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<mods:title id="FC4C7AA35B8449A572B3A19AAA27CB1A">The Higher Classification of the Ant Subfamily Ponerinae (Hymenoptera: Formicidae), with a Review of Ponerine Ecology and Behavior</mods:title>
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<subSubSection id="C3C4BB9BA6E82CB5FF17FE3A11D2FE0C" box="[151,439,476,502]" pageId="151" pageNumber="152" type="nomenclature">
<paragraph id="8B61E810A6E82CB5FF17FE3A11D2FE0C" blockId="151.[151,439,476,535]" box="[151,439,476,502]" pageId="151" pageNumber="152">
<heading id="D0295F7CA6E82CB5FF17FE3A11D2FE0C" bold="true" box="[151,439,476,502]" fontSize="11" level="2" pageId="151" pageNumber="152" reason="4">
<taxonomicName id="4CDE9393A6E82CB5FF17FE3A11D2FE0C" ID-CoL="6BRJ" ID-ENA="118887" authority="Smith, F." authorityName="Smith, F." box="[151,439,476,502]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FF17FE3A11D2FE0C" bold="true" box="[151,439,476,502]" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF17FE3A1125FE0C" bold="true" box="[151,320,476,502]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
Smith, F.
</emphasis>
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</heading>
</paragraph>
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<subSubSection id="C3C4BB9BA6E82CB5FF17FE181083FDED" box="[151,230,510,535]" pageId="151" pageNumber="152" type="description">
<paragraph id="8B61E810A6E82CB5FF17FE181083FDED" blockId="151.[151,439,476,535]" box="[151,230,510,535]" pageId="151" pageNumber="152">
<figureCitation id="13E5F495A6E82CB5FF17FE181083FDED" box="[151,230,510,535]" captionStart="FIGURE 31" captionStartId="152.[151,250,1774,1796]" captionTargetBox="[151,1436,323,1749]" captionTargetId="figure-216@152.[151,1436,323,1761]" captionTargetPageId="152" captionText="FIGURE 31. Worker caste of Pachycondyla crassinoda: lateral and dorsal view of body and full-face view of head (CASENT0249138, Ryan Perry and www.antweb.org); world distribution of Pachycondyla." figureDoi="http://doi.org/10.5281/zenodo.10120882" httpUri="https://zenodo.org/record/10120882/files/figure.png" pageId="151" pageNumber="152">Fig. 31</figureCitation>
</paragraph>
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<subSubSection id="C3C4BB9BA6E82CB5FF17FDA0112DFCD7" pageId="151" pageNumber="152" type="reference_group">
<paragraph id="8B61E810A6E82CB5FF17FDA012E6FD81" blockId="151.[151,1436,582,635]" pageId="151" pageNumber="152">
<treatmentCitationGroup id="ABCECF3EA6E82CB5FF17FDA012E6FD81" pageId="151" pageNumber="152">
<taxonomicName id="4CDE9393A6E82CB5FF17FDA01275FDA6" ID-CoL="6BRJ" authority="Smith, F., 1858: 105" authorityName="Smith, F." authorityPageNumber="105" authorityYear="1858" box="[151,528,582,604]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FF17FDA01143FDA6" box="[151,294,582,604]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
Smith, F., 1858: 105
</taxonomicName>
(as genus in Poneridae). Type-species: 
<treatmentCitation id="0A7FCE01A6E82CB5FC40FDA11516FDA6" author="Latreille, P. A." box="[960,1395,582,604]" page="198" pageId="151" pageNumber="152" year="1802">
<taxonomicName id="4CDE9393A6E82CB5FC40FDA11516FDA6" authority="Latreille, 1802 b: 198" authorityName="Latreille" authorityPageNumber="198" authorityYear="1802" box="[960,1395,582,604]" class="Insecta" family="Formicidae" genus="Ponera" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="crassinoda">
<emphasis id="B9AA3402A6E82CB5FC40FDA114E0FDA6" box="[960,1157,582,604]" italics="true" pageId="151" pageNumber="152">Ponera crassinoda</emphasis>
<bibRefCitation id="EF4F95E1A6E82CB5FB10FDA01516FDA6" author="Latreille, P. A." box="[1168,1395,582,604]" pageId="151" pageNumber="152" refId="ref150176" refString="Latreille, P. A. (1802 b) Histoire naturelle generale et particuliere des Crustaces et des insectes. Tome 3. Familles naturelles des genres. F. Dufart, Paris, xii + 467 pp." type="book" year="1802">Latreille, 1802b: 198</bibRefCitation>
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; by subsequent designation of 
<treatmentCitation id="0A7FCE01A6E82CB5FE54FD80121AFD81" author="Emery, C." box="[468,639,613,635]" page="42" pageId="151" pageNumber="152" year="1901">
<bibRefCitation id="EF4F95E1A6E82CB5FE54FD80121AFD81" author="Emery, C." box="[468,639,613,635]" pageId="151" pageNumber="152" pagination="32 - 54" refId="ref141230" refString="Emery, C. (1901) Notes sur les sous-familles des Dorylines et Ponerines (Famille des Formicides). Annales de la Societe Entomologique de Belgique, 45, 32 - 54." type="journal article" year="1901">Emery, 1901: 42</bibRefCitation>
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.
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</paragraph>
<paragraph id="8B61E810A6E82CB5FF17FD41112DFCD7" blockId="151.[151,1437,679,2037]" pageId="151" pageNumber="152">
<taxonomicName id="4CDE9393A6E82CB5FF17FD411153FD3A" ID-CoL="6BRJ" box="[151,310,679,704]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FF17FD411153FD3A" box="[151,310,679,704]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
</taxonomicName>
has been the focus of the greatest taxonomic confusion within 
<taxonomicName id="4CDE9393A6E82CB5FBB5FD4114CDFD3A" ID-CoL="K6Z" box="[1077,1192,679,704]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
, and was previously considered the senior synonym of numerous genera which are here considered distinct. 
<taxonomicName id="4CDE9393A6E82CB5FBE6FD2A1560FD1F" ID-CoL="6BRJ" box="[1126,1285,716,741]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FBE6FD2A1560FD1F" box="[1126,1285,716,741]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
</taxonomicName>
is in reality a small Neotropical genus (11 described species) that is closely related to 
<emphasis id="B9AA3402A6E82CB5FC60FD16140AFCFD" box="[992,1135,752,775]" italics="true" pageId="151" pageNumber="152">
<taxonomicName id="4CDE9393A6E82CB5FC60FD16140EFCFD" ID-CoL="45TH" box="[992,1131,752,775]" class="Insecta" family="Formicidae" genus="Dinoponera" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">Dinoponera</taxonomicName>
.
</emphasis>
Relatively little is known about its habits.
</paragraph>
</subSubSection>
<subSubSection id="C3C4BB9BA6E82CB5FF47FCD114A9F8E6" pageId="151" pageNumber="152" type="diagnosis">
<paragraph id="8B61E810A6E82CB5FF47FCD1110CFB42" blockId="151.[151,1437,679,2037]" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF47FCD11124FCAA" bold="true" box="[199,321,823,848]" pageId="151" pageNumber="152">Diagnosis.</emphasis>
<taxonomicName id="4CDE9393A6E82CB5FECAFCD1118CFCAA" box="[330,489,823,848]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FECAFCD1118CFCAA" box="[330,489,823,848]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
</taxonomicName>
workers are fairly generalized and lack any obvious autapomorphies, making their diagnosis more complicated than for most ponerine genera. They can most easily be identified by the following combination of characters: mandibles triangular, anterior clypeal margin without projecting teeth, metanotal groove at most present as a faint suture, propodeal spiracles slit-shaped, metapleural gland orifice with a posterior U-shaped cuticular lip, arolia not prominent, tarsal claws unarmed, petiole a thick block-like node, stridulitrum absent from pretergite of A4, and hypopygium with a row of stout spines on either side of the sting. 
<taxonomicName id="4CDE9393A6E82CB5FB7DFC0A15F9FBFF" box="[1277,1436,1004,1029]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FB7DFC0A15F9FBFF" box="[1277,1436,1004,1029]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
</taxonomicName>
is most likely to be confused with 
<taxonomicName id="4CDE9393A6E82CB5FDA4FBF612CEFBDD" box="[548,683,1040,1063]" class="Insecta" family="Formicidae" genus="Dinoponera" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FDA4FBF612CEFBDD" box="[548,683,1040,1063]" italics="true" pageId="151" pageNumber="152">Dinoponera</emphasis>
</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E82CB5FD3BFBF71352FBD2" box="[699,823,1041,1064]" class="Insecta" family="Formicidae" genus="Neoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FD3BFBF71352FBD2" box="[699,823,1041,1064]" italics="true" pageId="151" pageNumber="152">Neoponera</emphasis>
</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E82CB5FCC7FBF71390FBD2" box="[839,1013,1041,1064]" class="Insecta" family="Formicidae" genus="Ectomomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FCC7FBF71390FBD2" box="[839,1013,1041,1064]" italics="true" pageId="151" pageNumber="152">Ectomomyrmex</emphasis>
</taxonomicName>
, or 
<emphasis id="B9AA3402A6E82CB5FBA5FBE914ADFBD2" box="[1061,1224,1039,1064]" italics="true" pageId="151" pageNumber="152">
<taxonomicName id="4CDE9393A6E82CB5FBA5FBE914A1FBD2" baseAuthorityName="Brown" baseAuthorityYear="1973" box="[1061,1220,1039,1064]" class="Insecta" family="Formicidae" genus="Bothroponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">Bothroponera</taxonomicName>
,
</emphasis>
but 
<taxonomicName id="4CDE9393A6E82CB5FB7DFBE915F9FBD2" box="[1277,1436,1039,1064]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FB7DFBE915F9FBD2" box="[1277,1436,1039,1064]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
</taxonomicName>
differs from 
<taxonomicName id="4CDE9393A6E82CB5FEA9FBD311D5FBB6" box="[297,432,1077,1100]" class="Insecta" family="Formicidae" genus="Dinoponera" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FEA9FBD311D5FBB6" box="[297,432,1077,1100]" italics="true" pageId="151" pageNumber="152">Dinoponera</emphasis>
</taxonomicName>
in its smaller size, triangular mandibles, unarmed clypeal margin and tarsal claws, and block-like petiole; from 
<taxonomicName id="4CDE9393A6E82CB5FE26FBBF1246FB8A" box="[422,547,1113,1136]" class="Insecta" family="Formicidae" genus="Neoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FE26FBBF1246FB8A" box="[422,547,1113,1136]" italics="true" pageId="151" pageNumber="152">Neoponera</emphasis>
</taxonomicName>
in its lack of a stridulitrum on the pretergite of A4 and by its hypopygial spines; from 
<taxonomicName id="4CDE9393A6E82CB5FF54FB9B11E7FB6E" box="[212,386,1149,1172]" class="Insecta" family="Formicidae" genus="Ectomomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FF54FB9B11E7FB6E" box="[212,386,1149,1172]" italics="true" pageId="151" pageNumber="152">Ectomomyrmex</emphasis>
</taxonomicName>
in its complex metapleural gland orifice and hypopygial spines; and from 
<taxonomicName id="4CDE9393A6E82CB5FB42FB9A153BFB6F" baseAuthorityName="Brown" baseAuthorityYear="1973" box="[1218,1374,1148,1173]" class="Insecta" family="Formicidae" genus="Bothroponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FB42FB9A153BFB6F" box="[1218,1374,1148,1173]" italics="true" pageId="151" pageNumber="152">Bothroponera</emphasis>
</taxonomicName>
in its hypopygial spines.
