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<document id="4A9BA495763E20E3535ACB16814F2753" ENCODING="UTF8" ID-GBIF-Dataset="b542bb57-07e0-4b2e-952b-0bc5e9697698" ModsDocAuthor="Rabeling, Ch. &amp; Bacci, M." ModsDocDate="2010" ModsDocID="23066" ModsDocOrigin="http://antbase.org/ants/publications/23066/23066.pdf" ModsDocTitle="A new workerless inquiline in the Lower Attini (Hymenoptera: Formicidae), with a discussion of social parasitism in fungus-growing ants." checkinTime="1276014642463" checkinUser="donat" docAuthor="Rabeling, Ch. &amp; Bacci, M." docDate="2010" docId="0DA25152360E22A404D9ADD40B0ABC2C" docLanguage="en" docName="23066.htm" docOrigin="Systematic Entomology 35" docSource="http://antbase.org/ants/publications/23066/23066.pdf" docTitle="Mycocepurus castrator Rabeling &amp; Bacci, sp.n." docType="treatment" docVersion="11" lastPageNumber="391" masterDocId="7C2DE219932C33DE76A8E791BCAA74F7" masterDocTitle="A new workerless inquiline in the Lower Attini (Hymenoptera: Formicidae), with a discussion of social parasitism in fungus-growing ants." masterLastPageNumber="392" masterPageNumber="379" pageNumber="382" updateTime="1692406703124" updateUser="admin">
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<mods:title id="DB11D33EC3A5CF796EAB90D76868522B">A new workerless inquiline in the Lower Attini (Hymenoptera: Formicidae), with a discussion of social parasitism in fungus-growing ants.</mods:title>
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<mods:namePart id="797EFE910A04D54D2D01C410F9EBE5DC">Rabeling, Ch.</mods:namePart>
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<treatment id="0DA25152360E22A404D9ADD40B0ABC2C" ID-DOI="http://doi.org/10.5281/zenodo.6196567" ID-GBIF-Taxon="100116218" ID-Zenodo-Dep="6196567" LSID="urn:lsid:plazi:treatment:0DA25152360E22A404D9ADD40B0ABC2C" httpUri="http://treatment.plazi.org/id/0DA25152360E22A404D9ADD40B0ABC2C" lastPageNumber="391" pageNumber="382">
<subSubSection id="9DBE1B6A8F87554337C4435BC44A9C0A" type="nomenclature">
<paragraph id="0E5C8252B4F200D6930AD533D427E650" pageNumber="382">
<taxonomicName id="ACB4DA95DBAFFCE0B370FB8E1C6C6D40" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" authority="Rabeling &amp; Bacci" authorityName="Rabeling &amp; Bacci" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="382" phylum="Arthropoda" rank="species" species="castrator" status="sp.n.">Mycocepurus castrator Rabeling &amp; Bacci</taxonomicName>
,
<taxonomicNameLabel id="EB53373C26FC21501347AA1CB5159C26" rank="species" rankGroup="species" rankGroupNumber="3" rankNumber="3">sp.n.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="012E883F6FF36490DB9DC9A65DD04EB0" type="description">
<paragraph id="0BFE6E15126833D3254BCCEC1B857EF3" pageNumber="382">(Figs 1A, C, E, G; 2A, C, E, G)</paragraph>
</subSubSection>
<subSubSection id="2A6EE6F1559E4B468D92A6383C42CCDC" type="materials_examined">
<paragraph id="89E9BA4341D533EEC77B43BE49C59559" pageNumber="382">
<materialsCitation id="0696374036736E787859F2480A9D8F2B" ID-GBIF-Occurrence="923903319" collectingDate="2006-09-29" collectionCode="UNESP, MZSP" collectorName="C. Rabeling" country="BRAZIL" latitude="-22.3955" location="São Paulo, Rio Claro, Campus of São Paulo State University" longitude="-47.5424" specimenCode="CR 060929 - 14" typeStatus="Holotype">
<typeStatus id="F3C5571329D69C13B49E14FB70015584">Holotype</typeStatus>
, ♀,
<collectingCountry id="6E6D6785602D1619A7B803794031F9B7">BRAZIL</collectingCountry>
:
<location id="453BA97CE2EA0B802531AF3B1381F4D1" LSID="urn:lsid:plazi:treatment:0DA25152360E22A404D9ADD40B0ABC2C:453BA97CE2EA0B802531AF3B1381F4D1" country="BRAZIL" latitude="-22.3955" longitude="-47.5424" name="São Paulo, Rio Claro, Campus of São Paulo State University">São Paulo, Rio Claro, Campus of São Paulo State University</location>
(
<collectionCode id="0DEAE0B5E377FCA41E4409769C499E95" collectionName="UNESP">UNESP</collectionCode>
),
<geoCoordinate id="A18D980E0065F679F0B690F37F79AA59" direction="south" orientation="latitude" value="-22.3955">22.3955º S</geoCoordinate>
,
<geoCoordinate id="607EF24A85F17B659053C606FEB397DB" direction="west" orientation="longitude" value="-47.5424">047.5424ºW</geoCoordinate>
, elevation 608 m,
<date id="EB56E6127AC65B5DB64D34AC2183B479" day="29" month="09" value="2006-09-29" year="2006">
<collectingDate id="6A10B7106675B08E53805A04CC33CBD6" day="29" month="09" value="2006-09-29" year="2006">29.ix.2006</collectingDate>
</date>
,
<collectorName id="E1AE8CD546FD79E1F95949EC2B53CA5A">C. Rabeling</collectorName>
acc. no.
<specimenCode id="09178F41E91F0C6C20EE6E1C8D1F049F">
CR
<date id="D6190FBFE2F606F48A81EE80BCF32B46" day="06" month="09" value="29-09-06" year="29">060929</date>
- 14
</specimenCode>
, ex
<taxonomicName id="5C17FF8B7047BFEEF27AC4B643E4EBA5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="382" phylum="Arthropoda" rank="species" species="goeldii">Mycocepurus goeldii</taxonomicName>
nest.
<typeStatus id="2CFF5D6E95A64175FE19F8CF2E694A65">Holotype</typeStatus>
deposited at
<collectionCode id="70199E3906D08F725EEDB9962499CE5C" collectionName="Brazil, Sao Paulo, Sao Paulo, Museu de Zoologia da Universidade de Sao Paulo">MZSP</collectionCode>
. Measurements (in millimetres): HW 0.6, HL 0.64, SL 0.76, WL 1.07, PPW 0.62, PW 0.21, PL 0.24, PPL 0.19, CI 94, SI 127
</materialsCitation>
.
</paragraph>
<paragraph id="4E22547EF4948A4243974E07E4A20774" pageNumber="382">
<materialsCitation id="AEDD348ADA1C6CACAAEA8A1073DA5D34" ID-GBIF-Occurrence="923903320" collectingDate="2006-09-29" collectionCode="AMNH, BMEL, CRC, MCZC, MZSP, UCDC, USNM" collectorName="col. C. Rabeling" country="BRAZIL" location="BRAZIL" typeStatus="Paratypes">
<typeStatus id="B5A385C12E634BD1D1E39FCBF6E44052">Paratypes</typeStatus>
, 104 ♀, 78 ♂,
<collectingCountry id="4A464C2E073924DAD6FEDC5485F87F84">BRAZIL</collectingCountry>
: same nest as holotype,
<date id="824D1CCDDB18A0F7180028C9D5FD44A5" day="29" month="09" value="2006-09-29" year="2006">
<collectingDate id="04DFD32CB0E1662CE6EBD7939E94131D" day="29" month="09" value="2006-09-29" year="2006">29.ix.2006</collectingDate>
</date>
-
<date id="ADCB17D290A4FAC02BF7C099B21D17CD" day="02" month="10" value="2006-10-02" year="2006">
<collectingDate id="DD49FE9F8C96DBA75EB6C44568138CA9" day="02" month="10" value="2006-10-02" year="2006">02.x.2006</collectingDate>
</date>
,
<collectorName id="62E4D0AEDC4FB69867B755E522A62989">col. C. Rabeling</collectorName>
.
<typeStatus id="B6382AC36FF3C669127F5BB9D906EDF1">Paratypes</typeStatus>
deposited at:
<collectionCode id="0292FBBC538D8FA0E8C8630FAD2A663F" collectionName="USA, New York, New York, American Museum of Natural History">AMNH</collectionCode>
,
<collectionCode id="2B4EFA0CD743E7126414B90AC3E7DDD1" collectionName="BMEL">BMEL</collectionCode>
,
<collectionCode id="AE7AA03219D88ED188D523BB3AAD6712" collectionName="CRC">CRC</collectionCode>
,
<collectionCode id="ABF08926A6908540E0C128A8D0750EBF" collectionName="USA, Massachusetts, Cambridge, Harvard University, Museum of Comparative Zoology">MCZC</collectionCode>
,
<collectionCode id="DA71AE3A3D4D8E7868F0B9CC51E4F7D3" collectionName="Brazil, Sao Paulo, Sao Paulo, Museu de Zoologia da Universidade de Sao Paulo">MZSP</collectionCode>
,
<collectionCode id="6718222FE2F97CBC881354E5242E4231" collectionName="USA, California, Davis, University of California, R.M. Bohart Museum of Entomology">UCDC</collectionCode>
,
<collectionCode id="C53F710A97C627E848ACBB523B63EE0E" collectionName="USA, Washington D.C., National Museum of Natural History, [formerly, United States National Museum]">USNM</collectionCode>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="27D80C8EC8DFDE2EA5F346A1F375FA92" type="diagnosis">
<paragraph id="B93C38889202F6F074D65BB36A521ED5" pageNumber="382">
<typeStatus id="595686A38EB215E440C62A485663C97C">Holotype</typeStatus>
, ♀ (queen). Diagnosis. Small species (WL 1.07) with a unique morphology reflecting the parasitic life history. In full face view, head rectangular (CI 94); sides approximately parallel, slightly tapering above mandibular insertions; head widest directly above the eyes; posterior margin of the head heart shaped, with a slight but distinct median concavity; posterolateral corners rounded, in lateral view drawn out to form a short, rounded lobe forming the ventrolateral corner of the head. Antennae with 11 segments; antennal scapes extremely long (SL 0.76), surpassing the posterior margin of the head by nearly half their length (SI 127). In full face view, frontal carinae and antennal scrobes absent. Frontal lobes small and rounded, barely covering the antennal sockets in frontal view. Median triangular portion of clypeus raised
</paragraph>
<paragraph id="E41F537AE2F5AF3B036DFE51392BCCC0" pageNumber="382">
between the antennal insertions. Mandibles reduced, narrow, elongate, blade-like terminating in a pointed tooth; otherwise lacking teeth except for a small basal denticle. Maxillary palps reduced, with only three segments, labial palps with two segments. Ocelli slightly raised above the surface of vertex. Mesosoma with characteristic morphology related to wing bearing. Pronotal spines absent; propodeal spines well developed, stout, as wide as long at the base and sharply pointed; metapleural gland orifice very large and circular in oblique view, ventral margin forming small, vertical tooth. Petiole with a short peduncle; node triangular in side view, with sharp crest terminating in two thick pointed teeth. In dorsal view, postpetiole approximately 3× as wide as long (PPL 0.19, PPW 0.62); lateral borders tapering into pointed angles; translucent area near posterior margin forming broad u-shaped invagination. First gastric tergite strikingly concave in lateral view. Entire body surface more or less smooth and shiny, in most areas with hexagonal microsculpture resembling a honeycomb. Body sparsely covered with stiff setae; setae erect on vertex and frontal lobes, sub-decumbent on mesoscutum and scutellum, and appressed on postpetiole and metasoma. Wings infuscated with reduced venation, densely covered with setae; clear spot or fenestra in apical part of forewing absent; rsf1 faint, hardly visible. Colour: light to dark reddishbrown. -
<typeStatus id="E4CD1953E454FBA26C04F1759EC1448E">Paratype</typeStatus>
♀♀. Measurements (n = 15). HW 0.6-0.65, HL 0.63-0.64, SL 0.73-0.8, WL 1.07-1.23, PPW 0.62-0.65, PW 0.21-0.25, PL 0.24-0.28, PPL 0.18-0.2, CI 94-104, SI 115-128.
</paragraph>
<paragraph id="91C80D2CFFECC992A2648D7F6A6FA11F" lastPageNumber="385" pageNumber="382">
<typeStatus id="1E3F9493DA51F86387332FF869F92FE0">Paratype</typeStatus>
♂♂ (males). Diagnosis. Remarkably similar to female, not resembling any other
<taxonomicName id="D7C7C369932AC144A6A78E3D589A40D7" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2393" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus Forel" order="Hymenoptera" pageNumber="382" phylum="Arthropoda" rank="genus">Mycocepurus</taxonomicName>
male; characters as in female diagnosis with the following exceptions: head size of males smaller (HL 0.58-0.6, HW 0.58-0.6), whereas body length similar (WL 1.1-1.2). Mandibles reduced, narrow, elongate, blade-like, which do not terminate in a pointed tooth; otherwise lacking any teeth or denticles. Number of antennal segments reduced to 11; funicular segments approximately as long as broad, slowly increasing in length towards apex to 1.5× their width, only apical segment 5× as long as
<pageBreakToken id="637FC757ABDF246C05E4112467BA3901" pageNumber="385">wide</pageBreakToken>
. Mesosoma lower and narrower; tiny opening present at the metapleuron, corresponding to the position of a metapleural gland opening in the female. First gastric tergite flat to slightly concave; male genitalia projecting forward from tip of metasoma. Basal apodeme lobed, separated from aedeagus by a deep constriction; ventral border lacking serration. Wing colour: medium to dark brown. Measurements (n = 15). HW 0.58-0.6, HL 0.58-0.6, SL 0.73-0.75, WL 1.1-1.2, PPW 0.63-0.65, PW 0.23-0.3, PL 0.25-0.28, PPL 0.18-0.2, CI 96-104, SI 121-126.
</paragraph>
<footnote id="1CABDF2DD81B20C4C91AD9757F693AC6">
<paragraph id="D9F10EFB11C7202CB17742D19413ADA4" pageNumber="382">© 2010 The Authors Journal compilation © 2010 The Royal Entomological Society, Systematic Entomology, 35, 379-392</paragraph>
</footnote>
<caption id="CECC6B05BA3D584B0D338A9868DA8784">
<paragraph id="3021714DBA3D8070320F2FD5B41F7AA8" pageNumber="383">
Fig. 1. Queens of
<taxonomicName id="5F0B70F0071E294853F18447333083FA" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="383" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
(A, C, E, G) and
<taxonomicName id="CE99CECDE66C5277A7C3B18043B8A9FD" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="383" phylum="Arthropoda" rank="species" species="goeldii">Mycocepurus goeldii</taxonomicName>
(B, D, F, H). (A, B) Habitus in lateral view; scale bar represents 0.5 cm. (C, D) Posterior part of the mesosoma, petiole and postpetiole in lateral view; scale bar represents 0.5 cm. (E, F) Metapleural gland orifice in oblique view, metacoxa is visible in the lower left corner of the image; scale bar represents 50 μm. (G, H) Maxillary palp, scale bar represents 50 μm.
