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<mods:title id="F8A080D828ED7EE39A82C83716AA1010">The Higher Classification of the Ant Subfamily Ponerinae (Hymenoptera: Formicidae), with a Review of Ponerine Ecology and Behavior</mods:title>
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<treatment id="03775906A6182C48FF17F94214A7FAF7" ID-DOI="http://doi.org/10.5281/zenodo.5117488" ID-GBIF-Taxon="183709961" ID-Zenodo-Dep="5117488" LSID="urn:lsid:plazi:treatment:03775906A6182C48FF17F94214A7FAF7" httpUri="http://treatment.plazi.org/id/03775906A6182C48FF17F94214A7FAF7" lastPageId="106" lastPageNumber="107" pageId="103" pageNumber="104">
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<paragraph id="8B61E810A6182C45FF17F9421118F944" blockId="103.[151,381,1700,1759]" box="[151,381,1700,1726]" pageId="103" pageNumber="104">
<heading id="D0295F7CA6182C45FF17F9421118F944" bold="true" box="[151,381,1700,1726]" fontSize="11" level="2" pageId="103" pageNumber="104" reason="4">
<taxonomicName id="4CDE9393A6182C45FF17F9421118F944" authority="Mayr" authorityName="Mayr" box="[151,381,1700,1726]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6182C45FF17F9421118F944" bold="true" box="[151,381,1700,1726]" pageId="103" pageNumber="104">
<emphasis id="B9AA3402A6182C45FF17F9421155F944" bold="true" box="[151,304,1700,1726]" italics="true" pageId="103" pageNumber="104">Megaponera</emphasis>
Mayr
</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="C3C4BB9BA6182C45FF17F9201083F925" box="[151,230,1734,1759]" pageId="103" pageNumber="104" type="description">
<paragraph id="8B61E810A6182C45FF17F9201083F925" blockId="103.[151,381,1700,1759]" box="[151,230,1734,1759]" pageId="103" pageNumber="104">
<figureCitation id="13E5F495A6182C45FF17F9201083F925" box="[151,230,1734,1759]" captionStart="FIGURE 16" captionStartId="104.[151,250,1944,1966]" captionTargetBox="[151,1436,193,1911]" captionTargetId="figure-51@104.[151,1436,193,1923]" captionTargetPageId="104" captionText="FIGURE 16. Worker caste of Megaponera analis: lateral and dorsal view of body and full-face view of head (CASENT0249196, Estella Ortega and www.antweb.org); world distribution of Megaponera." figureDoi="http://doi.org/10.5281/zenodo.10120848" httpUri="https://zenodo.org/record/10120848/files/figure.png" pageId="103" pageNumber="104">Fig. 16</figureCitation>
</paragraph>
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<subSubSection id="C3C4BB9BA6182C45FF17F8E913AFF89B" pageId="103" pageNumber="104" type="reference_group">
<paragraph id="8B61E810A6182C45FF17F8E915F9F8DE" blockId="103.[151,1436,1806,1890]" box="[151,1436,1806,1828]" pageId="103" pageNumber="104">
<treatmentCitationGroup id="ABCECF3EA6182C45FF17F8E915F9F8DE" box="[151,1436,1806,1828]" pageId="103" pageNumber="104">
<treatmentCitation id="0A7FCE01A6182C45FF17F8E911A2F8DE" ID-DOI="http://doi.org/10.5281/zenodo.6295332" ID-Zenodo-Dep="6295332" LSID="urn:lsid:zoobank.org:act:1DD2BB63-DF69-4202-934F-E37F6075B162" author="Mayr, G." box="[151,455,1806,1828]" httpUri="http://treatment.plazi.org/id/F747F30B6EC8FB8E1C9AD10C1A856D78" page="714" pageId="103" pageNumber="104" year="1862">
<taxonomicName id="4CDE9393A6182C45FF17F8E911A2F8DE" authority="Mayr, 1862: 714" authorityName="Mayr" authorityPageNumber="714" authorityYear="1862" box="[151,455,1806,1828]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6182C45FF17F8E9117CF8DE" box="[151,281,1807,1828]" italics="true" pageId="103" pageNumber="104">Megaponera</emphasis>
<bibRefCitation id="EF4F95E1A6182C45FE9EF8E911A2F8DE" author="Mayr, G." box="[286,455,1806,1828]" pageId="103" pageNumber="104" pagination="649 - 776" refId="ref155085" refString="Mayr, G. (1862) Myrmecologische Studien. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 12, 649 - 776." type="journal article" year="1862">Mayr, 1862: 714</bibRefCitation>
</taxonomicName>
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(as genus in
<taxonomicName id="4CDE9393A6182C45FDCCF8E812D4F8DE" ID-CoL="K6Z" box="[588,689,1806,1828]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
[Poneridae]). Type-species:
<treatmentCitation id="0A7FCE01A6182C45FC54F8E91534F8DE" author="Fabricius, J. C." box="[980,1361,1806,1828]" page="354" pageId="103" pageNumber="104" year="1793">
<taxonomicName id="4CDE9393A6182C45FC54F8E91534F8DE" ID-CoL="6JGVR" authority="Fabricius, 1793: 354" authorityName="Fabricius" authorityPageNumber="354" authorityYear="1793" box="[980,1361,1806,1828]" class="Insecta" family="Formicidae" genus="Formica" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="species" species="foetens">
<emphasis id="B9AA3402A6182C45FC54F8E9141FF8DE" box="[980,1146,1806,1828]" italics="true" pageId="103" pageNumber="104">Formica foetens</emphasis>
<bibRefCitation id="EF4F95E1A6182C45FB00F8E81534F8DE" author="Fabricius, J. C." box="[1152,1361,1806,1828]" pageId="103" pageNumber="104" refId="ref141621" refString="Fabricius, J. C. (1793) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species, adjectis synonimis, locis observationibus, descriptionibus. Tome 2. C. G. Proft, Hafniae [= Copenhagen], 519 pp." type="book" year="1793">Fabricius, 1793: 354</bibRefCitation>
</taxonomicName>
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(junior
</treatmentCitationGroup>
</paragraph>
<paragraph id="8B61E810A6182C45FF47F8CB13AFF89B" blockId="103.[151,1436,1806,1890]" pageId="103" pageNumber="104">
primary homonym in
<taxonomicName id="4CDE9393A6182C45FE24F8C8119EF8B9" ID-CoL="4KS4" box="[420,507,1838,1859]" class="Insecta" family="Formicidae" genus="Formica" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6182C45FE24F8C8119EF8B9" box="[420,507,1838,1859]" italics="true" pageId="103" pageNumber="104">Formica</emphasis>
</taxonomicName>
, replaced by
<taxonomicName id="4CDE9393A6182C45FD04F8C81398F8B9" authority="Latreille, 1802 a: 282" authorityName="Latreille. The" authorityPageNumber="282" authorityYear="1802" box="[644,1021,1837,1859]" class="Insecta" family="Formicidae" genus="Formica" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="species" species="analis">
<emphasis id="B9AA3402A6182C45FD04F8C8137AF8B9" box="[644,799,1837,1859]" italics="true" pageId="103" pageNumber="104">Formica analis</emphasis>
<bibRefCitation id="EF4F95E1A6182C45FCA5F8CB1398F8B9" author="Latreille, P. A." box="[805,1021,1837,1859]" pageId="103" pageNumber="104" refId="ref150124" refString="Latreille, P. A. (1802 a) Histoire naturelle des fourmis, et recueil de memoires et d'observations sur les abeilles, les araignees, les faucheurs, et autres insectes. Impr. Crapelet (chez T. Barrois), Paris, xvi + 445 pp." type="book" year="1802">Latreille, 1802a: 282</bibRefCitation>
</taxonomicName>
); by monotypy.
