<documentID-CLB-Dataset="25143"ID-DOI="http://dx.doi.org/10.3897/phytokeys.134.37979"ID-GBIF-Dataset="c489096b-11b5-48b9-bd0b-b08af903a201"ID-PMC="PMC6821834"ID-Pensoft-Pub="1314-2003-134-1"ID-Pensoft-UUID="05B0E5AA9A175E549730B523BCD54C03"ID-PubMed="31686954"ModsDocID="1314-2003-134-1"checkinTime="1571900138529"checkinUser="pensoft"docAuthor="Santamaria-Aguilar, Daniel, Aguilar, Reinaldo & Lagomarsino, Laura P."docDate="2019"docId="8BF8C77C59735206B8A3A8C72DCE898F"docLanguage="en"docName="PhytoKeys 134: 1-82"docOrigin="PhytoKeys 134"docSource="http://dx.doi.org/10.3897/phytokeys.134.37979"docTitle="Virola allenii D. Santam. & Aguilar 2019, sp. nov."docType="treatment"docVersion="9"id="05B0E5AA9A175E549730B523BCD54C03"lastPageNumber="1"masterDocId="05B0E5AA9A175E549730B523BCD54C03"masterDocTitle="A taxonomic synopsis of Virola (Myristicaceae) in Mesoamerica, including six new species"masterLastPageNumber="82"masterPageNumber="1"pageNumber="1"updateTime="1732937537851"updateUser="ExternalLinkService">
<taxonomicNameid="3A5CAAB6A146D88F0393397B91B41DD8"ID-CoL="7Z6YJ"LSID="8BF8C77C-5973-5206-B8A3-A8C72DCE898F"authority="D. Santam. & Aguilar"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii"status="sp. nov.">Virola allenii D.Santam. & Aguilar</taxonomicName>
<figureCitationid="38B2D2273D222005E936703A2DA3A794"captionStart="Figure 5"captionStartId="F5"captionText="Figure 5. Virola allenii A leaves B fruit C partial inflorescences. Drawn by Pedro Juarez based on P. H. Allen 6727 (A), R. Aguilar 2224 (B) and R. Aguilar (C) from a photo, physical specimen not seen."figureDoi="10.3897/phytokeys.134.37979.figure5"httpUri="https://binary.pensoft.net/fig/349087"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure5">Figs 5</figureCitation>
<figureCitationid="35607EE10159AFB30F054ADD6338DE5B"captionStart="Figure 6"captionStartId="F6"captionText="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."figureDoi="10.3897/phytokeys.134.37979.figure6"httpUri="https://binary.pensoft.net/fig/349088"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure6">, 6</figureCitation>
<figureCitationid="2A759B21454F82B6B03C175A168043F0"captionStart="Figure 7"captionStartId="F7"captionText="Figure 7. Comparisons of Virola macrocarpa with similar species in Mesoamerica A V. allenii (P. H. Allen 5763, F; inset fruits, from P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9717, MO; inset fruits, from G. McPherson 9715, MO) C V. macrocarpa (A. E. Lawrance 675, MO) D V. otobifolia (J. P. Folsom 1440, MO). Image courtesy of Field Museum (A)."figureDoi="10.3897/phytokeys.134.37979.figure7"httpUri="https://binary.pensoft.net/fig/349089"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure7">, 7A</figureCitation>
in its leaf blades that are whitish on the abaxial side and covered with stellate, sessile trichomes with the centre reddish and contrasting with the hyaline branches to reddish-clear in colour and lateral veins that are not densely arranged, as well as large fruits that are covered with ferruginous trichomes. It differs in its narrow leaf blades (3.2-7.3 cm vs. 7-11 cm wide) with acute or obtuse to rounded bases (vs. broadly obtuse), fruits with thick pericarp (3.2-3.8 mm vs. 1.8-3 mm thick) and preference for humid lowland forests at 0-350 (-1350) m elevation (vs. montane forests in Andes of Colombia at around 1100 m elevation).
