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<document id="8AE6E78D0C9EEEF23AEA87BB5435A009" ID-CLB-Dataset="5350" ID-DOI="10.11646/zootaxa.3817.1.1" ID-GBIF-Dataset="d66f1b27-5891-4fa5-96e0-f75cb3ec2445" ID-ISSN="1175-5326" ID-Zenodo-Dep="10086256" ID-ZooBank="A3C10B34-7698-4C4D-94E5-DCF70B475603" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1623272878632" checkinUser="felipe" docAuthor="Schmidt, C. A. &amp; Shattuck, S. O." docDate="2014" docId="03775906A64B2C1AFF17F88D13C7FBCF" docLanguage="en" docName="zt03817p242.pdf.imf" docOrigin="Zootaxa 3817 (1)" docStyle="DocumentStyle:5EBBA59367AD13919D70D935FA04F6A3.14:Zootaxa.2013-.monograph" docStyleId="5EBBA59367AD13919D70D935FA04F6A3" docStyleName="Zootaxa.2013-.monograph" docStyleVersion="14" docTitle="Harpegnathos Jerdon" docType="treatment" docVersion="9" lastPageNumber="57" masterDocId="FF4E217EA67F2C22FF80FFE61065FFFA" masterDocTitle="The Higher Classification of the Ant Subfamily Ponerinae (Hymenoptera: Formicidae), with a Review of Ponerine Ecology and Behavior" masterLastPageNumber="242" masterPageNumber="1" pageNumber="53" updateTime="1699920028738" updateUser="ExternalLinkService">
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<mods:title id="B16A99C930858EEC182D8A8EE605C6FA">The Higher Classification of the Ant Subfamily Ponerinae (Hymenoptera: Formicidae), with a Review of Ponerine Ecology and Behavior</mods:title>
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<mods:namePart id="03DDCCECE6EEDF5B0F8A26E487957111">Schmidt, C. A.</mods:namePart>
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<treatment id="03775906A64B2C1AFF17F88D13C7FBCF" ID-DOI="http://doi.org/10.5281/zenodo.5117440" ID-GBIF-Taxon="183709985" ID-Zenodo-Dep="5117440" LSID="urn:lsid:plazi:treatment:03775906A64B2C1AFF17F88D13C7FBCF" httpUri="http://treatment.plazi.org/id/03775906A64B2C1AFF17F88D13C7FBCF" lastPageId="56" lastPageNumber="57" pageId="52" pageNumber="53">
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<paragraph id="8B61E810A64B2C16FF17F88D11C6F87F" blockId="52.[151,419,1899,1958]" box="[151,419,1899,1925]" pageId="52" pageNumber="53">
<heading id="D0295F7CA64B2C16FF17F88D11C6F87F" bold="true" box="[151,419,1899,1925]" fontSize="11" level="2" pageId="52" pageNumber="53" reason="4">
<taxonomicName id="4CDE9393A64B2C16FF17F88D11C6F87F" ID-CoL="6363H" ID-ENA="604375" authority="Jerdon" authorityName="Jerdon" box="[151,419,1899,1925]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="52" pageNumber="53" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A64B2C16FF17F88D11C6F87F" bold="true" box="[151,419,1899,1925]" pageId="52" pageNumber="53">
<emphasis id="B9AA3402A64B2C16FF17F88D1121F87F" bold="true" box="[151,324,1899,1925]" italics="true" pageId="52" pageNumber="53">Harpegnathos</emphasis>
Jerdon
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3C4BB9BA64B2C16FF17F86B10BDF85C" box="[151,216,1933,1958]" pageId="52" pageNumber="53" type="description">
<paragraph id="8B61E810A64B2C16FF17F86B10BDF85C" blockId="52.[151,419,1899,1958]" box="[151,216,1933,1958]" pageId="52" pageNumber="53">
<figureCitation id="13E5F495A64B2C16FF17F86B10BDF85C" box="[151,216,1933,1958]" captionStart="FIGURE 4" captionStartId="53.[151,250,1896,1918]" captionTargetBox="[151,1436,357,1862]" captionTargetId="figure-118@53.[151,1436,357,1874]" captionTargetPageId="53" captionText="FIGURE 4. Worker caste of Harpegnathos venator: lateral and dorsal view of body and full-face view of head (Philippines, G.Alpert, MCZC); world distribution of Harpegnathos." figureDoi="http://doi.org/10.5281/zenodo.10120820" httpUri="https://zenodo.org/record/10120820/files/figure.png" pageId="52" pageNumber="53">Fig. 4</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="C3C4BB9BA64B2C17FF17F8331549FECA" lastPageId="53" lastPageNumber="54" pageId="52" pageNumber="53" type="reference_group">
<paragraph id="8B61E810A64B2C17FF17F8331409FF36" blockId="52.[151,1321,2005,2027]" lastBlockId="53.[151,1437,152,204]" lastPageId="53" lastPageNumber="54" pageId="52" pageNumber="53">
<treatmentCitationGroup id="ABCECF3EA64B2C17FF17F8331409FF36" lastPageId="53" lastPageNumber="54" pageId="52" pageNumber="53">
<treatmentCitation id="0A7FCE01A64B2C16FF17F8331186F811" ID-DOI="http://doi.org/10.5281/zenodo.6297900" ID-Zenodo-Dep="6297900" LSID="urn:lsid:zoobank.org:act:2CC97E31-567E-4A7A-80E6-3E5CB4AF240E" author="Jerdon, T. C." box="[151,483,2005,2027]" httpUri="http://treatment.plazi.org/id/9B580F8D2A3E8B47552305C4DA36EAD6" page="116" pageId="52" pageNumber="53" year="1851">
<taxonomicName id="4CDE9393A64B2C16FF17F8331186F811" ID-CoL="6363H" authority="Jerdon, 1851: 116" authorityName="Jerdon" authorityPageNumber="116" authorityYear="1851" box="[151,483,2005,2027]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="52" pageNumber="53" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A64B2C16FF17F833114DF811" box="[151,296,2005,2027]" italics="true" pageId="52" pageNumber="53">Harpegnathos</emphasis>
<bibRefCitation id="EF4F95E1A64B2C16FEAEF8331186F811" author="Jerdon, T. C." box="[302,483,2005,2027]" pageId="52" pageNumber="53" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon, 1851: 116</bibRefCitation>
</taxonomicName>
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(as genus). Type-species:
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<taxonomicName id="4CDE9393A64B2C16FD6EF83314F6F811" ID-CoL="6LDGT" authority="Jerdon, 1851: 117" authorityName="Jerdon. Soon" authorityPageNumber="117" authorityYear="1851" box="[750,1171,2005,2027]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="52" pageNumber="53" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A64B2C16FD6EF83313B0F811" box="[750,981,2005,2027]" italics="true" pageId="52" pageNumber="53">Harpegnathos saltator</emphasis>
<bibRefCitation id="EF4F95E1A64B2C16FC5BF83314F6F811" author="Jerdon, T. C." box="[987,1171,2005,2027]" pageId="52" pageNumber="53" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon, 1851: 117</bibRefCitation>
</taxonomicName>
</treatmentCitation>
; by monotypy.
<taxonomicName id="4CDE9393A64A2C17FF17FF7E1240FF54" authority="Smith, F., 1858: 81" authorityName="Smith, F." authorityPageNumber="81" authorityYear="1858" box="[151,549,152,174]" class="Insecta" family="Formicidae" genus="Drepanognathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="53" pageNumber="54" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A64A2C17FF17FF7E1125FF54" box="[151,320,152,174]" italics="true" pageId="53" pageNumber="54">Drepanognathus</emphasis>
Smith, F., 1858: 81
</taxonomicName>
(in Poneridae [unnecessary replacement name for
<taxonomicName id="4CDE9393A64A2C17FBDBFF7E1489FF54" ID-CoL="6363H" box="[1115,1260,152,174]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="53" pageNumber="54" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A64A2C17FBDBFF7E1489FF54" box="[1115,1260,152,174]" italics="true" pageId="53" pageNumber="54">Harpegnathos</emphasis>
</taxonomicName>
]). Type-species:
<treatmentCitation id="0A7FCE01A64A2C17FF47FF50120EFF36" ID-DOI="http://doi.org/10.5281/zenodo.6297900" ID-Zenodo-Dep="6297900" LSID="urn:lsid:zoobank.org:act:2CC97E31-567E-4A7A-80E6-3E5CB4AF240E" author="Jerdon, T. C." box="[199,619,182,204]" httpUri="http://treatment.plazi.org/id/9B580F8D2A3E8B47552305C4DA36EAD6" page="117" pageId="53" pageNumber="54" year="1851">
<taxonomicName id="4CDE9393A64A2C17FF47FF50120EFF36" ID-CoL="6LDGT" authority="Jerdon, 1851: 117" authorityName="Jerdon. Soon" authorityPageNumber="117" authorityYear="1851" box="[199,619,182,204]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="53" pageNumber="54" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A64A2C17FF47FF5011C8FF36" box="[199,429,182,204]" italics="true" pageId="53" pageNumber="54">Harpegnathos saltator</emphasis>
<bibRefCitation id="EF4F95E1A64A2C17FE33FF50120EFF36" author="Jerdon, T. C." box="[435,619,182,204]" pageId="53" pageNumber="54" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon, 1851: 117</bibRefCitation>
</taxonomicName>
</treatmentCitation>
; by subsequent designation of
<treatmentCitation id="0A7FCE01A64A2C17FC23FF50140DFF36" author="Bingham, C. T." box="[931,1128,182,204]" page="49" pageId="53" pageNumber="54" year="1903">
<bibRefCitation id="EF4F95E1A64A2C17FC23FF50140DFF36" author="Bingham, C. T." box="[931,1128,182,204]" pageId="53" pageNumber="54" refId="ref132962" refString="Bingham, C. T. (1903) The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. Taylor and Francis, London, 506 pp." type="book" year="1903">Bingham, 1903: 49</bibRefCitation>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph id="8B61E810A64A2C17FF17FF121549FECA" blockId="53.[151,1436,244,304]" pageId="53" pageNumber="54">
<taxonomicName id="4CDE9393A64A2C17FF17FF121152FEF7" ID-CoL="6363H" box="[151,311,244,269]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="53" pageNumber="54" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A64A2C17FF17FF121152FEF7" box="[151,311,244,269]" italics="true" pageId="53" pageNumber="54">Harpegnathos</emphasis>
</taxonomicName>
is a small genus restricted to Southeast Asia. These ants are notable for their remarkable morphological traits, foraging behaviors, complex nest architecture, and unusual reproductive behaviors.
</paragraph>
</subSubSection>
<caption id="DFA1B898A64A2C17FF17F88E12A3F866" ID-DOI="http://doi.org/10.5281/zenodo.10120820" ID-Zenodo-Dep="10120820" httpUri="https://zenodo.org/record/10120820/files/figure.png" pageId="53" pageNumber="54" startId="53.[151,250,1896,1918]" targetBox="[151,1436,357,1862]" targetPageId="53">
<paragraph id="8B61E810A64A2C17FF17F88E12A3F866" blockId="53.[151,1436,1896,1948]" pageId="53" pageNumber="54">
<emphasis id="B9AA3402A64A2C17FF17F88E1171F887" bold="true" box="[151,276,1896,1918]" pageId="53" pageNumber="54">FIGURE 4.</emphasis>
Worker caste of
<taxonomicName id="4CDE9393A64A2C17FE46F88E12C9F884" baseAuthorityName="Crosland" baseAuthorityYear="1995" box="[454,684,1896,1918]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="53" pageNumber="54" phylum="Arthropoda" rank="species" species="venator">
<emphasis id="B9AA3402A64A2C17FE46F88E12C9F884" box="[454,684,1896,1918]" italics="true" pageId="53" pageNumber="54">Harpegnathos venator</emphasis>
</taxonomicName>
: lateral and dorsal view of body and full-face view of head (Philippines, G.Alpert, MCZC); world distribution of
<taxonomicName id="4CDE9393A64A2C17FDAFF86012A5F866" box="[559,704,1926,1948]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="53" pageNumber="54" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A64A2C17FDAFF86012A5F866" box="[559,704,1926,1948]" italics="true" pageId="53" pageNumber="54">Harpegnathos</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<subSubSection id="C3C4BB9BA64A2C14FF47F82F134BFC5F" lastPageId="54" lastPageNumber="55" pageId="53" pageNumber="54" type="diagnosis">
<paragraph id="8B61E810A64A2C14FF47F82F11FAFEBA" blockId="53.[199,1436,1993,2018]" lastBlockId="54.[151,1437,151,2013]" lastPageId="54" lastPageNumber="55" pageId="53" pageNumber="54">
<emphasis id="B9AA3402A64A2C17FF47F82F1124F818" bold="true" box="[199,321,1993,2018]" pageId="53" pageNumber="54">Diagnosis.</emphasis>
Workers of
<taxonomicName id="4CDE9393A64A2C17FE4EF82F120BF818" box="[462,622,1993,2018]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="53" pageNumber="54" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A64A2C17FE4EF82F120BF818" box="[462,622,1993,2018]" italics="true" pageId="53" pageNumber="54">Harpegnathos</emphasis>
</taxonomicName>
are virtually impossible to confuse with those of any other genus. They are among the most morphologically unique of all ponerines, and their scythe-shaped mandibles and huge anteriorly set eyes instantly identify them. Other diagnostic characters (in combination) include the presence of ocelli, an obsolete metanotal groove, a laterally opening metapleural gland orifice, toothed tarsal claws, prominent arolia, a long nodiform petiole, a short rounded anterior face of A3, and a strong girdling constriction between pre- and postsclerites of A4.
