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<document id="01E0382D85071E2333B750289241217D" ID-DOI="http://dx.doi.org/10.3897/zookeys.743.23029" ID-GBIF-Dataset="1e5fbbb7-a293-466f-b354-9eb70c8befe7" ID-PMC="PMC5904386" ID-Pensoft-Pub="1313-2970-743-67" ID-PubMed="29670437" ID-ZooBank="4D31DBBF07B1479C90489C9207B5103D" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-743-67" ModsDocOrigin="ZooKeys 743" ModsDocTitle="Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae)" checkinTime="1521107529766" checkinUser="pensoft" docAuthor="Kohlmann, Bert, Arriaga-Jimenez, Alfonsina &amp; Roes, Matthias" docDate="2018" docId="0D21E1174692236999186DBC687B22FA" docLanguage="en" docName="ZooKeys 743: 67-93" docOrigin="ZooKeys 743" docSource="http://dx.doi.org/10.3897/zookeys.743.23029" docTitle="Phanaeus (Notiophanaeus) malyi Arnaud 2002" docType="treatment" docVersion="5" lastPageNumber="76" masterDocId="FF813C19FFB8FFF8FFB7FFACFF851225" masterDocTitle="Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae)" masterLastPageNumber="93" masterPageNumber="67" pageNumber="73" updateTime="1701318031048" updateUser="plazi">
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<mods:title id="86E81BDC68C28DD534F4BBA1F9DAA1E8">Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae)</mods:title>
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<treatment id="0D21E1174692236999186DBC687B22FA" ID-GBIF-Taxon="142283990" LSID="urn:lsid:plazi:treatment:0D21E1174692236999186DBC687B22FA" httpUri="http://treatment.plazi.org/id/0D21E1174692236999186DBC687B22FA" lastPageId="9" lastPageNumber="76" pageId="6" pageNumber="73">
<subSubSection id="DB118AE91E387893FF03A53C965C7D1F" pageId="6" pageNumber="73" type="nomenclature">
<paragraph id="83AC13F4E3F050056FE68FAA68641E97" pageId="6" pageNumber="73">
<taxonomicName id="D326376030622F571FDC8205D7FF1A14" ID-CoL="4FSJS" authority="Arnaud, 2002" authorityName="Arnaud" authorityYear="2002" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus (Notiophanaeus) malyi" order="Coleoptera" pageId="6" pageNumber="73" phylum="Arthropoda" rank="species" species="malyi" subGenus="Notiophanaeus">Phanaeus (Notiophanaeus) malyi Arnaud, 2002</taxonomicName>
Fig. 8
<normalizedToken id="6FBFBFDF22D90EF73FB691E6CE4E3473" originalValue="ab">a-b</normalizedToken>
</paragraph>
</subSubSection>
<subSubSection id="2A71E6174C3C5A68C2E50A9832D9739F" pageId="7" pageNumber="74" type="reference_group">
<paragraph id="7E2273F9F2829CE97206E793927DE374" pageId="7" pageNumber="74">
<taxonomicName id="3453A8FD94D2B86674EB6E528AB45DF5" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus pyrois" order="Coleoptera" pageId="7" pageNumber="74" phylum="Arthropoda" rank="subspecies" species="pyrois" subspecies="malyi">
<pageBreakToken id="36853568FF1D4CC6E29B893E2EC016DB" pageId="7" pageNumber="74" start="start">Phanaeus</pageBreakToken>
pyrois malyi
</taxonomicName>
:
<bibRefCitation id="77E70050875B5606BD3E6226C7EB3FC2" author="Arnaud, P" journalOrPublisher="Hillside Books, Canterbury, Kent" pageId="19" pageNumber="86" title="Phanaeini. Les Coleopteres du Monde, vol. 28." year="2002">Arnaud (2002</bibRefCitation>
: 96-97).