</paragraph>
<paragraph id="8B61E810A6E82CB5FF47FB221489F9DA" blockId="151.[151,1437,679,2037]" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF47FB2211A3FB27" bold="true" box="[199,454,1220,1245]" pageId="151" pageNumber="152">Synoptic description.</emphasis>
<emphasis id="B9AA3402A6E82CB5FE50FB221243FB27" box="[464,550,1220,1245]" italics="true" pageId="151" pageNumber="152">Worker.</emphasis>
Medium to large (TL 
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) robust ants with the standard characters of 
<taxonomicName id="4CDE9393A6E82CB5FF17FB01109FFAFA" baseAuthorityName="Forel" baseAuthorityYear="1917" box="[151,250,1255,1280]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
. Mandibles triangular, sometimes with a faint basal groove. Anterior margin of clypeus convex and often medially emarginate. Frontal lobes moderately large. Eyes of moderate size and located anterior of head midline. Pronotum often with sharp lateral margins. Mesopleuron variable: fully, partially, or not at all divided by a transverse groove. Metanotal groove absent or at most present as a faint suture. Propodeum broad dorsally. Propodeal spiracles slit-shaped. Metapleural gland orifice with a posterior U-shaped cuticular lip and a lateral depression. Metatibial spur formula (1s, 1p). Petiole with a thick block-like node which widens posteriorly. Gaster with only a weak girdling constriction between pre- and postsclerites of A4. Hypopygium with a row of stout spines on either side of the sting. Head and body densely punctate to striate (rugoreticulate in at least one population of 
<taxonomicName id="4CDE9393A6E82CB5FEB4F9EF11FBF9DA" authorityName=": Longino" authorityYear="2013" box="[308,414,1543,1568]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E82CB5FEB4F9EF11FBF9DA" box="[308,414,1543,1568]" italics="true" pageId="151" pageNumber="152">P. harpax</emphasis>
</taxonomicName>
), with abundant pilosity and dense pubescence. Color dark brown to black.
</paragraph>
<paragraph id="8B61E810A6E82CB5FF47F9CA12C8F99D" blockId="151.[151,1437,679,2037]" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF47F9CA1172F9B9" box="[199,279,1580,1603]" italics="true" pageId="151" pageNumber="152">Queen.</emphasis>
Winged, with ocelli and the other modifications typical of ponerine queens, and slightly larger than the worker, but otherwise very similar to that caste.
</paragraph>
<paragraph id="8B61E810A6E82CB5FF47F9921401F977" blockId="151.[151,1437,679,2037]" box="[199,1124,1652,1677]" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF47F992116DF977" box="[199,264,1652,1677]" italics="true" pageId="151" pageNumber="152">Male.</emphasis>
<collectingRegion id="491A26F2A6E82CB5FE8EF9921153F977" box="[270,310,1652,1677]" country="South Sudan" name="Lakes" pageId="151" pageNumber="152">See</collectingRegion>
descriptions for individual species in 
<bibRefCitation id="EF4F95E1A6E82CB5FD61F9921310F977" author="Smith, F." box="[737,885,1652,1677]" pageId="151" pageNumber="152" refId="ref162573" refString="Smith, F. (1858) Catalogue of hymenopterous insects in the collection of the British Museum. Part VI. Formicidae. British Museum, London, 216 pp." type="book" year="1858">Smith (1858)</bibRefCitation>
and 
<bibRefCitation id="EF4F95E1A6E82CB5FC2CF9921405F977" author="Santschi, F." box="[940,1120,1652,1677]" pageId="151" pageNumber="152" pagination="81 - 103" refId="ref161277" refString="Santschi, F. (1921) Ponerinae, Dorylinae et quelques autres formicides neotropiques. Bulletin de la Societe Vaudoise des Sciences Naturelles, 54, 81 - 103." type="journal article" year="1921">Santschi (1921)</bibRefCitation>
.
</paragraph>
<paragraph id="8B61E810A6E82CB5FF47F97F1383F94A" blockId="151.[151,1437,679,2037]" box="[199,998,1687,1712]" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF47F97F1174F94A" box="[199,273,1689,1712]" italics="true" pageId="151" pageNumber="152">Larva.</emphasis>
Described for individual species by 
<bibRefCitation id="EF4F95E1A6E82CB5FD2BF9711384F94A" author="Wheeler, G. C. &amp; Wheeler, J." box="[683,993,1687,1712]" pageId="151" pageNumber="152" pagination="604 - 672" refId="ref166964" refString="Wheeler, G. C. &amp; Wheeler, J. (1952) The ant larvae of the subfamily Ponerinae - Part II. American Midland Naturalist, 48, 604 - 672. http: // dx. doi. org / 10.2307 / 2422200" type="journal article" year="1952">Wheeler &amp; Wheeler (1952)</bibRefCitation>
.
</paragraph>
<paragraph id="8B61E810A6E82CB5FF47F95A14A9F8E6" blockId="151.[151,1437,679,2037]" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF47F95A118BF92F" bold="true" box="[199,494,1724,1749]" pageId="151" pageNumber="152">Geographic distribution.</emphasis>
The range of 
<taxonomicName id="4CDE9393A6E82CB5FD0AF95A134CF92F" box="[650,809,1724,1749]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FD0AF95A134CF92F" box="[650,809,1724,1749]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
</taxonomicName>
extends from the southern 
<collectingCountry id="F3C9A880A6E82CB5FBD9F95A148BF92F" box="[1113,1262,1724,1749]" name="United States of America" pageId="151" pageNumber="152">United States</collectingCountry>
(
<collectingRegion id="491A26F2A6E82CB5FB7DF95A1508F92F" box="[1277,1389,1724,1749]" country="United States of America" name="Louisiana" pageId="151" pageNumber="152">Louisiana</collectingRegion>
and 
<collectingRegion id="491A26F2A6E82CB5FF17F93910BFF902" box="[151,218,1759,1784]" country="United States of America" name="Texas" pageId="151" pageNumber="152">Texas</collectingRegion>
) to northern 
<collectingCountry id="F3C9A880A6E82CB5FEEAF93911B9F902" box="[362,476,1759,1784]" name="Argentina" pageId="151" pageNumber="152">Argentina</collectingCountry>
, and includes some islands of the Caribbean (
<bibRefCitation id="EF4F95E1A6E82CB5FC5AF9391416F902" author="Kempf, W. W." box="[986,1139,1759,1784]" pageId="151" pageNumber="152" pagination="189 - 204" refId="ref149211" refString="Kempf, W. W. (1961) As formigas do genero Pachycondyla Fr. Smith no Brasil (Hymenoptera: Formicidae). Revista Brasileira de Entomologia, 10, 189 - 204." type="journal article" year="1961">Kempf, 1961</bibRefCitation>
). 
<taxonomicName id="4CDE9393A6E82CB5FB06F9071495F902" authorityName=": Longino" authorityYear="2013" box="[1158,1264,1759,1784]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E82CB5FB06F9071495F902" box="[1158,1264,1759,1784]" italics="true" pageId="151" pageNumber="152">P. harpax</emphasis>
</taxonomicName>
covers most of the range of the genus, but most other 
<taxonomicName id="4CDE9393A6E82CB5FDC5F8E2135EF8E7" box="[581,827,1796,1821]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B9AA3402A6E82CB5FDC5F8E21281F8E7" box="[581,740,1796,1821]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
species
</taxonomicName>
have a much more restricted range.
</paragraph>
</subSubSection>
<subSubSection id="C3C4BB9BA6E82CB8FF47F8C1122AFE0F" lastPageId="154" lastPageNumber="155" pageId="151" pageNumber="152" type="biology_ecology">
<paragraph id="8B61E810A6E82CBAFF47F8C1134BFEE7" blockId="151.[151,1437,679,2037]" lastBlockId="152.[151,1436,151,285]" lastPageId="152" lastPageNumber="153" pageId="151" pageNumber="152">
<emphasis id="B9AA3402A6E82CB5FF47F8C111B7F8BA" bold="true" box="[199,466,1831,1856]" pageId="151" pageNumber="152">Ecology and behavior.</emphasis>
Relatively little is known about the ecology and behavior of 
<taxonomicName id="4CDE9393A6E82CB5FB0FF8C11548F8BA" box="[1167,1325,1831,1856]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E82CB5FB0FF8C11548F8BA" box="[1167,1325,1831,1856]" italics="true" pageId="151" pageNumber="152">Pachycondyla</emphasis>
</taxonomicName>
. 
<bibRefCitation id="EF4F95E1A6E82CB5FABDF8C11084F89F" author="Longino, J. T." pageId="151" pageNumber="152" refId="ref152255" refString="Longino, J. T. (2013) Ants of Costa Rica. Available from: http: // academic. evergreen. edu / projects / ants / AntsofCostaRica. html (accessed 10 November 2013)" type="url" year="2013">Longino (2013)</bibRefCitation>
reports that 
<taxonomicName id="4CDE9393A6E82CB5FEEEF8AB11BCF89F" authorityName=": Longino" authorityYear="2013" box="[366,473,1868,1893]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E82CB5FEEEF8AB11BCF89F" box="[366,473,1868,1893]" italics="true" pageId="151" pageNumber="152">P. harpax</emphasis>
</taxonomicName>
and 
<taxonomicName id="4CDE9393A6E82CB5FD90F8AB12EAF89E" authorityName=": Ortius &amp; Lechner" authorityYear="1997" box="[528,655,1869,1892]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="impressa">
<emphasis id="B9AA3402A6E82CB5FD90F8AB12EAF89E" box="[528,655,1869,1892]" italics="true" pageId="151" pageNumber="152">P. impressa</emphasis>
</taxonomicName>
forage on the forest floor and are never observed foraging arboreally. They are presumably generalist predators and scavengers, though published accounts of their food preferences are scant. 
