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<mods:title id="2814EA93D64494E4BD7D2CA5219036E3">Taxonomic and nomenclatorial revision of the Neotropical genus Phaeoxantha Chaudoir (Coleoptera: Cicindelidae)</mods:title>
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<mods:affiliation id="BA27C47C65C02F3B5047709354AD15BD">Mendel University in Brno, Faculty of Forestry and Wood Technology, Department of Forest Ecology, Zemědělská 3, CZ- 613 00 Brno, Czech Republic. jirmor @ quick. cz; https: // orcid. org / 0000 - 0001 - 5294 - 6410 &amp; 5 place Jenson, F- 52200 Langres, France. charles. dheurle @ wanadoo. fr; https: // orcid. org / 0000 - 0002 - 2580 - 6962 Corresponding author &amp; Mendel University in Brno, Faculty of Forestry and Wood Technology, Department of Forest Ecology, Zemědělská 3, CZ- 613 00 Brno, Czech Republic. jirmor @ quick. cz; https: // orcid. org / 0000 - 0001 - 5294 - 6410</mods:affiliation>
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Genus
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<bibRefCitation id="EF9A4B73FFE0FFCDFE6E94073D359DDF" author="Chaudoir, M." box="[407,599,367,393]" pageId="6" pageNumber="7" pagination="3 - 21" refId="ref32376" refString="Chaudoir, M. (1850) Memoire sur la famille des Carabiques, 2 e Partie. Bulletin de la Societe Imperiale des Naturalistes de Moscou, 23, 3 - 21." type="journal article" year="1850">Chaudoir, 1850</bibRefCitation>
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<emphasis id="B97FEA90FFE0FFCDFF6E94DC3E7F9D9A" box="[151,285,436,460]" italics="true" pageId="6" pageNumber="7">Phaeoxantha</emphasis>
<bibRefCitation id="EF9A4B73FFE0FFCDFEDA94DC3EBE9D9D" author="Chaudoir, M." box="[291,476,435,460]" pageId="6" pageNumber="7" pagination="3 - 21" refId="ref32376" refString="Chaudoir, M. (1850) Memoire sur la famille des Carabiques, 2 e Partie. Bulletin de la Societe Imperiale des Naturalistes de Moscou, 23, 3 - 21." type="journal article" year="1850">Chaudoir, 1850: 7</bibRefCitation>
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species
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.
<taxonomicName id="4C0B4D01FFE0FFCDFEC094923DBD9E42" ID-CoL="738FL" authority="Perty, 1830: 2" authorityName="Perty" authorityPageNumber="2" authorityYear="1830" box="[313,735,506,532]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="laminata">
<emphasis id="B97FEA90FFE0FFCDFEC094923D5F9E42" box="[313,573,506,532]" italics="true" pageId="6" pageNumber="7">Megacephala laminata</emphasis>
<bibRefCitation id="EF9A4B73FFE0FFCDFDBA94923DBD9E42" author="Perty, M." box="[579,735,506,532]" pageId="6" pageNumber="7" pagination="125 - 224" refId="ref34474" refString="Perty, M. (1830 - 1833) Delectus animalium articulatorum in Brasilia. Insecta Brasiliensia, Coleoptera pentamera. Familia Cicindeletae. Munich, 224 pp. + 40 pls. [issued in three sections (only 1 and 3 cited here): Fascicle 1 (pp 1 - 60 + pls. 1 - 12, 1830). Fascicle 3 (pp 125 - 224 + pls. 25 - 40, 1833). The entire work along with the &quot; De insectorum in America meridionali &quot; was issued in 1834 yet with different titles of individual sections]." type="book chapter" year="1830">Perty, 1830: 2</bibRefCitation>
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subsequent
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designation by
<bibRefCitation id="EF9A4B73FFE0FFCDFB9894923A4A9E42" author="Horn, W." box="[1121,1320,506,533]" pageId="6" pageNumber="7" refId="ref33270" refString="Horn, W. (1915) Coleoptera Adephaga, Fam. Carabidae, Subfam. Cicindelinae. In: Wytsman, P. (Ed.), Genera Insectorum. Vol. 82. L. Desmet-Verteneuil, Bruxelles, pp. 209 - 487, pls. 16 - 23." type="book" year="1915">Horn (1915: 433)</bibRefCitation>
.
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<emphasis id="B97FEA90FFE0FFCDFF6E97563E439E00" box="[151,289,574,598]" italics="true" pageId="6" pageNumber="7">Megacephala</emphasis>
<bibRefCitation id="EF9A4B73FFE0FFCDFEDE97563EDF9E00" author="Latreille, P. A." box="[295,445,574,598]" pageId="6" pageNumber="7" refId="ref33633" refString="Latreille, P. A. (1802) Histoire naturelle generale et particuliere des Crustaces et des Insectes. Ouvrage faisant suite a l' histoire naturelle generale et particuliere, compose par Leclere de Buffon, et redigee par C. S. Sonnini, member de plusieurs societes savants. Familles naturelles des genres. Tome Troisieme. F. Dufart, Paris, xii + 13 - 447 + [1 (Errata)] pp." type="book" year="1802">Latreille, 1802</bibRefCitation>
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(partim)
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<bibRefCitation id="EF9A4B73FFE0FFCDFDD697563DBB9E00" author="Dejean, P. M. F. A." box="[559,729,574,598]" pageId="6" pageNumber="7" refId="ref32493" refString="Dejean, P. M. F. A. (1825) Species general des Coleopteres, de la collection de M. le Comte Dejean. Tome premier. Crevot, Paris, xxx + 463 pp. https: // doi. org / 10.5962 / bhl. title. 8863" type="book" year="1825">Dejean 1825: 14</bibRefCitation>
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<emphasis id="B97FEA90FFE0FFCDFF6E97353E7F9E23" box="[151,285,605,629]" italics="true" pageId="6" pageNumber="7">Phoeoxantha</emphasis>
(sic!): Lucas 1859: (wrong spelling of the genus-group name).