</paragraph>
<paragraph id="786FD929247671B725415AB3139B411A" pageId="7" pageNumber="74">
<taxonomicName id="363A7E771402FA0FC0125E30A91E1F72" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus malyi" order="Coleoptera" pageId="7" pageNumber="74" phylum="Arthropoda" rank="species" species="malyi">Phanaeus malyi</taxonomicName>
Arnaud,
<normalizedToken id="301890B60997CBA1CE0CAE97BDFB5464" originalValue="Solís">Solis</normalizedToken>
&amp; Kohlmann (2012: 10).
</paragraph>
<paragraph id="0E8E03BC152BBB5949E3580763FFEAFD" pageId="7" pageNumber="74">
<taxonomicName id="8172EA896322AB150C0BFCDAA93B462C" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus pyrois" order="Coleoptera" pageId="7" pageNumber="74" phylum="Arthropoda" rank="species" species="pyrois">Phanaeus pyrois</taxonomicName>
Bates, Edmonds &amp;
<normalizedToken id="0B22A4E31A39D6FE7BB3693C6B5F16CF" originalValue="Zídek">Zidek</normalizedToken>
(2012: 1, 3, 13, 52-53, 57)
</paragraph>
</subSubSection>
<subSubSection id="B67FD03053467AF7FCD4943EF22F8A20" lastPageId="9" lastPageNumber="76" pageId="7" pageNumber="74" type="remarks">
<paragraph id="47D022AD7899F21688F4563BE299EDCF" pageId="7" pageNumber="74">Remarks.</paragraph>
<paragraph id="24892F4B474DAA6ABBB3994B883B3583" lastPageId="8" lastPageNumber="75" pageId="7" pageNumber="74">
<bibRefCitation id="F9930D21BB2A9CA03FD848C34EFB86FC" author="Solis, A" journalOrPublisher="Zootaxa" pageId="21" pageNumber="88" pagination="1 - 32" title="Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica." volume="3482" year="2012">
<normalizedToken id="73B77E914333CFBB2F6EF1C836FBB7FE" originalValue="Solís">Solis</normalizedToken>
and Kohlmann (2012)
</bibRefCitation>
elevated
<taxonomicName id="7966687EBA37AB4ECBF40A92D85E6355" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus pyrois" order="Coleoptera" pageId="7" pageNumber="74" phylum="Arthropoda" rank="subspecies" species="pyrois" subspecies="malyi">Phanaeus pyrois malyi</taxonomicName>
Arnaud to species status based on the results of mitochondrial DNA studies of Costa Rican specimens that clearly separated Caribbean from Pacific populations at a genetic distance concordant with species level (average Kimura-2-parameter [K2P] = 3.8 %,
<bibRefCitation id="98266BCFEF147ACB5AEB8CAFB0E91CE9" author="Solis, A" journalOrPublisher="Zootaxa" pageId="21" pageNumber="88" pagination="1 - 32" title="Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica." volume="3482" year="2012">
<normalizedToken id="BE115286AD9D01505B993147E3443DD2" originalValue="Solís">Solis</normalizedToken>
and Kohlmann 2012
</bibRefCitation>
). Moreover,
<taxonomicName id="3C7F9CA9098CA312B21C1524C32EC3C3" lsidName="Ph. malyi" pageId="7" pageNumber="74" rank="species" species="malyi">Ph. malyi</taxonomicName>
has short and rounded postero-lateral pronotal angles (Fig. 8
<normalizedToken id="E967991DF175422EB6BB6C98A71FD2F3" originalValue="ab">a-b</normalizedToken>
), whereas they are long and slender in
<taxonomicName id="B7A694B510DA15CCD76DF0F70EA190DE" lsidName="Ph. pyrois" pageId="7" pageNumber="74" rank="species" species="pyrois">Ph. pyrois</taxonomicName>
(Fig. 8
<normalizedToken id="FD55DF3FCD3E2B980099A5CE0A842A76" originalValue="cd">c-d</normalizedToken>
). Subsequently,
<bibRefCitation id="F8D5433EEC8F4B8CE42343AA277510ED" author="Edmonds, WD" journalOrPublisher="Insecta Mundi" pageId="20" pageNumber="87" pagination="1 - 108" title="Taxonomy of Phanaeus revisited: Revised keys to and comments on species of the New World dung beetle genus Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)." volume="0274" year="2012">