<taxonomicNameid="892C601A64CFA1AA1D1B11BB166C4538"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<taxonomicNameid="C3061650EC58342605BD6C9FD307CDB4"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
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 reddish-clear, 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.
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.
1.5-2.5 cm, usually ellipsoid or rarely ovoid, stipitate, densely tomentose, 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.
<bibRefCitationid="AF351FCD45BD4F88FAF60F97C0EA1E94"author="Ley Lopez, JM"journalOrPublisher="Springer Verlag, Berlin"pageId="0"pageNumber="1"refId="B52"refString="Ley Lopez, JM, Chacon Madrigal, E, 2017. Las Plantulas de Arboles y Palmas de la Peninsula de Osa. San Jose, Costa Rica, 3-183."title="Las Plantulas de Arboles y Palmas de la Peninsula de Osa. San Jose, Costa Rica, 3 - 183."year="2017">
<taxonomicNameid="38FFFBB800DE0FA44E6CE3F8B005C47E"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<figureCitationid="C5B8C924745C10BC12D53BEF61685FE6"captionStart="Figure 8"captionStartId="F8"captionText="Figure 8. Pattern of the lateral veins in Mesoamerican Virola A Virola allenii (K. Thomsen 1284, NO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (B. Hammel et al. 16864, MO) D V. elongata (M. Correa & R. L. Dressler 1078, MO) E V. fosteri (G. de Nevers 7226, MO) F V. guatemalensis (G. Ibarra 957, MO) G V. koschnyi (J. Miller & J. C. Sandino 1110, MO) H V. laevigata (N. Zamora & T. D. Pennington 1583, MO) I V. megacarpa (G. de Nevers 5184, MO) J V. montana (E. Lepiz & J. F. Morales 284, MO) K V. multiflora (J. Manzanares 3561, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers et al. 7530, MO) N V. sebifera (T. B. Croat 5959, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO)."figureDoi="10.3897/phytokeys.134.37979.figure8"httpUri="https://binary.pensoft.net/fig/349090"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure8">Fig. 8A</figureCitation>
) 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 (
<figureCitationid="2088AF519B03B477EC5CD42F4AAA2E62"captionStart="Figure 3"captionStartId="F3"captionText="Figure 3. Trichomes in Mesoamerican Virola A Virola allenii (R. Aguilar 2224, MO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (O. Valverde 970, MO) D V. elongata (G. McPherson 13674, MO) E V. fosteri (G. McPherson 20913, MO) F V. guatemalensis (R. Cedillo 3349, MO) G V. koschnyi (R. E. Gereau et al. 3455, MO) H V. laevigata (R. Aguilar 2004, MO) I V. megacarpa (J. A. Duke 15261, MO) J V. montana (E. Bello 855, MO) K V. multiflora (R. Rueda et al. 8196, MO; inset, J. C. Sandino 3302, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers & H. Herrera 7056, MO) N V. sebifera (F. Almeda et al. 5553, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO)."figureDoi="10.3897/phytokeys.134.37979.figure3"httpUri="https://binary.pensoft.net/fig/349085"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure3">Figs 3A</figureCitation>
<figureCitationid="86DCD3B05FDFE229263F19AD54F32AB8"captionStart="Figure 6"captionStartId="F6"captionText="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."figureDoi="10.3897/phytokeys.134.37979.figure6"httpUri="https://binary.pensoft.net/fig/349088"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure6">6F</figureCitation>
); 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 (
