488 lines
57 KiB
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488 lines
57 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.881.38026" ID-GBIF-Dataset="b69834db-1e77-4c25-80c7-f1fd3be2f508" ID-PMC="PMC6813178" ID-Pensoft-Pub="1313-2970-881-23" ID-Pensoft-UUID="0B1397AFEA7B5910A468BA7A24DA14DD" ID-PubMed="31662610" ID-ZooBank="ADCA057E3E1B45D8B5C9665683F51B59" ModsDocID="1313-2970-881-23" checkinTime="1571384721580" checkinUser="pensoft" docAuthor="Kohlmann, Bert, Solis, Angel & Alvarado, Guillermo E." docDate="2019" docId="4E574931E2725135B5A689A4E7DD70E0" docLanguage="en" docName="ZooKeys 881: 23-51" docOrigin="ZooKeys 881" docSource="http://dx.doi.org/10.3897/zookeys.881.38026" docTitle="Onthophagus humboldti Kohlmann, Solis & Alvarado, 2019, sp. nov." docType="treatment" docUuid="C13D9441-5A22-4CA1-974F-88B6A97199AC" docUuidSource="ZooBank" docVersion="5" id="0B1397AFEA7B5910A468BA7A24DA14DD" lastPageNumber="29" masterDocId="0B1397AFEA7B5910A468BA7A24DA14DD" masterDocTitle="Description of Onthophagus humboldti and Uroxys bonplandi, two new scarab beetles (Coleoptera, Scarabaeidae, Scarabaeinae) from Costa Rica, with notes on tropical mountain brachyptery and endemicity" masterLastPageNumber="51" masterPageNumber="23" pageNumber="25" updateTime="1668167899117" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Description of Onthophagus humboldti and Uroxys bonplandi, two new scarab beetles (Coleoptera, Scarabaeidae, Scarabaeinae) from Costa Rica, with notes on tropical mountain brachyptery and endemicity</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Kohlmann, Bert</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Solis, Angel</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Alvarado, Guillermo E.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2019</mods:date>
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<mods:detail type="volume">
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<mods:number>881</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>23</mods:start>
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<mods:end>51</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.881.38026</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.881.38026</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-881-23</mods:identifier>
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<mods:identifier type="ZooBank">ADCA057E3E1B45D8B5C9665683F51B59</mods:identifier>
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<mods:identifier type="Pensoft-UUID">0B1397AFEA7B5910A468BA7A24DA14DD</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="159754711" LSID="urn:lsid:zoobank.org:act:C13D9441-5A22-4CA1-974F-88B6A97199AC" httpUri="http://treatment.plazi.org/id/4E574931E2725135B5A689A4E7DD70E0" lastPageId="6" lastPageNumber="29" pageId="2" pageNumber="25">
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<subSubSection pageId="2" pageNumber="25" type="nomenclature">
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<paragraph pageId="2" pageNumber="25">
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<taxonomicName LSID="4E574931-E272-5135-B5A6-89A4E7DD70E0" class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus humboldti" order="Coleoptera" pageId="2" pageNumber="25" phylum="Arthropoda" rank="species" species="humboldti">Onthophagus humboldti</taxonomicName>
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<taxonomicNameLabel pageId="2" pageNumber="25">sp. nov.</taxonomicNameLabel>
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal drawing of a male Onthophagus humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure1">Figures 1</figureCitation>
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,
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">2a, c, e</figureCitation>
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,
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<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Aedeagui of a O. humboldti sp. nov. b U. bonplandi sp. nov. c brachypterous wing of O. humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure3" httpUri="https://binary.pensoft.net/fig/347598" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure3">3a, c</figureCitation>
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,
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<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Known distribution of O. humboldti sp. nov. (orange circle) and U. bonplandi sp. nov. (red triangle). The distribution of the proposed sister species of these new taxa is also depicted, O. micropterus (blue rhombus) and U. dybasi (black square)." figureDoi="10.3897/zookeys.881.38026.figure5" httpUri="https://binary.pensoft.net/fig/347600" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure5">5</figureCitation>
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,
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<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Known distribution of brachypterous Scarabaeinae in Costa Rica." figureDoi="10.3897/zookeys.881.38026.figure6" httpUri="https://binary.pensoft.net/fig/347601" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure6">6</figureCitation>
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,
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<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Present day distribution of Onthophagus humboldti sp. nov. (blue dot) and O. micropterus (red dot) and lines indicating proposed localities (rhombi) depressed by 1500 m (14 km in straight line) generated by the last glacial maximum, ~ 25 - 23 ka, in the Cordillera de Talamanca. All mountain systems are 150 m lower than present day height and an estimated sea level descent of 120 m is depicted. Dotted black lines represent present-day sea levels." figureDoi="10.3897/zookeys.881.38026.figure7" httpUri="https://binary.pensoft.net/fig/347602" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure7">7</figureCitation>
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,
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<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. A cytochrome c-oxidase I (COI- 5 P) mitochondrial DNA sequence-based BOLD taxon ID tree of the nearest species of O. humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure9" httpUri="https://binary.pensoft.net/fig/347604" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure9">9</figureCitation>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="25" type="type locality">
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<paragraph pageId="2" pageNumber="25">Type locality.</paragraph>
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<paragraph pageId="2" pageNumber="25">Costa Rica. Prov. Puntarenas. Buenos Aires, P.N. La Amistad. Tres Colinas.</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="25" type="type deposition">
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<paragraph pageId="2" pageNumber="25">Type deposition.</paragraph>
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<paragraph pageId="2" pageNumber="25">Museo Nacional de Costa Rica, Santo Domingo de Heredia, Costa Rica.</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="25" type="type material">
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<paragraph pageId="2" pageNumber="25">Type material.</paragraph>
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<paragraph pageId="2" pageNumber="25">
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Holotype male, pinned, with genitalia in a separate microvial. Original label: "Costa Rica. Provincia Puntarenas. Buenos Aires, Parque Nacional La Amistad. Tres Colinas. 2100-2200 m. 27-29 Febrero 2008. A.