</paragraph>
</caption>
<footnote id="FFAD5319981964D84D673A24EA8E9513">
<paragraph id="BA86EC2E311106218B87746947B46D87" pageNumber="383">© 2010 The Authors</paragraph>
</footnote>
<footnote id="4ABF1A4865E9F60969EF28AC54841035">
<paragraph id="8863F1576475684DFD9DEDE119942FAD" pageNumber="383">Journal compilation © 2010 The Royal Entomological Society, Systematic Entomology, 35, 379-392</paragraph>
</footnote>
<caption id="1AE0CAFB3BB7C37AC3DF130B9C7E65B5">
<paragraph id="154EE0B3DCF3E4FF5CE89BC9A977694E" pageNumber="384">
Fig. 2. Males of
<taxonomicName id="B1055D1C17B8C033A5FD87F5BB18E8AF" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="384" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
(A, C, E, G) and
<taxonomicName id="CA87E954D3454CBECD684E1C9A231726" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="384" phylum="Arthropoda" rank="species" species="goeldii">Mycocepurus goeldii</taxonomicName>
(B, D, F, H). (A, B) Habitus in lateral view; scale bar represents 0.5 cm. (C, D) Posterior part of the mesosoma, petiole and postpetiole in lateral view; scale bar represents 0.5 cm. (E, F) Metapleural orifice in oblique view, metacoxa is visible in the lower left corner of the image; scale bar represents 50 μm. (G, H) Maxillary palp, scale bar represents 50 μm.
</paragraph>
</caption>
<footnote id="267807F09C5A59666A493334496E5584">
<paragraph id="3C77ABF0032A3DA3766BF843DF44766E" pageNumber="384">© 2010 The Authors Journal compilation © 2010 The Royal Entomological Society, Systematic Entomology, 35, 379-392</paragraph>
</footnote>
<paragraph id="683BC8506A8051164F2A5156D948F3E1" pageNumber="385">Worker. The worker caste is unknown and probably nonexistent.</paragraph>
</subSubSection>
<subSubSection id="FD7A6CEEE89134A7A59AE2E87BDDAFD1" type="materials_examined">
<paragraph id="E59CE8145FD5FC3AECC60D902AAB723B" pageNumber="385">
Additional material examined. BRAZIL: Sa˜o Paulo, Rio Claro, Campus of
<normalizedToken id="08F0404ADCCED01EFC6E349941A6530E" originalValue="São">Sao</normalizedToken>
Paulo State University (
<collectionCode id="C1087FD0C2436ED6A4D72557F8888CDD" collectionName="UNESP">UNESP</collectionCode>
), 22.3955◦S, 047.5424◦W, elevation 608 m,
<date id="09934E6CFE95693A9AF5FDD7933020C8" day="03" month="10" value="2008-10-03" year="2008">03.x.2008</date>
, C. Rabeling acc. no. CR
<date id="0B28F742FB575ACE47CF0F173E73CD04" day="08" month="10" value="03-10-08" year="03">081003</date>
-01, CR
<date id="D7F2F6D226BA236E99F9ACE4CE79D79A" day="08" month="10" value="03-10-08" year="03">081003</date>
-02, CR
<date id="789A92D56D90D425319C8B40526D8AE7" day="08" month="10" value="03-10-08" year="03">081003</date>
-03, CR
<date id="7B4B3192C1CC4A77704C80ED3CF4CDC5" day="08" month="10" value="03-10-08" year="03">081003</date>
-04, CR
<date id="BBAEBBFC2481007699592B86ABF6BB25" day="08" month="10" value="03-10-08" year="03">081003</date>
-05; ex
<taxonomicName id="1467F745BA253F4D13C4F6BD7A7ADA06" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">Mycocepurus goeldii</taxonomicName>
nest.
</paragraph>
</subSubSection>
<subSubSection id="DD57CCD5C977B14E6D7B1C50F46B9609" type="discussion">
<paragraph id="69C35A84A52147C8CDABBAB76BF01DF9" pageNumber="385">
Comments.
<taxonomicName id="1B243897939C2C882550A837999E4FEE" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
is an obligate, workerless social parasite of
<taxonomicName id="98DBB5D8B08E71C8318BADE62E433833" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
and is so far known only from Rio Claro, Sa˜o Paulo State, Brazil.
<taxonomicName id="0083B6B99E99B6C34137A6D44ECFA673" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
occurs sympatrically with
<taxonomicName id="92B2EFB8EF9EE9942859435CABE6A322" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31781" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus smithii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="smithii">M. smithii</taxonomicName>
and
<taxonomicName id="8383DC7CC912BCED0A697F11A5930D65" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31780" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus obsoletus Emery" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="obsoletus">M. obsoletus</taxonomicName>
, but cannot be confounded with any other
<taxonomicName id="974D954823F081F25AC938D75204B048" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2393" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus Forel" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="genus">Mycocepurus</taxonomicName>
species because of its multiple morphological adaptations for a parasitic lifestyle (Table 3).
<taxonomicName id="724265105FDB04F56E5AA28247AD0A92" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
can be recognized by the following characteristics: (i) the long antennal scapes surpassing the posterior margin of the head by half their length; (ii) reduced, blade-like mandibles lacking dentition of masticatory margin; (iii) concave shape of first gastric tergite; (iv) smooth and shiny body sculpture with hexagonal microsculpture; (v) reduced palpal formula (3,2); (vi) females and males with 11 antennal segments; (vii) males lacking serrated ventral border of aedeagus; (viii) absence of clear fenestra from forewings of queens and males; (ix) worker caste presumably absent; (x) metapleural gland orifice enlarged in females, and potentially present in males.
<taxonomicName id="46C3D2E898BAAE3EB50E05FBECDE43EE" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator males</taxonomicName>
and females look extremely similar to each other, and males are distinguished most easily from the females by the genitalia protruding from the tip of the metasoma and their darker brown colour (vs reddish brown in the queens).
</paragraph>
</subSubSection>
<subSubSection id="3692FF97B6C4CA6BC5A21424C5FD79C5" type="etymology">
<paragraph id="818AD889F3B46249B76C574A47DB6A3B" pageNumber="385">
Etymology. During collections of
<taxonomicName id="EED784948A440CB3A4FD960F459EB4A5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
, the host colonies were not observed to produce any alate queens and males, although sympatrically nesting
<taxonomicName id="058F5E2B9A078C6ECE93F58EDC9B6DE2" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colonies released alates. Therefore, we assume that the inquiline inhibits the host queens ' production of sexual offspring, allowing only for the production of the sterile worker caste. This is essentially social castration ', hence the specific name
<taxonomicName id="A5D6BD913739CD3A868CCF5D1BD27C5B" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="castrator">castrator</taxonomicName>
'.
</paragraph>
</subSubSection>
<subSubSection id="1C739941E4CA0B25D48F037ED986A9A3" type="biology_ecology">
<paragraph id="4A0300C1BC904248766397CE1A3BC8E1" pageNumber="385">
Host species.
<taxonomicName id="E2F262D3C92065D47ED254C7D233530D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
has been found only in nests of
<taxonomicName id="E2081FF205EE2759D395F8BDA773722D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
and is so far only known from the type locality (Rio Claro, SP).
<taxonomicName id="89873016B93CEF4C39AD4BF003EE74FC" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">Mycocepurus goeldii</taxonomicName>
is a conspicuous, widely distributed species ranging approximately from the 40th to the 67th meridian west and from the 2nd to the 31st latitude south, an area covering most of Brazil, parts of Bolivia, Paraguay and northern
<taxonomicName id="92F15E595939CD0C0C209560E4145D71" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">Argentina</taxonomicName>
. The range of habitats occupied by
<taxonomicName id="3242B32A20062E6A2D1D1E25A8A0870C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
is remarkably diverse and ranges from Amazon rainforest, savannahs (Cerrado) to the fertile South American lowlands (Pampas), and secondary habitats disturbed by human activities. It does not occur in elevated sites of the South American Cordilleras.
<taxonomicName id="BF90C16DF83C34A794985A122FE5A3D0" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">Mycocepurus goeldii</taxonomicName>
workers can be distinguished clearly from its congeners based on the size and spine pattern of the mesosoma: it is the largest species in the genus and has the most complete set of spine pairs on the mesosoma (
<bibRefCitation id="EA5C81960CC9D9633FE62E610D1EC8F7" author="Kempf, W. W." firstAuthor="Kempf" journalOrPublisher="Studia Entomologica" pagination="417 - 432" refId="ref11634" refString="Kempf, W. W. (1963) A review of the ant genus Mycocepurus Forel, 1893 (Hymenoptera: Formicidae). Studia Entomologica, 6, 417 - 432." title="A review of the ant genus Mycocepurus Forel, 1893 (Hymenoptera: Formicidae)" volume="6" year="1963">Kempf, 1963</bibRefCitation>
: figs 2, 3). The natural history of this species has been studied near Sa˜o Paulo City (
<bibRefCitation id="DF7CDC249A64C886C907402CCEA8D563" author="Luederwaldt, H." firstAuthor="Luederwaldt" journalOrPublisher="Revista Museu Paulista" pagination="29 - 64" refId="ref11773" refString="Luederwaldt, H. (1918) Notas myrmecologicas. Revista Museu Paulista, 10, 29 - 64." title="Notas myrmecologicas" volume="10" year="1918">Luederwaldt, 1918</bibRefCitation>
,
<bibRefCitation id="B7709B068E12354F0F932792090FECDB" author="Luederwaldt, H." firstAuthor="Luederwaldt" journalOrPublisher="Revista Museu Paulista" pagination="185 - 304" refId="ref11793" refString="Luederwaldt, H. (1926) Observac ¸ o ˜ es biolo ' gicas sobre formigas brasileiras, especialmente do estado de Sa ˜ o Paulo. Revista Museu Paulista, 14, 185 - 304." title="Observac ¸ o ˜ es biolo ' gicas sobre formigas brasileiras, especialmente do estado de Sa ˜ o Paulo" volume="14" year="1926">1926</bibRefCitation>
) and in the Manaus region of the Amazon Basin (
<bibRefCitation id="2ACA5149C362841EAC07BEBD5A4BD548" author="Rabeling, C." etAl="et al." firstAuthor="Rabeling" journalOrPublisher="Journal of Insect Science" refId="ref12022" refString="Rabeling, C., Verhaagh, M. &amp; Engels, W. (2007 b) Comparative study of nest architecture and colony structure of the fungus-growing ants, Mycocepurus goeldii and M. smithii. Journal of Insect Science, 7, 40. Available at: www. insectscience. org / 7.40" title="Comparative study of nest architecture and colony structure of the fungus-growing ants, Mycocepurus goeldii and M. smithii" volume="7, 40" year="2007" yearSuffix="b">Rabeling et al., 2007b</bibRefCitation>
), but these studies do not report the presence of a social parasite attacking
<taxonomicName id="7386C2208DD6B860A0B5D147EC499BD4" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
. Like most inquilines for which we have data,
<taxonomicName id="8FE8B23DB8D1983231BF2F2B3F7B72A3" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
probably has a patchy and locally restricted distribution. In addition, it is probable that
<taxonomicName id="1E1A3D90118639A582CE606EFBC0F88E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
is host specific, occurring only in nests of
<taxonomicName id="0014825A31B9F111DFB6D414E70A881B" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
. Despite extensive excavation of nests of sympatrically occurring
<taxonomicName id="7574BFF3F24A1434AC714722C8E04B07" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2393" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus Forel" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="genus">Mycocepurus</taxonomicName>
species, the parasite was never encountered in the nests of
<taxonomicName id="96600F13F99259CB2CDF3D8366EF763F" authority="(Rabeling et al., 2009)" baseAuthorityName="Rabeling" baseAuthorityYear="2009" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="smithii">
M. smithii (
<bibRefCitation id="538C4F7D1E3D2C9C3C145B90EE7D745B" author="Rabeling, C." etAl="et al." firstAuthor="Rabeling" journalOrPublisher="PLoS ONE" refId="ref12127" refString="Rabeling, C., Lino-Neto, J., Cappellari, S. C., Dos-Santos, I. A., Mueller, U. G. &amp; Bacci, M. (2009) Thelytokous parthenogenesis in the fungus-gardening ant Mycocepurus smithii (Hymenoptera: Formicidae). PLoS ONE, 4, e 6781. DOI: 10.1371 / journal. pone. 0006781." title="Thelytokous parthenogenesis in the fungus-gardening ant Mycocepurus smithii (Hymenoptera: Formicidae)" volume="4" year="2009">Rabeling et al., 2009</bibRefCitation>
)
</taxonomicName>
or any other
<taxonomicName id="1ACB6A6D84BC7BDB42E29B2833EEAE1E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2393" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus Forel" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="genus">Mycocepurus</taxonomicName>
species in Latin America (Rabeling, unpublished).
</paragraph>
<paragraph id="B6EF4BA1156A814D072E477BD26C6614" pageNumber="385">
Natural history and nest biology.
<taxonomicName id="3FD7EC7B2DBCF54CF8E67AB8F2031442" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
has been found twice in adjacent nests of
<taxonomicName id="7E0C360B3FE582F26CEE72B2D6F04531" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
. The two host nests had five and eight chambers, respectively, which were distributed between 5 and 190 cm depth (Table 1). The colony studied in 2006 contained 105 alate queens and 78 alate males of
<taxonomicName id="CA50E7BE7F71DE6CE5E371F93FC9B2CB" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
, and 771 workers of
<taxonomicName id="882B7731DA6542CA7AC47C7FE4849229" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
(Table 1). Dealate queens of either species could not be encountered, suggesting that the queenright chamber was either missed during the excavation or that the queens escaped into adjacent tunnels.
</paragraph>
<paragraph id="71C76D6B56DB1767ADA414F30C45C4EE" pageNumber="385">
The 2008 colony contained 15 dealate and 66 alate
<taxonomicName id="6F54D9C317A10C242944A7C82BF68525" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens, only six alate males, 1034
<taxonomicName id="C74463F6E786853700C45A21B795CB5D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers, a single dealate
<taxonomicName id="37C52C5BE473C0B764286A1C2741D614" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen and worker pupae (Table 1). The parasite 's numerical male/female sex ratio was strongly female biased (6/66 = 0.09). Twelve of the 15 dealate
<taxonomicName id="658F279B0A2843BF3F6A13DADF8691D8" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens were encountered in the same fungus garden chamber as the reproductively active female of
<taxonomicName id="D608B8D564D10D6F8458700A0A3395E7" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
. Thus,
<taxonomicName id="FE0F9ADC81B326F641F1904C75E1CDF9" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
is host-queen tolerant (Table 1). The other three dealate
<taxonomicName id="92D10A0EFBF491EFF3FF8826A9FBAF09" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens were found together in a separate fungus chamber (chamber 1; Table 1). The 12 queens encountered with the
<taxonomicName id="69B6BA9E8C5D2D0452260F51385729EB" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen showed different reproductive activities: three were active egg layers, showing developed ovaries, yellow bodies and sperm-filled spermathecae. Thus, the parasite can be polygynous. In contrast, the remaining nine queens were prereproductive with filled spermathecae, but the ovaries were still developing, and yellow bodies were absent. The three dealate queens from chamber 1 were also prereproductive. The single
<taxonomicName id="BDDD960B4E5E7340AFA78F56C6FDAED5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen was reproductively active.
