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<document ID-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 &amp; Lagomarsino, Laura P." docDate="2019" docId="CA381C53D596526F974179D43FC36DB2" docLanguage="en" docName="PhytoKeys 134: 1-82" docOrigin="PhytoKeys 134" docSource="http://dx.doi.org/10.3897/phytokeys.134.37979" docTitle="Virola laevigata Standl." 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">
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<mods:title id="CF095DC7D70AF54FA53C70C85D75AF2F">A taxonomic synopsis of Virola (Myristicaceae) in Mesoamerica, including six new species</mods:title>
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<mods:namePart id="61A27721FA6A44B600A4E3A350F6D225">Santamaria-Aguilar, Daniel</mods:namePart>
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<mods:namePart id="3B058EF42E4383A60F803E12978D3B69">Aguilar, Reinaldo</mods:namePart>
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<mods:namePart id="B30C53A402DDC7C362980B4C5FB0D3A2">Lagomarsino, Laura P.</mods:namePart>
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<treatment id="CA381C53D596526F974179D43FC36DB2" ID-GBIF-Taxon="159834608" LSID="urn:lsid:plazi:treatment:CA381C53D596526F974179D43FC36DB2" httpUri="http://treatment.plazi.org/id/CA381C53D596526F974179D43FC36DB2" lastPageNumber="1" pageId="0" pageNumber="1" scope_family="Myristicaceae">
<subSubSection id="124D368E223F43F23E39E960CBE8634A" pageId="0" pageNumber="1" type="nomenclature">
<paragraph id="42D211FB9E20DD3606DBFA4AC6A24942" pageId="0" pageNumber="1">
8.
<taxonomicName id="E100CB0DB6F72969C82EE96306069E05" ID-CoL="5BK53" LSID="CA381C53-D596-526F-9741-79D43FC36DB2" authority="Standl." class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="laevigata">Virola laevigata Standl.</taxonomicName>
<figureCitation id="38DBBEE8EFC41CD3505E5830A0C11E0A" captionStart="Figure 19" captionStartId="F19" captionText="Figure 19. Virola laevigata A branch with staminate inflorescences B leaf blades on abaxial surface, inset showing the glabrous midvein C twig, petiole and leaf on adaxial surface D close up of staminate inflorescences E leaf base F leaf apex, inset showing leaf punctations G branch with immature fruits H close-up of immature fruit I open fruit, showing the aril. Photos by Reinaldo Aguilar." figureDoi="10.3897/phytokeys.134.37979.figure19" httpUri="https://binary.pensoft.net/fig/349101" pageId="0" pageNumber="1" tableDoi="10.3897/phytokeys.134.37979.figure19">Fig. 19</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="7ABB40BDCD55B123C7F3C308141D588A" pageId="0" pageNumber="1" type="reference_group">
<paragraph id="E8385A6D5F17BCA2F875A7AC22A6A40E" pageId="0" pageNumber="1">
<taxonomicName id="96873DBAC465A3ADFA335F2BCD15DDE9" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="laevigata">Virola laevigata</taxonomicName>
Standl. Publ. Field Mus. Nat. Hist., Bot. Ser. 4(8): 209. 1929. Type. Panama. Province of
<normalizedToken id="E45DF218E4D5FF59F001595891E5BA28" originalValue="Chiriquí">Chiriqui</normalizedToken>
, Progreso, [
<normalizedToken id="CCABF832EDFDC2F76FDA60836393511A" originalValue="July–Aug">July-Aug</normalizedToken>
.] 1927 [♂ fl],
<emphasis id="7ED65B7F9057430E5EB78F30BA7A9974" italics="true" pageId="0" pageNumber="1">G. P. Cooper &amp; G. M. Slater 308</emphasis>
(holotype: F!*; isotypes: NY!*, WIS!*, US!*).
