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<document id="8FB249AE2F627222A7658B1F96606CF8" ID-CLB-Dataset="33255" ID-DOI="http://dx.doi.org/10.3897/zookeys.667.10716" ID-GBIF-Dataset="c2ffd9e3-afca-4942-9d2c-59758231b258" ID-PMC="PMC5523388" ID-Pensoft-Pub="1313-2970-667-95" ID-PubMed="28769637" ID-ZooBank="648D539FD99443188CD23F183172350C" ModsDocAuthor="" ModsDocDate="2017" ModsDocID="1313-2970-667-95" ModsDocOrigin="ZooKeys 667" ModsDocTitle="Phylogeny of the genus Yumtaax Boucher (Coleoptera, Passalidae, Proculini): Taxonomic and evolutionary implications with descriptions of three new species" checkinTime="1491908628330" checkinUser="pensoft" docAuthor="Beza-Beza, Cristian Fernando, Beck, James, Reyes-Castillo, Pedro &amp; Jameson, Mary Liz" docDate="2017" docId="5C592DF4EA8BC958B81B45901FAC1F67" docLanguage="en" docName="ZooKeys 667: 95-129" docOrigin="ZooKeys 667" docSource="http://dx.doi.org/10.3897/zookeys.667.10716" docTitle="Yumtaax recticornis Burmeister 1847" docType="treatment" docVersion="6" lastPageNumber="110" masterDocId="FF884937582ECA1CFFB79255C20EAF35" masterDocTitle="Phylogeny of the genus Yumtaax Boucher (Coleoptera, Passalidae, Proculini): Taxonomic and evolutionary implications with descriptions of three new species" masterLastPageNumber="129" masterPageNumber="95" pageNumber="106" updateTime="1701374162230" updateUser="ExternalLinkService">
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<mods:title id="8C21E81C474B172422F52142EDBAD812">Phylogeny of the genus Yumtaax Boucher (Coleoptera, Passalidae, Proculini): Taxonomic and evolutionary implications with descriptions of three new species</mods:title>
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<mods:namePart id="2F986942E9E9FFD681957645C255D285">Beza-Beza, Cristian Fernando</mods:namePart>
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<mods:namePart id="318F1C2E11C1EB533A37280DDC477E06">Beck, James</mods:namePart>
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<mods:namePart id="6E45F7F135071AE2A2A59BA25345AED7">Reyes-Castillo, Pedro</mods:namePart>
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<mods:namePart id="9961032F8ABF80DEF1BA0BEABC62E6CB">Jameson, Mary Liz</mods:namePart>
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<mods:date id="3321D193F6F56A9D80EB9DAC4399A3E3">2017</mods:date>
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<treatment id="5C592DF4EA8BC958B81B45901FAC1F67" ID-GBIF-Taxon="128345061" LSID="urn:lsid:plazi:treatment:5C592DF4EA8BC958B81B45901FAC1F67" httpUri="http://treatment.plazi.org/id/5C592DF4EA8BC958B81B45901FAC1F67" lastPageId="15" lastPageNumber="110" pageId="11" pageNumber="106">
<subSubSection id="DD13FB334FED8F06F2391DED20B0C00B" pageId="11" pageNumber="106" type="nomenclature">
<paragraph id="67750BDF369ECD8B2FCAEBC3EC3764DF" pageId="11" pageNumber="106">
<taxonomicName id="CCB1FE17BF0171E063081788DE515DAB" ID-CoL="5CTFK" authority="Burmeister, 1847" authorityName="Burmeister" authorityYear="1847" class="Insecta" family="Passalidae" genus="Yumtaax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yumtaax recticornis" order="Coleoptera" pageId="11" pageNumber="106" phylum="Arthropoda" rank="species" species="recticornis">Yumtaax recticornis (Burmeister, 1847)</taxonomicName>
</paragraph>
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<subSubSection id="63A0E38E9293FF1BE7B4CEE868041EF7" pageId="11" pageNumber="106" type="reference_group">
<paragraph id="9CCA664B246914E0BCF110EA47779E7D" pageId="11" pageNumber="106">
<taxonomicName id="BDFB74CC9BA89322B8A087E0CBF3BB3E" class="Insecta" family="Passalidae" genus="Passalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Passalus recticornis" order="Coleoptera" pageId="11" pageNumber="106" phylum="Arthropoda" rank="species" species="recticornis">Passalus recticornis</taxonomicName>
Burmeister, 1847: 508-509.
</paragraph>
<paragraph id="E084974863658CF98EC6537B62E55DCB" pageId="11" pageNumber="106">
<taxonomicName id="CEF27AF28866B515B416E86B23FAC4C5" class="Insecta" family="Passalidae" genus="Soranus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Soranus recticornis" order="Coleoptera" pageId="11" pageNumber="106" phylum="Arthropoda" rank="species" species="recticornis">Soranus recticornis</taxonomicName>
(Burmeister) [comb. n. by
<bibRefCitation id="EF0E38DDD338D1BF046D32F51D8EE79B" author="Kaup, JJ" journalOrPublisher="Berliner Entomologische zeitschrift," pageId="29" pageNumber="124" pagination="1 - 125" title="Monographie der Passaliden." url="https://doi.org/10.1002/mmnd.18710150306" volume="15" year="1871">Kaup 1871</bibRefCitation>
: 105, 108].
