<documentid="5805A158ED4B2C8818B8108DB2894D8E"ID-CLB-Dataset="22422"ID-DOI="10.5852/ejt.2015.120"ID-GBIF-Dataset="5fb91ffb-69b9-4e0f-9626-08d75ab0cba8"ID-ISSN="2118-9773"ID-Zenodo-Dep="3780152"ID-ZooBank="54714320-5726-44CB-8FF5-60E0B984873D"IM.illustrations_approvedBy="felipe"IM.materialsCitations_approvedBy="felipe"IM.metadata_approvedBy="felipe"IM.taxonomicNames_approvedBy="felipe"checkinTime="1588343872265"checkinUser="carolina"docAuthor="Boudinot, Brendon E."docDate="2015"docId="038E878CFF8CB148FD9CFCF6FBCF1986"docLanguage="en"docName="ejt-2015-120.pdf.imf"docOrigin="European Journal of Taxonomy 120"docStyle="DocumentStyle:EF2B578F1D15862ADE45B0C07C620911.14:EJT.2018-.journal_article.type1"docStyleId="EF2B578F1D15862ADE45B0C07C620911"docStyleName="EJT.2018-.journal_article.type1"docStyleVersion="14"docTitle="Martialis heureka Rabeling & Verhaagh 2008"docType="treatment"docVersion="7"lastPageNumber="46"masterDocId="FFB7FFF4FFA8B165FFB0FFEBFF941F34"masterDocTitle="Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages"masterLastPageNumber="62"masterPageNumber="1"pageNumber="37"updateTime="1732491263130"updateUser="ExternalLinkService"zenodo-license-document="CC-BY-4.0"zenodo-license-figures="CC-BY-4.0"zenodo-license-treatments="UNSPECIFIED">
<mods:titleid="D00A5E07B3C666177AF61BAB720E9232">Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages</mods:title>
<taxonomicNameid="4C274D19FF8CB141FD9CFCF6FBC01C0C"ID-CoL="CM3Y3"authority="Rabeling & Verhaagh, 2008"authorityName="Rabeling & Verhaagh"authorityYear="2008"box="[556,1108,797,824]"class="Insecta"family="Formicidae"genus="Martialis"higherTaxonomySource="GBIF"kingdom="Animalia"order="Hymenoptera"pageId="36"pageNumber="37"phylum="Arthropoda"rank="species"species="heureka">
<bibRefCitationid="EFB64B6BFF8CB141FCBCFCF5FBC01C0C"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[780,1108,798,824]"pageId="36"pageNumber="37"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">Rabeling & Verhaagh, 2008</bibRefCitation>
<figureCitationid="131C2A1FFF8CB141FD4EFCAAFC141C6F"box="[766,896,833,859]"captionStart-0="Fig"captionStart-1="Fig"captionStartId-0="40.[227,270,1966,1992]"captionStartId-1="41.[227,270,1896,1922]"captionTargetBox-0="[270,1395,304,1931]"captionTargetBox-1="[234,1428,302,1856]"captionTargetId-0="figure@40.[270,1395,302,1931]"captionTargetId-1="figure@41.[234,1428,302,1882]"captionTargetPageId-0="40"captionTargetPageId-1="41"captionText-0="Fig. 11. Martialis heureka Rabeling & Verhaagh, 2008, male, photomicrographs. A. Head, frontal view. B. Head, anteroventral oblique view. C. Body, lateral view. D. Body, dorsal view. Scale bars: A–B = 0.2 mm, C–D = 0.5 mm."captionText-1="Fig. 12. Martialis heureka Rabeling & Verhaagh, 2008, male, wing photomicrographs and genitalia illustrations, genital membranes not shown. A. Forewing. B. Hindwing. C. Abdominal sternum IX, ventral view. D. Genital capsule, dorsal view. E. Genital capsule, ventral view. F. Genital capsule, lateral view. G. Volsella and paramere, mesal view. H. Penisvalva in situ, mesal view. Scale bars: A–B = 0.5 mm, C–H = 0.1 mm. Abbreviations: see Material and Methods."figureDoi-0="http://doi.org/10.5281/zenodo.3780170"figureDoi-1="http://doi.org/10.5281/zenodo.3780172"httpUri-0="https://zenodo.org/record/3780170/files/figure.png"httpUri-1="https://zenodo.org/record/3780172/files/figure.png"pageId="36"pageNumber="37">Figs 11–12</figureCitation>
<taxonomicNameid="4C274D19FF8CB141FF53FC69FC751CA8"authority="Rabeling & Verhaagh, in Rabeling et al. 2008"authorityName="Rabeling & Verhaagh, in Rabeling"authorityYear="2008"box="[227,993,898,924]"class="Insecta"family="Formicidae"genus="Martialis"higherTaxonomySource="GBIF"kingdom="Animalia"order="Hymenoptera"pageId="36"pageNumber="37"phylum="Arthropoda"rank="species"species="heureka">
<bibRefCitationid="EFB64B6BFF8CB141FD5CFC69FC751CA8"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[748,993,898,924]"pageId="36"pageNumber="37"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">
INDICES. CI 0.94–1.02, CS 0.35–0.41, SEI 83.6–86.2, SI 55.6–58.4, EI 83.0–85.6, EYE 88.1–88.7, MI 26.7–27.8, OBI 80.1–82.8, OMI 3.46–4.74, MNI 2.00–2.26, MTI 79.9–83.2, FI 88.3–91.3, PTI 76.9– 83.2. Small, but body variable in overall size (
<figureCitationid="131C2A1FFF8CB141FCB1FAA6FCFF1A53"box="[769,875,1357,1383]"captionStart="Fig"captionStartId="40.[227,270,1966,1992]"captionTargetBox="[270,1395,304,1931]"captionTargetId="figure@40.[270,1395,302,1931]"captionTargetPageId="40"captionText="Fig. 11. Martialis heureka Rabeling & Verhaagh, 2008, male, photomicrographs. A. Head, frontal view. B. Head, anteroventral oblique view. C. Body, lateral view. D. Body, dorsal view. Scale bars: A–B = 0.2 mm, C–D = 0.5 mm."figureDoi="http://doi.org/10.5281/zenodo.3780170"httpUri="https://zenodo.org/record/3780170/files/figure.png"pageId="36"pageNumber="37">Fig. 11C</figureCitation>
<figureCitationid="131C2A1FFF8CB141FE85FA64FE1C1A9D"box="[309,392,1423,1449]"captionStart="Fig"captionStartId="40.[227,270,1966,1992]"captionTargetBox="[270,1395,304,1931]"captionTargetId="figure@40.[270,1395,302,1931]"captionTargetPageId="40"captionText="Fig. 11. Martialis heureka Rabeling & Verhaagh, 2008, male, photomicrographs. A. Head, frontal view. B. Head, anteroventral oblique view. C. Body, lateral view. D. Body, dorsal view. Scale bars: A–B = 0.2 mm, C–D = 0.5 mm."figureDoi="http://doi.org/10.5281/zenodo.3780170"httpUri="https://zenodo.org/record/3780170/files/figure.png"pageId="36"pageNumber="37">Fig. 11</figureCitation>
A–B). In full-face view head about as broad as long excluding eyes, broader than long including eyes. Palpal formula 2,1; palps short, not reaching hypostomal margin. Stipes simple, lacking carinae on medial surface. Labrum very small, medially emarginate, setose; lateral margins distant from mandibular bases by somewhat less than maximum lateromedial labrum length; labrum lacking basolateral trigger setae observable in workers. Mandibles linear, narrow; lateral and medial margins weakly tapering to apex; masticatory mandibular margin reduced, bidentate; apical tooth asymmetrical, larger than symmetrical basal tooth; mandalus enlarged, diameter equal to maximum mandible width. Clypeus reduced; anterior margin broadly emarginate; medial clypeal portion maximum anteroposterior length about 1.5 maximum antennal socket diameters; posterior clypeal margin produced between antennal toruli. Supraclypeal area arc-shaped, anteroposteriorly longer than maximum antennal socket diameter. Antennal toruli situated anteriorly, with anteriormost portion of torular arch anterad anterior tentorial pit. Frons and ocellar area bulging. Occipital carina present, weakly developed, obscured in full-face view by vertex, not enclosing occiput. Compound eyes bulging strongly; medial margin weakly convex; posterior margin weakly emarginate; compound eye narrower dorsally than ventrally. Ocelli small, situated distant from compound eye. Hypostomal margin reduced, lacking lamina. Antenna 13-merous; scape longer than maximum compound eye diameter and slightly more than 2 x pedicel length; pedicel cylindrical, long, about