</paragraph>
</subSubSection>
<subSubSection id="0E2A4AF3D78618F3980E6175B0D199A3" pageId="0" pageNumber="1" type="description">
<paragraph id="D045FEC519013F3DAAF963AD40CBF633" pageId="0" pageNumber="1">Distinctive characters.</paragraph>
<paragraph id="1210F8D31AD21103022C704E3AF4D867" pageId="0" pageNumber="1">
<taxonomicName id="0A0927E31D78FA13AD880E50A8E49859" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="laevigata">
<emphasis id="24E42FDAD5150B96D5B8B54FCC33CE5D" italics="true" pageId="0" pageNumber="1">Virola laevigata</emphasis>
</taxonomicName>
is distinguished by its glabrous or nearly glabrous vegetative parts (i.e. twigs, mature leaf blades on both surfaces [
<figureCitation id="D0E3835D9BDDA5391BD20162112A8322" 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 &amp; 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. 3H</figureCitation>
], petioles); when trichomes are present, they are primarily on the leaf buds or very new leaves. Additionally, leaves have 12-20 lateral veins and margins and bases that are slightly revolute, staminate flowers have a straight filament column that is longer (0.8-1.3 mm) than the anthers (0.5-0.7 mm) and relatively small fruits (1.8-2.9
<normalizedToken id="FDC7177C9BB7D371A6D69B3639A94F89" originalValue="×">x</normalizedToken>
1.5-1.8 cm) (
<figureCitation id="67EE7F4431E0758BBF4D1103F9B87DD8" 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 &amp; 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 &amp; K. Utley 5054, MO) I V. fosteri (S. Mori &amp; 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 &amp; 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. 4M</figureCitation>
) with pericarp that is 1.8-2.8 mm thick.
</paragraph>
</subSubSection>
<subSubSection id="B1806037BDDD582E3107E35DF3248545" pageId="0" pageNumber="1" type="distribution">
<paragraph id="0796B50BBEF9FD4BA926279310A0CCFC" pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph id="76AA02D77314039E2465008C40555B77" pageId="0" pageNumber="1">
<taxonomicName id="E2D5E98D7E5AFF62B9928F1BDC2870F3" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="laevigata">
<emphasis id="B9D5E41F11F83479050D69F1632D1B72" italics="true" pageId="0" pageNumber="1">Virola laevigata</emphasis>
</taxonomicName>
is known from Costa Rica (Puntarenas and San
<normalizedToken id="F88D40320C783DAD9267CE5F2D19657A" originalValue="José">Jose</normalizedToken>
) and Panama (
<normalizedToken id="E395684D3D0E451C4C94B5D9C37BD1C7" originalValue="Chiriquí">Chiriqui</normalizedToken>
) (
<figureCitation id="7822DEE50375711813C81C5D86A10AB4" captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Geographic distribution of Virola koschnyi (A), V. laevigata (B), V. megacarpa (C), V. montana (D), V. multiflora (E) and V. nobilis (F)." figureDoi="10.3897/phytokeys.134.37979.figure18" httpUri="https://binary.pensoft.net/fig/349100" pageId="0" pageNumber="1" tableDoi="10.3897/phytokeys.134.37979.figure18">Fig. 18B</figureCitation>
). It is found on the Pacific slope, where it has been recorded between 0-500 (1600?) m elevation.
<bibRefCitation id="44BA84E4474EDEB53E8490E6CC4B8070" 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">
<normalizedToken id="A5CA3B6A62E94336D7964B5008FFC71B" originalValue="Jiménez">Jimenez</normalizedToken>
(2007)
</bibRefCitation>
suggested that the maximum elevation for this species in Costa Rica is 1600 m, potentially based on
<emphasis id="EE988ADB0CB485DE7D8B9EF9C003500F" italics="true" pageId="0" pageNumber="1">
L.
<normalizedToken id="4D40FBDEF1617056CA34AC4803242AF8" originalValue="González">Gonzalez</normalizedToken>
3089
</emphasis>
from Cerro Turrubares (San
<normalizedToken id="0EB6F0F07FAA844BC67E132AA2F7466E" originalValue="José">Jose</normalizedToken>
province) (B. Hammel pers. comm., Aug. 2019); however, we have not found any specimen that occurs this high.
</paragraph>
</subSubSection>
<subSubSection id="897AFC4B137C107A2AA894B289B21451" pageId="0" pageNumber="1" type="common names">
<paragraph id="A3A24189E5F7D5A352E7305C9E52840E" pageId="0" pageNumber="1">Common names.</paragraph>
<paragraph id="F52C536CE7BD302508EB1676E3A6758F" pageId="0" pageNumber="1">Costa Rica: fruta dorada. Panama: bogamani.</paragraph>
</subSubSection>
<subSubSection id="85B36FEE0B7917FBA6675BC14DAFFB31" pageId="0" pageNumber="1" type="phenology">
<paragraph id="3D0E0CF197B2A66B9BB58122C6D2DB8E" pageId="0" pageNumber="1">Phenology.</paragraph>
<paragraph id="37F9532E495AF395A8EB89858537AF17" pageId="0" pageNumber="1">
Flowering of
<taxonomicName id="4290DF5A6AC9910633EB8FE45CB56812" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="laevigata">
<emphasis id="C8A4E98C644004809F27F428050D794B" italics="true" pageId="0" pageNumber="1">Virola laevigata</emphasis>
</taxonomicName>
has been recorded in January, May, July and November. Fruits are produced from December to February. Pistillate flowers were not present on herbarium sheets studied.