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<taxonomicName id="4C0B4D01FFE0FFCDFF6E97143EA29EC2" baseAuthorityName="W. Horn" baseAuthorityYear="1894" box="[151,448,636,660]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="subGenus" subGenus="Phaeoxantha">
<emphasis id="B97FEA90FFE0FFCDFF6E97143E439EC2" box="[151,289,636,660]" italics="true" pageId="6" pageNumber="7">Megacephala</emphasis>
(
<emphasis id="B97FEA90FFE0FFCDFEC997143ED49EC2" box="[304,438,636,660]" italics="true" pageId="6" pageNumber="7">Phaeoxantha</emphasis>
)
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:
<treatmentCitation id="0AAA1093FFE0FFCDFE3297143D0D9EC2" author="Horn, W." box="[459,623,636,660]" page="137" pageId="6" pageNumber="7" year="1910">
<bibRefCitation id="EF9A4B73FFE0FFCDFE3297143D0D9EC2" author="Horn, W." box="[459,623,636,660]" pageId="6" pageNumber="7" pagination="105 - 208" refId="ref33217" refString="Horn, W. (1910) Coleoptera Adephaga, Fam. Carabidae, Subfam. Cicindelinae. In: Wytsman, P. (Ed.), Genera Insectorum. Vol. 82 b. Desmet-Verteneuil, Bruxelles, pp. 105 - 208, pls. 6 - 15." type="book chapter" year="1910">Horn 1910: 137</bibRefCitation>
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<emphasis id="B97FEA90FFE0FFCDFF6E97F23E7F9EE4" box="[151,285,666,690]" italics="true" pageId="6" pageNumber="7">Phaeoxantha</emphasis>
</taxonomicName>
:
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<treatmentCitationGroup id="AB1B11ACFFE0FFCDFF6E97D13B069E87" box="[151,1124,697,721]" pageId="6" pageNumber="7">
<taxonomicName id="4C0B4D01FFE0FFCDFF6E97D13E079E87" ID-CoL="8LQTD" authority="Westwood" authorityName="Westwood" authorityYear="1852" box="[151,357,697,721]" class="Insecta" family="Carabidae" genus="Ammosia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="B97FEA90FFE0FFCDFF6E97D13F979E86" box="[151,245,697,720]" italics="true" pageId="6" pageNumber="7">Ammosia</emphasis>
Westwood
</taxonomicName>
[in Bates &amp; Westwood], 1852: 51, 57 synonymy by
<treatmentCitation id="0AAA1093FFE0FFCDFC7797D13B029E87" author="Chaudoir, M." box="[910,1120,697,721]" page="41" pageId="6" pageNumber="7" year="1865">
<bibRefCitation id="EF9A4B73FFE0FFCDFC7797D13B029E87" author="Chaudoir, M." box="[910,1120,697,721]" pageId="6" pageNumber="7" refId="ref32448" refString="Chaudoir, M. (1865) Catalogue de la collection des cicindel e t e s de M. le Baron Chaudoir. Nys, Bruxelles, 64 pp. https: // doi. org / 10.5962 / bhl. title. 134840" type="book" year="1865">Chaudoir (1865: 41)</bibRefCitation>
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.
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species of
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<emphasis id="B97FEA90FFE0FFCDFEA997973EDF9F4F" bold="true" box="[336,445,767,793]" italics="true" pageId="6" pageNumber="7">Ammosia</emphasis>
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.
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<taxonomicName id="4C0B4D01FFE0FFCDFE3297973C099F4F" ID-CoL="3YQRL" authority="Brulle, 1837" authorityName="Brulle" authorityYear="1837" box="[459,875,767,793]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="bifasciata">
<emphasis id="B97FEA90FFE0FFCDFE3297973DBA9F4F" box="[459,728,767,793]" italics="true" pageId="6" pageNumber="7">Megacephala bifasciata</emphasis>
<bibRefCitation id="EF9A4B73FFE0FFCDFD1997973C099F4F" author="Brulle, A." box="[736,875,767,793]" pageId="6" pageNumber="7" pagination="1 - 56" refId="ref32215" refString="Brulle, A. (1837) Insectes de l'Amerique meridionale recueillis par Alcide d'Orbigny et decrits par Emile Blanchard et Auguste Brulle. In: Voyage dans l'Amerique meridionale 6 (2 e Partie: Insectes). P. Bertrand, Paris [&amp;] V. Levrault, Strasbourg, pp. 1 - 56" type="book chapter" year="1837">Brullé, 1837</bibRefCitation>
</taxonomicName>
first
<typeStatus id="54B08820FFE0FFCDFC4296683C899F4C" box="[955,1003,768,794]" pageId="6" pageNumber="7">type</typeStatus>
designation by
<bibRefCitation id="EF9A4B73FFE0FFCDFB6597973AFE9F4F" author="Westwood, J. O." box="[1180,1436,767,794]" pageId="6" pageNumber="7" pagination="216 - 218" refId="ref34812" refString="Westwood, J. O. (1853) Observations on the dentition of the tiger beetles. The Annals and Magazine of Natural History, London, 12, Series 2, 216 - 218." type="journal article" year="1853">Westwood (1853: 218)</bibRefCitation>