Edmonds and
<normalizedToken id="67CACBAAA6CA453F1E59B83F055A89DA" originalValue="Zídek">Zidek</normalizedToken>
(2012)
</bibRefCitation>
synonymised
<taxonomicName id="4D0F9F70A8ACB0DCAD7680BF3500DF31" lsidName="Ph. pyrois" pageId="7" pageNumber="74" rank="subspecies" species="pyrois" subspecies="malyi">Ph. pyrois malyi</taxonomicName>
with
<taxonomicName id="4A0B702CDA13D2FB7B90A6911071C016" lsidName="Ph. pyrois" pageId="7" pageNumber="74" rank="species" species="pyrois">Ph. pyrois</taxonomicName>
, pending a comparative analysis of black varieties of Panamanian and South American populations.
<taxonomicName id="96659D4214161D66B51A9B5DE6462AE4" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus malyi" order="Coleoptera" pageId="7" pageNumber="74" phylum="Arthropoda" rank="species" species="malyi">Phanaeus malyi</taxonomicName>
was compared with material from J.
<normalizedToken id="BF54D333CB41D7A4535E62A84D382D6C" originalValue="Blackallers">Blackaller's</normalizedToken>
private collection and photos sent by
<normalizedToken id="F2D7B3000EDC60C4B00B53C9D346D36F" originalValue="François">Francois</normalizedToken>
<normalizedToken id="079B39D2FD32C402F76BE28EC347A70D" originalValue="Génier">Genier</normalizedToken>
from the Canadian Museum of Nature (Suppl. material 1: Figure S1) of black specimens of
<taxonomicName id="7141B62264A7EE3E9D5D3FC62D35FF24" lsidName="Ph. pyrois" pageId="7" pageNumber="74" rank="species" species="pyrois">Ph. pyrois</taxonomicName>
from Cerro Campana,
<pageBreakToken id="EF03698950E3464511ABA15FF4567EB2" pageId="8" pageNumber="75" start="start">Province</pageBreakToken>
of Panama, Panama (formerly treated as
<taxonomicName id="4A9A440314DB5855C79471C88D128039" lsidName="Ph. pyrois" pageId="8" pageNumber="75" rank="subspecies" species="pyrois" subspecies="olsoufieffi">Ph. pyrois olsoufieffi</taxonomicName>
Balthasar) and Playa de Oro, Esmeralda, Ecuador (formerly treated as
<taxonomicName id="C63A069E7D0620A16A92056E1966904D" lsidName="Ph. pyrois" pageId="8" pageNumber="75" rank="subspecies" species="pyrois" subspecies="funereus">Ph. pyrois funereus</taxonomicName>
Balthasar).
<taxonomicName id="2DAEC31E0095EBF18AE57B044AE4426A" lsidName="Ph. malyi" pageId="8" pageNumber="75" rank="species" species="malyi">Ph. malyi</taxonomicName>
and
<taxonomicName id="E235132848FE6EC14BC4F349D1A089FF" lsidName="Ph. p." pageId="8" pageNumber="75" rank="subspecies" species="p." subspecies="olsoufieffi">Ph. p. olsoufieffi</taxonomicName>
are different taxa. While
<taxonomicName id="B792C9F35F195D9020F183DD14AAE53D" lsidName="Ph. malyi" pageId="8" pageNumber="75" rank="species" species="malyi">Ph. malyi</taxonomicName>
has short and rounded posterolateral angles of the pronotum, similar to
<taxonomicName id="D37C730B6497722F8564C1F90862AD15" lsidName="Ph. endymion" pageId="8" pageNumber="75" rank="species" species="endymion">Ph. endymion</taxonomicName>
,
<taxonomicName id="BBB2C5AF93A27B776236179848AEC57E" lsidName="Ph. p." pageId="8" pageNumber="75" rank="subspecies" species="p." subspecies="olsoufieffi">Ph. p. olsoufieffi</taxonomicName>
has the typical
<taxonomicName id="38F18FDABEF27C459FFAFD7F38E3CB20" lsidName="pyrois" pageId="8" pageNumber="75" rank="species" species="pyrois">pyrois</taxonomicName>
long and slender posterolateral angles of the pronotum.
