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<mods:title id="20A340E9F380C3FFE75B1871DD4752C4">Comparative morphology of the genera with perpendicular parameres in Anomalini (Coleoptera, Scarabaeidae, Rutelinae) allows revalidation of the endemic Mexican genus Lamoana Casey, 1915 stat. rev.</mods:title>
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<mods:namePart id="8F490FA5D3F8E1600C63B46AC55E1AE6">Madrigal, Kevin</mods:namePart>
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<mods:affiliation id="08D65280D667B21901BE17B5CC35CB8C">Licenciatura en Biología, Universidad Autónoma de Tlaxcala, Km 10.5 San Martín Texmelucan- Tlaxcala s / n, Ixtacuixtla de Mariano Matamoros, Tlaxcala 90120, Mexico.</mods:affiliation>
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<mods:namePart id="07D4D51F235BFD3B549F22FBA5319C28">Zaragoza-Caballero, Santiago</mods:namePart>
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<mods:affiliation id="D0868412BB383B16CBB793CE54B0AC4B">Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70 - 153, 04510, Mexico City, Mexico.</mods:affiliation>
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<paragraph id="8BA6B26EFFD3FFB05FB98A0CD39AAD28" blockId="1.[189,958,1537,1564]" box="[189,958,1537,1564]" pageId="1" pageNumber="176">
<heading id="D0EE0502FFD3FFB05FB98A0CD39AAD28" bold="true" box="[189,958,1537,1564]" fontSize="11" level="3" pageId="1" pageNumber="176" reason="3">
<emphasis id="B96D6E7CFFD3FFB05FB98A0CD39AAD28" bold="true" box="[189,958,1537,1564]" pageId="1" pageNumber="176">
Taxonomic history of
<taxonomicName id="4C19C9EDFFD3FFB05EC98A0FD39AAD28" authority="(Blanchard, 1851)" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[461,958,1537,1564]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD3FFB05EC98A0FD2F2AD2F" bold="true" box="[461,726,1537,1564]" italics="true" pageId="1" pageNumber="176">Strigoderma villosella</emphasis>
(
<bibRefCitation id="EF88CF9FFFD3FFB05DE18A0FD391AD28" author="Blanchard E." box="[741,949,1537,1563]" pageId="1" pageNumber="176" refId="ref10253" refString="Blanchard E. 1851. Catalogue de la Collection Entomologique du Museum d'Histoire naturelle de Paris. Classe des Insectes. Ordre des Coleopteres. Tome 2. Gide et Baudry, Paris." type="book" year="1851">Blanchard, 1851</bibRefCitation>
)
</taxonomicName>
</emphasis>
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<paragraph id="8BA6B26EFFD3FFB05FB98A3ED1C2AD87" blockId="1.[189,1399,1584,1717]" pageId="1" pageNumber="176">
Historical taxonomic records are fundamental and pivotal tools to any taxonomic work, as they form a link between past ideas and modern changes (
<bibRefCitation id="EF88CF9FFFD3FFB05DF68A5AD3F8AD5D" author="Gravina M. F. &amp; Bonifazi A. &amp; Del Pasqua M. &amp; Giampaoletti J. &amp; Lezzi M. &amp; Ventura D. &amp; Giangrande A." box="[754,988,1620,1646]" pageId="1" pageNumber="176" pagination="480" refId="ref10582" refString="Gravina M. F., Bonifazi A., Del Pasqua M., Giampaoletti J., Lezzi M., Ventura D. &amp; Giangrande A. 2020. Perception of changes in marine benthic habitats: the relevance of taxonomic and ecology memory. Diversity 12 (12): 480. https: // doi. org / 10.3390 / d 12120480" type="journal article" year="2020">
Gravina
<emphasis id="B96D6E7CFFD3FFB05C5E8A5AD3B1AD5D" box="[858,917,1620,1646]" italics="true" pageId="1" pageNumber="176">et al.</emphasis>
2020
</bibRefCitation>
). For that reason, it is important to take a brief look at the historic background of
<taxonomicName id="4C19C9EDFFD3FFB05DEF8A79D352ADA2" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[747,886,1655,1681]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD3FFB05DEF8A79D352ADA2" box="[747,886,1655,1681]" italics="true" pageId="1" pageNumber="176">S. villosella</emphasis>
</taxonomicName>
so we can understand its present condition and justify our decisions.