<figureCitationid="9A5AA7C0F591CF658403257D8626A866"captionStart="Figure 4"captionStartId="F4"captionText="Figure 4. Fruits of Mesoameican Virola, organised in groups of morphologically similar species A V. allenii (P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9715, MO) C V. otobifolia (A. Gentry & S. Mori 14199, MO) D V. macrocarpa (A. E. Lawrance 675, MO) E V. chrysocarpa (B. Hammel et al. 16864, MO) F V. koschnyi (R. Robles 1587, MO) G V. guatemalensis (S. Sinaca 1542, MO) H V. montana (J. Utley & K. Utley 5054, MO) I V. fosteri (S. Mori & J. Kallunki 4891, MO) J V. multiflora (R. Rueda et a. l 9960, MO) K V. sebifera (T. B. Croat 5959, MO) L V. elongata (J. D. Dwyer & M. Correa 8420, MO) M V. laevigata (R. Aguilar 1585, MO) N V. nobilis from Barro Colorado (T. B. Croat 8090, MO) O V. nobilis from the Osa Peninsula (G. Herrera 4222, MO) P V. megacarpa (G. de Nevers 5184, MO). Scale bars: 1 cm."figureDoi="10.3897/phytokeys.134.37979.figure4"httpUri="https://binary.pensoft.net/fig/349086"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure4">Figs 4A</figureCitation>
<figureCitationid="409379F6DFA12CA5191DB6BA74677094"captionStart="Figure 6"captionStartId="F6"captionText="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."figureDoi="10.3897/phytokeys.134.37979.figure6"httpUri="https://binary.pensoft.net/fig/349088"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure6">
<figureCitationid="758269C84AEA2135E6E20EB111A6C880"captionStart="Figure 6"captionStartId="F6"captionText="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."figureDoi="10.3897/phytokeys.134.37979.figure6"httpUri="https://binary.pensoft.net/fig/349088"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure6">Fig. 6K</figureCitation>
<taxonomicNameid="966BA97C2E04E5A422F3253626A095D9"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
The specific epithet honours the collector of the type specimen, Paul H. Allen (1911-1963), who was probably the first person to collect this species 67 years ago (
<bibRefCitationid="5EC5C256BAF4D90D97E930B8A9171466"author="Grayum, MH"editor="Hammel, BE"journalOrPublisher="Monographs in Systematic Botany from the Missouri Botanical Garden"pageId="0"pageNumber="1"pagination="1 - 50"refId="B35"refString="Grayum, MH, Hammel, BE, Troyo, S, Zamora, N, 2004. Historia/History. In: Hammel, BE, Grayum, MH, Herrera, C, Zamora, N, Eds., Manual de Plantas de Costa Rica. Vol. I. Introduccion. Monographs in Systematic Botany from the Missouri Botanical Garden 97: 1 - 50"title="Historia / History."volume="97"volumeTitle="Manual de Plantas de Costa Rica. Vol. I. Introduccion."year="2004">Grayum et al. 2004</bibRefCitation>
<bibRefCitationid="DADA0DCF21E0D2DD2C1070FF9BBDD239"author="Allen, PH"journalOrPublisher="Monographs in Systematic Botany from the Missouri Botanical Garden 127 (1)"pageId="0"pageNumber="1"refId="B4"refString="Allen, PH, 1956. The Rain Forests of Golfo Dulce. University of Florida Press Gainesville, 1-417."title="The Rain Forests of Golfo Dulce. University of Florida Press Gainesville, 1 - 417."year="1956">Allen 1956</bibRefCitation>
<taxonomicNameid="64DF5320550BC37EF317A56B9E69456E"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<figureCitationid="2D4372D8C9F1DE7939A98444BD329D40"captionStart="Figure 9"captionStartId="F9"captionText="Figure 9. Geographic distribution of Virola allenii (A), V. amistadensis (B), V. chrysocarpa (C), V. elongata (D), V. fosteri (E) and V. guatemalensis (F)."figureDoi="10.3897/phytokeys.134.37979.figure9"httpUri="https://binary.pensoft.net/fig/349091"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure9">Fig. 9A</figureCitation>
<taxonomicNameid="857B8B10C226E76E8E40C53011EAB0FA"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<taxonomicNameid="CEC0E101486400C1188B4C00BC32F556"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="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 km2 and an AOO of 40 km2. 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.
<taxonomicNameid="B2AC5FAA031C0E40D65C5E57EB50D05C"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="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.