<emphasis id="B9AA3402A6182C45FB20F8CB1564F8B9" bold="true" box="[1184,1281,1837,1859]" pageId="103" pageNumber="104">Gen. rev.</emphasis>
<taxonomicName id="4CDE9393A6182C45FF17F8AA1184F898" authority="Roger, 1863 b: 17" authorityName="Roger" authorityPageNumber="17" authorityYear="1863" box="[151,481,1868,1890]" class="Insecta" family="Formicidae" genus="Megaloponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6182C45FF17F8AA1149F898" box="[151,300,1868,1890]" italics="true" pageId="103" pageNumber="104">Megaloponera</emphasis>
<bibRefCitation id="EF4F95E1A6182C45FEB2F8AA1184F898" author="Roger, J." box="[306,481,1868,1890]" pageId="103" pageNumber="104" pagination="1 - 65" refId="ref160986" refString="Roger, J. (1863 b) Verzeichniss der Formiciden-Gattungen und Arten. Berliner Entomologische Zeitschrift, 7 (Beilage), 1 - 65." type="journal article" year="1863">Roger, 1863b: 17</bibRefCitation>
</taxonomicName>
(incorrect subsequent spelling of
<taxonomicName id="4CDE9393A6182C45FCB9F8AA13DFF89B" box="[825,954,1868,1889]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6182C45FCB9F8AA13DFF89B" box="[825,954,1868,1889]" italics="true" pageId="103" pageNumber="104">Megaponera</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="C3C4BB9BA6182C45FF17F86915E2F831" pageId="103" pageNumber="104" type="description">
<paragraph id="8B61E810A6182C45FF17F86915E2F831" blockId="103.[151,1436,1934,1995]" pageId="103" pageNumber="104">
<taxonomicName id="4CDE9393A6182C45FF17F8691143F85C" box="[151,294,1935,1958]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6182C45FF17F8691143F85C" box="[151,294,1935,1958]" italics="true" pageId="103" pageNumber="104">Megaponera</emphasis>
</taxonomicName>
is a monotypic genus (with an additional five subspecies) widespread in Sub-Saharan Africa. It is notable for its ergatoid queens, polymorphic workers, obligate group foraging, and specialized termite predation.
</paragraph>
</subSubSection>
<caption id="DFA1B898A6172C4AFF17F87E142CF837" ID-DOI="http://doi.org/10.5281/zenodo.10120848" ID-Zenodo-Dep="10120848" httpUri="https://zenodo.org/record/10120848/files/figure.png" pageId="104" pageNumber="105" startId="104.[151,250,1944,1966]" targetBox="[151,1436,193,1911]" targetPageId="104">
<paragraph id="8B61E810A6172C4AFF17F87E142CF837" blockId="104.[151,1436,1944,1997]" pageId="104" pageNumber="105">
<emphasis id="B9AA3402A6172C4AFF17F87E114FF857" bold="true" box="[151,298,1944,1966]" pageId="104" pageNumber="105">FIGURE 16.</emphasis>
Worker caste of
<taxonomicName id="4CDE9393A6172C4AFD80F87F12B5F854" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[512,720,1944,1966]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="species" species="analis">
<emphasis id="B9AA3402A6172C4AFD80F87F12B5F854" box="[512,720,1944,1966]" italics="true" pageId="104" pageNumber="105">Megaponera analis</emphasis>
</taxonomicName>
: lateral and dorsal view of body and full-face view of head (CASENT0249196, Estella Ortega and www.antweb.org); world distribution of
<taxonomicName id="4CDE9393A6172C4AFC42F85E1426F837" box="[962,1091,1976,1997]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6172C4AFC42F85E1426F837" box="[962,1091,1976,1997]" italics="true" pageId="104" pageNumber="105">Megaponera</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<subSubSection id="C3C4BB9BA6162C4BFF47FF71154DFC5F" pageId="105" pageNumber="106" type="diagnosis">
<paragraph id="8B61E810A6162C4BFF47FF7112E8FEBA" blockId="105.[151,1437,151,2013]" pageId="105" pageNumber="106">
<emphasis id="B9AA3402A6162C4BFF47FF711124FF4A" bold="true" box="[199,321,151,176]" pageId="105" pageNumber="106">Diagnosis.</emphasis>
Diagnostic morphological apomorphies of
<taxonomicName id="4CDE9393A6162C4BFCACFF7F13DEFF4A" box="[812,955,153,176]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFCACFF7F13DEFF4A" box="[812,955,153,176]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
workers include the presence of preocular carinae and size polymorphism, which do not occur in combination in any other ponerine. Preocular carinae occur in
<taxonomicName id="4CDE9393A6162C4BFF38FF391139FF02" box="[184,348,223,248]" class="Insecta" family="Formicidae" genus="Odontoponera" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFF38FF391139FF02" box="[184,348,223,248]" italics="true" pageId="105" pageNumber="106">Odontoponera</emphasis>
</taxonomicName>
and many
<taxonomicName id="4CDE9393A6162C4BFE63FF0712DAFF02" box="[483,703,223,248]" class="Insecta" family="Formicidae" genus="Neoponera" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B9AA3402A6162C4BFE63FF071205FF02" box="[483,608,225,248]" italics="true" pageId="105" pageNumber="106">Neoponera</emphasis>
species
</taxonomicName>
, but
<taxonomicName id="4CDE9393A6162C4BFD7DFF0713E9FF02" box="[765,908,225,248]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFD7DFF0713E9FF02" box="[765,908,225,248]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
lacks
<emphasis id="B9AA3402A6162C4BFC5CFF3914F5FF02" box="[988,1168,223,248]" italics="true" pageId="105" pageNumber="106">Odontoponeras</emphasis>
striate sculpturing and denticulate clypeus, and
<emphasis id="B9AA3402A6162C4BFE2AFEE21252FEE1" box="[426,567,260,283]" italics="true" pageId="105" pageNumber="106">Neoponeras</emphasis>
prominent white projecting arolia, simple tarsal claws and U-shaped cuticular lip posterior to the metapleural gland orifice.