</paragraph>
<paragraph id="2D92ACD56B766E2CEE5DCDCDC12AB085" pageId="11" pageNumber="106">
<taxonomicName id="C6AE4ED72D972614A9CC6830104834D6" class="Insecta" family="Passalidae" genus="Popilius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Popilius recticornis" order="Coleoptera" pageId="11" pageNumber="106" phylum="Arthropoda" rank="species" species="recticornis">Popilius recticornis</taxonomicName>
(Burmeister) [comb. n. by
<bibRefCitation id="2629BCAA6F2C341E687D3F37A891E469" author="Gravely, FH" journalOrPublisher="Memoirs of the Indian Museum" pageId="29" pageNumber="124" pagination="1 - 144" title="A contribution towards the revision of the Passalidae of the World." url="https://doi.org/10.5962/bhl.title.9238" volume="7" year="1918">Gravely 1918</bibRefCitation>
: 24, 26].
</paragraph>
<paragraph id="4C9A3110F49DD361CAA312E7AB932438" pageId="11" pageNumber="106">
<taxonomicName id="5AA127892442898E4AE0E51A9AC33850" class="Insecta" family="Passalidae" genus="Petrejoides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Petrejoides recticornis" order="Coleoptera" pageId="11" pageNumber="106" phylum="Arthropoda" rank="species" species="recticornis">Petrejoides recticornis</taxonomicName>
(Burmeister) [comb. n. by
<bibRefCitation id="0372CD4F19C6FCC0EECD789F6B6F7ECC" pageId="11" pageNumber="106">Reyes-Castillo 1970</bibRefCitation>
: 125].
</paragraph>
<paragraph id="C721689416C45230424912B05CF7363B" pageId="11" pageNumber="106">
<taxonomicName id="07710C9D9479D86EC15AF95E561E2CBF" class="Insecta" family="Passalidae" genus="Yumtaax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yumtaax recticornis" order="Coleoptera" pageId="11" pageNumber="106" phylum="Arthropoda" rank="species" species="recticornis">Yumtaax recticornis</taxonomicName>
(Burmeister) [comb. n. by
<bibRefCitation id="F53117663DCB1FE1D17C23A7EA490D34" author="Boucher, S" journalOrPublisher="Annale de la Societe Entomologique de France" pageId="28" pageNumber="123" pagination="237 - 604" title="Evolution et phylogenie des coleopteres Passalidae (Scarabaeoidea)." url="https://doi.org/10.1080/00379271.2005.10697444" volume="41" year="2006">Boucher 2006</bibRefCitation>
: 348].
</paragraph>
</subSubSection>
<subSubSection id="DE117B761B2D37A0F12E499B7D101251" lastPageId="12" lastPageNumber="107" pageId="11" pageNumber="106" type="materials_examined">
<paragraph id="B88F0490B7B325DAFEB0F9E4C0FFCCAE" pageId="11" pageNumber="106">Material examined.</paragraph>
<paragraph id="015609AC6D34E8F46E44D02C0C3C51EF" pageId="11" pageNumber="106">105 specimens (lectotype, paralectotype, and 103 non-type specimens). Lectotype ♂. MEXICO: WB zoologie S. Nr. 812123. (MLU Halle). Paralectotype 1 ♀. MEXICO: no data (MLU Halle).</paragraph>
<paragraph id="A3185E0630517D8A9DB489A14E6A38B3" lastPageId="12" lastPageNumber="107" pageId="11" pageNumber="106">
Non-type specimens (103 total). MEXICO: 1 ♀, Oaxaca, Amatepec (1.6 km N), alt. 1840 m, bosque
<normalizedToken id="A963A2D5E4288578A4777474A8982E40" originalValue="mesófilo">mesofilo</normalizedToken>
de
<normalizedToken id="007ED26F67FDBA875D308067469CA047" originalValue="montaña">montana</normalizedToken>
, II-28-1988 (Reyes, Boucher, C. Castillo). 1 ♂, Carretera Tuxtepec-Oaxaca (87 km), III-21-1967 (R. MacGregor); 1 ♀, (88 km), alt. 2350 m, IV-4-1986 (A. Ibarra); 1 ♂, 1 ♀, (119 km), X-6-1973 (J. Mateu); 4 ♂, 2 ♀, (153 km), alt. 2800 m, X-7-1973 (J. Mateu). 1 ♀, Cerro
<normalizedToken id="9E7551F5265B7283A1953F463EFE3306" originalValue="Pelón">Pelon</normalizedToken>
(2.8 km), V-18-1980 (C. Castillo, M. L. Castillo, G. Quintero, E. Rivera); 3 ♀, (11.4 km NE), alt. 2170 m (P. Reyes et al.). 1 ♀, Comaltepec, Brecha 60 (unknown locality), V-18-1980 (C. Castillo, M. L. Castillo, G. Quintero, E. Rivera); 3 ♂, 2 ♀, (4.5 km), V-18-1980 (C. Castillo, M. L. Castillo, G. Quintero, E. Rivera). 1 ♂, 4 ♀, Comaltepec, Galera San Isidro (800 m), alt. 2000 m, V-17-1980 (C. Castillo, M. L. Castillo, G. Quintero, E. Rivera); 3 ♂, 5 ♀, (3.6 km), alt. 2160 m, V-17-1980 (C. Castillo, M. L. Castillo, G. Quintero, E. Rivera). 1 ♀, La Esperanza, alt. 1670 m, V-16-1980 (C. Castillo, G. Quintero, M. L. Castillo, E. Rivera); 4 ♂, 3 ♀, (3.5 km N), alt. 1670 m, bosque
<normalizedToken id="806431031673D4FE67AEE62FA1A09B37" originalValue="mesófilo">mesofilo</normalizedToken>
de