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<normalizedToken originalValue="Solís">Solis</normalizedToken>
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, M. Moraga. Trampa Foso. L S 343850 565700." "HOLOTYPE/
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<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus humboldti" order="Coleoptera" pageId="2" pageNumber="25" phylum="Arthropoda" rank="species" species="humboldti">Onthophagus humboldti</taxonomicName>
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Kohlmann,
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<normalizedToken originalValue="Solís">Solis</normalizedToken>
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, Alvarado [red printed label]".
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</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="25" type="materials_examined">
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<paragraph pageId="2" pageNumber="25">Other material.</paragraph>
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<paragraph pageId="2" pageNumber="25">
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Paratypes. (8 males, 4 females). "Costa Rica. Provincia Puntarenas. Buenos Aires, Parque Nacional La Amistad. Tres Colinas. 2100-2200 m. 27-29 Febrero 2008. A.
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<normalizedToken originalValue="Solís">Solis</normalizedToken>
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, M. Moraga. Trampa Foso. L S 343850 565700."
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</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="25" type="diagnosis">
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<paragraph pageId="2" pageNumber="25">Diagnosis.</paragraph>
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<paragraph pageId="2" pageNumber="25">
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Elytra as long as or shorter than pronotum (
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal drawing of a male Onthophagus humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure1">Fig. 1</figureCitation>
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), due to brachyptery (
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<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Aedeagui of a O. humboldti sp. nov. b U. bonplandi sp. nov. c brachypterous wing of O. humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure3" httpUri="https://binary.pensoft.net/fig/347598" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure3">Fig. 3c</figureCitation>
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). Broad clypeal horn bifurcation (
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2a</figureCitation>
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); pygidium and apex of elytra with evident setae; clypeal margin indented at junction with clypeo-genal suture (
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2e</figureCitation>
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).
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</paragraph>
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<caption doi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="2" pageNumber="25" start="Figure 1" startId="F1">
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<paragraph pageId="2" pageNumber="25">
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<emphasis bold="true" pageId="2" pageNumber="25">Figure 1.</emphasis>
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Dorsal drawing of a male
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<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus humboldti" order="Coleoptera" pageId="2" pageNumber="25" phylum="Arthropoda" rank="species" species="humboldti">
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<emphasis italics="true" pageId="2" pageNumber="25">Onthophagus humboldti</emphasis>
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</taxonomicName>
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sp. nov.
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</paragraph>
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</caption>
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<caption doi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" start="Figure 2" startId="F2">
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<paragraph pageId="2" pageNumber="25">
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<emphasis bold="true" pageId="2" pageNumber="25">Figure 2.</emphasis>
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Drawings of the clypeal horn of
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<emphasis bold="true" pageId="2" pageNumber="25">a</emphasis>
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<taxonomicName lsidName="O. humboldti" pageId="2" pageNumber="25" rank="species" species="humboldti">
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<emphasis italics="true" pageId="2" pageNumber="25">O. humboldti</emphasis>
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</taxonomicName>
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sp. nov. and
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<emphasis bold="true" pageId="2" pageNumber="25">b</emphasis>
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<taxonomicName lsidName="O. micropterus" pageId="2" pageNumber="25" rank="species" species="micropterus">
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<emphasis italics="true" pageId="2" pageNumber="25">O. micropterus</emphasis>
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</taxonomicName>
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; pronotum of
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<emphasis bold="true" pageId="2" pageNumber="25">c</emphasis>
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<taxonomicName lsidName="O. humboldti" pageId="2" pageNumber="25" rank="species" species="humboldti">
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<emphasis italics="true" pageId="2" pageNumber="25">O. humboldti</emphasis>
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</taxonomicName>
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sp. nov. and
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<emphasis bold="true" pageId="2" pageNumber="25">d</emphasis>
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<taxonomicName lsidName="O. micropterus" pageId="2" pageNumber="25" rank="species" species="micropterus">
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<emphasis italics="true" pageId="2" pageNumber="25">O. micropterus</emphasis>
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</taxonomicName>
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; head of
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<emphasis bold="true" pageId="2" pageNumber="25">e</emphasis>
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<taxonomicName lsidName="O. humboldti" pageId="2" pageNumber="25" rank="species" species="humboldti">
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<emphasis italics="true" pageId="2" pageNumber="25">O. humboldti</emphasis>
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</taxonomicName>
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sp. nov. and
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<emphasis bold="true" pageId="2" pageNumber="25">f</emphasis>
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<taxonomicName lsidName="O. micropterus" pageId="2" pageNumber="25" rank="species" species="micropterus">
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<emphasis italics="true" pageId="2" pageNumber="25">O. micropterus</emphasis>
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</taxonomicName>
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; and elytral apex of
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<emphasis bold="true" pageId="2" pageNumber="25">g</emphasis>
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<taxonomicName lsidName="U. bonplandi" pageId="2" pageNumber="25" rank="species" species="bonplandi">
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<emphasis italics="true" pageId="2" pageNumber="25">U. bonplandi</emphasis>
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</taxonomicName>
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sp. nov. and
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<emphasis bold="true" pageId="2" pageNumber="25">h</emphasis>
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<taxonomicName lsidName="U. dybasi" pageId="2" pageNumber="25" rank="species" species="dybasi">
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<emphasis italics="true" pageId="2" pageNumber="25">U. dybasi</emphasis>
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</taxonomicName>
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. Scale bars: 1 mm.
