treatments-xml/data/03/B9/87/03B987A9437DFF91FF46F992FBA7DEA0.xml

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<mods:title>Phylogeny, evolution, and classification of the ant genus Lasius, the tribe Lasiini and the subfamily Formicinae (Hymenoptera: Formicidae)</mods:title>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Boudinot, Brendon E.</mods:namePart>
<mods:affiliation>Department of Entomology and Nematology, University of California, Davis CA, U. S. A. &amp; Friedrich-Schiller-Universität Jena, Institut für Spezielle Zoologie, Jena, Germany</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Borowiec, Marek L.</mods:namePart>
<mods:affiliation>Department of Entomology and Nematology, University of California, Davis CA, U. S. A. &amp; Department of Plant Pathology, Entomology and Nematology, University of Idaho, Moscow ID, U. S. A. &amp; Institute for Bioinformatics and Evolutionary Studies, University of Idaho, Moscow ID, U. S. A.</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Prebus, Matthew M.</mods:namePart>
<mods:affiliation>Department of Entomology and Nematology, University of California, Davis CA, U. S. A. &amp; School of Life Sciences, Arizona State University, Tempe AZ, U. S. A.</mods:affiliation>
<mods:nameIdentifier type="email">mprebus@gmail.com</mods:nameIdentifier>
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<mods:title>Systematic Entomology</mods:title>
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<mods:date>2022</mods:date>
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<mods:number>2022-01-10</mods:number>
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<mods:number>47</mods:number>
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<mods:identifier type="DOI">10.1111/syen.12522</mods:identifier>
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Tribe 
<taxonomicName authority="Ashmead, 1905" authorityName="Ashmead" authorityYear="1905" box="[247,491,1543,1565]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="128" phylum="Arthropoda" rank="tribe" tribe="Lasiini">
Lasiini 
<bibRefCitation author="Ashmead, W. H." box="[330,491,1543,1565]" journalOrPublisher="The Canadian Entomologist" pageId="15" pageNumber="128" pagination="381 - 384" part="37" refId="ref30701" refString="Ashmead, W. H. (1905) A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea. The Canadian Entomologist, 37, 381 - 384." title="A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea" type="journal article" year="1905">Ashmead, 1905</bibRefCitation>
</taxonomicName>
</emphasis>
(
<figureCitation box="[510,579,1543,1564]" captionStart="Fig" captionStartId="16.[151,184,1221,1241]" captionTargetBox="[157,1422,174,1191]" captionTargetId="figure-8@16.[156,1423,172,1191]" captionTargetPageId="16" captionText="Fig. 6. Gross phenotypic synopsis of the Lasiini; all images in profile view; scale bars = 1.0 mm. (A) Cladomyrma hewitti (CASENT0173906); (B) Lasius lasioides (CASENT0906077); (C) Lasius citrinus (CASENT0103542); (D) Myrmecocystus melliger (CASENT0103518); (E) Metalasius myrmidon (CASENT0903666); (F) Euprenolepis procera (CASENT0906260); (G) Nylanderia amblyops (CASENT0007735); (H) Paraparatrechina albipes (CASENT0178759); (I) Paratrechina ankarana (CASENT0454372); (J) Prenolepis imparis (CASENT0179615); (K) Pseudolasius amaurops (CASENT0106005); (L) Zatania gibberosa (CASENT0281461). (Image credits, AntWeb: A, C, G, H = April Nobile; B = Shannon Hartman; D = Jen Fogarty; E, Will Ericson; F, Estella Ortega; I, Michele Esposito; J, Erin Prado; K, Michael Branstetter; L, Ziv Lieberman)." figureDoi="http://doi.org/10.5281/zenodo.5975364" httpUri="https://zenodo.org/record/5975364/files/figure.png" pageId="15" pageNumber="128">Figs 6</figureCitation>
, 
<figureCitation box="[595,609,1543,1564]" captionStart="Fig" captionStartId="17.[161,194,1432,1452]" captionTargetBox="[282,1315,173,1400]" captionTargetId="figure-8@17.[281,1316,172,1401]" captionTargetPageId="17" captionText="Fig. 7. Some key features of the major clades of the Lasiini. (A) third abdominal tergite modified with dorsoventrally-oriented groove for reception of entire petiole (Pseudolasius breviceps); black line indicates contour of tergum and black ellipse the spiracle; (B) third abdominal tergite receiving only posterior base of petiole (Myrmecocystus mimicus); white line indicates contour of tergum, and white ellipse the spiracle; (C) frontal protuberance and effaced carinae low relative to face (P. breviceps); (D) frontal protuberance and effaced carinae raised dorsally from face (M. mimicus); (E) metapleural gland atrium grossly enlarged (Lasius subumbratus); black lines show larger size of the metapleural gland bulla (lower indicator) as compared to F, and relative to the propodeal spiracle (upper indicator) and space between the spiracle and bulla (middle portion); (F) metapleural gland atrium small (Lasius turcicus); black lines show smaller size of metapleural gland bulla as compared to E, and relative to the propodeal spiracle and space between spiracle and bulla." figureDoi="http://doi.org/10.5281/zenodo.5975368" httpUri="https://zenodo.org/record/5975368/files/figure.png" pageId="15" pageNumber="128">7</figureCitation>
, 
<figureCitation box="[625,690,1543,1564]" captionStart="Fig" captionStartId="18.[151,184,1627,1647]" captionTargetBox="[157,1422,226,1596]" captionTargetId="figure-8@18.[156,1423,225,1597]" captionTargetPageId="18" captionText="Fig. 8. Gross phenotypic synopsis of the extant species groups of Lasius, plus two phenotypically similar fossil species; all images in full-face view; scale bars = 0.5 mm. (A) Lasius brunneus (CASENT0280440); (B) Lasius fuliginosus (CASENT0179898); (C) Lasius nr. niger (CASENT0106128); (D) Lasius nr. atopus (CASENT0234858); (E) Lasius carniolicus (CASENT0280471); (F) Lasius claviger (CASENT0103542); (G) Lasius brevicornis (CASENT0280456); (H) Lasius nearcticus (CASENT0104774); (I) Lasius pallitarsis (CASENT0005405); (J) Lasius aphidicola (CASENT0280468); (K) †Lasius schiefferdeckeri (MNHNB25217); (L) †Kyromyrma neffi (AMNH-NJ1029). (Image credits: D = the authors; AntWeb: A = Will Ericson; B = Erin Prado; C = Michael Branstetter; E, G, J = Shannon Hartman; F, H = April Nobile; I = unknown; K = Vincent Perrichot; L = Dave Grimaldi &amp; Vincent Perrichot.)" figureDoi="http://doi.org/10.5281/zenodo.5975370" httpUri="https://zenodo.org/record/5975370/files/figure.png" pageId="15" pageNumber="128">8A–K</figureCitation>