</paragraph>
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<bibRefCitation id="EF88CF9FFFD3FFB05FB98AD5D1AFADC5" author="Blanchard E." box="[189,395,1755,1782]" pageId="1" pageNumber="176" refId="ref10253" refString="Blanchard E. 1851. Catalogue de la Collection Entomologique du Museum d'Histoire naturelle de Paris. Classe des Insectes. Ordre des Coleopteres. Tome 2. Gide et Baudry, Paris." type="book" year="1851">Blanchard (1851)</bibRefCitation>
originally described
<taxonomicName id="4C19C9EDFFD3FFB05D858AD2D32EADC6" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[641,778,1755,1782]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD3FFB05D858AD2D32EADC6" box="[641,778,1755,1782]" italics="true" pageId="1" pageNumber="176">S. villosella</emphasis>
</taxonomicName>
in the genus
<emphasis id="B96D6E7CFFD3FFB05CA08AD5D461ADC6" box="[932,1093,1755,1781]" italics="true" pageId="1" pageNumber="176">
<taxonomicName id="4C19C9EDFFD3FFB05CA08AD5D465ADC6" authorityName="Stephens" authorityYear="1830" box="[932,1089,1755,1781]" class="Insecta" family="Scarabaeidae" genus="Phyllopertha" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="genus">Phyllopertha</taxonomicName>
.
</emphasis>
The information given by this author is somewhat simple and lacks certain important information such as the precise
<typeStatus id="54A20CCCFFD3FFB05BE08AF1D533AC2A" box="[1252,1303,1791,1817]" pageId="1" pageNumber="176">type</typeStatus>
locality and male genitalia.
<bibRefCitation id="EF88CF9FFFD3FFB05EA58B2CD21CAC0F" author="Bates H. W." box="[417,568,1826,1852]" pageId="1" pageNumber="176" pagination="216 - 413" refId="ref10141" refString="Bates H. W. 1888. Pectinicornia and Lamellicornia, family Rutelidae. In: Godman F. D. &amp; Salvin O. (eds) Biologia Centrali-Americana. Insecta Coleoptera, Vol. II, Part 2: 216 - 413. Taylor and Francis, London. https: // doi. org / 10.5962 / bhl. title. 730" type="book chapter" year="1888">Bates (1888)</bibRefCitation>
made the first change to the genus
<taxonomicName id="4C19C9EDFFD3FFB05CD68B2CD418AC0F" authorityName="Samouelle" authorityYear="1819" box="[978,1084,1826,1852]" class="Insecta" family="Scarabaeidae" genus="Anomala" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD3FFB05CD68B2CD418AC0F" box="[978,1084,1826,1852]" italics="true" pageId="1" pageNumber="176">Anomala</emphasis>
</taxonomicName>
, arguing that the legs were more robust than those of other species of
<taxonomicName id="4C19C9EDFFD3FFB05DAE8B4BD367AC6C" authorityName="Stephens" authorityYear="1830" box="[682,835,1861,1887]" class="Insecta" family="Scarabaeidae" genus="Phyllopertha" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD3FFB05DAE8B4BD367AC6C" box="[682,835,1861,1887]" italics="true" pageId="1" pageNumber="176">Phyllopertha</emphasis>
</taxonomicName>
. In that study, Bates also described
<taxonomicName id="4C19C9EDFFD3FFB05BE78B4BD132ACB1" authorityName="Bates" authorityYear="1888" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="species" species="presidii">
<emphasis id="B96D6E7CFFD3FFB05BE78B4BD132ACB1" italics="true" pageId="1" pageNumber="176">Strigoderma presidii</emphasis>
</taxonomicName>
but saw that its mesepimera did not rise near the elytral humeri, so he proposed that it might belong to
<emphasis id="B96D6E7CFFD3FFB05E2B8B85D185AC96" box="[303,417,1931,1957]" italics="true" pageId="1" pageNumber="176">
<taxonomicName id="4C19C9EDFFD3FFB05E2B8B85D1B9AC96" authorityName="Samouelle" authorityYear="1819" box="[303,413,1931,1957]" class="Insecta" family="Scarabaeidae" genus="Anomala" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="genus">Anomala</taxonomicName>
.