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</paragraph>
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</caption>
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<caption doi="10.3897/zookeys.881.38026.figure3" httpUri="https://binary.pensoft.net/fig/347598" pageId="2" pageNumber="25" start="Figure 3" startId="F3">
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<paragraph pageId="2" pageNumber="25">
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<emphasis bold="true" pageId="2" pageNumber="25">Figure 3.</emphasis>
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Aedeagui of
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<emphasis bold="true" pageId="2" pageNumber="25">a</emphasis>
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<taxonomicName lsidName="O. humboldti" pageId="2" pageNumber="25" rank="species" species="humboldti">
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<emphasis italics="true" pageId="2" pageNumber="25">O. humboldti</emphasis>
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</taxonomicName>
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sp. nov.
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<emphasis bold="true" pageId="2" pageNumber="25">b</emphasis>
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<taxonomicName lsidName="U. bonplandi" pageId="2" pageNumber="25" rank="species" species="bonplandi">
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<emphasis italics="true" pageId="2" pageNumber="25">U. bonplandi</emphasis>
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</taxonomicName>
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sp. nov.
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<emphasis bold="true" pageId="2" pageNumber="25">c</emphasis>
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brachypterous wing of
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<taxonomicName lsidName="O. humboldti" pageId="2" pageNumber="25" rank="species" species="humboldti">
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<emphasis italics="true" pageId="2" pageNumber="25">O. humboldti</emphasis>
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</taxonomicName>
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sp. nov.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection lastPageId="4" lastPageNumber="27" pageId="2" pageNumber="25" type="description">
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<paragraph pageId="2" pageNumber="25">Description.</paragraph>
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<paragraph pageId="2" pageNumber="25">
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<emphasis bold="true" italics="true" pageId="2" pageNumber="25">Holotype.</emphasis>
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Male (
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal drawing of a male Onthophagus humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure1">Fig. 1</figureCitation>
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), length 7.2 mm; maximum width 4.3 mm. Oval, shining reddish black. Centre of the clypeus projected forming a slender bifid horn (
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal drawing of a male Onthophagus humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure1">Figs 1</figureCitation>
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,
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">2c</figureCitation>
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); genae projected forming an angle (
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2e</figureCitation>
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), genal sutures almost effaced; head punctures coarse, regular, dense, becoming finer and sparser towards the center; clypeal carina absent, vertex carina substituted by two vertical asymmetric platelets, modestly developed, and obliquely oriented; eyes two times longer than wide and separated by eight times the eye width.
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</paragraph>
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<paragraph pageId="2" pageNumber="25">
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Pronotum (
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2c</figureCitation>
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) very convex, lateral margins with a small and irregular fovea, not lineal; lateral pronotal margins bordered by a deep sulcus, anterior and basal borders margined; pronotal surface reticulate and covered by dense, regular, coarse, annular, and deep punctures without setae; pronotal projection well-developed (
|
||
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal drawing of a male Onthophagus humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure1">Figs 1</figureCitation>
|
||
,
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">2c</figureCitation>
|
||
), forming a broad bilobed plate slightly bent downwards, with a depressed area antero-centrally, and having clear antero-lateral margins; anterior angles projected as long, slender, and curved projections (
|
||
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal drawing of a male Onthophagus humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure1">Figs 1</figureCitation>
|
||
,
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure2">2c</figureCitation>
|
||
); pronotal base with a sulcus extending forward one third its length; scutellum not visible between the base of the elytra.
|
||
</paragraph>
|
||
<paragraph pageId="2" pageNumber="25">
|
||
Elytra convex, with clear margins and without a humeral callus; with eight well-marked striae, fine and clearly impressed and with crenulating punctures; intervals clearly punctured, punctures big and dense, not aligned, bearing short, stiff setae along the lateral and apical margins; microsculpture reticulate and regular. Wing brachypterous, measuring 0.75 mm (
|
||
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Aedeagui of a O. humboldti sp. nov. b U. bonplandi sp. nov. c brachypterous wing of O. humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure3" httpUri="https://binary.pensoft.net/fig/347598" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure3">Fig. 3c</figureCitation>
|
||
). Pygidium moderately shiny and shagreen, margined border, with big, coarse, annular punctures bearing short and stiff setae. Aedeagus as
|
||
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Aedeagui of a O. humboldti sp. nov. b U. bonplandi sp. nov. c brachypterous wing of O. humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure3" httpUri="https://binary.pensoft.net/fig/347598" pageId="2" pageNumber="25" tableDoi="10.3897/zookeys.881.38026.figure3">Fig. 3a</figureCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph pageId="2" pageNumber="25">Mesosternum with evident annular punctures bearing no setae. Metasternum shagreen and finely punctured, more coarsely laterally, basal third with a sulcus. Abdominal segments shagreen and finely punctured.</paragraph>
|
||
<paragraph pageId="3" pageNumber="26">
|
||
<pageBreakToken pageId="3" pageNumber="26" start="start">Fore</pageBreakToken>
|
||
femur long, slender, and punctured; meso- and metafemur short and elongate, light yellow. Fore tibia long, slender and arched (
|
||
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Dorsal drawing of a male Onthophagus humboldti sp. nov." figureDoi="10.3897/zookeys.881.38026.figure1" httpUri="https://binary.pensoft.net/fig/347596" pageId="3" pageNumber="26" tableDoi="10.3897/zookeys.881.38026.figure1">Fig. 1</figureCitation>
|
||
); with four external teeth; tibial spur elongated, straight, pointed, deflexed anteriorly, extending to second tarsal segment. Middle- and hind femur light yellow at middle.