<pageBreakToken id="A2BCD6ACC52017A10000EACDFAF9A533" pageId="12" pageNumber="107" start="start">
<normalizedToken id="9E7A48DA3274D2F0BBF980CFFC17CA94" originalValue="montaña">montana</normalizedToken>
</pageBreakToken>
, III-1-1988 (Reyes, Boucher, Castillo); 1 ♀ (4 km), alt. 1800 m, V-20-1980 (C. Castillo, M. L. Castillo, G. Quintero, E. Rivera); 5 sex unknown, (6.8 km), alt. 1820 m, II-25-1984 (P. Reyes et al.); 1 sex unknown, (13.1 km), alt. 2030 m, II-25-1984 (P. Reyes et al.); 3 sex unknown, (14.1 km), alt. 1985 m, II-25-1984 (P. Reyes et al.); 2 sex unknown, (103.1 km), alt. 2030 m, II-25-1984 (P. Reyes et al.). 1 sex unknown, San Juan Lachao Viejo, km 85 de Sola de Vega (N 16°13.220' W 97°08.913), alt. 1858-1870 m, bosque
<normalizedToken id="322D1592627DC9D0439F7B67FEC03ED5" originalValue="mesófilo">mesofilo</normalizedToken>
de
<normalizedToken id="32B279293E084ABEF96CEF2DC4CF5559" originalValue="montaña">montana</normalizedToken>
, VIII-6-2004 (K. Araya). 30 sex unknown, San Miguel Talea de Castro (8 km SE) (N 17°19.620' W 96°17.403), alt. 2082 m, VII-22-2007 (O. Francke, H.
<normalizedToken id="20BAA97B4CE456E35E9212178869146C" originalValue="Montaño">Montano</normalizedToken>
, A.
<normalizedToken id="7A74CA20B0AEA1E998296B07CBBC1B3A" originalValue="Valdéz">Valdez</normalizedToken>
, C.
<normalizedToken id="70CDF3255FCCAB9D3274AB5F4023A95D" originalValue="Santibañez">Santibanez</normalizedToken>
, A. Ballesteros). 3 ♂, 4 ♀, Sierra de
<normalizedToken id="B6FEA5B75AF4C1B082F04EF1F44AEF15" originalValue="Juárez">Juarez</normalizedToken>
, alt. 2000 m, VI-2-1995 (G. Nogueira). 2 ♂, 1 ♀, Totontepec (3.4 km S), alt. 1940 m, bosque
<normalizedToken id="8557EFA386293AA4E21E635C924F1723" originalValue="mesófilo">mesofilo</normalizedToken>
de
<normalizedToken id="284DBEC40E4E7AD4361917C9B6F4BB45" originalValue="montaña">montana</normalizedToken>
, II-29-1988 (Reyes, Boucher, Castillo). 3 ♂, 3 ♀, Valle Nacional (32 miles S), V-21/24-1971 (H. Howden); 1 ♀, (105-117 km E), IV-1964 (C. R. Rotger).
</paragraph>
</subSubSection>
<subSubSection id="78D888F0B1583EF5DADB16D75D70F987" pageId="12" pageNumber="107" type="diagnosis">
<paragraph id="F165C2B85503DF6D8EE0792BE0C65A86" pageId="12" pageNumber="107">Diagnosis.</paragraph>
<paragraph id="AABDFDB3D2B59EB1A394A28312B57A57" pageId="12" pageNumber="107">
<taxonomicName id="8CDC4E79B04307EA1027BA516C195998" class="Insecta" family="Passalidae" genus="Yumtaax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yumtaax recticornis" order="Coleoptera" pageId="12" pageNumber="107" phylum="Arthropoda" rank="species" species="recticornis">Yumtaax recticornis</taxonomicName>
is a small (18.0-21.0 mm), macropterous species and is a member of the
<taxonomicName id="3BE9B329F4F9C7BB796A64B464710A25" lsidName="Y. imbellis" pageId="12" pageNumber="107" rank="species" species="imbellis">Y. imbellis</taxonomicName>
clade (Fig. 4). This species is diagnosed by the following character combination: the clypeus is inclined (shared with
<taxonomicName id="B4BC8799ACE3AFE13AF4F4FC4A73A9FC" lsidName="Y. jimenezi" pageId="12" pageNumber="107" rank="species" species="jimenezi">Y. jimenezi</taxonomicName>
,
<taxonomicName id="05C1C37D5823369A3608C83C1EF01721" lsidName="Y. imbellis" pageId="12" pageNumber="107" rank="species" species="imbellis">Y. imbellis</taxonomicName>
,
<taxonomicName id="2C2CD742AFCEB0A13CDC29989310F639" lsidName="Y. mazatecus" pageId="12" pageNumber="107" rank="species" species="mazatecus">Y. mazatecus</taxonomicName>
,
<taxonomicName id="D5C2AAFB4503A86629BF54062D34A3FD" lsidName="Y. nebulosus" pageId="12" pageNumber="107" rank="species" species="nebulosus">Y. nebulosus</taxonomicName>
,
<taxonomicName id="6834958D9BF0EDD1CC67B0CD0237EBAE" lsidName="Y. olmecae" pageId="12" pageNumber="107" rank="species" species="olmecae">Y. olmecae</taxonomicName>
; clypeus vertical in
<taxonomicName id="212FD71CC5B19984E05B3DCC661F1D8F" lsidName="Y. veracrucensis" pageId="12" pageNumber="107" rank="species" species="veracrucensis">Y. veracrucensis</taxonomicName>
,
<taxonomicName id="620461C3BC1C7EA17F8A468637236060" lsidName="Y. laticornis" pageId="12" pageNumber="107" rank="species" species="laticornis">Y. laticornis</taxonomicName>
,
<taxonomicName id="79FCFB88026AC615C3FEDAEEA62258E4" lsidName="Y. cameliae" pageId="12" pageNumber="107" rank="species" species="cameliae">Y. cameliae</taxonomicName>