</paragraph>
<paragraph id="8B61E810A6492C14FF47FEAA12ECFD0A" blockId="54.[151,1437,151,2013]" pageId="54" pageNumber="55">
<emphasis id="B9AA3402A6492C14FF47FEAA11ADFE9F" bold="true" box="[199,456,332,357]" pageId="54" pageNumber="55">Synoptic description.</emphasis>
<emphasis id="B9AA3402A6492C14FE55FEAA1248FE9F" box="[469,557,332,357]" italics="true" pageId="54" pageNumber="55">Worker.</emphasis>
Large to very large (TL
<quantity id="4C2645F5A6492C14FCE1FEAA1385FE9F" box="[865,992,332,357]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" metricValueMax="2.0" metricValueMin="1.4" pageId="54" pageNumber="55" unit="mm" value="17.0" valueMax="20.0" valueMin="14.0">1420 mm</quantity>
) ants with the standard characters of
<taxonomicName id="4CDE9393A6492C14FF17FE89109CFE72" box="[151,249,367,392]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
, except that the anterior face of A3 is short and rounded, not vertical. Mandibles long, upturned and scythe-shaped, with a small ventral edge bearing a few teeth and a long medial edge bearing two rows of small denticles. Clypeus shallow and with a nearly straight anterior margin. Frontal lobes very small. Eyes greatly enlarged, nearly half the length of the head, and located at the extreme anterolateral corners of the head. Ocelli present. Metanotal groove obsolete dorsally. Propodeum broad dorsally. Propodeal spiracles slit-shaped. Metapleural gland orifice opening laterally within a deep depression. Tarsal claws with a single preapical tooth. Arolia prominent and bright white. Metatibial spur formula (1s, 1p). Petiole longer than wide, becoming wider dorsally and posteriorly. Anterior face of A3 short and rounded, not vertical. Gaster with a strong girdling constriction between pre- and postsclerites of A4. Stridulitrum present on pretergite of A4. Head and mesosoma foveate and striate, the gaster punctate or foveolate. Head and body with abundant short pilosity and little to no pubescence. Color variable, orange to black.
</paragraph>
<paragraph id="8B61E810A6492C14FF47FD1A140BFCEF" blockId="54.[151,1437,151,2013]" box="[199,1134,764,789]" pageId="54" pageNumber="55">
<emphasis id="B9AA3402A6492C14FF47FD1A1172FCE9" box="[199,279,764,787]" italics="true" pageId="54" pageNumber="55">Queen.</emphasis>
Very similar to worker, but winged and slightly larger (
<bibRefCitation id="EF4F95E1A6492C14FC05FD1A1407FCEF" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[901,1122,764,789]" pageId="54" pageNumber="55" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6492C14FC5FFD1B1472FCEF" box="[991,1047,764,789]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 2000
</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6492C14FF47FCF9140EFCC2" blockId="54.[151,1437,151,2013]" box="[199,1131,799,824]" pageId="54" pageNumber="55">
<emphasis id="B9AA3402A6492C14FF47FCF9116DFCC2" box="[199,264,799,824]" italics="true" pageId="54" pageNumber="55">Male.</emphasis>
<collectingRegion id="491A26F2A6492C14FE8EFCF91153FCC2" box="[270,310,799,824]" country="South Sudan" name="Lakes" pageId="54" pageNumber="55">See</collectingRegion>
descriptions in
<bibRefCitation id="EF4F95E1A6492C14FE69FCF912C6FCC2" author="Bingham, C. T." box="[489,675,799,824]" pageId="54" pageNumber="55" refId="ref132962" refString="Bingham, C. T. (1903) The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. Taylor and Francis, London, 506 pp." type="book" year="1903">Bingham (1903)</bibRefCitation>
,
<bibRefCitation id="EF4F95E1A6492C14FD2FFCC61329FCC2" author="Emery, C." box="[687,844,799,824]" pageId="54" pageNumber="55" 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>
, and
<bibRefCitation id="EF4F95E1A6492C14FC08FCF91401FCC2" author="Donisthorpe, H." box="[904,1124,799,824]" pageId="54" pageNumber="55" pagination="64 - 72" refId="ref139595" refString="Donisthorpe, H. (1942) Descriptions of a few ants from the Philippine Islands, and a male of Polyrhachis bihamata Drury from India. Annals and Magazine of Natural History, 11 (9), 64 - 72. http: // dx. doi. org / 10.1080 / 03745481.1942.9755466" type="journal article" year="1942">Donisthorpe (1942)</bibRefCitation>
.
</paragraph>
<paragraph id="8B61E810A6492C14FF47FCA311D9FCA7" blockId="54.[151,1437,151,2013]" box="[199,444,836,861]" pageId="54" pageNumber="55">
<emphasis id="B9AA3402A6492C14FF47FCA31174FCA6" box="[199,273,837,860]" italics="true" pageId="54" pageNumber="55">Larva.</emphasis>
<collectingRegion id="491A26F2A6492C14FE98FCA21127FCA1" box="[280,322,836,859]" country="Cameroon" name="North" pageId="54" pageNumber="55">Not</collectingRegion>
described.
</paragraph>
<paragraph id="8B61E810A6492C14FF47FC81134BFC5F" blockId="54.[151,1437,151,2013]" pageId="54" pageNumber="55">
<emphasis id="B9AA3402A6492C14FF47FC81119CFC7A" bold="true" box="[199,505,871,896]" pageId="54" pageNumber="55">Geographic distribution.</emphasis>
The range of
<taxonomicName id="4CDE9393A6492C14FD3FFC81133AFC7A" box="[703,863,871,896]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FD3FFC81133AFC7A" box="[703,863,871,896]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
encompasses the area bounded by
<collectingCountry id="F3C9A880A6492C14FAA7FC811506FC7A" box="[1319,1379,871,896]" name="India" pageId="54" pageNumber="55">India</collectingCountry>
, the
<collectingCountry id="F3C9A880A6492C14FF17FC6A117DFC5F" box="[151,280,908,933]" name="Philippines" pageId="54" pageNumber="55">Philippines</collectingCountry>
, southern
<collectingCountry id="F3C9A880A6492C14FE0AFC6A11B5FC5F" box="[394,464,908,933]" name="China" pageId="54" pageNumber="55">China</collectingCountry>
, and Java (
<bibRefCitation id="EF4F95E1A6492C14FDCBFC6A1344FC5F" author="Donisthorpe, H." box="[587,801,908,933]" pageId="54" pageNumber="55" pagination="196 - 201" refId="ref139520" refString="Donisthorpe, H. (1937) A new species of Harpegnathos Jerd., with some remarks on the genus, and other known species (Hym. Formicidae). Entomologist's Monthly Magazine, 73, 196 - 201." type="journal article" year="1937">Donisthorpe, 1937</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C3C4BB9BA6492C1AFF47FC4913C7FBCF" lastPageId="56" lastPageNumber="57" pageId="54" pageNumber="55" type="biology_ecology">
<paragraph id="8B61E810A6492C14FF47FC49127AFB12" blockId="54.[151,1437,151,2013]" pageId="54" pageNumber="55">
<emphasis id="B9AA3402A6492C14FF47FC4911BFFC32" bold="true" box="[199,474,943,968]" pageId="54" pageNumber="55">Ecology and behavior.</emphasis>
<taxonomicName id="4CDE9393A6492C14FE66FC4912E3FC32" box="[486,646,943,968]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FE66FC4912E3FC32" box="[486,646,943,968]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
is morphologically and behaviorally one of the most unusual and interesting of all ponerines. Its bizarre scythe-shaped mandibles and huge eyes are instantly recognizable, and its jumping abilities and visual hunting strategy are unique within the
<taxonomicName id="4CDE9393A6492C14FC22FC111477FBEA" baseAuthorityName="sensu Bolton" baseAuthorityYear="2003" box="[930,1042,1015,1040]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
and matched among ants only by
<taxonomicName id="4CDE9393A6492C14FF17FBFB11D2FBCF" authority="(Myrmeciinae)" baseAuthorityName="Myrmeciinae" box="[151,439,1052,1077]" class="Insecta" family="Formicidae" genus="Myrmecia" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FF17FBFB116CFBCE" box="[151,265,1053,1076]" italics="true" pageId="54" pageNumber="55">Myrmecia</emphasis>
(Myrmeciinae)
</taxonomicName>
and
<taxonomicName id="4CDE9393A6492C14FE6EFBFA1366FBCF" authority="(Formicinae)" authorityName="Roger" authorityYear="1863" baseAuthorityName="Formicinae" box="[494,771,1052,1077]" class="Insecta" family="Formicidae" genus="Gigantiops" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FE6EFBFA120FFBC9" box="[494,618,1052,1075]" italics="true" pageId="54" pageNumber="55">Gigantiops</emphasis>
(Formicinae)
</taxonomicName>
(
<bibRefCitation id="EF4F95E1A6492C14FC92FBFA13ADFBCF" author="Wheeler, W. M." box="[786,968,1052,1077]" pageId="54" pageNumber="55" pagination="185 - 201" refId="ref167784" refString="Wheeler, W. M. (1922 a) Observations on Gigantiops destructor Fabricius and other leaping ants. Biological Bulletin (Woods Hole), 42, 185 - 201." type="journal article" year="1922">Wheeler, 1922a</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6492C14FC53FBFA14FFFBCF" author="Tautz, J. &amp; Holldobler, B. &amp; Danker, T." box="[979,1178,1052,1077]" pageId="54" pageNumber="55" pagination="1 - 6" refId="ref163773" refString="Tautz, J., Holldobler, B. &amp; Danker, T. (1994) The ants that jump - different techniques to take-off. Zoology, 98 (1), 1 - 6." type="journal article" year="1994">
Tautz
<emphasis id="B9AA3402A6492C14FB98FBFB1435FBCF" box="[1048,1104,1052,1077]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1994
</bibRefCitation>
).
<taxonomicName id="4CDE9393A6492C14FB2EFBFA152BFBCF" box="[1198,1358,1052,1077]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FB2EFBFA152BFBCF" box="[1198,1358,1052,1077]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
is also notable for its nest architecture, which is unusually complex for a ponerine, and for its unique social system in which reproduction is performed both by dealate queens and gamergates. Nearly everything known about the ecology and behavior of the genus comes from studies of the Indian species
<taxonomicName id="4CDE9393A6492C14FC66FB6F143BFB5A" authorityName="Jerdon. Soon" authorityYear="1851" box="[998,1118,1159,1184]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6492C14FC66FB6F143BFB5A" box="[998,1118,1159,1184]" italics="true" pageId="54" pageNumber="55">H. saltator</emphasis>
</taxonomicName>
, so the following discussion mainly concerns that species unless otherwise noted. The general applicability of these observations to other
<taxonomicName id="4CDE9393A6492C14FF17FB2911EAFB12" box="[151,399,1231,1256]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B9AA3402A6492C14FF17FB291152FB12" box="[151,311,1231,1256]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
species
</taxonomicName>
is unknown.