|
||
</paragraph>
|
||
<paragraph lastPageId="4" lastPageNumber="27" pageId="3" pageNumber="26">
|
||
Female, length 6.3 mm; maximum width 3.6 mm. It is similar to the male and varies in having a clypeus not forming a horn, clypeus shagreen, genae not projected as teeth, with a head frons keel, two small platelet projections at head vertex, no pronotal
|
||
<pageBreakToken pageId="4" pageNumber="27" start="start">projection</pageBreakToken>
|
||
, no projected pronotal anterior angles, fore tibia short, fore femur short, last abdominal sternite broad.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="4" pageNumber="27" type="variation">
|
||
<paragraph pageId="4" pageNumber="27">Variation.</paragraph>
|
||
<paragraph pageId="4" pageNumber="27">Length 5.6 to 7.2 mm. Width 3.2 to 4.3 mm. Small males do not have the bifid clypeal horn, just a small erect lamella; vertex platelets forming a small projection; anterior pronotal angles not projected, pronotal projection forming a small carina. Body color varying from black to piceous red.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="4" pageNumber="27" type="etymology">
|
||
<paragraph pageId="4" pageNumber="27">Etymology.</paragraph>
|
||
<paragraph pageId="4" pageNumber="27">This species is dedicated in honor of Friedrich Wilhelm Heinrich Alexander von Humboldt, Prussian geographer, explorer, and naturalist, commemorating the 250th anniversary of his birth. He is widely recognized for fathering the work on physical and plant geography, which laid the foundation for the development of modern biogeography.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="5" lastPageNumber="28" pageId="4" pageNumber="27" type="taxonomic considerations">
|
||
<paragraph pageId="4" pageNumber="27">Taxonomic considerations.</paragraph>
|
||
<paragraph pageId="4" pageNumber="27">
|
||
<bibRefCitation author="Kohlmann, B" journalOrPublisher="Giornale Italiano di Entomologia" pageId="22" pageNumber="45" pagination="159 - 261" refId="B33" refString="Kohlmann, B, Solis, A, 2001. El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia 9: 159 - 261" title="El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica." volume="9" year="2001">
|
||
Kohlmann and
|
||
<normalizedToken originalValue="Solís">Solis</normalizedToken>
|
||
(2001)
|
||
</bibRefCitation>
|
||
report the existence of 39 species of
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus" order="Coleoptera" pageId="4" pageNumber="27" phylum="Arthropoda" rank="genus">
|
||
<emphasis italics="true" pageId="4" pageNumber="27">Onthophagus</emphasis>
|
||
</taxonomicName>
|
||
for Costa Rica. This new species would increase their numbers to 40.
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus humboldti" order="Coleoptera" pageId="4" pageNumber="27" phylum="Arthropoda" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="4" pageNumber="27">Onthophagus humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov. belongs to the
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus dicranius" order="Coleoptera" pageId="4" pageNumber="27" phylum="Arthropoda" rank="species" species="dicranius">
|
||
<emphasis italics="true" pageId="4" pageNumber="27">Onthophagus dicranius</emphasis>
|
||
</taxonomicName>
|
||
Bates species group, as defined by
|
||
<bibRefCitation author="Kohlmann, B" journalOrPublisher="Giornale Italiano di Entomologia" pageId="22" pageNumber="45" pagination="159 - 261" refId="B33" refString="Kohlmann, B, Solis, A, 2001. El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia 9: 159 - 261" title="El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica." volume="9" year="2001">
|
||
Kohlmann and
|
||
<normalizedToken originalValue="Solís">Solis</normalizedToken>
|
||
(2001)
|
||
</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph pageId="5" pageNumber="28">
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus humboldti" order="Coleoptera" pageId="5" pageNumber="28" phylum="Arthropoda" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">
|
||
<pageBreakToken pageId="5" pageNumber="28" start="start">Onthophagus</pageBreakToken>
|
||
humboldti
|
||
</emphasis>
|
||
</taxonomicName>
|
||
sp. nov. will key out to
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
Zunino & Halffter, 1981, in Kohlmann and
|
||
<normalizedToken originalValue="Solís´">Solis'</normalizedToken>
|
||
key (
|
||
<bibRefCitation author="Kohlmann, B" journalOrPublisher="Giornale Italiano di Entomologia" pageId="22" pageNumber="45" pagination="159 - 261" refId="B33" refString="Kohlmann, B, Solis, A, 2001. El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia 9: 159 - 261" title="El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica." volume="9" year="2001">2001</bibRefCitation>
|
||
). It can be easily differentiated by the following characteristics: In males clypeal horn slender at middle and very bifurcated at apex (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2a</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. humboldti" pageId="5" pageNumber="28" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov.) versus broad at middle and notched at apex (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2b</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
); genae projected forming an angle (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2e</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. humboldti" pageId="5" pageNumber="28" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov.) versus genae projected forming a tooth (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2f</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
); vertex platelets forming a carina (
|
||
<taxonomicName lsidName="O. humboldti" pageId="5" pageNumber="28" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov.) versus a pointed projection (
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
