provinces in Ecuador (Fig. 18A). It grows in primary vegetation in premontane rain and very wet forest from (500-) 1600-1800 m.
Phenology. Only a single studied specimen of Virola alvaroperezii has pistillate flowers; it was collected in August. Staminate flowers were not observed in the studied material. Fruits were collected in August and September.
Preliminary conservation status. Virola alvaroperezii is Endangered following IUCN criteria B1a and B2a. It is known from three localities, has an EOO of 966 km 2 , and an AOO of 12 km 2 . Further justifying this status, this new species occurs in a region of very high rates of deforestation due to agricultural pressures (Kleemann et al. 2022). The only specimens of V. alvaroperezii that we were able to verify were collected on the Awá Reserve, and this species may be locally protected by sustainable forestry practices by the Awá indigenous community (Oviedo 2006).
Discussion. Herbarium specimens of Virola alvaroperezii have been confused with another montane species, V. macrocarpa from Colombia (1100 m elevation). It could be confused with another Colombian species, V. cogolloi (840-1500 m elevation), which is formally described here, as well with V. otobifolia D. Santam., recently described from Panama (50-850 m elevation) (Santamaría-Aguilar et al. 2019). All these species have relatively large leaf blades and fruits, with lateral veins that are well separated and the abaxial leaf surface covered with stellate, sessile trichomes. Differences among the three species are summarized in Table 2.
Notes. The holotype deposited at Missouri Botanical Garden (MO) represents a single collection mounted on two sheets that are clearly labeled (i.e., "Sheet 1 of 2" and "Sheet 2 of 2") as being parts of the same specimen (ICN Art. 8.3;Turland et al. 2018); one of the sheets carries pistillate flowers (MO-05005569) and the other, fruits (MO-05005570).
Leaf size and the number of lateral veins may be higher than presented here, as most of the material examined had broken leaves.
The collection C. Aulestia & M. Aulestia 1017 (MO) from Esmeraldas Province, Ecuador bears a fruit (possibly immature) that is smaller (ca. 3.7 × 2.6 cm) with thinner pericarp (ca. 2 mm) than other specimens of V. alvaroperezii. However, it otherwise matches very well including in its leaf morphology, tall tree habit, and occurrence Río Mira basin; for this reason, it is included in the concept we have adopted, although the fruit measurement is not included in the description.
The fruit specimen H. Vargas et al. 4603 (MO,QCNE [image!]), also from Esmeraldas Province, Ecuador, could not be confirmed to the species. It is similar to V. alvaroperezii in its leaf morphology (e.g. size, pubescence and trichomes, lateral vein spacing); however, its fruits are rounded and smaller (ca. 3.2-3.3 × 2.6-2.7 mm) with a conspicuous carina.
The two collections of V. macrocarpa from Carchi Province that are cited in Flora of Ecuador (Jaramillo et al. 2004) Fig. 8 Diagnosis. Virola bombuscaroensis differs morphologically from all other species by the combinations of abaxial leaf surface covered by stellate, sessile trichomes, perianth of staminate flowers densely pubescent on both faces, and the column of filaments that has scattered trichomes at the base.
Distinctive characters. Virola bombuscaroensis can be recognized by many pubescence characters. Dendritic trichomes cover its branches, petioles, inflorescence axes, and sometimes along the main veins on the abaxial sides of leaves; the abaxial leaf surface always is marked by stellate, sessile trichomes (Fig. 4C); its fruits are covered by ferruginous trichomes that fall very easily to the touch (Fig. 6C); the perianth of staminate flowers is densely pubescent on both sides; and the column of filaments has scattered trichomes at the base. Finally, the filament column is straight, not constricted at the apex, and is longer (ca. 1.4 mm) than the anthers (ca. 0.9 mm).
Etymology. The specific epithet makes reference to the Bombuscaro River in the Podocarpus National Park of southern Ecuador, where most of the collections of this new species were made. Distribution. Virola bombuscaroensis is known only from Ecuador (Zamora-Chinchipe Province) (Fig. 18A). It is found in premontane forest at elevations between 1200 to 1350 m, with a single collection from 1930 m in elevation.
Phenology. Staminate flowers of Virola bombuscaroensis have been recorded in April and fruits in January. Pistillate flowers were not seen in the studied material.