</paragraph>
<paragraph id="0313B070CE6117D80EB606C638DF595E" pageNumber="385">
The unparasitized
<taxonomicName id="8612186D19C1B7F01BCF54685049969D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="385" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colony studied in 2008 contained a single reproductively active queen, 33 alate queens,
</paragraph>
<paragraph id="BE538CD25B80B6E2A5454DE4BD53A2E9" pageNumber="385">Entomology, 35, 379-392</paragraph>
</subSubSection>
<subSubSection id="C65DBE4A66C6067B67F69D17D1C93B8B" type="multiple">
<footnote id="61B207CF7DAED90FAB982EDC0517B337">
<paragraph id="EEB1D489044B388523C18F534AA7F4D5" pageNumber="385">© 2010 The Authors</paragraph>
</footnote>
<footnote id="884610B502DEAE2408EC226DFCB2D138">
<paragraph id="B47D3F7D27D710B6B2201992B6DA42D3" pageNumber="385">Journal compilation © 2010 The Royal Entomological Society, Systematic Entomology, 35, 379-392</paragraph>
</footnote>
<caption id="D9A5BA448BB7CB464AA428961025D596">
<paragraph id="6E5ACCDA990C830DD729718AA8567DBD" pageNumber="386">
<pageBreakToken id="46043093CBE745D6C6FF4B58F811685B" pageNumber="386">Table</pageBreakToken>
3. Life history and morphological characters of inquiline parasites in fungus-gardening ants.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="299B79BF78D634FA2FE5C21C79C1512B" type="discussion">
<paragraph id="D7BF791668840A3A6D32EF659A804002">
<table id="65AF65D8DB50EE012B9DCBE17BABD6EE" border="0" frame="box" pageNumber="386" rules="all">
<tr id="76E86669834D889386C37B4726F4832C" pageNumber="386">
<td id="3DD88E6DE42F807D7EF9401E0314B604" pageNumber="386">Incipient inquiline parasites</td>
<td id="91B77018FE4D415939A29D02233113D0" pageNumber="386">Evolutionarily derived inquiline</td>
<td id="8B5CE569C80E9D2621CD7E5B80BAB61B" pageNumber="386">parasites</td>
</tr>
<tr id="F38A3F496B00504458DA225E0358BF17" pageNumber="386">
<td id="6A937CCE649661DB2C61690C63CEECD5" pageNumber="386">Life history and morphological traits</td>
<td id="E2FA22476C51397BED76D473A9A1C836" pageNumber="386">
<taxonomicName id="BE2384B33AEA7FA2243ACB4A1F7670AE" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:152541" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex insinuator Schultz, Bekkevold &amp; Boomsma" order="Hymenoptera" phylum="Arthropoda" rank="species" species="insinuator">Acromyrmex insinuator</taxonomicName>
</td>
<td id="6F3F7259BE16013C82D4511E88C6D37F" pageNumber="386">
<taxonomicName id="AECBFEB5FC90C6B2C97C3373BDF68187" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:222191" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex ameliae De Souza, Soares &amp; Della Lucia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="ameliae">Acromyrmex ameliae</taxonomicName>
</td>
<td id="C721EE12F9C139BF4586D15BEEDB6E6F" pageNumber="386">
<taxonomicName id="A297A35AF85DFB6D17DC3424E83CFD24" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
[3]
</td>
<td id="1661351383BC063DB833003BB0F1D815" pageNumber="386">
<taxonomicName id="3E99B09F32C24AD59D421AF984A54F7A" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">Pseudoatta argentina</taxonomicName>
</td>
</tr>
<tr id="C59297B2C8EE7EC9D83135F39BBAC754" pageNumber="386">
<td id="8D76EAAB2B026735FA679312D2B83BA0" pageNumber="386">Parasite association with host</td>
<td id="598BA62FFBFB8589668A0DEC9A8760BA" pageNumber="386">Obligate [1, 6]</td>
<td id="E366781254478C625544E8027C061FAE" pageNumber="386">Obligate [2]</td>
<td id="6A60DBDDA10D658C22300BFE2FA042B3" pageNumber="386">Obligate</td>
<td id="44BA18C8221521344EC932CBF2A4857A" pageNumber="386">Obligate [7,8]</td>
</tr>
<tr id="D5E97D59B33909AE30FD31734CF99F37" pageNumber="386">
<td id="6C476E73B54BEE6F58F7BA9CDC60F21C" pageNumber="386">Parasite tolerant/intolerant of host queen</td>
<td id="0AAB14E696AC4D919C0DACBCA0807368" pageNumber="386">Tolerant [1,6]</td>
<td id="67B82CF9AFDB311C624A088EA76BE212" pageNumber="386">Tolerant [2]</td>
<td id="11552A57B0C5CF708E5B0ECFF060A744" pageNumber="386">Tolerant</td>
<td id="117225B0D047B95CEF4F5838CB36A813" pageNumber="386">Intolerance suggested [8]</td>
</tr>
<tr id="31D7E1E23B26CF606CAC9CDA738EE0FB" pageNumber="386">
<td id="A58FF1882F32BBC0061318CE2E628ECB" pageNumber="386">Worker caste present/absent</td>
<td id="2A6EB168406E86B89CE387787C984BC6" pageNumber="386">Minors present, very rarely media or large workers [1,6]</td>
<td id="BDD802472A97DB65774826C293F481FB" pageNumber="386">Minors present [2]</td>
<td id="FB94ECF37434558B27E11692C0F4EF64" pageNumber="386">Absent</td>
<td id="D2CF54DD3555FFA46E73FCBF4AA00093" pageNumber="386">Absent [9]</td>
</tr>
<tr id="3777E6BC4BFF49AC017C65FA033B1DAA" pageNumber="386">
<td id="1B82698000F67177B04369BEED212387" pageNumber="386">Number of reproductively active parasite queens</td>
<td id="F346A59D20915E52626F254434E63FF4" pageNumber="386">Facultative polygyny [6]</td>
<td id="688B89669157C1153027355A59060220" pageNumber="386">Facultative polygyny [2]</td>
<td id="24A477312AA6F112713CD57E4C06BC16" pageNumber="386">Polygyny</td>
<td id="1DF26DD26635D8E115A9C9623326795F" pageNumber="386">Unknown</td>
</tr>
<tr id="D6B400C17A050E9E2CA577460C971D83" pageNumber="386">
<td id="181FE0C90837BAA6ED6E7DC374BE2D32" pageNumber="386">Production of host alates in parasitized nests</td>
<td id="B97CA6174761992849C135392563B7EC" pageNumber="386">Host and parasite alates co-occur [6]</td>
<td id="3A851ECBC2A521B8FEDAEF47C91AE37C" pageNumber="386">Parasite alates precede or follow host alates [2]</td>
<td id="DA2A7A89CACE02C4735E7ADC6B936698" pageNumber="386">Host alates absent when parasite alates are present</td>
<td id="F04F038E1A14E2CBEB5D3595C1B2C49D" pageNumber="386">Host alates absent when parasite alates are present [7,8]</td>
</tr>
<tr id="DA174053F38CBA1AECE6FF97053B5D95" pageNumber="386">
<td id="3A32CCDEC3FE9784B30DC4CACDB681A5" pageNumber="386">Mating flights (compared with host)</td>
<td id="0989F55AC9094672902351D3665DBD0E" pageNumber="386">Present (normal) [1,6]</td>
<td id="321E0656046B8C26B848BF3C9602026A" pageNumber="386">Present (normal) [2]</td>
<td id="E0F3A7E7DF56571466F3AA9DD70DD716" pageNumber="386">Absent</td>
<td id="AA19071959BA62CF848B7AF0B1CBD881" pageNumber="386">
Absent in cf;
<normalizedToken id="EB3C51D60EE1E0CC6AE3C73DB0546D75" originalValue="Ç">C</normalizedToken>
perform dispersal flight [7,8]
</td>
</tr>
<tr id="428B410A4F3AF342087A4DF6ED9CDDB0" pageNumber="386">
<td id="F63DF4F4EEE33EA27ACD76B2B02126E1" pageNumber="386">Nest founding strategy</td>
<td id="1C5789A72890B9A67E12335FE38E64EA" pageNumber="386">
Invasion of established nests; pleometrosis with host
<normalizedToken id="F724A0DAAF54CF64DDFD54565DE57A56" originalValue="Ç">C</normalizedToken>
suggested [1,6,16]
</td>
<td id="6CED3C338BCF1CE3D196E83447E3410E" pageNumber="386">Invasion of established host colonies suggested [2]</td>
<td id="73DE97A67791D523100687FFC6271AAB" pageNumber="386">Invasion of established host colonies</td>
<td id="A4813299251791978ABF6DEE58381CE0" pageNumber="386">Invasion of established host colonies suggested [7,8]</td>
</tr>
<tr id="00F483D932688097C9656473AF7F918B" pageNumber="386">
<td id="B6426C34426045D87E8D3BA130962689" pageNumber="386">Reproductive strategy (semelparity vs iteroparity)</td>
<td id="F295D03B8207D6EF94C631933822A48D" pageNumber="386">Semelparity [6]</td>
<td id="25CA31A1A43AB7926A3E654ECC3C9975" pageNumber="386">Unknown</td>
<td id="06800E2061E675D86B44A09D7F6872F9" pageNumber="386">Iteroparity suggested</td>
<td id="018B3DFC7094863FDB602C7BC0298CD7" pageNumber="386">Semelparity suggested (host colony died after nuptial flight) [7,8]</td>
</tr>
<tr id="E37262755E3F05FAB6C8776E71B4B09C" pageNumber="386">
<td id="AEDBBE20BF2E046D5D48D80EEB4AD8BD" pageNumber="386">Mating strategy (intranidal vs outside nest)</td>
<td id="96949861542DAA11558A2EA800291A76" pageNumber="386">Possibly mixed: inside and outside nest mating [6]</td>
<td id="3883A4D52037C5E776DDD29926C838E1" pageNumber="386">Unknown</td>
<td id="797AB1DFD7C2483DBA32453E903D07B4" pageNumber="386">Intranidal mating</td>
<td id="0455781AE21325DF5099DCDB47D9745B" pageNumber="386">Intranidal mating [7,8]</td>
</tr>
<tr id="64C1CE7A0F356C5886589AFDF5502078" pageNumber="386">
<td id="C21FD6B18F34C688256C1BF84D9F5403" pageNumber="386">Host species</td>
<td id="554E98ABAD823F215FC2D95C31F5B0F5" pageNumber="386">
<taxonomicName id="B80FF5CE73F0B5252923C771D17F2920" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:224141" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex echinatior (Forel)" order="Hymenoptera" phylum="Arthropoda" rank="species" species="echinatior">A. echinatior</taxonomicName>
[1,6]
</td>
<td id="D15C92E4C9D82ED5A668D5FEC7A2E7D2" pageNumber="386">
<taxonomicName id="3779D4C682D0CBF0B4E73F8D017EF92E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:25059" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex subterraneus subterraneus (Forel)" order="Hymenoptera" phylum="Arthropoda" rank="subSpecies" species="subterraneus" subSpecies="subterraneus">A. subterraneus subterraneus</taxonomicName>
,
<taxonomicName id="433A930D43699D07EDF603AF4BBFFF90" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:24719" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex subterraneus brunneus (Forel)" order="Hymenoptera" phylum="Arthropoda" rank="subSpecies" species="subterraneus" subSpecies="brunneus">A. subterraneus brunneus</taxonomicName>
[2]
</td>
<td id="2C907301F72D9A43A37403204C81E0AE" pageNumber="386">
<taxonomicName id="43A8E2C495B3E5DCED06212CFED5F064" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
</td>
<td id="691858F9DC5C995338637763A9169A7F" pageNumber="386">A. lundi [9]</td>
</tr>
<tr id="5D5C34B76F96ACF43EA51EB06840B71A" pageNumber="386">
<td id="F960CC1D337A8011610983FFD34502B5" pageNumber="386">Number of reproductively active host queens</td>
<td id="6CE12D1BC3ED9A5D69562992FEBA77AE" pageNumber="386">Facultative polygyny [12]</td>
<td id="AE22044106629D9E60CD504CADC457ED" pageNumber="386">Facultative polygyny [10,11]</td>
<td id="9C4CA09A7A4912AEC7AE8B84CF8870E3" pageNumber="386">Monogyny in Rio Claro [3]; facultative polygyny in Manaus [3,13]</td>
<td id="6A92B7DFFA4A94BB5F1E98B23C885C49" pageNumber="386">Unknown</td>
</tr>
<tr id="36070D959D388888E73B08CB03D31DA7" pageNumber="386">
<td id="638FE328234963134A020CC463FDDFDF" pageNumber="386">Reduction of parasite body size</td>
<td id="75B51DD4C0032F5BF4A74655C95BAD7E" pageNumber="386">Both sexes [1]</td>
<td id="9B74E9CF4BE4AA0C2948BBABA2754F68" pageNumber="386">Both sexes [2]</td>
<td id="CCA7F90E3F2FE58967AB5FCE1EA52AC2" pageNumber="386">Both sexes</td>
<td id="4F9BF4D0CAF025D3344238DCB3940EAC" pageNumber="386">Both sexes [9]</td>
</tr>
<tr id="3C588E3BFF74194BF3BA22EDDFEA0339" pageNumber="386">
<td id="DED723CA6277F8DD6397EF6B082A9618" pageNumber="386">Gynomorphism of parasite males</td>
<td id="953C1106CACD452E0688E4E8F4C0FE2E" pageNumber="386">Absent [1]</td>
<td id="B8DE1BE7FDD8833238A8A2CDD97D6850" pageNumber="386">Absent [2]</td>
<td id="2A7EA2313F66F183EF064CC367376D17" pageNumber="386">Present</td>
<td id="75EC55CD2FE36FE2515E16C4C044C501" pageNumber="386">Present [9]</td>
</tr>
<tr id="62DDD6E4584B597CD32E4180D0B168EB" pageNumber="386">
<td id="E7E8788DE87F1E5952686F946DF3AEE1" pageNumber="386">Integument sculpturing of parasite</td>
<td id="949A5D64D9B3BF53D89C14E690DB7967" pageNumber="386">Well developed, matt [1]</td>
<td id="62DC4BE5E9E2B0F0160E571C91FD7B1A" pageNumber="386">Well developed, matt [2]</td>
<td id="D4E2A5E80980A60B7F37F6D55F540D26" pageNumber="386">Smooth, shiny</td>
<td id="86D47A831FAEE273037F067D01CFA6C1" pageNumber="386">Smooth, shiny [9]</td>
</tr>
</table>
</paragraph>
<paragraph id="EB0AABF9D72D3AE2DF0A2679818196BE">
<table id="162E9F6D1EB2E62499DEA767C6151C43" border="0" frame="box" pageNumber="387" rules="all">
<tr id="A3D340D5A768C4AEFA2EF49C83DBF8D0" pageNumber="387">
<td id="E631C019F388FE66F3165C57014A55E5" pageNumber="387">Incipient inquiline parasites</td>
<td id="4DB7EB359FB91F3D190348ED52ED476E" pageNumber="387">Evolutionarily derived inquiline</td>
<td id="595A765177D36531E6075A6FE4259114" pageNumber="387">parasites</td>
</tr>
<tr id="60C9E99407691DCAC1F785B1D0C06788" pageNumber="387">
<td id="2AADEB071743083821B7EA9052711964" pageNumber="387">Life history and morphological traits</td>
<td id="0FE42F37F091D36A69F8F496D34FEFB2" pageNumber="387">
<taxonomicName id="DFFA29F6A14BBDDE3C38C23E442E931E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:152541" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex insinuator Schultz, Bekkevold &amp; Boomsma" order="Hymenoptera" phylum="Arthropoda" rank="species" species="insinuator">Acromyrmex insinuator</taxonomicName>
</td>
<td id="E50DB944E9C0CB9558DF46D8F8A1D3FA" pageNumber="387">
<taxonomicName id="7067FD7247B65C9E92993941B688453A" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:222191" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex ameliae De Souza, Soares &amp; Della Lucia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="ameliae">Acromyrmex ameliae</taxonomicName>
</td>
<td id="9C514AE71F2FE635FFF3A4BB9E3A2E3B" pageNumber="387">
<taxonomicName id="001A631D741DE36E7A4BE9D214AE519C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
[3]
</td>
<td id="2DFF3C6C9F61544BAA61634730D76D98" pageNumber="387">
<taxonomicName id="0D74C07B7E95F8EC3F254BB0C5598BB3" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">Pseudoatta argentina</taxonomicName>
</td>
</tr>
<tr id="80DEFA534670F826BD8A3905C5CEB453" pageNumber="387">
<td id="6DE607BFB5CD1AE16C200FAAD680AA29" pageNumber="387">Pilosity (relative to host)</td>
<td id="011334712950208CE87DE970477A5137" pageNumber="387">Similar [1]</td>