); anterior lateral angles of pronotum projected as long, slender, and curved projections (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2c</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. humboldti" pageId="5" pageNumber="28" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov.) versus a short, curved projection (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2d</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
); pronotal central forward projection well-developed, forming a broad bilobed plate slightly bent downwards (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2c</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. humboldti" pageId="5" pageNumber="28" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov.) versus a bilobed plate projecting forward (
|
||
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Drawings of the clypeal horn of a O. humboldti sp. nov. and b O. micropterus; pronotum of c O. humboldti sp. nov. and d O. micropterus; head of e O. humboldti sp. nov. and f O. micropterus; and elytral apex of g U. bonplandi sp. nov. and h U. dybasi. Scale bars: 1 mm." figureDoi="10.3897/zookeys.881.38026.figure2" httpUri="https://binary.pensoft.net/fig/347597" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure2">Fig. 2d</figureCitation>
|
||
) (
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
). In females: vertex platelets forming a carina (
|
||
<taxonomicName lsidName="O. humboldti" pageId="5" pageNumber="28" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov.) versus a pointed projection (
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="28" type="distribution">
|
||
<paragraph pageId="5" pageNumber="28">Geographical distribution.</paragraph>
|
||
<paragraph pageId="5" pageNumber="28">
|
||
This species is so far only known from the area of Tres Colinas, near Buenos Aires, in the province of Puntarenas (
|
||
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Known distribution of O. humboldti sp. nov. (orange circle) and U. bonplandi sp. nov. (red triangle). The distribution of the proposed sister species of these new taxa is also depicted, O. micropterus (blue rhombus) and U. dybasi (black square)." figureDoi="10.3897/zookeys.881.38026.figure5" httpUri="https://binary.pensoft.net/fig/347600" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure5">Fig. 5</figureCitation>
|
||
). It has been collected from 2100 to 2200 m altitude in the month of February in lower montane rain forest.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="5" pageNumber="28" type="chorological affinities">
|
||
<paragraph pageId="5" pageNumber="28">Chorological affinities.</paragraph>
|
||
<paragraph pageId="5" pageNumber="28">
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus humboldti" order="Coleoptera" pageId="5" pageNumber="28" phylum="Arthropoda" rank="species" species="humboldti">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">Onthophagus humboldti</emphasis>
|
||
</taxonomicName>
|
||
sp. nov. is endemic to the Cordillera de Talamanca and is the tenth known brachypterous
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus" order="Coleoptera" pageId="5" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">Onthophagus</emphasis>
|
||
</taxonomicName>
|
||
species to be described worldwide. A closely related species,
|
||
<taxonomicName lsidName="O. micropterus" pageId="5" pageNumber="28" rank="species" species="micropterus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. micropterus</emphasis>
|
||
</taxonomicName>
|
||
, is also distributed in the Cordillera de Talamanca (
|
||
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Known distribution of brachypterous Scarabaeinae in Costa Rica." figureDoi="10.3897/zookeys.881.38026.figure6" httpUri="https://binary.pensoft.net/fig/347601" pageId="5" pageNumber="28" tableDoi="10.3897/zookeys.881.38026.figure6">Fig. 6</figureCitation>
|
||
), from 2100 to 3000 m altitude in tropical mountain rainforest and has been collected from October to February.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="6" lastPageNumber="29" pageId="5" pageNumber="28" type="biogeography">
|
||
<paragraph pageId="5" pageNumber="28">Biogeography.</paragraph>
|
||
<paragraph pageId="5" pageNumber="28">
|
||
This species belongs to the
|
||
<taxonomicName lsidName="O. dicranius" pageId="5" pageNumber="28" rank="species" species="dicranius">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. dicranius</emphasis>
|
||
</taxonomicName>
|
||
species group, as established by
|
||
<bibRefCitation author="Kohlmann, B" journalOrPublisher="Giornale Italiano di Entomologia" pageId="22" pageNumber="45" pagination="159 - 261" refId="B33" refString="Kohlmann, B, Solis, A, 2001. El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia 9: 159 - 261" title="El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica." volume="9" year="2001">
|
||
Kohlmann and
|
||
<normalizedToken originalValue="Solís">Solis</normalizedToken>
|
||
(2001)
|
||
</bibRefCitation>
|
||
. This group of species has extra-American affinities, in which
|
||
<bibRefCitation DOI="https://doi.org/10.4039/Ent1251091-6" author="Howden, HF" journalOrPublisher="Canadian Entomologist" pageId="22" pageNumber="45" pagination="1091 - 1114" refId="B24" refString="Howden, HF, Gill, B, 1993. Mesoamerican Onthophagus Latreille in the dicranius and mirabilis species groups (Coleoptera: Scarabaeidae). Canadian Entomologist 125: 1091 - 1114" title="Mesoamerican Onthophagus Latreille in the dicranius and mirabilis species groups (Coleoptera: Scarabaeidae)." url="https://doi.org/10.4039/Ent1251091-6" volume="125" year="1993">Howden and Gill (1993)</bibRefCitation>
|
||
indicate that the American fauna of
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus" order="Coleoptera" pageId="5" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">Onthophagus</emphasis>
|
||
</taxonomicName>
|
||
is the result of invasive species from East Asia and that the
|
||
<taxonomicName lsidName="O. dicranius" pageId="5" pageNumber="28" rank="species" species="dicranius">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. dicranius</emphasis>
|
||
</taxonomicName>
|
||
group presents characters in common with New Guinea species. This agrees with the hypothesis originally proposed by