), surface concave (flat in other members of
<taxonomicName id="824A049C4DC7AE6B8BBCAEFA9F798261" class="Insecta" family="Passalidae" genus="Yumtaax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yumtaax" order="Coleoptera" pageId="12" pageNumber="107" phylum="Arthropoda" rank="genus">Yumtaax</taxonomicName>
), and the anterior border is concave (shared with
<taxonomicName id="B7ECB6BA5A3F7358DAAE98C254C34EA5" lsidName="Y. olmecae" pageId="12" pageNumber="107" rank="species" species="olmecae">Y. olmecae</taxonomicName>
; flat in other members of
<taxonomicName id="6AC93870DBAA7FD0E3214372B474AEC0" class="Insecta" family="Passalidae" genus="Yumtaax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yumtaax" order="Coleoptera" pageId="12" pageNumber="107" phylum="Arthropoda" rank="genus">Yumtaax</taxonomicName>
); mesofrontal structure (MFS) of the
<normalizedToken id="BAE6901B160A94DB95A7E62006737721" originalValue="“falsus”">&quot;falsus&quot;</normalizedToken>
type (see
<bibRefCitation id="6268B315220C4409EE84F9833D77D6F4" pageId="12" pageNumber="107">Reyes-Castillo 1970</bibRefCitation>
) (shared with all members of
<taxonomicName id="9B9F109F9AD0D795F9A9EAFB62B2EB84" class="Insecta" family="Passalidae" genus="Yumtaax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yumtaax" order="Coleoptera" pageId="12" pageNumber="107" phylum="Arthropoda" rank="genus">Yumtaax</taxonomicName>
except
<taxonomicName id="38171C20F14BC4CBF78C2456936D73E8" lsidName="Y. cameliae" pageId="12" pageNumber="107" rank="species" species="cameliae">Y. cameliae</taxonomicName>
that has a MFS of the
<normalizedToken id="0F852F8990BCCC10AF495D240AB96EB3" originalValue="“striatopunctatus”">&quot;striatopunctatus&quot;</normalizedToken>
type) with the central tooth largely free (shared with
<taxonomicName id="24CE43BC7B6F7BEB2C425B42182CDBC9" lsidName="Y. veracrucensis" pageId="12" pageNumber="107" rank="species" species="veracrucensis">Y. veracrucensis</taxonomicName>
,
<taxonomicName id="57B7A730BD8BBD1818F217C05BE270CC" lsidName="Y. laticornis" pageId="12" pageNumber="107" rank="species" species="laticornis">Y. laticornis</taxonomicName>
,
<taxonomicName id="B6B8B0CEB0E443614135F325DD5BBFC1" lsidName="Y. cameliae" pageId="12" pageNumber="107" rank="species" species="cameliae">Y. cameliae</taxonomicName>
,
<taxonomicName id="C7712674C906D5CC89D5EF79715F8E68" lsidName="Y. mazatecus" pageId="12" pageNumber="107" rank="species" species="mazatecus">Y. mazatecus</taxonomicName>
; fused with frontal ridges in
<taxonomicName id="F2DD2D5B52C211712DD83A164374BBBD" lsidName="Y. jimenezi" pageId="12" pageNumber="107" rank="species" species="jimenezi">Y. jimenezi</taxonomicName>
,
<taxonomicName id="B6952E1D770A541731A4130B93064D4E" lsidName="Y. imbellis" pageId="12" pageNumber="107" rank="species" species="imbellis">Y. imbellis</taxonomicName>
,
<taxonomicName id="96BC2407E4B7025A8752C4448FEC4498" lsidName="Y. nebulosus" pageId="12" pageNumber="107" rank="species" species="nebulosus">Y. nebulosus</taxonomicName>
,
<taxonomicName id="E38522AF6431A4525720D370F959175E" lsidName="Y. olmecae" pageId="12" pageNumber="107" rank="species" species="olmecae">Y. olmecae</taxonomicName>
), central tooth directed dorsally (shared with
<taxonomicName id="A56D1CCAD7FA93BC1F08D9B1622DE610" lsidName="Y. imbellis" pageId="12" pageNumber="107" rank="species" species="imbellis">Y. imbellis</taxonomicName>
,
<taxonomicName id="430737496D1E36AE9C79AC103DABCB2C" lsidName="Y. nebulosus" pageId="12" pageNumber="107" rank="species" species="nebulosus">Y. nebulosus</taxonomicName>
,
<taxonomicName id="B37B33BAFE8701C1B337BFB21791DEAA" lsidName="Y. olmecae" pageId="12" pageNumber="107" rank="species" species="olmecae">Y. olmecae</taxonomicName>
; directed dorsally and anteriorly in
<taxonomicName id="509FBCEBEE31BCD4C45EF4D19A98C868" lsidName="Y. jimenezi" pageId="12" pageNumber="107" rank="species" species="jimenezi">Y. jimenezi</taxonomicName>
,
<taxonomicName id="9BB8A0D1F74DC53E288733794854FC9A" lsidName="Y. mazatecus" pageId="12" pageNumber="107" rank="species" species="mazatecus">Y. mazatecus</taxonomicName>
; directed anteriorly in
<taxonomicName id="87E142B7BC3A4890C0148511057BFD42" lsidName="Y. veracrucensis" pageId="12" pageNumber="107" rank="species" species="veracrucensis">Y. veracrucensis</taxonomicName>