</paragraph>
<paragraph id="8B61E810A6492C14FF47FB12153EF9F2" blockId="54.[151,1437,151,2013]" pageId="54" pageNumber="55">
<taxonomicName id="4CDE9393A6492C14FF47FB121102FAF7" box="[199,359,1268,1293]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FF47FB121102FAF7" box="[199,359,1268,1293]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
are formidable hunters, and their excellent vision, long mandibles, venomous sting, and jumping capabilities make them superbly adapted to hunting fast moving prey (
<bibRefCitation id="EF4F95E1A6492C14FB91FAF114B2FACA" author="Maschwitz, U. &amp; Jessen, K. &amp; Maschwitz, E." box="[1041,1239,1303,1328]" pageId="54" pageNumber="55" pagination="79 - 81" refId="ref154031" refString="Maschwitz, U., Jessen, K. &amp; Maschwitz, E. (1981) Foaming in Pachycondyla - a new defense mechanism in ants. Behavioral Ecology and Sociobiology, 9 (1), 79 - 81. http: // dx. doi. org / 10.1007 / bf 00299857" type="journal article" year="1981">Maschwitz, 1981</bibRefCitation>
). Their eyes have the largest number of ommatidia of any ant examined and enable them to be visually oriented to a degree that is rare in ants (
<bibRefCitation id="EF4F95E1A6492C14FEA2FAB9122BFA82" author="Baroni Urbani, C. &amp; Boyan, G. S. &amp; Blarer, A. &amp; Billen, J. &amp; Musthak Ali, T. M." box="[290,590,1375,1400]" pageId="54" pageNumber="55" pagination="63 - 71" refId="ref132276" refString="Baroni Urbani, C., Boyan, G. S., Blarer, A., Billen, J. &amp; Musthak Ali, T. M. (1994) A novel mechanism for jumping in the Indian ant Harpegnathos saltator (Jerdon) (Formicidae, Ponerinae). Experientia, 50, 63 - 71. http: // dx. doi. org / 10.1007 / bf 01992052" type="journal article" year="1994">
Baroni Urbani
<emphasis id="B9AA3402A6492C14FE4CFA871261FA82" box="[460,516,1375,1400]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1994
</bibRefCitation>
). Their mandibles are designed for rapid closure, rather than extreme force, with a preponderance of fast muscle fibers controlling mandibular closure (
<bibRefCitation id="EF4F95E1A6492C14FC62FA62156CFA67" author="Paul, J. &amp; Gronenberg, W." box="[994,1289,1412,1437]" pageId="54" pageNumber="55" pagination="797 - 808" refId="ref158561" refString="Paul, J. &amp; Gronenberg, W. (1999) Optimizing force and velocity: mandible muscle fiber attachments in ants. Journal of Experimental Biology, 202, 797 - 808." type="journal article" year="1999">Paul &amp; Gronenberg, 1999</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6492C14FA94FA6215F5FA67" author="Paul, J." box="[1300,1424,1412,1437]" pageId="54" pageNumber="55" pagination="7 - 20" refId="ref158536" refString="Paul, J. (2001) Mandible movements in ants. Comparative Biochemistry and Physiology Part A, 131, 7 - 20." type="journal article" year="2001">Paul, 2001</bibRefCitation>
).
<taxonomicName id="4CDE9393A6492C14FF17FA411152FA3A" box="[151,311,1447,1472]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FF17FA411152FA3A" box="[151,311,1447,1472]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
workers have powerful paralyzing stings which rapidly subdue prey (
<bibRefCitation id="EF4F95E1A6492C14FB01FA4115F2FA3A" author="Maschwitz, U. &amp; Hahn, M. &amp; Schonegge, P." box="[1153,1431,1447,1472]" pageId="54" pageNumber="55" pagination="213 - 214" refId="ref153722" refString="Maschwitz, U., Hahn, M. &amp; Schonegge, P. (1979) Paralysis of prey in ponerine ants. Naturwissenschaften, 66, 213 - 214. http: // dx. doi. org / 10.1007 / bf 00366035" type="journal article" year="1979">
Maschwitz
<emphasis id="B9AA3402A6492C14FA8AFA4F152DFA3A" box="[1290,1352,1447,1472]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1979
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6492C14FF17FA2A113CFA1F" author="Maschwitz, U. &amp; Jessen, K. &amp; Maschwitz, E." box="[151,345,1484,1509]" pageId="54" pageNumber="55" pagination="79 - 81" refId="ref154031" refString="Maschwitz, U., Jessen, K. &amp; Maschwitz, E. (1981) Foaming in Pachycondyla - a new defense mechanism in ants. Behavioral Ecology and Sociobiology, 9 (1), 79 - 81. http: // dx. doi. org / 10.1007 / bf 00299857" type="journal article" year="1981">Maschwitz, 1981</bibRefCitation>
) and are also effectively used in defense (
<emphasis id="B9AA3402A6492C14FCB6FA2B1300FA1E" box="[822,869,1485,1508]" italics="true" pageId="54" pageNumber="55">e.g.,</emphasis>
<bibRefCitation id="EF4F95E1A6492C14FCEDFA2A1464FA1F" author="Jerdon, T. C." box="[877,1025,1484,1509]" pageId="54" pageNumber="55" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon, 1851</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6492C14FB8DFA2A14B2FA1F" author="Wroughton, R. C." box="[1037,1239,1484,1509]" pageId="54" pageNumber="55" pagination="13 - 60" refId="ref169119" refString="Wroughton, R. C. (1892) Our ants. Part I. Journal of the Bombay Natural History Society, 7, 13 - 60." type="journal article" year="1892">Wroughton, 1892</bibRefCitation>
). The paralysis is permanent, and paralyzed prey may be stored for short periods before consumption (
<bibRefCitation id="EF4F95E1A6492C14FBC9FA09152AF9F2" author="Maschwitz, U. &amp; Hahn, M. &amp; Schonegge, P." box="[1097,1359,1519,1544]" pageId="54" pageNumber="55" pagination="213 - 214" refId="ref153722" refString="Maschwitz, U., Hahn, M. &amp; Schonegge, P. (1979) Paralysis of prey in ponerine ants. Naturwissenschaften, 66, 213 - 214. http: // dx. doi. org / 10.1007 / bf 00366035" type="journal article" year="1979">
Maschwitz
<emphasis id="B9AA3402A6492C14FB4CFA171561F9F2" box="[1228,1284,1519,1544]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1979
</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6492C15FF47F9F21456FEBA" blockId="54.[151,1437,151,2013]" lastBlockId="55.[151,1437,151,2027]" lastPageId="55" lastPageNumber="56" pageId="54" pageNumber="55">
The jumping abilities of
<taxonomicName id="4CDE9393A6492C14FE63F9F212E6F9D7" box="[483,643,1556,1581]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FE63F9F212E6F9D7" box="[483,643,1556,1581]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
have been noted in the literature since the original description of the genus (
<bibRefCitation id="EF4F95E1A6492C14FF68F9D1111AF9AA" author="Jerdon, T. C." box="[232,383,1591,1616]" pageId="54" pageNumber="55" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon, 1851</bibRefCitation>
). Early observers (
<emphasis id="B9AA3402A6492C14FDD4F9DF12E6F9AA" box="[596,643,1593,1616]" italics="true" pageId="54" pageNumber="55">e.g.,</emphasis>
<bibRefCitation id="EF4F95E1A6492C14FD0CF9D11345F9AA" author="Jerdon, T. C." box="[652,800,1591,1616]" pageId="54" pageNumber="55" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon, 1851</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6492C14FCADF9D11393F9AA" author="Wroughton, R. C." box="[813,1014,1591,1616]" pageId="54" pageNumber="55" pagination="13 - 60" refId="ref169119" refString="Wroughton, R. C. (1892) Our ants. Part I. Journal of the Bombay Natural History Society, 7, 13 - 60." type="journal article" year="1892">Wroughton, 1892</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6492C14FB82F9D114D5F9AA" author="Bingham, C. T." box="[1026,1200,1591,1616]" pageId="54" pageNumber="55" refId="ref132962" refString="Bingham, C. T. (1903) The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. Taylor and Francis, London, 506 pp." type="book" year="1903">Bingham, 1903</bibRefCitation>
) generally described the jumping as an escape response, but more recent studies have demonstrated that
<taxonomicName id="4CDE9393A6492C14FBF9F9BA157CF98F" box="[1145,1305,1628,1653]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FBF9F9BA157CF98F" box="[1145,1305,1628,1653]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
utilizes its jumping for at least four distinct purposes: escape, normal locomotion, an unusual behavior called “group jumping”, and prey capture (
<bibRefCitation id="EF4F95E1A6492C14FE59F9421295F947" author="Musthak Ali, T. M. &amp; Baroni Urbani, C. &amp; Billen, J." box="[473,752,1700,1725]" pageId="54" pageNumber="55" pagination="374 - 376" refId="ref156519" refString="Musthak Ali, T. M., Baroni Urbani, C. &amp; Billen, J. (1992) Multiple jumping behaviors in the ant Harpegnathos saltator. Naturwissenschaften, 79, 374 - 376. http: // dx. doi. org / 10.1007 / bf 01140185" type="journal article" year="1992">
Musthak Ali
<emphasis id="B9AA3402A6492C14FDEFF94312C2F947" box="[623,679,1700,1725]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1992
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6492C14FD7CF942144DF947" author="Baroni Urbani, C. &amp; Boyan, G. S. &amp; Blarer, A. &amp; Billen, J. &amp; Musthak Ali, T. M." box="[764,1064,1700,1725]" pageId="54" pageNumber="55" pagination="63 - 71" refId="ref132276" refString="Baroni Urbani, C., Boyan, G. S., Blarer, A., Billen, J. &amp; Musthak Ali, T. M. (1994) A novel mechanism for jumping in the Indian ant Harpegnathos saltator (Jerdon) (Formicidae, Ponerinae). Experientia, 50, 63 - 71. http: // dx. doi. org / 10.1007 / bf 01992052" type="journal article" year="1994">
Baroni Urbani
<emphasis id="B9AA3402A6492C14FC25F94313B8F947" box="[933,989,1700,1725]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1994
</bibRefCitation>
). When disturbed,
<taxonomicName id="4CDE9393A6492C14FB7CF94215F9F947" box="[1276,1436,1700,1725]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FB7CF94215F9F947" box="[1276,1436,1700,1725]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
workers jump repeatedly and then hide under leaf litter (
<bibRefCitation id="EF4F95E1A6492C14FCACF921143DF91A" author="Shivashankar, T. &amp; Sharathchandra, H. C. &amp; Veeresh, G. K." box="[812,1112,1735,1760]" pageId="54" pageNumber="55" pagination="367 - 372" refId="ref162177" refString="Shivashankar, T., Sharathchandra, H. C. &amp; Veeresh, G. K. (1989) Foraging activity and temperature relations in the ponerine ant Harpegnathos saltator Jerdon (Formicidae). Proceedings of the Indian Academy Sciences - Animal Sciences, 98 (5), 367 - 372. http: // dx. doi. org / 10.1007 / bf 03179963" type="journal article" year="1989">
Shivashankar
<emphasis id="B9AA3402A6492C14FC4FF92F146EF91A" box="[975,1035,1735,1760]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1989
</bibRefCitation>
). Of all the jumps made by
<taxonomicName id="4CDE9393A6492C14FF17F90A1152F8FF" box="[151,311,1772,1797]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FF17F90A1152F8FF" box="[151,311,1772,1797]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
, these escape jumps cover the longest distance (
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;
<bibRefCitation id="EF4F95E1A6492C14FB8EF90A1550F8FF" author="Musthak Ali, T. M. &amp; Baroni Urbani, C. &amp; Billen, J." box="[1038,1333,1772,1797]" pageId="54" pageNumber="55" pagination="374 - 376" refId="ref156519" refString="Musthak Ali, T. M., Baroni Urbani, C. &amp; Billen, J. (1992) Multiple jumping behaviors in the ant Harpegnathos saltator. Naturwissenschaften, 79, 374 - 376. http: // dx. doi. org / 10.1007 / bf 01140185" type="journal article" year="1992">
Musthak Ali
<emphasis id="B9AA3402A6492C14FB2AF90B1482F8FF" box="[1194,1255,1772,1797]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1992
</bibRefCitation>
). During normal locomotion,
<taxonomicName id="4CDE9393A6492C14FE01F8E91244F8D2" box="[385,545,1807,1832]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FE01F8E91244F8D2" box="[385,545,1807,1832]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
typically make slow, deliberate movements (
<bibRefCitation id="EF4F95E1A6492C14FBAEF8E91484F8D2" author="Crosland, M. W. J." box="[1070,1249,1807,1832]" pageId="54" pageNumber="55" pagination="18 - 23" refId="ref136285" refString="Crosland, M. W. J. (1995) Nest and colony structure in the primitive ant, Harpegnathos venator (Smith) (Hymenoptera, Formicidae). Pan-Pacific Entomologist, 71 (1), 18 - 23." type="journal article" year="1995">Crosland, 1995</bibRefCitation>
), but apparently also employ jumping (Soans &amp; Soans, 1969;
<bibRefCitation id="EF4F95E1A6492C14FD25F8D213BCF8B7" author="Baroni Urbani, C. &amp; Boyan, G. S. &amp; Blarer, A. &amp; Billen, J. &amp; Musthak Ali, T. M." box="[677,985,1844,1869]" pageId="54" pageNumber="55" pagination="63 - 71" refId="ref132276" refString="Baroni Urbani, C., Boyan, G. S., Blarer, A., Billen, J. &amp; Musthak Ali, T. M. (1994) A novel mechanism for jumping in the Indian ant Harpegnathos saltator (Jerdon) (Formicidae, Ponerinae). Experientia, 50, 63 - 71. http: // dx. doi. org / 10.1007 / bf 01992052" type="journal article" year="1994">
Baroni Urbani
<emphasis id="B9AA3402A6492C14FCD2F8D313E8F8B7" box="[850,909,1844,1869]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1994
</bibRefCitation>
).
