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<mods:title id="C3E0C003827D53FA6453A055F3378248">The integration processes of the distributional patterns in the Mexican Transition Zone: Phyletic, paleogeographic and ecological factors of a case study</mods:title>
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<subSubSection id="8DD54F94FFA9D36D97B2FA99FC9B4B48" pageId="26" pageNumber="27" type="nomenclature">
<paragraph id="C5701C1FFFA9D36D97B2FA99FD5D4835" blockId="26.[151,707,1337,1364]" box="[151,707,1337,1364]" pageId="26" pageNumber="27">
<heading id="9E38AB73FFA9D36D97B2FA99FD5D4835" bold="true" box="[151,707,1337,1364]" fontSize="11" level="1" pageId="26" pageNumber="27" reason="1">
<emphasis id="F7BBC00DFFA9D36D97B2FA99FD5D4835" bold="true" box="[151,707,1337,1364]" pageId="26" pageNumber="27">
The 
<taxonomicName id="02CF679CFFA9D36D97EAFA9AFE684832" ID-CoL="7W5GJ" authorityName="Horn" authorityYear="1881" box="[207,502,1337,1363]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="F7BBC00DFFA9D36D97EAFA9AFE684832" bold="true" box="[207,502,1337,1363]" italics="true" pageId="26" pageNumber="27">Onthophagus brevifrons</emphasis>
</taxonomicName>
species complex
</emphasis>
</heading>
</paragraph>
<paragraph id="C5701C1FFFA9D36D97B2FA23FC9B4B48" blockId="26.[151,1437,1408,1937]" pageId="26" pageNumber="27">
Within the 
<taxonomicName id="02CF679CFFA9D36D963CFA22FE4348F8" authorityName="Harold" authorityYear="1869" box="[281,477,1408,1433]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="hippopotamus">
<emphasis id="F7BBC00DFFA9D36D963CFA22FE4348F8" box="[281,477,1408,1433]" italics="true" pageId="26" pageNumber="27">O. hippopotamus</emphasis>
</taxonomicName>
species line, the 
<taxonomicName id="02CF679CFFA9D36D9582FA22FCA348F8" authorityName="Horn" authorityYear="1881" box="[679,829,1408,1433]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="F7BBC00DFFA9D36D9582FA22FCA348F8" box="[679,829,1408,1433]" italics="true" pageId="26" pageNumber="27">O. brevifrons</emphasis>
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species complex has five species whose distribution forms an arch (
<figureCitation id="5DF4009AFFA9D36D9665FA07FE1A48DC" box="[320,388,1444,1469]" captionStart="FIGURE 4" captionStartId="6.[151,250,2010,2032]" captionTargetBox="[187,1405,240,1945]" captionTargetId="figure@6.[151,1436,240,1953]" captionTargetPageId="6" captionText="FIGURE 4. Onthophagus brevifrons, male dorsal view." httpUri="https://zenodo.org/record/2642580/files/figure.png" pageId="26" pageNumber="27">Fig. 4</figureCitation>
, 
<figureCitation id="5DF4009AFFA9D36D96B4FA07FE3F48DC" box="[401,417,1444,1469]" captionStart="FIGURE 5" captionStartId="7.[151,250,2004,2026]" captionTargetBox="[171,1416,273,1897]" captionTargetId="figure@7.[151,1436,273,1901]" captionTargetPageId="7" captionText="FIGURE 5. Onthophagus moroni, male dorsal view." httpUri="https://zenodo.org/record/2642582/files/figure.png" pageId="26" pageNumber="27">5</figureCitation>
, 
<figureCitation id="5DF4009AFFA9D36D9688FA06FE5548DC" box="[429,459,1445,1469]" captionStart="FIGURE 14" captionStartId="18.[151,250,1587,1609]" captionTargetBox="[154,1434,581,1562]" captionTargetId="figure@18.[151,1436,573,1566]" captionTargetPageId="18" captionText="FIGURE 14. Distribution of the O. brevifrons species complex." httpUri="https://zenodo.org/record/2642599/files/figure.png" pageId="26" pageNumber="27">14</figureCitation>
, 
<figureCitation id="5DF4009AFFA9D36D96F2FA07FE6A48DC" box="[471,500,1444,1469]" captionStart="FIGURE 15" captionStartId="19.[151,250,1524,1546]" captionTargetBox="[154,1434,527,1508]" captionTargetId="figure@19.[151,1436,519,1512]" captionTargetPageId="19" captionText="FIGURE 15. Location of the Cuetzalan cave associated fauna." httpUri="https://zenodo.org/record/2642601/files/figure.png" pageId="26" pageNumber="27">15</figureCitation>
) to the north of the Mexican High Plateau, from the Sierra Madre Occidental to the Sierra Madre Oriental. As mentioned, it is an equivalent line to the 
<taxonomicName id="02CF679CFFA9D36D94B3FA6AFBC64880" authorityName="Harold" authorityYear="1869" box="[918,1112,1480,1505]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="hippopotamus">
<emphasis id="F7BBC00DFFA9D36D94B3FA6AFBC64880" box="[918,1112,1480,1505]" italics="true" pageId="26" pageNumber="27">O. hippopotamus</emphasis>
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species complex, but it may have diversified in more recent geological times (see “The 
<taxonomicName id="02CF679CFFA9D36D9466FA4FFB1C4B64" authorityName="Harold" authorityYear="1869" box="[835,1154,1516,1541]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="hippopotamus">
<emphasis id="F7BBC00DFFA9D36D9466FA4FFB1C4B64" box="[835,1154,1516,1541]" italics="true" pageId="26" pageNumber="27">Onthophagus hippopotamus</emphasis>
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species line and species complex” and “Comparative analysis of the genitalia”).