[the inappropriate subsequent
<typeStatus id="54B08820FFE0FFCDFE1F964C3D749F68" box="[486,534,804,830]" pageId="6" pageNumber="7">type</typeStatus>
designation by
<bibRefCitation id="EF9A4B73FFE0FFCDFD3C964C3C339F6B" author="Horn, W." box="[709,849,803,830]" pageId="6" pageNumber="7" pagination="1 - 345" refId="ref33383" refString="Horn, W. (1926) Carabidae, Cicindelinae. In: Junk W. &amp; Schenkling S. (Eds.), Coleopterorum Catalogus, Volumen 30. W. Junk, Berlin, pp. 1 - 345." type="book chapter" year="1926">Horn (1926)</bibRefCitation>
, choosing
<taxonomicName id="4C0B4D01FFE0FFCDFC3F964C3B829F6B" ID-CoL="4FPYR" authority="Klug, 1834" authorityName="Klug" authorityYear="1834" box="[966,1248,803,829]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="testudinea">
<emphasis id="B97FEA90FFE0FFCDFC3F964C3B3F9F6B" box="[966,1117,803,829]" italics="true" pageId="6" pageNumber="7">M. testudinea</emphasis>
<bibRefCitation id="EF9A4B73FFE0FFCDFB9A964B3B829F6B" author="Klug, F." box="[1123,1248,803,829]" pageId="6" pageNumber="7" pagination="1 - 47" refId="ref33461" refString="Klug, F. (1834) Uebersicht der Cicindeletae der Sammlung. Jahrbucher der Insectenkunde, 1, 1 - 47, pl. 1. https: // doi. org / 10.1515 / 9783112606926 - 002" type="journal article" year="1834">Klug, 1834</bibRefCitation>
</taxonomicName>
as a
<typeStatus id="54B08820FFE0FFCDFAEC964C3A279F68" box="[1301,1349,804,830]" pageId="6" pageNumber="7">type</typeStatus>
species of
<taxonomicName id="4C0B4D01FFE0FFCDFF4F962F3E7C9F37" ID-CoL="8LQTD" authorityName="Westwood" authorityYear="1852" box="[182,286,839,865]" class="Insecta" family="Carabidae" genus="Ammosia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="B97FEA90FFE0FFCDFF4F962F3E7C9F37" box="[182,286,839,865]" italics="true" pageId="6" pageNumber="7">Ammosia</emphasis>
</taxonomicName>
, is unavailable].
</paragraph>
</subSubSection>
<subSubSection id="C3116509FFE0FFCDFF3E96033C599FA7" pageId="6" pageNumber="7" type="discussion">
<paragraph id="8BB43682FFE0FFCDFF3E96033C599FA7" blockId="6.[151,1437,767,2018]" pageId="6" pageNumber="7">
<emphasis id="B97FEA90FFE0FFCDFF3E96033F9D9FD3" bold="true" box="[199,255,875,901]" pageId="6" pageNumber="7">Note</emphasis>
.
<taxonomicName id="4C0B4D01FFE0FFCDFEF696033DF19FD3" ID-CoL="3YQV8" authority="(Perty, 1833)" baseAuthorityName="Perty" baseAuthorityYear="1833" box="[271,659,875,901]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="limata">
<emphasis id="B97FEA90FFE0FFCDFEF696033E969FD3" box="[271,500,875,901]" italics="true" pageId="6" pageNumber="7">Phaeoxantha limata</emphasis>
(Perty, 1833)
</taxonomicName>
, based on
<taxonomicName id="4C0B4D01FFE0FFCDFCF696033C9A9FD3" ID-CoL="3YQV8" authorityName="Perty" authorityYear="1830" box="[783,1016,875,901]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="limata">
<emphasis id="B97FEA90FFE0FFCDFCF696033C9A9FD3" box="[783,1016,875,901]" italics="true" pageId="6" pageNumber="7">Megacephala limata</emphasis>
</taxonomicName>
, is an unjustified emendation for
<taxonomicName id="4C0B4D01FFE0FFCDFA8796043EE69FFF" ID-CoL="738FL" authority="Perty, 1830" authorityName="Perty" authorityYear="1830" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="laminata">
<emphasis id="B97FEA90FFE0FFCDFA8796043F999FFF" italics="true" pageId="6" pageNumber="7">M. laminata</emphasis>
<bibRefCitation id="EF9A4B73FFE0FFCDFEFB96F83EE69FFF" author="Perty, M." box="[258,388,911,938]" pageId="6" pageNumber="7" pagination="125 - 224" refId="ref34474" refString="Perty, M. (1830 - 1833) Delectus animalium articulatorum in Brasilia. Insecta Brasiliensia, Coleoptera pentamera. Familia Cicindeletae. Munich, 224 pp. + 40 pls. [issued in three sections (only 1 and 3 cited here): Fascicle 1 (pp 1 - 60 + pls. 1 - 12, 1830). Fascicle 3 (pp 125 - 224 + pls. 25 - 40, 1833). The entire work along with the &quot; De insectorum in America meridionali &quot; was issued in 1834 yet with different titles of individual sections]." type="book chapter" year="1830">Perty, 1830</bibRefCitation>
</taxonomicName>