</paragraph>
<paragraph id="8B61E810A6162C4BFF47FEAA11B8FD9A" blockId="105.[151,1437,151,2013]" pageId="105" pageNumber="106">
<emphasis id="B9AA3402A6162C4BFF47FEAA11ACFE9F" bold="true" box="[199,457,332,357]" pageId="105" pageNumber="106">Synoptic description.</emphasis>
<emphasis id="B9AA3402A6162C4BFE57FEAA1248FE9F" box="[471,557,332,357]" italics="true" pageId="105" pageNumber="106">Worker.</emphasis>
Large to very large (TL
<quantity id="4C2645F5A6162C4BFCE8FEAA13BFFE9F" box="[872,986,332,357]" metricMagnitude="-2" metricUnit="m" metricValue="1.35" metricValueMax="1.8" metricValueMin="0.9" pageId="105" pageNumber="106" unit="mm" value="13.5" valueMax="18.0" valueMin="9.0">918 mm</quantity>
) ants with the standard characters of
<taxonomicName id="4CDE9393A6162C4BFF17FE89109FFE72" baseAuthorityName="Forel" baseAuthorityYear="1917" box="[151,250,367,392]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
. Workers polymorphic, varying principally in size and pubescence. Mandibles triangular. Eyes large, located near head midline, with distinct preocular carinae. Frontal lobes widely separated anteriorly and appearing flattened in frontal view. Scapes flattened, with distinct anterior margins. Metanotal groove shallow. Propodeum moderately narrowed dorsally. Propodeal spiracles slit-shaped. Tarsal claws with a preapical tooth. Metatibial spur formula (1s, 1p). Petiole nodiform. Constriction between pre- and postsclerites of A4 indistinct. Stridulitrum present on pretergite of A4. Head and body finely punctate, with scattered pilosity and dense pubescence (sparse in minor workers). Color black.
</paragraph>
<paragraph id="8B61E810A6162C4BFF47FD8A118CFD37" blockId="105.[151,1437,151,2013]" pageId="105" pageNumber="106">
<emphasis id="B9AA3402A6162C4BFF47FD8A1172FD79" box="[199,279,620,643]" italics="true" pageId="105" pageNumber="106">Queen.</emphasis>
Wingless, similar to a major worker but larger (TL
<quantity id="4C2645F5A6162C4BFCDFFD8A13A2FD7F" box="[863,967,620,645]" metricMagnitude="-2" metricUnit="m" metricValue="1.85" pageId="105" pageNumber="106" unit="mm" value="18.5">18.5 mm</quantity>
), with more extensive sculpturing, denser pilosity, a deeper metanotal groove, an anteriorly slanting squamiform petiole, and a more expansive gaster. (Adapted from
<bibRefCitation id="EF4F95E1A6162C4BFEC4FD5211B8FD37" author="Arnold, G." box="[324,477,692,717]" pageId="105" pageNumber="106" pagination="1 - 159" refId="ref131610" refString="Arnold, G. (1915) A monograph of the Formicidae of South Africa. Part I. Ponerinae; Dorylinae. Annals of the South African Museum, 14, 1 - 159." type="journal article" year="1915">Arnold, 1915</bibRefCitation>
.)
</paragraph>
<paragraph id="8B61E810A6162C4BFF47FD311423FD0A" blockId="105.[151,1437,151,2013]" box="[199,1094,727,752]" pageId="105" pageNumber="106">
<emphasis id="B9AA3402A6162C4BFF47FD31116DFD0A" box="[199,264,727,752]" italics="true" pageId="105" pageNumber="106">Male.</emphasis>
<collectingRegion id="491A26F2A6162C4BFE8EFD311153FD0A" box="[270,310,727,752]" country="South Sudan" name="Lakes" pageId="105" pageNumber="106">See</collectingRegion>
descriptions in
<bibRefCitation id="EF4F95E1A6162C4BFE69FD3E12F3FD0A" author="Emery, C." box="[489,662,727,752]" pageId="105" pageNumber="106" pagination="596 - 605" refId="ref141094" refString="Emery, C. (1897 b) Formiche raccolte da Don Eugenio dei Principi Ruspoli, durante l'ultimo suo viaggio nelle regioni dei Somali e dei Galla. Annali del Museo Civico di Storia Naturale di Genova, 38, 596 - 605." type="journal article" year="1897">Emery (1897b)</bibRefCitation>
,
<bibRefCitation id="EF4F95E1A6162C4BFD21FD311321FD0A" author="Arnold, G." box="[673,836,727,752]" pageId="105" pageNumber="106" pagination="1 - 159" refId="ref131610" refString="Arnold, G. (1915) A monograph of the Formicidae of South Africa. Part I. Ponerinae; Dorylinae. Annals of the South African Museum, 14, 1 - 159." type="journal article" year="1915">Arnold (1915)</bibRefCitation>
, and
<bibRefCitation id="EF4F95E1A6162C4BFCFFFD311424FD0A" author="Wheeler, W. M." box="[895,1089,727,752]" pageId="105" pageNumber="106" pagination="631 - 710" refId="ref167817" refString="Wheeler, W. M. (1922 b) Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bulletin of the American Museum of Natural History, 45, 631 - 710." type="journal article" year="1922">Wheeler (1922b)</bibRefCitation>
.
</paragraph>
<paragraph id="8B61E810A6162C4BFF47FD1B128BFCEF" blockId="105.[151,1437,151,2013]" box="[199,750,764,789]" pageId="105" pageNumber="106">
<emphasis id="B9AA3402A6162C4BFF47FD1B1174FCEE" box="[199,273,765,788]" italics="true" pageId="105" pageNumber="106">Larva.</emphasis>
Described by
<bibRefCitation id="EF4F95E1A6162C4BFE33FD1A128CFCEF" author="Wheeler, G. C. &amp; Wheeler, J." box="[435,745,764,789]" pageId="105" pageNumber="106" 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="8B61E810A6162C4BFF47FCF9154DFC5F" blockId="105.[151,1437,151,2013]" pageId="105" pageNumber="106">
<emphasis id="B9AA3402A6162C4BFF47FCF9118AFCC2" bold="true" box="[199,495,799,824]" pageId="105" pageNumber="106">Geographic distribution.</emphasis>
<taxonomicName id="4CDE9393A6162C4BFE75FCC712E1FCC2" box="[501,644,801,824]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFE75FCC712E1FCC2" box="[501,644,801,824]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
<materialsCitation id="3BB6E24DA6162C4BFD0BFCF91546FC5F" ID-GBIF-Occurrence="3325973302" collectionCode="USNM" country="India" latitude="-30.0" location="Africa" pageId="105" pageNumber="106" specimenCount="1">
is widespread in tropical Sub-Saharan Africa.