,
<taxonomicName id="1D11D0AF136DB2DACC9F9599DD8E3BA6" lsidName="Y. laticornis" pageId="12" pageNumber="107" rank="species" species="laticornis">Y. laticornis</taxonomicName>
; elevated in the posterior half bending abruptly forward in the anterior half in
<taxonomicName id="86CB07405634CB70868DC7B9A0E674D0" lsidName="Y. cameliae" pageId="12" pageNumber="107" rank="species" species="cameliae">Y. cameliae</taxonomicName>
), central tooth not reaching the anterior border of the frontoclypeal suture (shared with
<taxonomicName id="E16A8E6634683B3721C2468C4026A40D" lsidName="Y. jimenezi" pageId="12" pageNumber="107" rank="species" species="jimenezi">Y. jimenezi</taxonomicName>
,
<taxonomicName id="591A3AC8CFF77772347FC0370813E676" lsidName="Y. imbellis" pageId="12" pageNumber="107" rank="species" species="imbellis">Y. imbellis</taxonomicName>
,
<taxonomicName id="6D6DBE3B235B4C16B35FB43A20CAEA71" lsidName="Y. mazatecus" pageId="12" pageNumber="107" rank="species" species="mazatecus">Y. mazatecus</taxonomicName>
,
<taxonomicName id="6C9D9288F5F080A4F578808CF360C649" lsidName="Y. nebulosus" pageId="12" pageNumber="107" rank="species" species="nebulosus">Y. nebulosus</taxonomicName>
,
<taxonomicName id="7965296FE132527D73A7987D58E250B6" lsidName="Y. olmecae" pageId="12" pageNumber="107" rank="species" species="olmecae">Y. olmecae</taxonomicName>
; reaching the clypeus in
<taxonomicName id="AE8FACF829D7BFCB35F49050B5BF5678" lsidName="Y. laticornis" pageId="12" pageNumber="107" rank="species" species="laticornis">Y. laticornis</taxonomicName>
,
<taxonomicName id="3312DB036FDCDDDD2227E8A9648B47ED" lsidName="Y. cameliae" pageId="12" pageNumber="107" rank="species" species="cameliae">Y. cameliae</taxonomicName>
,
<taxonomicName id="51EAC7167864B60E9E76C4B5133139B1" lsidName="Y. veracrucensis" pageId="12" pageNumber="107" rank="species" species="veracrucensis">Y. veracrucensis</taxonomicName>
); and large eyes (shared with
<taxonomicName id="E87F8D7E948864373B5CF614F586533A" lsidName="Y. imbellis" pageId="12" pageNumber="107" rank="species" species="imbellis">Y. imbellis</taxonomicName>
; eyes moderately reduced in
<taxonomicName id="BEC042A66805719557990990F03A0684" lsidName="Y. veracrucensis" pageId="12" pageNumber="107" rank="species" species="veracrucensis">Y. veracrucensis</taxonomicName>
,
<taxonomicName id="9BD8A9B812D3A745F1483F941F36ABB6" lsidName="Y. cameliae" pageId="12" pageNumber="107" rank="species" species="cameliae">Y. cameliae</taxonomicName>
,
<taxonomicName id="2A8E7D7F0096D333172A9B615A575BD0" lsidName="Y. jimenezi" pageId="12" pageNumber="107" rank="species" species="jimenezi">Y. jimenezi</taxonomicName>
,
<taxonomicName id="00415696E1023A9DAD0774109B90255B" lsidName="Y. nebulosus" pageId="12" pageNumber="107" rank="species" species="nebulosus">Y. nebulosus</taxonomicName>
,
<taxonomicName id="599379CC8C14B50C763D32546973F620" lsidName="Y. olmecae" pageId="12" pageNumber="107" rank="species" species="olmecae">Y. olmecae</taxonomicName>
; strongly reduced in
<taxonomicName id="5FAB19DBD539AAD9996A25FDA914D4E2" lsidName="Y. laticornis" pageId="12" pageNumber="107" rank="species" species="laticornis">Y. laticornis</taxonomicName>
,
<taxonomicName id="0927D7559055723710439E497A5E4618" lsidName="Y. mazatecus" pageId="12" pageNumber="107" rank="species" species="mazatecus">Y. mazatecus</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="E9AA89E54BAEC5BBB687AFE901C0F6B2" pageId="12" pageNumber="107" type="description">
<paragraph id="CBDED8756EA06CB6100F9FA1A1599FDC" pageId="12" pageNumber="107">Dimensions</paragraph>
<paragraph id="E7B325384E5F109EEB223C66447E1681" pageId="12" pageNumber="107">
(mm) (n = 12). Total length 18.0-21.0 (
<normalizedToken id="B6EAC3238FE1E1DD8E480706DF144E0B" originalValue="χ">x</normalizedToken>
= 19.0); elytral length 10.0-11.5 (
<normalizedToken id="45EFAC386E3CBB515C322693987CA058" originalValue="χ">x</normalizedToken>
= 11.0); pronotal length 3.5-5.0 (
<normalizedToken id="CA4D44B31D7320BA68169C1243D94EBA" originalValue="χ">x</normalizedToken>
= 4.5); pronotal width 5.5-6.0 (
<normalizedToken id="CEE0E3F6B1AA45345A61F7142415DB76" originalValue="χ">x</normalizedToken>
= 5.5); humeral width 5.5-6.5 (
<normalizedToken id="F6EEC77E1FEE576C286A11510CC988F6" originalValue="χ">x</normalizedToken>