<figureCitationid="131C2A1FFF8DB140FEDDFEDAFE2D1E7F"box="[365,441,305,331]"captionStart="Fig"captionStartId="40.[227,270,1966,1992]"captionTargetBox="[270,1395,304,1931]"captionTargetId="figure@40.[270,1395,302,1931]"captionTargetPageId="40"captionText="Fig. 11. Martialis heureka Rabeling & Verhaagh, 2008, male, photomicrographs. A. Head, frontal view. B. Head, anteroventral oblique view. C. Body, lateral view. D. Body, dorsal view. Scale bars: A–B = 0.2 mm, C–D = 0.5 mm."figureDoi="http://doi.org/10.5281/zenodo.3780170"httpUri="https://zenodo.org/record/3780170/files/figure.png"pageId="37"pageNumber="38">Fig. 11</figureCitation>
C–D). Pronotal neck continuous with remainder of sclerite in dorsal view;main portion of pronotum swollen, muscular; anteromedian pronotal face convex in profile view, short, dorsoventral height of pronotum from pronotal neck about
x mesoscutum height in profile view; lateral pronotal face concave. Mesoscutum broader than long in dorsal view (length 0.80–0.83 x width); anterior and posterolateral areas swollen. Notauli distinct, crossribbed, meeting at body midline, not extending to transscutal line although narrow longitudinal line present from notauli to transscutal line. Parapsidal lines impressed, slightly divergent. Parascutal carinae nearly linear; weakly sinuate. Scutoscutellar sulcus unimpressed. Axillae small and widely situated. Mesoscutellum high and convex in profile view; not modified. Metascutellum small, lateromedial width slightly less than one half anteroposterior length; in profile view metascutellum strongly produced. Metanotal trough deep, small, circular. Mesopectus with oblique longitudinal sulcus, anterior terminus of sulcus nearly contacting posterolateral pronotal corner. Spiracular sclerite inconspicuous. Lower metapleural area strongly offset from upper metapleural area by deep, broad, margined sulcus. Metapleural gland orifice occluded; presence of internal metapleural gland not visible through metapleural sclerite. Propodeum parabolic in profile view, dorsal face about as long as and continuous with posterior face; propodeal spiracle circular, small; propodeal lobe weakly developed, carinate, clearly visible in anterolateral oblique view.
<figureCitationid="131C2A1FFF8DB140FEC3FC69FE741CA8"box="[371,480,898,925]"captionStart="Fig"captionStartId="40.[227,270,1966,1992]"captionTargetBox="[270,1395,304,1931]"captionTargetId="figure@40.[270,1395,302,1931]"captionTargetPageId="40"captionText="Fig. 11. Martialis heureka Rabeling & Verhaagh, 2008, male, photomicrographs. A. Head, frontal view. B. Head, anteroventral oblique view. C. Body, lateral view. D. Body, dorsal view. Scale bars: A–B = 0.2 mm, C–D = 0.5 mm."figureDoi="http://doi.org/10.5281/zenodo.3780170"httpUri="https://zenodo.org/record/3780170/files/figure.png"pageId="37"pageNumber="38">Fig. 11C</figureCitation>
). Petiole nodiform, pedunculate; anteriormost portion of petiolar tergum offset by parabolic carina; petiolar tergum and sternum fused, longitudinal lateral carinae not suggestive of suture; petiolar node shallow, in profile view anterodorsal face nearly linear, dorsum weakly convex, posterior face very weak; petiolar sternum linear for most of length, posteriorly narrowed, ventral petiolar surface with paired diverging carinulae; subpetiolar process absent. Abdominal segment III slightly reduced and differentiated from segment IV; helcium axial, sternal presclerite visible in profile view, not obscured by tergal presclerite; abdominal posttergite and poststernite III not fused; abdominal sternum III prora present as anterolateral bosses subtending helcium, anteromedian area of sternum concave. Abdominal terga IV–VIII and abdominal sterna IV–IX normally developed, not reduced or obscured
<figureCitationid="131C2A1FFF8DB140FED7FAC8FE411A0A"box="[359,469,1315,1342]"captionStart="Fig"captionStartId="41.[227,270,1896,1922]"captionTargetBox="[234,1428,302,1856]"captionTargetId="figure@41.[234,1428,302,1882]"captionTargetPageId="41"captionText="Fig. 12. Martialis heureka Rabeling & Verhaagh, 2008, male, wing photomicrographs and genitalia illustrations, genital membranes not shown. A. Forewing. B. Hindwing. C. Abdominal sternum IX, ventral view. D. Genital capsule, dorsal view. E. Genital capsule, ventral view. F. Genital capsule, lateral view. G. Volsella and paramere, mesal view. H. Penisvalva in situ, mesal view. Scale bars: A–B = 0.5 mm, C–H = 0.1 mm. Abbreviations: see Material and Methods."figureDoi="http://doi.org/10.5281/zenodo.3780172"httpUri="https://zenodo.org/record/3780172/files/figure.png"pageId="37"pageNumber="38">Fig. 12A</figureCitation>
). Tegulum reduced, subrectangular, longer than broad. Wings weakly infuscated, completely covered in fine setose layer. Pterostigma poorly-developed, only anterior enclosing abscissa tubular. Wing venation Ogata
x length of and meeting Mf1 obliquely. Rs+M continuous with undifferentiated Mf2-3 until meeting very short 2rsm. Rsf2+3 absent. 2r-rs very long, longer than combined lengths of Rsf1 and Mf1; 2r-rs directed posteroapically, not orthogonal with anterior wing margin. Rsf4–6 tubular to Rf, enclosing marginal cell. Mf4–6 absent. Crossvein cu-a incompletely tubular, situated basad Mf1. Cuf divergent with respect to Rs+M+Mf2+3. 1A extending only slightly beyond cu-a, not enclosing subdiscal cell 1.
<figureCitationid="131C2A1FFF8DB140FED7F995FE4619AD"box="[359,466,1662,1689]"captionStart="Fig"captionStartId="41.[227,270,1896,1922]"captionTargetBox="[234,1428,302,1856]"captionTargetId="figure@41.[234,1428,302,1882]"captionTargetPageId="41"captionText="Fig. 12. Martialis heureka Rabeling & Verhaagh, 2008, male, wing photomicrographs and genitalia illustrations, genital membranes not shown. A. Forewing. B. Hindwing. C. Abdominal sternum IX, ventral view. D. Genital capsule, dorsal view. E. Genital capsule, ventral view. F. Genital capsule, lateral view. G. Volsella and paramere, mesal view. H. Penisvalva in situ, mesal view. Scale bars: A–B = 0.5 mm, C–H = 0.1 mm. Abbreviations: see Material and Methods."figureDoi="http://doi.org/10.5281/zenodo.3780172"httpUri="https://zenodo.org/record/3780172/files/figure.png"pageId="37"pageNumber="38">Fig. 12B</figureCitation>
). Hindwing venation reduced, only R+Rs and 1A tubular; R not reaching anterior wing margin; 1A short, weakly indicated. Three hamuli present. Claval region poorly developed.