, 
<figureCitation box="[706,762,1543,1564]" captionStart="Fig" captionStartId="19.[161,194,1192,1212]" captionTargetBox="[185,1412,173,1160]" captionTargetId="figure-8@19.[184,1413,172,1161]" captionTargetPageId="19" captionText="Fig. 9. Gross phenotypic synopsis of the species groups of Lasius, plus two phenotypically similar fossil species; all images in profile view; scale bars = 1.0 mm; scale bar unavailable for K. (A) Lasius brunneus (CASENT0280440); (B) Lasius fuliginosus (CASENT0179898); (C) Lasius nr. niger (CASENT0106128); (D) Lasius nr. atopus (CASENT0234858); (E) Lasius carniolicus (CASENT0280471); (F) Lasius claviger (CASENT0103542); (G) Lasius brevicornis (CASENT0280456); (H) Lasius nearcticus (CASENT0104774); (I) Lasius pallitarsis (CASENT0005405); (J) Lasius aphidicola (CASENT0280468); (K) †Lasius schiefferdeckeri (HJF013); (L) †Kyromyrma neffi (AMNH-NJ1029). (Image credits: D = the authors; AntWeb: A = Will Ericson; B = Erin Prado; C = Michael Branstetter; E, G, J = Shannon Hartman; F, H = April Nobile; I = unknown; K = Vincent Perrichot; L = Dave Grimaldi &amp; Vincent Perrichot.)" figureDoi="http://doi.org/10.5281/zenodo.5975374" httpUri="https://zenodo.org/record/5975374/files/figure.png" pageId="15" pageNumber="128">9A–J</figureCitation>
, 
<figureCitation box="[161,237,1573,1594]" captionStart="Fig" captionStartId="20.[151,184,1169,1189]" captionTargetBox="[307,1271,173,1137]" captionTargetId="figure-8@20.[306,1272,172,1138]" captionTargetPageId="20" captionText="Fig. 10. Habitus images for the three new genera recognized in this study; scale bars = 0.25 mm. (A, B) Metalasius myrmidon gen.nov. comb.nov. (CASENT0906115); (C, D) †Metalasius pumilus gen.nov. comb.nov. (SMFBE1226); (E, F) †Sussudio gen.nov. comb.nov. (GZG-BST04646). Views: A = full-face; B, E = profile; C = laterally oblique facial; D, F = dorsal. Observe: (1) The compound eyes of Metalasius myrmidon and †M. pumilus are mid-set, and (2) †Sussudio gen.nov. has a distinctly-carinate propodeal foramen, vertical petiolar node and nongrooved (lateromedially convex) third abdominal tergite." figureDoi="http://doi.org/10.5281/zenodo.5975378" httpUri="https://zenodo.org/record/5975378/files/figure.png" pageId="15" pageNumber="128">10A–D</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection box="[179,366,1603,1624]" pageId="15" pageNumber="128" type="type_taxon">
<paragraph blockId="15.[161,767,1364,1714]" box="[179,366,1603,1624]" pageId="15" pageNumber="128">
<emphasis box="[179,360,1603,1624]" italics="true" pageId="15" pageNumber="128">
Type genus. 
<taxonomicName authorityName="Fabricius" authorityYear="1804" box="[298,360,1603,1624]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="128" phylum="Arthropoda" rank="genus">Lasius</taxonomicName>
</emphasis>
.
</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="128" type="discussion">
<paragraph blockId="15.[161,767,1364,1714]" pageId="15" pageNumber="128">
<emphasis box="[179,538,1633,1654]" italics="true" pageId="15" pageNumber="128">
Included genus groups. 
<taxonomicName authorityName="Wheeler" authorityYear="1920" box="[413,538,1633,1654]" class="Insecta" family="Formicidae" genus="Cladomyrma" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="128" phylum="Arthropoda" rank="genus">Cladomyrma</taxonomicName>
</emphasis>
, 
<taxonomicName authorityName="Fabricius" authorityYear="1804" box="[551,613,1633,1654]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="128" phylum="Arthropoda" rank="genus">
<emphasis box="[551,613,1633,1654]" italics="true" pageId="15" pageNumber="128">Lasius</emphasis>
</taxonomicName>
group, 
<taxonomicName authorityName="Boudinot &amp; Borowiec &amp; Prebus" authorityYear="2022" class="Insecta" family="Formicidae" genus="Metalasius" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="128" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="15" pageNumber="128">Metalasius</emphasis>
</taxonomicName>
, 
<taxonomicName authorityName="Mayr" authorityYear="1861" box="[210,313,1663,1684]" class="Insecta" family="Formicidae" genus="Prenolepis" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="128" phylum="Arthropoda" rank="genus">
<emphasis box="[210,313,1663,1684]" italics="true" pageId="15" pageNumber="128">Prenolepis</emphasis>
</taxonomicName>
group.
</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="128" type="description">
<paragraph blockId="15.[161,767,1364,1714]" box="[179,370,1693,1714]" pageId="15" pageNumber="128">
<emphasis box="[179,364,1693,1714]" italics="true" pageId="15" pageNumber="128">Definition (worker)</emphasis>
.
</paragraph>
<paragraph blockId="15.[174,767,1752,1863]" pageId="15" pageNumber="128">
1. With characters of 
<taxonomicName baseAuthorityName="Lepeletier de Saint-Fargeau" box="[410,519,1752,1773]" class="Insecta" family="Formicidae" genus="Formicinae" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="150" phylum="Arthropoda" rank="subFamily" subFamily="Formicinae">Formicinae</taxonomicName>
(see 
<bibRefCitation author="Bolton, B." box="[585,718,1752,1773]" journalOrPublisher="Memoirs of the American Entomological Institute" pageId="15" pageNumber="128" pagination="1 - 370" part="71" refId="ref31502" refString="Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370." title="Synopsis and classification of Formicidae" type="journal article" year="2003">Bolton, 2003</bibRefCitation>
and Note 1).