<td id="D93D91F2C0C381DDB11ECABA6353D246" pageNumber="387">Similar [2]</td>
<td id="FDD076C2976181EC06111866358B1013" pageNumber="387">Reduced</td>
<td id="747FF44C31DACE631A8DFDAD493AD9E4" pageNumber="387">Reduced [9]</td>
</tr>
<tr id="F71D9E0D7F4756C1F301358C164AA90F" pageNumber="387">
<td id="85EBE87548B7247F59E5D0C2BD78F132" pageNumber="387">Mandibular dentition</td>
<td id="E06F28C719730481A092E4EDC93C09D9" pageNumber="387">Well developed [1]</td>
<td id="8C054EA9426AE8029927272D1D46E7C2" pageNumber="387">7-8 denticles in cf [2]</td>
<td id="1D298EE432096D02239BE36C4BA8A2C1" pageNumber="387">Reduced</td>
<td id="D10899C9CDC7EB5F86CE44EE73040CDE" pageNumber="387">7-8 denticles in $ [9]</td>
</tr>
<tr id="30CBCBD4B896CD966C203930213BA15D" pageNumber="387">
<td id="76406295A797E0729395CC552CCE3B6C" pageNumber="387">Number of antennal segments</td>
<td id="0D1C47F7BF972A0403B2B4C53D68344F" pageNumber="387">J: 11; cf: 13; funicular segments 4 and 5 partly fused in some cf individuals [1]</td>
<td id="76FA116775996E5789CC1E6C8D031602" pageNumber="387">5: 11; cf: 13; funicular segments 4 and 5 partly fused in some cf individuals [2,14,15]</td>
<td id="ED99A2DC25F350FD638395383D8BD12B" pageNumber="387">5: 11; cf: 11</td>
<td id="600DE8B4F7DACEF61364B1A7D8B0F694" pageNumber="387">?: 11; cf: 11 [9]</td>
</tr>
<tr id="238891CE895238EA99FB352EC817F7A8" pageNumber="387">
<td id="6551632484455EF51004FCE286E01A71" pageNumber="387">Palp formula (maxillary, labial palp)</td>
<td id="10538544EF5DCB80270AE4872F00E6F2" pageNumber="387">4,2 [1]</td>
<td id="D904AE2FF605C33EFB0FAEF07F9266C2" pageNumber="387">4,2 [2]</td>
<td id="9EF851528B973F67797D1C61AE08F879" pageNumber="387">3,2</td>
<td id="15F97397B85FB266CCEC6B2AB10AC8FA" pageNumber="387">3,2 [1,14,15]</td>
</tr>
<tr id="A4560F06CB0AC39AF3B8CD8C6E268577" pageNumber="387">
<td id="83ACC2980416FA3E9CB0C7F4C2C56127" pageNumber="387" rowspan="2">
Size and shape of metapleural gland opening in
<normalizedToken id="CA8F811C8E2F14C99EBE3511F134E8BE" originalValue="Ç">C</normalizedToken>
</td>
<td id="3C3F751B671AF178B141AEA1DFF507BA" pageNumber="387">Similar to host [4]</td>
<td id="44F5AF27AEDAA6B0037109FE1F2A1576" pageNumber="387">Similar to host [5]</td>
<td id="13C313EE17D1DBEBEA311EE3865AE2E6" pageNumber="387">Enlarged, circular, pointing backwards</td>
<td id="F583D8C58D4E0D2D44ADB092B985F800" pageNumber="387">Enlarged, circular, pointing backwards [4]</td>
</tr>
<tr id="950A0D220EC8033E8AE71DD82CF8C174" pageNumber="387">
<td id="AEDA392957D33B61E48434F253CBCCE6" pageNumber="387">Size and shape of</td>
<td id="37C174EA7D017B2C5E78EB122547B7AA" pageNumber="387">Absent [17]</td>
<td id="7C373925D8EBF5D3214B2BF164243541" pageNumber="387">Absent [17]</td>
<td id="BB63F6ECF595F701665200BE7832F05B" pageNumber="387">Tiny, circular opening on</td>
<td id="D67D8E6A454862AB54E6E95070CD4C0C" pageNumber="387">Enlarged opening, similar to</td>
</tr>
<tr id="FB165E80E009DCD4E4FE7E043E9001CE" pageNumber="387">
<td id="906390240102071DED42DC080CD5EE5F" pageNumber="387">metapleural opening in cf (existence of gland not confirmed with histological methods)</td>
<td id="784AB91A56D424BD0377B1BB928197B9" pageNumber="387">
metapleuron, congruent with position of
<collectionCode id="16EFBAE1039B898AF335A72393BF513B" collectionName="MPG">MPG</collectionCode>
opening in female
</td>
<td id="985DD32BCD5121C04E3EE14086F2BC3C" pageNumber="387">the queen [17]</td>
</tr>
<tr id="8F2FC9C6F3DB80615E57D79892E85309" pageNumber="387">
<td id="C2CEB0E1075F0F02FCC7699C53CFAF98" pageNumber="387">Postpetiole width</td>
<td id="23D352833550FCC8D2E434EEF4B23875" pageNumber="387">Similar to host [1]</td>
<td id="8373C48385A3B2B3E4E4F9332575491C" pageNumber="387">Similar to host [2]</td>
<td id="5ACC468024EC6260BAEFE33BA2E3B40D" pageNumber="387">Enlarged</td>
<td id="25E917E1750B7F89A656CB6DFC96563D" pageNumber="387">Enlarged [9]</td>
</tr>
<tr id="2716C874739A38C9DD2838AB97A7261B" pageNumber="387">
<td id="8036C753493AF06B164F8694BEAF2727" pageNumber="387">First gastral tergite of $ with concave, median impression</td>
<td id="B906CF9B1AF2D9E25D57D89A05DFF99D" pageNumber="387">Absent [1]</td>
<td id="26AA84E90B0762B3AF3A293811F71937" pageNumber="387">Absent [2]</td>
<td id="A3E5D96A565B9B71DA29F49E3DFFD6A9" pageNumber="387">Present</td>
<td id="CB087BEE72E9A6560381BFD042FEA308" pageNumber="387">Present [9]</td>
</tr>
</table>
</paragraph>
<caption id="8F84F9ECC8727A3588DB801A4621F588">
<paragraph id="C6CAB20ADBBED18600DF5506D030704C" pageNumber="387">
<pageBreakToken id="EE3E65DB3C42F7FBA054B840EDA2805F" pageNumber="387">Table</pageBreakToken>
3. Continued
</paragraph>
</caption>
<paragraph id="A434020340FCA7074F7C0802BF0FF9FF" pageNumber="387">
All four species originated independently, but evolved similar traits convergently, allowing for classifying them as incipient and evolutionarily derived social parasites. A second, hitherto undescribed
<taxonomicName id="D3BEFDF3D7A8D7A6AF4BE17306FF1CB5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2395" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" lsidName-HNS="Pseudoatta Gallardo" order="Hymenoptera" pageNumber="387" phylum="Arthropoda" rank="genus">Pseudoatta</taxonomicName>
species is not considered in this table, because only few morphological characteristics are described in the literature (
<bibRefCitation id="6AF6D46372C8D8A6CBE56BB2F4298028" author="Delabie, J. H. C." etAl="et al." firstAuthor="Delabie" journalOrPublisher="Fourth International Symposium on Pest Ants / XI Encontro de Mirmecologia, Belo Horizonte, Minas Gerais, 21 - 24 November 1993" refId="ref11151" refString="Delabie, J. H. C., Fowler H. G. &amp; Schlindwein M. N. (1993) Ocorreˆncia do parasita social Pseudoatta sp. nova em ninhos de Acromyrmex rugosus em Ilhe ' us, Bahia: primeiro registro para os tro ' picos. Fourth International Symposium on Pest Ants / XI Encontro de Mirmecologia, Belo Horizonte, Minas Gerais, 21 - 24 November 1993." title="Ocorre ˆ ncia do parasita social Pseudoatta sp. nova em ninhos de Acromyrmex rugosus em Ilhe ' us, Bahia: primeiro registro para os tro ' picos" year="1993">Delabie et al., 1993</bibRefCitation>
; see also
<bibRefCitation id="9CD313F21BAB72629F0DAAB3FA282A3B" author="Schultz, T. R." etAl="et al." firstAuthor="Schultz" journalOrPublisher="Insectes Sociaux" pagination="457 - 471" refId="ref12339" refString="Schultz, T. R., Bekkevold, D. &amp; Boomsma, J. J. (1998) Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants. Insectes Sociaux, 45, 457 - 471." title="Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants" volume="45" year="1998">Schultz et al., 1998</bibRefCitation>
). Some life history information summarized here is derived only from single observations or stems from indirect evidence, and therefore should be considered tentative.
</paragraph>
<paragraph id="F415EB39BDE8FA55A232D91224377AD2" pageNumber="387">
Numbered references in this table refer to the following sources (please see reference list for complete citations): [1]
<bibRefCitation id="FE818EFA63C2F54A12C8B9C828423F81" author="Schultz, T. R." etAl="et al." firstAuthor="Schultz" journalOrPublisher="Insectes Sociaux" pagination="457 - 471" refId="ref12339" refString="Schultz, T. R., Bekkevold, D. &amp; Boomsma, J. J. (1998) Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants. Insectes Sociaux, 45, 457 - 471." title="Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants" volume="45" year="1998">Schultz et al. (1998)</bibRefCitation>
; [2]
<bibRefCitation id="1DF5AC0F31B95754670E55AB9100E511" author="de Souza, D. J." etAl="et al." firstAuthor="Souza" journalOrPublisher="Insect Science" pagination="251 - 257" refId="ref12382" refString="de Souza, D. J., Soares, I. M. F. &amp; Della Lucia, T. M. C. (2007) Acromyrmex ameliae sp. n. (Hymenoptera: Formicidae): a new social parasite of leaf-cutting ants in Brazil. Insect Science, 14, 251 - 257." title="Acromyrmex ameliae sp. n. (Hymenoptera: Formicidae): a new social parasite of leaf-cutting ants in Brazil" volume="14" year="2007">Souza et al. (2007)</bibRefCitation>
; [3] this study; [4] S.H. Yek &amp; U.G. Mueller (personal communication); [5] Rabeling (personal observation); [6]
<bibRefCitation id="DC33A59F9D4B8E3BF895C8C4B69C897E" author="Bekkevold, D." firstAuthor="Bekkevold" journalOrPublisher="Journal of Evolutionary Biology" pagination="615 - 623" refId="ref10761" refString="Bekkevold, D. &amp; Boomsma, J. J. (2000) Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator. Journal of Evolutionary Biology, 13, 615 - 623." title="Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator" volume="13" year="2000">Bekkevold &amp; Boomsma (2000)</bibRefCitation>
; [7]
<bibRefCitation id="772B21564E69068BF94DC41043401EDF" author="Gallardo, A." firstAuthor="Gallardo" journalOrPublisher="Revista de la Sociedad Entomológica Argentina" pagination="197 - 202" refId="ref11526" refString="Gallardo, A. (1929) Note sur les moeurs de la fourmi Pseudoatta argentina. Revista de la Sociedad Entomológica Argentina, 2, 197 - 202." title="Note sur les moeurs de la fourmi Pseudoatta argentina" volume="2" year="1929">Gallardo (1929)</bibRefCitation>
; [8]
<bibRefCitation id="F76C9F4D38CA3C8256460814AF1DAAC1" author="Bruch, C." firstAuthor="Bruch" journalOrPublisher="Buenos Aires" pagination="341 - 360" refId="ref10978" refString="Bruch, C. (1928) Estudios mirmecolo ' gicos. Anales del Museo Nacional de Historia Natural, Buenos Aires, 34, 341 - 360." title="Estudios mirmecolo ' gicos. Anales del Museo Nacional de Historia Natural" volume="34" year="1928">Bruch (1928)</bibRefCitation>
; [9]
<bibRefCitation id="FBA75558F0C1FFBAB401F33947E11972" author="Gallardo, A." firstAuthor="Gallardo" journalOrPublisher="Buenos Aires" pagination="317 - 344" refId="ref11483" refString="Gallardo, A. (1916) Notes syste ' matiques et e ' thologiques sur les fourmis attines de la Re ' publique Argentine. Anales del Museo Nacional de Historia Natural, Buenos Aires, 28, 317 - 344." title="Notes syste ' matiques et e ' thologiques sur les fourmis attines de la Re ' publique Argentine. Anales del Museo Nacional de Historia Natural" volume="28" year="1916">Gallardo (1916)</bibRefCitation>
; [10]
<bibRefCitation id="65E28044C88CA62AC4122EC08C7A2F4D" author="Della Lucia, T. M. C." firstAuthor="Della Lucia" journalOrPublisher="Hymenoptera" refId="ref11226" refString="Della Lucia, T. M. C. &amp; Vilela, E. F. (1986) Ocorrência de poliginia em Acromyrmex subterraneus subterraneus Forel, 1893 (Hymenoptera:" title="Ocorrência de poliginia em Acromyrmex subterraneus subterraneus Forel" volume="1893" year="1986">Della Lucia &amp; Vilela (1986)</bibRefCitation>
; [11]
<bibRefCitation id="75735B060DE0BDD6CDF2BD714110101F" author="Delabie, J. H. C." firstAuthor="Delabie" journalOrPublisher="Anais da Sociedade Entomolo ' gica do Brasil" pagination="193 - 197" refId="ref11100" refString="Delabie, J. H. C. (1989) Observações sobre a ocorrência de poliginia em colônias de Acromyrmex subterraneus brunneus Forel, 1893 (Formicidae, Myrmcinae, Attini) em cacauais. Anais da Sociedade Entomolo ' gica do Brasil, 18, 193 - 197." title="Observações sobre a ocorrência de poliginia em colônias de Acromyrmex subterraneus brunneus Forel, 1893 (Formicidae, Myrmcinae, Attini) em cacauais" volume="18" year="1989">Delabie (1989)</bibRefCitation>
; [12]
<bibRefCitation id="60108A6B0D88185D857DDFA5D0B58D88" author="Bekkevold, D." etAl="et al." firstAuthor="Bekkevold" journalOrPublisher="Behavioral Ecology and Sociobiology" pagination="103 - 109" refId="ref10801" refString="Bekkevold, D., Frydenberg, J. &amp; Boomsma, J. J. (1999) Facultative polygyny and multiple mating in the Panamanian leafcutter ant Acromyrmex echinatior (Hymenoptera: Formicidae). Behavioral Ecology and Sociobiology, 46, 103 - 109." title="Facultative polygyny and multiple mating in the Panamanian leafcutter ant Acromyrmex echinatior (Hymenoptera: Formicidae)" volume="46" year="1999">Bekkevold et al. (1999)</bibRefCitation>
; [13]
<bibRefCitation id="D614FBCD35BDC19AE080B4F4F325FBF0" author="Rabeling, C." etAl="et al." firstAuthor="Rabeling" journalOrPublisher="Journal of Insect Science" refId="ref12022" refString="Rabeling, C., Verhaagh, M. &amp; Engels, W. (2007 b) Comparative study of nest architecture and colony structure of the fungus-growing ants, Mycocepurus goeldii and M. smithii. Journal of Insect Science, 7, 40. Available at: www. insectscience. org / 7.40" title="Comparative study of nest architecture and colony structure of the fungus-growing ants, Mycocepurus goeldii and M. smithii" volume="7, 40" year="2007" yearSuffix="b">Rabeling et al. (2007b)</bibRefCitation>
; [14]
<bibRefCitation id="7CD47564F1A5C161FFD9CDD5799C672F" author="Kusnezov, N." firstAuthor="Kusnezov" journalOrPublisher="Universidad de Cochabamba" pagination="62 - 70" refId="ref11668" refString="Kusnezov, N. (1951) Los segmentos palpales en hormigas. Folia Universitaria, Universidad de Cochabamba, 5, 62 - 70." title="Los segmentos palpales en hormigas. Folia Universitaria" volume="5" year="1951">Kusnezov (1951)</bibRefCitation>
; [15]
<bibRefCitation id="64EF6916D750C97F0C69FD817B132293" author="Kusnezov, N." firstAuthor="Kusnezov" journalOrPublisher="Zoologischer Anzeiger" pagination="28 - 38" refId="ref11694" refString="Kusnezov, N. (1954) Phyletische Bedeutung der Maxillar- und Labialtaster der Ameisen. Zoologischer Anzeiger, 153, 28 - 38." title="Phyletische Bedeutung der Maxillar- und Labialtaster der Ameisen" volume="153" year="1954">Kusnezov (1954)</bibRefCitation>
; [16] J. J. Boomsma &amp; V. Nehring (personal communication); [17] T.R. Schultz (personal communication).