|
||
<bibRefCitation author="Zunino, M" journalOrPublisher="Elytron" pageId="22" pageNumber="45" pagination="137 - 142" refId="B81" refString="Zunino, M, Halffter, G, 1988. Una nueva especie braquiptera de Onthophagus de Mexico (Coleoptera, Scarabaeidae). Elytron 2: 137 - 142" title="Una nueva especie braquiptera de Onthophagus de Mexico (Coleoptera, Scarabaeidae)." volume="2" year="1988">Zunino and Halffter (1988)</bibRefCitation>
|
||
, which points out for the supraspecific groups of American
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus" order="Coleoptera" pageId="5" pageNumber="28" phylum="Arthropoda" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">Onthophagus</emphasis>
|
||
</taxonomicName>
|
||
, an origin of its lineages, which in the case of the current representatives is distributed in East or Southeast Asia; and for this case, the Asian representation of the ancestral line, like the American one, has its distribution present in the humid tropics. On the other hand, the
|
||
<taxonomicName lsidName="O. dicranius" pageId="5" pageNumber="28" rank="species" species="dicranius">
|
||
<emphasis italics="true" pageId="5" pageNumber="28">O. dicranius</emphasis>
|
||
</taxonomicName>
|
||
species group has its present-day center of diversity in tropical North America and relatives in South America (
|
||
<bibRefCitation author="Zunino, M" journalOrPublisher="Elytron" pageId="22" pageNumber="45" pagination="157 - 178" refId="B82" refString="Zunino, M, Halffter, G, 1997. Sobre Onthophagus Latreille, 1802 americanos (Coleoptera: Scarabaeidae: Scarabaeinae). Elytron 11: 157 - 178" title="Sobre Onthophagus Latreille, 1802 americanos (Coleoptera: Scarabaeidae: Scarabaeinae)." volume="11" year="1997">Zunino and Halffter 1997</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Kohlmann, B" journalOrPublisher="Giornale Italiano di Entomologia" pageId="22" pageNumber="45" pagination="159 - 261" refId="B33" refString="Kohlmann, B, Solis, A, 2001. El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia 9: 159 - 261" title="El Genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica." volume="9" year="2001">
|
||
Kohlmann and
|
||
<normalizedToken originalValue="Solís">Solis</normalizedToken>
|
||
2001
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph lastPageId="6" lastPageNumber="29" pageId="5" pageNumber="28">
|
||
This situation seems to be in congruence with the boreotropical distribution hypothesis (
|
||
<bibRefCitation pageId="5" pageNumber="28" refId="B74">Wang 1961</bibRefCitation>
|
||
;
|
||
<bibRefCitation DOI="https://doi.org/10.2307/2395198" author="Wolfe, JA" journalOrPublisher="Annals of the Missouri Botanical Garden" pageId="22" pageNumber="45" pagination="264 - 279" refId="B76" refString="Wolfe, JA, 1975. Some aspects of plant geography in the Northern Hemisphere during the late Cretaceous and Tertiary. Annals of the Missouri Botanical Garden 62: 264 - 279" title="Some aspects of plant geography in the Northern Hemisphere during the late Cretaceous and Tertiary." url="https://doi.org/10.2307/2395198" volume="62" year="1975">Wolfe 1975</bibRefCitation>
|
||
;
|
||
<bibRefCitation DOI="https://doi.org/10.1002/j.1537-2197.1993.tb13761.x" author="Lavin, M" journalOrPublisher="American Journal of Botany" pageId="22" pageNumber="45" pagination="1 - 14" refId="B40" refString="Lavin, M, Luckow, M, 1993. Origins and relationships of tropical North America in the context of the boreotropics hypothesis. American Journal of Botany 80: 1 - 14" title="Origins and relationships of tropical North America in the context of the boreotropics hypothesis." url="https://doi.org/10.1002/j.1537-2197.1993.tb13761.x" volume="80" year="1993">Lavin and Luckow 1993</bibRefCitation>
|
||
;
|
||
<bibRefCitation DOI="https://doi.org/10.1086/323332" author="Xiang, Q-Y" journalOrPublisher="International Journal of Plant Science" pageId="22" pageNumber="45" pagination="29 - 39" refId="B77" refString="Xiang, Q-Y, Soltis, DE, 2001. Dispersal-vicariance analyses of intercontinental disjuncts: historical biogeographical implications for angiosperms in the Northern hemisphere. International Journal of Plant Science 162: 29 - 39" title="Dispersal-vicariance analyses of intercontinental disjuncts: historical biogeographical implications for angiosperms in the Northern hemisphere." url="https://doi.org/10.1086/323332" volume="162" year="2001">Xiang and Soltis 2001</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Davis, CC" journalOrPublisher="Systematic Botany" pageId="22" pageNumber="45" pagination="289 - 302" refId="B10" refString="Davis, CC, Fritsch, PW, Li, J, Donohue, MJ, 2002. Phylogeny and biogeography of Cercis (Fabaceae): Evidence from nuclear ribosomal ITS and chloroplast nhdF sequence data. Systematic Botany 27: 289 - 302" title="Phylogeny and biogeography of Cercis (Fabaceae): Evidence from nuclear ribosomal ITS and chloroplast nhdF sequence data." volume="27" year="2002">Davis et al. 2002</bibRefCitation>
|
||
), where current flora groups show a tropical disjunct distribution, generally centered in America, Africa, and tropical Asia. This hypothesis is based on the observation of the existence of tropical broadleaf forests during the Early Paleogene (in old Stratigraphy terminology, Early Tertiary) at high latitudes in regions that are currently temperate, directed by a Late Paleocene-Early Eocene thermal maxima (ca. 52 ma,
|
||
<bibRefCitation author="Zachos, J" journalOrPublisher="Science" pageId="22" pageNumber="45" pagination="686 - 693" refId="B79" refString="Zachos, J, Pagani, M, Sloan, L, Thomas, E, Billups, K, 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292: 686 - 693" title="Trends, rhythms, and aberrations in global climate 65 Ma to present." volume="292" year="2001">Zachos et al. 2001</bibRefCitation>
|
||
) and that many current angiosperm temperate taxa have evergreen relatives in subtropical rainforests (
|
||