</paragraph>
<caption id="E5DA014F5FE14A2872802952D536DEC2" doi="10.3897/phytokeys.134.37979.figure19" httpUri="https://binary.pensoft.net/fig/349101" pageId="0" pageNumber="1" start="Figure 19" startId="F19">
<paragraph id="A564A9614C5C76B44AD2E5A4E706F23E" pageId="0" pageNumber="1">
<emphasis id="7D0CB2FF8F60FC146A39ECB40BA3B43D" bold="true" pageId="0" pageNumber="1">Figure 19.</emphasis>
<taxonomicName id="4BBA8643C4DEF01FDBA6C77D29C37C0F" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="laevigata">
<emphasis id="ECB175D901A0F0F831FC18D9DD3C68AD" italics="true" pageId="0" pageNumber="1">Virola laevigata</emphasis>
</taxonomicName>
<emphasis id="69E156EE7970EE7C1F73EC97CA91E0AC" bold="true" pageId="0" pageNumber="1">A</emphasis>
branch with staminate inflorescences
<emphasis id="0B53B144C734FDE5276F49D4F21E7FE5" bold="true" pageId="0" pageNumber="1">B</emphasis>
leaf blades on abaxial surface, inset showing the glabrous midvein
<emphasis id="AFC50FD464B3D4CB70AD6AE56A2021DF" bold="true" pageId="0" pageNumber="1">C</emphasis>
twig, petiole and leaf on adaxial surface
<emphasis id="F5955E5B034129D11F538CE395652CC5" bold="true" pageId="0" pageNumber="1">D</emphasis>
close up of staminate inflorescences
<emphasis id="D5E20615EB7495BBBDBD7A67FBB5231F" bold="true" pageId="0" pageNumber="1">E</emphasis>
leaf base
<emphasis id="E57EAEB415C8373FC6EE9BF98992EDDA" bold="true" pageId="0" pageNumber="1">F</emphasis>
leaf apex, inset showing leaf punctations
<emphasis id="2FBAD852EA0EC5CD5BC01DA183723590" bold="true" pageId="0" pageNumber="1">G</emphasis>
branch with immature fruits
<emphasis id="DD9CCF3CC46676DBB3EFB4825ABF95B3" bold="true" pageId="0" pageNumber="1">H</emphasis>
close-up of immature fruit
<emphasis id="E3411083D78239901771499B600A5892" bold="true" pageId="0" pageNumber="1">I</emphasis>
open fruit, showing the aril. Photos by Reinaldo Aguilar.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="9C7754A26559B4CF8FC8910F8335976B" pageId="0" pageNumber="1" type="description">
<paragraph id="98DA1BB790A3240B0C2E5F9BEF830DFF" pageId="0" pageNumber="1">Field characters.</paragraph>
<paragraph id="D700C50A3AB3A76366D2621A56D9AD4B" pageId="0" pageNumber="1">
Plants are trees 9-40 m tall and 35-60 cm DBH with a straight trunk and small (ca. 20 cm tall), triangular buttresses. The bark is described as finally grooved, smooth, flaking in vertical strips or scaly and is grey, blackish or reddish in colour, with exudate that is reddish or colourless and oxidising to reddish-cream. The leaves are bright green on both sides and have numerous pellucid dots that are most visible against the light. Flowers have yellow, yellow-brown or yellowish perianth, sometimes with a slight aroma in staminate flowers (
<emphasis id="557BF1F67A8DDBE2947FB5352C8FFD72" italics="true" pageId="0" pageNumber="1">N.</emphasis>
Z
<emphasis id="2FB732603B5898F04B6ABDABE5B0AAEE" italics="true" pageId="0" pageNumber="1">amora &amp; T. D. Pennington 1583</emphasis>
, but the specimen label states pistillate flower). The mature fruit is yellow with a red aril (when immature, it is white). In the Osa Peninsula, where this species is frequent, it prefers riparian habitats.