= 6.0).
</paragraph>
</subSubSection>
<subSubSection id="83109DC481BE4F6CEB1F53DB424407DA" lastPageId="14" lastPageNumber="109" pageId="12" pageNumber="107" type="redescription">
<paragraph id="6172BC96EF54FA855A99A429A8E55433" pageId="12" pageNumber="107">Redescription of lectotype</paragraph>
<paragraph id="6964306A4081FEB62A648DAAE05ED45E" lastPageId="14" lastPageNumber="109" pageId="12" pageNumber="107">
(Fig. 5). Head (Fig. 5C). Labrum: anterior border concave, dorsal surface smooth and glabrous medially, punctate and setose apicolaterally, apically, and basally; anterior edge excavated. Clypeus: inclined, rectangular, concave, shiny, and smooth. Frontoclypeal suture: concave and opaque; external tubercles rounded, directed anteriorly and laterally. Frontal area: inclined, concave, smooth and shiny; frontal ridges present without inner tubercles. Frontal fossae: punctate and setose. Mesofrontal structure (MFS): of the
<normalizedToken id="D978E759BEAA00C684B8D9681EAC3137" originalValue="“falsus”">&quot;falsus&quot;</normalizedToken>
type (see
<bibRefCitation id="EB4B907B45E28355F655C9FE6B0A96C0" pageId="12" pageNumber="107">Reyes-Castillo 1970</bibRefCitation>
); base subparallel and as wide as the lateral ridge of MFS; center horn short with apex rounded, largely free and directed anteriorly and dorsally (Fig. 5D), not reaching the
<pageBreakToken id="626CDEA4B658518308CD497B4D4CC50C" pageId="13" pageNumber="108" start="start">posterior</pageBreakToken>
margin of clypeus, dorsally without micro-punctures; base of center horn wide, narrowing gradually until apex; dorsal fossa present at the base of MFS. Occipital fossa: shallow posteriorly and deeper laterally, connected with the frontal fossae. Posterior occipital sulcus biconcave. Supraorbital ridge: bituberculate, anterior tubercle larger than posterior tubercle; posterior half of supraorbital ridge not bifurcated. Canthus: with apex rounded, covering less than 1/3 of eye, not expanded distally. Eyes large (distal edge of the eye surpassing distal edge of the canthus), width = 0.5 mm (each eye).
<pageBreakToken id="C937186BF18CD495960B980BE532A39B" pageId="14" pageNumber="109" start="start">Head</pageBreakToken>
width (between posterior tubercles of the supraorbital ridge) = 3.0 mm. Ratio of sums of both eye widths/total head width = 0.36; postocular area punctate and setose. Ligula: tridentate, central tooth surpassing apex of lateral teeth, lateral teeth rounded; setose punctures present in discal area; posterior border convex. Mentum: lateral lobes rounded and wide, with setose punctures. Basomedial portion protruding ventrally; anterior border at middle convex; basal fossae present and rugose. Hypostomal process: with lateral depression; separated from mentum by a distance shorter than the width of the anterior width of hypostomal process. Infraocular ridge (e.g., Fig. 3): short and narrow anteriorly. Mandible: with 3 apical teeth; internal tooth of left mandible bidentate (teeth 1 and 2 of internal tooth fused); dorsal tooth occupies less than half length of the mandible. Pronotum: anterior angles rounded. Anterior fossae of marginal sulcus punctate. Lateral fossae with scattered punctures. Marginal groove lacking punctures. Prosternum: opaque; prosternellum with anterior half and lateral edges opaque and posterior half and middle area shiny. Scutellum: smooth and glabrous. Mesosternum: with lateral areas opaque. Metasternum: with setae anterolaterally, lacking punctures in lateral margins of metasternal disc. Lateral fossae wide across metathorax, with setose punctures. Elytra: anterior border straight. Meeting point of striae 1-10 (see
<bibRefCitation id="D9D957E9AEF844A1A83D6A673B225B67" pageId="14" pageNumber="109">Reyes-Castillo 1970</bibRefCitation>
) with one line of punctures. Wings: well developed. Legs (Fig. 5E): femur I with longitudinal anteroventral groove weakly developed, not reaching distal end of femur, posteroventral half pubescent; setae long, dense, reddish. Abdomen: last sternite with marginal groove complete (Fig. 5B). Aedeagus (Fig. 5F, G, and H) (description based on non-type material): dorsal view phallus elongated (longer than wider). In ventral view distal edges of the phallus more or less at the same level of distal edges of parameres.