</paragraph>
<paragraph blockId="15.[174,767,1752,1863]" pageId="15" pageNumber="128">2. Mandible triangular, with 4–11 teeth; third tooth from apex reduced.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">3. Dorsal mandibular groove, when present, running along lateral mandibular margin as seen in dorsal view (Note 2).</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[844,1253,232,253]" pageId="15" pageNumber="128">4. Palp formula usually 6,4, less often 3,4.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">5. Frontal protuberance mediad antennal toruli with frontal carinae effaced, becoming broadly rounded posteriorly until continuous with face (Note 3).</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[844,1326,351,372]" pageId="15" pageNumber="128">6. Frontal protuberance raised above toruli or not.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[844,1422,381,402]" pageId="15" pageNumber="128">7. Antennal toruli near or abutting posterior clypeal margin.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[844,1171,411,432]" pageId="15" pageNumber="128">8. Antenna 12-, 11-, or 8-merous.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">9. Compound eyes not enormous, that is, not longer than half the length of the head; eyes may be reduced to absent; long axis of eyes subparallel.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[832,1193,530,551]" pageId="15" pageNumber="128">10. Ocelli present or absent (Note 4).</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[832,1187,560,581]" pageId="15" pageNumber="128">11. Metanotum differentiated or not.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">12. Metapleural gland present, dorsal rim of metapleural gland curved inward.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">13. Propodeal spiracle at or near posterolateral margin of propodeum.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[832,1403,709,730]" pageId="15" pageNumber="128">14. Propodeal spiracle circular to elliptical, not slit-shaped.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">15. Metacoxae wideset: Distance between mesocoxal bases less than between that between bases of metacoxae with mesosoma in ventral view and coxae oriented at right-angles to long axis of body.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[832,1398,858,879]" pageId="15" pageNumber="128">16. Metatibiae without double row of ventral (inner) setae.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">17. Petiolar foramen inprofile view low, not to barelyexceeding dorsal margin of metapleural gland, with or without dorsal margin or lip, but lip, when present, inconspicuous (Note 5).</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">18. Petiolar foramen in ventral view long, anterior margin exceeding metacoxal cavities anteriorly.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">19. Petiolar node conspicuously shorter than propodeum, not reaching dorsal surface of propodeum (Note 6).</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">20. Petiolar apodeme (situated anteriorly on tergum) contiguous or nearly contiguous with petiolar node (Note 7).</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[832,1376,1157,1178]" pageId="15" pageNumber="128">21. Petiolar sternum U-shaped in cross-section (Note 8).</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[832,1339,1186,1207]" pageId="15" pageNumber="128">22. Abdominal segment III transverse sulcus absent.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" pageId="15" pageNumber="128">23. Base of abdominal segment III with or without complete tergosternal fusion lateral to helcium, free sclerites commencing distantly up segment or near helcium; tergum III overhanging petiole or not.</paragraph>
<paragraph blockId="15.[832,1436,172,1356]" box="[832,1193,1335,1356]" pageId="15" pageNumber="128">24. Proventriculus sepalous (Note 9).</paragraph>
</subSubSection>
<subSubSection lastPageId="19" lastPageNumber="132" pageId="15" pageNumber="128" type="discussion">
<paragraph blockId="15.[830,1437,1394,1863]" box="[830,1029,1394,1416]" pageId="15" pageNumber="128">
<heading bold="true" box="[830,1029,1394,1416]" fontSize="9" level="1" pageId="15" pageNumber="128" reason="6">
<emphasis bold="true" box="[830,1022,1394,1416]" pageId="15" pageNumber="128">Notes on definition</emphasis>
:
</heading>
</paragraph>
<paragraph blockId="15.[830,1437,1394,1863]" pageId="15" pageNumber="128">
<emphasis box="[830,903,1424,1445]" italics="true" pageId="15" pageNumber="128">Note 1.</emphasis>
The generic composition of the 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[1240,1306,1424,1445]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
has recently been reassessed following 
<bibRefCitation author="Blaimer, B. B. &amp; Brady, S. G. &amp; Schultz, T. R. &amp; Lloyd, M. W. &amp; Fisher, B. L. &amp; Ward, P. S." box="[1127,1334,1454,1475]" journalOrPublisher="BMC Evolutionary Biology" pageId="15" pageNumber="128" pagination="257" part="15" refId="ref31116" refString="Blaimer, B. B., Brady, S. G., Schultz, T. R., Lloyd, M. W., Fisher, B. L. &amp; Ward, P. S. (2015) Phylogenomic methods outperform traditional multi-locus approaches in resolving deep evolutionary history: a case study of formicine ants. BMC Evolutionary Biology, 15, 257." title="Phylogenomic methods outperform traditional multi-locus approaches in resolving deep evolutionary history: a case study of formicine ants" type="journal article" year="2015">
Blaimer 
<emphasis box="[1212,1263,1454,1475]" italics="true" pageId="15" pageNumber="128">et al.</emphasis>
(2015)
</bibRefCitation>
and 
<bibRefCitation author="Ward, P. S. &amp; Blaimer, B. B. &amp; Fisher, B. L." journalOrPublisher="Zootaxa" pageId="15" pageNumber="128" pagination="343 - 357" part="4072" refId="ref39006" refString="Ward, P. S., Blaimer, B. B. &amp; Fisher, B. L. (2016) A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinoponera. Zootaxa, 4072, 343 - 357. https: // doi. org / 10.11646 / Zootaxa. 4072.3.4." title="A revised phylogenetic classification of the ant subfamily Formicinae (Hymenoptera: Formicidae), with resurrection of the genera Colobopsis and Dinoponera" type="journal article" year="2016">
Ward 
<emphasis box="[866,919,1484,1505]" italics="true" pageId="15" pageNumber="128">et al.</emphasis>
(2016)
</bibRefCitation>
. The tribal definition proposed here is thus revised relative to 
<bibRefCitation author="Bolton, B." box="[1041,1179,1514,1535]" journalOrPublisher="Memoirs of the American Entomological Institute" pageId="15" pageNumber="128" pagination="1 - 370" part="71" refId="ref31502" refString="Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370." title="Synopsis and classification of Formicidae" type="journal article" year="2003">Bolton (2003)</bibRefCitation>
. It includes new characters and recognizes that several former plagiolepidines belong in the 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[901,971,1573,1594]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
.