<bibRefCitation id="EF4F95E1A6492C14FC6FF8D21572F8B7" author="Musthak Ali, T. M. &amp; Baroni Urbani, C. &amp; Billen, J." box="[1007,1303,1844,1869]" pageId="54" pageNumber="55" pagination="374 - 376" refId="ref156519" refString="Musthak Ali, T. M., Baroni Urbani, C. &amp; Billen, J. (1992) Multiple jumping behaviors in the ant Harpegnathos saltator. Naturwissenschaften, 79, 374 - 376. http: // dx. doi. org / 10.1007 / bf 01140185" type="journal article" year="1992">
Musthak Ali
<emphasis id="B9AA3402A6492C14FB08F8D314A6F8B7" box="[1160,1219,1844,1869]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
(1992)
</bibRefCitation>
observed a strange behavior in
<taxonomicName id="4CDE9393A6492C14FEFEF8BF119EF88A" authorityName="Jerdon. Soon" authorityYear="1851" box="[382,507,1879,1904]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6492C14FEFEF8BF119EF88A" box="[382,507,1879,1904]" italics="true" pageId="54" pageNumber="55">H. saltator</emphasis>
</taxonomicName>
which they termed “group jumping”: an individual worker will sometimes start jumping in an uncoordinated fashion for several minutes, and eventually other workers in the vicinity follow suit. The function of this behavior is unknown, but it may serve to flush out prey or to prevent attacks by parasitoids (
<bibRefCitation id="EF4F95E1A6492C14FF1EF82211D2F827" author="Musthak Ali, T. M. &amp; Baroni Urbani, C. &amp; Billen, J." box="[158,439,1988,2013]" pageId="54" pageNumber="55" pagination="374 - 376" refId="ref156519" refString="Musthak Ali, T. M., Baroni Urbani, C. &amp; Billen, J. (1992) Multiple jumping behaviors in the ant Harpegnathos saltator. Naturwissenschaften, 79, 374 - 376. http: // dx. doi. org / 10.1007 / bf 01140185" type="journal article" year="1992">
Musthak Ali
<emphasis id="B9AA3402A6492C14FEB4F8231108F827" box="[308,365,1988,2013]" italics="true" pageId="54" pageNumber="55">et al.</emphasis>
, 1992
</bibRefCitation>
).
<taxonomicName id="4CDE9393A6492C14FE4BF822120EF827" box="[459,619,1988,2013]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="54" pageNumber="55" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6492C14FE4BF822120EF827" box="[459,619,1988,2013]" italics="true" pageId="54" pageNumber="55">Harpegnathos</emphasis>
</taxonomicName>
workers also jump to capture fast prey and can even catch flying insects in midair, but do not jump at slow prey such as termites (
<bibRefCitation id="EF4F95E1A6482C15FC93FF711451FF4A" author="Shivashankar, T. &amp; Sharathchandra, H. C. &amp; Veeresh, G. K." box="[787,1076,151,176]" pageId="55" pageNumber="56" pagination="367 - 372" refId="ref162177" refString="Shivashankar, T., Sharathchandra, H. C. &amp; Veeresh, G. K. (1989) Foraging activity and temperature relations in the ponerine ant Harpegnathos saltator Jerdon (Formicidae). Proceedings of the Indian Academy Sciences - Animal Sciences, 98 (5), 367 - 372. http: // dx. doi. org / 10.1007 / bf 03179963" type="journal article" year="1989">
Shivashankar
<emphasis id="B9AA3402A6482C15FC33FF7F138EFF4A" box="[947,1003,151,176]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1989
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FBC0FF711531FF4A" author="Musthak Ali, T. M. &amp; Baroni Urbani, C. &amp; Billen, J." box="[1088,1364,151,176]" pageId="55" pageNumber="56" pagination="374 - 376" refId="ref156519" refString="Musthak Ali, T. M., Baroni Urbani, C. &amp; Billen, J. (1992) Multiple jumping behaviors in the ant Harpegnathos saltator. Naturwissenschaften, 79, 374 - 376. http: // dx. doi. org / 10.1007 / bf 01140185" type="journal article" year="1992">
Musthak Ali
<emphasis id="B9AA3402A6482C15FB53FF7F156EFF4A" box="[1235,1291,151,176]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1992
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FAE0FF7E109AFF2F" author="Ke, Y. &amp; Zhuang, T. &amp; Wang, C. &amp; Zhao, S. &amp; Tian, W." pageId="55" pageNumber="56" pagination="199 - 206" refId="ref148970" refString="Ke, Y., Zhuang, T., Wang, C., Zhao, S. &amp; Tian, W. (2008) Agonistic behavior of six ant species to Coptotermes formosanus (Isoptera: Rhinotermitidae) in laboratory assays. Sociobiology, 51 (1), 199 - 206." type="journal article" year="2008">
Ke
<emphasis id="B9AA3402A6482C15FA08FF7F10D1FF2F" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2008
</bibRefCitation>
). These predatory jumps tend to cover shorter distances than the escape jumps (
<bibRefCitation id="EF4F95E1A6482C15FBF8FF5A15F5FF2F" author="Musthak Ali, T. M. &amp; Baroni Urbani, C. &amp; Billen, J." box="[1144,1424,188,213]" pageId="55" pageNumber="56" pagination="374 - 376" refId="ref156519" refString="Musthak Ali, T. M., Baroni Urbani, C. &amp; Billen, J. (1992) Multiple jumping behaviors in the ant Harpegnathos saltator. Naturwissenschaften, 79, 374 - 376. http: // dx. doi. org / 10.1007 / bf 01140185" type="journal article" year="1992">
Musthak Ali
<emphasis id="B9AA3402A6482C15FA8EFF5B1523FF2F" box="[1294,1350,188,213]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1992
</bibRefCitation>
).
<bibRefCitation id="EF4F95E1A6482C15FF17FF391131FF02" author="Wheeler, W. M." box="[151,340,223,248]" pageId="55" pageNumber="56" pagination="185 - 201" refId="ref167784" refString="Wheeler, W. M. (1922 a) Observations on Gigantiops destructor Fabricius and other leaping ants. Biological Bulletin (Woods Hole), 42, 185 - 201." type="journal article" year="1922">Wheeler (1922a)</bibRefCitation>
hypothesized that the mandibles must be involved in generating the forces necessary for leaping, but recent studies of
<taxonomicName id="4CDE9393A6482C15FE06FEE21243FEE7" box="[390,550,260,285]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FE06FEE21243FEE7" box="[390,550,260,285]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
jumping have confirmed that it is the legs which drive the jumps (though the biomechanics involved are unclear:
<bibRefCitation id="EF4F95E1A6482C15FDABFEC11331FEBA" author="Baroni Urbani, C. &amp; Boyan, G. S. &amp; Blarer, A. &amp; Billen, J. &amp; Musthak Ali, T. M." box="[555,852,295,320]" pageId="55" pageNumber="56" pagination="63 - 71" refId="ref132276" refString="Baroni Urbani, C., Boyan, G. S., Blarer, A., Billen, J. &amp; Musthak Ali, T. M. (1994) A novel mechanism for jumping in the Indian ant Harpegnathos saltator (Jerdon) (Formicidae, Ponerinae). Experientia, 50, 63 - 71. http: // dx. doi. org / 10.1007 / bf 01992052" type="journal article" year="1994">
Baroni Urbani
<emphasis id="B9AA3402A6482C15FD53FECF136EFEBA" box="[723,779,295,320]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1994
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FCE0FECE1442FEBA" author="Tautz, J. &amp; Holldobler, B. &amp; Danker, T." box="[864,1063,295,320]" pageId="55" pageNumber="56" pagination="1 - 6" refId="ref163773" refString="Tautz, J., Holldobler, B. &amp; Danker, T. (1994) The ants that jump - different techniques to take-off. Zoology, 98 (1), 1 - 6." type="journal article" year="1994">
Tautz
<emphasis id="B9AA3402A6482C15FC24FECF13B8FEBA" box="[932,989,295,320]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1994
</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6482C15FF47FEAA1484FDC1" blockId="55.[151,1437,151,2027]" pageId="55" pageNumber="56">
<taxonomicName id="4CDE9393A6482C15FF47FEAA1102FE9F" box="[199,359,332,357]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FF47FEAA1102FE9F" box="[199,359,332,357]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
are crepuscular foragers, requiring daylight to locate prey but preferring to avoid the heat of midday (
<bibRefCitation id="EF4F95E1A6482C15FF7AFE89127AFE72" author="Shivashankar, T. &amp; Sharathchandra, H. C. &amp; Veeresh, G. K." box="[250,543,367,392]" pageId="55" pageNumber="56" pagination="367 - 372" refId="ref162177" refString="Shivashankar, T., Sharathchandra, H. C. &amp; Veeresh, G. K. (1989) Foraging activity and temperature relations in the ponerine ant Harpegnathos saltator Jerdon (Formicidae). Proceedings of the Indian Academy Sciences - Animal Sciences, 98 (5), 367 - 372. http: // dx. doi. org / 10.1007 / bf 03179963" type="journal article" year="1989">
Shivashankar
<emphasis id="B9AA3402A6482C15FE1BFE9711B1FE72" box="[411,468,367,392]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1989
</bibRefCitation>
). They hunt individually in leaf litter and among low vegetation for a diversity of arthropods (
<bibRefCitation id="EF4F95E1A6482C15FEBFFE7211B2FE57" author="Jerdon, T. C." box="[319,471,404,429]" pageId="55" pageNumber="56" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon, 1851</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FE64FE721288FE57" author="Maschwitz, U. &amp; Hahn, M. &amp; Schonegge, P." box="[484,749,404,429]" pageId="55" pageNumber="56" pagination="213 - 214" refId="ref153722" refString="Maschwitz, U., Hahn, M. &amp; Schonegge, P. (1979) Paralysis of prey in ponerine ants. Naturwissenschaften, 66, 213 - 214. http: // dx. doi. org / 10.1007 / bf 00366035" type="journal article" year="1979">
Maschwitz
<emphasis id="B9AA3402A6482C15FDE8FE7312C6FE57" box="[616,675,404,429]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1979
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FD7BFE7213A4FE57" author="Maschwitz, U. &amp; Jessen, K. &amp; Maschwitz, E." box="[763,961,404,429]" pageId="55" pageNumber="56" pagination="79 - 81" refId="ref154031" refString="Maschwitz, U., Jessen, K. &amp; Maschwitz, E. (1981) Foaming in Pachycondyla - a new defense mechanism in ants. Behavioral Ecology and Sociobiology, 9 (1), 79 - 81. http: // dx. doi. org / 10.1007 / bf 00299857" type="journal article" year="1981">Maschwitz, 1981</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FC4FFE721496FE57" author="Shivashankar, T. &amp; Sharathchandra, H. C. &amp; Veeresh, G. K." box="[975,1267,404,429]" pageId="55" pageNumber="56" pagination="367 - 372" refId="ref162177" refString="Shivashankar, T., Sharathchandra, H. C. &amp; Veeresh, G. K. (1989) Foraging activity and temperature relations in the ponerine ant Harpegnathos saltator Jerdon (Formicidae). Proceedings of the Indian Academy Sciences - Animal Sciences, 98 (5), 367 - 372. http: // dx. doi. org / 10.1007 / bf 03179963" type="journal article" year="1989">
Shivashankar
<emphasis id="B9AA3402A6482C15FBEEFE7314CCFE57" box="[1134,1193,404,429]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1989
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FA81FE7210B6FE2A" author="Peeters, C. &amp; Holldobler, B. &amp; Moffett, M. &amp; Musthak Ali, T. M." pageId="55" pageNumber="56" pagination="211 - 218" refId="ref159548" refString="Peeters, C., Holldobler, B., Moffett, M. &amp; Musthak Ali, T. M. (1994) &quot; Wall-papering &quot; and elaborate nest architecture in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 41, 211 - 218. http: // dx. doi. org / 10.1007 / bf 01240479" type="journal article" year="1994">
Peeters
<emphasis id="B9AA3402A6482C15FADBFE7315F0FE57" box="[1371,1429,404,429]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1994
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FF60FE511192FE2A" author="Liebig, J. &amp; Poethke, H." box="[224,503,439,464]" pageId="55" pageNumber="56" pagination="203 - 207" refId="ref151554" refString="Liebig, J. &amp; Poethke, H. (2004) Queen lifespan and colony longevity in the ant Harpegnathos saltator. Ecological Entomology, 29, 203 - 207. http: // dx. doi. org / 10.1111 / j. 1365 - 2311.2004.00583. x" type="journal article" year="2004">Liebig &amp; Poethke, 2004</bibRefCitation>
), though a substantial proportion of their diet consists of fast-running or jumping prey such as crickets, cockroaches, cicadas, flies, and spiders (
<bibRefCitation id="EF4F95E1A6482C15FCE8FE3A1455FE0F" author="Maschwitz, U. &amp; Jessen, K. &amp; Maschwitz, E." box="[872,1072,476,501]" pageId="55" pageNumber="56" pagination="79 - 81" refId="ref154031" refString="Maschwitz, U., Jessen, K. &amp; Maschwitz, E. (1981) Foaming in Pachycondyla - a new defense mechanism in ants. Behavioral Ecology and Sociobiology, 9 (1), 79 - 81. http: // dx. doi. org / 10.1007 / bf 00299857" type="journal article" year="1981">Maschwitz, 1981</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FBBEFE3A1500FE0F" author="Shivashankar, T. &amp; Sharathchandra, H. C. &amp; Veeresh, G. K." box="[1086,1381,476,501]" pageId="55" pageNumber="56" pagination="367 - 372" refId="ref162177" refString="Shivashankar, T., Sharathchandra, H. C. &amp; Veeresh, G. K. (1989) Foraging activity and temperature relations in the ponerine ant Harpegnathos saltator Jerdon (Formicidae). Proceedings of the Indian Academy Sciences - Animal Sciences, 98 (5), 367 - 372. http: // dx. doi. org / 10.1007 / bf 03179963" type="journal article" year="1989">
Shivashankar
<emphasis id="B9AA3402A6482C15FB5EFE3B157CFE0F" box="[1246,1305,476,501]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1989
</bibRefCitation>
). By enabling them to utilize such prey, which might normally be difficult for an ant to capture, the specialized adaptations of
<taxonomicName id="4CDE9393A6482C15FEBCFDC211B9FDC7" box="[316,476,548,573]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FEBCFDC211B9FDC7" box="[316,476,548,573]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
allow it to fill a niche which is otherwise not fully exploited by ants.