<figureCitationid="131C2A1FFF8DB140FED4F908FE2019C9"box="[356,436,1763,1790]"captionStart="Fig"captionStartId="41.[227,270,1896,1922]"captionTargetBox="[234,1428,302,1856]"captionTargetId="figure@41.[234,1428,302,1882]"captionTargetPageId="41"captionText="Fig. 12. Martialis heureka Rabeling & Verhaagh, 2008, male, wing photomicrographs and genitalia illustrations, genital membranes not shown. A. Forewing. B. Hindwing. C. Abdominal sternum IX, ventral view. D. Genital capsule, dorsal view. E. Genital capsule, ventral view. F. Genital capsule, lateral view. G. Volsella and paramere, mesal view. H. Penisvalva in situ, mesal view. Scale bars: A–B = 0.5 mm, C–H = 0.1 mm. Abbreviations: see Material and Methods."figureDoi="http://doi.org/10.5281/zenodo.3780172"httpUri="https://zenodo.org/record/3780172/files/figure.png"pageId="37"pageNumber="38">Fig. 12</figureCitation>
C–H). Pygostyles absent.Abdominal sternum IX spiculum short; anterior margin linear, curving posterolaterally near lateral margins; lateral margins short, slightly divergent; posterolateral margins weakly concave, tapering strongly to acute, narrowly rounded apex. Cupula dorsal and lateral faces about as broad as telomeral base; lateral face narrowing ventrally to narrow bar-like ventral face. Basimere and telomere more-or-less continuous, basimere weakly shouldered dorsomedially anterad telomeral base; dorsomedian margins of basimeres parallel for about half length of paramere; telomere acutely triangular in profile view; basimere and telomere with ventrolateral layer of posteroventrallydirected setae. Basivolsella lateromedially broad; base transversely connected with basimere; cuspis
valviceps height; valviceps linear, dorsoventrally short; valviceps dorsomedially fused for most of length; in profile view dorsal valviceps margin weakly convex, ventral margin concave, edentate, valviceps apex weakly convex, subrectangular, produced ventrally; dorsolateral valviceps face convex, margined by lateral apodeme which extends almost to apex before curving ventrad and contacting ventral margin; ventrolateral valviceps face concave; phallotreme situated at aedeagal apex, sclerotic aperture formed by valviceps circular.
</paragraph>
<paragraphid="8B98369AFF8EB143FF53FDADFAE11D54"blockId="38.[227,1397,582,608]"box="[227,1397,582,608]"pageId="38"pageNumber="39">COLORATION. Body almost uniformly brown to brownish yellow; extremities slightly lighter colored.</paragraph>
<paragraphid="8B98369AFF8EB143FF53FD6CFB5A1CA3"blockId="38.[227,1437,647,920]"pageId="38"pageNumber="39">SCULPTURATION. Body weakly sculptured overall; head with fine piligerous punctae; mesonotal piligerous punctae coarser, posterolateral mesoscutal area above parascutal carina roughened; dorsomedian scutoscutellar area finely and densely anteroposteriorly striate; mesoscutellum weakly roughened; metascutellum with fine transverse carina subtending posteriorly produced portion of disc; mesopectus and metapleuron smooth, shining, slightly rough; propodeum finely striate, striae extending from anterior margin down along lateral faces, dorsal propodeal face weakly rugose, posterior face mostly smooth; petiole mostly smooth and shining, with lateral longitudinal carinulae; abdominal segment III mostly smooth and shining; abdominal segments posterior to segment III weakly sclerotized.</paragraph>
<paragraphid="8B98369AFF8EB143FF53FC54FCE51BE4"blockId="38.[227,1437,959,1232]"pageId="38"pageNumber="39">SETATION. Head, median pronotal portion including pronotal neck, mesonotum, and procoxae covered by dense layer of somewhat short, uniform, weakly curved, erect to suberect setae, longer setae present on these areas very sparsely; clypeus lacking clypeal brush of worker, although setal layer denser than on remainder of head capsule; setae sparse and subdecumbent to nearly appressed on pronotal lateral face, mesopectus, metapleuron, and propodeum; setae somewhat denser on metasoma, but not as dense as on head and mesonotal dorsum; petiolar setae elongate, linear, setae on remaining segments shorter and curved; setae on legs, including meso- and metacoxae, about as dense as on metasoma, mostly subdecumbent with a few longer suberect setae present.</paragraph>
<taxonomicNameid="4C274D19FF8EB143FB7AFA7EFDB71AE7"authority="Rabeling & Verhaagh, 2008"authorityName="Rabeling & Verhaagh"authorityYear="2008"class="Insecta"family="Formicidae"genus="Martialis"higherTaxonomySource="GBIF"kingdom="Animalia"order="Hymenoptera"pageId="38"pageNumber="39"phylum="Arthropoda"rank="species"species="heureka">
<bibRefCitationid="EFB64B6BFF8EB143FF53FA52FDB71AE7"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[227,547,1465,1491]"pageId="38"pageNumber="39"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">Rabeling & Verhaagh, 2008</bibRefCitation>
described in recent years. Displaying a bizarre mixture of pleisiomorphic and autapomorphic traits, the species was attributed to its own subfamily, the
. This decision was supported by further morphological study (
<bibRefCitationid="EFB64B6BFF8EB143FA87FA14FEF61908"author="Brandao C. R. F. & Diniz J. L. M. & Feitosa R. S. M."pageId="38"pageNumber="39"pagination="413 - 423"refId="ref30401"refString="Brandao C. R. F., Diniz J. L. M. & Feitosa R. S. M. 2010. The venom apparatus and other morphological characters of the ant Martialis heureka (Hymenoptera, Formicidae, Martialinae). Papeis Avulsos de Zoologia 50: 413 - 423."type="journal article"year="2010">
) and multi-locus molecular phylogenetic reconstruction (
<bibRefCitationid="EFB64B6BFF8EB143FBB6F9C9FB6F1908"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[1030,1275,1570,1596]"pageId="38"pageNumber="39"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">
<bibRefCitationid="EFB64B6BFF8EB143FF53F9AEFE3A1954"author="Kuck P. & Hita Garcia F. & Misof B. & Meusemann K."box="[227,430,1605,1632]"pageId="38"pageNumber="39"pagination="21031"refId="ref31727"refString="Kuck P., Hita Garcia F., Misof B. & Meusemann K. 2011. Improved phylogenetic analyses corroborate a plausible position of Martialis heureka in the ant tree of life. PLoS ONE 6: e 21031. http: // dx. doi. org / 10.1371 / journal. pone. 0021031"type="journal article"year="2011">
<bibRefCitationid="EFB64B6BFF8EB143FE73F9AEFD5C1954"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[451,712,1605,1632]"pageId="38"pageNumber="39"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">
<bibRefCitationid="EFB64B6BFF8EB143FBF3F982FAB419B7"author="Kuck P. & Hita Garcia F. & Misof B. & Meusemann K."box="[1091,1312,1641,1667]"pageId="38"pageNumber="39"pagination="21031"refId="ref31727"refString="Kuck P., Hita Garcia F., Misof B. & Meusemann K. 2011. Improved phylogenetic analyses corroborate a plausible position of Martialis heureka in the ant tree of life. PLoS ONE 6: e 21031. http: // dx. doi. org / 10.1371 / journal. pone. 0021031"type="journal article"year="2011">
is described for the first time based on material from the Biological Dynamics of Forest Fragments Project (BDFFP [English], PDBFF [Portuguese]) study region, about
differs from the worker by standard intercaste dimorphism, e.g., eyes welldeveloped, ocelli and notauli present, alate, flight sclerites developed, mesosoma musculated for flight, but the male also differs notably in several specific characters: mandibles far shorter, reduced relative to worker; labral trigger setae absent; clypeal brush absent; antennal toruli situated more anteriorly; forelegs weak; metatibial gland absent; petiolar node weakly developed; helcium broader; abdominal tergum and sternum III of equal length; cinctus between pre- and postsclerites IV not developed. On the other hand, the male of
displays numerous similarities with the worker, such as linear mandibles, reduced palpal count (although worker count unconfirmed, but certainly less than 3,3), clypeus poorly developed, anterior tentorial pits posteriorly-situated, antennal toruli well-developed (but not quite as cup-like as in worker), scapes long, pedicel elongate, frontal carinae absent, propodeal lobes weakly developed (contra the initial worker diagnosis), tibial spur formula 1,1, basipetiolar carina present, petiolar tergum and sternum fused, abdominal segment III differentiated from IV, and sculpture and setation remarkably similar, although the somatic sclerites of the male are generally less strongly sclerotized. The two castes are similar in several other specifics, but this brief list captures most of the significant features.