Common name and uses. None recorded. Preliminary conservation status. Virola bombuscaroensis is Critically Endangered following IUCN criterion B2a. While its very small range from only a single locality justifies this status, V. bombuscaroensis benefits from its occurrence in Podocarpus National Park and surrounding areas, which are highly protected within Ecuador (Kleemann et al. 2022).
Discussion. Virola bombuscaroensis is difficult to confuse with other species of Virola described to date. However, it shares the combination of a cordate leaf base, well-separated lateral veins, sessile trichomes on abaxial leaf surface, and fruits with a conspicuous layer of trichomes with V. excisa (described here). Other species with similar leaf morphology, but with fruits with less conspicuous trichomes include: V. calophylla (Fig. 6E), V. peruviana (Fig. 6K), and V. schultesii A. C. Sm.; they can be distinguished by the characters in Table 3.
Other species with similar leaf morphology (i.e., a usually deeply cordate base and well-separated lateral veins) are: V. divergens Ducke, V. mollissima (Poepp. ex A. DC.) Warb., and V. sebifera. However, these species differ in having pediculate trichomes on the abaxial leaf surface (vs. sessile in V. bombuscaroensis), internally puberulent staminate perianth (vs. densely pubescent), a glabrous filament column (vs. with scattered trichomes at the base; Fig. 10D) that is shorter (vs. longer) than the anthers. In fruiting material, V. divergens, and V. mollissima are covered with a thick layer of long hairs (at least 1.4 mm long) that differ from V. bombuscaroensis. While the layer of trichomes on the fruit are somewhat similar to the new species in V. sebifera, the fruits are notably smaller (1-1.9 [-2.1] × 0.7-1.4 [-1.7] vs. 2.4 × 1.8-1.9 cm), and V. sebifera usually occurs at lower elevations.
Additionally, compared to the species with with it can be confused, V. bombuscaroensis tends to have longer staminate inflorescences and more obviously pubescent twigs, petioles, and abaxial leaf surface, very often with dendritic trichomes.
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 km 2 , and an AOO of 12 km 2 . 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 (Vélez et al. 2020) and proximity to the Awá 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.
Notes. The first collection of Virola calimensis was made 76 years ago by José Cuatrecasas (J. Cuatrecasas 17540;5-15 May 1944) in Río 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 Chocó region (5-50 m elevations) respectively. 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).
Distinctive characters. Virola cogolloi is best distinguished by the combination of its wide leaf blades that are abaxially covered with dense but inconspicuous stellate and sessile trichomes and with lateral veins that are well-separated (Fig. 4G); wide staminate inflorescences with few flowers per fascicle; and staminate flowers with fleshy perianth lobes that are internally covered by a dense layer of trichomes (especially on the lobes) and filament columns that are longer (1.2-1.4 mm long) than the anthers (1.1-1.2 mm long). It is also distinctive for its densely tomentose fruits with relatively thin pericarp (1.2-1.6 mm thick, the thinnest side) (Fig. 6G) and a seed covered by a thin aril that is laciniate almost to the base.
Etymology. The specific epithet honors Álvaro Cogollo Pacheco, the Colombian botanist who collected most known specimens of this new species as well for his valuable contribution to our knowledge of the Colombian flora. We celebrate his important contributions to botany, epitomized by his numerous collections, ~48 of which now represent type specimens (Tropicos 2021). Álvaro is author of the Myristicaceae treatment for the Catálogo de las Plantas Vasculares de Antioquia (Cogollo 2011), among others.
Distribution. Virola cogolloi is known only from Colombia (Antioquia department) (Fig. 18C). It is found between 840-1500 m elevation in premontane forest in Las Orquídeas National Park in the Western Cordillera of Colombia.
Phenology. Specimens with staminate flowers of Virola cogolloi were collected in February and May. Fruits have been observed in February, April to July. Pistillate flowers were not seen in the studied material.
Common name and uses. Sebo (Colombia; Á. Cogollo et al. 4195). Preliminary conservation status. Virola cogolloi is Endangered following IUCN criteria B1a and B2a. It is known from three localities, has an EOO of 24 km 2 , and an AOO of 24 km 2 . While V. cogolloi benefits from its occurrence within Las Orquídeas National Park, this region (including within the national park) is still vulnerable to deforestation to expand human activities, including agriculture and livestock grazing (González-Caro and Vásquez 2017; Pedraza-Peñalosa 2015).