</paragraph>
<paragraph id="A65C4B02B5F49F06202CFA9E484AACB6" pageNumber="388">
<pageBreakToken id="1C3E72B4B06D9D64A630EBFA984C30E4" pageNumber="388">496</pageBreakToken>
workers and no males (Table 2). During the excavation, males and queens were leaving the maternal colony for their nuptial flight, which started on
<date id="5D6BF9164314E73D7108183D57E204E9" day="07" month="10">7 October</date>
.
</paragraph>
<paragraph id="D3E2780A5122D8824772752A86734526" pageNumber="388">
A natural history study of
<taxonomicName id="AE3CADD7350AA61F1B0F5F1A0226A430" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
in the Amazon Basin (
<bibRefCitation id="A14570A19B9E4723A85651EE8E9A1564" author="Rabeling, C." etAl="et al." firstAuthor="Rabeling" journalOrPublisher="Journal of Insect Science" refId="ref12022" refString="Rabeling, C., Verhaagh, M. &amp; Engels, W. (2007 b) Comparative study of nest architecture and colony structure of the fungus-growing ants, Mycocepurus goeldii and M. smithii. Journal of Insect Science, 7, 40. Available at: www. insectscience. org / 7.40" title="Comparative study of nest architecture and colony structure of the fungus-growing ants, Mycocepurus goeldii and M. smithii" volume="7, 40" year="2007" yearSuffix="b">Rabeling et al., 2007b</bibRefCitation>
) showed that some colonies had a single queen, whereas others were occupied by as many as four queens. Dissection of eight individuals from three separate colonies revealed that all of them were inseminated and had fully developed ovaries, demonstrating that these colonies were functionally polygynous.
</paragraph>
</subSubSection>
<subSubSection id="EE1B42FF6F13F63E4D4B5305D8ACB4BA" type="biology_ecology">
<paragraph id="1F32A83DC880B697921681A38E697796" pageNumber="388">
The observations on nesting biology and colony counts suggest that
<taxonomicName id="82F708060F63BDAA9503E9EF954C4965" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
is polygynous, host tolerant and allows for the production of sterile
<taxonomicName id="8BDE19350FD0B3565D6E104ABDC455C4" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers, whereas the production of host sexual offspring is suppressed in the presence of the parasite. The host,
<taxonomicName id="50F6D4CD0805FD6A4A79670309B76DA3" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
, appears to be monogynous in the Rio Claro population, but both mono- and polygynous colonies co-occur in the Brazilian Amazon.
</paragraph>
<paragraph id="59A5C65638D68FB897763A3C4FABC53C" pageNumber="388">
Behaviour. In the late afternoon of
<date id="0B4B1B8003E99149D3B73061D7C2FF2B" day="29" month="09" value="2006-09-29" year="2006">29 September 2006</date>
,
<taxonomicName id="56374EABFE6D1D5471EFADD35B276A9C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
was discovered when 31 queens and a single male left the host colony to aggregate on the nest mound. The dispersal activity was interrupted by rain, but continued on
<date id="83E2C8C9E030FC3B5A967F7C0153464B" day="02" month="10">2 October</date>
, when 24 queens and 72 males emerged. No further behavioural observations were made that year.
</paragraph>
<paragraph id="D24407B49BF54C179CF21BF9438CF209" pageNumber="388">
In 2008,
<taxonomicName id="91B563873168EA4F2738983A81E6421B" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colonies were excavated at the end of the dry season in order to study parasitized colonies before the nuptial flight. An approaching mating flight is easily identified in
<taxonomicName id="50BE12B9BD022C07885B9176C29F7072" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colonies, because the workers increase the number of nest entrances per soil mound to maximally 30 entrances, giving the nest mound a sponge-like appearance (
<bibRefCitation id="12F907E0A680ADEADEEFC875604B76E4" author="Rabeling, C." etAl="et al." firstAuthor="Rabeling" journalOrPublisher="PLoS ONE" refId="ref12127" refString="Rabeling, C., Lino-Neto, J., Cappellari, S. C., Dos-Santos, I. A., Mueller, U. G. &amp; Bacci, M. (2009) Thelytokous parthenogenesis in the fungus-gardening ant Mycocepurus smithii (Hymenoptera: Formicidae). PLoS ONE, 4, e 6781. DOI: 10.1371 / journal. pone. 0006781." title="Thelytokous parthenogenesis in the fungus-gardening ant Mycocepurus smithii (Hymenoptera: Formicidae)" volume="4" year="2009">Rabeling et al., 2009</bibRefCitation>
). Until
<date id="B54BC538B7B4B439C862C1540D8B548D" day="03" month="10">3 October</date>
, when a
<taxonomicName id="3F08DA0DD87723368007F66D47E5A85B" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colony parasitized by
<taxonomicName id="5712191450CE401C6F6EE46C0EBE0596" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
was encountered, the
<taxonomicName id="19283E67920E2778FD4560E4ABE247C8" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers did not modify the nest mounds for mating flights. Upon excavating the parasitized colony, all individuals from a total of five nest chambers were transferred to artificial nest chambers for behavioural studies.
</paragraph>
<paragraph id="B215424C88090DBD4EC061F48F20752A" pageNumber="388">
Parasite mating behaviour. As soon as the uppermost chamber(CR
<date id="614156109894CAE2FDF8A6F75446EF6C" day="08" month="10" value="03-10-08" year="03">081003</date>
-01, Table 1) was opened during excavation, and the ants were transferred to the artificial nest chambers,
<taxonomicName id="9B013804BEAE06DD44D881BACBAA9A92" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="subSpecies" species="castrator">M. castrator males</taxonomicName>
started copulating with
<taxonomicName id="4F2DCC3AB10CE88CD28EA21ED0F452C6" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
females inside the artificial chamber (
<taxonomicName id="37CBA3D2636D4E399F4FC7773FE82897" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii alates</taxonomicName>
from an adjacent nest, placed into a laboratory nest, were never observed to copulate). During this time of ongoing mating activities, females and males ran erratically in jerky movements, and males mounted females seemingly at random. Observed copulations lasted between 18 and 27 s (n = 4). Single males attempted to copulate more than once. It is unknown whether repeated copulations resulted in successful transfer of sperm. Within 3 h after transfer to the artificial nest, three queens shed their wings, and subsequent dissections demonstrated that these females had been inseminated. However, their ovaries were still developing (i.e. ripe oocytes and yellow bodies were absent), indicating recent insemination (note: queens were preserved for dissection2 weeks after copulation). Postcopulatory females did not tolerate mating attempts of males, and walked faster to outdistance their pursuer. After copulation, and wing shedding,
</paragraph>
<paragraph id="5FC2E584A66B45C2A48EFC05C9395730" pageNumber="388">the recently mated queens gathered and engaged in allogrooming, frequently licked each others meso- and metasomas, and wings for extensive periods of time. The first dead males were found 12 h after the mating event. Alates from other chambers did not copulate after transfer to the artificial nest. Potentially, the individuals in the topmost chamber were anticipating the upcoming nuptial flight and the opening of their nest chamber triggered the mating behaviour.</paragraph>
<paragraph id="437316E7CAFEC7A36AE4C1D36D5F2EE6" pageNumber="388">
Host worker -parasite interactions. Host workers and parasite alates frequently antennated and interacted nonaggressively.
<taxonomicName id="D375A248084A0875CF2AC78873659272" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator alates</taxonomicName>
did not require grooming by host workers because individuals cleaned themselves (i.e. licking appendages, cleaning antennae), and females groomed each other. Dealate
<taxonomicName id="70E8D10B44FC9BB20228C850468AFCB9" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens groomed
<taxonomicName id="24F8BFA7BDEC1AC1CF9026573BF98983" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers, and were groomed by them also. On several occasions,
<taxonomicName id="0C698A90E1DB3DE0C819B06566BD1427" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers licked the tip of a
<taxonomicName id="78A01239EBA6899FF2B5CFFB5F41BD57" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
metasoma for several minutes; it is not clear if the workers removed fecal droplets, or
<taxonomicName id="642F5C8C3A21DB341846F8D094394206" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens laid either fertile or trophic eggs.
<taxonomicName id="98810656450119DC4BCC41CBFC8AC0BA" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">Mycocepurus goeldii</taxonomicName>
workers fed the parasite queens via trophallaxis. To be fed,
<taxonomicName id="BEDE14761477F5BD9FD427AD77F4E4D4" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
females frequently climbed onto the host workers ' backs, antennated the host 's antennae and head, until it bent its head backwards, regurgitated liquid, which was then consumed by the parasite. In addition to being fed,
<taxonomicName id="88D0B7569730174C82E05659603D8075" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="subSpecies" species="castrator">M. castrator males</taxonomicName>
and females actively licked the fungus garden.
</paragraph>
<paragraph id="83F5B3CCD50BABCC2A4AC33F44D12DB9" pageNumber="388">
Three days after insemination, the host workers aggressively attacked one dealate queen from the topmost chamber (CR
<date id="618E17A773979C0CC5D051F9EBF62F19" day="08" month="10" value="03-10-08" year="03">081003</date>
-01); six to eight workers secured her by the antennae, legs, head and petiole, until she died. Approximately 24 h after her death, three workers continued to carry around her corpse in the nest chamber. Six days after insemination, the host workers had attacked and killed several
<taxonomicName id="4DB8E5862422E1E2133C590AECE02C6C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens, and had placed them on the refuse dump. Three dealate queens remained unmolested by hiding together in the fungus garden.
</paragraph>
<paragraph id="B77376A308337E7FDE192741CAB5AE57" lastPageNumber="389" pageNumber="388">
Host queen -parasite interactions. To observe the interactions between the host queen and the dealate
<taxonomicName id="447B980D19E02A27A0415458E444D917" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens (n = 12), we placed the queens in a smaller nest chamber, after
<taxonomicName id="3AF96322B43314546992B46EB016831D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers had arranged the fungus garden. The
<taxonomicName id="0DAFA40D5BC8617F72099BE60A21F29E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens were much more agile than the
<taxonomicName id="BBCF1A55914C5D28A7B7726C5628393E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen and initially walked around the nest chamber until they encountered a suitable spot; there they aggregated and started licking each other. When first placed in the chamber, the
<taxonomicName id="C708F694E46C022371FA7DC61E937C19" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen crawled under an adjacent piece of fungus garden and remained motionless; a worker then picked her up by the metasoma and moved the queen to a different position. During the carrying, the
<taxonomicName id="FCE22A666EC9D71135E708E95D7830BB" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen remained motionless. After several minutes, one
<taxonomicName id="81925C4D9AC1B01015F24CFE9AF43090" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen left the aggregation, searching ' for the
<taxonomicName id="F92BA26705A81EE19BE4561D21A107C1" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen. When the host queen was encountered, she was surrounded by host workers antennating her. Regardless, the parasite climbed on the host queen 's back (Fig. 3), and started licking her mesosoma, petiole, postpetiole and metasoma. Shortly afterwards, a second
<taxonomicName id="3BE1DD8E544C47754636DCBF2803D1A5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen joined the first; the
<taxonomicName id="9A641F4289557C6648E4224A48056D03" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="388" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
<pageBreakToken id="F54AE15BA6AA9A93A1D8637DDCBB9295" pageNumber="389">queen</pageBreakToken>
continued to remain motionless. The remaining
<taxonomicName id="3B0E3EDD95747CEB5CBB45E4B8AEC3E1" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens eventually joined the grooming cluster, and alternated grooming themselves by pulling their legs and antennae through the tibio-tarsal cleaning apparatus of the foreleg, with grooming the host queen. When the first
<taxonomicName id="20DE313A56FD1BC24E7157209D48A561" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen climbed on the host queen 's back, the attending workers left and resumed fungus-gardening activities. They did not react aggressively to the parasite queens and often returned to antennate and to feed the host queen via trophallaxis. Once a worker carried the host queen to a different part of the fungus garden, and a
<taxonomicName id="220E53323EC38A49D68E6C17B231C8BB" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen rode on her back during the location, licking her, and was not chased away. Either workers or
<taxonomicName id="622C80B27A9B2499091DD615C67748AF" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens attended the host queen for most of time. Rarely and then for very short periods of time, she sat by herself. The
<taxonomicName id="CD757567F40952CECEA0A7985F42577B" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens attempted constantly to climb on the backs of either other
<taxonomicName id="FCD053838F753497D8226BD533522F1A" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens, the host queen or host workers.