. Consequently, despite the fact that
<bibRefCitation id="EF9A4B73FFE0FFCDFCEF96F83CC39FFC" author="Horn, W." box="[790,929,911,938]" pageId="6" pageNumber="7" refId="ref33270" refString="Horn, W. (1915) Coleoptera Adephaga, Fam. Carabidae, Subfam. Cicindelinae. In: Wytsman, P. (Ed.), Genera Insectorum. Vol. 82. L. Desmet-Verteneuil, Bruxelles, pp. 209 - 487, pls. 16 - 23." type="book" year="1915">Horn (1915)</bibRefCitation>
used the name
<taxonomicName id="4C0B4D01FFE0FFCDFBA996F83B9C9FFC" ID-CoL="3YQV8" authority="Perty" authorityName="Perty" authorityYear="1830" box="[1104,1278,911,938]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="limata">
<emphasis id="B97FEA90FFE0FFCDFBA996F83BDF9FFF" box="[1104,1213,911,937]" italics="true" pageId="6" pageNumber="7">M. limata</emphasis>
Perty
</taxonomicName>
for his subsequent
<typeStatus id="54B08820FFE0FFCDFF2596DC3E6E9F98" box="[220,268,948,974]" pageId="6" pageNumber="7">type</typeStatus>
designation, according to the Articles 67.6 and 69.2, 1 of the
<bibRefCitation id="EF9A4B73FFE0FFCDFC4D96DB3B2A9F9B" author="ICZN" box="[948,1096,947,973]" pageId="6" pageNumber="7" refId="ref33426" refString="ICZN (1999) International Code of Zoological Nomenclature, fourth edition, adopted by the International Union of Biological Sciences. International Trust for Zoological Nomenclature, London, xxix + 306 pp." type="book" year="1999">ICZN (1999)</bibRefCitation>
, the
<typeStatus id="54B08820FFE0FFCDFB8296DC3BC99F98" box="[1147,1195,948,974]" pageId="6" pageNumber="7">type</typeStatus>
species must be cited in its correct original spelling, thus
<taxonomicName id="4C0B4D01FFE0FFCDFDDC96B03C559FA7" ID-CoL="738FL" authority="Perty, 1830" authorityName="Perty" authorityYear="1830" box="[549,823,983,1010]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="laminata">
<emphasis id="B97FEA90FFE0FFCDFDDC96B03DCC9FA7" box="[549,686,983,1009]" italics="true" pageId="6" pageNumber="7">M. laminata</emphasis>
<bibRefCitation id="EF9A4B73FFE0FFCDFD4D96B03C559FA7" author="Perty, M." box="[692,823,983,1010]" pageId="6" pageNumber="7" pagination="125 - 224" refId="ref34474" refString="Perty, M. (1830 - 1833) Delectus animalium articulatorum in Brasilia. Insecta Brasiliensia, Coleoptera pentamera. Familia Cicindeletae. Munich, 224 pp. + 40 pls. [issued in three sections (only 1 and 3 cited here): Fascicle 1 (pp 1 - 60 + pls. 1 - 12, 1830). Fascicle 3 (pp 125 - 224 + pls. 25 - 40, 1833). The entire work along with the &quot; De insectorum in America meridionali &quot; was issued in 1834 yet with different titles of individual sections]." type="book chapter" year="1830">Perty, 1830</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C3116509FFE0FFCDFF3E96933CF998BB" pageId="6" pageNumber="7" type="diagnosis">
<paragraph id="8BB43682FFE0FFCDFF3E96933AF198F3" blockId="6.[151,1437,767,2018]" pageId="6" pageNumber="7">
<emphasis id="B97FEA90FFE0FFCDFF3E96933EAC9843" bold="true" box="[199,462,1019,1045]" pageId="6" pageNumber="7">Differential diagnosis.</emphasis>
Due to their pale ivory-yellow (sometimes partly almost translucent), ochraceous, cinnamon-ochre to rusty-brownish basic body coloration (mostly with more or less expanded dark brown or black elytral areas) and absence of any metallic-green-blue coloration, the 12 species and one subspecies of the genus immediately differ from all other genera of the tribe
<taxonomicName id="4C0B4D01FFE0FFCDFD63910F3B1998D7" authority="Laporte de Castelnau, 1834" authorityName="Hope" authorityYear="1838" box="[666,1147,1127,1154]" class="Insecta" family="Carabidae" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="tribe" tribe="Megacephalini">
Megacephalini
<bibRefCitation id="EF9A4B73FFE0FFCDFCBE91003B1998D7" author="Laporte de Castelnau, F. L." box="[839,1147,1127,1154]" pageId="6" pageNumber="7" pagination="27 - 38" refId="ref33606" refString="Laporte de Castelnau, F. L. (1834) Observations sur la tribu des Cicindeletes. Revue Entomologique, 2, 27 - 38." type="journal article" year="1834">Laporte de Castelnau, 1834</bibRefCitation>
</taxonomicName>
. Legs mostly concolorous with body. Buccal appendages, particularly both maxillary and labial palpi very long, ivory-yellow to ochraceous.