</paragraph>
<paragraph blockId="15.[830,1437,1394,1863]" lastBlockId="16.[151,758,1444,1863]" lastPageId="16" lastPageNumber="129" pageId="15" pageNumber="128">
<emphasis box="[830,906,1603,1624]" italics="true" pageId="15" pageNumber="128">Note 2.</emphasis>
Previous studies have not focused attention on the dorsal mandibular groove. Here it was observed that the groove, as seen with the mandible in dorsal view, runs along the outer margin of the mandible toward the mandibular apex in the Myrmelachistini, 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[1146,1212,1722,1743]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
and most Melophorini. This contrasts with the state observed in the ‘formicoform radiation’ (i.e., the clade that includes Camponotini, 
<taxonomicName authorityName="Latreille" authorityYear="1802" box="[866,960,1812,1833]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="15" pageNumber="147" phylum="Arthropoda" rank="tribe" tribe="Formicini">Formicini</taxonomicName>
and Melophorini, along with the monotypic tribes Gesomyrmecini, Gigantiopini, Oecophyllini, and Santschiellini; see the section † 
<taxonomicName authorityName="Grimaldi &amp; Agosti" authorityYear="2000" box="[509,621,1444,1465]" class="Insecta" family="Formicidae" genus="Kyromyrma" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="genus">
<emphasis box="[509,621,1444,1465]" italics="true" pageId="16" pageNumber="129">Kyromyrma</emphasis>
</taxonomicName>
, an ancestral formicine, under the section ‘ 
<emphasis box="[472,606,1474,1495]" italics="true" pageId="16" pageNumber="129">Incertae sedis</emphasis>
in the Formicinae’ below). When the groove is present in the ‘formicoform radiation’, it is very close to the basal mandibular margin and is shortened, except in various plagiolepidines. The medial and shortened state may have arisen multiple times in the 
<taxonomicName authorityName="Latreille" authorityYear="1802" box="[247,343,1627,1648]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="147" phylum="Arthropoda" rank="tribe" tribe="Formicini">Formicini</taxonomicName>
, Plagiolepidini, and Camponotini clade.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5975364" ID-Zenodo-Dep="5975364" httpUri="https://zenodo.org/record/5975364/files/figure.png" pageId="16" pageNumber="129" startId="16.[151,184,1221,1241]" targetBox="[157,1422,174,1191]" targetPageId="16">
<paragraph blockId="16.[151,1428,1221,1374]" pageId="16" pageNumber="129">
<emphasis bold="true" box="[151,214,1221,1241]" pageId="16" pageNumber="129">Fig. 6.</emphasis>
Gross phenotypic synopsis of the 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[524,586,1222,1241]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
; all images in profile view; scale bars = 1.0 mm. (A) 
<taxonomicName baseAuthorityName="Wheeler" baseAuthorityYear="1910" box="[1066,1242,1222,1241]" class="Insecta" family="Formicidae" genus="Cladomyrma" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="hewitti">
<emphasis box="[1066,1242,1222,1241]" italics="true" pageId="16" pageNumber="129">Cladomyrma hewitti</emphasis>
</taxonomicName>
(CASENT0173906); (B) 
<taxonomicName baseAuthorityName="Emery" baseAuthorityYear="1869" box="[186,324,1249,1268]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="lasioides">
<emphasis box="[186,324,1249,1268]" italics="true" pageId="16" pageNumber="129">Lasius lasioides</emphasis>
</taxonomicName>
(CASENT0906077); (C) 
<taxonomicName authorityName="Emery" authorityYear="1922" box="[548,675,1249,1268]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="citrinus">
<emphasis box="[548,675,1249,1268]" italics="true" pageId="16" pageNumber="129">Lasius citrinus</emphasis>
</taxonomicName>
(CASENT0103542); (D) 
<taxonomicName baseAuthorityName="Llave" baseAuthorityYear="1832" box="[900,1110,1249,1268]" class="Insecta" family="Formicidae" genus="Myrmecocystus" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="melliger">
<emphasis box="[900,1110,1249,1268]" italics="true" pageId="16" pageNumber="129">Myrmecocystus melliger</emphasis>
</taxonomicName>
(CASENT0103518); (E) 
<taxonomicName authorityName="Boudinot &amp; Borowiec &amp; Prebus" authorityYear="2022" baseAuthorityName="Mei" baseAuthorityYear="1998" class="Insecta" family="Formicidae" genus="Metalasius" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="myrmidon">
<emphasis italics="true" pageId="16" pageNumber="129">Metalasius myrmidon</emphasis>
</taxonomicName>
(CASENT0903666); (F) 
<taxonomicName baseAuthorityName="Emery" baseAuthorityYear="1900" box="[459,644,1276,1295]" class="Insecta" family="Formicidae" genus="Euprenolepis" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="procera">
<emphasis box="[459,644,1276,1295]" italics="true" pageId="16" pageNumber="129">Euprenolepis procera</emphasis>
</taxonomicName>
(CASENT0906260); (G) 
<emphasis box="[869,1051,1276,1295]" italics="true" pageId="16" pageNumber="129">Nylanderia amblyops</emphasis>
(CASENT0007735); (H) 
<emphasis italics="true" pageId="16" pageNumber="129">Paraparatrechina albipes</emphasis>
(CASENT0178759); (I) 
<taxonomicName authorityName="LaPolla &amp; Fisher" authorityYear="2014" box="[428,624,1302,1321]" class="Insecta" family="Formicidae" genus="Paratrechina" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="ankarana">
<emphasis box="[428,624,1302,1321]" italics="true" pageId="16" pageNumber="129">Paratrechina ankarana</emphasis>
</taxonomicName>
(CASENT0454372); (J) 
<taxonomicName baseAuthorityName="Say" baseAuthorityYear="1836" box="[840,1002,1302,1321]" class="Insecta" family="Formicidae" genus="Prenolepis" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="imparis">
<emphasis box="[840,1002,1302,1321]" italics="true" pageId="16" pageNumber="129">Prenolepis imparis</emphasis>
</taxonomicName>
(CASENT0179615); (K) 
<taxonomicName authorityName="Emery" authorityYear="1922" box="[1225,1427,1302,1321]" class="Insecta" family="Formicidae" genus="Pseudolasius" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="species" species="amaurops">
<emphasis box="[1225,1427,1302,1321]" italics="true" pageId="16" pageNumber="129">Pseudolasius amaurops</emphasis>
</taxonomicName>
(CASENT0106005); (L) 
<emphasis box="[366,521,1329,1348]" italics="true" pageId="16" pageNumber="129">Zatania gibberosa</emphasis>
(CASENT0281461). (Image credits, AntWeb: A, C, G, H = April Nobile; B = Shannon Hartman; D = Jen Fogarty; E, Will Ericson; F, Estella Ortega; I, Michele Esposito; J, Erin Prado; K, Michael Branstetter; L, Ziv Lieberman).