</paragraph>
</subSubSection>
<subSubSection id="D411459AC275CC2D755434B8148A7FB3" pageId="14" pageNumber="109" type="variation">
<paragraph id="A148DC105F4EBA0517A3EB9A5191804C" pageId="14" pageNumber="109">Variation.</paragraph>
<paragraph id="6FEBBE8A82D4102FE23A50895ED37976" pageId="14" pageNumber="109">The paralectotype and other specimens vary from the lectotype by the following characteristics: frontoclypeal suture varies from opaque to shiny, from concave to almost straight; frons and clypeus inclined to nearly vertical (always concave); concavity of frons and clypeus vary from strongly developed to nearly flat. Internal tubercles strongly developed or absent; frontal ridges always present, but not always terminating in internal tubercles; ratio of eyes and head width varies from 0.32-0.57; supraocular ridge weakly developed or absent. Occipital sulcus biconcave to concave in a few individuals. Small portion of individuals with frontal ridges fused to the base of the central horn of MFS (apex of the central horn always free).</paragraph>
</subSubSection>
<subSubSection id="1F12A3706F76010ACA0A6AADC4191279" pageId="14" pageNumber="109" type="distribution">
<paragraph id="135F93B161632B919CCB0D0819195BD2" pageId="14" pageNumber="109">Distribution.</paragraph>
<paragraph id="4E260004E5A6A7B6FEA88B0FE93CDE0C" pageId="14" pageNumber="109">
The lectotype and paralectotype are labeled
<normalizedToken id="9680F9549165CE641796D3ADC15EAE21" originalValue="“Mexico”">&quot;Mexico&quot;</normalizedToken>
(
<bibRefCitation id="DFDFED157CD233F567ACFCA04F624A37" pageId="14" pageNumber="109">Burmeister 1847</bibRefCitation>
).
<bibRefCitation id="0ED2B78A722414F799F5B26252B07D09" author="Castillo, C" journalOrPublisher="Acta Zoologica Mexicana (nueva serie)" pageId="29" pageNumber="124" pagination="1 - 84" title="Biosistematica del genero Petrejoides Kuwert (Coleoptera, Lamellicornia, Passalidae)." volume="4" year="1984">Castillo and Reyes-Castillo (1984)</bibRefCitation>
suggested the Sierra de Juarez in Oaxaca, Mexico, as the possible type locality. The species is known only from cloud forest (1424-2986 m elevation) in Oaxaca, Mexico.
</paragraph>
</subSubSection>
<subSubSection id="CB0D30F51039786949AABB844D60C59B" lastPageId="15" lastPageNumber="110" pageId="14" pageNumber="109" type="remarks">
<paragraph id="489B16A69C396BEAEFB99DE2E691A0D6" pageId="14" pageNumber="109">Remarks.</paragraph>
<paragraph id="D9042365C08DB928004EA18E68245A78" lastPageId="15" lastPageNumber="110" pageId="14" pageNumber="109">
Originally, this species was thought to be widely distributed across the Sierra Madre Oriental, the Mexican Transvolcanic Belt, and Sierra Madre del Sur (
<bibRefCitation id="5D0C3D08501E381AFE2050C0799244C3" pageId="14" pageNumber="109">Reyes-Castillo 1970</bibRefCitation>
,
<bibRefCitation id="E7B47EF68B0C685E236D0D2388D077D1" author="Castillo, C" journalOrPublisher="Acta Zoologica Mexicana (nueva serie)" pageId="29" pageNumber="124" pagination="1 - 84" title="Biosistematica del genero Petrejoides Kuwert (Coleoptera, Lamellicornia, Passalidae)." volume="4" year="1984">Castillo and Reyes-Castillo 1984</bibRefCitation>
,
<bibRefCitation id="6B8922884DF462C299CBDB8C6A304989" author="Boucher, S" journalOrPublisher="Annale de la Societe Entomologique de France" pageId="28" pageNumber="123" pagination="237 - 604" title="Evolution et phylogenie des coleopteres Passalidae (Scarabaeoidea)." url="https://doi.org/10.1080/00379271.2005.10697444" volume="41" year="2006">Boucher 2006</bibRefCitation>
). Our phylogenetic analysis (Fig. 4), as well as close examination of morphological characters, provide evidence that
<taxonomicName id="8B2C8651D4B373E779B8C714F84AEF36" lsidName="Y. recticornis" pageId="14" pageNumber="109" rank="species" species="recticornis">Y. recticornis</taxonomicName>
s. l. is composed of at least two cryptic species:
<taxonomicName id="F76570E30C190B6D9B2F993C3958F79B" lsidName="Y. recticornis" pageId="14" pageNumber="109" rank="species" species="recticornis">Y. recticornis</taxonomicName>
(&quot;
<taxonomicName id="559865B49303D3747DABCD16667C6377" lsidName="Y. recticornis" pageId="14" pageNumber="109" rank="species" species="recticornis">Y. recticornis</taxonomicName>
OM&quot; in the
<taxonomicName id="622A17C1FA6FDB2D0440471B666A2E1D" lsidName="Y. imbellis" pageId="14" pageNumber="109" rank="species" species="imbellis">Y. imbellis</taxonomicName>
clade, Fig. 4 and Suppl. materials 2-4) and
<taxonomicName id="2B471674E6420EE7E8964FA1F124759B" lsidName="Y. jimenezi" pageId="14" pageNumber="109" rank="species" species="jimenezi">Y. jimenezi</taxonomicName>
sp. n.