<bibRefCitation id="EF4F95E1A6E82CB5FF5FF87211F8F857" author="Wheeler, W. M." box="[223,413,1940,1965]" pageId="151" pageNumber="152" pagination="1 - 31" refId="ref167551" refString="Wheeler, W. M. (1900 b) A study of some Texan Ponerinae. Biological Bulletin, 11 (1), 1 - 31." type="journal article" year="1900">Wheeler (1900b)</bibRefCitation>
reported that 
<taxonomicName id="4CDE9393A6E82CB5FDBBF87312C3F857" authorityName=": Longino" authorityYear="2013" box="[571,678,1940,1965]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E82CB5FDBBF87312C3F857" box="[571,678,1940,1965]" italics="true" pageId="151" pageNumber="152">P. harpax</emphasis>
</taxonomicName>
workers in captivity would feed on egg yolk and sugar but ignored termites, 
<bibRefCitation id="EF4F95E1A6E82CB5FE84F85E11C3F82B" author="Overal, W. L." box="[260,422,1976,2001]" pageId="151" pageNumber="152" pagination="268 - 269" refId="ref158140" refString="Overal, W. L. (1987) Defensive chemical weaponry in the ant Pachycondyla harpax (Formicidae, Ponerinae). Journal of Entomological Science, 22 (3), 268 - 269." type="journal article" year="1987">Overal (1987)</bibRefCitation>
stated that 
<taxonomicName id="4CDE9393A6E82CB5FDB4F85F12C6F82B" authorityName=": Longino" authorityYear="2013" box="[564,675,1976,2001]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E82CB5FDB4F85F12C6F82B" box="[564,675,1976,2001]" italics="true" pageId="151" pageNumber="152">P. harpax</emphasis>
</taxonomicName>
eats soft-bodied insects (including termites) and myriapods, and 
<bibRefCitation id="EF4F95E1A6E82CB5FF17F83A11DDF80F" author="Garcia-Perez, J. A. &amp; Blanco-Pinon, A. &amp; Mercado-Hernandez, R. &amp; Badii, M." box="[151,440,2012,2037]" pageId="151" pageNumber="152" pagination="345 - 353" refId="ref143901" refString="Garcia-Perez, J. A., Blanco-Pinon, A., Mercado-Hernandez, R. &amp; Badii, M. (1997) El comportamiento depredador de Pachycondyla harpax Fabr. sobre Gnathamitermes tubiformans Buckley en condiciones de cautiverio. Southwestern Entomology, 22 (3), 345 - 353." type="journal article" year="1997">
Garcia-Pérez 
<emphasis id="B9AA3402A6E82CB5FEAFF83B1102F80F" box="[303,359,2012,2037]" italics="true" pageId="151" pageNumber="152">et al.</emphasis>
(1997)
</bibRefCitation>
observed 
<taxonomicName id="4CDE9393A6E82CB5FDAAF83B12F1F80F" authorityName=": Longino" authorityYear="2013" box="[554,660,2012,2037]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E82CB5FDAAF83B12F1F80F" box="[554,660,2012,2037]" italics="true" pageId="151" pageNumber="152">P. harpax</emphasis>
</taxonomicName>
preying predominantly on termites. 
<taxonomicName id="4CDE9393A6E82CB5FBABF83B14EAF80E" box="[1067,1167,2013,2036]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="151" pageNumber="152" phylum="Arthropoda" rank="species" species="striata">
<emphasis id="B9AA3402A6E82CB5FBABF83B14EAF80E" box="[1067,1167,2013,2036]" italics="true" pageId="151" pageNumber="152">P. striata</emphasis>
</taxonomicName>
will readily harvest and consume fruits and the arils of seeds from the forest floor, though it is unknown if this behavior occurs in other members of the genus (
<bibRefCitation id="EF4F95E1A6E72CBAFE1BFF5A12F2FF2F" author="Pizo, M. A. &amp; Oliveira, P. S." box="[411,663,188,213]" pageId="152" pageNumber="153" pagination="669 - 674" refId="ref160187" refString="Pizo, M. A. &amp; Oliveira, P. S. (1998) Interaction between ants and seeds of a nonmyrmecochorous neotropical tree, Cabralea canjerana (Meliaceae), in the Atlantic forest of southeast Brazil. American Journal of Botany, 85 (5), 669 - 674. http: // dx. doi. org / 10.2307 / 2446536" type="journal article" year="1998">Pizo &amp; Oliveira, 1998</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E72CBAFD22FF5A12BBFF2F" author="Pizo, M. A. &amp; Oliveira, P. S." box="[674,734,188,213]" pageId="152" pageNumber="153" pagination="37 - 52" refId="ref160254" refString="Pizo, M. A. &amp; Oliveira, P. S. (2001) Size and lipid content of nonmyrmecochorous diaspores: effects on the interaction with litter-foraging ants in the Atlantic rain forest of Brazil. Plant Ecology, 157, 37 - 52. http: // dx. doi. org / 10.1023 / a: 1013735305100" type="journal article" year="2001">2001</bibRefCitation>
; 
<bibRefCitation id="EF4F95E1A6E72CBAFD68FF5A139EFF2F" author="Passos, L. &amp; Oliveira, P. S." box="[744,1019,188,213]" pageId="152" pageNumber="153" pagination="517 - 528" refId="ref158277" refString="Passos, L. &amp; Oliveira, P. S. (2002) Ants affect the distribution and performance of seedlings of Clusia criuva, a primarily birddispersed rain forest tree. Journal of Ecology, 90 (3), 517 - 528. http: // dx. doi. org / 10.1046 / j. 1365 - 2745.2002.00687. x" type="journal article" year="2002">Passos &amp; Oliveira, 2002</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E72CBAFB86FF5A1424FF2F" author="Passos, L. &amp; Oliveira, P. S." box="[1030,1089,188,213]" pageId="152" pageNumber="153" pagination="261 - 270" refId="ref158341" refString="Passos, L. &amp; Oliveira, P. S. (2003) Interactions between ants, fruits and seeds in a restinga forest in south-eastern Brazil. Journal of Tropical Ecology, 19, 261 - 270. http: // dx. doi. org / 10.1017 / s 0266467403003298" type="journal article" year="2003">2003</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E72CBAFBCCFF5A14E2FF2F" author="Passos, L. &amp; Oliveira, P. S." box="[1100,1159,188,213]" pageId="152" pageNumber="153" pagination="376 - 382" refId="ref158394" refString="Passos, L. &amp; Oliveira, P. S. (2004) Interaction between ants and fruits of Guapira opposita (Nyctaginaceae) in a Brazilian sandy plain rainforest: ant effects on seeds and seedlings. Oecologia, 139, 376 - 382. http: // dx. doi. org / 10.1007 / s 00442 - 004 - 1531 - 5" type="journal article" year="2004">2004</bibRefCitation>
; 
<bibRefCitation id="EF4F95E1A6E72CBAFB12FF5A15F4FF2F" author="Raimundo, R. L. G. &amp; Guimaraes, P. R. &amp; Almeida-Neto, M. &amp; Pizo, M. A." box="[1170,1425,188,213]" pageId="152" pageNumber="153" pagination="261 - 270" refId="ref160548" refString="Raimundo, R. L. G., Guimaraes, P. R., Almeida-Neto, M. &amp; Pizo, M. A. (2004) The influence of fruit morphology and habitat structure on ant-seed interactions: a study with artificial fruits. Sociobiology, 44 (2), 261 - 270." type="journal article" year="2004">
Raimundo 
<emphasis id="B9AA3402A6E72CBAFA8EFF5B1523FF2F" box="[1294,1350,188,213]" italics="true" pageId="152" pageNumber="153">et al.</emphasis>
, 2004
</bibRefCitation>
). Nestmates are apparently recruited to food sources via tandem running (observed in 
<taxonomicName id="4CDE9393A6E72CBAFBD2FF0714D8FF02" authorityName=": Longino" authorityYear="2013" box="[1106,1213,223,248]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E72CBAFBD2FF0714D8FF02" box="[1106,1213,223,248]" italics="true" pageId="152" pageNumber="153">P. harpax</emphasis>
</taxonomicName>
and 
<taxonomicName id="4CDE9393A6E72CBAFB75FF071513FF0D" authorityName=": Ortius &amp; Lechner" authorityYear="1997" box="[1269,1398,224,248]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="impressa">
<emphasis id="B9AA3402A6E72CBAFB75FF071513FF0D" box="[1269,1398,224,248]" italics="true" pageId="152" pageNumber="153">P. impressa</emphasis>
</taxonomicName>
; S. Levings, pers. comm. cited in 
<bibRefCitation id="EF4F95E1A6E72CBAFE6AFEE21344FEE7" author="Holldobler, B. &amp; Wilson, E. O." box="[490,801,260,285]" pageId="152" pageNumber="153" refId="ref146902" refString="Holldobler, B. &amp; Wilson, E. O. (1990) The ants. Harvard University Press, Cambridge, Mass., xii + 732 pp." type="book" year="1990">Hölldobler &amp; Wilson, 1990</bibRefCitation>
).