</paragraph>
<paragraph id="8B61E810A6482C15FF47FDA1117DFCEF" blockId="55.[151,1437,151,2027]" pageId="55" pageNumber="56">
<bibRefCitation id="EF4F95E1A6482C15FF47FDA11194FD9A" author="Liebig, J. &amp; Poethke, H." box="[199,497,583,608]" pageId="55" pageNumber="56" pagination="203 - 207" refId="ref151554" refString="Liebig, J. &amp; Poethke, H. (2004) Queen lifespan and colony longevity in the ant Harpegnathos saltator. Ecological Entomology, 29, 203 - 207. http: // dx. doi. org / 10.1111 / j. 1365 - 2311.2004.00583. x" type="journal article" year="2004">Liebig &amp; Poethke (2004)</bibRefCitation>
found no evidence of intercolony aggression in
<taxonomicName id="4CDE9393A6482C15FBBCFDAF14DEFD9A" authorityName="Jerdon. Soon" authorityYear="1851" box="[1084,1211,583,608]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FBBCFDAF14DEFD9A" box="[1084,1211,583,608]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
despite potentially overlapping foraging ranges, and
<bibRefCitation id="EF4F95E1A6482C15FD94FD8A1284FD7F" author="Maschwitz, U. &amp; Jessen, K. &amp; Maschwitz, E." box="[532,737,620,645]" pageId="55" pageNumber="56" pagination="79 - 81" refId="ref154031" refString="Maschwitz, U., Jessen, K. &amp; Maschwitz, E. (1981) Foaming in Pachycondyla - a new defense mechanism in ants. Behavioral Ecology and Sociobiology, 9 (1), 79 - 81. http: // dx. doi. org / 10.1007 / bf 00299857" type="journal article" year="1981">Maschwitz (1981)</bibRefCitation>
found no evidence of chemical recruitment in either foraging or emigrations of
<taxonomicName id="4CDE9393A6482C15FEE3FD7711B8FD52" authorityName="Jerdon. Soon" authorityYear="1851" box="[355,477,655,680]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FEE3FD7711B8FD52" box="[355,477,655,680]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
(emigrations occur via social carrying). The chemical ecology of
<taxonomicName id="4CDE9393A6482C15FB4EFD69150BFD52" box="[1230,1390,655,680]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FB4EFD69150BFD52" box="[1230,1390,655,680]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
has not been extensively studied, though
<bibRefCitation id="EF4F95E1A6482C15FDB9FD521335FD37" author="Nascimento, R. R. do &amp; Billen, J. &amp; Morgan, E. D." box="[569,848,692,717]" pageId="55" pageNumber="56" pagination="505 - 508" refId="ref156910" refString="Nascimento, R. R. do, Billen, J. &amp; Morgan, E. D. (1993) The exocrine secretions of the jumping ant Harpegnathos saltator. Comparative Biochemistry and Physiology, 104 B (3), 505 - 508. http: // dx. doi. org / 10.1016 / 0305 - 0491 (93) 90274 - 9" type="journal article" year="1993">
Nascimento
<emphasis id="B9AA3402A6482C15FD46FD53129AFD37" box="[710,767,692,717]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
(1993)
</bibRefCitation>
examined the exocrine secretions of
<taxonomicName id="4CDE9393A6482C15FB73FD531509FD37" authorityName="Jerdon. Soon" authorityYear="1851" box="[1267,1388,692,717]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FB73FD531509FD37" box="[1267,1388,692,717]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
and
<bibRefCitation id="EF4F95E1A6482C15FF17FD3111E4FD0A" author="Gobin, B. &amp; Ito, F. &amp; Billen, J." box="[151,385,727,752]" pageId="55" pageNumber="56" pagination="285 - 291" refId="ref144054" refString="Gobin, B., Ito, F. &amp; Billen, J. (2003 b) The subepithelial gland in ants: a novel exocrine gland closely associated with the cuticle surface. Acta Zoologica, 84, 285 - 291. http: // dx. doi. org / 10.1046 / j. 1463 - 6395.2003.00149. x" type="journal article" year="2003">
Gobin
<emphasis id="B9AA3402A6482C15FF66FD3F1145FD0A" box="[230,288,727,752]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
(2003b)
</bibRefCitation>
discovered a unique epidermal gland in
<taxonomicName id="4CDE9393A6482C15FCD6FD3F13B7FD0A" authorityName="Jerdon. Soon" authorityYear="1851" box="[854,978,727,752]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FCD6FD3F13B7FD0A" box="[854,978,727,752]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
which may be involved in hydrocarbon production.
</paragraph>
<paragraph id="8B61E810A6482C15FF47FCC6117BFB5A" blockId="55.[151,1437,151,2027]" pageId="55" pageNumber="56">
<bibRefCitation id="EF4F95E1A6482C15FF47FCC611D5FCC2" author="Peeters, C. &amp; Holldobler, B. &amp; Moffett, M. &amp; Musthak Ali, T. M." box="[199,432,799,824]" pageId="55" pageNumber="56" pagination="211 - 218" refId="ref159548" refString="Peeters, C., Holldobler, B., Moffett, M. &amp; Musthak Ali, T. M. (1994) &quot; Wall-papering &quot; and elaborate nest architecture in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 41, 211 - 218. http: // dx. doi. org / 10.1007 / bf 01240479" type="journal article" year="1994">
Peeters
<emphasis id="B9AA3402A6482C15FEA1FCC71139FCC2" box="[289,348,799,824]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
(1994)
</bibRefCitation>
studied the nests of
<taxonomicName id="4CDE9393A6482C15FD22FCC7137BFCC2" authorityName="Jerdon. Soon" authorityYear="1851" box="[674,798,799,824]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FD22FCC7137BFCC2" box="[674,798,799,824]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
and discovered that they are exceptionally complex by ponerine standards. In a mature colony the nest consists of a series of stacked chambers forming a nearly spherical structure, surmounted by a thick vaulted roof and separated from the surrounding soil by a hollow space. A tunnel leads down to a separate refuse chamber.
<bibRefCitation id="EF4F95E1A6482C15FDE8FC6A132CFC5F" author="Peeters, C. &amp; Holldobler, B. &amp; Moffett, M. &amp; Musthak Ali, T. M." box="[616,841,908,933]" pageId="55" pageNumber="56" pagination="211 - 218" refId="ref159548" refString="Peeters, C., Holldobler, B., Moffett, M. &amp; Musthak Ali, T. M. (1994) &quot; Wall-papering &quot; and elaborate nest architecture in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 41, 211 - 218. http: // dx. doi. org / 10.1007 / bf 01240479" type="journal article" year="1994">
Peeters
<emphasis id="B9AA3402A6482C15FD3FFC6B129DFC5F" box="[703,760,908,933]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
(1994)
</bibRefCitation>
hypothesized that this nest design is an adaptation to survive periodic flooding, which is common in the areas inhabited by
<taxonomicName id="4CDE9393A6482C15FC24FC571479FC32" authorityName="Jerdon. Soon" authorityYear="1851" box="[932,1052,943,968]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FC24FC571479FC32" box="[932,1052,943,968]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
. This idea is supported by the fact that
<taxonomicName id="4CDE9393A6482C15FF49FC331121FC17" authorityName="Jerdon. Soon" authorityYear="1851" box="[201,324,980,1005]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FF49FC331121FC17" box="[201,324,980,1005]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
“wallpapers” the surfaces of its nests with discarded cocoons, insect parts, and plant matter. This wallpaper is glued down and probably helps keep the chambers dry.
<bibRefCitation id="EF4F95E1A6482C15FC2BFC1114BDFBEA" author="Shivashankar, T. &amp; Sharathchandra, H. C. &amp; Veeresh, G. K." box="[939,1240,1015,1040]" pageId="55" pageNumber="56" pagination="367 - 372" refId="ref162177" refString="Shivashankar, T., Sharathchandra, H. C. &amp; Veeresh, G. K. (1989) Foraging activity and temperature relations in the ponerine ant Harpegnathos saltator Jerdon (Formicidae). Proceedings of the Indian Academy Sciences - Animal Sciences, 98 (5), 367 - 372. http: // dx. doi. org / 10.1007 / bf 03179963" type="journal article" year="1989">
Shivashankar
<emphasis id="B9AA3402A6482C15FBCBFC1F14E0FBEA" box="[1099,1157,1015,1040]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
(1989)
</bibRefCitation>
further observed that
<taxonomicName id="4CDE9393A6482C15FF49FBFB1126FBCF" authorityName="Jerdon. Soon" authorityYear="1851" box="[201,323,1052,1077]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FF49FBFB1126FBCF" box="[201,323,1052,1077]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
</taxonomicName>
nest entrances are purposefully plugged with rodent feces or plant matter. The nests of
<taxonomicName id="4CDE9393A6482C15FAA3FBFB15FEFBCE" baseAuthorityName="Crosland" baseAuthorityYear="1995" box="[1315,1435,1053,1076]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="venator">
<emphasis id="B9AA3402A6482C15FAA3FBFB15FEFBCE" box="[1315,1435,1053,1076]" italics="true" pageId="55" pageNumber="56">H. venator</emphasis>
</taxonomicName>
are comparatively simple, consisting of two disc-shaped chambers separated by a unique funnel-like opening (
<bibRefCitation id="EF4F95E1A6482C15FF1EFB821136FB87" author="Crosland, M. W. J." box="[158,339,1124,1149]" pageId="55" pageNumber="56" pagination="18 - 23" refId="ref136285" refString="Crosland, M. W. J. (1995) Nest and colony structure in the primitive ant, Harpegnathos venator (Smith) (Hymenoptera, Formicidae). Pan-Pacific Entomologist, 71 (1), 18 - 23." type="journal article" year="1995">Crosland, 1995</bibRefCitation>
); this funnel could make the inner chamber accessible only via jumping, an effective defense mechanism.