</emphasis>
<bibRefCitation id="EF88CF9FFFD3FFB05EAC8B85D243AC95" author="Nonfried A. F." box="[424,615,1931,1958]" pageId="1" pageNumber="176" pagination="279 - 297" refId="ref11757" refString="Nonfried A. F. 1893. Monographische Beitrage zur Kaferfauna von Central-Amerika. I. Rutelidae: Epectinaspis und Strigoderma. Berliner entomologische Zeitschrift 38: 279 - 297. https: // doi. org / 10.1002 / mmnd. 18940380304" type="journal article" year="1893">Nonfried (1893)</bibRefCitation>
made the revision of the genera
<taxonomicName id="4C19C9EDFFD3FFB05CDB8B82D451AC96" authorityName="Blanchard" authorityYear="1851" box="[991,1141,1932,1957]" class="Insecta" family="Scarabaeidae" genus="Epectinaspis" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD3FFB05CDB8B82D451AC96" box="[991,1141,1932,1957]" italics="true" pageId="1" pageNumber="176">Epectinaspis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C19C9EDFFD3FFB05BAF8B85D51AAC96" authorityName="Burmeister" authorityYear="1844" box="[1195,1342,1931,1957]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD3FFB05BAF8B85D51AAC96" box="[1195,1342,1931,1957]" italics="true" pageId="1" pageNumber="176">Strigoderma</emphasis>
</taxonomicName>
, and described
<taxonomicName id="4C19C9EDFFD3FFB05E308BA1D180ACFA" authorityName="Nonfried" authorityYear="1983" box="[308,420,1967,1993]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="species" species="hirsuta">
<emphasis id="B96D6E7CFFD3FFB05E308BA1D180ACFA" box="[308,420,1967,1993]" italics="true" pageId="1" pageNumber="176">S. hirsuta</emphasis>
</taxonomicName>
.
<bibRefCitation id="EF88CF9FFFD3FFB05EB68BA1D274ACFA" author="Casey T. L." box="[434,592,1966,1993]" pageId="1" pageNumber="176" pagination="1460" refId="ref10287" refString="Casey T. L. 1915. A review of the American species of Rutelinae, Dynastinae and Cetoniinae. Memoirs on the Coleoptera 6: 1460. https: // doi. org / 10.5962 / bhl. title. 15993" type="journal article" year="1915">Casey (1915)</bibRefCitation>
established
<taxonomicName id="4C19C9EDFFD3FFB05DDB8BA1D36AACFB" authorityName="Casey" authorityYear="1915" box="[735,846,1967,1992]" class="Insecta" family="Scarabaeidae" genus="Lamoana" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD3FFB05DDB8BA1D36AACFB" box="[735,846,1967,1992]" italics="true" pageId="1" pageNumber="176">Lamoana</emphasis>
</taxonomicName>
, a monospecific genus created for
<emphasis id="B96D6E7CFFD3FFB05BE78BA1D553ACFA" box="[1251,1399,1967,1993]" italics="true" pageId="1" pageNumber="176">
<taxonomicName id="4C19C9EDFFD3FFB05BE78BA1D557ACFA" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[1251,1395,1967,1993]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="176" phylum="Arthropoda" rank="species" species="villosella">A. villosella</taxonomicName>
.