</paragraph>
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<subSubSection id="8DD54F94FFA9D36D97E2F997FB114AF0" pageId="26" pageNumber="27" type="description">
<paragraph id="C5701C1FFFA9D36D97E2F997FB114AF0" blockId="26.[151,1437,1408,1937]" pageId="26" pageNumber="27">
There is a series of morphological and ecological differences that support the idea of conserving both species complexes. The supposedly plesiomorphic characters present in the 
<taxonomicName id="02CF679CFFA9D36D94ABF9FAFBBE4B10" authorityName="Horn" authorityYear="1881" box="[910,1056,1624,1649]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="F7BBC00DFFA9D36D94ABF9FAFBBE4B10" box="[910,1056,1624,1649]" italics="true" pageId="26" pageNumber="27">O. brevifrons</emphasis>
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species complex include the midkeel of the lamella copulatrix lacking a sagittal suture and the orientation of the axis of the apexes of the parameres. Supposedly synapomorphic characters are the introflexion of the left lateral wall of the copulatory lamella, as well as the development of the caudal branches of the sclerotization of the vaginal support that is much larger than the cephalic branches. In this species complex, 
<taxonomicName id="02CF679CFFA9D36D95B6F94AFCF24A61" authority="Horn" authorityName="Horn" authorityYear="1881" box="[659,876,1768,1793]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="F7BBC00DFFA9D36D95B6F94AFCB74A60" box="[659,809,1768,1793]" italics="true" pageId="26" pageNumber="27">O. brevifrons</emphasis>
Horn
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and 
<taxonomicName id="02CF679CFFA9D36D948CF94AFB3D4A60" authority="Howden" authorityName="Howden" authorityYear="1973" box="[937,1187,1768,1793]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="cuevensis">
<emphasis id="F7BBC00DFFA9D36D948CF94AFBA74A61" box="[937,1081,1769,1792]" italics="true" pageId="26" pageNumber="27">O. cuevensis</emphasis>
Howden
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show the supposedly plesiomorphic character states. The supposedly most apomorphic character states are present in 
<taxonomicName id="02CF679CFFA9D36D93C2F8AEFE074A29" authority="Howden &amp; Cartwright." authorityName="Howden &amp; Cartwright." class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="cavernicollis">
<emphasis id="F7BBC00DFFA9D36D93C2F8AEFA024A44" box="[1255,1436,1804,1829]" italics="true" pageId="26" pageNumber="27">O. cavernicollis</emphasis>
Howden &amp; Cartwright.
</taxonomicName>
<taxonomicName id="02CF679CFFA9D36D96BAF893FD134A29" authorityName="Zunino &amp; Halffter" authorityYear="1988" box="[415,653,1840,1865]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="moroni">
<emphasis id="F7BBC00DFFA9D36D96BAF893FD134A29" box="[415,653,1840,1865]" italics="true" pageId="26" pageNumber="27">Onthophagus moroni</emphasis>
</taxonomicName>
(
<figureCitation id="5DF4009AFFA9D36D95B9F893FD434A28" box="[668,733,1840,1865]" captionStart="FIGURE 5" captionStartId="7.[151,250,2004,2026]" captionTargetBox="[171,1416,273,1897]" captionTargetId="figure@7.[151,1436,273,1901]" captionTargetPageId="7" captionText="FIGURE 5. Onthophagus moroni, male dorsal view." httpUri="https://zenodo.org/record/2642582/files/figure.png" pageId="26" pageNumber="27">Fig. 5</figureCitation>
) is supposed to be a closely related species to 
<taxonomicName id="02CF679CFFA9D36D93C7F892FA0B4A28" box="[1250,1429,1840,1865]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="cavernicollis">
<emphasis id="F7BBC00DFFA9D36D93C7F892FA0B4A28" box="[1250,1429,1840,1865]" italics="true" pageId="26" pageNumber="27">O. cavernicollis</emphasis>
</taxonomicName>
. The relationship of 
<taxonomicName id="02CF679CFFA9D36D965FF8F6FCB74A0C" authority="Howden &amp; Cartwright" authorityName="Howden &amp; Cartwright" authorityYear="1963" box="[378,809,1876,1901]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="subtropicus">
<emphasis id="F7BBC00DFFA9D36D965FF8F6FDBE4A0C" box="[378,544,1876,1901]" italics="true" pageId="26" pageNumber="27">O. subtropicus</emphasis>
Howden &amp; Cartwright
</taxonomicName>
with the other species in the 
<taxonomicName id="02CF679CFFA9D36D93A4F8F6FA884A0C" authorityName="Horn" authorityYear="1881" box="[1153,1302,1876,1901]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="species" species="brevifrons">
<emphasis id="F7BBC00DFFA9D36D93A4F8F6FA884A0C" box="[1153,1302,1876,1901]" italics="true" pageId="26" pageNumber="27">O. brevifrons</emphasis>
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complex is unclear, but this is the single species that occurs near sea level in tropical lowland habitats.
</paragraph>
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</treatment>
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