Specimens were examined from collecting events in January, February, April, and October of 1985. The BDFFP study region displays weak seasonality of rainfall and day length, with the months of June through December roughly representing the “dry season”, July through September being the driest (Bierregaard Jr.
flights occur year-round, although the sampling of BDFFP material examined for this study is too small to confidently assert the flight phenology. While the range of
may be recovered via alates more readily than workers. The use of Malaise traps and flight intercept traps should be encouraged for studies of ant diversity, particularly for species with cryptic habits. Although our knowledge of flight phenology is poor, tropical rainforests may be particularly amenable to these studies due to the relatively more year-round flights of Neotropical (
<bibRefCitationid="EFB64B6BFF8FB142FB49FC07FEB91B1D"author="Kaspari M. & Pickering J. & Longino J. T. & Windsor D."pageId="39"pageNumber="40"pagination="382 - 390"refId="ref31567"refString="Kaspari M., Pickering J., Longino J. T. & Windsor D. 2001 a. The phenology of a Neotropical ant assemblage: Evidence for continuous and overlapping production. Behavioral Ecology and Sociobiology 50: 382 - 390. http: // dx. doi. org / 10.1007 / s 002650100378"type="journal article"year="2001">
<bibRefCitationid="EFB64B6BFF8FB142FE89FBE4FE101B1E"author="Kaspari M. & Pickering J. & Windsor D."box="[313,388,1039,1066]"pageId="39"pageNumber="40"pagination="245 - 257"refId="ref31625"refString="Kaspari M., Pickering J. & Windsor D. 2001 b. The reproductive flight phenology of a Neotropical ant assemblage. Ecological Entomology 26: 245 - 257. http: // dx. doi. org / 10.1046 / j. 1365 - 2311.2001.00320. x"type="journal article"year="2001">2001b</bibRefCitation>
) than of Nearctic ants (
<bibRefCitationid="EFB64B6BFF8FB142FD28FBFBFCF71B1D"author="Dunn R. R. & Parker C. R. & Geraghty M. & Sanders N. J."box="[664,867,1039,1066]"pageId="39"pageNumber="40"pagination="135 - 142"refId="ref31054"refString="Dunn R. R., Parker C. R., Geraghty M. & Sanders N. J. 2007. Reproductive phenologies in a diverse temperate ant fauna. Ecological Entomology 32: 135 - 142. http: // dx. doi. org / 10.1111 / j. 1365 - 2311.2006.00839. x"type="journal article"year="2007">
in the BDFFP plots have survived the intervening 30 years. This may not be the case, as ant communities have been observed to hemorrhage in BDFFP study plots (
<bibRefCitationid="EFB64B6BFF8FB142FC3AFB36FB371BCC"author="Vasconcelos H. L. & Carvalho K. S. & Delabie J. H. C."box="[906,1187,1245,1272]"pageId="39"pageNumber="40"pagination="199 - 207"refId="ref32761"refString="Vasconcelos H. L., Carvalho K. S. & Delabie J. H. C. 2001. Landscape modifications and ant communities. In: Bierregaard Jr., Gascon C., Lovejoy T. E. & Mesquita R. C. G. (eds) Lessons from Amazonia: The Ecology and Conservation of a Fragmented Forest: 199 - 207. Yale University Press, New Haven."type="book chapter"year="2001">
), although hypogaeic ants may be less sensitive to habitat changes than epigaeic ants. The placement of the Malaise samples relative to the edges of the study plots is unknown. As well, the gyne of
remains unknown. It is possible that this caste will be ergatoid, but the presence of alate gynes cannot be ruled out. In general, the male of a given ant species is more frequently collected via Malaise traps than females; thus it is possible that although only males were encountered, alate gynes could still be present. Regardless, the natural history of
<collectionCodeid="ED36AE5FFF8FB142FDE6F9CAFD37190F"box="[598,675,1569,1595]"collectionName="Brazil, Sao Paulo, Sao Paulo, Museu de Zoologia da Universidade de Sao Paulo"country="Brazil"httpUri="http://grbio.org/cool/9yp6-zxp9"name="Sao Paulo, Museu de Zoologia da Universidade de Sao Paulo"pageId="39"pageNumber="40"type="Museum">MZSP</collectionCode>
. Specimen was cleared by non-destructive extraction of DNA, which allowed for examination of internal characters.
<locationid="8EF86041FF8FB142FCC6F9AFFC0F196A"LSID="urn:lsid:plazi:treatment:038E878CFF8CB148FD9CFCF6FBCF1986:8EF86041FF8FB142FCC6F9AFFC0F196A"box="[886,923,1604,1630]"country="Brazil"county="Plot"latitude="-2.38897"longLatPrecision="1"longitude="-59.85012"municipality="Biological Dynamics of Forest Fragments Project"name="All"pageId="39"pageNumber="40"stateProvince="Amazonas">All</location>
males were examined from the following collecting events at the
<locationid="8EF86041FF8FB142FDB7F98CFD2919B5"LSID="urn:lsid:plazi:treatment:038E878CFF8CB148FD9CFCF6FBCF1986:8EF86041FF8FB142FDB7F98CFD2919B5"box="[519,701,1639,1665]"country="Brazil"county="Plot"latitude="-2.38897"longLatPrecision="1"longitude="-59.85012"municipality="Biological Dynamics of Forest Fragments Project"name="Fazenda Esteio"pageId="39"pageNumber="40"stateProvince="Amazonas">Fazenda Esteio</location>
study area of the
<collectingMunicipalityid="6BFCACE0FF8FB142FC11F98CFEA21990"pageId="39"pageNumber="40">Biological Dynamics of Forest Fragments Project</collectingMunicipality>
<materialsCitationid="3B4F3CC7FF8FB142FF53F93BFCF0183A"ID-GBIF-Occurrence="2609658203"collectingDate="1922-10"collectingDateMax="1926-02"collectingDateMin="1922-10"collectingMethod="via Malaise trap"collectionCode="MZSP, PDBFF, INPA"collectorName="Bert Klein & Amazonian & Note & Bierregaard Jr. & Google Earth"country="Brazil"county="Plot"elevation="250"latitude="-2.38897"location="All"longLatPrecision="1"longitude="-59.85012"municipality="Biological Dynamics of Forest Fragments Project"pageId="39"pageNumber="40"specimenCode="INPA-SI 2014"specimenCount="6"stateProvince="Amazonas"typeStatus="holotype">
<materialsCitationid="3B4F3CC7FF8FB142FCDDF91FFC97188A"ID-GBIF-Occurrence="2609658202"collectingDate="1923-01"collectingDateMax="1924-04"collectingDateMin="1923-01"collectingMethod="via Malaise trap"collectionCode="PDBFF, INPA"collectorName="Amazonian & Note & Bierregaard Jr. & Google Earth"country="Brazil"county="Plot"elevation="500"latitude="-2.38897"location="All"longLatPrecision="1"longitude="-59.85012"municipality="Biological Dynamics of Forest Fragments Project"pageId="39"pageNumber="40"specimenCode="INPA-SI 2014"specimenCount="19"stateProvince="Amazonas"typeStatus="holotype">
with error estimates to account for uncertainty of exact plot location; half the male material examined remains at
<collectionCodeid="ED36AE5FFF8FB142FD00F84FFD63188A"box="[688,759,1956,1982]"collectionName="Brazil, Amazonas, Manaus, Instituto Nacional de Pesquisas da Amazoonia, Colecao Sistematica da Entomologia"country="Brazil"httpUri="http://grbio.org/cool/rm88-499z"name="Instituto Nacional de Pesquisas da Amazonia"pageId="39"pageNumber="40">INPA</collectionCode>
<taxonomicNameid="4C274D19FF80B14DFEF3F844FCF018FD"authority="Rabeling & Verhaagh, 2008"authorityName="Rabeling & Verhaagh"authorityYear="2008"box="[323,868,1966,1993]"class="Insecta"family="Formicidae"genus="Martialis"higherTaxonomySource="GBIF"kingdom="Animalia"order="Hymenoptera"pageId="40"pageNumber="41"phylum="Arthropoda"rank="species"species="heureka">
<bibRefCitationid="EFB64B6BFF80B14DFDACF845FCF018FD"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[540,868,1966,1993]"pageId="40"pageNumber="41"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">Rabeling & Verhaagh, 2008</bibRefCitation>
<taxonomicNameid="4C274D19FF81B14CFEFDF883FC1618B6"authority="Rabeling & Verhaagh, 2008"authorityName="Rabeling & Verhaagh"authorityYear="2008"box="[333,898,1896,1922]"class="Insecta"family="Formicidae"genus="Martialis"higherTaxonomySource="GBIF"kingdom="Animalia"order="Hymenoptera"pageId="41"pageNumber="42"phylum="Arthropoda"rank="species"species="heureka">
<bibRefCitationid="EFB64B6BFF81B14CFD9DF883FC1618B6"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[557,898,1896,1922]"pageId="41"pageNumber="42"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">Rabeling & Verhaagh, 2008</bibRefCitation>
</taxonomicName>
, male, wing photomicrographs and genitalia illustrations, genital membranes not shown.