</emphasis>
Caseys observations were quite complete, considering the standards of that time, but he did not make a morphological comparison of the male genitalia. It must be said that the specimens that Casey evaluated came from a different locality than those that Bates and Blanchard studied, so it is uncertain whether they belonged to the same species or not.
<bibRefCitation id="EF88CF9FFFD0FFB35DB68D7CD370AABE" author="Ohaus F." box="[690,852,370,397]" pageId="2" pageNumber="177" pagination="1 - 241" refId="ref11894" refString="Ohaus F. 1918. Scarabaeidae: Euchirinae, Phaenomerinae, Rutelinae. In: Junk W. &amp; Schenkling S. (eds) Coleopterorum Catalogus 66: 1 - 241. Available from https: // www. biodiversitylibrary. org / page / 10439622 [accessed 23 Oct. 2023]." type="journal article" year="1918">Ohaus (1918)</bibRefCitation>
synonymized
<taxonomicName id="4C19C9EDFFD0FFB35B058D7DD456AABF" authorityName="Nonfried" authorityYear="1983" box="[1025,1138,370,397]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="hirsuta">
<emphasis id="B96D6E7CFFD0FFB35B058D7DD456AABF" box="[1025,1138,370,397]" italics="true" pageId="2" pageNumber="177">S. hirsuta</emphasis>
</taxonomicName>
with
<taxonomicName id="4C19C9EDFFD0FFB35BB38D7DD561AABF" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[1207,1349,370,397]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD0FFB35BB38D7DD561AABF" box="[1207,1349,370,397]" italics="true" pageId="2" pageNumber="177">A. villosella</emphasis>
</taxonomicName>
and retained it in
<taxonomicName id="4C19C9EDFFD0FFB35E528D98D1E4AA83" authorityName="Samouelle" authorityYear="1819" box="[342,448,406,432]" class="Insecta" family="Scarabaeidae" genus="Anomala" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35E528D98D1E4AA83" box="[342,448,406,432]" italics="true" pageId="2" pageNumber="177">Anomala</emphasis>
</taxonomicName>
. This nomenclatural change was made without considering all the morphological criteria that Casey established for this taxon three years earlier. We must say that there is no explanation why Ohaus ignored Caseys decision.
</paragraph>
<paragraph id="8BA6B26EFFD0FFB35FB98E10D320AF7B" blockId="2.[188,1400,542,1096]" pageId="2" pageNumber="177">
In 1944, Blackwelder published a checklist of beetles of
<collectingCountry id="F30EF2FEFFD0FFB35C538E10D390A90B" box="[855,948,542,568]" name="Mexico" pageId="2" pageNumber="177">Mexico</collectingCountry>
, Central America, South America, and the West Indies. In this study, he retained the synonymy between
<emphasis id="B96D6E7CFFD0FFB35CB08E4FD499A968" box="[948,1213,577,603]" italics="true" pageId="2" pageNumber="177">
<taxonomicName id="4C19C9EDFFD0FFB35CB08E4FD41BA968" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[948,1087,577,603]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="villosella">A. villosella</taxonomicName>
<taxonomicName id="4C19C9EDFFD0FFB35B548E4FD499A968" authorityName="Nonfried" authorityYear="1983" box="[1104,1213,577,603]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="hirsuta">S. hirsuta</taxonomicName>
</emphasis>
and recognized
<taxonomicName id="4C19C9EDFFD0FFB35FB88E6BD10FA94D" authorityName="Casey" authorityYear="1915" box="[188,299,613,638]" class="Insecta" family="Scarabaeidae" genus="Lamoana" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35FB88E6BD10FA94D" box="[188,299,613,638]" italics="true" pageId="2" pageNumber="177">Lamoana</emphasis>
</taxonomicName>
as a subgenus. Almost 50 years later,