<pageBreakToken id="3FC4F85447C933553FB5F4D6D8C50CB4" pageId="15" pageNumber="110" start="start">(</pageBreakToken>
&quot;
<taxonomicName id="53EE3E6D1266432804DD83FD08046F07" lsidName="Y. recticornis" pageId="15" pageNumber="110" rank="species" species="recticornis">Y. recticornis</taxonomicName>
VM&quot; in the
<taxonomicName id="69E1EEEF9580A9071F0DF5043161175A" lsidName="Y. laticornis" pageId="15" pageNumber="110" rank="species" species="laticornis">Y. laticornis</taxonomicName>
clade, Fig. 4 and Suppl. materials 2-4). Based on comparison with the lectotype and one paralectotype of
<taxonomicName id="BF532F71B7C666CE75532653ACC1D28F" lsidName="Y. recticornis" pageId="15" pageNumber="110" rank="species" species="recticornis">Y. recticornis</taxonomicName>
, this name should be applied to
<taxonomicName id="A4E9805E2B09BCCCD693638250D36420" lsidName="Y. recticornis" pageId="15" pageNumber="110" rank="species" species="recticornis">Y. recticornis</taxonomicName>
OM. The following character states provide evidence that
<taxonomicName id="3CC7DB72D64BB2644620540487FE8951" lsidName="Y. recticornis" pageId="15" pageNumber="110" rank="species" species="recticornis">Y. recticornis</taxonomicName>
OM is conspecific with
<normalizedToken id="39754DE1DDC46D12B63F92318B404CFA" originalValue="Burmeister’s">Burmeister's</normalizedToken>
concept of
<taxonomicName id="7A37FD252CFB5C0AA94493E78DED6DED" lsidName="Y. recticornis" pageId="15" pageNumber="110" rank="species" species="recticornis">Y. recticornis</taxonomicName>
: large eye size (distal edge of the eye surpassing distal edge of the canthus), shape of the central tooth of the MFS (base subparallel and as wide as the lateral ridge of the MFS; center horn short with apex rounded, largely free and directed anteriorly and dorsally [Fig. 5D]), and shape of the frons and clypeus concave (rather than straight as in
<taxonomicName id="E9EA615AB51E44C72DF1A45F5471B0F9" lsidName="Y. jimenezi" pageId="15" pageNumber="110" rank="species" species="jimenezi">Y. jimenezi</taxonomicName>
).
<bibRefCitation id="55F0BE56FF52763EDD905F0CB2D5D475" author="Castillo, C" journalOrPublisher="Acta Zoologica Mexicana (nueva serie)" pageId="29" pageNumber="124" pagination="1 - 84" title="Biosistematica del genero Petrejoides Kuwert (Coleoptera, Lamellicornia, Passalidae)." volume="4" year="1984">Castillo and Reyes-Castillo (1984)</bibRefCitation>
suggested that eye size variation among
<taxonomicName id="DB44ED00262B5125F99BD9CB7DA8F6B6" lsidName="Y. recticornis" pageId="15" pageNumber="110" rank="species" species="recticornis">Y. recticornis</taxonomicName>
s. l. was dependent upon the locality of the population (populations in Oaxaca possess large eyes; populations in Veracruz possess small eyes). Characters of the internal tooth of the left mandible and aedeagus are described based on the paralectotype and non-type material because the mandibles of the lectotype are closed (thus making it impossible to determine the state of this character in this specimen).
</paragraph>
<caption id="264F02E65BBEA589D0D29F8D02774659" pageId="15" pageNumber="110">
<paragraph id="3540075701983362DF0751D3E42A44EB" pageId="15" pageNumber="110">
Figure 5.
<taxonomicName id="2EE84D2396D8F0BDAA97D2C68C4CD72C" class="Insecta" family="Passalidae" genus="Yumtaax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yumtaax recticornis" order="Coleoptera" pageId="15" pageNumber="110" phylum="Arthropoda" rank="species" species="recticornis">Yumtaax recticornis</taxonomicName>
(Burmeister), lectotype: A dorsal habitus B ventral habitus C dorsal view of pronotum and head D lateral view of pronotum and head E lateral view of right mesotibia F lateral view of aedeagus G ventral view of aedeagus H dorsal view of aedeagus.
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>
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