<taxonomicName id="9B325F2FC514F399E650203783D3BC58" lsidName="Ph. p." pageId="8" pageNumber="75" rank="subspecies" species="p." subspecies="olsoufieffi">Ph. p. olsoufieffi</taxonomicName>
also exists in Costa Rica, distributed along the foot of the mountains and not in the lowlands like the typical metallic-red
<taxonomicName id="9E82D59FE27C87998BFA42824EE5586C" lsidName="pyrois" pageId="8" pageNumber="75" rank="species" species="pyrois">pyrois</taxonomicName>
. A more detailed analysis would be required in order to determine if it should again be resurrected into subspecific status. In relation to
<taxonomicName id="84DF817E95723EEA1CF9A23B79FD662E" lsidName="Ph. p." pageId="8" pageNumber="75" rank="subspecies" species="p." subspecies="funereus">Ph. p. funereus</taxonomicName>
, these specimens have an intermediate form of the postero-lateral angles of the pronotum between
<taxonomicName id="7498E377E634D7BE1B2FA99FD5AB7797" lsidName="malyi" pageId="8" pageNumber="75" rank="species" species="malyi">malyi</taxonomicName>
and
<taxonomicName id="D421295B5282639C48B26F709AFF7584" lsidName="olsoufieffi" pageId="8" pageNumber="75" rank="species" species="olsoufieffi">olsoufieffi</taxonomicName>
. Moreover
<taxonomicName id="F530D90F8C2A2C9D063BA4B88554A998" lsidName="funereus" pageId="8" pageNumber="75" rank="species" species="funereus">funereus</taxonomicName>
is distinctly dull black whereas
<taxonomicName id="3264BD4BA546644D97B956A0B8FA118E" lsidName="malyi" pageId="8" pageNumber="75" rank="species" species="malyi">malyi</taxonomicName>
is a shiny black. A more detailed analysis is required for resurrecting
<taxonomicName id="500A7AAA5F342BF4588D352902C8AE95" lsidName="funereus" pageId="8" pageNumber="75" rank="species" species="funereus">funereus</taxonomicName>
, but it is a different taxon from
<taxonomicName id="A88997D97B14ADE6522FAF9BF6E0B6E9" lsidName="malyi" pageId="8" pageNumber="75" rank="species" species="malyi">malyi</taxonomicName>
.
</paragraph>
<caption id="31BD86CC734A47D4CBCD31F89874F85B" pageId="8" pageNumber="75">
<paragraph id="C7C826AD4A96ABED564ED81B022A5FE5" pageId="8" pageNumber="75">
Figure 8. Major male habitus in a dorsal and b lateral view of
<taxonomicName id="DB592EEE1E05462052F103700A85EBEC" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus malyi" order="Coleoptera" pageId="8" pageNumber="75" phylum="Arthropoda" rank="species" species="malyi">Phanaeus malyi</taxonomicName>
, and in c dorsal and d lateral view of
<taxonomicName id="78F2F0C0A51AEC7B0F230F4DE7B7C9EE" lsidName="Ph. pyrois." pageId="8" pageNumber="75" rank="species" species="pyrois.">Ph. pyrois.</taxonomicName>
Scale bar= 5mm.