Discussion. Prior to our study, most of the specimens with fruits of Virola cogolloi were identified as V. macrocarpa. Both species grow in Colombian montane forests, respectively in the departments of Antioquia (840-1500 m elevation) and Boyacá (1100 m elevation). In addition to a shared habitat, these species have similar size and shape of the fruits and, like other species described here, abaxial leaf blades with stellate, sessile trichomes with a reddish central portion. However, V. cogolloi is distinguished by the size of its leaf blades, petiole thickness, separation of the lateral veins, and other features summarized in Table 6.
A second Virola species (V. tuckerae, formally described below) occurs with V. cogolloi. In addition to their similar distributions, these species share leaf blades that are densely pubescent abaxially (Fig. 4G, N). However, V. tuckerae, has narrower leaf blades with more lateral veins, a filament column that is shorter than the anthers, and a fruit that is covered by a dense layer of trichomes. A comparison between these two species is presented in Table 7.
Distinctive characters. The most distinctive characteristic of Virola cumala is the dense indument of dendritic brown to ferruginous trichomes that cover almost all parts of the plant. Vegetatively, this species also has large leaves with numerous lateral veins that are prominent (including the tertiary veins, especially below), thick twigs, and bark that sometimes cracks and flakes into small pieces. Further, the staminate flowers have a subcarnose perianth that is densely and uniformly pubescent outside and glabrous inside with narrow lobes, a filament column that is straight and similar in length to the anthers, and large fruits that are covered with a thick layer of trichomes, thick pericarp, and a seed covered by an aril that is laciniate in very narrow bands almost to the base (Fig. 12D). Distribution. Virola cumala is known only from Peru (in the Amazonas, Loreto, and Pasco departments) (Fig. 18C). It occurs in primary forests in non-inundated, lateritic soil. It ranges in elevation from 120-320 m, although one collection from Pasco reaches 700-800 m (H. van der Werff 20095). It is possible that this species also occurs in Ecuador, as some collections were made close to the border of Peru and Ecuador.
Phenology. Staminate flowers of Virola cumala have been recorded in July; pistillate flowers were not seen in the studied material. Fruits have been collected in May, August to October, and December.
Common name and uses. Cumala negra (Peru; R. Vásquez & N. Jaramillo 7879,MO). Preliminary conservation status. Virola cumala is Endangered following IUCN criterion B2a. It is known from five localities, has an EOO of 219,024 km 2 , and an AOO of 20 km 2 . Justifying its status, the western Amazonian region where this species occurs is modeled to be modestly impacted by new road construction, a threat that is likely to be superseded by the development of oil palm plantations in the region (Arima 2016).
Discussion. Virola cumala is found among a morphological group of ca. 17 species (personal count) that correspond to the group Rugolosae of Smith and Wodehouse (1938). Generally, this group is characterized by leaves with numerous, closely adjacent lateral veins, percurrent tertiary veins that are usually conspicuous, cordate bases, and dendritic trichomes; staminate inflorescences often with large bracts that cover flowers; staminate flowers that are small and usually densely aggregated with membranous to sub membranous perianth, the tube almost split to the base, and a slender filament column. Within this group, the new species is most similar to V. decorticans Ducke and V. multinervia Ducke, both from South America, and V. megacarpa A. H. Gentry from Panama. All four species have large leaves, numerous lateral veins, dense indument of dendritic brown to ferruginous trichomes in almost all parts of the plant, large fruits with a thick layer of trichomes (thin in V. megacarpa), and thick pericarp. Virola cumala further shares bark that cracks and flakes into small pieces and the thick young terminal bud of leaves. These species are distinguished by the characteristics presented in Table 8.
Distinctive characters. Virola excisa can be recognized by its long, lanceolate to obovate leaves that are truncate to subcordate at the base with an underside that is usually white grayish when dry and sparsely pubescent with stellate sessile trichomes (sometimes with dendritic trichomes along the veins) (Fig. 14B); lateral veins that are well separated, with the marginal and tertiary veins not very conspicuous; staminate flowers with the external perianth covered with dense pubescence of dendritic trichomes and a glabrous internal surface, a fleshy filament column that is similar in length ([0.4-] 0.5-0.8 mm long) to the anthers (0.5-0.7 mm long); fruits that are densely pubescent with trichomes that fall easily like dust; and the seed with a bullate, sulcate testa sulcate (Fig. 6I). Additionally, herbarium specimens are usually aromatic.