</paragraph>
<paragraph id="8B61E810A6482C15FF47FB4A13FEF98F" blockId="55.[151,1437,151,2027]" pageId="55" pageNumber="56">
Like most ponerines,
<taxonomicName id="4CDE9393A6482C15FE3CFB4A1239FB3F" box="[444,604,1196,1221]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FE3CFB4A1239FB3F" box="[444,604,1196,1221]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
have small colonies, with an average of 65 to
<specimenCount id="9DD82399A6482C15FBF0FB4A1499FB3F" box="[1136,1276,1196,1221]" pageId="55" pageNumber="56" type="worker">225 workers</specimenCount>
in
<taxonomicName id="4CDE9393A6482C15FAA1FB4B111AFB12" authority="(Peeters et al., 2000)" baseAuthorityName="Peeters" baseAuthorityYear="2000" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6482C15FAA1FB4B15F9FB3F" box="[1313,1436,1196,1221]" italics="true" pageId="55" pageNumber="56">H. saltator</emphasis>
(
<bibRefCitation id="EF4F95E1A6482C15FF1EFB291112FB12" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[158,375,1231,1256]" pageId="55" pageNumber="56" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6482C15FF78FB371155FB12" box="[248,304,1231,1256]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
)
</taxonomicName>
and
<specimenCount id="9DD82399A6482C15FE35FB291257FB12" box="[437,562,1231,1256]" pageId="55" pageNumber="56" type="worker">35 workers</specimenCount>
in
<taxonomicName id="4CDE9393A6482C15FDD6FB3713F0FB12" authority="(Crosland, 1995)" baseAuthorityName="Crosland" baseAuthorityYear="1995" box="[598,917,1231,1256]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="venator">
<emphasis id="B9AA3402A6482C15FDD6FB3712ABFB12" box="[598,718,1233,1256]" italics="true" pageId="55" pageNumber="56">H. venator</emphasis>
(
<bibRefCitation id="EF4F95E1A6482C15FD5DFB2913E8FB12" author="Crosland, M. W. J." box="[733,909,1231,1256]" pageId="55" pageNumber="56" pagination="18 - 23" refId="ref136285" refString="Crosland, M. W. J. (1995) Nest and colony structure in the primitive ant, Harpegnathos venator (Smith) (Hymenoptera, Formicidae). Pan-Pacific Entomologist, 71 (1), 18 - 23." type="journal article" year="1995">Crosland, 1995</bibRefCitation>
)
</taxonomicName>
. New colonies are founded by a single dealate queen (
<bibRefCitation id="EF4F95E1A6482C15FF68FB121245FAF7" author="Peeters, C. &amp; Holldobler, B." box="[232,544,1268,1293]" pageId="55" pageNumber="56" pagination="10977 - 10979" refId="ref159484" refString="Peeters, C. &amp; Holldobler, B. (1995) Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest. Proceedings of the National Academy of Sciences (USA), 92, 10977 - 10979. http: // dx. doi. org / 10.1073 / pnas. 92.24.10977" type="journal article" year="1995">Peeters &amp; Hölldobler, 1995</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FDACFB121360FAF7" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[556,773,1268,1293]" pageId="55" pageNumber="56" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6482C15FD04FB1312D9FAF7" box="[644,700,1268,1293]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
; some polygynous queenright colonies have been observed in
<taxonomicName id="4CDE9393A6482C15FF35FAFF118AFACA" authority=": Crosland, 1995" authorityName="Crosland" authorityYear="1995" box="[181,495,1303,1328]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="species" species="venator">
<emphasis id="B9AA3402A6482C15FF35FAFF114BFACA" box="[181,302,1305,1328]" italics="true" pageId="55" pageNumber="56">H. venator</emphasis>
:
<bibRefCitation id="EF4F95E1A6482C15FEBEFAF1118AFACA" author="Crosland, M. W. J." box="[318,495,1303,1328]" pageId="55" pageNumber="56" pagination="18 - 23" refId="ref136285" refString="Crosland, M. W. J. (1995) Nest and colony structure in the primitive ant, Harpegnathos venator (Smith) (Hymenoptera, Formicidae). Pan-Pacific Entomologist, 71 (1), 18 - 23." type="journal article" year="1995">Crosland, 1995</bibRefCitation>
</taxonomicName>
). Like most ponerines, virgin alate queens leave their natal nests, mate, found new colonies, and continue to forage until the first crop of workers are mature, though from this point on the colonys reproductive life cycle is highly unusual (
<bibRefCitation id="EF4F95E1A6482C15FDFAFAB913D9FA82" author="Peeters, C. &amp; Holldobler, B." box="[634,956,1375,1400]" pageId="55" pageNumber="56" pagination="10977 - 10979" refId="ref159484" refString="Peeters, C. &amp; Holldobler, B. (1995) Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest. Proceedings of the National Academy of Sciences (USA), 92, 10977 - 10979. http: // dx. doi. org / 10.1073 / pnas. 92.24.10977" type="journal article" year="1995">Peeters &amp; Hölldobler, 1995</bibRefCitation>
).
<taxonomicName id="4CDE9393A6482C15FC52FAB91417FA82" box="[978,1138,1375,1400]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FC52FAB91417FA82" box="[978,1138,1375,1400]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
workers retain functional spermathecae (J. Billen, pers. comm. in
<bibRefCitation id="EF4F95E1A6482C15FDD7FA621355FA67" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[599,816,1412,1437]" pageId="55" pageNumber="56" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6482C15FD2EFA631282FA67" box="[686,743,1412,1437]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
), and many of them will mate with their brothers inside the nest (
<bibRefCitation id="EF4F95E1A6482C15FF7EFA4111BAFA3A" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[254,479,1447,1472]" pageId="55" pageNumber="56" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6482C15FEDAFA4F11F6FA3A" box="[346,403,1447,1472]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
). These mated workers do not begin to lay eggs until the colonys short-lived queen senesces or dies (
<bibRefCitation id="EF4F95E1A6482C15FED7FA2A12EBFA1F" author="Peeters, C. &amp; Holldobler, B." box="[343,654,1484,1509]" pageId="55" pageNumber="56" pagination="10977 - 10979" refId="ref159484" refString="Peeters, C. &amp; Holldobler, B. (1995) Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest. Proceedings of the National Academy of Sciences (USA), 92, 10977 - 10979. http: // dx. doi. org / 10.1073 / pnas. 92.24.10977" type="journal article" year="1995">Peeters &amp; Hölldobler, 1995</bibRefCitation>
). In newly orphaned colonies, mated workers aggressively compete in order to achieve reproductive status (
<bibRefCitation id="EF4F95E1A6482C15FDC4FA0912BEF9F2" author="Liebig, J. &amp; Holldobler, B. &amp; Peeters, C." box="[580,731,1519,1544]" pageId="55" pageNumber="56" pagination="133 - 135" refId="ref151369" refString="Liebig, J., Holldobler, B. &amp; Peeters, C. (1998) Are ant workers capable of colony foundation? Naturwissenschaften, 85, 133 - 135. http: // dx. doi. org / 10.1007 / s 001140050470" type="journal article" year="1998">Liebig, 1998</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FD69FA0913A0F9F2" author="Liebig, J. &amp; Peeters, C. &amp; Oldham, N. J. &amp; Markstadter, C. &amp; Holldobler, B." box="[745,965,1519,1544]" pageId="55" pageNumber="56" pagination="4124 - 4131" refId="ref151471" refString="Liebig, J., Peeters, C., Oldham, N. J., Markstadter, C. &amp; Holldobler, B. (2000) Are variations in cuticular hydrocarbons of queens and workers a reliable signal of fertility in the ant Harpegnathos saltator? Proceedings of the National Academy of Sciences (USA), 97 (8), 4124 - 4131. http: // dx. doi. org / 10.1073 / pnas. 97.8.4124" type="journal article" year="2000">
Liebig
<emphasis id="B9AA3402A6482C15FCBCFA171312F9F2" box="[828,887,1519,1544]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
). Those that are successful develop their ovaries and begin laying eggs (they are now considered gamergates;
<bibRefCitation id="EF4F95E1A6482C15FC16F9F21403F9D7" author="Liebig, J. &amp; Holldobler, B. &amp; Peeters, C." box="[918,1126,1556,1581]" pageId="55" pageNumber="56" pagination="133 - 135" refId="ref151369" refString="Liebig, J., Holldobler, B. &amp; Peeters, C. (1998) Are ant workers capable of colony foundation? Naturwissenschaften, 85, 133 - 135. http: // dx. doi. org / 10.1007 / s 001140050470" type="journal article" year="1998">
Liebig
<emphasis id="B9AA3402A6482C15FC65F9F31478F9D7" box="[997,1053,1556,1581]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1998
</bibRefCitation>
, 2000;
<bibRefCitation id="EF4F95E1A6482C15FB37F9F215F5F9D7" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[1207,1424,1556,1581]" pageId="55" pageNumber="56" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6482C15FA8EF9F31523F9D7" box="[1294,1350,1556,1581]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
). As the active gamergates gradually lose fecundity or die, new gamergates arise to take their place, and as a result a
<taxonomicName id="4CDE9393A6482C15FF17F9BA1152F98F" box="[151,311,1628,1653]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FF17F9BA1152F98F" box="[151,311,1628,1653]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
colony is theoretically immortal (
<bibRefCitation id="EF4F95E1A6482C15FD33F9BA13F5F98F" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[691,912,1628,1653]" pageId="55" pageNumber="56" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6482C15FC8CF9BB1320F98F" box="[780,837,1628,1653]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6482C15FF47F99915F8F878" blockId="55.[151,1437,151,2027]" pageId="55" pageNumber="56">
This complex reproductive system has recently been examined by several research groups. Whereas queenright colonies are generally monogynous, queenless colonies typically contain a large number of mated workers, of which a significant fraction are reproductively active gamergates (
<bibRefCitation id="EF4F95E1A6482C15FB9BF924153EF921" author="Peeters, C. &amp; Holldobler, B." box="[1051,1371,1730,1755]" pageId="55" pageNumber="56" pagination="10977 - 10979" refId="ref159484" refString="Peeters, C. &amp; Holldobler, B. (1995) Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest. Proceedings of the National Academy of Sciences (USA), 92, 10977 - 10979. http: // dx. doi. org / 10.1073 / pnas. 92.24.10977" type="journal article" year="1995">Peeters &amp; Hölldobler, 1995</bibRefCitation>
). The reproductive status of a
<taxonomicName id="4CDE9393A6482C15FE24F9021221F907" box="[420,580,1764,1789]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FE24F9021221F907" box="[420,580,1764,1789]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
worker is communicated via its cuticular hydrocarbon profile (
<bibRefCitation id="EF4F95E1A6482C15FA8BF90210B6F8E4" author="Liebig, J. &amp; Peeters, C. &amp; Oldham, N. J. &amp; Markstadter, C. &amp; Holldobler, B." pageId="55" pageNumber="56" pagination="4124 - 4131" refId="ref151471" refString="Liebig, J., Peeters, C., Oldham, N. J., Markstadter, C. &amp; Holldobler, B. (2000) Are variations in cuticular hydrocarbons of queens and workers a reliable signal of fertility in the ant Harpegnathos saltator? Proceedings of the National Academy of Sciences (USA), 97 (8), 4124 - 4131. http: // dx. doi. org / 10.1073 / pnas. 97.8.4124" type="journal article" year="2000">
Liebig
<emphasis id="B9AA3402A6482C15FADCF90315F0F907" box="[1372,1429,1764,1789]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2000
</bibRefCitation>
), allowing low-ranking workers to detect sexual maturation by their nestmates and potentially prevent it through aggression (
<bibRefCitation id="EF4F95E1A6482C15FE01F8C01238F8C5" author="Liebig, J. &amp; Peeters, C. &amp; Holldobler, B." box="[385,605,1830,1855]" pageId="55" pageNumber="56" pagination="1865 - 1870" refId="ref151415" refString="Liebig, J., Peeters, C. &amp; Holldobler, B. (1999) Worker policing limits the number of reproductives in a ponerine ant. Proceedings of the Royal Society B, 266, 1865 - 1870. http: // dx. doi. org / 10.1098 / rspb. 1999.0858" type="journal article" year="1999">
Liebig
<emphasis id="B9AA3402A6482C15FE56F8C11274F8C5" box="[470,529,1830,1855]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 1999
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6482C15FDEBF8C11321F8C5" author="Hoyer, S. C. &amp; Liebig, J. &amp; Rossler, W." box="[619,836,1830,1855]" pageId="55" pageNumber="56" pagination="429 - 440" refId="ref147099" refString="Hoyer, S. C., Liebig, J. &amp; Rossler, W. (2005) Biogenic amines in the ponerine ant Harpegnathos saltator: serotonin and dopamine immunoreactivity in the brain. Arthropod Structure and Development, 34, 429 - 440. http: // dx. doi. org / 10.1016 / j. asd. 2005.03.003" type="journal article" year="2005">
Hoyer
<emphasis id="B9AA3402A6482C15FD3CF8C11292F8C5" box="[700,759,1830,1855]" italics="true" pageId="55" pageNumber="56">et al.</emphasis>