</paragraph>
<paragraph id="8BB43682FFE0FFCDFF3E91C73CF998BB" blockId="6.[151,1437,767,2018]" pageId="6" pageNumber="7">The genus is here subdivided into two distinctly differentiated subgenera (which due to their distinct differentiating characters may alternatively be treated as two separate genera).</paragraph>
</subSubSection>
<subSubSection id="C3116509FFE0FFCDFF3E919F3AFE9BB7" pageId="6" pageNumber="7" type="distribution">
<paragraph id="8BB43682FFE0FFCDFF3E919F3C549B5C" blockId="6.[151,1437,767,2018]" pageId="6" pageNumber="7">
<emphasis id="B97FEA90FFE0FFCDFF3E919F3D3B9947" bold="true" box="[199,601,1271,1297]" pageId="6" pageNumber="7">Distribution, ecology and biology</emphasis>
. Species of the genus
<taxonomicName id="4C0B4D01FFE0FFCDFCA6919F3C919947" authorityName="Chaudoir" authorityYear="1850" box="[863,1011,1271,1297]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="B97FEA90FFE0FFCDFCA6919F3C919947" box="[863,1011,1271,1297]" italics="true" pageId="6" pageNumber="7">Phaeoxantha</emphasis>
</taxonomicName>
are widely distributed in most countries of South America (none of the species penetrates from
<collectingCountry id="F31C7612FFE0FFCDFCC090733CC59963" box="[825,935,1307,1333]" name="Colombia" pageId="6" pageNumber="7">Colombia</collectingCountry>
to Central America through the Panamanian Isthmus); the countries of their occurrence are mentioned in each individual species below. Majority of them occur throughout the large Amazon Basin and partly also in the area of the Rio de la Plata Basin. Both adults and larvae prefer sandy beaches, semidry or dry riverbeds, but also marshlands. Adults are nocturnal, some of them flightless, several others with only restricted flight ability due to sexually dimorphic development of flight muscle in some species, as observed by
<bibRefCitation id="EF9A4B73FFE0FFCDFE5D90B83DEC99BF" author="Zerm, M. &amp; Adis, J." box="[420,654,1487,1514]" pageId="6" pageNumber="7" pagination="491 - 500" refId="ref35148" refString="Zerm, M. &amp; Adis, J. (2002) Flight ability in nocturnal tiger beetles (Coleoptera: Cicindelidae) from Central Amazonian floodplains (Brazil). Coleopterist Bulletin, 56 (4), 491 - 500. https: // doi. org / 10.1649 / 0010 - 065 X (2002) 056 [0491: FAINTB] 2.0. CO; 2" type="journal article" year="2002">Zerm &amp; Adis (2002)</bibRefCitation>
. Life cycles of adults and larvae of species occurring in open areas of Brazilian Central Amazonian floodplains in the region of Manaus, including larval development (also reared in the laboratory), were presented by Adis
<emphasis id="B97FEA90FFE0FFCDFDD493703D079A67" box="[557,613,1559,1585]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(1998),
<bibRefCitation id="EF9A4B73FFE0FFCDFD3D93703CCC9A67" author="Zerm, M. &amp; Adis, J." box="[708,942,1559,1586]" pageId="6" pageNumber="7" pagination="141 - 155" refId="ref34992" refString="Zerm, M. &amp; Adis, J. (2001 a) On the life cycles of Phaeoxantha species (Coleoptera: Cicindelidae) from Central Amazonian floodplains (Brazil). Ecotropica, 6, 141 - 155." type="journal article" year="2001">Zerm &amp; Adis (2001a</bibRefCitation>
, b, c) and Zerm
<emphasis id="B97FEA90FFE0FFCDFB9D93703BFE9A67" box="[1124,1180,1559,1585]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(2001). Their biotopes in
<collectingCountry id="F31C7612FFE0FFCDFF4A93533E679A03" box="[179,261,1595,1621]" name="Bolivia" pageId="6" pageNumber="7">Bolivia</collectingCountry>
were described by
<bibRefCitation id="EF9A4B73FFE0FFCDFE2093543DAF9A03" author="Pearson, D. L. &amp; Guerra, J. F. &amp; Brzoska, D. W." box="[473,717,1595,1622]" pageId="6" pageNumber="7" pagination="379 - 524" refId="ref34364" refString="Pearson, D. L., Guerra, J. F. &amp; Brzoska, D. W. (1999 a) The Tiger beetles of Bolivia: their Identification, Distribution and Natural History (Coleoptera: Cicindelidae). Contributions on Entomology, International, 3 (4), 379 - 524." type="journal article" year="1999">
Pearson
<emphasis id="B97FEA90FFE0FFCDFDC193543D0D9A03" box="[568,623,1595,1621]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(1999a)
</bibRefCitation>
, and examined species from
<collectingCountry id="F31C7612FFE0FFCDFBF293533B3F9A03" box="[1035,1117,1595,1621]" name="Bolivia" pageId="6" pageNumber="7">Bolivia</collectingCountry>
preferred the same biotopes, mostly open sandy beaches or riverbeds of periodical rivers which are usually dry during the day because of strong evaporation in hot sunny days (Ondřej Šafranek. pers. com.). For details see under individual species below. The larvae were partly or with more characters described and illustrated by Arndt
<emphasis id="B97FEA90FFE0FFCDFBF893C03B589A97" box="[1025,1082,1703,1729]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(1996), with a key to larvae by Adis
<emphasis id="B97FEA90FFE0FFCDFF2D93A43E6E9AB3" box="[212,268,1739,1765]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(1998) and Arndt
<emphasis id="B97FEA90FFE0FFCDFE2493A43D749AB3" box="[477,534,1739,1765]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(2002) yet no entire habitus of a larval instar was illustrated in the cited papers. Pupal morphology was presented by
<bibRefCitation id="EF9A4B73FFE0FFCDFDCF93873C509B5C" author="Cardenas, A. M. &amp; Zerm, M. &amp; Adis, J." box="[566,818,1775,1802]" pageId="6" pageNumber="7" pagination="113 - 121" refId="ref32273" refString="Cardenas, A. M., Zerm, M. &amp; Adis, J. (2005) Pupal morphology of several species of tiger beetle (Coleoptera: Cicindelidae) from Central Amazonian floodplains (Brazil). Studies on Neotropical Fauna and Environment, 40 (2) 113 - 121. https: // doi. org / 10.1080 / 01650520500138449" type="journal article" year="2005">
Cárdenas
<emphasis id="B97FEA90FFE0FFCDFD5C93983DBF9B5F" box="[677,733,1775,1801]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(2005)
</bibRefCitation>
.