Etymology. The specific epithet of the new species comes from the word excisus, meaning cut out (Stearn 1992). The makes reference to the fact that several herbarium specimens of this new species come from felled trees that were killed to make way for oil pipes (e.g. V. Zak & S. Espinoza 4733, 5149;F. Hurtado 2980). This has created irreparable damage to nature, destroying the habitat of this and many other species, potentially paving the path toward their extinction. Distribution. Virola excisa is known from Colombia (Amazonas, Putumayo departments), Ecuador (Napo, Pastaza, Morona-Santiago provinces), Peru (Amazonas, Huánuco, Loreto, Pasco, Ucayali departments), and Brazil (Acre, Amazonas state) (Fig.  18B), where it occurs in primarily moist tropical forest. Some herbarium labels mention that it grows on hills with red soils, sandy clay soil, near black water or várzea. It ranges in elevation from 130-580 m, with two collections reaching 700 and 830 m.
Phenology. Staminate flowers of Virola excisa have been collected in February, May to October, and pistillate flowers in May, July to September. Fruits have been collected in January to April, August to December.
Common name and uses.  (M. Silveira et al. 718). The trunk of V. excisa is used for construction of houses, while the aril attracts birds, rodents, and monkeys (Pérez et al. 2014, as V. obovata) or the wood used for firewood (M. Silveira et al. 840).
Preliminary conservation status. Virola excisa is Not Threatened following IUCN criteria B1a and B2a. It is known from 25 localities, has an EOO of 810,234 km 2 , and an AOO of 112 km 2 . While this species grows in regions of the world that are threatened by human landscape modification (Antonelli 2022), its distribution is wide enough that this species does not need to be considered threatened currently.
Discussion. Of morphology apparent in herbarium specimens, the leaf blades and some flower and fruit features of V. excisa are similar to V. obovata, and most of the studied specimens were identified as such in Flora of Ecuador (Jaramillo et al. 2004). Both species share similarly shaped leaves that appear to be glabrous abaxially, but actually bear sparsely stellate and/or dendritic trichomes, or both kinds; staminate flowers with perianth that is nearly glabrous internally and the filament column of similar size to the anthers; and fruits that are covered with a conspicuous layer of ferruginous trichomes. Despite these morphological similarities, V. excisa differs from V. obovata in traits related to the leaf base, the perianth of staminate flowers, and fruit, pericarp, and seed size. Differences between these species are summarized in Table 9. In addition to the features in Table 9, there are distinguishing traits that are difficult to describe properly; compared to V. obovata, herbarium specimens of V. excisa tend to have more elongate leaves and staminate perianth, fruits usually have more conspicuous carina, and trichomes are denser and fall more easily.
Virola excisa resembles V. calophylla and especially V. peruviana in the shape of its leaves (including the base and lateral vein pattern) and long staminate inflorescences with perianth that is glabrous internally. In addition, V. calophylla and the new species have similar fruit morphology (at least when young). Virola excisa can be distinguished from V. calophylla by its abaxial leaf blades that are sparsely pubescent and puberulent (vs. densely pubescent and appearing squamose; see Fig. 14A), shorter, narrower staminate inflorescences, shorter perianth, and filament column that is similar in length to the anthers (vs. filament column longer than anthers). Immature fruits of V. calophylla are densely pubescent (e.g. D. Daly et al. 6773,INPA [image!], MO!, NY!) and appear similar to those of V. excisa; however, at maturity, its fruits tend to be minutely tomentelous to glabrescent (vs. densely tomentose; see Fig. 6E, I). Additionally, abaxial leaf blades of herbarium specimens of V. calophylla tend to be silver to golden.
Virola excisa can distinguished from V. peruviana by its staminate flowers with short anthers (0.5-0.7 mm vs. 1.1-1.6 mm long, from Smith and Wodehouse 1938) and fruits that are slightly carinate (vs. conspicuously carinate) and densely pubescent at maturity (vs. glabrescent). For a detailed comparison of the indument and carina of fruits between these two species see Figs 6I, K (fruits) and 14B, D (indument). Although, some specimens of V. excisa have a cordate leaf base (e.g. V. Huashikat 2267, MO; D. Rubio & T. Coba 839, MO), they are never as deeply cordate or as narrow as in V. peruviana, whose lobes also sometimes overlap or cover the twig (e.g. W. H. Lewis et al. 10074, MO).