<bibRefCitation id="EF88CF9FFFD0FFB35DF08E6AD3B4A94D" author="Bader A." box="[756,912,612,638]" pageId="2" pageNumber="177" pagination="269 - 335" refId="ref10106" refString="Bader A. 1992. A review of the North and Central American Strigoderma (Coleoptera: Scarabaeidae). Transactions of the American Entomological Society 118 (2): 269 - 335." type="journal article" year="1992">Bader (1992)</bibRefCitation>
redescribed
<taxonomicName id="4C19C9EDFFD0FFB35B2E8E6AD486A94D" authorityName="Bates" authorityYear="1888" box="[1066,1186,612,638]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="presidii">
<emphasis id="B96D6E7CFFD0FFB35B2E8E6AD486A94D" box="[1066,1186,612,638]" italics="true" pageId="2" pageNumber="177">S. presidii</emphasis>
</taxonomicName>
and made several notes about its position. He remarked on the similarity of
<taxonomicName id="4C19C9EDFFD0FFB35C788E86D42FA992" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[892,1035,647,674]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD0FFB35C788E86D42FA992" box="[892,1035,647,674]" italics="true" pageId="2" pageNumber="177">A. villosella</emphasis>
</taxonomicName>
with
<taxonomicName id="4C19C9EDFFD0FFB35B568E86D553A992" authority="Bates, 1888" authorityName="Bates" authorityYear="1888" box="[1106,1399,647,674]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="teapensis">
<emphasis id="B96D6E7CFFD0FFB35B568E86D4FBA992" box="[1106,1247,648,674]" italics="true" pageId="2" pageNumber="177">S. teapensis</emphasis>
<bibRefCitation id="EF88CF9FFFD0FFB35BED8E86D553A992" author="Bates H. W." box="[1257,1399,647,674]" pageId="2" pageNumber="177" pagination="216 - 413" refId="ref10141" refString="Bates H. W. 1888. Pectinicornia and Lamellicornia, family Rutelidae. In: Godman F. D. &amp; Salvin O. (eds) Biologia Centrali-Americana. Insecta Coleoptera, Vol. II, Part 2: 216 - 413. Taylor and Francis, London. https: // doi. org / 10.5962 / bhl. title. 730" type="book chapter" year="1888">Bates, 1888</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C19C9EDFFD0FFB35FF68EA5D149A9F7" authorityName="Bates" authorityYear="1888" box="[242,365,682,709]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="presidii">
<emphasis id="B96D6E7CFFD0FFB35FF68EA5D149A9F7" box="[242,365,682,709]" italics="true" pageId="2" pageNumber="177">S. presidii</emphasis>
</taxonomicName>
in both general habitus and the shape of the male genitalia and noted that the latter presented an intermediate morphological condition between
<taxonomicName id="4C19C9EDFFD0FFB35C9C8EC0D40FA9DB" authorityName="Burmeister" authorityYear="1844" box="[920,1067,718,744]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35C9C8EC0D40FA9DB" box="[920,1067,718,744]" italics="true" pageId="2" pageNumber="177">Strigoderma</emphasis>
</taxonomicName>
and
<taxonomicName id="4C19C9EDFFD0FFB35B6F8EC0D4F1A9DB" authorityName="Samouelle" authorityYear="1819" box="[1131,1237,718,744]" class="Insecta" family="Scarabaeidae" genus="Anomala" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35B6F8EC0D4F1A9DB" box="[1131,1237,718,744]" italics="true" pageId="2" pageNumber="177">Anomala</emphasis>
</taxonomicName>
, arguing that it must be placed in a new genus or be changed to
<emphasis id="B96D6E7CFFD0FFB35C198EFFD3ABA838" box="[797,911,753,779]" italics="true" pageId="2" pageNumber="177">
<taxonomicName id="4C19C9EDFFD0FFB35C198EFFD3AFA838" authorityName="Samouelle" authorityYear="1819" box="[797,907,753,779]" class="Insecta" family="Scarabaeidae" genus="Anomala" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">Anomala</taxonomicName>
.