</paragraph>
<caption id="DFA1B898A6E72CBAFF17F908145CF8D8" ID-DOI="http://doi.org/10.5281/zenodo.10120882" ID-Zenodo-Dep="10120882" httpUri="https://zenodo.org/record/10120882/files/figure.png" pageId="152" pageNumber="153" startId="152.[151,250,1774,1796]" targetBox="[151,1436,323,1749]" targetPageId="152">
<paragraph id="8B61E810A6E72CBAFF17F908145CF8D8" blockId="152.[151,1436,1774,1826]" pageId="152" pageNumber="153">
<emphasis id="B9AA3402A6E72CBAFF17F9081143F8F9" bold="true" box="[151,294,1774,1796]" pageId="152" pageNumber="153">FIGURE 31.</emphasis>
Worker caste of 
<taxonomicName id="4CDE9393A6E72CBAFE6DF908129CF8FE" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[493,761,1774,1796]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="crassinoda">
<emphasis id="B9AA3402A6E72CBAFE6DF908129CF8FE" box="[493,761,1774,1796]" italics="true" pageId="152" pageNumber="153">Pachycondyla crassinoda</emphasis>
</taxonomicName>
: lateral and dorsal view of body and full-face view of head (CASENT0249138, Ryan Perry and www.antweb.org); world distribution of 
<taxonomicName id="4CDE9393A6E72CBAFC24F8EA1456F8D8" box="[932,1075,1804,1826]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E72CBAFC24F8EA1456F8D8" box="[932,1075,1804,1826]" italics="true" pageId="152" pageNumber="153">Pachycondyla</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8B61E810A6E72CBBFF47F8AC11FFFE51" blockId="152.[151,1436,1866,2035]" lastBlockId="153.[151,1437,151,2013]" lastPageId="153" lastPageNumber="154" pageId="152" pageNumber="153">
When reported, nests are constructed in the ground (
<taxonomicName id="4CDE9393A6E72CBAFC92F8AD1319F899" authorityName=": Longino" authorityYear="2013" box="[786,892,1866,1891]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E72CBAFC92F8AD1319F899" box="[786,892,1866,1891]" italics="true" pageId="152" pageNumber="153">P. harpax</emphasis>
</taxonomicName>
and 
<taxonomicName id="4CDE9393A6E72CBAFC33F8AD1457F898" authorityName=": Ortius &amp; Lechner" authorityYear="1997" box="[947,1074,1867,1890]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="impressa">
<emphasis id="B9AA3402A6E72CBAFC33F8AD1457F898" box="[947,1074,1867,1890]" italics="true" pageId="152" pageNumber="153">P. impressa</emphasis>
</taxonomicName>
; 
<bibRefCitation id="EF4F95E1A6E72CBAFBC1F8AC1492F899" author="Wheeler, W. M." box="[1089,1271,1866,1891]" pageId="152" pageNumber="153" pagination="1 - 31" refId="ref167551" refString="Wheeler, W. M. (1900 b) A study of some Texan Ponerinae. Biological Bulletin, 11 (1), 1 - 31." type="journal article" year="1900">Wheeler, 1900b</bibRefCitation>
; 
<bibRefCitation id="EF4F95E1A6E72CBAFA82F8AC15F2F899" author="Overal, W. L." box="[1282,1431,1866,1891]" pageId="152" pageNumber="153" pagination="268 - 269" refId="ref158140" refString="Overal, W. L. (1987) Defensive chemical weaponry in the ant Pachycondyla harpax (Formicidae, Ponerinae). Journal of Entomological Science, 22 (3), 268 - 269." type="journal article" year="1987">Overal, 1987</bibRefCitation>
; 
<bibRefCitation id="EF4F95E1A6E72CBAFF17F8881125F87D" author="Longino, J. T." box="[151,320,1902,1927]" pageId="152" pageNumber="153" refId="ref152255" refString="Longino, J. T. (2013) Ants of Costa Rica. Available from: http: // academic. evergreen. edu / projects / ants / AntsofCostaRica. html (accessed 10 November 2013)" type="url" year="2013">Longino, 2013</bibRefCitation>
) or in soil collected in the crowns of palms (
<taxonomicName id="4CDE9393A6E72CBAFCCBF88913DDF87D" authorityName=": Longino" authorityYear="2013" box="[843,952,1902,1927]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E72CBAFCCBF88913DDF87D" box="[843,952,1902,1927]" italics="true" pageId="152" pageNumber="153">P. harpax</emphasis>
</taxonomicName>
; 
<bibRefCitation id="EF4F95E1A6E72CBAFC4AF8881407F87D" author="Overal, W. L." box="[970,1122,1902,1927]" pageId="152" pageNumber="153" pagination="268 - 269" refId="ref158140" refString="Overal, W. L. (1987) Defensive chemical weaponry in the ant Pachycondyla harpax (Formicidae, Ponerinae). Journal of Entomological Science, 22 (3), 268 - 269." type="journal article" year="1987">Overal, 1987</bibRefCitation>
). Mating occurs via typical nuptial flights (
<taxonomicName id="4CDE9393A6E72CBAFEC4F8751202F851" authority=": Longino, 2013" authorityName=": Longino" authorityYear="2013" box="[324,615,1938,1963]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E72CBAFEC4F87511CAF851" box="[324,431,1938,1963]" italics="true" pageId="152" pageNumber="153">P. harpax</emphasis>
: 
<bibRefCitation id="EF4F95E1A6E72CBAFE3EF8741202F851" author="Longino, J. T." box="[446,615,1938,1963]" pageId="152" pageNumber="153" refId="ref152255" refString="Longino, J. T. (2013) Ants of Costa Rica. Available from: http: // academic. evergreen. edu / projects / ants / AntsofCostaRica. html (accessed 10 November 2013)" type="url" year="2013">Longino, 2013</bibRefCitation>
</taxonomicName>
; 
<taxonomicName id="4CDE9393A6E72CBAFDF3F8751474F851" authority=": Ortius &amp; Lechner, 1997" authorityName=": Ortius &amp; Lechner" authorityYear="1997" box="[627,1041,1938,1963]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="impressa">
<emphasis id="B9AA3402A6E72CBAFDF3F8751296F850" box="[627,755,1939,1962]" italics="true" pageId="152" pageNumber="153">P. impressa</emphasis>
: 
<bibRefCitation id="EF4F95E1A6E72CBAFC82F8741474F851" author="Ortius, D. &amp; Lechner, K." box="[770,1041,1938,1963]" pageId="152" pageNumber="153" pagination="227 - 229" refId="ref157852" refString="Ortius, D. &amp; Lechner, K. (1997) Nuptial flight in two ponerine ants, Pachycondyla impressa and P. fauveli, overlooking Machu Picchu, Peru. Studies on Neotropical Fauna and Environment, 32, 227 - 229." type="journal article" year="1997">Ortius &amp; Lechner, 1997</bibRefCitation>
</taxonomicName>
). 
<bibRefCitation id="EF4F95E1A6E72CBAFBA5F8741481F851" author="Wheeler, W. M." box="[1061,1252,1938,1963]" pageId="152" pageNumber="153" pagination="1 - 31" refId="ref167551" refString="Wheeler, W. M. (1900 b) A study of some Texan Ponerinae. Biological Bulletin, 11 (1), 1 - 31." type="journal article" year="1900">Wheeler (1900b)</bibRefCitation>
reported on egg production by workers of 
<taxonomicName id="4CDE9393A6E72CBAFE3EF851124CF835" authorityName=": Longino" authorityYear="2013" box="[446,553,1974,1999]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E72CBAFE3EF851124CF835" box="[446,553,1974,1999]" italics="true" pageId="152" pageNumber="153">P. harpax</emphasis>
</taxonomicName>
, which he interpreted as being ergatoid queens but which are more likely just normal workers laying haploid eggs, as is common in 
<taxonomicName id="4CDE9393A6E72CBAFC9AF83C13EBF809" box="[794,910,2010,2035]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="152" pageNumber="153" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
. The mandibular, Dufour’s and venom gland secretions of 
<taxonomicName id="4CDE9393A6E62CBBFEAFFF7F11F0FF55" box="[303,405,152,176]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="striata">
<emphasis id="B9AA3402A6E62CBBFEAFFF7F11F0FF55" box="[303,405,152,176]" italics="true" pageId="153" pageNumber="154">P. striata</emphasis>
</taxonomicName>
were studied by 
<bibRefCitation id="EF4F95E1A6E62CBBFDDBFF7E1327FF4A" author="Morgan, E. D. &amp; do Nascimento, R. R. &amp; Keegans, S. J. &amp; Billen, J." box="[603,834,151,176]" pageId="153" pageNumber="154" pagination="1395 - 1409" refId="ref156388" refString="Morgan, E. D., do Nascimento, R. R., Keegans, S. J. &amp; Billen, J. (1999) Comparative study of mandibular gland secretions of workers of ponerine ants. Journal of Chemical Ecology, 25 (6), 1395 - 1409. http: // dx. doi. org / 10.1023 / a: 1020987028163" type="journal article" year="1999">
Morgan 
<emphasis id="B9AA3402A6E62CBBFD3CFF7F1290FF4A" box="[700,757,151,176]" italics="true" pageId="153" pageNumber="154">et al.</emphasis>
(1999
</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E62CBBFCCEFF7113EFFF4A" author="Morgan, E. D. &amp; Jungnickel, H. &amp; Keegans, S. J. &amp; do Nascimento, R. R. &amp; Billen, J. &amp; Gobin, B. &amp; Ito, F." box="[846,906,151,176]" pageId="153" pageNumber="154" pagination="95 - 114" refId="ref156304" refString="Morgan, E. D., Jungnickel, H., Keegans, S. J., do Nascimento, R. R., Billen, J., Gobin, B. &amp; Ito, F. (2003) Comparative survey of abdominal gland secretions of the ant subfamily Ponerinae. Journal of Chemical Ecology, 29 (1), 95 - 114. http: // dx. doi. org / 10.1023 / a: 1021928630441" type="journal article" year="2003">2003</bibRefCitation>
; the mandibular gland of this species was also studied by 
<bibRefCitation id="EF4F95E1A6E62CBBFE93FF5A1269FF2F" author="Tomotake, M. E. M. &amp; Mathias, M. I. C. &amp; Yabuki, A. T. &amp; Caetano, F. H." box="[275,524,188,213]" pageId="153" pageNumber="154" pagination="1 - 6" refId="ref164494" refString="Tomotake, M. E. M., Mathias, M. I. C., Yabuki, A. T. &amp; Caetano, F. H. (1992) Scanning electron microscopy of mandibular glands of workers and queens of the ants Pachycondyla striata (Hymenoptera: Ponerinae). Journal of Advanced Zoology, 13 (1 - 2), 1 - 6." type="journal article" year="1992">
Tomotake 
<emphasis id="B9AA3402A6E62CBBFE0AFF5B11A6FF2F" box="[394,451,188,213]" italics="true" pageId="153" pageNumber="154">et al.</emphasis>
, 1992
</bibRefCitation>
, and Mathias 
<emphasis id="B9AA3402A6E62CBBFD29FF5B1284FF2F" box="[681,737,188,213]" italics="true" pageId="153" pageNumber="154">et al.</emphasis>
, 1995), the ovaries and corpora allata of 
<taxonomicName id="4CDE9393A6E62CBBFB32FF5B1573FF2E" box="[1202,1302,189,212]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="striata">
<emphasis id="B9AA3402A6E62CBBFB32FF5B1573FF2E" box="[1202,1302,189,212]" italics="true" pageId="153" pageNumber="154">P. striata</emphasis>
</taxonomicName>
queens and workers were compared by 
<bibRefCitation id="EF4F95E1A6E62CBBFE51FF39128EFF02" author="Thiele, E. &amp; Mathias, M. I. C." box="[465,747,223,248]" pageId="153" pageNumber="154" pagination="51 - 64" refId="ref164232" refString="Thiele, E. &amp; Mathias, M. I. C. (1999) Morphology, ultramorphology and histology of ovaries of workers and mated queens of Pachycondyla striata ants (Hymenoptera: Ponerinae). Biocell, 23 (1), 51 - 64." type="journal article" year="1999">Thiele &amp; Mathias (1999)</bibRefCitation>