, 2005
</bibRefCitation>
). Since gamergates do not work as much as normal workers (as demonstrated by their reduced brain volumes:
<bibRefCitation id="EF4F95E1A6482C15FCA9F8A11400F89A" author="Gronenberg, W. &amp; Liebig, J." box="[809,1125,1863,1888]" pageId="55" pageNumber="56" pagination="343 - 345" refId="ref144811" refString="Gronenberg, W. &amp; Liebig, J. (1999) Smaller brains and optic lobes in reproductive workers of the ant Harpegnathos. Naturwissenschaften, 86, 343 - 345. http: // dx. doi. org / 10.1007 / s 001140050631" type="journal article" year="1999">Gronenberg &amp; Liebig, 1999</bibRefCitation>
), the number of gamergates in a queenless colony is tightly regulated to optimize reproduction versus productivity (
<bibRefCitation id="EF4F95E1A6482C15FBE6F88F15F5F878" author="Monnin, T. &amp; Ratnieks, F. L. W." box="[1126,1424,1897,1922]" pageId="55" pageNumber="56" pagination="97 - 108" refId="ref156027" refString="Monnin, T. &amp; Ratnieks, F. L. W. (2001) Policing in queenless ponerine ants. Behavioral Ecology and Sociobiology, 50, 97 - 108. http: // dx. doi. org / 10.1007 / s 002650100351" type="journal article" year="2001">Monnin &amp; Ratnieks, 2001</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6482C1AFF47F86C1515FE72" blockId="55.[151,1437,151,2027]" lastBlockId="56.[151,1437,151,1077]" lastPageId="56" lastPageNumber="57" pageId="55" pageNumber="56">
Colonies of most ponerine species die along with their queens, and new colonies arise via independent foundation by a fertilized alate queen (exceptions include species with ergatoid queens). In contrast,
<taxonomicName id="4CDE9393A6482C15FB7CF84815F9F83D" box="[1276,1436,1966,1991]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="55" pageNumber="56" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6482C15FB7CF84815F9F83D" box="[1276,1436,1966,1991]" italics="true" pageId="55" pageNumber="56">Harpegnathos</emphasis>
</taxonomicName>
emphasizes long-term stability (even immortality) of existing colonies. This likely stems from two related factors: the high mortality rates of new
<taxonomicName id="4CDE9393A6472C1AFE7AFF7112FFFF4A" box="[506,666,151,176]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFE7AFF7112FFFF4A" box="[506,666,151,176]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
colonies (
<bibRefCitation id="EF4F95E1A6472C1AFC8EFF711446FF4A" author="Liebig, J. &amp; Poethke, H." box="[782,1059,151,176]" pageId="56" pageNumber="57" pagination="203 - 207" refId="ref151554" refString="Liebig, J. &amp; Poethke, H. (2004) Queen lifespan and colony longevity in the ant Harpegnathos saltator. Ecological Entomology, 29, 203 - 207. http: // dx. doi. org / 10.1111 / j. 1365 - 2311.2004.00583. x" type="journal article" year="2004">Liebig &amp; Poethke, 2004</bibRefCitation>
), and their elaborate nests, which represent major investments of time and energy. Successful
<taxonomicName id="4CDE9393A6472C1AFCCBFF5A138EFF2F" box="[843,1003,188,213]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFCCBFF5A138EFF2F" box="[843,1003,188,213]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
queens typically only live for two or three years (
<bibRefCitation id="EF4F95E1A6472C1AFEA1FF39123EFF02" author="Peeters, C. &amp; Holldobler, B." box="[289,603,223,248]" pageId="56" pageNumber="57" pagination="10977 - 10979" refId="ref159484" refString="Peeters, C. &amp; Holldobler, B. (1995) Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest. Proceedings of the National Academy of Sciences (USA), 92, 10977 - 10979. http: // dx. doi. org / 10.1073 / pnas. 92.24.10977" type="journal article" year="1995">Peeters &amp; Hölldobler, 1995</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6472C1AFDE7FF39131FFF02" author="Liebig, J. &amp; Poethke, H." box="[615,890,223,248]" pageId="56" pageNumber="57" pagination="203 - 207" refId="ref151554" refString="Liebig, J. &amp; Poethke, H. (2004) Queen lifespan and colony longevity in the ant Harpegnathos saltator. Ecological Entomology, 29, 203 - 207. http: // dx. doi. org / 10.1111 / j. 1365 - 2311.2004.00583. x" type="journal article" year="2004">Liebig &amp; Poethke, 2004</bibRefCitation>
), but by extending the colonys lifespan through multiple generations of gamergates the colony can utilize the nests for longer periods and produce many more alate queens (
<bibRefCitation id="EF4F95E1A6472C1AFF78FEC11186FEBA" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[248,483,295,320]" pageId="56" pageNumber="57" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6472C1AFED6FECF11F6FEBA" box="[342,403,295,320]" italics="true" pageId="56" pageNumber="57">et al.</emphasis>
, 2000
</bibRefCitation>
). Colony emigrations and colony fissions are unknown in
<taxonomicName id="4CDE9393A6472C1AFB19FEC1155CFEBA" box="[1177,1337,295,320]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFB19FEC1155CFEBA" box="[1177,1337,295,320]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
and are unlikely, given the major investment represented by the nest and given that gamergate colonies emphasize production of new queens over workers (
<bibRefCitation id="EF4F95E1A6472C1AFDE1FE891359FE72" author="Peeters, C. &amp; Holldobler, B. &amp; Moffett, M. &amp; Musthak Ali, T. M." box="[609,828,367,392]" pageId="56" pageNumber="57" pagination="211 - 218" refId="ref159548" refString="Peeters, C., Holldobler, B., Moffett, M. &amp; Musthak Ali, T. M. (1994) &quot; Wall-papering &quot; and elaborate nest architecture in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 41, 211 - 218. http: // dx. doi. org / 10.1007 / bf 01240479" type="journal article" year="1994">
Peeters
<emphasis id="B9AA3402A6472C1AFD3BFE971296FE72" box="[699,755,367,392]" italics="true" pageId="56" pageNumber="57">et al.</emphasis>
, 1994
</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6472C1AFCC8FE96141BFE72" author="Peeters, C. &amp; Holldobler, B." box="[840,1150,367,392]" pageId="56" pageNumber="57" pagination="10977 - 10979" refId="ref159484" refString="Peeters, C. &amp; Holldobler, B. (1995) Reproductive cooperation between queens and their mated workers: the complex life history of an ant with a valuable nest. Proceedings of the National Academy of Sciences (USA), 92, 10977 - 10979. http: // dx. doi. org / 10.1073 / pnas. 92.24.10977" type="journal article" year="1995">Peeters &amp; Hölldobler, 1995</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6472C1AFB09FE961501FE72" author="Peeters, C. &amp; Liebig, J. &amp; Holldobler, B." box="[1161,1380,367,392]" pageId="56" pageNumber="57" pagination="325 - 332" refId="ref159644" refString="Peeters, C., Liebig, J. &amp; Holldobler, B. (2000) Sexual reproduction by both queens and workers in the ponerine ant Harpegnathos saltator. Insectes Sociaux, 47, 325 - 332. http: // dx. doi. org / 10.1007 / pl 00001724" type="journal article" year="2000">
Peeters
<emphasis id="B9AA3402A6472C1AFB61FE97157CFE72" box="[1249,1305,367,392]" italics="true" pageId="56" pageNumber="57">et al.</emphasis>
, 2000
</bibRefCitation>
).
</paragraph>
<paragraph id="8B61E810A6472C1AFF47FE721574FD37" blockId="56.[151,1437,151,1077]" pageId="56" pageNumber="57">
<emphasis id="B9AA3402A6472C1AFF47FE7212B5FE57" bold="true" box="[199,720,404,429]" pageId="56" pageNumber="57">Phylogenetic and taxonomic considerations.</emphasis>
<taxonomicName id="4CDE9393A6472C1AFD57FE721312FE57" box="[727,887,404,429]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFD57FE721312FE57" box="[727,887,404,429]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
was erected by
<bibRefCitation id="EF4F95E1A6472C1AFBAAFE7214A0FE57" author="Jerdon, T. C." box="[1066,1221,404,429]" pageId="56" pageNumber="57" pagination="103 - 127" refId="ref148048" refString="Jerdon, T. C. (1851) A catalogue of the species of ants found in Southern India. Madras Journal of Literature and Science, 17, 103 - 127." type="journal article" year="1851">Jerdon (1851)</bibRefCitation>
to house the single species
<taxonomicName id="4CDE9393A6472C1AFF6EFE5F1264FE2A" authority="Jerdon. Soon" authorityName="Jerdon. Soon" authorityYear="1851" box="[238,513,439,464]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="species" species="saltator">
<emphasis id="B9AA3402A6472C1AFF6EFE5F1102FE2A" box="[238,359,439,464]" italics="true" pageId="56" pageNumber="57">H. saltator</emphasis>
Jerdon. Soon
</taxonomicName>
afterward,
<bibRefCitation id="EF4F95E1A6472C1AFD01FE511373FE2A" author="Smith, F." box="[641,790,439,464]" pageId="56" pageNumber="57" 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>
replaced the genus name with
<taxonomicName id="4CDE9393A6472C1AFBF0FE51154EFE2A" box="[1136,1323,439,464]" class="Insecta" family="Formicidae" genus="Drepanognathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFBF0FE51154EFE2A" box="[1136,1323,439,464]" italics="true" pageId="56" pageNumber="57">Drepanognathus</emphasis>
</taxonomicName>
, based on some confusion over possible homonymy with a genus of staphilinid beetles (discussed by
<bibRefCitation id="EF4F95E1A6472C1AFB39FE3A15F5FE0F" author="Donisthorpe, H." box="[1209,1424,476,501]" pageId="56" pageNumber="57" pagination="196 - 201" refId="ref139520" refString="Donisthorpe, H. (1937) A new species of Harpegnathos Jerd., with some remarks on the genus, and other known species (Hym. Formicidae). Entomologist's Monthly Magazine, 73, 196 - 201." type="journal article" year="1937">Donisthorpe, 1937</bibRefCitation>
). Subsequent authors alternated between treating
<taxonomicName id="4CDE9393A6472C1AFD35FE191330FDE2" box="[693,853,511,536]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFD35FE191330FDE2" box="[693,853,511,536]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
or
<taxonomicName id="4CDE9393A6472C1AFCFDFE19145DFDE2" box="[893,1080,511,536]" class="Insecta" family="Formicidae" genus="Drepanognathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFCFDFE19145DFDE2" box="[893,1080,511,536]" italics="true" pageId="56" pageNumber="57">Drepanognathus</emphasis>
</taxonomicName>
as the valid name (
<emphasis id="B9AA3402A6472C1AFA9AFDE7152CFDE2" box="[1306,1353,513,536]" italics="true" pageId="56" pageNumber="57">e.g.,</emphasis>
<bibRefCitation id="EF4F95E1A6472C1AFAD2FDE610BBFDC7" author="Roger, J." pageId="56" pageNumber="57" pagination="131 - 214" refId="ref160939" refString="Roger, J. (1863 a) Die neu aufgefuhrten Gattungen und Arten meines Formiciden-Verzeichnisses nebst Erganzung einiger fruher gegebenen Beschreibungen. Berliner Entomologische Zeitschrift, 7, 131 - 214. http: // dx. doi. org / 10.1002 / mmnd. 47918630116" type="journal article" year="1863">Roger, 1863a</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6472C1AFF71FDC211EDFDC7" author="Forel, A." box="[241,392,548,573]" pageId="56" pageNumber="57" pagination="161 - 167" refId="ref142534" refString="Forel, A. (1893 a) Sur la classification de la famille des Formicides, avec remarques synonymiques. Annales de la Societe Entomologique de Belgique, 37, 161 - 167." type="journal article" year="1893">Forel, 1893a</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6472C1AFE1AFDC21215FDC7" author="Dalla Torre, K. W. von" box="[410,624,548,573]" pageId="56" pageNumber="57" refId="ref136742" refString="Dalla Torre, K. W. von (1893) Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 7. Formicidae (Heterogyna). W. Engelmann, Leipzig, 289 pp." type="book" year="1893">Dalla Torre, 1893</bibRefCitation>
;
<bibRefCitation id="EF4F95E1A6472C1AFD02FDC2135DFDC7" author="Bingham, C. T." box="[642,824,548,573]" pageId="56" pageNumber="57" refId="ref132962" refString="Bingham, C. T. (1903) The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps. Taylor and Francis, London, 506 pp." type="book" year="1903">Bingham, 1903</bibRefCitation>
), but eventually the proper consensus was reached recognizing the validity of the original name
<taxonomicName id="4CDE9393A6472C1AFD54FDA11416FD9A" authority="(Donisthorpe, 1937)" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[724,1139,583,608]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFD54FDA11311FD9A" box="[724,884,583,608]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
(
<bibRefCitation id="EF4F95E1A6472C1AFC0BFDA1140EFD9A" author="Donisthorpe, H." box="[907,1131,583,608]" pageId="56" pageNumber="57" pagination="196 - 201" refId="ref139520" refString="Donisthorpe, H. (1937) A new species of Harpegnathos Jerd., with some remarks on the genus, and other known species (Hym. Formicidae). Entomologist's Monthly Magazine, 73, 196 - 201." type="journal article" year="1937">Donisthorpe, 1937</bibRefCitation>
)
</taxonomicName>
.