<figureCitationid="3890AD5A58B6367E06F783AD8C529359"captionStart="Figure 6"captionStartId="F6"captionText="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."figureDoi="10.3897/phytokeys.134.37979.figure6"httpUri="https://binary.pensoft.net/fig/349088"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure6">Fig. 6I</figureCitation>
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.
<taxonomicNameid="9EC13844EA78030B829358984E9C3095"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<bibRefCitationid="1551B6107005923FFBE9B6F0122D56A8"author="Jimenez, Q"editor="Hammel, BE"journalOrPublisher="Monographs in Systematic Botany from the Missouri Botanical Garden"pageId="0"pageNumber="1"pagination="684 - 691"refId="B46"refString="Jimenez, Q, 2007. Myristicaceae. In: Hammel, BE, Grayum, MH, Herrera, C, Zamora, N, Eds., Manual de Plantas de Costa Rica. Vol. VI. Monographs in Systematic Botany from the Missouri Botanical Garden 111: 684 - 691"title="Myristicaceae."volume="111"volumeTitle="Manual de Plantas de Costa Rica. Vol. VI."year="2007">
<bibRefCitationid="62D3992E36997E5FA4116F8F49C9E87D"DOI="https://doi.org/10.1007/s12228-011-9194-0"author="Cornejo, X"journalOrPublisher="Brittonia"pageId="0"pageNumber="1"pagination="76 - 101"refId="B16"refString="Cornejo, X, Mori, SA, Aguilar, R, Stevens, H, Douwes, F, 2012. Phytogeography of the trees of the Osa Peninsula, Costa Rica. Brittonia 64 (1): 76 - 101, DOI: https://doi.org/10.1007/s12228-011-9194-0"title="Phytogeography of the trees of the Osa Peninsula, Costa Rica."url="https://doi.org/10.1007/s12228-011-9194-0"volume="64"year="2012">Cornejo et al. 2012</bibRefCitation>
<bibRefCitationid="36600BBB30320FD18E1CCAE6986CBBF2"author="Aguilar, R"journalOrPublisher="Monographs in Systematic Botany from the Missouri Botanical Garden 127 (1)"pageId="0"pageNumber="1"publicationUrl="http://sweetgum.nybg.org/osa/"refId="B3"refString="Aguilar, R, Cornejo, X, Santamaria-Aguilar, D, Tulig, M, Bainbridge, C, Mori, SA, 2017 onward. Vascular Plants of the Osa Peninsula, Costa Rica., http://sweetgum.nybg.org/osa/"title="Vascular Plants of the Osa Peninsula, Costa Rica."url="http://sweetgum.nybg.org/osa/"year="2017 onward">Aguilar et al. 2017 onward</bibRefCitation>
<tableCitationid="76E7F989ABCE3575C23B044C7A35E168"captionStart="Table 2"captionStartId="T2"captionText="Table 2. Comparison of Virola allenii with the morphologically most similar species."httpUri="http://table.plazi.org/id/1A2AE0917E035E6F6ACDB17A2E796F2C"pageId="0"pageNumber="1"tableUuid="1A2AE0917E035E6F6ACDB17A2E796F2C">2</tableCitation>
<figureCitationid="38F7E1C54CFC29D60917CF1A41F6D8D2"captionStart="Figure 7"captionStartId="F7"captionText="Figure 7. Comparisons of Virola macrocarpa with similar species in Mesoamerica A V. allenii (P. H. Allen 5763, F; inset fruits, from P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9717, MO; inset fruits, from G. McPherson 9715, MO) C V. macrocarpa (A. E. Lawrance 675, MO) D V. otobifolia (J. P. Folsom 1440, MO). Image courtesy of Field Museum (A)."figureDoi="10.3897/phytokeys.134.37979.figure7"httpUri="https://binary.pensoft.net/fig/349089"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure7">Fig. 7C</figureCitation>
) 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 (