</paragraph>
<paragraph id="8BB43682FFE0FFCDFF3E927C3AFE9BB7" blockId="6.[151,1437,767,2018]" pageId="6" pageNumber="7">
<bibRefCitation id="EF9A4B73FFE0FFCDFF3E927C3EFD9B7B" author="Zerm, M. &amp; Wiesner, J. &amp; Ledezma, J. &amp; Brzoska, D. &amp; Drechsel, U. &amp; Cicchino, A. C. &amp; RodrIguez, J. P. &amp; Martinsen, L. &amp; Adis, J." box="[199,415,1811,1838]" pageId="6" pageNumber="7" pagination="211 - 219" refId="ref35361" refString="Zerm, M., Wiesner, J., Ledezma, J., Brzoska, D., Drechsel, U., Cicchino, A. C., RodrIguez, J. P., Martinsen, L., Adis, J. &amp; Bach- mann, L. (2007) Molecular phylogeny of Megacephalina Horn, 1910, tiger beetles (Coleoptera: Cicindelidae). Studies on Neotropical Fauna and Environment, 42, 211 - 219. https: // doi. org / 10.1080 / 01650520701409235" type="journal article" year="2007">
Zerm
<emphasis id="B97FEA90FFE0FFCDFEF4927C3E2A9B7B" box="[269,328,1811,1837]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(2007)
</bibRefCitation>
, in their phylogenetic analysis, based on the nuclear 18S and the mitochondrial 16S and cytochrome oxidase III genes, have concluded that
<taxonomicName id="4C0B4D01FFE0FFCDFD26925F3CAE9B07" box="[735,972,1847,1873]" pageId="6" pageNumber="7">
<emphasis id="B97FEA90FFE0FFCDFD26925F3C119B07" box="[735,883,1847,1873]" italics="true" pageId="6" pageNumber="7">Phaeoxantha</emphasis>
species
</taxonomicName>
form a monophyletic group of three distinct clades: 1)
<emphasis id="B97FEA90FFE0FFCDFEBA92333D2E9B23" box="[323,588,1883,1909]" italics="true" pageId="6" pageNumber="7">
<taxonomicName id="4C0B4D01FFE0FFCDFEBA92333E839B23" authorityName="sensu Dejean" authorityYear="1825" box="[323,481,1883,1909]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="aequinoctialis">aequinoctialis</taxonomicName>
<taxonomicName id="4C0B4D01FFE0FFCDFE0992333D2E9B23" baseAuthorityName="Brulle" baseAuthorityYear="1837" box="[496,588,1883,1909]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="cruciata">cruciata</taxonomicName>
</emphasis>
; 2)
<taxonomicName id="4C0B4D01FFE0FFCDFD8392333DA09B23" authorityName="Perty" authorityYear="1830" box="[634,706,1883,1909]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="limata">
<emphasis id="B97FEA90FFE0FFCDFD8392333DA09B23" box="[634,706,1883,1909]" italics="true" pageId="6" pageNumber="7">limata</emphasis>
</taxonomicName>
; 3)
<emphasis id="B97FEA90FFE0FFCDFD1792333B5F9B23" box="[750,1085,1883,1909]" italics="true" pageId="6" pageNumber="7">
<taxonomicName id="4C0B4D01FFE0FFCDFD1792333C4E9B23" baseAuthorityName="Chaudoir" baseAuthorityYear="1850" box="[750,812,1883,1909]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="klugii">klugii</taxonomicName>
<taxonomicName id="4C0B4D01FFE0FFCDFCC292333CAE9B23" baseAuthorityName="Mandl" baseAuthorityYear="1964" box="[827,972,1883,1909]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="lindemannae">lindemannae</taxonomicName>
<taxonomicName id="4C0B4D01FFE0FFCDFC2292333B5F9B23" baseAuthorityName="Mandl" baseAuthorityYear="1958" box="[987,1085,1883,1909]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="wimmeri">wimmeri</taxonomicName>
</emphasis>
. This is in accordance with our concept based on morphology, as well as with our division into the two subgenera as presented here. It must be, however, mentioned here that under the name
<taxonomicName id="4C0B4D01FFE0FFCDFD6092CB3C5A9BEB" authorityName="sensu Dejean" authorityYear="1825" box="[665,824,1955,1981]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="aequinoctialis">
<emphasis id="B97FEA90FFE0FFCDFD6092CB3C5A9BEB" box="[665,824,1955,1981]" italics="true" pageId="6" pageNumber="7">aequinoctialis</emphasis>
</taxonomicName>
the authors studied in fact
<taxonomicName id="4C0B4D01FFE0FFCDFB9B92CC3A429BEB" authorityName="Brulle" authorityYear="1837" box="[1122,1312,1955,1982]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="9" phylum="Arthropoda" rank="species" species="bifasciata" subGenus="Phaeoxantha">
<emphasis id="B97FEA90FFE0FFCDFB9B92CC3B119BEB" box="[1122,1139,1956,1981]" italics="true" pageId="6" pageNumber="7">P</emphasis>
. (
<emphasis id="B97FEA90FFE0FFCDFB7392CC3BF99BEB" box="[1162,1179,1956,1981]" italics="true" pageId="6" pageNumber="7">P</emphasis>
.)
<emphasis id="B97FEA90FFE0FFCDFB4B92CB3A429BEB" box="[1202,1312,1955,1981]" italics="true" pageId="6" pageNumber="7">bifasciata</emphasis>
</taxonomicName>
, and under the name
<taxonomicName id="4C0B4D01FFE0FFCDFEFD92AF3E2E9BB7" authorityName="Perty" authorityYear="1830" box="[260,332,1991,2017]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="limata">
<emphasis id="B97FEA90FFE0FFCDFEFD92AF3E2E9BB7" box="[260,332,1991,2017]" italics="true" pageId="6" pageNumber="7">limata</emphasis>
</taxonomicName>
studied either
<taxonomicName id="4C0B4D01FFE0FFCDFE0892A03DC59BB7" baseAuthorityName="Dejean" baseAuthorityYear="1831" box="[497,679,1991,2018]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="4" phylum="Arthropoda" rank="species" species="nocturna" subGenus="Phaeoxantha">
<emphasis id="B97FEA90FFE0FFCDFE0892A03D609BB7" box="[497,514,1992,2017]" italics="true" pageId="6" pageNumber="7">P</emphasis>
. (
<emphasis id="B97FEA90FFE0FFCDFDE392A03D499BB7" box="[538,555,1992,2017]" italics="true" pageId="6" pageNumber="7">P</emphasis>
.)