<bibRefCitation id="EF4F95E1A6162C4BFB10FCF9152AFCC2" author="Wheeler, W. M." box="[1168,1359,799,824]" pageId="105" pageNumber="106" pagination="631 - 710" refId="ref167817" refString="Wheeler, W. M. (1922 b) Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bulletin of the American Museum of Natural History, 45, 631 - 710." type="journal article" year="1922">Wheeler (1922b)</bibRefCitation>
shows the range as encompassing the African continent from roughly
<geoCoordinate id="EEEA8ED7A6162C4BFCE1FCA213C1FCA1" box="[865,932,836,861]" degrees="10" direction="north" orientation="latitude" pageId="105" pageNumber="106" precision="55555" value="10.0">10° N</geoCoordinate>
latitude to
<geoCoordinate id="EEEA8ED7A6162C4BFBA6FCA21400FCA7" box="[1062,1125,836,861]" degrees="30" direction="south" orientation="latitude" pageId="105" pageNumber="106" precision="55555" value="-30.0">30° S</geoCoordinate>
latitude. We have seen
<specimenCount id="9DD82399A6162C4BFAF2FCA21169FC7A" pageId="105" pageNumber="106" type="generic">two specimens</specimenCount>
of
<taxonomicName id="4CDE9393A6162C4BFEB0FC8F11DAFC7A" box="[304,447,873,896]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFEB0FC8F11DAFC7A" box="[304,447,873,896]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
in the
<collectionCode id="EDCF70D5A6162C4BFD8BFC811239FC7A" box="[523,604,871,896]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="105" pageNumber="106" type="Museum">USNM</collectionCode>
labelled as coming from
<collectingCountry id="F3C9A880A6162C4BFCF7FC8113D5FC7A" box="[887,944,871,896]" name="India" pageId="105" pageNumber="106">India</collectingCountry>
(with no additional collection information), but this is almost certainly in error as we know of no other collections of this genus from outside
<location id="8E01BECBA6162C4BFB58FC6A1546FC5F" LSID="urn:lsid:plazi:treatment:03775906A6182C48FF17F94214A7FAF7:8E01BECBA6162C4BFB58FC6A1546FC5F" box="[1240,1315,908,933]" country="India" latitude="-30.0" name="Africa" pageId="105" pageNumber="106">Africa</location>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C3C4BB9BA6162C48FF47FC4914A7FAF7" lastPageId="106" lastPageNumber="107" pageId="105" pageNumber="106" type="biology_ecology">
<paragraph id="8B61E810A6162C4BFF47FC491248FBA2" blockId="105.[151,1437,151,2013]" pageId="105" pageNumber="106">
<emphasis id="B9AA3402A6162C4BFF47FC4911BAFC32" bold="true" box="[199,479,943,968]" pageId="105" pageNumber="106">Ecology and behavior.</emphasis>
<taxonomicName id="4CDE9393A6162C4BFE6EFC5712B5FC32" baseAuthorityName="Latreille" baseAuthorityYear="1802" box="[494,720,943,968]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="analis">
<emphasis id="B9AA3402A6162C4BFE6EFC5712B5FC32" box="[494,720,943,968]" italics="true" pageId="105" pageNumber="106">Megaponera analis</emphasis>
</taxonomicName>
has an unusual suite of ecological, behavioral, and social adaptations relative to most ponerines. The typical ponerine species has small colonies (usually fewer than 100 individuals), alate queens, and monomorphic workers which forage singly.
<taxonomicName id="4CDE9393A6162C4BFC64FC1F1416FBEA" box="[996,1139,1017,1040]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFC64FC1F1416FBEA" box="[996,1139,1017,1040]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
has diverged significantly from this basic pattern, having relatively large colonies, flightless queens, and polymorphic workers which are specialized mass raiders of termites.
</paragraph>
<paragraph id="8B61E810A6162C4BFF47FB8312CBFA1F" blockId="105.[151,1437,151,2013]" pageId="105" pageNumber="106">
<taxonomicName id="4CDE9393A6162C4BFF47FB831133FB86" box="[199,342,1125,1148]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFF47FB831133FB86" box="[199,342,1125,1148]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
nests in the ground, in deserted termite mounds, or even partially in rotting logs (
<bibRefCitation id="EF4F95E1A6162C4BFB5FFB8215E0FB87" author="Levieux, J." box="[1247,1413,1124,1149]" pageId="105" pageNumber="106" pagination="23 - 33" refId="ref150919" refString="Levieux, J. (1976) Etude de la structure du nid de quelques especes terricoles de fourmis tropicales. Annales de l'Universite d' Abidjan, sec. C, 12, 23 - 33." type="journal article" year="1976">Lèvieux, 1976</bibRefCitation>
B;
<bibRefCitation id="EF4F95E1A6162C4BFF17FB6111BBFB5A" author="Longhurst, C. &amp; Howse, P. E." box="[151,478,1159,1184]" pageId="105" pageNumber="106" pagination="204 - 215" refId="ref152031" refString="Longhurst, C. &amp; Howse, P. E. (1979 a) Foraging, recruitment and emigration in Megaponera-foetens (Fab) (Hymenoptera, Formicidae) from the Nigerian Guinea Savanna. Insectes Sociaux, 26 (3), 204 - 215. http: // dx. doi. org / 10.1007 / bf 02223798" type="journal article" year="1979">Longhurst &amp; Howse, 1979a</bibRefCitation>
), and has relatively large colonies by ponerine standards, with from 500 to 2,000 workers (
<bibRefCitation id="EF4F95E1A6162C4BFE82FB4A11CBFB3F" author="Peeters, C." box="[258,430,1196,1221]" pageId="105" pageNumber="106" pagination="1 - 15" refId="ref158848" refString="Peeters, C. (1991 b) Ergatoid queens and intercastes in ants - 2 distinct adult forms which look morphologically intermediate between workers and winged queens. Insectes Sociaux, 38 (1), 1 - 15. http: // dx. doi. org / 10.1007 / bf 01242708" type="journal article" year="1991">Peeters, 1991b</bibRefCitation>
; Hölldobler
<emphasis id="B9AA3402A6162C4BFDBFFB4B121CFB3F" box="[575,633,1196,1221]" italics="true" pageId="105" pageNumber="106">et al.</emphasis>
, 1994). The nests are host to a diverse assemblage of myrmecophiles (Arnold, reprinted in
<bibRefCitation id="EF4F95E1A6162C4BFE17FB291236FB12" author="Wheeler, W. M." box="[407,595,1231,1256]" pageId="105" pageNumber="106" pagination="631 - 710" refId="ref167817" refString="Wheeler, W. M. (1922 b) Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bulletin of the American Museum of Natural History, 45, 631 - 710." type="journal article" year="1922">Wheeler, 1922b</bibRefCitation>
). Colonies have a single highly fecund ergatoid queen, which in most respects resembles a large major worker but is endowed with a rich array of glands for chemical communication with the workers (
<bibRefCitation id="EF4F95E1A6162C4BFEE1FAF1126FFACA" author="Peeters, C." box="[353,522,1303,1328]" pageId="105" pageNumber="106" pagination="1 - 15" refId="ref158848" refString="Peeters, C. (1991 b) Ergatoid queens and intercastes in ants - 2 distinct adult forms which look morphologically intermediate between workers and winged queens. Insectes Sociaux, 38 (1), 1 - 15. http: // dx. doi. org / 10.1007 / bf 01242708" type="journal article" year="1991">Peeters, 1991b</bibRefCitation>
; Hölldobler
<emphasis id="B9AA3402A6162C4BFD15FAFF12A8FACA" box="[661,717,1303,1328]" italics="true" pageId="105" pageNumber="106">et al.</emphasis>
, 1994). Males locate virgin queens by following the recruitment trails of workers back to their nests (