<pageTitleid="CBB8EEFDFF81B14CFF53F81DFC5E173B"box="[227,970,2037,2064]"pageId="41"pageNumber="42">mm, C–H = 0.1 mm. Abbreviations: see Material and Methods.</pageTitle>
<emphasisid="B953EA88FF82B14FFF53FEDBFD401E79"bold="true"box="[227,724,304,333]"italics="true"pageId="42"pageNumber="43">Shared apomorphies of the basal ants</emphasis>
Of particular interest for the “basal ant” problem (
<bibRefCitationid="EFB64B6BFF82B14FFCA6FE8EFC4A1E4B"author="Brady S. G. & Schultz T. R. & Fisher B. L. & Ward P. S."box="[790,990,357,383]"pageId="42"pageNumber="43"pagination="18172 - 18177"refId="ref30264"refString="Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103"type="journal article"year="2006">
<bibRefCitationid="EFB64B6BFF82B14FFC57FE8EFB441E4B"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[999,1232,357,383]"pageId="42"pageNumber="43"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">
<bibRefCitationid="EFB64B6BFF82B14FFB69FE8EFA001E4B"author="Kuck P. & Hita Garcia F. & Misof B. & Meusemann K."box="[1241,1428,357,383]"pageId="42"pageNumber="43"pagination="21031"refId="ref31727"refString="Kuck P., Hita Garcia F., Misof B. & Meusemann K. 2011. Improved phylogenetic analyses corroborate a plausible position of Martialis heureka in the ant tree of life. PLoS ONE 6: e 21031. http: // dx. doi. org / 10.1371 / journal. pone. 0021031"type="journal article"year="2011">
<taxonomicNameid="4C274D19FF82B14FFEF9FE25FE4E1EDD"authorityName="Lepeletier de Saint-Fargeau"authorityYear="1835"box="[329,474,462,489]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="60"phylum="Arthropoda"rank="subFamily"subFamily="Myrmicinae">Myrmicinae</taxonomicName>
) have crystallized in the past decade (
<bibRefCitationid="EFB64B6BFF82B14FFC23FE25FBE21EDC"author="Moreau C. S. & Bell C. D. & Vila R. & Archibald S. B. & Pierce N. E."box="[915,1142,462,488]"pageId="42"pageNumber="43"pagination="101 - 104"refId="ref32067"refString="Moreau C. S., Bell C. D., Vila R., Archibald S. B. & Pierce N. E. 2006. Phylogeny of the ants: diversification in the age of angiosperms. Science 312: 101 - 104. http: // dx. doi. org / 10.1126 / science. 1124891"type="journal article"year="2006">
<bibRefCitationid="EFB64B6BFF82B14FFB31FE25FADB1EDC"author="Brady S. G. & Schultz T. R. & Fisher B. L. & Ward P. S."box="[1153,1359,462,488]"pageId="42"pageNumber="43"pagination="18172 - 18177"refId="ref30264"refString="Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103"type="journal article"year="2006">
<bibRefCitationid="EFB64B6BFF82B14FFAE9FE25FA031EDC"author="Brady S. G. & Schultz T. R. & Fisher B. L. & Ward P. S."box="[1369,1431,462,488]"pageId="42"pageNumber="43"pagination="93"refId="ref30334"refString="Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2014. The rise of army ants and their relatives: diversification of specialized predatory doryline ants. BMC Evolutionary Biology 14: 93. http: // dx. doi. org / 10.1186 / 1471 - 2148 - 14 - 93."type="journal article"year="2014">2014</bibRefCitation>
;
<bibRefCitationid="EFB64B6BFF82B14FFF53FE1AFE3E1D38"author="Ward P. S. & Brady S. G. & Fisher B. L. & Schultz T. R."box="[227,426,497,524]"pageId="42"pageNumber="43"pagination="61 - 81"refId="ref33114"refString="Ward P. S., Brady S. G., Fisher B. L. & Schultz T. R. 2015. The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40: 61 - 81. http: // dx. doi. org / 10.1111 / syen. 12090"type="journal article"year="2015">
<taxonomicNameid="4C274D19FF82B14FFEF7FDFEFE2B1D1B"authorityName="Lepeletier de Saint-Fargeau"authorityYear="1835"box="[327,447,533,559]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="subFamily"subFamily="Ponerinae">Ponerinae</taxonomicName>
<bibRefCitationid="EFB64B6BFF82B14FFC24FDFEFB881D1B"author="Ward P. S."box="[916,1052,533,559]"pageId="42"pageNumber="43"refId="ref32956"refString="Ward P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 2.1 - 2.21. http: // dx. doi. org / 10.1146 / annurev-ecolsys- 120213 - 091824"type="journal volume"year="2014">Ward 2014</bibRefCitation>
). The poneroids may constitute a clade or a grade, depending on placement of the root on the ant tree of life (
<bibRefCitationid="EFB64B6BFF82B14FFBE8FDD3FABF1D67"author="Brady S. G. & Schultz T. R. & Fisher B. L. & Ward P. S."box="[1112,1323,568,595]"pageId="42"pageNumber="43"pagination="18172 - 18177"refId="ref30264"refString="Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103"type="journal article"year="2006">
, which has been recovered as sister to the remainder of the extant
<taxonomicNameid="4C274D19FF82B14FFC11FD76FAAC1D83"authority="(Rabeling et al. 2008)"baseAuthorityName="Rabeling"baseAuthorityYear="2008"box="[929,1336,669,695]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="family">
Formicidae (
<bibRefCitationid="EFB64B6BFF82B14FFB87FD76FAA41D83"author="Rabeling C. & Brown J. M. & Verhaagh M."box="[1079,1328,669,695]"pageId="42"pageNumber="43"pagination="14913 - 14917"refId="ref32411"refString="Rabeling C., Brown J. M. & Verhaagh M. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences B 105: 14913 - 14917. http: // dx. doi. org / 10.1073 / pnas. 0806187105"type="journal article"year="2008">
<bibRefCitationid="EFB64B6BFF82B14FFECFFD2BFDCF1DEE"author="Kuck P. & Hita Garcia F. & Misof B. & Meusemann K."box="[383,603,704,730]"pageId="42"pageNumber="43"pagination="21031"refId="ref31727"refString="Kuck P., Hita Garcia F., Misof B. & Meusemann K. 2011. Improved phylogenetic analyses corroborate a plausible position of Martialis heureka in the ant tree of life. PLoS ONE 6: e 21031. http: // dx. doi. org / 10.1371 / journal. pone. 0021031"type="journal article"year="2011">
. These results not only disagree in topology, but may be subject to the biases of long-branch attraction and CG-bias (
<bibRefCitationid="EFB64B6BFF82B14FFEE5FCEDFE4B1C14"author="Ward P. S."box="[341,479,774,800]"pageId="42"pageNumber="43"refId="ref32956"refString="Ward P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 2.1 - 2.21. http: // dx. doi. org / 10.1146 / annurev-ecolsys- 120213 - 091824"type="journal volume"year="2014">Ward 2014</bibRefCitation>
). As summarized by