</emphasis>
<bibRefCitation id="EF88CF9FFFD0FFB35C918EFFD53DA838" author="Ramirez-Ponce A. &amp; Moron M. A." box="[917,1305,753,779]" pageId="2" pageNumber="177" pagination="357 - 394" refId="ref12303" refString="Ramirez-Ponce A. &amp; Moron M. A. 2009. Relaciones filogeneticas del genero Anomala (Coleoptera: Melolonthidae: Rutelinae). Revista mexicana de Biodiversidad 80: 357 - 394. https: // doi. org / 10.22201 / ib. 20078706 e. 2009.002.610" type="journal article" year="2009">Ramírez-Ponce &amp; Morón (2009)</bibRefCitation>
made a phylogenetic study to evaluate the relationships between the genera of
<taxonomicName id="4C19C9EDFFD0FFB35CFF8F1AD458A81D" authorityName="Peringuey" authorityYear="1902" box="[1019,1148,788,814]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="196" phylum="Arthropoda" rank="tribe" tribe="Anomalini">Anomalini</taxonomicName>
. They concluded that the principal difficulty in delimiting genera with precision within the tribe was that some of genera had been established on questionable criteria. They revalidated
<taxonomicName id="4C19C9EDFFD0FFB35C878F54D438A847" authorityName="Blanchard" authorityYear="1851" box="[899,1052,858,884]" class="Insecta" family="Scarabaeidae" genus="Pachystethus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35C878F54D438A847" box="[899,1052,858,884]" italics="true" pageId="2" pageNumber="177">Pachystethus</emphasis>
</taxonomicName>
; demonstrated the paraphyly of
<taxonomicName id="4C19C9EDFFD0FFB35FDF8F70D143A8AB" authorityName="Blanchard" authorityYear="1851" box="[219,359,894,920]" class="Insecta" family="Scarabaeidae" genus="Callistethus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35FDF8F70D143A8AB" box="[219,359,894,920]" italics="true" pageId="2" pageNumber="177">Callistethus</emphasis>
</taxonomicName>
; validated the synonymy between
<taxonomicName id="4C19C9EDFFD0FFB35DFC8F70D439A8AB" authority="Casey, 1915" authorityName="Casey" authorityYear="1915" box="[760,1053,893,920]" class="Insecta" family="Scarabaeidae" genus="Anomalacra" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35DFC8F70D3ADA8AB" box="[760,905,894,920]" italics="true" pageId="2" pageNumber="177">Anomalacra</emphasis>
<bibRefCitation id="EF88CF9FFFD0FFB35C8B8F70D439A8AB" author="Casey T. L." box="[911,1053,893,920]" pageId="2" pageNumber="177" pagination="1460" refId="ref10287" refString="Casey T. L. 1915. A review of the American species of Rutelinae, Dynastinae and Cetoniinae. Memoirs on the Coleoptera 6: 1460. https: // doi. org / 10.5962 / bhl. title. 15993" type="journal article" year="1915">Casey, 1915</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C19C9EDFFD0FFB35B568F70D4C1A8AB" authorityName="Casey" authorityYear="1915" box="[1106,1253,894,920]" class="Insecta" family="Scarabaeidae" genus="Paranomala" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35B568F70D4C1A8AB" box="[1106,1253,894,920]" italics="true" pageId="2" pageNumber="177">Paranomala</emphasis>
</taxonomicName>
; revalidated
<taxonomicName id="4C19C9EDFFD0FFB35FB88FAFD16BA888" authorityName="Casey" authorityYear="1915" box="[188,335,929,955]" class="Insecta" family="Scarabaeidae" genus="Paranomala" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35FB88FAFD16BA888" box="[188,335,929,955]" italics="true" pageId="2" pageNumber="177">Paranomala</emphasis>
</taxonomicName>
, and thus
<taxonomicName id="4C19C9EDFFD0FFB35ED58FAFD244A888" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[465,608,929,955]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD0FFB35ED58FAFD244A888" box="[465,608,929,955]" italics="true" pageId="2" pageNumber="177">A. villosella</emphasis>
</taxonomicName>
was transferred to
<emphasis id="B96D6E7CFFD0FFB35C4D8FAFD3C7A888" box="[841,995,929,955]" italics="true" pageId="2" pageNumber="177">
<taxonomicName id="4C19C9EDFFD0FFB35C4D8FAFD3C4A888" authorityName="Casey" authorityYear="1915" box="[841,992,929,955]" class="Insecta" family="Scarabaeidae" genus="Paranomala" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">Paranomala</taxonomicName>
.