<figureCitationid="0AD1C2C78B0B6AB334485295E3E62235"captionStart="Figure 4"captionStartId="F4"captionText="Figure 4. Fruits of Mesoameican Virola, organised in groups of morphologically similar species A V. allenii (P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9715, MO) C V. otobifolia (A. Gentry & S. Mori 14199, MO) D V. macrocarpa (A. E. Lawrance 675, MO) E V. chrysocarpa (B. Hammel et al. 16864, MO) F V. koschnyi (R. Robles 1587, MO) G V. guatemalensis (S. Sinaca 1542, MO) H V. montana (J. Utley & K. Utley 5054, MO) I V. fosteri (S. Mori & J. Kallunki 4891, MO) J V. multiflora (R. Rueda et a. l 9960, MO) K V. sebifera (T. B. Croat 5959, MO) L V. elongata (J. D. Dwyer & M. Correa 8420, MO) M V. laevigata (R. Aguilar 1585, MO) N V. nobilis from Barro Colorado (T. B. Croat 8090, MO) O V. nobilis from the Osa Peninsula (G. Herrera 4222, MO) P V. megacarpa (G. de Nevers 5184, MO). Scale bars: 1 cm."figureDoi="10.3897/phytokeys.134.37979.figure4"httpUri="https://binary.pensoft.net/fig/349086"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure4">Fig. 4D</figureCitation>
]) 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
(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
<tableCitationid="6D8D4026C7CE405557B9661AC2D50E33"captionStart="Table 2"captionStartId="T2"captionText="Table 2. Comparison of Virola allenii with the morphologically most similar species."httpUri="http://table.plazi.org/id/1A2AE0917E035E6F6ACDB17A2E796F2C"pageId="0"pageNumber="1"tableUuid="1A2AE0917E035E6F6ACDB17A2E796F2C">2</tableCitation>
<figureCitationid="6B92A5A147DF40020660538442C85ABC"captionStart="Figure 10"captionStartId="F10"captionText="Figure 10. Virola calophylla A branch with leaves, showing both sides B branch with inflorescences. Virola elongata C leafy branch with inflorescences, inset fruit D inflorescences. Virola guatemalensis E, F leaf blades showing adaxial (E) and abaxial (F) surface. G young twigs and petioles. Photos by Robin Foster (A, B), from https: // plantidtools. fieldmuseum. org / en / nlp; Steven Paton (C, D), Rolando Perez (C inset), from https: // stricollections. org / portal / index. php and Angela Rojas (E-G)."figureDoi="10.3897/phytokeys.134.37979.figure10"httpUri="https://binary.pensoft.net/fig/349092"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure10">Fig. 10A, B</figureCitation>
<taxonomicNameid="0DDC6BAED8A9B943D61392098691ECFF"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="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
<tableCitationid="DAE9DFE5E78684860078CCD1C8B0F38C"captionStart="Table 2"captionStartId="T2"captionText="Table 2. Comparison of Virola allenii with the morphologically most similar species."httpUri="http://table.plazi.org/id/1A2AE0917E035E6F6ACDB17A2E796F2C"pageId="0"pageNumber="1"tableUuid="1A2AE0917E035E6F6ACDB17A2E796F2C">2</tableCitation>
<taxonomicNameid="B12C930B8EA6E986A074D051E735D875"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<figureCitationid="2B623797EBA4A44860FACB99D3C10893"captionStart="Figure 8"captionStartId="F8"captionText="Figure 8. Pattern of the lateral veins in Mesoamerican Virola A Virola allenii (K. Thomsen 1284, NO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (B. Hammel et al. 16864, MO) D V. elongata (M. Correa & R. L. Dressler 1078, MO) E V. fosteri (G. de Nevers 7226, MO) F V. guatemalensis (G. Ibarra 957, MO) G V. koschnyi (J. Miller & J. C. Sandino 1110, MO) H V. laevigata (N. Zamora & T. D. Pennington 1583, MO) I V. megacarpa (G. de Nevers 5184, MO) J V. montana (E. Lepiz & J. F. Morales 284, MO) K V. multiflora (J. Manzanares 3561, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers et al. 7530, MO) N V. sebifera (T. B. Croat 5959, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO)."figureDoi="10.3897/phytokeys.134.37979.figure8"httpUri="https://binary.pensoft.net/fig/349090"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure8">Fig. 8A, B, M</figureCitation>