<emphasis id="B97FEA90FFE0FFCDFDBB92A03DC59BB7" box="[578,679,1992,2017]" italics="true" pageId="6" pageNumber="7">nocturna</emphasis>
</taxonomicName>
or
<emphasis id="B97FEA90FFE0FFCDFD3592A03DBF9BB7" box="[716,733,1992,2017]" italics="true" pageId="6" pageNumber="7">P</emphasis>
.
<taxonomicName id="4C0B4D01FFE0FFCDFD1592AF3CE19BB7" authorityName="Perty" authorityYear="1830" box="[748,899,1991,2017]" class="Insecta" family="Carabidae" genus="Megacephala" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="laminata" subGenus="Phaeoxantha">
(
<emphasis id="B97FEA90FFE0FFCDFD0C92A03C649BB7" box="[757,774,1992,2017]" italics="true" pageId="6" pageNumber="7">P</emphasis>
.)
<emphasis id="B97FEA90FFE0FFCDFCE492AF3CE19BB7" box="[797,899,1991,2017]" italics="true" pageId="6" pageNumber="7">laminata</emphasis>
</taxonomicName>
<emphasis id="B97FEA90FFE0FFCDFC7D92AF3CEA9BB7" box="[900,904,1991,2017]" italics="true" pageId="6" pageNumber="7">,</emphasis>
which were confused at the time of their study.
</paragraph>
</subSubSection>
<subSubSection id="C3116509FFE1FFCCFF6E95FF3AFE9D0B" pageId="7" pageNumber="8" type="key">
<paragraph id="8BB43682FFE1FFCCFF6E95FF3EBE9CE7" blockId="7.[151,476,151,177]" box="[151,476,151,177]" pageId="7" pageNumber="8">
<emphasis id="B97FEA90FFE1FFCCFF6E95FF3EBE9CE7" bold="true" box="[151,476,151,177]" pageId="7" pageNumber="8">Key to the two subgenera.</emphasis>
</paragraph>
<keyStep id="30FF2E27FFE1FFCCFF6E95BB3AFE9D0B" pageId="7" pageNumber="8">
<key id="FF9A61B3FFE1FFCCFF6E95BB3AFE9D0B" pageId="7" pageNumber="8">
<paragraph id="8BB43682FFE1FFCCFF6E95BB3AFE9D50" blockId="7.[151,1437,211,350]" pageId="7" pageNumber="8">
<keyLead id="30FA95B7FFE1FFCCFF6E95BB3AFE9D50" pageId="7" pageNumber="8">
1 Labrum with lateral margins arcuate (
<figureCitation id="13302A07FFE1FFCCFDB595BB3DC69CBE" box="[588,676,211,233]" captionStart="FIGURES 112" captionStartId="8.[151,264,1770,1794]" captionTargetBox="[200,1389,190,1734]" captionTargetId="figure-30@8.[189,1398,179,1744]" captionTargetPageId="8" captionText="FIGURES 112. Basic diagnostic characters of two subgenera of Phaeoxantha. 17—P. (Phaeoxantha). 13—male right mandible: 1—P. (P.) nocturna (Dejean); 2—P. (P.) laminata (Perty); 3—P. (P.) bifasciata (Brullé); 47—labrum: 4—P. (P.) nocturna (♂); 5—P. (P.) laminata (♂); 6—P. (P.) tremolerasi (W. Horn); 7—P. (P.) bifasciata (Brullé). 8—12—P. (Euphae- oxantha). 89—male right mandible: 8—P. (E.) wimmeri (Mandl); 9—P. (E.) klugii (Chaudoir); 1012—labrum: 1011—P. (E.) wimmeri (♂, ♀); 12—P. (E.) klugii." figureDoi="http://doi.org/10.5281/zenodo.10376590" httpUri="https://zenodo.org/record/10376590/files/figure.png" pageId="7" pageNumber="8">Figs 47</figureCitation>
). Mandibles normally shaped in both sexes (
<figureCitation id="13302A07FFE1FFCCFBA995BB3BC59CBE" box="[1104,1191,211,233]" captionStart="FIGURES 112" captionStartId="8.[151,264,1770,1794]" captionTargetBox="[200,1389,190,1734]" captionTargetId="figure-30@8.[189,1398,179,1744]" captionTargetPageId="8" captionText="FIGURES 112. Basic diagnostic characters of two subgenera of Phaeoxantha. 17—P. (Phaeoxantha). 13—male right mandible: 1—P. (P.) nocturna (Dejean); 2—P. (P.) laminata (Perty); 3—P. (P.) bifasciata (Brullé); 47—labrum: 4—P. (P.) nocturna (♂); 5—P. (P.) laminata (♂); 6—P. (P.) tremolerasi (W. Horn); 7—P. (P.) bifasciata (Brullé). 8—12—P. (Euphae- oxantha). 89—male right mandible: 8—P. (E.) wimmeri (Mandl); 9—P. (E.) klugii (Chaudoir); 1012—labrum: 1011—P. (E.) wimmeri (♂, ♀); 12—P. (E.) klugii." figureDoi="http://doi.org/10.5281/zenodo.10376590" httpUri="https://zenodo.org/record/10376590/files/figure.png" pageId="7" pageNumber="8">Figs 13</figureCitation>
). Body small to medium-sized (length
<quantity id="4CF39B67FFE1FFCCFE9A95983EB09D53" box="[355,466,240,261]" metricMagnitude="-2" metricUnit="m" metricValue="1.365" metricValueMax="1.9" metricValueMin="0.83" pageId="7" pageNumber="8" unit="mm" value="13.65" valueMax="19.0" valueMin="8.3">8.319 mm</quantity>
)...........................................