</paragraph>
</caption>
<paragraph id="157DDCBBDC67CD0DD15ECD3FFAC9ABF5" lastPageId="9" lastPageNumber="76" pageId="8" pageNumber="75">
We therefore revalidate
<taxonomicName id="46228C2D1EAC3C4541EB9829160C8FDF" lsidName="Ph. malyi" pageId="8" pageNumber="75" rank="species" species="malyi">Ph. malyi</taxonomicName>
to full species status based on the previously mentioned mitochondrial DNA analysis (
<bibRefCitation id="9EF3EA5A680BF8E5D964F4FA59A74AAC" author="Solis, A" journalOrPublisher="Zootaxa" pageId="21" pageNumber="88" pagination="1 - 32" title="Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica." volume="3482" year="2012">
<normalizedToken id="C6CC29450B398E587CBA5AB594C7040A" originalValue="Solís">Solis</normalizedToken>
and Kohlmann 2012
</bibRefCitation>
); the characteristic ecology and distribution of this species in tropical rainforest along the southern Pacific coast
<pageBreakToken id="F0E9387EF0C486EE41A597A62D140E9F" pageId="9" pageNumber="76" start="start">of</pageBreakToken>
Costa Rica, probably extending to the northern Panamanian Pacific coast (
<bibRefCitation id="6C9F982F3F27BA3E0D3761B75EB3A799" author="Solis, A" journalOrPublisher="Zootaxa" pageId="21" pageNumber="88" pagination="1 - 32" title="Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica." volume="3482" year="2012">
<normalizedToken id="FB4F38217BE0E7CDB9D606FEFF65CC39" originalValue="Solís">Solis</normalizedToken>
and Kohlmann 2012
</bibRefCitation>
); and the different shapes of the pronota (Suppl. material 1: Figure S1).
</paragraph>
<paragraph id="5B0F8EDDEF7CB6ACD656E676CEE83476" pageId="9" pageNumber="76">
<taxonomicName id="369056A422787AC6F642181705A75C6D" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus malyi" order="Coleoptera" pageId="9" pageNumber="76" phylum="Arthropoda" rank="species" species="malyi">Phanaeus malyi</taxonomicName>
follows a characteristic distribution pattern described by
<bibRefCitation id="E20A117082B8345A0A58D9FEF46430DF" author="Kohlmann, B" journalOrPublisher="Giornale Italiano di Entomologia" pageId="20" pageNumber="87" pagination="1 - 30" title="The Tarcoles line: biogeographic effects of the Talamanca Range in lower Central America." volume="12" year="2007">Kohlmann and Wilkinson (2007)</bibRefCitation>
and by
<bibRefCitation id="20A56C68AFD2C0A0FD855E52780FF1CC" author="Solis, A" journalOrPublisher="Zootaxa" pageId="21" pageNumber="88" pagination="1 - 32" title="Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica." volume="3482" year="2012">
<normalizedToken id="210C7EA9E8520B2A9F9161DFE34E47B7" originalValue="Solís">Solis</normalizedToken>
and Kohlmann (2012)
</bibRefCitation>
, where many species living in the Costa Rican southern Pacific tropical forest are sister species of taxa found along the tropical forest of the Caribbean coast. This vicariant speciation pattern (apparently mediated by the emergence of the Talamanca Cordillera that has divided a formerly continuous tropical forest stretching from the Caribbean to the Pacific) has been observed to occur in species of dung beetles, fishes, amphibians, reptiles, and birds (
<bibRefCitation id="6C1B61872070AF9608A60C617990FB4B" author="Kohlmann, B" journalOrPublisher="Giornale Italiano di Entomologia" pageId="20" pageNumber="87" pagination="1 - 30" title="The Tarcoles line: biogeographic effects of the Talamanca Range in lower Central America." volume="12" year="2007">Kohlmann and Wilkinson 2007</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
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