</paragraph>
</caption>
<paragraph blockId="16.[151,758,1444,1863]" lastBlockId="16.[821,1427,1443,1863]" pageId="16" pageNumber="129">
<emphasis box="[151,225,1658,1679]" italics="true" pageId="16" pageNumber="129">Note 3.</emphasis>
The ‘frontal protuberance’ is the region between the antennal toruli, which is raised relative to the regions of the face laterad the toruli. To some degree, development of this condition accounts for the ‘laterally directed’ torular condition of 
<taxonomicName authority="(Boudinot et al., 2020)" baseAuthorityName="Boudinot" baseAuthorityYear="2020" box="[316,666,1780,1802]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="family">
Formicidae (
<bibRefCitation author="Boudinot, B. E. &amp; Perrichot, V. &amp; Chaul, J. C. M." box="[441,659,1780,1802]" journalOrPublisher="ZooKeys" pageId="16" pageNumber="129" pagination="21 - 55" part="1005" refId="ref31850" refString="Boudinot, B. E., Perrichot, V. &amp; Chaul, J. C. M. (2020) † Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicidae). ZooKeys, 1005, 21 - 55." title="† Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicidae)" type="journal article" year="2020">
Boudinot 
<emphasis box="[541,593,1781,1802]" italics="true" pageId="16" pageNumber="129">et al.</emphasis>
, 2020
</bibRefCitation>
)
</taxonomicName>
. In other formicine tribes, including the Melophorini (with 
<taxonomicName authorityName="Forel" authorityYear="1892" box="[667,758,1811,1832]" class="Insecta" family="Formicidae" genus="Prolasius" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="genus">
<emphasis box="[667,758,1811,1832]" italics="true" pageId="16" pageNumber="129">Prolasius</emphasis>
</taxonomicName>
as an exception), the frontal protuberance is carinate above the toruli (
<emphasis box="[969,976,1443,1464]" italics="true" pageId="16" pageNumber="129">i</emphasis>
. 
<emphasis box="[982,993,1443,1464]" italics="true" pageId="16" pageNumber="129">e</emphasis>
., ‘frontal carinae are present’), with these carinae continuing as a sharp ridge until their posterior terminus. Within the 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[1058,1127,1505,1526]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
, the frontal carinae are effaced; they are long in all groups except 
<taxonomicName authorityName="Wheeler" authorityYear="1920" box="[1205,1330,1535,1556]" class="Insecta" family="Formicidae" genus="Cladomyrma" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="genus">
<emphasis box="[1205,1330,1535,1556]" italics="true" pageId="16" pageNumber="129">Cladomyrma</emphasis>
</taxonomicName>
, while in the Plagiolepidini the carinae are short in all genera except 
<taxonomicName authorityName="Santschi" box="[857,971,1597,1618]" class="Insecta" family="Formicidae" genus="Anoplolepis" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="genus">
<emphasis box="[857,971,1597,1618]" italics="true" pageId="16" pageNumber="129">Anoplolepis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="16.[821,1427,1443,1863]" lastBlockId="17.[197,767,1712,1863]" lastPageId="17" lastPageNumber="130" pageId="16" pageNumber="129">
<emphasis box="[821,895,1627,1648]" italics="true" pageId="16" pageNumber="129">Note 4.</emphasis>
Ocellus presence–absence is arelatively weak character as expression of ocelli may be variable for genera in which ocelli are observed, such as 
<taxonomicName authorityName="Fabricius" authorityYear="1804" box="[1200,1262,1689,1710]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="genus">
<emphasis box="[1200,1262,1689,1710]" italics="true" pageId="16" pageNumber="129">Lasius</emphasis>
</taxonomicName>
and 
<emphasis box="[1313,1421,1689,1710]" italics="true" pageId="16" pageNumber="129">Nylanderia</emphasis>
. However, this statement is included as ocellus expression is a traditional character which is easy to evaluate and may have value for future works defining tribes wherein ocelli may be consistently present or absent. Within the 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[1354,1423,1811,1832]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
, ocelli are always absent in 
<taxonomicName authorityName="Wheeler" authorityYear="1920" box="[1155,1280,1842,1863]" class="Insecta" family="Formicidae" genus="Cladomyrma" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="genus">
<emphasis box="[1155,1280,1842,1863]" italics="true" pageId="16" pageNumber="129">Cladomyrma</emphasis>
</taxonomicName>
, 
<taxonomicName authorityName="Emery" box="[1300,1427,1842,1863]" class="Insecta" family="Formicidae" genus="Euprenolepis" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="129" phylum="Arthropoda" rank="genus">
<emphasis box="[1300,1427,1842,1863]" italics="true" pageId="16" pageNumber="129">Euprenolepis</emphasis>
</taxonomicName>
Emery, and 
<taxonomicName authority="Emery" authorityName="Emery" authorityYear="1887" box="[320,521,1712,1733]" class="Insecta" family="Formicidae" genus="Pseudolasius" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="genus">
<emphasis box="[320,446,1712,1733]" italics="true" pageId="17" pageNumber="130">Pseudolasius</emphasis>
Emery
</taxonomicName>
, variably present among 
<taxonomicName authority=", Paraparatrechina Donisthorpe" authorityName="Paraparatrechina Donisthorpe" box="[197,590,1745,1766]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="genus">
<emphasis box="[197,259,1745,1766]" italics="true" pageId="17" pageNumber="130">Lasius</emphasis>
, 
<emphasis box="[283,452,1745,1766]" italics="true" pageId="17" pageNumber="130">Paraparatrechina</emphasis>
Donisthorpe
</taxonomicName>
, and 
<taxonomicName pageId="17" pageNumber="130">
<emphasis box="[664,767,1745,1766]" italics="true" pageId="17" pageNumber="130">Prenolepis</emphasis>
Mayr species
</taxonomicName>
, and consistently present in 
<taxonomicName authorityName="Wasmael" box="[612,761,1777,1798]" class="Insecta" family="Formicidae" genus="Myrmecocystus" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="genus">
<emphasis box="[612,761,1777,1798]" italics="true" pageId="17" pageNumber="130">Myrmecocystus</emphasis>
</taxonomicName>
, 
<taxonomicName authority="Motschoulsky" authorityName="Motschoulsky" authorityYear="1863" box="[197,462,1809,1831]" class="Insecta" family="Formicidae" genus="Paratrechina" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="genus">
<emphasis box="[197,321,1810,1831]" italics="true" pageId="17" pageNumber="130">Paratrechina</emphasis>
Motschoulsky
</taxonomicName>
(except 
<taxonomicName authorityName="Boudinot &amp; Borowiec &amp; Prebus" authorityYear="2022" baseAuthorityName="Boudinot &amp; Borowiec &amp; Prebus" baseAuthorityYear="2022" box="[546,618,1810,1831]" class="Insecta" family="Formicidae" genus="Paratrechina" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="species" species="kohli">
<emphasis box="[546,618,1810,1831]" italics="true" pageId="17" pageNumber="130">P. kohli</emphasis>
</taxonomicName>
; see comments under 
<taxonomicName authorityName="Mayr" authorityYear="1861" box="[257,360,1842,1863]" class="Insecta" family="Formicidae" genus="Prenolepis" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="genus">