<taxonomicName id="4CDE9393A6472C1AFB08FDA1154DFD9A" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[1160,1320,583,608]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFB08FDA1154DFD9A" box="[1160,1320,583,608]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
has also experienced some instability at the tribal level, being variously placed in
<taxonomicName id="4CDE9393A6472C1AFC60FD8A145AFD7F" box="[992,1087,620,645]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
(
<emphasis id="B9AA3402A6472C1AFBD1FD8B14E5FD7E" box="[1105,1152,621,644]" italics="true" pageId="56" pageNumber="57">e.g.,</emphasis>
<bibRefCitation id="EF4F95E1A6472C1AFB09FD8A1556FD7F" author="Wheeler, W. M." box="[1161,1331,620,645]" pageId="56" pageNumber="57" refId="ref167610" refString="Wheeler, W. M. (1910) Ants: their structure, development and behavior. Columbia University Press, New York, xxv + 663 pp." type="book" year="1910">Wheeler, 1910</bibRefCitation>
), its own tribe
<taxonomicName id="4CDE9393A6472C1AFF51FD691106FD52" box="[209,355,655,680]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Harpegnathii">Harpegnathii</taxonomicName>
(
<bibRefCitation id="EF4F95E1A6472C1AFEF2FD691263FD52" author="Forel, A." box="[370,518,655,680]" pageId="56" pageNumber="57" pagination="52 - 65" refId="ref142801" refString="Forel, A. (1900 b) Les Formicides de l'Empire des Indes et de Ceylan. Part VI. Journal of the Bombay Natural History Society, 13, 52 - 65." type="journal article" year="1900">Forel, 1900b</bibRefCitation>
;
<taxonomicName id="4CDE9393A6472C1AFD91FD6912B7FD52" authorityName="Ashmead" authorityYear="1905" box="[529,722,655,680]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Drepanognathini">Drepanognathini</taxonomicName>
:
<bibRefCitation id="EF4F95E1A6472C1AFD5CFD6913EBFD52" author="Ashmead, W. H." box="[732,910,655,680]" pageId="56" pageNumber="57" pagination="381 - 384" refId="ref131809" refString="Ashmead, W. H. (1905) A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea. Canadian Entomologist, 37, 381 - 384. http: // dx. doi. org / 10.4039 / ent 37381 - 11" type="journal article" year="1905">Ashmead, 1905</bibRefCitation>
), or its own subtribe
<taxonomicName id="4CDE9393A6472C1AFBFAFD69157FFD52" box="[1146,1306,655,680]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Harpegnathini">Harpegnathini</taxonomicName>
in
<taxonomicName id="4CDE9393A6472C1AFABDFD6915F9FD52" box="[1341,1436,655,680]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
(
<emphasis id="B9AA3402A6472C1AFF20FD5310AAFD36" box="[160,207,693,716]" italics="true" pageId="56" pageNumber="57">e.g.,</emphasis>
<bibRefCitation id="EF4F95E1A6472C1AFF56FD52110DFD37" author="Emery, C." box="[214,360,692,717]" pageId="56" pageNumber="57" 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>
).
<bibRefCitation id="EF4F95E1A6472C1AFEFCFD521275FD37" author="Smith, F." box="[380,528,692,717]" pageId="56" pageNumber="57" pagination="285 - 348" refId="ref162750" refString="Smith, F. (1871) A catalogue of the Aculeate Hymenoptera and Ichneumonidae of India and the Eastern Archipelago. With introductory remarks by A. R. Wallace. [part]. Journal of the Proceedings of the Linnean Society. Zoology, 11, 285 - 348. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.1871. tb 02225. x" type="journal article" year="1871">Smith (1871)</bibRefCitation>
even grouped it with
<taxonomicName id="4CDE9393A6472C1AFC85FD5213CAFD37" box="[773,943,692,717]" class="Insecta" family="Formicidae" genus="Odontomachus" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFC85FD5213CAFD37" box="[773,943,692,717]" italics="true" pageId="56" pageNumber="57">Odontomachus</emphasis>
</taxonomicName>
in his family
<taxonomicName id="4CDE9393A6472C1AFBCAFD521568FD37" authorityName="Mayr" authorityYear="1862" box="[1098,1293,692,717]" class="Insecta" family="Odontomachidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="family">Odontomachidae</taxonomicName>
.
</paragraph>
<paragraph id="8B61E810A6472C1AFF47FD3113C7FBCF" blockId="56.[151,1437,151,1077]" pageId="56" pageNumber="57">
Both molecular and morphological evidence support the placement of
<taxonomicName id="4CDE9393A6472C1AFC5CFD311419FD0A" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[988,1148,727,752]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFC5CFD311419FD0A" box="[988,1148,727,752]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
in its own “genus group” within
<taxonomicName id="4CDE9393A6472C1AFF6BFD1A1128FCEF" box="[235,333,764,789]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
.
<bibRefCitation id="EF4F95E1A6472C1AFEDCFD1A1244FCEF" author="Schmidt, C." box="[348,545,764,789]" pageId="56" pageNumber="57" 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="4CDE9393A6472C1AFCFDFD1A1388FCEF" baseAuthorityName="sensu Bolton" baseAuthorityYear="2003" box="[893,1005,764,789]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
confirms that
<taxonomicName id="4CDE9393A6472C1AFB1FFD1A155AFCEF" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[1183,1343,764,789]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFB1FFD1A155AFCEF" box="[1183,1343,764,789]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
is very closely related to the rest of
<taxonomicName id="4CDE9393A6472C1AFE51FCF91256FCC2" box="[465,563,799,824]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
, if not deeply nested within the tribe. The exact placement of
<taxonomicName id="4CDE9393A6472C1AFB5FFCF9151AFCC2" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[1247,1407,799,824]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFB5FFCF9151AFCC2" box="[1247,1407,799,824]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
in the phylogeny of
<taxonomicName id="4CDE9393A6472C1AFEDDFCA211A8FCA7" baseAuthorityName="sensu Bolton" baseAuthorityYear="2003" box="[349,461,836,861]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="subFamily" subFamily="Ponerinae">Ponerinae</taxonomicName>
is unresolved, possibly due to long branch issues (
<bibRefCitation id="EF4F95E1A6472C1AFB89FCA214D6FCA7" author="Schmidt, C." box="[1033,1203,836,861]" pageId="56" pageNumber="57" 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>
). The best supported phylogenetic position of
<taxonomicName id="4CDE9393A6472C1AFE2DFC811228FC7A" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[429,589,871,896]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFE2DFC811228FC7A" box="[429,589,871,896]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
is as sister to the remainder of the
<taxonomicName id="4CDE9393A6472C1AFC54FC811453FC7A" box="[980,1078,871,896]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
, but other placements cannot be statistically rejected. Commonly resolved alternative placements include as sister to
<emphasis id="B9AA3402A6472C1AFBCAFC6B14B8FC5E" box="[1098,1245,909,932]" italics="true" pageId="56" pageNumber="57">
<taxonomicName id="4CDE9393A6472C1AFBCAFC6B14BCFC5E" box="[1098,1241,909,932]" class="Insecta" family="Formicidae" genus="Hypoponera" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">Hypoponera</taxonomicName>
,
</emphasis>
the
<taxonomicName id="4CDE9393A6472C1AFA8EFC6A15F9FC5F" box="[1294,1436,908,933]" class="Insecta" family="Formicidae" genus="Plectroctena" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFA8EFC6A15F9FC5F" box="[1294,1436,908,933]" italics="true" pageId="56" pageNumber="57">Plectroctena</emphasis>
</taxonomicName>
group, or
<taxonomicName id="4CDE9393A6472C1AFE86FC5711F7FC32" box="[262,402,945,968]" class="Insecta" family="Formicidae" genus="Hypoponera" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFE86FC5711F7FC32" box="[262,402,945,968]" italics="true" pageId="56" pageNumber="57">Hypoponera</emphasis>
</taxonomicName>
plus the
<taxonomicName id="4CDE9393A6472C1AFE7BFC4912ECFC32" box="[507,649,943,968]" class="Insecta" family="Formicidae" genus="Plectroctena" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFE7BFC4912ECFC32" box="[507,649,943,968]" italics="true" pageId="56" pageNumber="57">Plectroctena</emphasis>
</taxonomicName>
group. Morphological data suggests that
<taxonomicName id="4CDE9393A6472C1AFBDEFC49149BFC32" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[1118,1278,943,968]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFBDEFC49149BFC32" box="[1118,1278,943,968]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
may be sister to the remainder of
<taxonomicName id="4CDE9393A6472C1AFE00FC321187FC17" box="[384,482,980,1005]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="tribe" tribe="Ponerini">Ponerini</taxonomicName>
, though it is still an open question. The phylogenetic position of
<taxonomicName id="4CDE9393A6472C1AFB5FFC32151AFC17" baseAuthorityName="Donisthorpe" baseAuthorityYear="1937" box="[1247,1407,980,1005]" class="Insecta" family="Formicidae" genus="Harpegnathos" kingdom="Animalia" order="Hymenoptera" pageId="56" pageNumber="57" phylum="Arthropoda" rank="genus">
<emphasis id="B9AA3402A6472C1AFB5FFC32151AFC17" box="[1247,1407,980,1005]" italics="true" pageId="56" pageNumber="57">Harpegnathos</emphasis>
</taxonomicName>
is arguably the single most important outstanding question in ponerine phylogenetics, given its implications for reconstructions of ancestral ponerine structure, ecology and behavior.
</paragraph>
</subSubSection>
</treatment>
</document>