<tableCitationid="A5816E5D0850BE62F76FA74702B53A39"captionStart="Table 3"captionStartId="T3"captionText="Table 3. Comparison of Virola allenii with the two other morphologically most similar species in Mesoamerica."httpUri="http://table.plazi.org/id/F0B1B2423AD169B5D41C14685AACA9A1"pageId="0"pageNumber="1"tableUuid="F0B1B2423AD169B5D41C14685AACA9A1">3</tableCitation>
<figureCitationid="1ED5C191D18B194373069018689F6E0B"captionStart="Figure 3"captionStartId="F3"captionText="Figure 3. Trichomes in Mesoamerican Virola A Virola allenii (R. Aguilar 2224, MO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (O. Valverde 970, MO) D V. elongata (G. McPherson 13674, MO) E V. fosteri (G. McPherson 20913, MO) F V. guatemalensis (R. Cedillo 3349, MO) G V. koschnyi (R. E. Gereau et al. 3455, MO) H V. laevigata (R. Aguilar 2004, MO) I V. megacarpa (J. A. Duke 15261, MO) J V. montana (E. Bello 855, MO) K V. multiflora (R. Rueda et al. 8196, MO; inset, J. C. Sandino 3302, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers & H. Herrera 7056, MO) N V. sebifera (F. Almeda et al. 5553, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO)."figureDoi="10.3897/phytokeys.134.37979.figure3"httpUri="https://binary.pensoft.net/fig/349085"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure3">Fig. 3A, N</figureCitation>
, F, GH) with the central part of the trichome colourless (vs. reddish) and with more compressed lateral veins (0.6-1.1 cm vs. 1.2-1.8 cm of separation between veins) (
<figureCitationid="45EACAD8E1F1E95476D8268D93A3B0DD"captionStart="Figure 3"captionStartId="F3"captionText="Figure 3. Trichomes in Mesoamerican Virola A Virola allenii (R. Aguilar 2224, MO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (O. Valverde 970, MO) D V. elongata (G. McPherson 13674, MO) E V. fosteri (G. McPherson 20913, MO) F V. guatemalensis (R. Cedillo 3349, MO) G V. koschnyi (R. E. Gereau et al. 3455, MO) H V. laevigata (R. Aguilar 2004, MO) I V. megacarpa (J. A. Duke 15261, MO) J V. montana (E. Bello 855, MO) K V. multiflora (R. Rueda et al. 8196, MO; inset, J. C. Sandino 3302, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers & H. Herrera 7056, MO) N V. sebifera (F. Almeda et al. 5553, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO)."figureDoi="10.3897/phytokeys.134.37979.figure3"httpUri="https://binary.pensoft.net/fig/349085"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure3">Figs 3A</figureCitation>
<figureCitationid="1B35602011F0401118380A2A4B87B26A"captionStart="Figure 8"captionStartId="F8"captionText="Figure 8. Pattern of the lateral veins in Mesoamerican Virola A Virola allenii (K. Thomsen 1284, NO) B V. amistadensis (G. McPherson 8703, MO) C V. chrysocarpa (B. Hammel et al. 16864, MO) D V. elongata (M. Correa & R. L. Dressler 1078, MO) E V. fosteri (G. de Nevers 7226, MO) F V. guatemalensis (G. Ibarra 957, MO) G V. koschnyi (J. Miller & J. C. Sandino 1110, MO) H V. laevigata (N. Zamora & T. D. Pennington 1583, MO) I V. megacarpa (G. de Nevers 5184, MO) J V. montana (E. Lepiz & J. F. Morales 284, MO) K V. multiflora (J. Manzanares 3561, MO) L V. nobilis from Barro Colorado (R. J. Schmalzel 320, MO) M V. otobifolia (G. de Nevers et al. 7530, MO) N V. sebifera (T. B. Croat 5959, MO) O V. nobilis from Osa Peninsula (R. Aguilar 11186, MO)."figureDoi="10.3897/phytokeys.134.37979.figure8"httpUri="https://binary.pensoft.net/fig/349090"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure8">8A, F</figureCitation>