</paragraph>
</subSubSection>
<subSubSection id="740DBF4E7973804999038BC591782F8D" pageId="0" pageNumber="1" type="discussion">
<paragraph id="7F80565766E0A60E31F906E7C27463E9" pageId="0" pageNumber="1">Discussion.</paragraph>
<paragraph id="F3011EF778C6F6B7C2DB184ECD9DEA43" pageId="0" pageNumber="1">
<taxonomicName id="8B14B9AE81A6FA5756847CB81A897729" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="laevigata">
<emphasis id="4535E101013C29092AFB0E79DA3CCFFE" italics="true" pageId="0" pageNumber="1">Virola laevigata</emphasis>
</taxonomicName>
has traditionally been considered a synonym of
<taxonomicName id="7E19BA7DFC23A39AFA59AE948341D3FF" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. guatemalensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="guatemalensis">
<emphasis id="FFB6BCDFCCE3F7E1783E57DC609C64F1" italics="true" pageId="0" pageNumber="1">V. guatemalensis</emphasis>
</taxonomicName>
(e.g.
<bibRefCitation id="C217881A5DE5A0EC03835E9088145D4C" DOI="https://doi.org/10.2307/2804799" author="Smith, AC" journalOrPublisher="Brittonia" pageId="0" pageNumber="1" pagination="393 - 510" refId="B90" refString="Smith, AC, Wodehouse, RP, 1938. The American species of Myristicaceae. Brittonia 2 (5): 393 - 510, DOI: https://doi.org/10.2307/2804799" title="The American species of Myristicaceae." url="https://doi.org/10.2307/2804799" volume="2" year="1938">Smith and Wodehouse 1938</bibRefCitation>
;
<bibRefCitation id="E637DB2F403092A71CB4F9E2A9B2C22E" author="Standley, PC" editor="Standley, PC" journalOrPublisher="Fieldiana: Botany 24 (4)" pageId="0" pageNumber="1" pagination="294 - 299" refId="B99" refString="Standley, PC, Steyermark, JA, 1946. Myristicaceae. In: Standley, PC, Steyermark, JA, Eds., Flora of Guatemala - Part IV. Fieldiana: Botany 24(4): 294 - 299" title="Myristicaceae." volumeTitle="Flora of Guatemala - Part IV." year="1946">Standley and Steyermark 1946</bibRefCitation>
;
<bibRefCitation id="BB3C32D8ABD0D6981A880258D0920C43" DOI="https://doi.org/10.2307/2394708" author="Duke, JA" journalOrPublisher="Annals of the Missouri Botanical Garden" pageId="0" pageNumber="1" pagination="214 - 225" refId="B24" refString="Duke, JA, 1962. Myristicaceae. Flora of Panama, Part IV. Fascicle 5. Annals of the Missouri Botanical Garden 49 (3-4): 214 - 225, DOI: https://doi.org/10.2307/2394708" title="Myristicaceae. Flora of Panama, Part IV. Fascicle 5." url="https://doi.org/10.2307/2394708" volume="49" year="1962">Duke 1962</bibRefCitation>
), likely due to limited material. However, with new herbarium specimens, both species can be clearly distinguished vegetatively and with fruit characters. Vegetative material of
<taxonomicName id="5F93761ECFF594A2CE8CD94EAD654130" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="03395FF94CF9C2E1DD6AFED705D202A5" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
can be distinguished by petioles and mature leaf blades that are abaxially glabrous (vs. diminutively pubescent with tiny stellate, sessile trichomes in
<taxonomicName id="3911201F823575CE7477EB6D3DC9CDCD" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. guatemalensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="guatemalensis">
<emphasis id="3930176D0BF21BF31C0159F3ABC30CA6" italics="true" pageId="0" pageNumber="1">V. guatemalensis</emphasis>
</taxonomicName>
). Their fruits also differ in size, shape and thickness of the pericarp; in
<taxonomicName id="C76CE2A1B39487E2CEAE98807EE7B6D7" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="CC985207670EBCFA6916DCAE2C8F631D" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
, they are smaller (1.8-2.9
<normalizedToken id="A0F8FE0D82825C243F1A01AEC6A10E4E" originalValue="×">x</normalizedToken>
1.5-1.8 cm), ovoid or subglobose and with thick pericarp (1.8-2.8 mm); while in
<taxonomicName id="12BE23F65427AE4856C9B101DBD508F7" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. guatemalensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="guatemalensis">
<emphasis id="4888287AF9FF273204A3FEA92CE0EB1E" italics="true" pageId="0" pageNumber="1">V. guatemalensis</emphasis>
</taxonomicName>
, they are large (vs. 2.7-3.4
<normalizedToken id="44EC14F1A474D9279D7FB1988BBF8746" originalValue="×">x</normalizedToken>
1.7-2.3 cm), ellipsoid and with a thin pericarp [0.4-1 (-2.5) mm].