<bibRefCitation id="EF4F95E1A6162C4BFDB2FADA1316FAAF" author="Longhurst, C. &amp; Howse P. E." box="[562,883,1340,1365]" pageId="105" pageNumber="106" pagination="85 - 91" refId="ref152092" refString="Longhurst, C. &amp; Howse P. E. (1979 b) Some aspects of the biology of the males of Megaponera-foetens (Fab) (Hymenoptera, Formicidae). Insectes Sociaux, 26 (2), 85 - 91. http: // dx. doi. org / 10.1007 / bf 02223798" type="journal article" year="1979">Longhurst &amp; Howse, 1979b</bibRefCitation>
). Workers retain spermathecae and ovarioles, but do not appear to mate and do not produce viable haploid eggs (Villet, 1990a;
<bibRefCitation id="EF4F95E1A6162C4BFC7DFAB91564FA82" author="Villet, M. H. &amp; Duncan, F. D." box="[1021,1281,1375,1400]" pageId="105" pageNumber="106" pagination="280 - 282" refId="ref165994" refString="Villet, M. H. &amp; Duncan, F. D. (1992) Reproductive abilities of orphaned workers of two ponerine ant species (Hymenoptera: Formicidae). Journal of the Entomological Society of Southern Africa, 55 (2), 280 - 282." type="journal article" year="1992">Villet &amp; Duncan, 1992</bibRefCitation>
). Workers are continuously polymorphic for size (
<bibRefCitation id="EF4F95E1A6162C4BFDA5FA621305FA67" author="Longhurst, C. &amp; Howse, P. E." box="[549,864,1412,1437]" pageId="105" pageNumber="106" pagination="204 - 215" refId="ref152031" refString="Longhurst, C. &amp; Howse, P. E. (1979 a) Foraging, recruitment and emigration in Megaponera-foetens (Fab) (Hymenoptera, Formicidae) from the Nigerian Guinea Savanna. Insectes Sociaux, 26 (3), 204 - 215. http: // dx. doi. org / 10.1007 / bf 02223798" type="journal article" year="1979">Longhurst &amp; Howse, 1979a</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6162C4BFCEBFA621459FA67" author="Crewe, R. M. &amp; Peeters, C. &amp; Villet, M." box="[875,1084,1412,1437]" pageId="105" pageNumber="106" pagination="247 - 248" refId="ref136107" refString="Crewe, R. M., Peeters, C. &amp; Villet, M. (1984) Frequency distribution of worker sizes in Megaponera foetens (Fabricius). South African Journal of Zoology, 19, 247 - 248." type="journal article" year="1984">
Crewe
<emphasis id="B9AA3402A6162C4BFC3BFA631396FA67" box="[955,1011,1412,1437]" italics="true" pageId="105" pageNumber="106">et al.</emphasis>
, 1984
</bibRefCitation>
), though they are often referred to as “majors”, “medias” and “minors”. They are known to emit a strong odor, to stridulate loudly when disturbed, and to have a very painful sting (
<bibRefCitation id="EF4F95E1A6162C4BFD87FA2A12C7FA1F" author="Arnold, G." box="[519,674,1484,1509]" pageId="105" pageNumber="106" pagination="1 - 159" refId="ref131610" refString="Arnold, G. (1915) A monograph of the Formicidae of South Africa. Part I. Ponerinae; Dorylinae. Annals of the South African Museum, 14, 1 - 159." type="journal article" year="1915">Arnold, 1915</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6162C4BFF47FA17140EF869" blockId="105.[151,1437,151,2013]" pageId="105" pageNumber="106">
<taxonomicName id="4CDE9393A6162C4BFF47FA171133F9F2" box="[199,342,1521,1544]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFF47FA171133F9F2" box="[199,342,1521,1544]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
are specialized mass raiders of termites (
<bibRefCitation id="EF4F95E1A6162C4BFCB5FA09138BF9F2" author="Wheeler, W. M." box="[821,1006,1519,1544]" pageId="105" pageNumber="106" pagination="631 - 710" refId="ref167817" refString="Wheeler, W. M. (1922 b) Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bulletin of the American Museum of Natural History, 45, 631 - 710." type="journal article" year="1922">Wheeler, 1922b</bibRefCitation>
,
<bibRefCitation id="EF4F95E1A6162C4BFC7CFA09145DF9F2" author="Wheeler, W. M." box="[1020,1080,1519,1544]" pageId="105" pageNumber="106" pagination="159 - 243" refId="ref168101" refString="Wheeler, W. M. (1936) Ecological relations of ponerine and other ants to termites. Proceedings of the American Academy of Arts and Sciences, 71, 159 - 243. http: // dx. doi. org / 10.2307 / 20023221" type="journal article" year="1936">1936</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6162C4BFBC6FA0914BCF9F2" author="Weber, N. A." box="[1094,1241,1519,1544]" pageId="105" pageNumber="106" pagination="197 - 204" refId="ref166770" refString="Weber, N. A. (1964) Termite prey of some African ants. Entomological News, 75, 197 - 204." type="journal article" year="1964">Weber, 1964</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6162C4BFB67FA0915F5F9F2" author="Levieux, J." box="[1255,1424,1519,1544]" pageId="105" pageNumber="106" pagination="117 - 126" refId="ref150872" refString="Levieux, J. (1966) Note preliminaire sur les colonnes de chasse de Megaponera foetens F. (Hymenoptera: Formicidae). Insectes Sociaux, 13, 117 - 126. http: // dx. doi. org / 10.1007 / bf 02223567" type="journal article" year="1966">Lévieux, 1966</bibRefCitation>
).
<bibRefCitation id="EF4F95E1A6162C4BFF17F9F21184F9D7" author="Longhurst, C. &amp; Howse, P. E." box="[151,481,1556,1581]" pageId="105" pageNumber="106" pagination="204 - 215" refId="ref152031" refString="Longhurst, C. &amp; Howse, P. E. (1979 a) Foraging, recruitment and emigration in Megaponera-foetens (Fab) (Hymenoptera, Formicidae) from the Nigerian Guinea Savanna. Insectes Sociaux, 26 (3), 204 - 215. http: // dx. doi. org / 10.1007 / bf 02223798" type="journal article" year="1979">Longhurst &amp; Howse (1979a)</bibRefCitation>
studied their foraging behavior in detail and described the sequence of events in a typical raid (see also the account by
<bibRefCitation id="EF4F95E1A6162C4BFDC6F9D112BEF9AA" author="Weber, N. A." box="[582,731,1591,1616]" pageId="105" pageNumber="106" pagination="197 - 204" refId="ref166770" refString="Weber, N. A. (1964) Termite prey of some African ants. Entomological News, 75, 197 - 204." type="journal article" year="1964">Weber, 1964</bibRefCitation>
). Raids begin when a solitary major worker locates foraging termites. This scout returns directly to its nest, laying a trail of poison gland-derived pheromones exuded through its sting (Longhurst
<emphasis id="B9AA3402A6162C4BFEFCF96711D3F962" box="[380,438,1663,1688]" italics="true" pageId="105" pageNumber="106">et al.</emphasis>
, 1979A), and recruits up to several hundred of its nestmates (
<bibRefCitation id="EF4F95E1A6162C4BFBF3F99915F4F962" author="Corbara, B. &amp; Dejean, A." box="[1139,1425,1663,1688]" pageId="105" pageNumber="106" pagination="465 - 483" refId="ref135918" refString="Corbara, B. &amp; Dejean, A. (2000) Adaptive behavioral flexibility of the ant Pachycondyla analis (Megaponera foetens) (Formicidae: Ponerinae) during prey capture. Sociobiology, 36, 465 - 483." type="journal article" year="2000">Corbara &amp; Déjean, 2000</bibRefCitation>
). They proceed as a column to the termites protected foraging tunnels, which the major workers tear open. Minor workers then haul out termites. When this process is complete, the major workers stack the termites in their mandibles and the ants return as a column to their nest.