<bibRefCitationid="EFB64B6BFF82B14FFD6AFCEDFCE61C14"author="Ward P. S."box="[730,882,774,800]"pageId="42"pageNumber="43"refId="ref32956"refString="Ward P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 2.1 - 2.21. http: // dx. doi. org / 10.1146 / annurev-ecolsys- 120213 - 091824"type="journal volume"year="2014">Ward (2014)</bibRefCitation>
, one of the major questions of ant systematics is whether
or whether they are highly derived poneroids. Unpublished molecular phylogenetic analyses by P.S. Ward (discussed in
<bibRefCitationid="EFB64B6BFF82B14FFF53FC9BFEFF1CBE"author="Ward P. S."box="[227,363,880,906]"pageId="42"pageNumber="43"refId="ref32956"refString="Ward P. S. 2014. The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics 45: 2.1 - 2.21. http: // dx. doi. org / 10.1146 / annurev-ecolsys- 120213 - 091824"type="journal volume"year="2014">Ward 2014</bibRefCitation>
. Considerable uncertainty thus exists, even with molecular data. The present work seeks, in part, to render this problem more tractable by providing novel morphological characters that are shared by
<taxonomicNameid="4C274D19FF82B14FFDBAFC16FD1F1B23"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[522,651,1021,1047]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFF53FB8AFECF1B4F"baseAuthorityName="Holldobler & Wilson"baseAuthorityYear="1990"box="[227,347,1121,1147]"class="Insecta"family="Formicidae"genus="Protanilla"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
by the following characters: mandibles linear, meeting at head midline, bidentate apically; antennal toruli situated posterad anterior portion of antennal torulus; lower metapleuron dorsoventrally longer than anteroposteriorly broad; metanotal trough pit-like (vs. oblong); propodeal lobe present; first submarginal cell enclosed by tubular abscissa; marginal cell 1 closed; petiole pedunculate; genitalia more generalized. The worker of
<taxonomicNameid="4C274D19FF82B14FFA95FAFAFA091A1F"baseAuthorityName="Holldobler & Wilson"baseAuthorityYear="1990"box="[1317,1437,1297,1323]"class="Insecta"family="Formicidae"genus="Protanilla"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFBFFFA9DFB441AA4"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[1103,1232,1398,1424]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFAABFA56FA081AE3"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[1307,1436,1469,1495]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
by the following characters: mandibles linear (rather than curved and subfalcate); clypeus reduced; antennal toruli strongly developed, conspicuous (vs. inconspicuous); metascutellar trough pit-like; petiolar tergum and sternum fused; helcium axial; and helcial sternite projecting ventrad helcial tergite in profile view.
<taxonomicNameid="4C274D19FF82B14FFDFCF960FD591991"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[588,717,1675,1701]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFBA3F945FB0019FC"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[1043,1172,1710,1736]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFB02F91EFAA7183B"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[1202,1331,1781,1807]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFC12F8D0FBB71861"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[930,1059,1851,1877]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFC47F8B5FBEC184C"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[1015,1144,1886,1912]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF82B14FFE0AF828FDAF18E9"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[442,571,1987,2013]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="42"pageNumber="43"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF83B14EFF53FEDAFE241E7F"authorityName="Wilson & Brown"authorityYear="1967"box="[227,432,305,331]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="43"pageNumber="44"phylum="Arthropoda"rank="subFamily"subFamily="Sphecomyrminae">Sphecomyrminae</taxonomicName>
males are superficially similar to males tentatively identified as
<taxonomicNameid="4C274D19FF83B14EFCCCFE93FABF1EA6"authority="(Grimaldi et al. 1997)"baseAuthorityName="Grimaldi"baseAuthorityYear="1997"box="[892,1323,375,402]"class="Insecta"family="Formicidae"genus="Sphecomyrma"kingdom="Animalia"order="Hymenoptera"pageId="43"pageNumber="44"phylum="Arthropoda"rank="genus">
<bibRefCitationid="EFB64B6BFF83B14EFB81FE9CFAB61EA6"author="Grimaldi D. & Agosti D. & Carpenter J. M."box="[1073,1314,375,402]"pageId="43"pageNumber="44"pagination="1 - 43"refId="ref31309"refString="Grimaldi D., Agosti D. & Carpenter J. M. 1997. New and rediscovered primitive ants (Hymenoptera: Formicidae) in Cretaceous amber from New Jersey, and their phylogenetic relationships. American Museum Novitates 3208: 1 - 43."type="journal article"year="1997">
, differing mainly in having antennae situated anteriorly, reduced venation, tergosternal petiolar fusion, a tibial spur formula of 1,1, and having abdominal segment III reduced. The extinct genus
differs similarly, except the mandibles are curved and palps are longer; the petiole is not visible in the specimen illustrated by
<bibRefCitationid="EFB64B6BFF83B14EFEB7FDEFFD981D2A"author="Grimaldi D. & Agosti D. & Carpenter J. M."box="[263,524,516,542]"pageId="43"pageNumber="44"pagination="1 - 43"refId="ref31309"refString="Grimaldi D., Agosti D. & Carpenter J. M. 1997. New and rediscovered primitive ants (Hymenoptera: Formicidae) in Cretaceous amber from New Jersey, and their phylogenetic relationships. American Museum Novitates 3208: 1 - 43."type="journal article"year="1997">
<taxonomicNameid="4C274D19FF83B14EFEDAFD82FE7F1DB7"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[362,491,617,643]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="43"pageNumber="44"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF83B14EFCD1FD82FC761DB7"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[865,994,617,643]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="43"pageNumber="44"phylum="Arthropoda"rank="genus">
, with groups presented by increasing qualitative similarity. Pleisiomorphic conditions are presented in brackets next to the respective apomorphic conditions.