</paragraph>
<paragraph id="4551F013BA6B2F77FD482A4930D33B2B" pageId="0" pageNumber="1">
In addition to
<taxonomicName id="75C428586D61A912D06B4D59E25B24CC" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola guatemalensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="species" species="guatemalensis">
<emphasis id="96A703C28B817CB2C57CB7E9370BAD3B" italics="true" pageId="0" pageNumber="1">Virola guatemalensis</emphasis>
</taxonomicName>
, herbarium specimens of
<taxonomicName id="6C4F1FF0637314A0137DC329353BDDC7" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="139EF37C7410D8716E83415AC1608D8A" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
have been determined as
<taxonomicName id="4152DE05849592CC719C2DFAB8628BA9" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. surinamensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="surinamensis">
<emphasis id="CD32B7F18B74AAC2D9884A3BD605DA13" italics="true" pageId="0" pageNumber="1">V. surinamensis</emphasis>
</taxonomicName>
(interpreted here as
<taxonomicName id="48D6DDD230951C185944496E0E56CB1D" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. nobilis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="nobilis">
<emphasis id="B55A53C557DD2423180F08DE5B7EEFC5" italics="true" pageId="0" pageNumber="1">V. nobilis</emphasis>
</taxonomicName>
). However,
<taxonomicName id="4FB25035C008FB517EF92485502E8303" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="04C05AF6C25F10A35860B6A1299A5FBB" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
is distinguished by its glabrous or almost glabrous abaxial leaf surface (
<figureCitation id="2D22A49DA6F88DE4CF1FAA2D35BCB200" 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 &amp; 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. 3H, O</figureCitation>
) and mature fruits (vs. pubescent) and its tendency towards thinner pericarp [1.8-2.8 mm vs. 2.5-3.5 (-4.2) mm thick]. In the Osa Peninsula, where these two species grow together, they can be easily distinguished in the field:
<taxonomicName id="CD664AE34E6EEF453681B7D9C8098F5F" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="C010A670E4E14B28806E567E9DD1691F" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
prefers riparian habitats, does not usually develop tall buttresses and the external bark has a greenish tone, while
<taxonomicName id="08D190E407E2DD6DB323979003AAFE70" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. nobilis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="nobilis">
<emphasis id="D2FAA5EB600FA4BE64B04CB5532CC7C9" italics="true" pageId="0" pageNumber="1">V. nobilis</emphasis>
</taxonomicName>
grows far from bodies of water, has tall buttresses and the external bark is reddish to brown. For a description and comparison of the bark of these two species, see
<bibRefCitation id="4EE184F68C147F3A7951318C626C04A6" author="Moya Roque, R" journalOrPublisher="Editorial Tecnologico de Costa Rica, Cartago" pageId="0" pageNumber="1" refId="B59" refString="Moya Roque, R, Rodriguez Gonzalez, A, Olivares Gutierrez, C, 2014. Arboles maderables de la Peninsula de Osa, madera y corteza. Editorial Tecnologico de Costa Rica, Cartago" title="Arboles maderables de la Peninsula de Osa, madera y corteza." year="2014">Moya Roque et al. (2014)</bibRefCitation>
, as
<taxonomicName id="3DE427D605E7E7025604FC85E6E4DEA0" authorityName="Aublet" authorityYear="1775" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="genus">
<emphasis id="3ADC0C62097EFABE12BF74820F9BD0C4" italics="true" pageId="0" pageNumber="1">Virola</emphasis>