</emphasis>
Some years later, Ramírez-Ponce (2012), studied the supraspecific systematics of
<taxonomicName id="4C19C9EDFFD0FFB35DF98FCAD3B4A8ED" authorityName="Casey" authorityYear="1915" box="[765,912,964,990]" class="Insecta" family="Scarabaeidae" genus="Paranomala" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35DF98FCAD3B4A8ED" box="[765,912,964,990]" italics="true" pageId="2" pageNumber="177">Paranomala</emphasis>
</taxonomicName>
, and found that
<taxonomicName id="4C19C9EDFFD0FFB35B5E8FCBD4C2A8ED" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[1114,1254,964,990]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD0FFB35B5E8FCBD4C2A8ED" box="[1114,1254,964,990]" italics="true" pageId="2" pageNumber="177">P. villosella</emphasis>
</taxonomicName>
was near to the
<emphasis id="B96D6E7CFFD0FFB35FE98FE9D23BAF32" box="[237,543,999,1025]" italics="true" pageId="2" pageNumber="177">Strigoderma-Epectinaspis</emphasis>
clade, so he decided to place it in
<taxonomicName id="4C19C9EDFFD0FFB35CDA8FE9D455AF32" authorityName="Burmeister" authorityYear="1844" box="[990,1137,999,1025]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35CDA8FE9D455AF32" box="[990,1137,999,1025]" italics="true" pageId="2" pageNumber="177">Strigoderma</emphasis>
</taxonomicName>
. Notwithstanding his decision, he remarked on the similarity in terms of male genitalia to some species of
<taxonomicName id="4C19C9EDFFD0FFB35BAB8805D561AF17" authorityName="Blanchard" authorityYear="1851" box="[1199,1349,1035,1060]" class="Insecta" family="Scarabaeidae" genus="Epectinaspis" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35BAB8805D561AF17" box="[1199,1349,1035,1060]" italics="true" pageId="2" pageNumber="177">Epectinaspis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C19C9EDFFD0FFB35FBA8820D171AF7B" authorityName="Paucar-Cabrera" authorityYear="2003" box="[190,341,1070,1096]" class="Insecta" family="Scarabaeidae" genus="Balanogonia" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35FBA8820D171AF7B" box="[190,341,1070,1096]" italics="true" pageId="2" pageNumber="177">Balanogonia</emphasis>
</taxonomicName>
, just as Paucar-Cabrera had in 2003.