<taxonomicName id="4C0B4D01FFE1FFCCFC1695983AFE9D50" authority="Chaudoir, 1850" authorityName="Chaudoir" authorityYear="1850" box="[1007,1436,240,262]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="subGenus" subGenus="Phaeoxantha">
<emphasis id="B97FEA90FFE1FFCCFC1695983B0C9D53" box="[1007,1134,240,261]" italics="true" pageId="7" pageNumber="8">Phaeoxantha</emphasis>
(
<emphasis id="B97FEA90FFE1FFCCFB8295983B989D53" box="[1147,1274,240,261]" italics="true" pageId="7" pageNumber="8">Phaeoxantha</emphasis>
)
<bibRefCitation id="EF9A4B73FFE1FFCCFAF195983AFE9D50" author="Chaudoir, M." box="[1288,1436,240,262]" pageId="7" pageNumber="8" pagination="3 - 21" refId="ref32376" refString="Chaudoir, M. (1850) Memoire sur la famille des Carabiques, 2 e Partie. Bulletin de la Societe Imperiale des Naturalistes de Moscou, 23, 3 - 21." type="journal article" year="1850">Chaudoir, 1850</bibRefCitation>
</taxonomicName>
</keyLead>
</paragraph>
<paragraph id="8BB43682FFE1FFCCFF6E94663AFE9D0B" blockId="7.[151,1437,211,350]" pageId="7" pageNumber="8">
<keyLead id="30FA95B7FFE1FFCCFF6E94663AFE9D0B" pageId="7" pageNumber="8">
Labrum with lateral margins dorsally excised in either side, forming large lateral teeth (
<figureCitation id="13302A07FFE1FFCCFBC094653BC89D75" box="[1081,1194,269,291]" captionStart="FIGURES 112" captionStartId="8.[151,264,1770,1794]" captionTargetBox="[200,1389,190,1734]" captionTargetId="figure-30@8.[189,1398,179,1744]" captionTargetPageId="8" captionText="FIGURES 112. Basic diagnostic characters of two subgenera of Phaeoxantha. 17—P. (Phaeoxantha). 13—male right mandible: 1—P. (P.) nocturna (Dejean); 2—P. (P.) laminata (Perty); 3—P. (P.) bifasciata (Brullé); 47—labrum: 4—P. (P.) nocturna (♂); 5—P. (P.) laminata (♂); 6—P. (P.) tremolerasi (W. Horn); 7—P. (P.) bifasciata (Brullé). 8—12—P. (Euphae- oxantha). 89—male right mandible: 8—P. (E.) wimmeri (Mandl); 9—P. (E.) klugii (Chaudoir); 1012—labrum: 1011—P. (E.) wimmeri (♂, ♀); 12—P. (E.) klugii." figureDoi="http://doi.org/10.5281/zenodo.10376590" httpUri="https://zenodo.org/record/10376590/files/figure.png" pageId="7" pageNumber="8">Figs 1012</figureCitation>
). Male mandibles abnormally shaped, particularly right mandible conspicuously aberrant (
<figureCitation id="13302A07FFE1FFCCFCA094433CD29D16" box="[857,944,299,321]" captionStart="FIGURES 112" captionStartId="8.[151,264,1770,1794]" captionTargetBox="[200,1389,190,1734]" captionTargetId="figure-30@8.[189,1398,179,1744]" captionTargetPageId="8" captionText="FIGURES 112. Basic diagnostic characters of two subgenera of Phaeoxantha. 17—P. (Phaeoxantha). 13—male right mandible: 1—P. (P.) nocturna (Dejean); 2—P. (P.) laminata (Perty); 3—P. (P.) bifasciata (Brullé); 47—labrum: 4—P. (P.) nocturna (♂); 5—P. (P.) laminata (♂); 6—P. (P.) tremolerasi (W. Horn); 7—P. (P.) bifasciata (Brullé). 8—12—P. (Euphae- oxantha). 89—male right mandible: 8—P. (E.) wimmeri (Mandl); 9—P. (E.) klugii (Chaudoir); 1012—labrum: 1011—P. (E.) wimmeri (♂, ♀); 12—P. (E.) klugii." figureDoi="http://doi.org/10.5281/zenodo.10376590" httpUri="https://zenodo.org/record/10376590/files/figure.png" pageId="7" pageNumber="8">Figs 89</figureCitation>
). Body large to very large and mostly robust (length
<quantity id="4CF39B67FFE1FFCCFF1B94203E299D0B" box="[226,331,328,349]" metricMagnitude="-2" metricUnit="m" metricValue="2.35" metricValueMax="2.7" metricValueMin="2.0" pageId="7" pageNumber="8" unit="mm" value="23.5" valueMax="27.0" valueMin="20.0">2027 mm</quantity>
)........................................................
<taxonomicName id="4C0B4D01FFE1FFCCFC0A94203A7C9D0B" authorityName="Moravec &amp; Dheurle" authorityYear="2023" box="[1011,1310,328,349]" class="Insecta" family="Carabidae" genus="Phaeoxantha" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="8" phylum="Arthropoda" rank="subGenus" subGenus="Euphaeoxantha">
<emphasis id="B97FEA90FFE1FFCCFC0A94203B109D0B" box="[1011,1138,328,349]" italics="true" pageId="7" pageNumber="8">Phaeoxantha</emphasis>
(
<emphasis id="B97FEA90FFE1FFCCFB7994203A759D0B" box="[1152,1303,328,349]" italics="true" pageId="7" pageNumber="8">Euphaeoxantha</emphasis>
)
</taxonomicName>
<emphasis id="B97FEA90FFE1FFCCFADD94203AFE9D0B" bold="true" box="[1316,1436,328,349]" pageId="7" pageNumber="8">subgen nov.</emphasis>
</keyLead>
</paragraph>
</key>
</keyStep>
</subSubSection>
</treatment>
</document>