</taxonomicName>
sp. A and
<taxonomicName id="B06CAB28B39595A081AD2C465C8DEE3A" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. surinamensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="surinamensis">
<emphasis id="532A908B3CA0C0580C92EC47533D6853" italics="true" pageId="0" pageNumber="1">V. surinamensis</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="B0B9DB0F732CFDFF848BB9EF66FCA71B" pageId="0" pageNumber="1" type="notes">
<paragraph id="770DC48075B0EFAF0B71EF5A0EBAC1B1" pageId="0" pageNumber="1">Notes.</paragraph>
<paragraph id="9871B3FFB8FCF629BA771017A6FDAAAF" pageId="0" pageNumber="1">
The seedlings of
<taxonomicName id="4F7574156893F0CE5356DF3861F8AAA7" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="2778F0117AFFF132FC2C4C75551FB607" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
are described by
<bibRefCitation id="5078C678EE753A6F63048F0694C24588" 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">
Ley
<normalizedToken id="FBF4968EE8CEC389F9FD90AEE979463A" originalValue="López">Lopez</normalizedToken>
and
<normalizedToken id="E33D0116B439DCE119CBEC3A6AA5C175" originalValue="Chacón">Chacon</normalizedToken>
Madrigal (2017)
</bibRefCitation>
(though as &quot;
<taxonomicName id="20DAA9206BAB97B2483AF35E79A91CB2" authorityName="Aublet" authorityYear="1775" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="genus">Virola</taxonomicName>
sp. A&quot;). Additionally, the species presented as &quot;
<taxonomicName id="84EE38583864448D39F9A81A0535B430" authorityName="Aublet" authorityYear="1775" class="Magnoliopsida" family="Myristicaceae" genus="Virola" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Virola" order="Magnoliales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="genus">Virola</taxonomicName>
sp. A&quot; in
<bibRefCitation id="1F7BCF6714E18C75BDD3440AE7EA32B7" 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">
<normalizedToken id="A7C4AD78B060D38F1F848E03F9620ECE" originalValue="Jiménez">Jimenez</normalizedToken>
(2007)
</bibRefCitation>
and as
<taxonomicName id="AA71A25A7638066137D59481810C4281" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. surinamensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="surinamensis">
<emphasis id="0A426CEADA7CEB60FF0B0AA38F2CA5CB" italics="true" pageId="0" pageNumber="1">V. surinamensis</emphasis>
</taxonomicName>
in
<bibRefCitation id="DA60BF2DA50B2A1CB3156F84A3341734" author="Jimenez Madrigal, Q" journalOrPublisher="Monographs in Systematic Botany from the Missouri Botanical Garden" pageId="0" pageNumber="1" refId="B47" refString="Jimenez Madrigal, Q, Grayum, MH, 2002. Vegetacion del Parque Nacional Carara, Costa Rica. Brenesia 57-58: 25-66." title="Vegetacion del Parque Nacional Carara, Costa Rica. Brenesia 57 - 58: 25 - 66." year="2002">
<normalizedToken id="82714CB00F444D900F0F104EDA14B97A" originalValue="Jiménez">Jimenez</normalizedToken>
Madrigal and Grayum (2002
</bibRefCitation>
; RZ [=
<emphasis id="759F6ED21954093CC2EE407A31132520" italics="true" pageId="0" pageNumber="1">
R.
<normalizedToken id="81A1893EFC12B32FAC43B22E14BF0BA1" originalValue="Zuñiga">Zuniga</normalizedToken>
</emphasis>
] 459) corresponds with
<taxonomicName id="148DE47FB3E31F8D6E456800BBD568A0" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="6761B9C5B3BCADD5DD0FA7D17861FB25" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
. The illustration in