<bibRefCitation id="EF4F95E1A6162C4BFC8AF90A144FF8FF" author="Corbara, B. &amp; Dejean, A." box="[778,1066,1772,1797]" pageId="105" pageNumber="106" pagination="465 - 483" refId="ref135918" refString="Corbara, B. &amp; Dejean, A. (2000) Adaptive behavioral flexibility of the ant Pachycondyla analis (Megaponera foetens) (Formicidae: Ponerinae) during prey capture. Sociobiology, 36, 465 - 483." type="journal article" year="2000">Corbara &amp; Déjean (2000)</bibRefCitation>
compared the behavior of minor and major workers during prey capture and found that though they are generally similar, major workers are more likely to attack termite soldiers than are minor workers.
<bibRefCitation id="EF4F95E1A6162C4BFCA2F8D21409F8B7" author="Longhurst, C. &amp; Howse, P. E." box="[802,1132,1844,1869]" pageId="105" pageNumber="106" pagination="204 - 215" refId="ref152031" refString="Longhurst, C. &amp; Howse, P. E. (1979 a) Foraging, recruitment and emigration in Megaponera-foetens (Fab) (Hymenoptera, Formicidae) from the Nigerian Guinea Savanna. Insectes Sociaux, 26 (3), 204 - 215. http: // dx. doi. org / 10.1007 / bf 02223798" type="journal article" year="1979">Longhurst &amp; Howse (1979a)</bibRefCitation>
reviewed the literature on
<taxonomicName id="4CDE9393A6162C4BFF17F8BF1143F88A" box="[151,294,1881,1904]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFF17F8BF1143F88A" box="[151,294,1881,1904]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
foraging behavior and found significant regional variation in how raids are conducted.
<bibRefCitation id="EF4F95E1A6162C4BFA81F8B115F9F88A" author="Taylor, B." box="[1281,1436,1879,1904]" pageId="105" pageNumber="106" refId="ref163811" refString="Taylor, B. (2008) The Ants of (sub-Saharan) Africa. Available from: http: // antbase. org / ants / africa / (accessed 1 October 2008)" type="url" year="2008">Taylor (2008)</bibRefCitation>
hints that this variation supports his hypothesis of multiple species within
<taxonomicName id="4CDE9393A6162C4BFC54F89B1406F86E" box="[980,1123,1917,1940]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFC54F89B1406F86E" box="[980,1123,1917,1940]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B61E810A6162C48FF47F84613DAFE51" blockId="105.[151,1437,151,2013]" lastBlockId="106.[151,1437,151,1293]" lastPageId="106" lastPageNumber="107" pageId="105" pageNumber="106">
<bibRefCitation id="EF4F95E1A6162C4BFF47F8461261F843" author="Holldobler, B. &amp; Wilson, E. O." box="[199,516,1952,1977]" pageId="105" pageNumber="106" 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>
; see also
<bibRefCitation id="EF4F95E1A6162C4BFDF3F846137CF843" author="Wilson, E. O." box="[627,793,1952,1977]" pageId="105" pageNumber="106" pagination="24 - 31" refId="ref168604" refString="Wilson, E. O. (1958 a) The beginnings of nomadic and group-predatory behavior in the ponerine ants. Evolution, 12 (1), 24 - 31." type="journal article" year="1958">Wilson, 1958a</bibRefCitation>
) hypothesized that the
<taxonomicName id="4CDE9393A6162C4BFBA3F84714D7F842" box="[1059,1202,1953,1976]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6162C4BFBA3F84714D7F842" box="[1059,1202,1953,1976]" italics="true" pageId="105" pageNumber="106">Megaponera</emphasis>
</taxonomicName>
style of foraging, in which scouts lead columns of workers on raids of other social insects, could be the first step in the evolution of true legionary behavior, such as that of the dorylines (see also
<bibRefCitation id="EF4F95E1A6152C48FCB3FF711456FF4A" author="Oster, G. F. &amp; Wilson, E. O." box="[819,1075,151,176]" pageId="106" pageNumber="107" refId="ref158002" refString="Oster, G. F. &amp; Wilson, E. O. (1978) Caste and Ecology in the Social Insects. Princeton University Press, Princeton, New Jersey, 352 pp." type="book" year="1978">
<collectingRegion id="491A26F2A6152C48FCB3FF71130AFF4A" box="[819,879,151,176]" country="Cameroon" name="East" pageId="106" pageNumber="107">Oster</collectingRegion>
&amp; Wilson, 1978
</bibRefCitation>
). Of the hallmarks of legionary behavior identified by
<bibRefCitation id="EF4F95E1A6152C48FE28FF5A1222FF2F" author="Brady, S. G." box="[424,583,188,213]" pageId="106" pageNumber="107" pagination="6575 - 6579" refId="ref134048" refString="Brady, S. G. (2003) Evolution of the army ant syndrome: The origin and long-term evolutionary stasis of a complex of behavioral and reproductive adaptations. Proceedings of the National Academy of Sciences (USA), 100 (11), 6575 - 6579. http: // dx. doi. org / 10.1073 / pnas. 1137809100" type="journal article" year="2003">Brady (2003)</bibRefCitation>
,
<taxonomicName id="4CDE9393A6152C48FDD9FF5B128DFF2E" box="[601,744,189,212]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FDD9FF5B128DFF2E" box="[601,744,189,212]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
exhibits obligate collective foraging and a weak form of nomadism, by which colonies frequently emigrate to new nest sites. This latter behavior was described in detail by Arnold (reprinted in
<bibRefCitation id="EF4F95E1A6152C48FE10FEE2122DFEE7" author="Wheeler, W. M." box="[400,584,260,285]" pageId="106" pageNumber="107" pagination="631 - 710" refId="ref167817" refString="Wheeler, W. M. (1922 b) Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bulletin of the American Museum of Natural History, 45, 631 - 710." type="journal article" year="1922">Wheeler, 1922b</bibRefCitation>
) and
<bibRefCitation id="EF4F95E1A6152C48FD12FEE2138DFEE7" author="Longhurst, C. &amp; Howse, P. E." box="[658,1000,260,285]" pageId="106" pageNumber="107" pagination="204 - 215" refId="ref152031" refString="Longhurst, C. &amp; Howse, P. E. (1979 a) Foraging, recruitment and emigration in Megaponera-foetens (Fab) (Hymenoptera, Formicidae) from the Nigerian Guinea Savanna. Insectes Sociaux, 26 (3), 204 - 215. http: // dx. doi. org / 10.1007 / bf 02223798" type="journal article" year="1979">Longhurst &amp; Howse (1979a)</bibRefCitation>
.