</paragraph>
<paragraph id="8BA6B26EFFD0FFB35FB9887ED219AEB3" blockId="2.[188,1399,1135,1408]" pageId="2" pageNumber="177">
In recent studies, the internal and external morphology have been carefully compared among the genera of
<taxonomicName id="4C19C9EDFFD0FFB35FDE889CD17FAF9F" authorityName="Peringuey" authorityYear="1902" box="[218,347,1170,1196]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="196" phylum="Arthropoda" rank="tribe" tribe="Anomalini">Anomalini</taxonomicName>
, which has allowed us to recognize patterns of variation and homologies at different levels, thus proposing taxonomic arrangements within the tribe, finding that the parameres in a perpendicular position with respect to the tectum presented in
<taxonomicName id="4C19C9EDFFD0FFB35C1988D7D40CAFC0" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[797,1064,1241,1267]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD0FFB35C1988D7D40CAFC0" box="[797,1064,1241,1267]" italics="true" pageId="2" pageNumber="177">Strigoderma villosella</emphasis>
</taxonomicName>
, is shared with the genera
<taxonomicName id="4C19C9EDFFD0FFB35FB988F2D171AE25" authorityName="Paucar-Cabrera" authorityYear="2003" box="[189,341,1276,1302]" class="Insecta" family="Scarabaeidae" genus="Balanogonia" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35FB988F2D171AE25" box="[189,341,1276,1302]" italics="true" pageId="2" pageNumber="177">Balanogonia</emphasis>
</taxonomicName>
,
<taxonomicName id="4C19C9EDFFD0FFB35E6C88F2D1C9AE25" authorityName="Blanchard" authorityYear="1851" box="[360,493,1276,1302]" class="Insecta" family="Scarabaeidae" genus="Callirhinus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35E6C88F2D1C9AE25" box="[360,493,1276,1302]" italics="true" pageId="2" pageNumber="177">Callirhinus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C19C9EDFFD0FFB35D0488F2D2BFAE25" authorityName="Bates" authorityYear="1888" box="[512,667,1276,1302]" class="Insecta" family="Scarabaeidae" genus="Dilophochila" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35D0488F2D2BFAE25" box="[512,667,1276,1302]" italics="true" pageId="2" pageNumber="177">Dilophochila</emphasis>
</taxonomicName>
,
<taxonomicName id="4C19C9EDFFD0FFB35DAA88F2D315AE25" authorityName="Ramirez-Ponce" authorityYear="2015" box="[686,817,1276,1302]" class="Insecta" family="Scarabaeidae" genus="Moroniella" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35DAA88F2D315AE25" box="[686,817,1276,1302]" italics="true" pageId="2" pageNumber="177">Moroniella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C19C9EDFFD0FFB35C7088F2D40AAE25" authorityName="Moron &amp; Nogueira" authorityYear="1998" box="[884,1070,1276,1302]" class="Insecta" family="Scarabaeidae" genus="Mazahuapertha" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35C7088F2D40AAE25" box="[884,1070,1276,1302]" italics="true" pageId="2" pageNumber="177">Mazahuapertha</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF88CF9FFFD0FFB35B4588F2D0DFAE0A" author="Ramirez-Ponce A. &amp; Moron M. A." pageId="2" pageNumber="177" pagination="1 - 24" refId="ref12352" refString="Ramirez-Ponce A. &amp; Moron M. A. 2012. Revision of the genus Pachystethus Blanchard with description of three new species from Mexico (Coleoptera: Scarabaeidae: Rutelinae: Anomalini). Zootaxa 3394 (1): 1 - 24. https: // doi. org / 10.11646 / zootaxa. 3394.1.1" type="journal article" year="2012">Morón &amp; Ramírez-Ponce 2012</bibRefCitation>
; Ramírez-Ponce 2012, 2015). An overall comparative study of male genitalia within these genera has led us to recognize clear differences in
<taxonomicName id="4C19C9EDFFD0FFB35DD1894CD3FAAE6F" baseAuthorityName="Blanchard" baseAuthorityYear="1851" box="[725,990,1346,1372]" class="Insecta" family="Scarabaeidae" genus="Strigoderma" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="species" species="villosella">
<emphasis id="B96D6E7CFFD0FFB35DD1894CD3FAAE6F" box="[725,990,1346,1372]" italics="true" pageId="2" pageNumber="177">Strigoderma villosella</emphasis>
</taxonomicName>
thus reinforcing the evidence for revalidating the genus
<taxonomicName id="4C19C9EDFFD0FFB35EC38968D212AE4C" authorityName="Casey" authorityYear="1915" box="[455,566,1382,1407]" class="Insecta" family="Scarabaeidae" genus="Lamoana" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="177" phylum="Arthropoda" rank="genus">
<emphasis id="B96D6E7CFFD0FFB35EC38968D212AE4C" box="[455,566,1382,1407]" italics="true" pageId="2" pageNumber="177">Lamoana</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>