<bibRefCitation DOI="https://doi.org/10.1111/j.1558-5646.1966.tb03339.x" author="Axelrod, DI" journalOrPublisher="Evolution" pageId="22" pageNumber="45" pagination="1 - 15" refId="B4" refString="Axelrod, DI, 1966. Origin of deciduous and evergreen habitats in temperate forests. Evolution 20: 1 - 15" title="Origin of deciduous and evergreen habitats in temperate forests." url="https://doi.org/10.1111/j.1558-5646.1966.tb03339.x" volume="20" year="1966">Axelrod 1966</bibRefCitation>
|
||
). This proposal then suggests the existence of northern bridges that were once at lower latitudes, such as the Bering
|
||
<pageBreakToken pageId="6" pageNumber="29" start="start">Bridge</pageBreakToken>
|
||
during the Early Paleogene and the North Atlantic Bridge during the Eocene, which may have served as migration routes for groups of organisms that currently present intercontinental disjunct distributions. This hypothesis suggests that a taxon with a present-day center of diversity in tropical North America, and with an early Paleogene fossil record from any region there, has a high probability of having sister-group relatives in the Paleotropics and derived relatives in South America (
|
||
<bibRefCitation DOI="https://doi.org/10.1002/j.1537-2197.1993.tb13761.x" author="Lavin, M" journalOrPublisher="American Journal of Botany" pageId="22" pageNumber="45" pagination="1 - 14" refId="B40" refString="Lavin, M, Luckow, M, 1993. Origins and relationships of tropical North America in the context of the boreotropics hypothesis. American Journal of Botany 80: 1 - 14" title="Origins and relationships of tropical North America in the context of the boreotropics hypothesis." url="https://doi.org/10.1002/j.1537-2197.1993.tb13761.x" volume="80" year="1993">Lavin and Luckow 1993</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="6" pageNumber="29">
|
||
This pattern of distribution would clarify those proposed by Halffter (
|
||
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4226.1.1" author="Halffter, G" journalOrPublisher="Zootaxa" pageId="22" pageNumber="45" pagination="1 - 46" refId="B19" refString="Halffter, G, Morrone, JJ, 2017. An analytical review of Halffter's Mexican transition Zone, and its relevance for evolutionary biogeography, ecology and biogeographical regionalization. Zootaxa 4226: 1 - 46" title="An analytical review of Halffter's Mexican transition Zone, and its relevance for evolutionary biogeography, ecology and biogeographical regionalization." url="https://doi.org/10.11646/zootaxa.4226.1.1" volume="4226" year="2017">Halffter and Morrone 2017</bibRefCitation>
|
||
) for the "Mexican Transition Zone" in particular one of them, the so-called "Paleoamerican Dispersion Pattern" (
|
||
<bibRefCitation author="Halffter, G" journalOrPublisher="Folia Entomologica Mexicana" pageId="22" pageNumber="45" pagination="1 - 108" refId="B18" refString="Halffter, G, 1964. La entomofauna americana, ideas acerca de su origen y distribucion. Folia Entomologica Mexicana 6: 1 - 108" title="La entomofauna americana, ideas acerca de su origen y distribucion." volume="6" year="1964">Halffter 1964</bibRefCitation>
|
||
). This pattern of dispersion corresponds to northern taxa that arrived in North America from Eurasia, and has been subdivided by
|
||
<bibRefCitation author="Halffter, G" journalOrPublisher="Elytron" pageId="22" pageNumber="45" pagination="151 - 185" refId="B20" refString="Halffter, G, Favila, ME, Arellano, L, 1995. Spatial distribution of three groups of Coleoptera along an altitudinal transect in the Mexican Transition Zone and its biogeographical implications. Elytron 9: 151 - 185" title="Spatial distribution of three groups of Coleoptera along an altitudinal transect in the Mexican Transition Zone and its biogeographical implications." volume="9" year="1995">Halffter et al. (1995)</bibRefCitation>
|
||
into four variants, where one of them, called the "Paleoamerican Tropical Pattern", corresponds to species found in the lowlands of the tropics and at medium altitudes, their distribution being very similar to that of the Neotropical pattern, but their affinities are with the Old World taxa.
|
||
<bibRefCitation author="Halffter, G" journalOrPublisher="Elytron" pageId="22" pageNumber="45" pagination="151 - 185" refId="B20" refString="Halffter, G, Favila, ME, Arellano, L, 1995. Spatial distribution of three groups of Coleoptera along an altitudinal transect in the Mexican Transition Zone and its biogeographical implications. Elytron 9: 151 - 185" title="Spatial distribution of three groups of Coleoptera along an altitudinal transect in the Mexican Transition Zone and its biogeographical implications." volume="9" year="1995">Halffter et al. (1995</bibRefCitation>
|
||
,
|
||
<bibRefCitation DOI="https://doi.org/10.4081/fe.2008.99" author="Halffter, G" journalOrPublisher="Fragmenta entomologica" pageId="22" pageNumber="45" pagination="273 - 332" refId="B21" refString="Halffter, G, Verdu, JR, Marquez, J, Moreno, CE, 2008. Biogeographical analysis of Scarabaeinae and Geotrupinae along a transect in Central Mexico. Fragmenta entomologica 40: 273 - 332" title="Biogeographical analysis of Scarabaeinae and Geotrupinae along a transect in Central Mexico." url="https://doi.org/10.4081/fe.2008.99" volume="40" year="2008">2008</bibRefCitation>
|
||
) placed the
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus clypeatus" order="Coleoptera" pageId="6" pageNumber="29" phylum="Arthropoda" rank="species" species="clypeatus">
|
||
<emphasis italics="true" pageId="6" pageNumber="29">Onthophagus clypeatus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis italics="true" pageId="6" pageNumber="29">
|
||
>
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus dicranius" order="Coleoptera" pageId="6" pageNumber="29" phylum="Arthropoda" rank="species" species="dicranius">Onthophagus dicranius</taxonomicName>
|
||
</emphasis>
|
||
species groups of the genus
|
||
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus" order="Coleoptera" pageId="6" pageNumber="29" phylum="Arthropoda" rank="genus">
|
||
<emphasis italics="true" pageId="6" pageNumber="29">Onthophagus</emphasis>
|
||
</taxonomicName>
|
||
within this pattern.