<bibRefCitation id="EF4F95E1A6152C48FC78FEE214D9FEE7" author="Wheeler, W. M." box="[1016,1212,260,285]" pageId="106" pageNumber="107" pagination="631 - 710" refId="ref167817" refString="Wheeler, W. M. (1922 b) Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bulletin of the American Museum of Natural History, 45, 631 - 710." type="journal article" year="1922">Wheeler (1922b)</bibRefCitation>
also discussed the adaptations of
<taxonomicName id="4CDE9393A6152C48FEC3FECF11B7FEBA" box="[323,466,297,320]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FEC3FECF11B7FEBA" box="[323,466,297,320]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
brood to emigration. The reasons for these frequent emigrations are unknown, but
<bibRefCitation id="EF4F95E1A6152C48FF17FEAA1186FE9F" author="Longhurst, C. &amp; Howse, P. E." box="[151,483,332,357]" pageId="106" pageNumber="107" pagination="204 - 215" refId="ref152031" refString="Longhurst, C. &amp; Howse, P. E. (1979 a) Foraging, recruitment and emigration in Megaponera-foetens (Fab) (Hymenoptera, Formicidae) from the Nigerian Guinea Savanna. Insectes Sociaux, 26 (3), 204 - 215. http: // dx. doi. org / 10.1007 / bf 02223798" type="journal article" year="1979">Longhurst &amp; Howse (1979a)</bibRefCitation>
argue that they are not likely caused by prey shortages and may be a response to predation by
<taxonomicName id="4CDE9393A6152C48FEADFE8911E2FE72" authorityName="Fabricius" authorityYear="1793" box="[301,391,367,392]" class="Insecta" family="Formicidae" genus="Dorylus" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FEADFE8911E2FE72" box="[301,391,367,392]" italics="true" pageId="106" pageNumber="107">Dorylus</emphasis>
</taxonomicName>
driver ants. In an interesting observation,
<bibRefCitation id="EF4F95E1A6152C48FCECFE891438FE72" author="Beck, J. &amp; Kunz, B. K." box="[876,1117,367,392]" pageId="106" pageNumber="107" pagination="27 - 28" refId="ref132554" refString="Beck, J. &amp; Kunz, B. K. (2007) Cooperative self-defence: Matabele ants (Pachycondyla analis) against African driver ants (Dorylus sp.; Hymenoptera: Formicidae). Myrmecological News, 10, 27 - 28." type="journal article" year="2007">Beck &amp; Kunz (2007)</bibRefCitation>
found cooperative altruistic defensive actions among
<taxonomicName id="4CDE9393A6152C48FE33FE731227FE56" box="[435,578,405,428]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FE33FE731227FE56" box="[435,578,405,428]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
workers under attack by
<taxonomicName id="4CDE9393A6152C48FCDDFE7213D2FE57" authorityName="Fabricius" authorityYear="1793" box="[861,951,404,429]" class="Insecta" family="Formicidae" genus="Dorylus" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FCDDFE7213D2FE57" box="[861,951,404,429]" italics="true" pageId="106" pageNumber="107">Dorylus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B61E810A6152C48FF47FE511141FD9A" blockId="106.[151,1437,151,1293]" pageId="106" pageNumber="107">
<emphasis id="B9AA3402A6152C48FF47FE5112B5FE2A" bold="true" box="[199,720,439,464]" pageId="106" pageNumber="107">Phylogenetic and taxonomic considerations.</emphasis>
The genus
<taxonomicName id="4CDE9393A6152C48FCD0FE5F13BAFE2A" box="[848,991,441,464]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FCD0FE5F13BAFE2A" box="[848,991,441,464]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
was erected by
<bibRefCitation id="EF4F95E1A6152C48FB11FE5E157AFE2A" author="Mayr, G." box="[1169,1311,439,464]" pageId="106" pageNumber="107" pagination="649 - 776" refId="ref155085" refString="Mayr, G. (1862) Myrmecologische Studien. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 12, 649 - 776." type="journal article" year="1862">Mayr (1862)</bibRefCitation>
to hold the single species
<taxonomicName id="4CDE9393A6152C48FEB9FE3B12E1FE0F" authority="Latreille. The" authorityName="Latreille. The" authorityYear="1802" box="[313,644,476,501]" class="Insecta" family="Formicidae" genus="Formica" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="analis">
<emphasis id="B9AA3402A6152C48FEB9FE3B1181FE0F" box="[313,484,476,501]" italics="true" pageId="106" pageNumber="107">Formica analis</emphasis>
Latreille. The
</taxonomicName>
original specific epithet
<emphasis id="B9AA3402A6152C48FC1AFE3A146CFE0F" box="[922,1033,476,501]" italics="true" pageId="106" pageNumber="107">
<taxonomicName id="4CDE9393A6152C48FC22FE3A1464FE0F" authorityName="Fabricius" authorityYear="1793" box="[930,1025,476,501]" class="Insecta" family="Formicidae" genus="Formica" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="foetens">foetens</taxonomicName>
</emphasis>
was a junior primary homonym but has been used incorrectly in much of the extensive literature on this species.
<taxonomicName id="4CDE9393A6152C48FB9DFDE714C9FDE2" box="[1053,1196,513,536]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FB9DFDE714C9FDE2" box="[1053,1196,513,536]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
experienced relative taxonomic stability until W. L. Brown (in
<bibRefCitation id="EF4F95E1A6152C48FDEEFDC21364FDC7" author="Bolton, B." box="[622,769,548,573]" pageId="106" pageNumber="107" 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="491A26F2A6152C48FDEEFDC212D8FDC7" box="[622,701,548,573]" country="United Kingdom" name="Bolton" pageId="106" pageNumber="107">Bolton</collectingRegion>
, 1994
</bibRefCitation>
) synonymized it under
<taxonomicName id="4CDE9393A6152C48FB89FDC214CDFDC7" baseAuthorityName="Emery" baseAuthorityYear="1900" box="[1033,1192,548,573]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FB89FDC214CDFDC7" box="[1033,1192,548,573]" italics="true" pageId="106" pageNumber="107">Pachycondyla</emphasis>
</taxonomicName>
without phylogenetic justification.
</paragraph>
<paragraph id="8B61E810A6152C48FF47FD8A1309FCC2" blockId="106.[151,1437,151,1293]" pageId="106" pageNumber="107">
We are reviving
<taxonomicName id="4CDE9393A6152C48FE11FD8B1245FD7E" box="[401,544,621,644]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FE11FD8B1245FD7E" box="[401,544,621,644]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
to full genus status based on both morphological and molecular evidence.