<bibRefCitation id="D31A217100B5157546261A7E028D3884" author="Quesada Quesada, FJ" journalOrPublisher="Economic Botany" pageId="0" pageNumber="1" refId="B71" refString="Quesada Quesada, FJ, Jimenez Madrigal, Q, Zamora Villalobos, N, Aguilar Fernandez, F, Gonzalez Ramirez, J, 1997. Arboles de la Peninsula de Osa. INBio, Santo Domingo de Heredia, 1-411." title="Arboles de la Peninsula de Osa. INBio, Santo Domingo de Heredia, 1 - 411." year="1997">Quesada Quesada et al. (1997)</bibRefCitation>
as
<taxonomicName id="0F6B5E9825E541CE8B848BB50F96CC61" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. guatemalensis" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="guatemalensis">
<emphasis id="DF883A06C0E19EF1CC8047AADE2F658D" italics="true" pageId="0" pageNumber="1">V. guatemalensis</emphasis>
</taxonomicName>
is potentially also
<taxonomicName id="099C177303C2BCE1FE64AF3982DA3347" class="Dicotyledoneae" family="Myristicaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="V. laevigata" order="Magnoliales" pageId="0" pageNumber="1" phylum="Angiospermae" rank="species" species="laevigata">
<emphasis id="DAE9A0B85C952E7B598BF35F79EF1EED" italics="true" pageId="0" pageNumber="1">V. laevigata</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="EEE421A1345F15DB14268E230E465777" pageId="0" pageNumber="1" type="selected specimens seen">
<paragraph id="C1830ED6E9C47BE90DE17C56E00F2EC8" pageId="0" pageNumber="1">Selected specimens seen.</paragraph>
<paragraph id="C5189AAC6CEFABEE04A620014F53A1CB" pageId="0" pageNumber="1">
<emphasis id="0C2B10CEEFCDBF7E549099FA9B01BA64" bold="true" pageId="0" pageNumber="1">Costa Rica. Puntarenas</emphasis>
: Golfito, 1 km antes de llegar a La Palma, 8 m elev., 16 Jan 1993 (fr),
<emphasis id="1AFFC50C6DD2380EBC6655B01B2478D4" italics="true" pageId="0" pageNumber="1">R. Aguilar 1585</emphasis>
(CR-2 sheets!, LSU!, MO!); fila Carbonera, cabo Matapalo, 300 m elev., 16 Jul 1993 (♂ fl),
<emphasis id="2212201088AF744B6BA2279E46CD305A" italics="true" pageId="0" pageNumber="1">R. Aguilar 2004</emphasis>
(CR!, MO!);
<normalizedToken id="91FDBDA7F2CF4A501F86181EB1D47398" originalValue="estación">estacion</normalizedToken>
Sirena, 10 m elev., 12 Oct 1993 (fl bud, fr),
<emphasis id="3D29ABBDD7B9D5A41D20EDCEA29C8490" italics="true" pageId="0" pageNumber="1">R. Aguilar 2490</emphasis>
(CR-2 sheets!, LSU!, MO!); Golfito, orillas del camino a las torres de
<normalizedToken id="595DCDB3D7C47CCFE11F6AC11BE461F1" originalValue="comunicación">comunicacion</normalizedToken>
, 450-500 m elev., 12 Jan 1999 (fr),
<emphasis id="0BF8508BEF205E259A3175BFD4A0FEB2" italics="true" pageId="0" pageNumber="1">
J.
<normalizedToken id="F0AD070A2E8CC8E621CC2A837627D621" originalValue="Gómez-Laurito">Gomez-Laurito</normalizedToken>
&amp; V. Mora 13191
</emphasis>
(CR!); Playa Cacao, Fila entre quebrada Nazanero y el mar en Punta Voladora, 200 m elev., 26 May 1994 (♂ fl),
<emphasis id="D0A51B442D4FBEED0CDCB55D3775264C" italics="true" pageId="0" pageNumber="1">G. Herrera &amp; G. Rivera 7076</emphasis>
(CR!); Parque Nacional Corcovado, Sirena, 10 m elev., 28 Feb 1989 (fr),
<emphasis id="BFC791060E05C7D2BB98B6DFCBFB44E7" italics="true" pageId="0" pageNumber="1">C. Kernan &amp; P. Phillips 962</emphasis>
(CR-2 sheets!, MO!); Parque Nacional Corcovado, Sirena, Ollas Trail, 1-20 m elev., 19 Oct 1989 (fr),
<emphasis id="5488CD725C17E90723D2F1BB55FB80CD" italics="true" pageId="0" pageNumber="1">C. Kernan &amp; G. Fonseca 1288</emphasis>
(CR-2 sheets!, INPA!*, MO!);
<normalizedToken id="58D57A78D78D7052898703ED6CFC4D6D" originalValue="Jiménez">Jimenez</normalizedToken>
, Piro, camino a Laguna Silvestre, [0-100 m elev.], 01 Feb 2012 (fr),
<emphasis id="4B1A65571BD7FFD6EBEF2D0AC595C6EF" italics="true" pageId="0" pageNumber="1">