</paragraph>
<caption id="A296622BF71F1C59D46244340959F8B2">
<paragraph id="60F9F31BBC84233A8DF600BE25A1C48D" pageNumber="389">
Fig. 3. A queen of the social parasite
<taxonomicName id="61796507B8188C8A7C4F41C70F0EE1F7" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
standing on the host queen 's back. The photograph was taken in a laboratory nest, set up at Sa˜o Paulo State University in Rio Claro, Brazil in
<date id="8740042E9615340E33E6E3252CB01862" month="10" value="2008-10" year="2008">October 2008</date>
. The photograph was taken by Scott Solomon.
</paragraph>
</caption>
<paragraph id="16B4961AF71A77AF54AFE93FBB161D1A" pageNumber="389">
Introduction of parasite queen into a field colony. Tw o inseminated
<taxonomicName id="B473269998F46199322B24F5C60BA9F5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens from the topmost chamber (CR
<date id="4751EED5DDD0047A34EAE89304413FB1" day="08" month="10" value="03-10-08" year="03">081003</date>
-01) were introduced to a
<taxonomicName id="1D2C462A016AF07D2B1764EC7CE9267C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colony, which opened its nest mound in preparation for the nuptial flight the previous day. The
<taxonomicName id="A6A18ABCE66E385D7CFB333DD274C215" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen was placed next to the nest mound. After orienting briefly, she immediately walked towards one of the entrance holes, and within a few seconds she disappeared into one of the entrances. The
<taxonomicName id="F6B4DEEAB39A97F461E48481FBD0129C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers, which guarded the entrances, were not seen to attack, catch or struggle with the invading parasite. After 3 h the observation was stopped, and until then,
<taxonomicName id="7A71818C1F3475E299A9AB3475A5CC7A" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers had not expelled the
<taxonomicName id="B22C48D7C8D36953F7EA00481796E1EE" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen.
</paragraph>
<paragraph id="6F18D995CF513DC50C53041A0EC0F190" pageNumber="389">
A second parasite queen was placed next to a
<taxonomicName id="809A14EC54E6D8A9D0247BE72E781CAD" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colony, which had closed the supernumerary nest entrances after the nuptial flight. In contrast, the parasite did not start searching for the nest entrance and we repeatedly (five times) placed her on the side of the nest mound before she finally, perhaps by chance, walked over the nest entrance. When crossing the entrance,
<taxonomicName id="547C2B16C7FFE54FD3142F283991885D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers attacked the parasite immediately. We collected the parasite queen and a dissection identified her as recently inseminated with developing ovaries.
</paragraph>
<paragraph id="0A272A1DEF709392F95AB5CC0E7266D8" pageNumber="389">
Introduction of parasite queen into a laboratory colony. To observe how
<taxonomicName id="9B7BD501FA8983F27CC5D98163E89E7D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens invade a
<taxonomicName id="3D13490CB373AD6650BA273774F6DF58" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colony, we maintained a nonparasitized
<taxonomicName id="EC2A8534C0B49889DD120DEB53F5F576" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
colony in the laboratory. Upon transfer to the artificial nest,
<taxonomicName id="69EAC38F4C8C7ABA47EC597DCB166BE5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers immediately covered the host queen with mycelial tufts, until she was completely hidden some 5 min later. To introduce the parasite queen, she was placed in a tube, which was connected to the fungus chamber. Quickly, she found her way out, headed directly towards the fungus garden, and immediately encountered the host queen. Then she started running in circles on the piece of fungus garden, under which the host queen was hidden. At that point, the host workers started chasing her, until one worker got hold of her petiole, and a second worker grabbed an antenna. The trio remained motionless for about 10 min, until both workers suddenly released the captive. Immediately, the
<taxonomicName id="C54EF28A1A56E0731B90C0FAC6DC7DA4" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen made a beeline for the host queen 's hiding place, where she was captured again. When being captured, the
<taxonomicName id="D7AFCAA808CCD228A3E6E666E4764321" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen assumed a characteristic position: she tucked her metasoma under the mesosoma, with the first gastric sternite touching the coxae. The combination of smooth body surface and broad, concave first gastric tergite provided little contact surface for attacking host workers, and their mandibles repeatedly slipped, upon which the host workers turned around and left. While being held by workers, the parasite queen was often antennated and licked at the base of the mandibles. After she was dumped ' on the refuse pile, she promptly aimed for the host queen. This cycle of capture and release was repeated overnight. Meanwhile, the
<taxonomicName id="CD52029C220CE943D158D752D5B9BDDF" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
worker removed the fungus garden fragments from the host queen, releasing her from her mycelium prison. Unfortunately, the first contact of host and parasite queen was not observed, but early the next morning, the
<taxonomicName id="5A0EEBD98D3496C2287FE6A40DD42406" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen was riding ' on the
<taxonomicName id="DC36FC91A96173EFE9047213752BA62B" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen, licking her mesosoma, petiole, postpetiole and metasoma. The
<taxonomicName id="31E61BC85848EFCFA5D65682A2C01578" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers frequently antennated and licked both queens, and the parasite queen was not attacked anymore. Subsequent observations revealed that the
<taxonomicName id="0CB2FC99040EB7C190AB11E7AE0D81AE" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queen was always riding ' on top of the
<taxonomicName id="57885EA4639B61AC8D972304FCD6D3DF" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
queen, and
<taxonomicName id="BAF0CC5002D3CE42FF534FA6936238AC" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
workers seemed to attend both queens equally.
</paragraph>
</subSubSection>
<subSubSection id="3F079BEFDAC82F35D4E925729CBA66FC" type="discussion">
<paragraph id="F63B4ACC3F1466251D7D3A874E5C4393" pageNumber="389">Discussion</paragraph>
<paragraph id="41EEE8BDE89AC825581A43E3EB78283F" lastPageNumber="390" pageNumber="389">
Several lines of evidence from natural history, behaviour and morphology identify
<taxonomicName id="614C99050AD6CB3AD3792AE7A4BFCA88" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
as an evolutionarily derived inquiline parasite of
<taxonomicName id="71C5512375E78B2519DBC0CC57BBF8E4" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
.
<taxonomicName id="03313FC27D84DBDF195821AF0F8A41C6" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
is functionally polygynous and host-queen tolerant. It does not seem to produce a worker caste. Parasite queens apparently suppress the production of host sexual offspring, effectively castrating the infested host colony. Mating takes place inside the host nest and mating flights of the parasite have not been observed and probably do not occur.
<taxonomicName id="43021DF2D6EA60650AF19E264B337D5C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="389" phylum="Arthropoda" rank="species" species="castrator">Alates of M. castrator</taxonomicName>
, although
<pageBreakToken id="240147C64D3A078E4D047A3F581FE944" pageNumber="390">fully</pageBreakToken>
alate and seemingly capable, do not fly. After mating, we opened artificial laboratory nests to test if queens and/or males disperse via flight and none did. Alates were encouraged to climb objects that could serve as potential launch pads (i.e. pencils), but they plunged down after reaching the tip, without becoming airborne. Thus, dispersing queens must walk to new host nests, severely limiting their dispersal.
</paragraph>
<paragraph id="361CCFF75CD457AFF6D3ACE567C0F43C" pageNumber="390">
Recently inseminated
<taxonomicName id="9C3B0090CE3D5580E5CF1A77366D480D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
queens found new colonies by invading host nests and showed elaborate behaviours related to securing adoption by the hosts. Compared with the host, parasite queens and males are reduced in body size and exhibit several morphological specializations known as the inquiline or anatomical parasite syndrome (
<bibRefCitation id="CFF6D3BD5C272A1DB210F7D45EA74CA9" author="Kutter, H." firstAuthor="Kutter" journalOrPublisher="Neujahrsblatt der Naturforschenden Gesellschaft in Zu ¨ rich" pagination="1 - 62" refId="ref11745" refString="Kutter, H. (1969) Die sozialparasitischen Ameisen der Schweiz. Neujahrsblatt der Naturforschenden Gesellschaft in Zu ¨ rich, 171, 1 - 62." title="Die sozialparasitischen Ameisen der Schweiz" volume="171" year="1969">Kutter, 1969</bibRefCitation>
;
<bibRefCitation id="109F1F2DEDFFC94672FE287991779564" author="Wilson, E. O." firstAuthor="Wilson" journalOrPublisher="Belknap Press of Harvard University Press, Cambridge, MA" refId="ref12619" refString="Wilson, E. O. (1971) The Insect Societies. Belknap Press of Harvard University Press, Cambridge, MA." title="The Insect Societies" year="1971">Wilson, 1971</bibRefCitation>
,
<bibRefCitation id="19E64C074C2D181CE286848F6AB26F23" author="Wilson, E. O." firstAuthor="Wilson" journalOrPublisher="Insectes Sociaux" pagination="316 - 334" refId="ref12643" refString="Wilson, E. O. (1984) Tropical social parasites in the ant genus Pheidole, with an analysis of the anatomical parasitic syndrome (Hymenoptera: Formicidae). Insectes Sociaux, 31, 316 - 334." title="Tropical social parasites in the ant genus Pheidole, with an analysis of the anatomical parasitic syndrome (Hymenoptera: Formicidae)" volume="31" year="1984">1984</bibRefCitation>
;
<bibRefCitation id="FB012550C8203175CFB18B31D33B16B9" author="Hölldobler, B" firstAuthor="Holldobler" journalOrPublisher="The Belknap Press of Harvard University Press, Cambridge, MA" refId="ref11556" refString="Holldobler, B. &amp; Wilson, E. O. (1990) The Ants. The Belknap Press of Harvard University Press, Cambridge, MA." title="The Ants" year="1990">Holldobler &amp; Wilson, 1990</bibRefCitation>
). The new parasite appears to have only one host,
<taxonomicName id="AB7AF927C4F408093BAD8CA835509295" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:31779" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus goeldii (Forel)" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="goeldii">M. goeldii</taxonomicName>
, even though other free-living congeners are present in the type locality. Lastly, even though the host ant is widespread and abundant throughout much of southern South America,
<taxonomicName id="B62E0FC084DFDEA9380354F1625ABA30" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
has been collected only twice, both times at the type locality. This suggests that parasite populations are probably few in number, small in size and patchily distributed. Intranidal mating, limited dispersal and small, isolated populations imply minimal gene flow between populations, and high levels of inbreeding within populations, conditions that have been postulated to exist for a number of other inquilines (
<bibRefCitation id="087CFB1365E66A856FA7BF80ACE0FA38" author="Kutter, H." firstAuthor="Kutter" journalOrPublisher="Neujahrsblatt der Naturforschenden Gesellschaft in Zu ¨ rich" pagination="1 - 62" refId="ref11745" refString="Kutter, H. (1969) Die sozialparasitischen Ameisen der Schweiz. Neujahrsblatt der Naturforschenden Gesellschaft in Zu ¨ rich, 171, 1 - 62." title="Die sozialparasitischen Ameisen der Schweiz" volume="171" year="1969">Kutter, 1969</bibRefCitation>
;
<bibRefCitation id="EF1EF6E70E262BC4092528205B87C2D9" author="Wilson, E. O." firstAuthor="Wilson" journalOrPublisher="Belknap Press of Harvard University Press, Cambridge, MA" refId="ref12619" refString="Wilson, E. O. (1971) The Insect Societies. Belknap Press of Harvard University Press, Cambridge, MA." title="The Insect Societies" year="1971">Wilson, 1971</bibRefCitation>
;
<bibRefCitation id="80A5CDFB8157215EC4A52B7901F37C37" author="Zamora-Munoz, C." etAl="et al." firstAuthor="Zamora-Munoz" journalOrPublisher="Sociobiology" pagination="1 - 19" title="Coevolution in the slave-parasite system Proformica longiseta - Rossomyrmex minuchae (Hymenoptera: Formicidae)" volume="42" year="2003">Zamora-Munoz et al., 2003</bibRefCitation>
;
<bibRefCitation id="E51CBC1FADE0201D002CD3AD59AF04B2" author="Buschinger, A." firstAuthor="Buschinger" journalOrPublisher="Myrmecological News" pagination="219 - 235" refId="ref11071" refString="Buschinger, A. (2009) Social parasitism among ants: a review (Hymenoptera: Formicidae). Myrmecological News, 12, 219 - 235." title="Social parasitism among ants: a review (Hymenoptera: Formicidae)" volume="12" year="2009">Buschinger, 2009</bibRefCitation>
). Inbreeding, however, has not yet been documented genetically in
<taxonomicName id="F8BB12DA362A27AF082BA1F6D2A019EE" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
(Rabeling, in preparation). Alternatively, it might be possible that the queens from a single nest are not necessarily closely related, because
<taxonomicName id="AE87F7B6F4BE1C0071FA2ED306984955" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
is functionally polygynous, and parasite queens seem to invade host colonies independently.
</paragraph>
<paragraph id="A9243D28509FC82FBB07FBE6DB23C040" pageNumber="390">
<taxonomicName id="FCB786C1FFB3742069EC0FEDFBDCD24D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
shows several additional morphological features related to its parasitic lifestyle (Table 3).