<emphasis box="[257,360,1842,1863]" italics="true" pageId="17" pageNumber="130">Prenolepis</emphasis>
</taxonomicName>
genus group), and 
<emphasis box="[544,618,1842,1863]" italics="true" pageId="17" pageNumber="130">Zatania</emphasis>
LaPolla 
<emphasis box="[704,753,1842,1863]" italics="true" pageId="17" pageNumber="130">et al.</emphasis>
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5975368" ID-Zenodo-Dep="5975368" httpUri="https://zenodo.org/record/5975368/files/figure.png" pageId="17" pageNumber="130" startId="17.[161,194,1432,1452]" targetBox="[282,1315,173,1400]" targetPageId="17">
<paragraph blockId="17.[161,1437,1432,1638]" pageId="17" pageNumber="130">
<emphasis bold="true" box="[161,223,1432,1452]" pageId="17" pageNumber="130">Fig. 7.</emphasis>
Some key features of the major clades of the 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[613,675,1433,1452]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
. (A) third abdominal tergite modified with dorsoventrally-oriented groove for reception of entire petiole (
<taxonomicName authorityName="Emery" authorityYear="1887" box="[284,482,1460,1479]" class="Insecta" family="Formicidae" genus="Pseudolasius" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="species" species="breviceps">
<emphasis box="[284,482,1460,1479]" italics="true" pageId="17" pageNumber="130">Pseudolasius breviceps</emphasis>
</taxonomicName>
); black line indicates contour of tergum and black ellipse the spiracle; (B) third abdominal tergite receiving only posterior base of petiole (
<taxonomicName authorityName="Wheeler" authorityYear="1908" box="[377,586,1486,1505]" class="Insecta" family="Formicidae" genus="Myrmecocystus" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="species" species="mimicus">
<emphasis box="[377,586,1486,1505]" italics="true" pageId="17" pageNumber="130">Myrmecocystus mimicus</emphasis>
</taxonomicName>
); white line indicates contour of tergum, and white ellipse the spiracle; (C) frontal protuberance and effaced carinae low relative to face (
<taxonomicName authorityName="Emery" authorityYear="1887" box="[466,566,1513,1532]" class="Insecta" family="Formicidae" genus="Pseudolasius" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="species" species="breviceps">
<emphasis box="[466,566,1513,1532]" italics="true" pageId="17" pageNumber="130">P. breviceps</emphasis>
</taxonomicName>
); (D) frontal protuberance and effaced carinae raised dorsally from face (
<taxonomicName authorityName="Wheeler" authorityYear="1908" box="[1187,1286,1513,1532]" class="Insecta" family="Formicidae" genus="Myrmecocystus" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="species" species="mimicus">
<emphasis box="[1187,1286,1513,1532]" italics="true" pageId="17" pageNumber="130">M. mimicus</emphasis>
</taxonomicName>
); (E) metapleural gland atrium grossly enlarged (
<taxonomicName authorityName="Viereck" authorityYear="1903" box="[431,602,1539,1558]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="species" species="subumbratus">
<emphasis box="[431,602,1539,1558]" italics="true" pageId="17" pageNumber="130">Lasius subumbratus</emphasis>
</taxonomicName>
); black lines show larger size of the metapleural gland bulla (lower indicator) as compared to F, and relative to the propodeal spiracle (upper indicator) and space between the spiracle and bulla (middle portion); (F) metapleural gland atrium small (
<taxonomicName authorityName="Santschi" authorityYear="1921" box="[168,297,1592,1611]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="130" phylum="Arthropoda" rank="species" species="turcicus">
<emphasis box="[168,297,1592,1611]" italics="true" pageId="17" pageNumber="130">Lasius turcicus</emphasis>
</taxonomicName>
); black lines show smaller size of metapleural gland bulla as compared to E, and relative to the propodeal spiracle and space between spiracle and bulla.
</paragraph>
</caption>
<paragraph blockId="17.[830,1436,1712,1863]" lastBlockId="19.[161,767,1423,1863]" lastPageId="19" lastPageNumber="132" pageId="17" pageNumber="130">
<emphasis box="[830,903,1712,1733]" italics="true" pageId="17" pageNumber="130">Note 5.</emphasis>
The conformation of the dorsal region of the petiolar foramen is newly described here. There is complex variation of the form across the subfamily, but it appears at least that the form observed in the 
<taxonomicName authorityName="Ashmead" authorityYear="1905" box="[1182,1248,1809,1830]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="17" pageNumber="143" phylum="Arthropoda" rank="tribe" tribe="Lasiini">Lasiini</taxonomicName>
is consistent; this form is also observed in the Myrmelachistini. In various lineages within the formicoform radiation, a raised and conspicuously carinate dorsal margin is observed.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5975370" ID-Zenodo-Dep="5975370" httpUri="https://zenodo.org/record/5975370/files/figure.png" pageId="18" pageNumber="131" startId="18.[151,184,1627,1647]" targetBox="[157,1422,226,1596]" targetPageId="18">
<paragraph blockId="18.[151,1428,1627,1806]" pageId="18" pageNumber="131">
<emphasis bold="true" box="[151,214,1627,1647]" pageId="18" pageNumber="131">Fig. 8.</emphasis>
Gross phenotypic synopsis of the extant species groups of 
<taxonomicName authorityName="Fabricius" box="[723,778,1628,1647]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">
<emphasis box="[723,778,1628,1647]" italics="true" pageId="18" pageNumber="131">Lasius</emphasis>
</taxonomicName>
, plus two phenotypically similar fossil species; all images in full-face view; scale bars = 0.5 mm. (A) 
<taxonomicName baseAuthorityName="Latreille" baseAuthorityYear="1798" box="[370,511,1655,1674]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="brunneus">
<emphasis box="[370,511,1655,1674]" italics="true" pageId="18" pageNumber="131">Lasius brunneus</emphasis>
</taxonomicName>
(CASENT0280440); (B) 
<taxonomicName baseAuthorityName="Latreille" baseAuthorityYear="1798" box="[733,887,1655,1674]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="fuliginosus">
<emphasis box="[733,887,1655,1674]" italics="true" pageId="18" pageNumber="131">Lasius fuliginosus</emphasis>
</taxonomicName>
(CASENT0179898); (C) 
<taxonomicName authorityName="Fabricius" box="[1110,1166,1655,1674]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">
<emphasis box="[1110,1166,1655,1674]" italics="true" pageId="18" pageNumber="131">Lasius</emphasis>
</taxonomicName>
nr. 
<taxonomicName authorityName="" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1199,1244,1655,1674]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="niger">
<emphasis box="[1199,1244,1655,1674]" italics="true" pageId="18" pageNumber="131">niger</emphasis>
</taxonomicName>
(CASENT0106128); (D) 
<taxonomicName authorityName="Fabricius" box="[186,242,1681,1700]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="genus">
<emphasis box="[186,242,1681,1700]" italics="true" pageId="18" pageNumber="131">Lasius</emphasis>
</taxonomicName>
nr. 