Notes. The majority of specimens cited here as V. excisa, including the source of illustrations in Pérez et al. (2014) and Jaramillo et al. (2004) were previously identified as V. obovata. In addition to the list of paratypes of V. excisa below, we provide a list of specimens that correspond to V. obovata. Our comparison between these two species is based on these specimens, as well as images of the original material of V. obovata collected and described by the Italian-Brazilian botanist, Adolpho Ducke (A. Ducke 1509, A, F, IAN, MG, NY!, RB-2 sheets, US-2 sheets).
Distinctive characters. The most striking character of Virola parkeri is its fruit, which is reminiscent of Saturn's form: it is oblate with a conspicuous "wing" along its dehiscence line (Fig. 6L, 15F). The fruit is further bullate and inconspicuously pubescent. Other characteristics that distinguish this new species are its leaf blades that are covered abaxially with inconspicuous, usually colorless stellate and sessile trichomes and lateral veins that are evenly spaced; staminate flowers with perianth that is densely pubescent outside and the inner side moderately pubescent, the lobes are relatively thin, the column of filaments are shorter (0.2-0.4 mm long) than the anthers (0.5-0.7 mm long); and the thin, laciniate aril that covers the seed almost to the base.
Etymology. The specific epithet honors Theodore A. Parker III (1 Apr. 1953-3 Aug. 1993), renowned and talented ornithologist and research associate of the LSU Museum of Natural Science. Parker died in a plane crash on August 3, 1993 while surveying a remote forest in Ecuador, along with three other people: Raul Mortensen (the pilot), Eduardo Aspiazu (ecologist), and Alwyn H. Gentry (botanist). Virola parkeri occurs in Peru, a country where Parker spent much of his life studying birds and protecting natural resources. In this country, Parker together with Scott Robinson, established a Big Day record (331 bird species). He also contributed to a field guide to the birds of Peru, an important resource for birdwatching ecotourism. Finally, Parker was part of the Rapid Assessment Program (RAP) at Conservation International, which involved teams of scientists conducting biological surveys in remote areas of the tropics to determine their level of biodiversity and potential for conservation. Thanks to Parker's efforts, Bolivia established Madidi National Park, one of the most important centers for biodiversity.
Distribution. Virola parkeri is known only from Peru (Pasco Department) (Fig. 18C). It is found in primary forest between 400 and 620 m in elevation.
Phenology. Staminate flowers and fruits of Virola parkeri have been recorded in October. Pistillate flowers were not seen in the studied material. Preliminary conservation status. Virola parkeri is Critically Endangered following IUCN criterion B2a. Justifying this status, it is known from only a single specimen, collected in 2008. The only locality of this species is within the Loma Linda community of the Yanesha indigenous community, and so may benefit from an ongoing compensated community-based forest monitoring program (Kowler et al. 2020).
Discussion. Virola parkeri is morphologically similar to V. yasuniana from Ecuador (which is formally described below) in various leaf traits, including overall shape, trichomes on the abaxial surface, lateral vein separation, and color when dry; additionally, both species share staminate perianth that is moderately pubescent internally and fruits with a markedly carinate dehiscence line, appearing like a wing. Differences among the two species are summarized in Table 10.