<taxonomicName id="435E40D8A26FD0768F99DEDC22C870FE" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
is the only inquiline known from the Lower
<taxonomicName id="626CE8BB202AF389967D52FDCA9FA0F9" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2269" class="Insecta" family="Formicidae" kingdom="Animalia" lsidName-HNS="Attini Smith" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="tribe" tribe="Attini">Attini</taxonomicName>
and, interestingly, it shares convergently evolved derived morphological characters with another distantly related attine workerless inquiline,
<taxonomicName id="D7D55EAFC19903A6264FB4E4B4A59897" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">P. argentina</taxonomicName>
(Table 3). In both species, males and females exhibit a reduced palp formula of 3,2 (Figs 1G, 2G), whereas the plesiotypic attine palp formula is 4,2 (Figs 1H, 2H) (
<bibRefCitation id="CD23EEA9331EC7B9BB15662047726790" author="Gallardo, A." firstAuthor="Gallardo" journalOrPublisher="Buenos Aires" pagination="317 - 344" refId="ref11483" refString="Gallardo, A. (1916) Notes syste ' matiques et e ' thologiques sur les fourmis attines de la Re ' publique Argentine. Anales del Museo Nacional de Historia Natural, Buenos Aires, 28, 317 - 344." title="Notes syste ' matiques et e ' thologiques sur les fourmis attines de la Re ' publique Argentine. Anales del Museo Nacional de Historia Natural" volume="28" year="1916">Gallardo, 1916</bibRefCitation>
;
<bibRefCitation id="F04FF756FC8A3916227C4033F02067FC" author="Schultz, T. R." etAl="et al." firstAuthor="Schultz" journalOrPublisher="Insectes Sociaux" pagination="457 - 471" refId="ref12339" refString="Schultz, T. R., Bekkevold, D. &amp; Boomsma, J. J. (1998) Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants. Insectes Sociaux, 45, 457 - 471." title="Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants" volume="45" year="1998">Schultz et al., 1998</bibRefCitation>
). Elsewhere in the
<taxonomicName id="F788583A926B5550ED198563D8A94D2D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2269" class="Insecta" family="Formicidae" kingdom="Animalia" lsidName-HNS="Attini Smith" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="tribe" tribe="Attini">Attini</taxonomicName>
, the reduced palp formula is found only in the free-living genus
<taxonomicName id="63DA71DEEDD190E68B94CD94A2E2DC38" authority="Mayr 1865" authorityName="Mayr" authorityYear="1865" class="Insecta" family="Formicidae" genus="Apterostigma" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="genus">Apterostigma Mayr 1865</taxonomicName>
(
<bibRefCitation id="7D8B4BD49238C48CDB5B82643CA3015D" author="Kusnezov, N." firstAuthor="Kusnezov" journalOrPublisher="Universidad de Cochabamba" pagination="62 - 70" refId="ref11668" refString="Kusnezov, N. (1951) Los segmentos palpales en hormigas. Folia Universitaria, Universidad de Cochabamba, 5, 62 - 70." title="Los segmentos palpales en hormigas. Folia Universitaria" volume="5" year="1951">Kusnezov, 1951</bibRefCitation>
,
<bibRefCitation id="EB2EA9EA6F29F20817623B75043D2A91" author="Kusnezov, N." firstAuthor="Kusnezov" journalOrPublisher="Zoologischer Anzeiger" pagination="28 - 38" refId="ref11694" refString="Kusnezov, N. (1954) Phyletische Bedeutung der Maxillar- und Labialtaster der Ameisen. Zoologischer Anzeiger, 153, 28 - 38." title="Phyletische Bedeutung der Maxillar- und Labialtaster der Ameisen" volume="153" year="1954">1954</bibRefCitation>
;
<bibRefCitation id="BD37FC83FF890B559882FF49C31DEC95" author="Schultz, T. R." firstAuthor="Schultz" journalOrPublisher="Memories of the American Entomological Institute" pagination="425 - 436" refId="ref12222" refString="Schultz, T. R. (2007) The fungus-growing ant genus Apterostigma in Dominican amber. Memories of the American Entomological Institute, 80, 425 - 436." title="The fungus-growing ant genus Apterostigma in Dominican amber" volume="80" year="2007">Schultz, 2007</bibRefCitation>
), where it evolved independently. In addition, in
<taxonomicName id="3E6A07D037658FD85A8BC5F5592E9B6B" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
the number of antennal segments is reduced from the plesiotypic attine condition of 11 segments in the females and 13 in the males to 11 segments in both sexes. The reduction of segments in
<taxonomicName id="AD9C66708417BED7230E6C84824B3C99" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
and
<taxonomicName id="67AA4523D3CBD7FA5107A9F60430D052" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">P. argentina</taxonomicName>
suggests that both species are evolutionarily derived inquilines. According to Wilson 's (1984) character analysis of nine presumably independently evolved inquilines in the genus
<taxonomicName id="61919C28FD5CB4CFB6DA757BB029E28E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:24885" authority="Westwood" authorityName="Westwood" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" lsidName-HNS="Pheidole Westwood" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="genus">Pheidole Westwood</taxonomicName>
, the reduction of antennal segmentation occurred secondarily during parasite evolution, preceded by the loss of the worker caste and other morphological reductions, like the reduction in size and body sculpture.
</paragraph>
<paragraph id="6D582E5B10CC59A8020007B7AF347542" pageNumber="390">
The wing morphology of
<taxonomicName id="1916C950FF07429E5DA27BD6574907EA" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
provides indirect support for the functional wing morphology hypothesis of the
</paragraph>
<paragraph id="E4526A89E6A865CB18E2325EF991EBF6" pageNumber="390">
<taxonomicName id="317E9FD75A9E336E5F3CA2E7D03D6B98" pageNumber="390" rank="tribe" tribe="Paleoattini">Paleoattini</taxonomicName>
(sensu
<bibRefCitation id="76A6E27A8FF790B146C0A3B97B604101" author="Kusnezov, N." firstAuthor="Kusnezov" journalOrPublisher="Acta Zoologica Lilloana" pagination="25 - 186" refId="ref11719" refString="Kusnezov, N. (1963) Zoogeografia de las hormigas en Sudame ' rica. Acta Zoologica Lilloana, 19, 25 - 186." title="Zoogeografia de las hormigas en Sudame ' rica" volume="19" year="1963">Kusnezov, 1963</bibRefCitation>
;
<bibRefCitation id="DAB6504B03AA1ED7C3FAED4649044F68" author="Schultz, T. R." firstAuthor="Schultz" journalOrPublisher="Systematic Entomology" pagination="337 - 370" refId="ref12294" refString="Schultz, T. R. &amp; Meier, R. (1995) A phylogenetic analysis of the fungus-growing ants (Hymenoptera: Formicidae: Attini) based on morphological characters of the larvae. Systematic Entomology, 20, 337 - 370." title="A phylogenetic analysis of the fungus-growing ants (Hymenoptera: Formicidae: Attini) based on morphological characters of the larvae" volume="20" year="1995">Schultz &amp; Meier, 1995</bibRefCitation>
) (
<bibRefCitation id="64BAB1C1CAF1CC20152551F9E6377F14" author="Fernandez-Marin, H." etAl="et al." firstAuthor="Fernandez-Marin" journalOrPublisher="Journal of Natural History" pagination="1735 - 1743" refId="ref11422" refString="Fernandez-Marin, H., Zimmerman, J. K., Wcislo, W. T. &amp; Rehner, S. A. (2005) Colony foundation, nest architecture, and demography of a basal fungus-growing ant, Mycocepurus smithii (Hymenoptera: Formicidae). Journal of Natural History, 39, 1735 - 1743." title="Colony foundation, nest architecture, and demography of a basal fungus-growing ant, Mycocepurus smithii (Hymenoptera: Formicidae)" volume="39" year="2005">Fernandez-Marin et al., 2005</bibRefCitation>
).
<bibRefCitation id="55931F18AE40A16B9BA6C430294E9269" author="Emery, C." firstAuthor="Emery" journalOrPublisher="Annales de la Société Entomologique de Belgique" pagination="250 - 262" refId="ref11326" refString="Emery, C. (1913) Études sur les Myrmicinae V. Les genres des Attini; descriptions de nouvelles formes de Mycocepurus et de Myrmicocrypta. Annales de la Société Entomologique de Belgique, 57, 250 - 262." title="Études sur les Myrmicinae V. Les genres des Attini; descriptions de nouvelles formes de Mycocepurus et de Myrmicocrypta" volume="57" year="1913">Emery (1913)</bibRefCitation>
first noted that the forewings of queens of
<taxonomicName id="B4229944CCC79802990E75F051CB2B9B" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2394" authority="F. Smith" authorityName="F. Smith" class="Insecta" family="Formicidae" genus="Myrmicocrypta" kingdom="Animalia" lsidName-HNS="Myrmicocrypta Smith" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="genus">Myrmicocrypta F. Smith</taxonomicName>
bear a small clear spot, or fenestra, which is not covered with fine setae and lacks pigmentation. This fenestra is also present in other paleoattines (
<bibRefCitation id="38920F9F7747003E217F9170EFA536F1" author="Fernandez-Marin, H." etAl="et al." firstAuthor="Fernandez-Marin" journalOrPublisher="Journal of Natural History" pagination="1735 - 1743" refId="ref11422" refString="Fernandez-Marin, H., Zimmerman, J. K., Wcislo, W. T. &amp; Rehner, S. A. (2005) Colony foundation, nest architecture, and demography of a basal fungus-growing ant, Mycocepurus smithii (Hymenoptera: Formicidae). Journal of Natural History, 39, 1735 - 1743." title="Colony foundation, nest architecture, and demography of a basal fungus-growing ant, Mycocepurus smithii (Hymenoptera: Formicidae)" volume="39" year="2005">Fernandez-Marin et al., 2005</bibRefCitation>
;
<bibRefCitation id="3C2F6ACA6EEEF8018AD0A9E52D401E64" author="Schultz, T. R." firstAuthor="Schultz" journalOrPublisher="Memories of the American Entomological Institute" pagination="425 - 436" refId="ref12222" refString="Schultz, T. R. (2007) The fungus-growing ant genus Apterostigma in Dominican amber. Memories of the American Entomological Institute, 80, 425 - 436." title="The fungus-growing ant genus Apterostigma in Dominican amber" volume="80" year="2007">Schultz, 2007</bibRefCitation>
). During colony founding, queens of
<taxonomicName id="817158140D3FA6DFE8D69DBF4A5ACFE2" pageNumber="390" rank="tribe" tribe="Paleoattini">Paleoattini</taxonomicName>
inoculate their fungus garden on their shed forewings, and fix the forewing under the ceiling of the new nest chamber to grow the fungus cultivar (
<bibRefCitation id="3D0FACB0CF841027DA88E046CF82238D" author="Fernandez-Marin, H." etAl="et al." firstAuthor="Fernandez-Marin" journalOrPublisher="Biological Journal of the Linnean Society" pagination="39 - 48" refId="ref11368" refString="Fernandez-Marin, H., Zimmerman, J. K. &amp; Wcislo, W. T. (2004) Ecological traits and evolutionary sequence of the nest establishment in fungus-growing ants (Hymenoptera, Formicidae, Attini). Biological Journal of the Linnean Society, 81, 39 - 48." title="Ecological traits and evolutionary sequence of the nest establishment in fungus-growing ants (Hymenoptera, Formicidae, Attini)" volume="81" year="2004">Fernandez-Marin et al., 2004</bibRefCitation>
). The fenestra was hypothesized to serve some unspecified function during nest founding (
<bibRefCitation id="BE475769D9D52C9E4A8A4A4EF74CCF78" author="Fernandez-Marin, H." etAl="et al." firstAuthor="Fernandez-Marin" journalOrPublisher="Journal of Natural History" pagination="1735 - 1743" refId="ref11422" refString="Fernandez-Marin, H., Zimmerman, J. K., Wcislo, W. T. &amp; Rehner, S. A. (2005) Colony foundation, nest architecture, and demography of a basal fungus-growing ant, Mycocepurus smithii (Hymenoptera: Formicidae). Journal of Natural History, 39, 1735 - 1743." title="Colony foundation, nest architecture, and demography of a basal fungus-growing ant, Mycocepurus smithii (Hymenoptera: Formicidae)" volume="39" year="2005">Fernandez-Marin et al., 2005</bibRefCitation>
). Interestingly, the clear spot is absent in
<taxonomicName id="E5B30C5C22A9C64EDBD1105958BA188C" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
wings, whereas it is present in all
<taxonomicName id="4BAB99B8CDAEAF2316F9FA5BCB628230" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2393" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus Forel" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="genus">Mycocepurus</taxonomicName>
species. The loss of the fenestra in obligate social parasites, which do not found colonies independently, suggests that it indeed has an adaptive function during independent nest founding, because otherwise the fenestra would be maintained by natural selection in obligate inquilines. Potentially, the clear area is easier to clean before inoculation with a novel cultivar, or it may provide a tactile or even visual cue to the queen where to place the inoculum. It would be interesting to test if parasite queens contribute a fungal inoculum to the new host colony [suggested by
<bibRefCitation id="307038181583D7FA60B4CED6A19CF4DD" author="Schultz, T. R." etAl="et al." firstAuthor="Schultz" journalOrPublisher="Insectes Sociaux" pagination="457 - 471" refId="ref12339" refString="Schultz, T. R., Bekkevold, D. &amp; Boomsma, J. J. (1998) Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants. Insectes Sociaux, 45, 457 - 471." title="Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants" volume="45" year="1998">Schultz et al. (1998)</bibRefCitation>
], and if the infrabuccal cavity of socially parasitic attines experienced morphological modifications or reductions.
</paragraph>
<paragraph id="DA83E279294DA96BAA2290BB817AF1E1" pageNumber="390">
Comparing
<taxonomicName id="F7FF530F018C4C9E8DE2BF9C28CF8F0D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
with other attine inquilines is instructive and provides insight into the life history evolution of social parasites (Table 3). Like
<taxonomicName id="BA04B4BA72840734E76F4FF436D93BCF" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
,
<taxonomicName id="86AA7AE6020F7D977FF36A5D1A1346BC" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">P. argentina</taxonomicName>
appears to be a phylogenetically derived, workerless inquiline and shares some morphological and life history traits with
<taxonomicName id="6B198D74E9C852EE1044EDD83B32210E" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
(see Table 3).