and 
<bibRefCitation id="EF4F95E1A6E62CBBFCA3FF391430FF02" author="Figueira, C. R. F. &amp; Mathias, M. I. C." box="[803,1109,223,248]" pageId="153" pageNumber="154" pagination="77 - 87" refId="ref142001" refString="Figueira, C. R. F. &amp; Mathias, M. I. C. (2002) Histological, histochemical and morphometric study of female corpora allata of Pachycondyla striata ants (Hymenoptera: Ponerinae). Sociobiology, 39 (1), 77 - 87." type="journal article" year="2002">Figueira &amp; Mathias (2002)</bibRefCitation>
, respectively, the fat body of 
<taxonomicName id="4CDE9393A6E62CBBFF17FEE31098FEE6" box="[151,253,261,284]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="striata">
<emphasis id="B9AA3402A6E62CBBFF17FEE31098FEE6" box="[151,253,261,284]" italics="true" pageId="153" pageNumber="154">P. striata</emphasis>
</taxonomicName>
queens was studied by 
<bibRefCitation id="EF4F95E1A6E62CBBFD8DFEE2134EFEE7" author="Thiele, E. &amp; Mathias, M. I. C." box="[525,811,260,285]" pageId="153" pageNumber="154" pagination="243 - 254" refId="ref164281" refString="Thiele, E. &amp; Mathias, M. I. C. (2003) Morphology, ultramorphology and morphometry of the fat body of virgin females and queens of the ants Pachycondyla striata (Hymenoptera: Formicidae). Sociobiology, 42 (2), 243 - 254." type="journal article" year="2003">Thiele &amp; Mathias (2003)</bibRefCitation>
, and the structure of the venom gland in 
<taxonomicName id="4CDE9393A6E62CBBFA82FEE3150DFEE6" box="[1282,1384,261,284]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="striata">
<emphasis id="B9AA3402A6E62CBBFA82FEE3150DFEE6" box="[1282,1384,261,284]" italics="true" pageId="153" pageNumber="154">P. striata</emphasis>
</taxonomicName>
was described by 
<bibRefCitation id="EF4F95E1A6E62CBBFEB1FEC1125EFEBA" author="Ortiz, G. &amp; Mathias, M. I. C." box="[305,571,295,320]" pageId="153" pageNumber="154" pagination="103 - 116" refId="ref157897" refString="Ortiz, G. &amp; Mathias, M. I. C. (2003) Morphohistological study of the venom gland in workers of the ant Pachycondyla striata F. Smith (Hymenoptera: Ponerinae). Sociobiology, 42 (1), 103 - 116." type="journal article" year="2003">Ortiz &amp; Mathias (2003</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E62CBBFDC8FEC112E0FEBA" author="Ortiz, G. &amp; Mathias, M. I. C." box="[584,645,295,320]" pageId="153" pageNumber="154" pagination="243 - 248" refId="ref157946" refString="Ortiz, G. &amp; Mathias, M. I. C. (2006) Venom gland of Pachycondyla striata worker ants (Hymenoptera: Ponerinae). Ultrastructural characterization. Micron, 37, 243 - 248. http: // dx. doi. org / 10.1016 / j. micron. 2005.10.008" type="journal article" year="2006">2006</bibRefCitation>
). 
<bibRefCitation id="EF4F95E1A6E62CBBFD1AFEC1135FFEBA" author="Overal, W. L." box="[666,826,295,320]" pageId="153" pageNumber="154" pagination="268 - 269" refId="ref158140" refString="Overal, W. L. (1987) Defensive chemical weaponry in the ant Pachycondyla harpax (Formicidae, Ponerinae). Journal of Entomological Science, 22 (3), 268 - 269." type="journal article" year="1987">Overal (1987)</bibRefCitation>
observed that 
<taxonomicName id="4CDE9393A6E62CBBFC65FECF1437FEBA" authorityName=": Longino" authorityYear="2013" box="[997,1106,295,320]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E62CBBFC65FECF1437FEBA" box="[997,1106,295,320]" italics="true" pageId="153" pageNumber="154">P. harpax</emphasis>
</taxonomicName>
produces a foamy defensive secretion from the tip of the abdomen, similar to the behavior exhibited by 
<taxonomicName id="4CDE9393A6E62CBBFC6EFEAA14DDFE9F" box="[1006,1208,332,357]" class="Insecta" family="Formicidae" genus="Pseudoneoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFC6EFEAA14DDFE9F" box="[1006,1208,332,357]" italics="true" pageId="153" pageNumber="154">Pseudoneoponera</emphasis>
</taxonomicName>
(see the description under that genus). 
<taxonomicName id="4CDE9393A6E62CBBFEE9FE9711B1FE72" authorityName=": Longino" authorityYear="2013" box="[361,468,367,392]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="harpax">
<emphasis id="B9AA3402A6E62CBBFEE9FE9711B1FE72" box="[361,468,367,392]" italics="true" pageId="153" pageNumber="154">P. harpax</emphasis>
</taxonomicName>
also injects venom from the sting, and 
<bibRefCitation id="EF4F95E1A6E62CBBFC14FE8914C7FE72" author="Orivel, J. &amp; Dejean, A." box="[916,1186,367,392]" pageId="153" pageNumber="154" pagination="195 - 201" refId="ref157656" refString="Orivel, J. &amp; Dejean, A. (2001) Comparative effect of the venoms of ants of the genus Pachycondyla (Hymenoptera: Ponerinae). Toxicon, 39 (2 - 3), 195 - 201. http: // dx. doi. org / 10.1016 / s 0041 - 0101 (00) 00113 - 6" type="journal article" year="2001">Orivel &amp; Déjean (2001)</bibRefCitation>
measured the toxicity of this species’ venom.
</paragraph>
<paragraph id="8B61E810A6E62CBBFF47FE5114E9FD5D" blockId="153.[151,1437,151,2013]" pageId="153" pageNumber="154">
<emphasis id="B9AA3402A6E62CBBFF47FE51128EFE2A" bold="true" box="[199,747,439,464]" pageId="153" pageNumber="154">Phylogenetic and taxonomic considerations.</emphasis>
The taxonomic status of 
<taxonomicName id="4CDE9393A6E62CBBFBB2FE5114B4FE2A" box="[1074,1233,439,464]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFBB2FE5114B4FE2A" box="[1074,1233,439,464]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
and its putative synonyms has been one of the central problems in ponerine systematics for many years. In fact, the taxonomic chaos represented by 
<taxonomicName id="4CDE9393A6E62CBBFE07FE191240FDE2" box="[391,549,511,536]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFE07FE191240FDE2" box="[391,549,511,536]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
was the initial motivation for this revision of the 
<taxonomicName id="4CDE9393A6E62CBBFBCDFE1914A4FDE2" box="[1101,1217,511,536]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
. From the very first description of 
<taxonomicName id="4CDE9393A6E62CBBFEBAFDC2121BFDC7" authority="(Smith, 1858)" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[314,638,548,573]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFEBAFDC211BCFDC7" box="[314,473,548,573]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
(
<bibRefCitation id="EF4F95E1A6E62CBBFE68FDC21213FDC7" author="Smith, F." box="[488,630,548,573]" pageId="153" pageNumber="154" refId="ref162573" refString="Smith, F. (1858) Catalogue of hymenopterous insects in the collection of the British Museum. Part VI. Formicidae. British Museum, London, 216 pp." type="book" year="1858">Smith, 1858</bibRefCitation>
)
</taxonomicName>
, and continuing until the present day, the true boundaries of this genus have been obscured by excessive synonymy and a lack of serious phylogenetic consideration. We will briefly review the taxonomic history of 
<taxonomicName id="4CDE9393A6E62CBBFDA5FD8A12A1FD7F" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[549,708,620,645]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFDA5FD8A12A1FD7F" box="[549,708,620,645]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
before discussing the results of 
<bibRefCitation id="EF4F95E1A6E62CBBFBD6FD8A157BFD7F" author="Schmidt, C." box="[1110,1310,620,645]" pageId="153" pageNumber="154" pagination="201 - 250" refId="ref161753" refString="Schmidt, C. (2013) Molecular phylogenetics of ponerine ants (Hymenoptera: Formicidae: Ponerinae). Zootaxa, 3647 (2), 201 - 250. http: // dx. doi. org / 10.11646 / zootaxa. 3647.2.1" type="journal article" year="2013">Schmidt’s (2013)</bibRefCitation>
molecular phylogenetic analyses and our morphological observations as they relate to 
<taxonomicName id="4CDE9393A6E62CBBFC66FD6914E1FD52" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[998,1156,655,680]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFC66FD6914E1FD52" box="[998,1156,655,680]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B61E810A6E62CBBFF47FD5213DEFC11" blockId="153.[151,1437,151,2013]" pageId="153" pageNumber="154">
<bibRefCitation id="EF4F95E1A6E62CBBFF47FD52113AFD37" author="Smith, F." box="[199,351,692,717]" pageId="153" pageNumber="154" refId="ref162573" refString="Smith, F. (1858) Catalogue of hymenopterous insects in the collection of the British Museum. Part VI. Formicidae. British Museum, London, 216 pp." type="book" year="1858">Smith (1858)</bibRefCitation>
erected 
<taxonomicName id="4CDE9393A6E62CBBFE46FD521201FD37" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[454,612,692,717]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFE46FD521201FD37" box="[454,612,692,717]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
based on a suite of morphological traits now shown to be of dubious phylogenetic value, including a subquadrate head, a convex anterior clypeal margin, large denticulate mandibles, clavate antennae (this character is mysterious, as the species that Smith included in 
<taxonomicName id="4CDE9393A6E62CBBFBDCFD1A149EFCEF" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[1116,1275,764,789]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFBDCFD1A149EFCEF" box="[1116,1275,764,789]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
have more or less filiform antennae), medium-sized eyes that are located anteriorly on the sides of the head, four-segmented labial and maxillary palps, pectinate tibial spurs (presumably he meant only the larger spur), a thick and wide petiolar node, and an elongate abdomen. While this long list of characters may seem to be a reasonable basis for defining a genus, most or all of these characters are probably plesiomorphic within the 
<taxonomicName id="4CDE9393A6E62CBBFB23FC6A1563FC5F" baseAuthorityName="Forel" baseAuthorityYear="1917" box="[1187,1286,908,933]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
. The species originally included in 
<taxonomicName id="4CDE9393A6E62CBBFE1FFC49125BFC32" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[415,574,943,968]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFE1FFC49125BFC32" box="[415,574,943,968]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
by Smith are now dispersed into several distinct genera: 