Tree (12-) 18-30 m × 17-30.1 cm diameter, inner bark not described. Exudate red, location of exudate on plant not stated, or in the fruit hyaline and oxidizing red. Twigs 0.21-0.37 cm thick, terete or slightly compressed, tomentose, trichomes dendritic, sessile, ferruginous, without lenticels or lenticels very small and scattered.  . 16805), tomentose, the trichomes dendritic; leaf blades 21-29.5 (-33.5) × 5-7.2 (-11.7) cm, narrowly oblong or rarely elliptical; adaxial surface when drying on mature leaves brown to blackish brown, the surface smooth, sometimes shiny, glabrous; abaxial surface when drying pale brown to grayish brown, densely pubescent, the trichomes stellate, ca. 0.1 mm diameter, sessile, the central part of the trichome pale reddish, sometimes a little darker, the branches pale reddish and not contrasting much in color with the central part of the trichome; lateral veins 16-19 per side, 3-4 (-6) veins per 5 cm, spaced 1.1-1.8 (-2.1) cm, on adaxial side, the same color as the adaxial surface or slightly darker, flat to slightly raised, on abaxial surface blackish to brown reddish, raised, puberulent 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 very slightly visible on both sides, but especially above; midvein adaxially flat to slightly elevated, abaxially raised, rounded, tomentose to puberulent, more pubescent to the sides; base cuneate, not revolute, flat; margin flat; apex acute. Staminate inflorescence 5-9.5 cm long, axes flattened, tomentose, trichomes dendritic, ferruginous; peduncle 0.6-1.8 (-2.8) × 0.2-0.38 cm; main axes with (3-) 5-9 (-12) ramifications, the first pair opposite to subopposite, the others alternate; bracts not seen. Staminate flowers (in bud) in dense terminal fascicles of 9-20 flowers, on a receptacle 2-3 mm wide; perianth 3.5-5 mm long, oblong, fleshy, ferruginous when fresh (probably by the trichomes), connate to 1.5-2.5 mm in length, external surface densely pubescent with ferruginous and dendritic trichomes, internal surface densely pubescent (especially in the lobes); lobes 3, 2-2.5 × 1.2-1.7 mm, and 0.2-0.5 mm thick, without resinous punctuations when rehydrate); stamens 3, the filament column 0.6-0.7 (-0.9) mm long and 0.2 mm wide, glabrous, straight or sometimes a little wider at the base, not constricted at the apex; anthers 1.2-1.6 mm long, and 0.2-0.4 mm wide; apiculus 0.1-0.2 mm long, acuminate, slightly separated or connate. Pistillate inflorescence and flowers unknown. Infructescence unknown. Fruit 2.7 × 2.5 cm (only one seen, and that immature; Á. Cogollo et al. 4147), when fresh green and covered with brown trichomes, globose, densely tomentose, the trichomes dendritic (ca. 0.1-0.2 mm long), sessile, ferruginous, that fall as easily as dust, the surface probably smooth, the line of dehiscence slightly carinate, the base and the apex obtuse; pericarp ca. 2.4 mm thick; pedicel unknown. Seed length unknown × ca. 1.4 cm, the testa brown when dry, slightly ribbed distally; aril color not described when fresh, blackish when dry, the texture dry and thin, laciniate almost to the base, in narrow bands distally. Distinctive characters. Virola tuckerae can be recognized by its narrow, oblong leaves with relatively close lateral veins (3-4 [-6] veins per 5 cm) and a cuneate base; its short-pedunculate staminate inflorescence with flowers organized in dense fascicles; its staminate flowers with fleshy perianth that is densely pubescent on both sides and a straight filament column that is shorter (0.6-0.7 [-0.9]) mm long) than the anthers (1.2-1.6 mm long); and its globe fruit that is densely tomentose with dendritic and ferruginous trichomes that fall like dust (Fig. 6N). Like other species described here, the new species is covered with stellate and sessile trichomes on the abaxial side of the leaf blades.
Etymology. It is a pleasure to name a species of Virola in honor of Dr. Shirley Cotter Tucker, a botanist, lichenologist, and Professor Emeritus at Louisiana State University (LSU). Despite many challenges she faced as a woman in science, Shirley has had an illustrious career marked by many honors, including a Boyd Professorship, the most prestigious academic rank granted at LSU to internationally renowned scholars.
Shirley is an important leader in botany, and served as president of two of the USA's most prominent botanical societies, the Botanical Society of America and the American Society of Plant Taxonomists. Shirley's intellectual contributions to botany are lasting, providing the foundation framework from which current research on floral morphology and evolution builds. Much of Shirley's academic research (which includes more than 100 published articles) has focused on floral morphology and anatomy, especially of legumes and magnoliids-including studies within Myristicaceae (Armstrong and Tucker 1986), making it particularly special to name a species of Virola in her honor. She maintains her passion for lichens into retirement, and actively curates loans of lichen specimens from her home in Santa Barbara, California. The generous philanthropy of Shirley and her late husband, Kenneth Tucker, have greatly benefitted the botanical community. At LSU, where two of the authors work, donations from the Tucker family established the Shirley C. Tucker Endowed Chair in Plant Systematics. The LSU herbarium is named in her honor.
Distribution. Virola tuckerae is only know from Antioquia, Colombia (Fig. 18A). It has been collected in premontane wet forest at 1300-1420 m elevation.
Phenology. Virola tuckerae was collected with flowers in December and fruit specimens collected in February. Pistillate flowers were not seen in the studied material.
Common name and uses. Sebo cordillero (Colombia; Á. Cogollo et al. 4147). Preliminary conservation status. Virola tuckerae is Endangered following IUCN criterion B2a. Justifying this status, it is known from two localities and has an AOO of 4 km 2 . While V. tuckerae benefits from its occurrence within Las Orquídeas National Park, this region (including within the national park) is still vulnerable to deforestation to expand human activities, including agriculture and livestock grazing (Pedraza-Peñalosa 2015; González-Caro and Vásquez 2017).
Virola tuckerae shares similarities with V. yasuniana, including its leaf shape, sessile, stellate trichomes on the abaxial leaf surface, and the color of dried herbarium specimens. Virola yasuniana is a species primarily from the lowlands of Ecuador (200-480 [1000] m elevations) that is formally described below. However, V. tuckerae differs from it in its densely pubescent abaxial leaf surface (vs. sparsely pubescent to glabrescent in V. yasuniana) (Fig. 4N, O), staminate flowers with perianth that is densely pubescent internally (vs. moderately pubescent) and long anthers (1.2-1.6 mm vs. 0.5-0.6 mm long), and densely pubescent fruits (vs. puberulent).
Finally, Virola tuckerae and V. cogolloi, grow closely in the same region (Urrao, sector Calles, Colombia); the differences and similarities between these species are discussed under V. cogolloi.
Notes. As mentioned above, collections with flower have been previously identified as V. sebifera. The specimen with fruit (Á. Cogollo et al. 4147) was previously identified as V. elongata. Duplicates may have been distributed under these names.
Distinctive characters. The very distinctive fruits of Virola yasuniana, which are large with very conspicuous wings in the line of dehiscence (Fig. 6M), make it almost impossible to confuse with any other species of Virola already described. Other characteristics that distinguish this new species include leaf blades that are abaxially covered with inconspicuous stellate and sessile trichomes, evenly spaced lateral veins that are comparatively spaced, and moderately pubescent internal perianth in staminate flowers, and a filaments column that is wide at the base, constricted at the apex, and usually shorter (0.3-0.5 mm long) than the anthers (0.5-0.6 mm long).
Etymology. The specific epithet refers to Parque Nacional Yasuní, where most of the collections of this new species come from. At present, 2700 vascular plant species are documented from Yasuní, including a high diversity of lianas, epiphytes, and ferns; it is estimated its flora comprises approximately 3213 species; Yasuní is also home to ca. 1570 species of birds, fishes, mammals, amphibians and reptiles (Bass et al. 2010;Pérez et al. 2014). The description of Virola yasuniana is one step closer to a full documentation of this region.
Distribution. Virola yasuniana is known from the Napo and Pastaza provinces of Ecuador, where it occurs in primary forests, on hills with red soil, or soils composed of sedimentary rocks. Is also located in Acre state of Brazil, where it grows in terra firme on poorly drained terrace (Fig. 18A). It ranges from 200-480 m in elevation, with a single collection reaching 1000 m (C. Cerón & F. Hurtado 6560).
Phenology. Staminate flowers of Virola yasuniana have been collected in January, February and November, while pistillate flowers have been collected in April, May, and July. Fruits are known from February, March, June, and September. Preliminary Conservation Status. Virola yasuniana is Vulnerable following IUCN criterion B2a. It is known from two localities, has an EOO of 117,581 km 2 , and an AOO of 32 km 2 . This species benefits from its occurrence in the Yasuní National Park of Ecuador, which experiences very low rates of deforestation (Bass et al. 2010;van der Hoek 2017), even while the broader region is experiencing land use changes (Heredia-R et al. 2021).
Discussion. It is possible to confuse Virola yasuniana with V. parkeri from Peru due to their similar leaves (i.e. shape, color when dry, base, venations, and trichomes), staminate perianth that is pubescents on both surfaces, and markedly carinate fruit. Differences among the two species are summarized in Table 11.
Herbarium specimens of Virola yasuniana were previously identified as V. calophylla, V. obovata, or V. peruviana, all of which have leaf blades with lateral veins that are well separated and stellate, sessile trichomes on abaxial leaf surface. Additionally, the new species shares fruits with a conspicuous carina with V. peruviana. Differences among these species are summarized in Table 10.  Notes. Several collections of Virola yasuniana were treated as V. obovata in Flora of Ecuador (H. Vargas & J. Cerda 678, V. Zak & S. Espinoza 4871, 5039, 5049, 5149;Jaramillo et al. 2004). Additionally, studied with "aberrant fruits" that is discussed under V. peruviana in Flora of Ecuador (Jaramillo et al. 2004) (Mitchell 2002).
Etymology. The specific epithet alludes to the Spider monkeys, or ateles (Ateles panisczis), locally called kwata, who consume the fruits of this species (Sabatier 1997).
Note. Virola kwatae was previously known from French Guiana. However, in a recent study of herbarium specimens at New York Botanical Garden, a collection with fruit made in Amapá state, Brazil was located. To our knowledge this represents the first record of this species among the Flora of Brazil.
Specimens examined. brazil. Amapá: Município Macapá, Rio Falsino, approx. 10 km upstream of confluence with Rio Araguari, 00°50'S, 51°45'W, 13 Dec 1984 (fr), D. C. Daly et al. 3865 (MO [n.v.] Distribution. Endemic to Colombia (Boyacá). Notes. Virola macrocarpa was described by Smith based on a specimen collected by the Scottish immigrant Alexander E. Lawrance in the Andes of Boyacá, Colombia (Smith and Wodehouse 1938). In the protologue, Smith mentions that the type is deposited in the herbarium of the Botanisches Museum, Berlin-Dahlem (B), and F, M, S, US; there are additional, duplicates at A, G, K, and MO. Because the material at B was destroyed during World War II, we designate the material deposited at Missouri Botanical Garden (MO) as lectotype for this name, according to Art. 9.11 of ICN (Turland et al. 2018). This specimen is mounted on two sheets that are properly labelled as being parts of the same specimen (i.e., "Sheet 1 of 2," "Sheet 2 of 2) according to ICN Art. 8.3 (Turland et al. 2018). One of these sheets shows the adaxial side of leaves and the infrutescence axes (MO-1068404; sheet 1 of 2), while the other (MO-1068405; sheet 2 of 2) includes fragments of both adaxial and abaxial leaf surfaces and a single dehiscent fruit that is attached, with six additional fruits inside the packet. This second sheet is annotated in Smith's handwriting. These specimens were distributed under the name of V. sebifera.
The name Virola macrocarpa was applied to collections from Costa Rica and Panama that now correspond to V. allenii, V. amistadensis, and V. otobifolia, as well some collections from Colombia and Ecuador that now correspond to V. alvaroperezii, V. calimensis, and V. cogolloi.
It is important to mention that, following our species concept, this name has been widely used to identify specimens, and is thus frequent in the literature and specimen databases. As circumscribed here, V. macrocarpa is known only from the type collection and is restricted to Colombia. We believe that the identity of following and others specimens identified as V. macrocarpa should be evaluated when more collections are available, including: 1a Mature leaf blades densely pubescent on abaxial surface ( Fig. 4C-G, N) ....2a 2a Staminate flowers with the filament column shorter than the anthers ....... 3a 3a Leaf blades widely oblong to elliptical (7.3-11 cm wide), base obtuse; staminate inflorescence broadly paniculate; staminate flowers in lax terminal fascicles, the filament column 0.4-0.6 (-0.8) mm long, anthers 0.6-1 (- Twigs and inflorescence axes covered with appressed trichomes; fruits 2.7-3.5 long, covered by an inconspicuous layer of trichomes (Fig. 3B) Leaf blades 25-60 cm long; staminate perianth submembranous; mature fruits covered by a conspicuous layer of trichomes (Fig. 12E)  Leaf blades on abaxial side densely tomentose, with pediculate, dendritic trichomes; staminate flowers with perianth 2-2.5 mm long, carnose; androecium 1.2-1.6 mm long; fruits on the line of dehiscence smooth or slightly carinate, apex obtuse (Fig. 12F)  Leaf blades with 49-65 lateral veins; staminate flowers with filament column 0.5-0.7 mm long; fruits rounded to ellipsoid, covered by a conspicuous