J. M.
<normalizedToken id="BCA0FCCC1FEB962D3A45D54C6AA49A2A" originalValue="Ley-López">Ley-Lopez</normalizedToken>
74
</emphasis>
(USJ); Osa, fila Esquinas, 200 m elev., 26 May 1993 (♂ fl),
<emphasis id="5AAC4E53719AF7BED0BD71FBCC571079" italics="true" pageId="0" pageNumber="1">M. Segura &amp; F. Quesada 69</emphasis>
(CR!, LSU!, MO!);
<normalizedToken id="103FD5D670FDDE810E46A5E2F63C21CB" originalValue="Rincón">Rincon</normalizedToken>
de Osa, in vicinity of airstrip, 40 m elev., 25 Jul 1974 (fl bud),
<emphasis id="E774DA3A18DBE18685720743A24CECC1" italics="true" pageId="0" pageNumber="1">J. Utley &amp; K. Utley 1236</emphasis>
(CR!); Garabito, por Carara, cerca de la toma de agua, 97 m elev., 22 Nov 2006 (fl bud),
<emphasis id="ACA22B3239EA021E78A4E89F8C6DDC7B" italics="true" pageId="0" pageNumber="1">L. D. Vargas &amp; D. Castillo 1870</emphasis>
(CR!); Osa, Uvita, cerca de Dominical, 200 m elev., 28 Jan 1991 (♂ fl),
<emphasis id="CD2C63D38D31035793EBAFD1AB34F768" italics="true" pageId="0" pageNumber="1">N. Zamora &amp; T. D. Pennington 1583</emphasis>
(CR-2 sheets!, INPA!*, MEXU!*, MO!); Puntarenas, Montezuma, camino a
<normalizedToken id="D90CE3AD049E06E011D315DA92002D9E" originalValue="Cóbano">Cobano</normalizedToken>
por el
<normalizedToken id="E0FEEB1C5D1E40B661BC26FF2D71AF5F" originalValue="río">rio</normalizedToken>
Montezuma, ca. 1.5 km oeste de la
<normalizedToken id="39AD027FB90EC57929EC44A71C66184B" originalValue="intersección">interseccion</normalizedToken>
con camino a Cabuya, 101 m elev., 11 Dec 2005 (fr),
<emphasis id="D1DB3CCCC478E0CA57116EBCF55389FB" italics="true" pageId="0" pageNumber="1">
B. Hammel &amp; I.
<normalizedToken id="C66D209FA0028A80182455F86E4A02FC" originalValue="Pérez">Perez</normalizedToken>
23944
</emphasis>
(CR!); Montezuma, por el Canopy, 100 m elev., 08 Jul 2006 (fl bud),
<emphasis id="DA0D97CA23B3637C22FF8C7407B4D492" italics="true" pageId="0" pageNumber="1">
B. Hammel &amp; I.
<normalizedToken id="8DBAE3E5578AE04AC8433A7A2A9801C5" originalValue="Pérez">Perez</normalizedToken>
24149
</emphasis>
(CR!).
<emphasis id="ABFF1DA60C41FF1723C5517DA157DA4D" bold="true" pageId="0" pageNumber="1">
San
<normalizedToken id="FD397C4FE5D6406A800A8187D39F7800" originalValue="José">Jose</normalizedToken>
</emphasis>
: Carara, sector Agrominas, sitio Carretera Costanera, 100 m elev., 20 Sep 1991 (fr),
<emphasis id="60ABF41AFEABABC00C7D717A36B439A2" italics="true" pageId="0" pageNumber="1">
R.
<normalizedToken id="AAE0B0F0D88EE5CAA37DDF83F5C9010E" originalValue="Zuñiga">Zuniga</normalizedToken>
459
</emphasis>
(CR-sheets!, LSU!).
<emphasis id="1E7C1D83404AE7EAC2D98B0E059C5D29" bold="true" pageId="0" pageNumber="1">
Panama.
<normalizedToken id="251490087FBBEAFC86579DF956D1CBE4" originalValue="Chiriquí">Chiriqui</normalizedToken>
</emphasis>
:
<normalizedToken id="12B476E5ADE492B9A22806018DB57E72" originalValue="Río">Rio</normalizedToken>
Platanal-Bugaba, 10 Dec 1975 (fr),
<emphasis id="B56C2372E35B65A72B9A48BB77E7EF50" italics="true" pageId="0" pageNumber="1">
M. M.
<normalizedToken id="3B71C51657AC0EE1C41F8E9A694FA83C" originalValue="Gutiérrez">Gutierrez</normalizedToken>
21
</emphasis>
(MO!).
</paragraph>
</subSubSection>
</treatment>
</document>
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