<taxonomicName authorityName="Cole" authorityYear="1958" box="[272,329,1681,1700]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="atopus">
<emphasis box="[272,329,1681,1700]" italics="true" pageId="18" pageNumber="131">atopus</emphasis>
</taxonomicName>
(CASENT0234858); (E) 
<taxonomicName authorityName="Mayr" authorityYear="1861" box="[547,703,1681,1700]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="carniolicus">
<emphasis box="[547,703,1681,1700]" italics="true" pageId="18" pageNumber="131">Lasius carniolicus</emphasis>
</taxonomicName>
(CASENT0280471); (F) 
<taxonomicName baseAuthorityName="Roger" baseAuthorityYear="1862" box="[920,1049,1681,1700]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="claviger">
<emphasis box="[920,1049,1681,1700]" italics="true" pageId="18" pageNumber="131">Lasius claviger</emphasis>
</taxonomicName>
(CASENT0103542); (G) 
<taxonomicName authorityName="Emery" authorityYear="1893" box="[1271,1427,1681,1700]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="brevicornis">
<emphasis box="[1271,1427,1681,1700]" italics="true" pageId="18" pageNumber="131">Lasius brevicornis</emphasis>
</taxonomicName>
(CASENT0280456); (H) 
<taxonomicName authorityName="Wheeler" authorityYear="1906" box="[368,517,1708,1727]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="nearcticus">
<emphasis box="[368,517,1708,1727]" italics="true" pageId="18" pageNumber="131">Lasius nearcticus</emphasis>
</taxonomicName>
(CASENT0104774); (I) 
<taxonomicName baseAuthorityName="Provancher" baseAuthorityYear="1881" box="[731,878,1708,1727]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="pallitarsis">
<emphasis box="[731,878,1708,1727]" italics="true" pageId="18" pageNumber="131">Lasius pallitarsis</emphasis>
</taxonomicName>
(CASENT0005405); (J) 
<taxonomicName baseAuthorityName="Walsh" baseAuthorityYear="1863" box="[1093,1244,1708,1727]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="aphidicola">
<emphasis box="[1093,1244,1708,1727]" italics="true" pageId="18" pageNumber="131">Lasius aphidicola</emphasis>
</taxonomicName>
(CASENT0280468); (K) † 
<taxonomicName authorityName="Mayr" authorityYear="1868" box="[197,392,1734,1753]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="schiefferdeckeri">
<emphasis box="[197,392,1734,1753]" italics="true" pageId="18" pageNumber="131">Lasius schiefferdeckeri</emphasis>
</taxonomicName>
(MNHNB25217); (L) † 
<taxonomicName authority="(AMNH-NJ 1029)" baseAuthorityName="AMNH-NJ" baseAuthorityYear="1029" box="[599,901,1734,1753]" class="Insecta" family="Formicidae" genus="Kyromyrma" kingdom="Animalia" order="Hymenoptera" pageId="18" pageNumber="131" phylum="Arthropoda" rank="species" species="neffi">
<emphasis box="[599,740,1734,1753]" italics="true" pageId="18" pageNumber="131">Kyromyrma neffi</emphasis>
(AMNH-NJ1029)
</taxonomicName>
. (Image credits: D = the authors; AntWeb: A = Will Ericson; B = Erin Prado; C = Michael Branstetter; E, G, J = Shannon Hartman; F, H = April Nobile; I = unknown; K = Vincent Perrichot; L = Dave Grimaldi &amp; Vincent Perrichot.)
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5975374" ID-Zenodo-Dep="5975374" httpUri="https://zenodo.org/record/5975374/files/figure.png" pageId="19" pageNumber="132" startId="19.[161,194,1192,1212]" targetBox="[185,1412,173,1160]" targetPageId="19">
<paragraph blockId="19.[161,1438,1192,1371]" pageId="19" pageNumber="132">
<emphasis bold="true" box="[161,223,1192,1212]" pageId="19" pageNumber="132">Fig. 9.</emphasis>
Gross phenotypic synopsis of the species groups of 
<taxonomicName authorityName="Fabricius" box="[686,741,1193,1212]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[686,741,1193,1212]" italics="true" pageId="19" pageNumber="132">Lasius</emphasis>
</taxonomicName>
, plus two phenotypically similar fossil species; all images in profile view; scale bars = 1.0 mm; scale bar unavailable for K. (A) 
<taxonomicName baseAuthorityName="Latreille" baseAuthorityYear="1798" box="[568,708,1219,1238]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="brunneus">
<emphasis box="[568,708,1219,1238]" italics="true" pageId="19" pageNumber="132">Lasius brunneus</emphasis>
</taxonomicName>
(CASENT0280440); (B) 
<taxonomicName baseAuthorityName="Latreille" baseAuthorityYear="1798" box="[929,1082,1219,1238]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="fuliginosus">
<emphasis box="[929,1082,1219,1238]" italics="true" pageId="19" pageNumber="132">Lasius fuliginosus</emphasis>
</taxonomicName>
(CASENT0179898); (C) 
<taxonomicName authorityName="Fabricius" box="[1304,1360,1219,1238]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[1304,1360,1219,1238]" italics="true" pageId="19" pageNumber="132">Lasius</emphasis>
</taxonomicName>
nr. 
<taxonomicName authorityName="" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1391,1436,1219,1238]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="niger">
<emphasis box="[1391,1436,1219,1238]" italics="true" pageId="19" pageNumber="132">niger</emphasis>
</taxonomicName>
(CASENT0106128); (D) 
<taxonomicName authorityName="Fabricius" box="[379,435,1246,1265]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[379,435,1246,1265]" italics="true" pageId="19" pageNumber="132">Lasius</emphasis>
</taxonomicName>
nr. 