<taxonomicName id="4CDE9393A6E62CBBFB76FC4915F0FC32" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[1270,1429,943,968]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFB76FC4915F0FC32" box="[1270,1429,943,968]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFF17FC321104FC17" box="[151,353,980,1005]" class="Insecta" family="Formicidae" genus="Pseudoneoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFF17FC321104FC17" box="[151,353,980,1005]" italics="true" pageId="153" pageNumber="154">Pseudoneoponera</emphasis>
</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFEEFFC331278FC16" box="[367,541,981,1004]" class="Insecta" family="Formicidae" genus="Ectomomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFEEFFC331278FC16" box="[367,541,981,1004]" italics="true" pageId="153" pageNumber="154">Ectomomyrmex</emphasis>
</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFDABFC3212DCFC17" box="[555,697,980,1005]" class="Insecta" family="Formicidae" genus="Paltothyreus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFDABFC3212DCFC17" box="[555,697,980,1005]" italics="true" pageId="153" pageNumber="154">Paltothyreus</emphasis>
</taxonomicName>
, and even 
<taxonomicName id="4CDE9393A6E62CBBFCB2FC3213D1FC17" authorityName="Roger. These" authorityYear="1863" box="[818,948,980,1005]" class="Insecta" family="Formicidae" genus="Platythyrea" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFCB2FC3213D1FC17" box="[818,948,980,1005]" italics="true" pageId="153" pageNumber="154">Platythyrea</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B61E810A6E62CBBFF47FC111455FB3E" blockId="153.[151,1437,151,2013]" pageId="153" pageNumber="154">
Smith did not designate a 
<typeStatus id="546556B2A6E62CBBFE71FC1E1244FBEA" box="[497,545,1016,1040]" pageId="153" pageNumber="154">type</typeStatus>
species for 
<taxonomicName id="4CDE9393A6E62CBBFD2BFC11132FFBEA" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[683,842,1015,1040]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFD2BFC11132FFBEA" box="[683,842,1015,1040]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
, but 
<bibRefCitation id="EF4F95E1A6E62CBBFC05FC1E1447FBEA" author="Emery, C." box="[901,1058,1015,1040]" pageId="153" pageNumber="154" pagination="32 - 54" refId="ref141230" refString="Emery, C. (1901) Notes sur les sous-familles des Dorylines et Ponerines (Famille des Formicides). Annales de la Societe Entomologique de Belgique, 45, 32 - 54." type="journal article" year="1901">Emery (1901)</bibRefCitation>
later designated 
<taxonomicName id="4CDE9393A6E62CBBFB65FC1F1519FBEA" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[1253,1404,1015,1040]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="crassinoda">
<emphasis id="B9AA3402A6E62CBBFB65FC1F1519FBEA" box="[1253,1404,1015,1040]" italics="true" pageId="153" pageNumber="154">P. crassinoda</emphasis>
</taxonomicName>
as the 
<typeStatus id="546556B2A6E62CBBFF43FBFA1093FBCE" box="[195,246,1052,1076]" pageId="153" pageNumber="154">type</typeStatus>
, presumably because it was the first species listed by Smith under 
<taxonomicName id="4CDE9393A6E62CBBFB86FBFA14C1FBCF" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[1030,1188,1052,1077]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFB86FBFA14C1FBCF" box="[1030,1188,1052,1077]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
. 
<bibRefCitation id="EF4F95E1A6E62CBBFB36FBFA150CFBCF" author="Latreille, P. A." box="[1206,1385,1052,1077]" pageId="153" pageNumber="154" pagination="129 - 200" refId="ref150252" refString="Latreille, P. A. (1804) Tableau methodique des insectes. In: Societe de Naturalistes et d'Agriculteurs. Nouveau dictionnaire d'histoire naturelle. Tome 24. Deterville, Paris, pp. 129 - 200." type="book chapter" year="1804">Latreille (1804)</bibRefCitation>
had previously designated 
<taxonomicName id="4CDE9393A6E62CBBFE14FBA71249FBA2" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[404,556,1087,1112]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="crassinoda">
<emphasis id="B9AA3402A6E62CBBFE14FBA71249FBA2" box="[404,556,1087,1112]" italics="true" pageId="153" pageNumber="154">P. crassinoda</emphasis>
</taxonomicName>
as the 
<typeStatus id="546556B2A6E62CBBFDFDFBA612C8FBA2" box="[637,685,1088,1112]" pageId="153" pageNumber="154">type</typeStatus>
species of 
<taxonomicName id="4CDE9393A6E62CBBFCADFBA7131AFBA2" baseAuthorityName="Emery" baseAuthorityYear="1887" box="[813,895,1089,1112]" class="Insecta" family="Formicidae" genus="Ponera" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFCADFBA7131AFBA2" box="[813,895,1089,1112]" italics="true" pageId="153" pageNumber="154">Ponera</emphasis>
</taxonomicName>
, but 
<bibRefCitation id="EF4F95E1A6E62CBBFC3AFBD914E7FBA2" author="Westwood, J. O." box="[954,1154,1087,1112]" pageId="153" pageNumber="154" pagination="81 - 89" refId="ref166795" refString="Westwood, J. O. (1840) Observations on the genus Typhlopone, with descriptions of several exotic species of ants. Annals and Magazine of Natural History, 6, 81 - 89." type="journal article" year="1840">Westwood (1840)</bibRefCitation>
subsequently designated 
<taxonomicName id="4CDE9393A6E62CBBFF17FB83113FFB86" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[151,346,1125,1148]" class="Insecta" family="Formicidae" genus="Ponera" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="coarctata">
<emphasis id="B9AA3402A6E62CBBFF17FB83113FFB86" box="[151,346,1125,1148]" italics="true" pageId="153" pageNumber="154">Ponera coarctata</emphasis>
</taxonomicName>
as the 
<typeStatus id="546556B2A6E62CBBFE29FB8211BCFB86" box="[425,473,1124,1148]" pageId="153" pageNumber="154">type</typeStatus>
species of 
<taxonomicName id="4CDE9393A6E62CBBFDD7FB8312CCFB86" baseAuthorityName="Emery" baseAuthorityYear="1887" box="[599,681,1125,1148]" class="Insecta" family="Formicidae" genus="Ponera" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFDD7FB8312CCFB86" box="[599,681,1125,1148]" italics="true" pageId="153" pageNumber="154">Ponera</emphasis>
</taxonomicName>
. We continue to recognize the traditional application of the generic names 
<taxonomicName id="4CDE9393A6E62CBBFF66FB6F115DFB5A" baseAuthorityName="Emery" baseAuthorityYear="1887" box="[230,312,1161,1184]" class="Insecta" family="Formicidae" genus="Ponera" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFF66FB6F115DFB5A" box="[230,312,1161,1184]" italics="true" pageId="153" pageNumber="154">Ponera</emphasis>
</taxonomicName>
and 
<emphasis id="B9AA3402A6E62CBBFEF2FB61127DFB5A" box="[370,536,1159,1184]" italics="true" pageId="153" pageNumber="154">
<taxonomicName id="4CDE9393A6E62CBBFEF2FB611271FB5A" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[370,532,1159,1184]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Pachycondyla</taxonomicName>
,
</emphasis>
which has been nearly universal since Emery’s designation of 
<taxonomicName id="4CDE9393A6E62CBBFB64FB6F1519FB5A" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[1252,1404,1159,1184]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="species" species="crassinoda">
<emphasis id="B9AA3402A6E62CBBFB64FB6F1519FB5A" box="[1252,1404,1159,1184]" italics="true" pageId="153" pageNumber="154">P. crassinoda</emphasis>
</taxonomicName>
as the 
<typeStatus id="546556B2A6E62CBBFF40FB4A1095FB3E" box="[192,240,1196,1220]" pageId="153" pageNumber="154">type</typeStatus>
species of 
<taxonomicName id="4CDE9393A6E62CBBFEEDFB4A126EFB3F" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[365,523,1196,1221]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFEEDFB4A126EFB3F" box="[365,523,1196,1221]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
. 
<collectingRegion id="491A26F2A6E62CBBFD9AFB4A1227FB3F" box="[538,578,1196,1221]" country="South Sudan" name="Lakes" pageId="153" pageNumber="154">See</collectingRegion>
<bibRefCitation id="EF4F95E1A6E62CBBFDC9FB4A1281FB3F" author="Taylor, R. W." box="[585,740,1196,1221]" pageId="153" pageNumber="154" pagination="1 - 112" refId="ref163846" refString="Taylor, R. W. (1967) A monographic revision of the ant genus Ponera Latreille (Hymenoptera: Formicidae). Pacific Insects Monographs, 13, 1 - 112." type="journal article" year="1967">Taylor (1967)</bibRefCitation>
for more on this controversy.
</paragraph>
<paragraph id="8B61E810A6E62CBBFF47FB29115FFA3A" blockId="153.[151,1437,151,2013]" pageId="153" pageNumber="154">
After Smith’s original description, 
<taxonomicName id="4CDE9393A6E62CBBFDE5FB291361FB12" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[613,772,1231,1256]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFDE5FB291361FB12" box="[613,772,1231,1256]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
experienced more than a century of relative taxonomic stability, with the most significant change being 
<bibRefCitation id="EF4F95E1A6E62CBBFD33FB121304FAF7" author="Emery, C." box="[691,865,1268,1293]" pageId="153" pageNumber="154" pagination="32 - 54" refId="ref141230" refString="Emery, C. (1901) Notes sur les sous-familles des Dorylines et Ponerines (Famille des Formicides). Annales de la Societe Entomologique de Belgique, 45, 32 - 54." type="journal article" year="1901">Emery’s (1901)</bibRefCitation>
designation of 
<emphasis id="B9AA3402A6E62CBBFB8EFB121500FAF6" box="[1038,1381,1268,1293]" italics="true" pageId="153" pageNumber="154">
<taxonomicName id="4CDE9393A6E62CBBFB8EFB1214C8FAF7" baseAuthorityName="Brown" baseAuthorityYear="1973" box="[1038,1197,1268,1293]" class="Insecta" family="Formicidae" genus="Bothroponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Bothroponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFB37FB131500FAF6" box="[1207,1381,1269,1292]" class="Insecta" family="Formicidae" genus="Ectomomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Ectomomyrmex</taxonomicName>
</emphasis>
, and 
<taxonomicName id="4CDE9393A6E62CBBFF17FAF1115CFACA" box="[151,313,1303,1328]" class="Liliopsida" family="Orchidaceae" genus="Pseudoponera" higherTaxonomySource="CoL" kingdom="Plantae" order="Asparagales" pageId="153" pageNumber="154" phylum="Tracheophyta" rank="genus">
<emphasis id="B9AA3402A6E62CBBFF17FAF1115CFACA" box="[151,313,1303,1328]" italics="true" pageId="153" pageNumber="154">Pseudoponera</emphasis>
</taxonomicName>
as subgenera of 
<taxonomicName id="4CDE9393A6E62CBBFD85FAF112C6FACA" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[517,675,1303,1328]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFD85FAF112C6FACA" box="[517,675,1303,1328]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
. 