<taxonomicName id="21F2A615C40E9A0F89C0F40FAF19A50D" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:222191" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex ameliae De Souza, Soares &amp; Della Lucia" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="ameliae">Acromyrmex ameliae</taxonomicName>
and
<taxonomicName id="D9B835A961B540636B510AD9E54689A7" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:152541" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex insinuator Schultz, Bekkevold &amp; Boomsma" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="insinuator">A. insinuator</taxonomicName>
, however, are much different. Like
<taxonomicName id="6E462FEEC600276A3F562E746C734B9A" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
and
<taxonomicName id="4865826A8E1CD82F6BE152D708F010C1" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">P. argentina</taxonomicName>
, the
<taxonomicName id="4C22FF44F7DD8E7EA0F151A9D5D8CFA6" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2323" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex Mayr" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species">Acromyrmex inquilines</taxonomicName>
are each associated with a single host species. However, they show much less morphological and life history specialization (
<bibRefCitation id="87997F0692075E5A49BA6253FBBF8582" author="Schultz, T. R." etAl="et al." firstAuthor="Schultz" journalOrPublisher="Insectes Sociaux" pagination="457 - 471" refId="ref12339" refString="Schultz, T. R., Bekkevold, D. &amp; Boomsma, J. J. (1998) Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants. Insectes Sociaux, 45, 457 - 471." title="Acromyrmex insinuator new species: an incipient social parasite of fungus-growing ants" volume="45" year="1998">Schultz et al., 1998</bibRefCitation>
;
<bibRefCitation id="69600B38E2015DCC367D347BE3851E04" author="de Souza, D. J." etAl="et al." firstAuthor="de Souza" journalOrPublisher="Insect Science" pagination="251 - 257" refId="ref12382" refString="de Souza, D. J., Soares, I. M. F. &amp; Della Lucia, T. M. C. (2007) Acromyrmex ameliae sp. n. (Hymenoptera: Formicidae): a new social parasite of leaf-cutting ants in Brazil. Insect Science, 14, 251 - 257." title="Acromyrmex ameliae sp. n. (Hymenoptera: Formicidae): a new social parasite of leaf-cutting ants in Brazil" volume="14" year="2007">de Souza et al., 2007</bibRefCitation>
). For example,
<taxonomicName id="6DEC176C2D829A0CE50B1E000E82987A" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:222191" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex ameliae De Souza, Soares &amp; Della Lucia" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="ameliae">A. ameliae</taxonomicName>
and
<taxonomicName id="5ED808BEFFCA711D42EB0CDA6117DAA5" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:152541" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex insinuator Schultz, Bekkevold &amp; Boomsma" order="Hymenoptera" pageNumber="390" phylum="Arthropoda" rank="species" species="insinuator">A. insinuator</taxonomicName>
lack the palpal and antennal segment reductions and other morphological features defining the inquiline syndrome(Table 3). Both species have mating flights, sometimes synchronously with host alates. Both species produce a worker caste, and parasitize host colonies at higher frequencies than most inquilines for which we have data (
<bibRefCitation id="070C737B4C36F353DC4C8C648A61307F" author="Bekkevold, D." firstAuthor="Bekkevold" journalOrPublisher="Journal of Evolutionary Biology" pagination="615 - 623" refId="ref10761" refString="Bekkevold, D. &amp; Boomsma, J. J. (2000) Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator. Journal of Evolutionary Biology, 13, 615 - 623." title="Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator" volume="13" year="2000">Bekkevold &amp; Boomsma, 2000</bibRefCitation>
;
<bibRefCitation id="5D4DE1B25D1C8CD38A3ABD2816EA7144" author="de Souza, D. J." etAl="et al." firstAuthor="de Souza" journalOrPublisher="Insect Science" pagination="251 - 257" refId="ref12382" refString="de Souza, D. J., Soares, I. M. F. &amp; Della Lucia, T. M. C. (2007) Acromyrmex ameliae sp. n. (Hymenoptera: Formicidae): a new social parasite of leaf-cutting ants in Brazil. Insect Science, 14, 251 - 257." title="Acromyrmex ameliae sp. n. (Hymenoptera: Formicidae): a new social parasite of leaf-cutting ants in Brazil" volume="14" year="2007">de Souza et al., 2007</bibRefCitation>
). Because mating flights are retained, dispersal is probably much less restricted, and inbreeding may be slight or nonexistent, as was demonstrated for
<taxonomicName id="02CBC84DBFC8C13D190CD9DC234515D5" authority="(Sumner et al., 2004a)" baseAuthorityName="Sumner" baseAuthorityYear="2004" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="insinuator">
A. insinuator (
<bibRefCitation id="7C2409246EAC640F47A7D3EF06F86773" author="Sumner, S." etAl="et al." firstAuthor="Sumner" journalOrPublisher="Nature" pagination="35 - 36" refId="ref12442" refString="Sumner, S., Hughes, W. O. H., Pedersen, J. &amp; Boomsma, J. J. (2004 a) Social parasite queens abandon multiple mating. Nature, 428, 35 - 36." title="Social parasite queens abandon multiple mating" volume="428" year="2004" yearSuffix="a">Sumner et al., 2004a</bibRefCitation>
)
</taxonomicName>
. These traits and other aspects of their biology (summarized in Table 3) strongly suggest that these species are recently evolved inquiline social parasites that have yet to develop the morphological inquiline syndrome and full workerless parasitism.
</paragraph>
<paragraph id="2575011C70A2F8C7041039EC4ABF100D" lastPageNumber="391" pageNumber="390">
It should be stressed that the attine inquilines are not closely related to each other (
<bibRefCitation id="FDE31023642EFC8D26C5C54DF7762178" author="Sumner, S." etAl="et al." firstAuthor="Sumner" journalOrPublisher="Insectes Sociaux" pagination="37 - 42" refId="ref12486" refString="Sumner, S., Aanen, D. K., Delabie, J. &amp; Boomsma, J. J. (2004 b) The evolution of social parasitism in leaf-cutting ants. Insectes Sociaux, 51, 37 - 42." title="The evolution of social parasitism in leaf-cutting ants" volume="51" year="2004" yearSuffix="b">Sumner et al., 2004b</bibRefCitation>
;
<bibRefCitation id="1E745F559CE63F28D3433F65F3FFE9C2" author="Schultz, T. R." firstAuthor="Schultz" journalOrPublisher="Proceedings of the National Academy of Sciences United States of America" pagination="5435 - 5440" refId="ref12253" refString="Schultz, T. R. &amp; Brady, S. G. (2008) Major evolutionary transitions in ant agriculture. Proceedings of the National Academy of Sciences United States of America, 105, 5435 - 5440." title="Major evolutionary transitions in ant agriculture" volume="105" year="2008">Schultz &amp; Brady, 2008</bibRefCitation>
) and what is known of their life histories provides additional evidence of separate and idiosyncratic evolutionary trajectories. For instance, limited evidence suggests that the derived inquiline
<taxonomicName id="0DD6D1585C514C8A1A3043D99F80861B" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">P. argentina</taxonomicName>
may be host-queen intolerant(
<bibRefCitation id="F35A6AD20B0AD3C4924FC122366F1F11" author="Bruch, C." firstAuthor="Bruch" journalOrPublisher="Buenos Aires" pagination="341 - 360" refId="ref10978" refString="Bruch, C. (1928) Estudios mirmecolo ' gicos. Anales del Museo Nacional de Historia Natural, Buenos Aires, 34, 341 - 360." title="Estudios mirmecolo ' gicos. Anales del Museo Nacional de Historia Natural" volume="34" year="1928">Bruch, 1928</bibRefCitation>
). In addition, there is some indication that
<pageBreakToken id="8766582F80CE193094B64FD91D86D278" pageNumber="391">the</pageBreakToken>
fungus garden in
<taxonomicName id="9C7D045EF05A8D624F67AE5BCDB19654" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" order="Hymenoptera" phylum="Arthropoda" rank="species" species="argentina">P. argentina</taxonomicName>
colonies breaks down after a parasite brood has been reared by the host workers (
<bibRefCitation id="9EF12E4950D4429E7F779EF419E826EE" author="Bruch, C." firstAuthor="Bruch" journalOrPublisher="Buenos Aires" pagination="341 - 360" refId="ref10978" refString="Bruch, C. (1928) Estudios mirmecolo ' gicos. Anales del Museo Nacional de Historia Natural, Buenos Aires, 34, 341 - 360." title="Estudios mirmecolo ' gicos. Anales del Museo Nacional de Historia Natural" volume="34" year="1928">Bruch, 1928</bibRefCitation>
). Thus,
<taxonomicName id="2330022AA17028A07EE6C87CC464C25F" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2395" class="Insecta" family="Formicidae" genus="Pseudoatta" kingdom="Animalia" lsidName-HNS="Pseudoatta Gallardo" order="Hymenoptera" pageNumber="391" phylum="Arthropoda" rank="genus">Pseudoatta</taxonomicName>
colonies may be quite short lived, perhaps even semelparous. Curiously, evidence has accumulated that the recently evolved inquiline
<taxonomicName id="27666F03FA80EBFF8B812C1E8379E055" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:152541" class="Insecta" family="Formicidae" genus="Acromyrmex" kingdom="Animalia" lsidName-HNS="Acromyrmex insinuator Schultz, Bekkevold &amp; Boomsma" order="Hymenoptera" pageNumber="391" phylum="Arthropoda" rank="species" species="insinuator">A. insinuator</taxonomicName>
may also be semelparous (
<bibRefCitation id="1057B29798BDAE91175456CEAB0B8150" author="Bekkevold, D." firstAuthor="Bekkevold" journalOrPublisher="Journal of Evolutionary Biology" pagination="615 - 623" refId="ref10761" refString="Bekkevold, D. &amp; Boomsma, J. J. (2000) Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator. Journal of Evolutionary Biology, 13, 615 - 623." title="Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator" volume="13" year="2000">Bekkevold &amp; Boomsma, 2000</bibRefCitation>
), even though parasitized colonies retain a fertile host queen and could theoretically last as long as she lives and lays eggs. Nonetheless,
<bibRefCitation id="906BF7A292BE7D8C4FC93AE8A1776775" author="Bekkevold, D." firstAuthor="Bekkevold" journalOrPublisher="Journal of Evolutionary Biology" pagination="615 - 623" refId="ref10761" refString="Bekkevold, D. &amp; Boomsma, J. J. (2000) Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator. Journal of Evolutionary Biology, 13, 615 - 623." title="Evolutionary transition to a semelparous life history in the socially parasitic ant Acromyrmex insinuator" volume="13" year="2000">Bekkevold &amp; Boomsma (2000)</bibRefCitation>
provided evidence that the colony 's fungus garden breaks down after a generation of parasite sexuals is produced. In contrast, in
<taxonomicName id="F2E1110A850DFD9105F4B28081D16391" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="391" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
colonies the host queen is retained and the fungus garden appears to remain healthy after the production of parasite sexuals. Thus, the colonies may last for several parasite reproductive cycles. Although one might expect that colony longevity should be favoured by natural selection under a broad range of conditions, such counterintuitive examples remind us that we are far from fully understanding the complex and diverse life history phenomena grouped under the term inquiline social parasitism '.
</paragraph>
<paragraph id="E269C3C041BB89AE8E1CE330785E1C29" pageNumber="391">
In conclusion, obligate social parasites are prime examples for the study of convergent evolution. Over 80 inquiline species are known currently (
<bibRefCitation id="C09BA59AF4E97132AFD20228E68345CF" author="Holldobler, B" firstAuthor="Holldobler" journalOrPublisher="The Belknap Press of Harvard University Press, Cambridge, MA" refId="ref11556" refString="Holldobler, B. &amp; Wilson, E. O. (1990) The Ants. The Belknap Press of Harvard University Press, Cambridge, MA." title="The Ants" year="1990">Holldobler &amp; Wilson, 1990</bibRefCitation>
;
<bibRefCitation id="2A853C4381F1F1CD30A3E52010A101B3" author="Huang, M. H." firstAuthor="Huang" journalOrPublisher="Ecological Entomology" pagination="589 - 596" refId="ref11585" refString="Huang, M. H. &amp; Dornhaus, A. (2008) Are ant social parasites always closely related to their hosts? Host characteristics in different parasitism types and a test of Emery ' s rule. Ecological Entomology, 33, 589 - 596." title="Are ant social parasites always closely related to their hosts? Host characteristics in different parasitism types and a test of Emery ' s rule" volume="33" year="2008">Huang &amp; Dornhaus, 2008</bibRefCitation>
), and mapping parasite lineages onto recent phylogenies (
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;
<bibRefCitation id="E5D53CFB853DA9CCC97208CB859628A6" author="Moreau, C. S." etAl="et al." firstAuthor="Moreau" journalOrPublisher="Science" pagination="101 - 104" refId="ref11857" refString="Moreau, C. S., Bell, C. D., Vila, R., Archibald, S. B. &amp; Pierce, N. E. (2006) Phylogeny of the ants: diversification in the age of angiosperms. Science, 312, 101 - 104." title="Phylogeny of the ants: diversification in the age of angiosperms" volume="312" year="2006">Moreau et al., 2006</bibRefCitation>
;
<bibRefCitation id="120C096A2C1E4AE3495FADAB1F5EC981" author="Rabeling, C." etAl="et al." firstAuthor="Rabeling" journalOrPublisher="Proceedings of the National Academy of Sciences United States of America" pagination="14913 - 14917" refId="ref12079" refString="Rabeling, C., Brown, J. M. &amp; Verhaagh, M. (2008) Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences United States of America, 105, 14913 - 14917." title="Newly discovered sister lineage sheds light on early ant evolution" volume="105" year="2008">Rabeling et al., 2008</bibRefCitation>
) illustrates that inquilines evolved at least 30 times convergently. A higher taxonomic resolution of speciose groups will probably increase the number of independent parasite origins. In fungus-gardening ants, two groups of inquilines evolved: incipient and evolutionarily derived social parasites.
<taxonomicName id="E9C573B96D99614216336BDDEBFBC21A" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="391" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
is a derived parasite, showing multiple morphological and life history traits of the inquiline syndrome (
<bibRefCitation id="60AA36B22CB21BEE3C58FA4F8AA04FC7" author="Kutter, H." firstAuthor="Kutter" journalOrPublisher="Neujahrsblatt der Naturforschenden Gesellschaft in Zu ¨ rich" pagination="1 - 62" refId="ref11745" refString="Kutter, H. (1969) Die sozialparasitischen Ameisen der Schweiz. Neujahrsblatt der Naturforschenden Gesellschaft in Zu ¨ rich, 171, 1 - 62." title="Die sozialparasitischen Ameisen der Schweiz" volume="171" year="1969">Kutter, 1969</bibRefCitation>
;
<bibRefCitation id="5756E5796C2ED0898EBA694F52BD3B6E" author="Wilson, E. O." firstAuthor="Wilson" journalOrPublisher="Belknap Press of Harvard University Press, Cambridge, MA" refId="ref12619" refString="Wilson, E. O. (1971) The Insect Societies. Belknap Press of Harvard University Press, Cambridge, MA." title="The Insect Societies" year="1971">Wilson, 1971</bibRefCitation>
,
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).
<taxonomicName id="AB5ECF8204E25BD566A9E92DB7135DE6" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="391" phylum="Arthropoda" rank="species" species="castrator">Mycocepurus castrator</taxonomicName>
is the first social parasite of the Lower
<taxonomicName id="71CCC0F83B5E957E9D33BB85825CA3D1" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:2269" class="Insecta" family="Formicidae" kingdom="Animalia" lsidName-HNS="Attini Smith" order="Hymenoptera" pageNumber="391" phylum="Arthropoda" rank="tribe" tribe="Attini">Attini</taxonomicName>
and an additional social parasite from a subtropical habitat, a geographical region from which social parasites are little known. A phylogenetic and population genetic study is underway and will answer additional questions about the evolutionary origin of
<taxonomicName id="43879B7AA758D0EF800F7E8AC1076CD7" LSID-HNS="urn:lsid:biosci.ohio-state.edu:osuc_concepts:268617" class="Insecta" family="Formicidae" genus="Mycocepurus" kingdom="Animalia" lsidName-HNS="Mycocepurus castrator Rabeling &amp; Bacci" order="Hymenoptera" pageNumber="391" phylum="Arthropoda" rank="species" species="castrator">M. castrator</taxonomicName>
(Rabeling, in preparation).
</paragraph>
</subSubSection>
</treatment>
</document>