<paragraphid="8B98369AFF83B14EFF53FD1AFD521C3F"blockId="43.[227,1437,753,885]"box="[227,710,753,779]"pageId="43"pageNumber="44">Apomorphies shared among all five taxa:</paragraph>
<taxonomicNameid="4C274D19FF83B14EFEA0FC08FE051CC9"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[272,401,995,1021]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="43"pageNumber="44"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF83B14EFEA0FBC2FE051B77"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[272,401,1065,1091]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="43"pageNumber="44"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF83B14EFCA1FB80FC061BB1"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[785,914,1131,1157]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="43"pageNumber="44"phylum="Arthropoda"rank="genus">
2. Antennal toruli directed dorsally or anterodorsally rather than laterally (female castes) (note 1). [Antennal toruli directed more-or-less laterally in
6. Parascutal carina situated very low on mesoscutum, almost completely obscured by wing base and thus inconspicuous (male). [Parascutal carina raised on mesoscutum, near height of mesoscutal dorsum, only partially obscured by wing base, conspicuous; in
<paragraphid="8B98369AFF83B14EFF53F92AFE011815"blockId="43.[227,1437,1130,2036]"pageId="43"pageNumber="44">9. Valviceps linear to arched, with concave ventral and convex dorsal margin in ectal view (male). [Valviceps more-or-less elliptical, with both dorsal and ventral margins convex in ectal view in other poneroids.]</paragraph>
<paragraphid="8B98369AFF83B14EFF53F8C1FDE3189A"blockId="43.[227,1437,1130,2036]"pageId="43"pageNumber="44">10. Valviceps lateral apodeme evenly linear, extending almost to valviceps apex before curving ventrad, delimiting convex dorsolateral and concave ventrolateral face (male). [Lateral apodeme sinuate, wavy, extending to valviceps apex but not curving ventrad; dorsolateral and ventrolateral faces more or less flat in other poneroids.]</paragraph>
<paragraphid="8B98369AFF83B14EFF53F85CFE0118C0"blockId="43.[227,1437,1130,2036]"pageId="43"pageNumber="44">11. Valviceps ventral margin edentate (male) (note 5). [Valviceps ventral margin dentate in other poneroids.]</paragraph>
<paragraphid="8B98369AFF84B149FF53FE93FC3B1EA6"blockId="44.[227,1436,305,402]"box="[227,943,375,402]"pageId="44"pageNumber="45">13. Characters of the sting apparatus (worker, gyne) (note 6).</paragraph>
<taxonomicNameid="4C274D19FF84B149FE21FDEBFD861D2E"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[401,530,512,538]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF84B149FEA0FDADFE0B1D54"authorityName="Lepeletier de Saint-Fargeau"authorityYear="1835"box="[272,415,582,608]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="60"phylum="Arthropoda"rank="subFamily"subFamily="Myrmicinae">Myrmicinae</taxonomicName>
and
<taxonomicNameid="4C274D19FF84B149FE68FDADFDD51D54"authorityName="Brown & Kempf"authorityYear="1968"box="[472,577,582,608]"class="Insecta"family="Formicidae"genus="Tatuidris"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF84B149FC47FD67FBFB1D92"authorityName="Lepeletier de Saint-Fargeau"authorityYear="1835"box="[1015,1135,652,678]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="subFamily"subFamily="Ponerinae">Ponerinae</taxonomicName>
<taxonomicNameid="4C274D19FF84B149FBA6FCF2FB381C07"baseAuthorityName="Yoshimura & Fisher"baseAuthorityYear="2012"box="[1046,1196,793,819]"class="Insecta"family="Formicidae"genus="Adetomyrma"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<bibRefCitationid="EFB64B6BFF84B149FD05FCB4FCCF1C4D"author="Bolton B."box="[693,859,863,889]"pageId="44"pageNumber="45"pagination="1339 - 1364"refId="ref29952"refString="Bolton B. 1990 c. Army ants reassessed: the phylogeny and classification of the doryline section (Hymenoptera, Formicidae). Journal of Natural History 24: 1339 - 1364. http: // dx. doi. org / 10.1080 / 00222939000770811"type="journal article"year="1990">Bolton 1990c</bibRefCitation>
)
</taxonomicName>
. The bizarre, elongate, filamentous structures of some unassociated leptanilline males are, amazingly, appendages of the basimere (
<figureCitationid="131C2A1FFF84B149FB68FC69FAD51CA9"box="[1240,1345,898,925]"captionStart="Fig"captionStartId="31.[227,270,1858,1884]"captionTargetBox="[228,1436,601,1807]"captionTargetId="figure@31.[228,1436,601,1818]"captionTargetPageId="31"captionText="Fig. 10. Representative males of Leptanillinae, lateral view A. Protanilla “TH01” (Thailand, CASENT0119776, A. Nobile), arrow indicates loss of abdominal segment II petiolation. B. Protanilla “TH03” (Thailand, CASENT0119791, E. Prado). C. Leptanilla swani (Australia, CASENT0172318, A. Nobile). D. Protanilla sp. (Indonesia, CASENT0178838, A. Nobile), arrow indicates basolateral basimeral process. E. Scyphodon sp. (Indonesia, MCZ155112w, A. Nobile). F. Noonilla sp., used with permission from Petersen (1968). Scale bars: A, C, E–F = 0.2 mm, D = 0.5 mm, B = 1.0 mm."figureDoi="http://doi.org/10.5281/zenodo.3780168"httpUri="https://zenodo.org/record/3780168/files/figure.png"pageId="44"pageNumber="45">Fig. 10D</figureCitation>
5. Absence of ventral valviceps teeth has evolved several times, including the amblyoponine genus
<taxonomicNameid="4C274D19FF84B149FEA0FC22FC971CD0"authority="(Yoshimura & Fisher 2012 b)"baseAuthorityName="Yoshimura & Fisher"baseAuthorityYear="2012"box="[272,771,969,996]"class="Insecta"family="Formicidae"genus="Adetomyrma"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<bibRefCitationid="EFB64B6BFF84B149FE05FC22FD6F1CD0"author="Yoshimura M. & Fisher B. L."box="[437,763,969,996]"pageId="44"pageNumber="45"pagination="1 - 31"refId="ref33711"refString="Yoshimura M. & Fisher B. L. 2012 b. A revision of the Malagasy endemic genus Adetomyrma (Hymenoptera: Formicidae: Amblyoponinae). Zootaxa 3341: 1 - 31."type="journal article"year="2012">Yoshimura & Fisher 2012b</bibRefCitation>
<bibRefCitationid="EFB64B6BFF84B149FEA0FC07FE211B32"author="Kugler C."box="[272,437,1004,1030]"pageId="44"pageNumber="45"pagination="103 - 115"refId="ref31785"refString="Kugler C. 1992. Stings of ants of the Leptanillinae (Hymenoptera: Formicidae). Psyche 99: 103 - 115."type="journal article"year="1992">Kugler (1992)</bibRefCitation>
indicated as synapomorphic a set of sting apparatus characters applying to
<taxonomicNameid="4C274D19FF84B149FAA4FC06FA011B33"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[1300,1429,1005,1031]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF84B149FEA0FBE4FE1C1B1D"baseAuthorityName="Holldobler & Wilson"baseAuthorityYear="1990"box="[272,392,1039,1065]"class="Insecta"family="Formicidae"genus="Protanilla"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
. These characters are, in short, lack of medial lobe on quadrate plate, fulcral arm short and lacking lateral extensions. Notably, the anal plate of
<taxonomicNameid="4C274D19FF84B149FDE5FBBDFD421B44"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[597,726,1110,1136]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF84B149FCA7FBBDFC1B1B44"baseAuthorityName="Holldobler & Wilson"baseAuthorityYear="1990"box="[791,911,1110,1136]"class="Insecta"family="Formicidae"genus="Protanilla"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<taxonomicNameid="4C274D19FF84B149FEA0FB92FD1F1BA7"authority="(Brandao et al. 2010)"baseAuthorityName="Brandao"baseAuthorityYear="2010"box="[272,651,1145,1171]"class="Insecta"family="Formicidae"genus="Martialis"higherTaxonomySource="GBIF"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
<bibRefCitationid="EFB64B6BFF84B149FE3FFB92FD171BA7"author="Brandao C. R. F. & Diniz J. L. M. & Feitosa R. S. M."box="[399,643,1145,1171]"pageId="44"pageNumber="45"pagination="413 - 423"refId="ref30401"refString="Brandao C. R. F., Diniz J. L. M. & Feitosa R. S. M. 2010. The venom apparatus and other morphological characters of the ant Martialis heureka (Hymenoptera, Formicidae, Martialinae). Papeis Avulsos de Zoologia 50: 413 - 423."type="journal article"year="2010">
<bibRefCitationid="EFB64B6BFF84B149FAF9FB77FEDB1BED"author="Kugler C."pageId="44"pageNumber="45"pagination="103 - 115"refId="ref31785"refString="Kugler C. 1992. Stings of ants of the Leptanillinae (Hymenoptera: Formicidae). Psyche 99: 103 - 115."type="journal article"year="1992">Kugler 1992</bibRefCitation>
)
</taxonomicName>
. The present author admittedly has insufficient expertise with the sting apparatus to critically evaluate these characters, however. Future researchers are encouraged to examine the sting apparati of the poneroids, as knowledge of this group is poor (
<bibRefCitationid="EFB64B6BFF84B149FC3DFAEDFBB01A14"author="Kugler C."box="[909,1060,1286,1312]"pageId="44"pageNumber="45"pagination="103 - 115"refId="ref31785"refString="Kugler C. 1992. Stings of ants of the Leptanillinae (Hymenoptera: Formicidae). Psyche 99: 103 - 115."type="journal article"year="1992">Kugler 1992</bibRefCitation>
) and as understanding the sting apparatus might assist in providing a more detailed picture of the evolution of the early branching ant lineages.