<taxonomicNameid="2D0948ECBE11C06263FB722C9447AC3F"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<taxonomicNameid="52D5F5C22CE4EB6A5DE0A5429C2ABBCC"authorityName="D. Santam. & Aguilar"authorityYear="2019"class="Magnoliopsida"family="Myristicaceae"genus="Virola"higherTaxonomySource="CoL"kingdom="Plantae"lsidName="Virola allenii"order="Magnoliales"pageId="0"pageNumber="1"phylum="Tracheophyta"rank="species"species="allenii">
<figureCitationid="DD2F2BFF43D7CFA749A567DFC3D3906E"captionStart="Figure 4"captionStartId="F4"captionText="Figure 4. Fruits of Mesoameican Virola, organised in groups of morphologically similar species A V. allenii (P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9715, MO) C V. otobifolia (A. Gentry & S. Mori 14199, MO) D V. macrocarpa (A. E. Lawrance 675, MO) E V. chrysocarpa (B. Hammel et al. 16864, MO) F V. koschnyi (R. Robles 1587, MO) G V. guatemalensis (S. Sinaca 1542, MO) H V. montana (J. Utley & K. Utley 5054, MO) I V. fosteri (S. Mori & J. Kallunki 4891, MO) J V. multiflora (R. Rueda et a. l 9960, MO) K V. sebifera (T. B. Croat 5959, MO) L V. elongata (J. D. Dwyer & M. Correa 8420, MO) M V. laevigata (R. Aguilar 1585, MO) N V. nobilis from Barro Colorado (T. B. Croat 8090, MO) O V. nobilis from the Osa Peninsula (G. Herrera 4222, MO) P V. megacarpa (G. de Nevers 5184, MO). Scale bars: 1 cm."figureDoi="10.3897/phytokeys.134.37979.figure4"httpUri="https://binary.pensoft.net/fig/349086"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure4">Fig. 4A</figureCitation>
<figureCitationid="4E23E2E4A2B24025217A52C966EA7415"captionStart="Figure 4"captionStartId="F4"captionText="Figure 4. Fruits of Mesoameican Virola, organised in groups of morphologically similar species A V. allenii (P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9715, MO) C V. otobifolia (A. Gentry & S. Mori 14199, MO) D V. macrocarpa (A. E. Lawrance 675, MO) E V. chrysocarpa (B. Hammel et al. 16864, MO) F V. koschnyi (R. Robles 1587, MO) G V. guatemalensis (S. Sinaca 1542, MO) H V. montana (J. Utley & K. Utley 5054, MO) I V. fosteri (S. Mori & J. Kallunki 4891, MO) J V. multiflora (R. Rueda et a. l 9960, MO) K V. sebifera (T. B. Croat 5959, MO) L V. elongata (J. D. Dwyer & M. Correa 8420, MO) M V. laevigata (R. Aguilar 1585, MO) N V. nobilis from Barro Colorado (T. B. Croat 8090, MO) O V. nobilis from the Osa Peninsula (G. Herrera 4222, MO) P V. megacarpa (G. de Nevers 5184, MO). Scale bars: 1 cm."figureDoi="10.3897/phytokeys.134.37979.figure4"httpUri="https://binary.pensoft.net/fig/349086"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure4">Fig. 4B</figureCitation>