|
||
</paragraph>
|
||
<paragraph pageId="6" pageNumber="29">
|
||
Actually, the groups of species mentioned above are congruent with the typical characteristics of the so-called boreotropical distribution. Therefore, the aforementioned distribution variant, the "Paleoamerican Tropical Pattern", seems to be the same with the boreotropical distribution and it is proposed here to use the term boreotropical distribution from now on as it is a more complete and well-founded concept, besides being an older one. This pattern has been studied and characterized at very fine phylogenetic and biogeographic analysis levels in animal and plants (
|
||
<bibRefCitation DOI="https://doi.org/10.1017/S009483730000974X" author="Lidgard, S" journalOrPublisher="Paleobiology" pageId="22" pageNumber="45" pagination="77 - 93" refId="B42" refString="Lidgard, S, Crane, PR, 1990. Angiosperm diversification and Cretaceous floristic trends: A comparison of palynofloras and leaf macrofloras. Paleobiology 16: 77 - 93" title="Angiosperm diversification and Cretaceous floristic trends: A comparison of palynofloras and leaf macrofloras." url="https://doi.org/10.1017/S009483730000974X" volume="16" year="1990">Lidgard and Crane 1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation DOI="https://doi.org/10.1086/323332" author="Xiang, Q-Y" journalOrPublisher="International Journal of Plant Science" pageId="22" pageNumber="45" pagination="29 - 39" refId="B77" refString="Xiang, Q-Y, Soltis, DE, 2001. Dispersal-vicariance analyses of intercontinental disjuncts: historical biogeographical implications for angiosperms in the Northern hemisphere. International Journal of Plant Science 162: 29 - 39" title="Dispersal-vicariance analyses of intercontinental disjuncts: historical biogeographical implications for angiosperms in the Northern hemisphere." url="https://doi.org/10.1086/323332" volume="162" year="2001">Xiang and Soltis 2001</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Davis, CC" journalOrPublisher="Systematic Botany" pageId="22" pageNumber="45" pagination="289 - 302" refId="B10" refString="Davis, CC, Fritsch, PW, Li, J, Donohue, MJ, 2002. Phylogeny and biogeography of Cercis (Fabaceae): Evidence from nuclear ribosomal ITS and chloroplast nhdF sequence data. Systematic Botany 27: 289 - 302" title="Phylogeny and biogeography of Cercis (Fabaceae): Evidence from nuclear ribosomal ITS and chloroplast nhdF sequence data." volume="27" year="2002">Davis et al. 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation DOI="https://doi.org/10.1016/j.ympev.2009.01.017" author="Feng, C-M" journalOrPublisher="Molecular and Phylogenetic Evolution" pageId="22" pageNumber="45" pagination="201 - 214" refId="B12" refString="Feng, C-M, Manchester, SR, Xiang, Q-Y, 2009. Phylogeny and biogeography of Alangiaceae (Cornales) inferred from DNA sequences, morphology, and fossils. Molecular and Phylogenetic Evolution 51: 201 - 214" title="Phylogeny and biogeography of Alangiaceae (Cornales) inferred from DNA sequences, morphology, and fossils." url="https://doi.org/10.1016/j.ympev.2009.01.017" volume="51" year="2009">Feng et al. 2009</bibRefCitation>
|
||
;
|
||
<bibRefCitation DOI="https://doi.org/10.1371/journal.pone.0038788" author="Guo, Y-Y" journalOrPublisher="Fabreries" pageId="22" pageNumber="45" refId="B16" refString="Guo, Y-Y, Luo, Y-B, Liu, Z-J, Wang, X-Q, 2012. . https://doi.org/10.1371/journal.pone.0038788" url="https://doi.org/10.1371/journal.pone.0038788" year="2012">Guo et al. 2012</bibRefCitation>
|
||
;
|
||
<bibRefCitation DOI="https://doi.org/10.1002/ece3.2688" author="Ye, Z" journalOrPublisher="Ecology and Evolution" pageId="22" pageNumber="45" pagination="793 - 802" refId="B78" refString="Ye, Z, Zhen, Y, Zhou, Y, Bu, W, 2016. Out of Africa: Biogeography and diversification of the pantropical pond skater genus Limnogonus Stal, 1868 (Hemiptera: Gerridae). Ecology and Evolution 7: 793 - 802" title="Out of Africa: Biogeography and diversification of the pantropical pond skater genus Limnogonus Stal, 1868 (Hemiptera: Gerridae)." url="https://doi.org/10.1002/ece3.2688" volume="7" year="2016">Ye et al. 2016</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |