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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.891.36027" ID-GBIF-Dataset="dc0e74d5-f9a0-4ae1-93f9-c0f60431eb90" ID-PMC="PMC6882928" ID-Pensoft-Pub="1313-2970-891-71" ID-Pensoft-UUID="76F1823B3926587AB20B249DB0DD1D1B" ID-PubMed="31802973" ID-ZooBank="F3F32E940AB749C4A108162690F122B4" ModsDocID="1313-2970-891-71" checkinTime="1574358824618" checkinUser="pensoft" docAuthor="Dehon, Manuel, Engel, Michael S., Gerard, Maxence, Aytekin, A. Murat, Ghisbain, Guillaume, Williams, Paul H., Rasmont, Pierre &amp; Michez, Denis" docDate="2019" docId="81A479F7619E50A88342300A233BDFB5" docLanguage="en" docName="ZooKeys 891: 71-118" docOrigin="ZooKeys 891" docSource="http://dx.doi.org/10.3897/zookeys.891.36027" docTitle="Bombus (Melanobombus) cerdanyensis Dehon, De Meulemeester &amp; Engel 2014" docType="treatment" docVersion="4" id="76F1823B3926587AB20B249DB0DD1D1B" lastPageNumber="71" masterDocId="76F1823B3926587AB20B249DB0DD1D1B" masterDocTitle="Morphometric analysis of fossil bumble bees (Hymenoptera, Apidae, Bombini) reveals their taxonomic affinities" masterLastPageNumber="118" masterPageNumber="71" pageNumber="71" updateTime="1668168126927" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Morphometric analysis of fossil bumble bees (Hymenoptera, Apidae, Bombini) reveals their taxonomic affinities</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Dehon, Manuel</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Engel, Michael S.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Gerard, Maxence</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Aytekin, A. Murat</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ghisbain, Guillaume</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Williams, Paul H.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rasmont, Pierre</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Michez, Denis</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2019</mods:date>
<mods:detail type="volume">
<mods:number>891</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>71</mods:start>
<mods:end>118</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.891.36027</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.891.36027</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-891-71</mods:identifier>
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<treatment ID-GBIF-Taxon="160511255" LSID="urn:lsid:plazi:treatment:81A479F7619E50A88342300A233BDFB5" httpUri="http://treatment.plazi.org/id/81A479F7619E50A88342300A233BDFB5" lastPageNumber="71" pageId="0" pageNumber="71">
<subSubSection pageId="0" pageNumber="71" type="nomenclature">
<paragraph pageId="0" pageNumber="71">
<taxonomicName LSID="81A479F7-619E-50A8-8342-300A233BDFB5" authority="Dehon, De Meulemeester &amp; Engel, 2014" authorityName="Dehon, De Meulemeester &amp; Engel" authorityYear="2014" class="Insecta" family="Apidae" genus="Bombus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bombus (Melanobombus) cerdanyensis" order="Hymenoptera" pageId="0" pageNumber="71" phylum="Arthropoda" rank="species" species="cerdanyensis" subGenus="Melanobombus">Bombus (Melanobombus) cerdanyensis Dehon, De Meulemeester &amp; Engel, 2014</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="71" type="holotype">
<paragraph pageId="0" pageNumber="71">Holotype.</paragraph>
<paragraph pageId="0" pageNumber="71">
Sex unknown. Conserved in the Paleontology department collection,
<normalizedToken originalValue="Muséum">Museum</normalizedToken>
national
<normalizedToken originalValue="dHistoire">d'Histoire</normalizedToken>
naturelle, Paris, France. The fossil consists of a part and counterpart. Type specimen has been located and revised (
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Representative fossil bumble bees A Bombus (Cullumanobombus) trophonius (photograph by Jakup Prokop) B B. (Cullumanobombus) randeckensis (photograph by Torsten Wappler) C B. vetustus (photograph by Alexandr P. Rasnitsyn) D B. (Cullumanobombus) pristinus (photograph by Irene Zorn and Monika Brueggeman-Ledolter) E B. (Melanobombus) cerdanyensis (photograph by Thibaut De Meulemeester)." figureDoi="10.3897/zookeys.891.36027.figure2" httpUri="https://binary.pensoft.net/fig/358114" pageId="0" pageNumber="71" tableDoi="10.3897/zookeys.891.36027.figure2">Figs 2E</figureCitation>
,
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Forewing drawings of the fossil bumble bees studied herein. Some forewings were mirrored to enable comparison across all specimens A Oligobombus cuspidatus (mirrored) B Holotype of Calyptapis florissantensis (mirrored) C C. florissantensis D Bombus (Paraelectrobombus) patriciae (mirrored) E B. (Mendacibombus) beskonakensis F B. (Cullumanobombus) trophonius (mirrored) G B. (Cullumanobombus) randeckensis (mirrored) H B. vetustus I B. (Cullumanobombus) pristinus (mirrored) J B. (Melanobombus) cerdanyensis." figureDoi="10.3897/zookeys.891.36027.figure3" httpUri="https://binary.pensoft.net/fig/358115" pageId="0" pageNumber="71" tableDoi="10.3897/zookeys.891.36027.figure3">3I</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="71" type="type strata and locality">
<paragraph pageId="0" pageNumber="71">Type strata and locality.</paragraph>
<paragraph pageId="0" pageNumber="71">Late Miocene (i.e., 10.0 Ma), lacustrine beds of Cerdanya, Spain.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="71" type="diagnosis">
<paragraph pageId="0" pageNumber="71">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="71">
Forewing membrane with alar papillae beyond apical crossveins; membrane infuscate, particularly in area beyond apical crossveins and along anterior borders of radial and marginal cells; pterostigma small, trapezoidal, not larger relative to prestigma and width not much shorter than length; marginal cell longer than distance from apex to forewing tip, tapering in width across its length, with apex acutely rounded and slightly offset from forewing margin; three submarginal cells of approximately same size, anterior borders of second and third submarginal cells subequal; 1m-cu angulate anteriorly, meeting second submarginal cell near midpoint; 2m-cu slightly arched, meeting third submarginal cell in apical fifth; mesotibia five times longer than wide; transector vein visible in the first submarginal cell. See
<bibRefCitation DOI="https://doi.org/10.1371/journal.pone.0108865" author="Dehon, M" journalOrPublisher="Annals of the Entomological Society of America" pageId="0" pageNumber="71" refId="B23" refString="Dehon, M, Michez, D, Nel, A, Engel, MS, De Meulemeester, T, 2014. .https://doi.org/10.1371/journal.pone.0108865" url="https://doi.org/10.1371/journal.pone.0108865" year="2014">Dehon et al. (2014)</bibRefCitation>
for original diagnosis.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="71" type="description">
<paragraph pageId="0" pageNumber="71">Description.</paragraph>
<paragraph pageId="0" pageNumber="71">
Fossil compressed in apparently dorsal oblique view, with left forewing outstretched; right forewing not preserved; hind wings not preserved; prosoma not preserved; mesosoma and metasoma incomplete and damaged; mid and hind legs preserved, partially overlapping forewing; right profemur length 1.4 m, width 0.8 mm as preserved; left mesofemur length 3.6 mm, width 0.9 mm; mesotibia length 3.0 mm, width 0.6 mm; mesobasitarsus length 3.2 mm, width 0.9 mm; remaining tarsomeres and pretarsal claws well preserved; pretarsal claws apparently not toothed as preserved; right mesofemur length 3.5 mm, width 0.5 mm; mesotibia length 2.0 mm, width 0.4 mm as preserved; left forewing length 13.3 mm, maximum width 4.6 mm; three submarginal cells of similar size; first submarginal cell length 1.5 mm (as measured from origin of Rs+M to juncture of r-rs and Rs), heigth 0.7 mm (as measured from Rs+M to pterostigma); second submarginal cell length 1.5 mm (as measured from juncture of Rs+M and M to juncture of Rs and 1rs-m), height 0.8 mm (as measured from midpoint on M between 1m-cu and 1rs-m to juncture of r-rs and Rs); third submarginal cell length 1.3 mm (as measured from juncture of 1rs-m and M to juncture of M and 2rs-m), height 1.1 mm (as measured from juncture of M and 2m-cu to juncture of 2rs-m and Rs); first medial cell length 3.4 mm (as measured from juncture of M+Cu and Cu to juncture of 1m-cu and M), height 1.2 mm (as measured from juncture of M and Rs+M to midpoint on Cu between M+Cu and 1m-cu); pterostigma length 0.9 mm; marginal cell length 3.4 mm with apex rounded, offset from anterior wing margin, not appendiculate; 1m-cu strongly curved, meeting second submarginal cell near midpoint; 2m-cu slightly arched, meeting third submarginal cell in apical fifth; metasoma width 5.8 mm as preserved; first two segments visible, first segment length 1.8 mm, second segment length 1.2 mm as preserved. See
<bibRefCitation DOI="https://doi.org/10.1371/journal.pone.0108865" author="Dehon, M" journalOrPublisher="Annals of the Entomological Society of America" pageId="0" pageNumber="71" refId="B23" refString="Dehon, M, Michez, D, Nel, A, Engel, MS, De Meulemeester, T, 2014. .https://doi.org/10.1371/journal.pone.0108865" url="https://doi.org/10.1371/journal.pone.0108865" year="2014">Dehon et al. (2014)</bibRefCitation>
for original description.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="71" type="comments">
<paragraph pageId="0" pageNumber="71">Comments.</paragraph>
<paragraph pageId="0" pageNumber="71">
The attribution based on geometric morphometric analysis (i.e.,
<taxonomicName class="Insecta" family="Apidae" genus="Melanobombus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Melanobombus" order="Hymenoptera" pageId="0" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="71">Melanobombus</emphasis>
</taxonomicName>
) is consistent with the timing and geographic origin of the subgenus proposed by
<bibRefCitation DOI="https://doi.org/10.1080/10635150801898912" author="Hines, HM" journalOrPublisher="Systematic Biology" pageId="0" pageNumber="71" pagination="58 - 75" refId="B55" refString="Hines, HM, 2008. Historical biogeography, divergence times, and diversification patterns of bumblebees (Hymenoptera: Apidae: Bombus). Systematic Biology 57: 58 - 75" title="Historical biogeography, divergence times, and diversification patterns of bumblebees (Hymenoptera: Apidae: Bombus)." url="https://doi.org/10.1080/10635150801898912" volume="57" year="2008">Hines (2008)</bibRefCitation>
. Indeed, the fossil was found in the Upper Miocene (i.e., 10.0 Ma) deposit of La Cerdanya in Spain, while
<taxonomicName class="Insecta" family="Apidae" genus="Melanobombus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Melanobombus" order="Hymenoptera" pageId="0" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="71">Melanobombus</emphasis>
</taxonomicName>
is estimated to have originated between 20.0-15.0 Ma in the Old World. The relative sizes of the prestigma and pterostigma exclude a placement in the
<taxonomicName lsidName="" pageId="0" pageNumber="71" rank="tribe" tribe="Electrobombini">Electrobombini</taxonomicName>
(although the presence or absence of a jugal lobe in the hind wing cannot be determined in the holotype). The forewing is apically papillate (as in
<taxonomicName lsidName="" pageId="0" pageNumber="71" rank="tribe" tribe="Bombini">Bombini</taxonomicName>
), and the marginal cell is not appendiculate and 1m-cu is strongly angulate together suggesting the species does not belong to the
<taxonomicName lsidName="" pageId="0" pageNumber="71" rank="tribe" tribe="Electrapini">Electrapini</taxonomicName>
or
<taxonomicName lsidName="" pageId="0" pageNumber="71" rank="tribe" tribe="Melikertini">Melikertini</taxonomicName>
(although some melikertines have 1m-cu more angulate, such as
<taxonomicName class="Insecta" family="Apidae" genus="Melissites" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Melissites trigona" order="Hymenoptera" pageId="0" pageNumber="71" phylum="Arthropoda" rank="species" species="trigona">
<emphasis italics="true" pageId="0" pageNumber="71">Melissites trigona</emphasis>
</taxonomicName>
Engel, 1m-cu is always much shorter and not as long as in
<taxonomicName lsidName="" pageId="0" pageNumber="71" rank="tribe" tribe="Bombini">Bombini</taxonomicName>
or
<taxonomicName lsidName="" pageId="0" pageNumber="71" rank="tribe" tribe="Euglossini">Euglossini</taxonomicName>
; a long 1m-cu is more plesiomorphic among Corbiculata). Indeed, the forewings of the present fossil are distinctly
<taxonomicName class="Insecta" family="Apidae" genus="Bombus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bombus" order="Hymenoptera" pageId="0" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="71">Bombus</emphasis>
</taxonomicName>
-like: presence of papillae, general infuscation of the membrane, three submarginal cells of relatively similar size (albeit the latter character is assuredly plesiomorphic). Based on the specimen morphology and forewing shape affinities, the fossil is likely an extinct species of
<taxonomicName class="Insecta" family="Apidae" genus="Melanobombus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Melanobombus" order="Hymenoptera" pageId="0" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="71">Melanobombus</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>