<taxonomicNameid="4C274D19FF84B149FD25FA65FC821A9C"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[661,790,1422,1448]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="44"pageNumber="45"phylum="Arthropoda"rank="genus">
1. Lateral bases of mandibles set deep in pits, suggestive of a trap-jaw mechanism (female castes). [Pleurostoma posterad mandibular insertion without a deep pit for reception of mandible; mandibles without trap-jaw mechanism in other poneroids, excepting
3. Entire clypeus anteriorly produced (not just median portion), with rectangular anterolateral margins (worker, gyne) (note 1). [Clypeus not produced anteriorly; anterolateral corner obtuse in other
<paragraphid="8B98369AFF84B149FF53F8FAFBFF187A"blockId="44.[227,1437,1422,2011]"pageId="44"pageNumber="45">5. Thickened setae present on ventromedial mandibular face (worker, gyne). [Thin setae present on ventromedial mandibular face in other poneroids, with some exceptions.]</paragraph>
<paragraphid="8B98369AFF84B149FF53F875FB5218EF"blockId="44.[227,1437,1422,2011]"pageId="44"pageNumber="45">7. Basivolsella lateromedially narrow in ventral view (male) (note 4). [Basivolsella lateromedially broad in ventral view in other poneroids, excepting various proceratiine species.]</paragraph>
9. Larva elongate, slender, and club-shaped (“leptanilloid” in terminology of
<bibRefCitationid="EFB64B6BFF85B148FB17FE9CFED81E82"author="Wheeler G. C. & Wheeler E. W."pageId="45"pageNumber="46"pagination="1 - 108"refId="ref33207"refString="Wheeler G. C. & Wheeler E. W. 1976. Ant larvae: Review and synthesis. Memoirs of the Entomological Society of Washington 7: 1 - 108."type="journal article"year="1976">Wheeler & Wheeler 1976</bibRefCitation>
) (note 5). [Larva pogonomyrmecoid, myrmecioid, or platythyreoid in other poneroids; see
<bibRefCitationid="EFB64B6BFF85B148FEA0FE55FDAA1EEC"author="Wheeler G. C. & Wheeler E. W."box="[272,574,446,472]"pageId="45"pageNumber="46"pagination="1 - 108"refId="ref33207"refString="Wheeler G. C. & Wheeler E. W. 1976. Ant larvae: Review and synthesis. Memoirs of the Entomological Society of Washington 7: 1 - 108."type="journal article"year="1976">Wheeler & Wheeler 1976</bibRefCitation>
<taxonomicNameid="4C274D19FF85B148FD0FFCB0FCD41C41"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[703,832,859,885]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="45"pageNumber="46"phylum="Arthropoda"rank="genus">
<paragraphid="8B98369AFF85B148FF53FC95FE041C8F"blockId="45.[227,1436,858,1131]"pageId="45"pageNumber="46">1. Promesonotal flexion extreme (worker, gyne?) (note 1). [Promesonotal articulation flexible, but not enhanced.]</paragraph>
<paragraphid="8B98369AFF85B148FF53FC2FFD581B36"blockId="45.[227,1436,858,1131]"pageId="45"pageNumber="46">2. Propleurae bulging in profile view (worker, gyne?) (note 2). [Propleurae flat, only weakly produced anteriorly beyond pronotum, if at all.]</paragraph>
3. The tergal-tergal fusion “collar” around the petiolar base is not present in the male of
<taxonomicNameid="4C274D19FF85B148FAA4FAF4FA011A0D"authorityName="Brown, Gotwald Jr. & Levieux"authorityYear="1971"box="[1300,1429,1311,1337]"class="Insecta"family="Formicidae"genus="Apomyrma"kingdom="Animalia"order="Hymenoptera"pageId="45"pageNumber="46"phylum="Arthropoda"rank="genus">
<emphasisid="B953EA88FF85B148FF53FA6EFC301A96"bold="true"box="[227,932,1413,1442]"italics="true"pageId="45"pageNumber="46">Brief global diagnoses of subfamilies, based on males</emphasis>
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;
<bibRefCitationid="EFB64B6BFF85B148FD13F9E0FCF01912"author="Ward P. S. & Brady S. G. & Fisher B. L. & Schultz T. R."box="[675,868,1547,1574]"pageId="45"pageNumber="46"pagination="61 - 81"refId="ref33114"refString="Ward P. S., Brady S. G., Fisher B. L. & Schultz T. R. 2015. The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40: 61 - 81. http: // dx. doi. org / 10.1111 / syen. 12090"type="journal article"year="2015">
). The subfamilies are organized by the systematic results of
<bibRefCitationid="EFB64B6BFF85B148FEEAF9C5FDAB197D"author="Brady S. G. & Schultz T. R. & Fisher B. L. & Ward P. S."box="[346,575,1582,1609]"pageId="45"pageNumber="46"pagination="18172 - 18177"refId="ref30264"refString="Brady S. G., Schultz T. R., Fisher B. L. & Ward P. S. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the United States of America 103: 18172 - 18177. http: // dx. doi. org / 10.1073 / pnas. 0605858103"type="journal article"year="2006">
<bibRefCitationid="EFB64B6BFF85B148FDCBF9C5FCCE197D"author="Ward P. S. & Brady S. G. & Fisher B. L. & Schultz T. R."box="[635,858,1582,1609]"pageId="45"pageNumber="46"pagination="61 - 81"refId="ref33114"refString="Ward P. S., Brady S. G., Fisher B. L. & Schultz T. R. 2015. The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology 40: 61 - 81. http: // dx. doi. org / 10.1111 / syen. 12090"type="journal article"year="2015">
<taxonomicNameid="4C274D19FF85B148FF53F99FFE5A19BB"authorityName="M. R. Smith"authorityYear="1952"box="[227,462,1652,1679]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="45"pageNumber="46"phylum="Arthropoda"rank="subFamily"subFamily="Pseudomyrmecinae">Pseudomyrmecinae</taxonomicName>
<taxonomicNameid="4C274D19FF85B148FC74F973FBC31986"authorityName="Lepeletier de Saint-Fargeau"authorityYear="1835"box="[964,1111,1688,1714]"class="Insecta"family="Formicidae"kingdom="Animalia"order="Hymenoptera"pageId="45"pageNumber="60"phylum="Arthropoda"rank="subFamily"subFamily="Myrmicinae">Myrmicinae</taxonomicName>