<bibRefCitation id="EF4F95E1A6E62CBBFD36FAFE1330FACA" author="Emery, C." box="[694,853,1303,1328]" pageId="153" pageNumber="154" pagination="1 - 125" refId="ref141346" refString="Emery, C. (1911) Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum, 118, 1 - 125." type="journal article" year="1911">Emery (1911)</bibRefCitation>
later removed 
<taxonomicName id="4CDE9393A6E62CBBFB8BFAF114C8FACA" box="[1035,1197,1303,1328]" class="Liliopsida" family="Orchidaceae" genus="Pseudoponera" higherTaxonomySource="CoL" kingdom="Plantae" order="Asparagales" pageId="153" pageNumber="154" phylum="Tracheophyta" rank="genus">
<emphasis id="B9AA3402A6E62CBBFB8BFAF114C8FACA" box="[1035,1197,1303,1328]" italics="true" pageId="153" pageNumber="154">Pseudoponera</emphasis>
</taxonomicName>
from the genus and rediagnosed 
<taxonomicName id="4CDE9393A6E62CBBFEA4FADA11A6FAAF" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[292,451,1340,1365]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFEA4FADA11A6FAAF" box="[292,451,1340,1365]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
by a presence of “subtriangular” toothed mandibles, convex anterior clypeal margin, the location of the eyes, a lack of preocular carinae, an obsolete metanotal groove, and a thick petiole. Subsequent authors variously treated 
<taxonomicName id="4CDE9393A6E62CBBFE35FA621234FA67" baseAuthorityName="Brown" baseAuthorityYear="1973" box="[437,593,1412,1437]" class="Insecta" family="Formicidae" genus="Bothroponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFE35FA621234FA67" box="[437,593,1412,1437]" italics="true" pageId="153" pageNumber="154">Bothroponera</emphasis>
</taxonomicName>
and 
<taxonomicName id="4CDE9393A6E62CBBFD0AFA63135DFA66" box="[650,824,1413,1436]" class="Insecta" family="Formicidae" genus="Ectomomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFD0AFA63135DFA66" box="[650,824,1413,1436]" italics="true" pageId="153" pageNumber="154">Ectomomyrmex</emphasis>
</taxonomicName>
as valid genera or as subgenera of 
<taxonomicName id="4CDE9393A6E62CBBFB48FA621502FA67" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[1224,1383,1412,1437]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFB48FA621502FA67" box="[1224,1383,1412,1437]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
(see 
<bibRefCitation id="EF4F95E1A6E62CBBFF17FA41114BFA3A" author="Bolton, B. &amp; Alpert, G. &amp; Ward, P. S. &amp; Naskrecki, P." box="[151,302,1447,1472]" pageId="153" pageNumber="154" refId="ref133459" refString="Bolton, B., Alpert, G., Ward, P. S. &amp; Naskrecki, P. (2006) Bolton's Catalogue of Ants of the World: 1758 - 2005. Harvard University Press, Cambridge, Mass., CD-ROM." type="book" year="2006">
<collectingRegion id="491A26F2A6E62CBBFF17FA411083FA3A" box="[151,230,1447,1472]" country="United Kingdom" name="Bolton" pageId="153" pageNumber="154">Bolton</collectingRegion>
, 2006
</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6E62CBBFF47FA2A1195F86F" blockId="153.[151,1437,151,2013]" pageId="153" pageNumber="154">
W. L. Brown worked for decades on a broad revision of 
<emphasis id="B9AA3402A6E62CBBFCC8FA2A138BFA1F" box="[840,1006,1484,1509]" italics="true" pageId="153" pageNumber="154">
<taxonomicName id="4CDE9393A6E62CBBFCC8FA2A138FFA1F" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[840,1002,1484,1509]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Pachycondyla</taxonomicName>
,
</emphasis>
and though he died before publishing his formal revision, his planned taxonomic changes became entrenched in the myrmecology community through various publications (principally 
<bibRefCitation id="EF4F95E1A6E62CBBFD91F9F212CDF9D7" author="Brown, W. L. Jr." box="[529,680,1556,1581]" pageId="153" pageNumber="154" pagination="161 - 185" refId="ref134603" refString="Brown, W. L. Jr. (1973) A comparison of the Hylean and Congo-West African rain forest ant faunas. In: Meggers, B. J., Ayensu, E. S. &amp; Duckworth, W. D. (Eds.), Tropical forest ecosystems in Africa and South America: a comparative review. Smithsonian Institution Press, Washington, D. C., pp. 161 - 185." type="book chapter" year="1973">Brown, 1973</bibRefCitation>
; 
<bibRefCitation id="EF4F95E1A6E62CBBFD36F9F2133AF9D7" author="Snelling, R. R." box="[694,863,1556,1581]" pageId="153" pageNumber="154" pagination="369 - 453" refId="ref162858" refString="Snelling, R. R. (1981) Systematics of social Hymenoptera. In: Hermann, H. R. (Ed.), Social insects. Vol. 2. Academic Press, New York, pp. 369 - 453." type="book chapter" year="1981">Snelling, 1981</bibRefCitation>
; 
<bibRefCitation id="EF4F95E1A6E62CBBFCEDF9F214CFF9D7" author="Holldobler, B. &amp; Wilson, E. O." box="[877,1194,1556,1581]" pageId="153" pageNumber="154" refId="ref146902" refString="Holldobler, B. &amp; Wilson, E. O. (1990) The ants. Harvard University Press, Cambridge, Mass., xii + 732 pp." type="book" year="1990">Hölldobler &amp; Wilson, 1990</bibRefCitation>
; 
<bibRefCitation id="EF4F95E1A6E62CBBFB38F9F21535F9D7" author="Bolton, B." box="[1208,1360,1556,1581]" pageId="153" pageNumber="154" refId="ref133375" refString="Bolton, B. (1994) Identification guide to the ant genera of the world. Harvard University Press, Cambridge, Mass., 222 pp." type="book" year="1994">
<collectingRegion id="491A26F2A6E62CBBFB38F9F21562F9D7" box="[1208,1287,1556,1581]" country="United Kingdom" name="Bolton" pageId="153" pageNumber="154">Bolton</collectingRegion>
, 1994
</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E62CBBFADDF9F215FDF9D7" author="Bolton, B." box="[1373,1432,1556,1581]" pageId="153" pageNumber="154" refId="ref133404" refString="Bolton, B. (1995) A new general catalogue of the ants of the world. Harvard University Press, Cambridge, Mass., 504 pp." type="book" year="1995">1995</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E62CBBFF17F9D110B7F9AA" author="Bolton, B." box="[151,210,1591,1616]" pageId="153" pageNumber="154" refId="ref133434" refString="Bolton, B. (2003) Synopsis and Classification of Formicidae. Memoirs of the American Entomological Institute, 71, 370 pp." type="book" year="2003">2003</bibRefCitation>
, 
<bibRefCitation id="EF4F95E1A6E62CBBFF5CF9D1117CF9AA" author="Bolton, B. &amp; Alpert, G. &amp; Ward, P. S. &amp; Naskrecki, P." box="[220,281,1591,1616]" pageId="153" pageNumber="154" refId="ref133459" refString="Bolton, B., Alpert, G., Ward, P. S. &amp; Naskrecki, P. (2006) Bolton's Catalogue of Ants of the World: 1758 - 2005. Harvard University Press, Cambridge, Mass., CD-ROM." type="book" year="2006">2006</bibRefCitation>
). These changes amounted to a broad synonymization of no fewer than 18 distinct generic names under 
<emphasis id="B9AA3402A6E62CBBFF17F9BA1159F98F" box="[151,316,1628,1653]" italics="true" pageId="153" pageNumber="154">
<taxonomicName id="4CDE9393A6E62CBBFF17F9BA115CF98F" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[151,313,1628,1653]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Pachycondyla</taxonomicName>
,
</emphasis>
without phylogenetic justification: 
<emphasis id="B9AA3402A6E62CBBFD61F9BA1388F947" italics="true" pageId="153" pageNumber="154">
<taxonomicName id="4CDE9393A6E62CBBFD61F9BA13E5F98F" baseAuthorityName="Brown" baseAuthorityYear="1973" box="[737,896,1628,1653]" class="Insecta" family="Formicidae" genus="Bothroponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Bothroponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFC10F9BA1456F98F" box="[912,1075,1628,1653]" class="Insecta" family="Formicidae" genus="Brachyponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Brachyponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFBC3F9BB1491F98E" box="[1091,1268,1629,1652]" class="Insecta" family="Formicidae" genus="Ectomomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Ectomomyrmex</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFA84F9BB15FDF98E" box="[1284,1432,1629,1652]" class="Insecta" family="Formicidae" genus="Neoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="150" phylum="Arthropoda" rank="subGenus" subGenus="Eumecopone">Eumecopone</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFF17F967116DF962" baseAuthorityName="Emery" baseAuthorityYear="1901" box="[151,264,1665,1688]" class="Insecta" family="Formicidae" genus="Euponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Euponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFE98F96611E3F96D" box="[280,390,1664,1687]" class="Insecta" family="Formicidae" genus="Hagensia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Hagensia</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFE15F9671242F962" box="[405,551,1665,1688]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Megaponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFDB7F96712A3F962" box="[567,710,1665,1688]" class="Insecta" family="Formicidae" genus="Mesoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Mesoponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFD55F9671331F962" box="[725,852,1665,1688]" class="Insecta" family="Formicidae" genus="Neoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Neoponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFCE4F999147EF962" box="[868,1051,1663,1688]" class="Insecta" family="Formicidae" genus="Ophthalmopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Ophthalmopone</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFBAAF99914DEF962" box="[1066,1211,1663,1688]" class="Insecta" family="Formicidae" genus="Paltothyreus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Paltothyreus</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFB4BF99915FDF962" box="[1227,1432,1663,1688]" class="Insecta" family="Formicidae" genus="Pseudoneoponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Pseudoneoponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFF17F942115EF947" box="[151,315,1700,1725]" class="Liliopsida" family="Orchidaceae" genus="Pseudoponera" higherTaxonomySource="CoL" kingdom="Plantae" order="Asparagales" pageId="153" pageNumber="154" phylum="Tracheophyta" rank="genus">Pseudoponera</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFED0F9421266F941" box="[336,515,1700,1723]" class="Insecta" family="Formicidae" genus="Syntermitopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Syntermitopone</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFD98F94312C2F946" box="[536,679,1701,1724]" class="Insecta" family="Formicidae" genus="Termitopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Termitopone</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFD3CF9421308F947" authorityName="Emery" authorityYear="1911" box="[700,877,1700,1725]" class="Insecta" family="Formicidae" genus="Trachymesopus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Trachymesopus</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E62CBBFC02F942138CF947" authorityName="Weber" authorityYear="1939" box="[898,1001,1700,1725]" class="Insecta" family="Formicidae" genus="Wadeura" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">Wadeura</taxonomicName>
,
</emphasis>
and 
<emphasis id="B9AA3402A6E62CBBFBB6F94214D4F947" box="[1078,1201,1700,1725]" italics="true" pageId="153" pageNumber="154">Xiphopelta</emphasis>
. Brown apparently included in 
<taxonomicName id="4CDE9393A6E62CBBFEA1F92111A5F91A" baseAuthorityName="Smith" baseAuthorityYear="1858" box="[289,448,1735,1760]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFEA1F92111A5F91A" box="[289,448,1735,1760]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
any ponerine species whose workers have triangular mandibles, eyes, two metatibial spurs, and the absence of any apomorphies extreme enough to justify a separate genus. These characters are almost certainly plesiomorphic, rendering them uninformative about the monophyly of the group as thus defined. Given its extensive morphological diversity and the apparent close relationship between some of its species and other recognized genera, 
<taxonomicName id="4CDE9393A6E62CBBFEF8F8B112ADF88A" authority="(sensu Brown)" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[376,712,1879,1904]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFEF8F8B11272F88A" box="[376,535,1879,1904]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
(
<emphasis id="B9AA3402A6E62CBBFDAAF8BF120DF88A" box="[554,616,1881,1904]" italics="true" pageId="153" pageNumber="154">sensu</emphasis>
Brown)
</taxonomicName>
was almost certainly bound to be non-monophyletic based on morphological evidence alone.
</paragraph>
<paragraph id="8B61E810A6E62CB8FF47F846122AFE0F" blockId="153.[151,1437,151,2013]" lastBlockId="154.[151,1436,151,501]" lastPageId="154" lastPageNumber="155" pageId="153" pageNumber="154">
<bibRefCitation id="EF4F95E1A6E62CBBFF47F84611EDF843" author="Schmidt, C." box="[199,392,1952,1977]" pageId="153" pageNumber="154" pagination="201 - 250" refId="ref161753" refString="Schmidt, C. (2013) Molecular phylogenetics of ponerine ants (Hymenoptera: Formicidae: Ponerinae). Zootaxa, 3647 (2), 201 - 250. http: // dx. doi. org / 10.11646 / zootaxa. 3647.2.1" type="journal article" year="2013">Schmidt's (2013)</bibRefCitation>
molecular phylogeny of the 
<taxonomicName id="4CDE9393A6E62CBBFD54F8461321F843" box="[724,836,1952,1977]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
confirms the vast non-monophyly of 
<taxonomicName id="4CDE9393A6E62CBBFB76F84615F1F843" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[1270,1428,1952,1977]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFB76F84615F1F843" box="[1270,1428,1952,1977]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
. 
<bibRefCitation id="EF4F95E1A6E62CBBFF17F8221122F827" author="Schmidt, C." box="[151,327,1988,2013]" pageId="153" pageNumber="154" pagination="201 - 250" refId="ref161753" refString="Schmidt, C. (2013) Molecular phylogenetics of ponerine ants (Hymenoptera: Formicidae: Ponerinae). Zootaxa, 3647 (2), 201 - 250. http: // dx. doi. org / 10.11646 / zootaxa. 3647.2.1" type="journal article" year="2013">Schmidt (2013)</bibRefCitation>
sampled representatives from nearly all junior synonyms of 
<taxonomicName id="4CDE9393A6E62CBBFC7CF82214FFF827" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[1020,1178,1988,2013]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="153" pageNumber="154" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E62CBBFC7CF82214FFF827" box="[1020,1178,1988,2013]" italics="true" pageId="153" pageNumber="154">Pachycondyla</emphasis>
</taxonomicName>
, and none of them are inferred to even be the sister group of true 
<emphasis id="B9AA3402A6E52CB8FDFAFF71137AFF4A" box="[634,799,151,176]" italics="true" pageId="154" pageNumber="155">
<taxonomicName id="4CDE9393A6E52CB8FDFAFF71137EFF4A" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[634,795,151,176]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">Pachycondyla</taxonomicName>
,
</emphasis>
which is represented in the phylogeny by 
<emphasis id="B9AA3402A6E52CB8FB7EFF7F1161FF2F" italics="true" pageId="154" pageNumber="155">
<taxonomicName id="4CDE9393A6E52CB8FB7EFF7F15FDFF4A" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[1278,1432,151,176]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="species" species="crassinoda">P. crassinoda</taxonomicName>
, 
<taxonomicName id="4CDE9393A6E52CB8FF17FF5B1161FF2F" authorityName=": Longino" authorityYear="2013" box="[151,260,188,213]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="species" species="harpax">P. harpax</taxonomicName>
</emphasis>
and 
<taxonomicName id="4CDE9393A6E52CB8FEC1FF5B11A6FF2E" authorityName=": Ortius &amp; Lechner" authorityYear="1997" box="[321,451,189,212]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="species" species="impressa">
<emphasis id="B9AA3402A6E52CB8FEC1FF5B11A6FF2E" box="[321,451,189,212]" italics="true" pageId="154" pageNumber="155">P. impressa</emphasis>
</taxonomicName>
. From morphology we are also confident that those junior synonyms which 
<bibRefCitation id="EF4F95E1A6E52CB8FABFFF5A1084FF02" author="Schmidt, C." pageId="154" pageNumber="155" pagination="201 - 250" refId="ref161753" refString="Schmidt, C. (2013) Molecular phylogenetics of ponerine ants (Hymenoptera: Formicidae: Ponerinae). Zootaxa, 3647 (2), 201 - 250. http: // dx. doi. org / 10.11646 / zootaxa. 3647.2.1" type="journal article" year="2013">Schmidt (2013)</bibRefCitation>
did not sample (
<taxonomicName id="4CDE9393A6E52CB8FE27FF39126EFF02" authorityName="Weber" authorityYear="1939" box="[423,523,223,248]" class="Insecta" family="Formicidae" genus="Wadeura" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E52CB8FE27FF39126EFF02" box="[423,523,223,248]" italics="true" pageId="154" pageNumber="155">Wadeura</emphasis>
</taxonomicName>
and 
<emphasis id="B9AA3402A6E52CB8FDC6FF3912A4FF02" box="[582,705,223,248]" italics="true" pageId="154" pageNumber="155">Xiphopelta</emphasis>
) are also not closely related to 
<taxonomicName id="4CDE9393A6E52CB8FBB1FF3914AAFF02" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[1073,1231,223,248]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E52CB8FBB1FF3914AAFF02" box="[1073,1231,223,248]" italics="true" pageId="154" pageNumber="155">Pachycondyla</emphasis>
</taxonomicName>
. The actual sister group of 
<taxonomicName id="4CDE9393A6E52CB8FF7DFEE211FEFEE7" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[253,411,260,285]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E52CB8FF7DFEE211FEFEE7" box="[253,411,260,285]" italics="true" pageId="154" pageNumber="155">Pachycondyla</emphasis>
</taxonomicName>
is 
<taxonomicName id="4CDE9393A6E52CB8FE3DFEE31221FEE6" box="[445,580,261,284]" class="Insecta" family="Formicidae" genus="Dinoponera" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E52CB8FE3DFEE31221FEE6" box="[445,580,261,284]" italics="true" pageId="154" pageNumber="155">Dinoponera</emphasis>
</taxonomicName>
, which ironically has never been considered its junior synonym. Their close relationship is supported by strong molecular and morphological evidence (see the discussion under 
<taxonomicName id="4CDE9393A6E52CB8FA94FECE15FEFEC5" box="[1300,1435,296,319]" class="Insecta" family="Formicidae" genus="Dinoponera" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E52CB8FA94FECE15FEFEC5" box="[1300,1435,296,319]" italics="true" pageId="154" pageNumber="155">Dinoponera</emphasis>
</taxonomicName>
for more on their morphological synapomorphies). True 
<taxonomicName id="4CDE9393A6E52CB8FC99FEAA13D2FE9F" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[793,951,332,357]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E52CB8FC99FEAA13D2FE9F" box="[793,951,332,357]" italics="true" pageId="154" pageNumber="155">Pachycondyla</emphasis>
</taxonomicName>
is a small Neotropical genus consisting of the 
<typeStatus id="546556B2A6E52CB8FF42FE961097FE72" box="[194,242,368,392]" pageId="154" pageNumber="155">type</typeStatus>
species 
<taxonomicName id="4CDE9393A6E52CB8FED5FE97118BFE72" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[341,494,367,392]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="species" species="crassinoda">
<emphasis id="B9AA3402A6E52CB8FED5FE97118BFE72" box="[341,494,367,392]" italics="true" pageId="154" pageNumber="155">P. crassinoda</emphasis>
</taxonomicName>
and its close relatives. 
<taxonomicName id="4CDE9393A6E52CB8FD7CFE8913FEFE72" baseAuthorityName="sensu Brown" baseAuthorityYear="1973" box="[764,923,367,392]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="154" pageNumber="155" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6E52CB8FD7CFE8913FEFE72" box="[764,923,367,392]" italics="true" pageId="154" pageNumber="155">Pachycondyla</emphasis>
</taxonomicName>
workers are united morphologically by their combination of triangular mandibles, unadorned anterior clypeal margin, medium-sized eyes, unarmed tarsal claws, block-like petiole, absence of a stridulitrum from the pretergite of A4, and a row of stout spines on the hypopygium on either side of the sting.
</paragraph>
</subSubSection>
</treatment>
</document>