<figureCitationid="33C638E2C591C821E69DAEB89DD4DA68"captionStart="Figure 4"captionStartId="F4"captionText="Figure 4. Fruits of Mesoameican Virola, organised in groups of morphologically similar species A V. allenii (P. H. Allen 6727, GH) B V. amistadensis (G. McPherson 9715, MO) C V. otobifolia (A. Gentry & S. Mori 14199, MO) D V. macrocarpa (A. E. Lawrance 675, MO) E V. chrysocarpa (B. Hammel et al. 16864, MO) F V. koschnyi (R. Robles 1587, MO) G V. guatemalensis (S. Sinaca 1542, MO) H V. montana (J. Utley & K. Utley 5054, MO) I V. fosteri (S. Mori & J. Kallunki 4891, MO) J V. multiflora (R. Rueda et a. l 9960, MO) K V. sebifera (T. B. Croat 5959, MO) L V. elongata (J. D. Dwyer & M. Correa 8420, MO) M V. laevigata (R. Aguilar 1585, MO) N V. nobilis from Barro Colorado (T. B. Croat 8090, MO) O V. nobilis from the Osa Peninsula (G. Herrera 4222, MO) P V. megacarpa (G. de Nevers 5184, MO). Scale bars: 1 cm."figureDoi="10.3897/phytokeys.134.37979.figure4"httpUri="https://binary.pensoft.net/fig/349086"pageId="0"pageNumber="1"tableDoi="10.3897/phytokeys.134.37979.figure4">Fig. 4C</figureCitation>
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) (
<bibRefCitationid="A932FCD0A342735CDA63124CBC9F3F48"DOI="https://doi.org/10.12705/Code.2018"author="Turland, NJ"journalOrPublisher="Scientific Reports"pageId="0"pageNumber="1"refId="B105"refString="Turland, NJ, Wiersema, JH, Barrie, FR, Greuter, W, Hawksworth, DL, Herendeen, PS, Knapp, S, Kusber, W-H, Li, D-Z, Marhold, K, May, TW, McNeill, J, Monro, AM, Prado, J, Price, MJ, Smith, GF, 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashuetten. https://doi.org/10.12705/Code.2018"title="International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashuetten."url="https://doi.org/10.12705/Code.2018"year="2018">Turland et al. 2018</bibRefCitation>
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
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
) 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
<bibRefCitationid="4BDC50CB8EB89DFA37E2A89E6D13B7F1"author="Jimenez, Q"editor="Hammel, BE"journalOrPublisher="Monographs in Systematic Botany from the Missouri Botanical Garden"pageId="0"pageNumber="1"pagination="684 - 691"refId="B46"refString="Jimenez, Q, 2007. Myristicaceae. In: Hammel, BE, Grayum, MH, Herrera, C, Zamora, N, Eds., Manual de Plantas de Costa Rica. Vol. VI. Monographs in Systematic Botany from the Missouri Botanical Garden 111: 684 - 691"title="Myristicaceae."volume="111"volumeTitle="Manual de Plantas de Costa Rica. Vol. VI."year="2007">
<bibRefCitationid="CDC830E1B5845573D04A26A31814E501"DOI="https://doi.org/10.1038/s41598-019-40101-y"author="ter Steege, H"journalOrPublisher="Scientific Reports"pageId="0"pageNumber="1"pagination=": 3501"refId="B103"refString="ter Steege, H, Mota de Oliveira, S, Pitman, NCA, Sabatier, D, Antonelli, A, Guevara Andino, JE, Aymard, GA, Salomao, RP, 2019. Towards a dynamic list of Amazonian tree species. Scientific Reports 9 (1): 3501, DOI: https://doi.org/10.1038/s41598-019-40101-y"title="Towards a dynamic list of Amazonian tree species."url="https://doi.org/10.1038/s41598-019-40101-y"volume="9"year="2019">ter Steege et al. 2019</bibRefCitation>
). However, here, it is treated as a morphologically distinct species. This is due to its smaller leaves, more compact staminate and pistillate inflorescences and infructescences, staminate flowers with the filament column longer than the anthers and smaller fruits (see Table
<tableCitationid="64DAA915CCA82382C594DB82BA76E50B"captionStart="Table 2"captionStartId="T2"captionText="Table 2. Comparison of Virola allenii with the morphologically most similar species."httpUri="http://table.plazi.org/id/1A2AE0917E035E6F6ACDB17A2E796F2C"pageId="0"pageNumber="1"tableUuid="1A2AE0917E035E6F6ACDB17A2E796F2C">2</tableCitation>