<bibRefCitation id="EF4F95E1A6152C48FF17FD691138FD52" author="Schmidt, C." box="[151,349,655,680]" pageId="106" pageNumber="107" 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="4CDE9393A6152C48FD41FD691354FD52" box="[705,817,655,680]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
places
<taxonomicName id="4CDE9393A6152C48FC12FD771444FD52" box="[914,1057,657,680]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FC12FD771444FD52" box="[914,1057,657,680]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
with strong support within the
<taxonomicName id="4CDE9393A6152C48FF17FD521124FD37" baseAuthorityName="Brown" baseAuthorityYear="1978" box="[151,321,692,717]" class="Insecta" family="Formicidae" genus="Odontomachus" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FF17FD521124FD37" box="[151,321,692,717]" italics="true" pageId="106" pageNumber="107">Odontomachus</emphasis>
</taxonomicName>
group as sister to
<taxonomicName id="4CDE9393A6152C48FD8DFD5212A4FD37" box="[525,705,692,717]" class="Insecta" family="Formicidae" genus="Ophthalmopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FD8DFD5212A4FD37" box="[525,705,692,717]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
</taxonomicName>
, and not at all close to
<taxonomicName id="4CDE9393A6152C48FC45FD521401FD37" baseAuthorityName="Emery" baseAuthorityYear="1900" box="[965,1124,692,717]" class="Insecta" family="Formicidae" genus="Pachycondyla" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FC45FD521401FD37" box="[965,1124,692,717]" italics="true" pageId="106" pageNumber="107">Pachycondyla</emphasis>
</taxonomicName>
. A sister group relationship with
<taxonomicName id="4CDE9393A6152C48FF50FD3111E0FD0A" box="[208,389,727,752]" class="Insecta" family="Formicidae" genus="Ophthalmopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FF50FD3111E0FD0A" box="[208,389,727,752]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
</taxonomicName>
is also strongly supported by morphology. Workers of the two genera are remarkably similar, differing significantly only in
<emphasis id="B9AA3402A6152C48FD81FD1A12C4FCE9" box="[513,673,764,787]" italics="true" pageId="106" pageNumber="107">Megaponeras</emphasis>
preocular carinae and size polymorphism and
<emphasis id="B9AA3402A6152C48FB57FD1A15F9FCEF" box="[1239,1436,764,789]" italics="true" pageId="106" pageNumber="107">Ophthalmopones</emphasis>
slender build, huge posteriorly set eyes, and armed hypopygium.
</paragraph>
<paragraph id="8B61E810A6152C48FF47FCA21426FB87" blockId="106.[151,1437,151,1293]" pageId="106" pageNumber="107">
These genera also share derived ecological and behavioral attributes. Both are specialist termite predators, with
<taxonomicName id="4CDE9393A6152C48FF17FC8F1143FC7A" box="[151,294,873,896]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FF17FC8F1143FC7A" box="[151,294,873,896]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
and some
<taxonomicName id="4CDE9393A6152C48FE27FC8112D3FC7A" box="[423,694,871,896]" class="Insecta" family="Formicidae" genus="Ophthalmopone" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B9AA3402A6152C48FE27FC811239FC7A" box="[423,604,871,896]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
species
</taxonomicName>
raiding termites in large groups. It is possible that the common ancestor of these genera was a mass raider of termites, with a reversal to solitary foraging being favored in some
<taxonomicName id="4CDE9393A6152C48FF17FC4911D6FC32" box="[151,435,943,968]" class="Insecta" family="Formicidae" genus="Ophthalmopone" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B9AA3402A6152C48FF17FC491129FC32" box="[151,332,943,968]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
species.
</taxonomicName>
Alternatively, mass raiding may have evolved independently in
<taxonomicName id="4CDE9393A6152C48FB54FC571506FC32" box="[1236,1379,945,968]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FB54FC571506FC32" box="[1236,1379,945,968]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
and
<taxonomicName id="4CDE9393A6152C48FF17FC32112EFC17" box="[151,331,980,1005]" class="Insecta" family="Formicidae" genus="Ophthalmopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FF17FC32112EFC17" box="[151,331,980,1005]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
</taxonomicName>
. Both genera also lack a winged queen caste, though reproduction in
<taxonomicName id="4CDE9393A6152C48FBD5FC331481FC16" box="[1109,1252,981,1004]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FBD5FC331481FC16" box="[1109,1252,981,1004]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
is performed by ergatoid queens and in
<taxonomicName id="4CDE9393A6152C48FE2FFC111201FBEA" box="[431,612,1015,1040]" class="Insecta" family="Formicidae" genus="Ophthalmopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FE2FFC111201FBEA" box="[431,612,1015,1040]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
</taxonomicName>
it is performed by gamergate workers. The reproductive caste in the ancestor of these genera could conceivably have been either ergatoid or gamergate (or neither). The close relationship of these taxa make them excellent models for studying the selective forces driving the evolution of alternative reproductive strategies in ants, as well as the evolution of mass foraging.
</paragraph>
<paragraph id="8B61E810A6152C48FF47FB6114A7FAF7" blockId="106.[151,1437,151,1293]" pageId="106" pageNumber="107">
While
<taxonomicName id="4CDE9393A6152C48FE99FB6F11CDFB5A" box="[281,424,1161,1184]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FE99FB6F11CDFB5A" box="[281,424,1161,1184]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
and
<taxonomicName id="4CDE9393A6152C48FE6BFB6112FAFB5A" box="[491,671,1159,1184]" class="Insecta" family="Formicidae" genus="Ophthalmopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FE6BFB6112FAFB5A" box="[491,671,1159,1184]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
</taxonomicName>
could arguably be synonymized, the age of their divergence is consistent with that of other ponerine genera (
<bibRefCitation id="EF4F95E1A6152C48FD1EFB4A1320FB3F" author="Schmidt, C." box="[670,837,1196,1221]" pageId="106" pageNumber="107" 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>
) and we feel that their morphological and behavioral distinctiveness warrant separate generic status. The sister group of
<taxonomicName id="4CDE9393A6152C48FC4FFB37143BFB12" box="[975,1118,1233,1256]" class="Insecta" family="Formicidae" genus="Megaponera" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FC4FFB37143BFB12" box="[975,1118,1233,1256]" italics="true" pageId="106" pageNumber="107">Megaponera</emphasis>
</taxonomicName>
+
<taxonomicName id="4CDE9393A6152C48FB0CFB291524FB12" box="[1164,1345,1231,1256]" class="Insecta" family="Formicidae" genus="Ophthalmopone" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FB0CFB291524FB12" box="[1164,1345,1231,1256]" italics="true" pageId="106" pageNumber="107">Ophthalmopone</emphasis>
</taxonomicName>
is still unknown, but a close relationship with
<taxonomicName id="4CDE9393A6152C48FDD0FB1312D9FAF6" box="[592,700,1269,1292]" class="Insecta" family="Formicidae" genus="Hagensia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6152C48FDD0FB1312D9FAF6" box="[592,700,1269,1292]" italics="true" pageId="106" pageNumber="107">Hagensia</emphasis>
</taxonomicName>
is plausible (see discussion under that genus).
</paragraph>
</subSubSection>
</treatment>
</document>