<taxonomicName authorityName="Cole" authorityYear="1958" box="[467,524,1246,1265]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="atopus">
<emphasis box="[467,524,1246,1265]" italics="true" pageId="19" pageNumber="132">atopus</emphasis>
</taxonomicName>
(CASENT0234858); (E) 
<taxonomicName authorityName="Mayr" authorityYear="1861" box="[745,902,1246,1265]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="carniolicus">
<emphasis box="[745,902,1246,1265]" italics="true" pageId="19" pageNumber="132">Lasius carniolicus</emphasis>
</taxonomicName>
(CASENT0280471); (F) 
<taxonomicName baseAuthorityName="Roger" baseAuthorityYear="1862" box="[1122,1252,1246,1265]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="claviger">
<emphasis box="[1122,1252,1246,1265]" italics="true" pageId="19" pageNumber="132">Lasius claviger</emphasis>
</taxonomicName>
(CASENT0103542); (G) 
<taxonomicName authorityName="Emery" authorityYear="1893" box="[194,350,1273,1292]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="brevicornis">
<emphasis box="[194,350,1273,1292]" italics="true" pageId="19" pageNumber="132">Lasius brevicornis</emphasis>
</taxonomicName>
(CASENT0280456); (H) 
<taxonomicName authorityName="Wheeler" authorityYear="1906" box="[569,717,1273,1292]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="nearcticus">
<emphasis box="[569,717,1273,1292]" italics="true" pageId="19" pageNumber="132">Lasius nearcticus</emphasis>
</taxonomicName>
(CASENT0104774); (I) 
<taxonomicName baseAuthorityName="Provancher" baseAuthorityYear="1881" box="[928,1073,1273,1292]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="pallitarsis">
<emphasis box="[928,1073,1273,1292]" italics="true" pageId="19" pageNumber="132">Lasiuspallitarsis</emphasis>
</taxonomicName>
(CASENT0005405); (J) 
<taxonomicName baseAuthorityName="Walsh" baseAuthorityYear="1863" box="[1285,1436,1273,1292]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="aphidicola">
<emphasis box="[1285,1436,1273,1292]" italics="true" pageId="19" pageNumber="132">Lasius aphidicola</emphasis>
</taxonomicName>
(CASENT0280468); (K) † 
<taxonomicName authorityName="Mayr" authorityYear="1868" box="[395,593,1299,1318]" class="Insecta" family="Formicidae" genus="Lasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="schiefferdeckeri">
<emphasis box="[395,593,1299,1318]" italics="true" pageId="19" pageNumber="132">Lasius schiefferdeckeri</emphasis>
</taxonomicName>
(HJF013); (L) † 
<taxonomicName authority="(AMNH-NJ 1029)" baseAuthorityName="AMNH-NJ" baseAuthorityYear="1029" box="[743,1051,1299,1318]" class="Insecta" family="Formicidae" genus="Kyromyrma" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="species" species="neffi">
<emphasis box="[743,887,1299,1318]" italics="true" pageId="19" pageNumber="132">Kyromyrma neffi</emphasis>
(AMNH-NJ1029)
</taxonomicName>
. (Image credits: D = the authors; AntWeb: A = Will Ericson; B = Erin Prado; C = Michael Branstetter; E, G, J = Shannon Hartman; F, H = April Nobile; I = unknown; K = Vincent Perrichot; L = Dave Grimaldi &amp; Vincent Perrichot.)
</paragraph>
</caption>
<paragraph blockId="19.[161,767,1423,1863]" pageId="19" pageNumber="132">
<emphasis box="[161,248,1488,1509]" italics="true" pageId="19" pageNumber="132">Note 6.</emphasis>
Short petiolar nodes are also observed in the Myrmelachistini and Plagiolepidini (excluding 
<taxonomicName authorityName="Santschi" box="[197,311,1552,1573]" class="Insecta" family="Formicidae" genus="Anoplolepis" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[197,311,1552,1573]" italics="true" pageId="19" pageNumber="132">Anoplolepis</emphasis>
</taxonomicName>
). The node height is variable in the Melophorini, being short in 
<taxonomicName authorityName="Forel" authorityYear="1892" box="[334,425,1585,1606]" class="Insecta" family="Formicidae" genus="Prolasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[334,425,1585,1606]" italics="true" pageId="19" pageNumber="132">Prolasius</emphasis>
</taxonomicName>
and 
<taxonomicName authority="Andre." authorityName="Andre" box="[471,722,1584,1606]" class="Insecta" family="Formicidae" genus="Myrmecorhynchus" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[471,649,1585,1606]" italics="true" pageId="19" pageNumber="132">Myrmecorhynchus</emphasis>
André.
</taxonomicName>
</paragraph>
<paragraph blockId="19.[161,767,1423,1863]" pageId="19" pageNumber="132">
<emphasis box="[161,230,1617,1638]" italics="true" pageId="19" pageNumber="132">Note 7.</emphasis>
The apodeme is clearly separated from the node in most Melophorini, except 
<taxonomicName authorityName="Forel" authorityYear="1892" box="[396,487,1649,1670]" class="Insecta" family="Formicidae" genus="Prolasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[396,487,1649,1670]" italics="true" pageId="19" pageNumber="132">Prolasius</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="19.[161,767,1423,1863]" pageId="19" pageNumber="132">
<emphasis box="[161,229,1681,1702]" italics="true" pageId="19" pageNumber="132">Note 8.</emphasis>
A U-shaped petiolar sternum was used by 
<bibRefCitation author="Bolton, B." box="[633,767,1681,1702]" journalOrPublisher="Memoirs of the American Entomological Institute" pageId="19" pageNumber="132" pagination="1 - 370" part="71" refId="ref31502" refString="Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370." title="Synopsis and classification of Formicidae" type="journal article" year="2003">Bolton (2003)</bibRefCitation>
to diagnose the lasiine tribe group. This trait also occurs in the Myrmelachistini, Myrmoteratini Emery, Plagiolepidini, and four Melophorini (
<taxonomicName authorityName="Emery" authorityYear="1895" box="[418,539,1778,1799]" class="Insecta" family="Formicidae" genus="Lasiophanes" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[418,539,1778,1799]" italics="true" pageId="19" pageNumber="132">Lasiophanes</emphasis>
</taxonomicName>
, 
<taxonomicName authorityName="Forel" authorityYear="1892" box="[551,642,1778,1799]" class="Insecta" family="Formicidae" genus="Prolasius" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[551,642,1778,1799]" italics="true" pageId="19" pageNumber="132">Prolasius</emphasis>
</taxonomicName>
, 
<taxonomicName authorityName="Forel" authorityYear="1902" box="[654,760,1778,1799]" class="Insecta" family="Formicidae" genus="Stigmacros" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[654,760,1778,1799]" italics="true" pageId="19" pageNumber="132">Stigmacros</emphasis>
</taxonomicName>
, 
<taxonomicName authorityName="McAreavey" authorityYear="1957" box="[197,333,1810,1831]" class="Insecta" family="Formicidae" genus="Teratomyrmex" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[197,333,1810,1831]" italics="true" pageId="19" pageNumber="132">Teratomyrmex</emphasis>
</taxonomicName>
); see ‘Ancestral state estimation’ results above and Figure S15.
</paragraph>
<paragraph blockId="19.[830,1437,1423,1687]" pageId="19" pageNumber="132">
<emphasis box="[830,900,1424,1445]" italics="true" pageId="19" pageNumber="132">Note 9.</emphasis>
The proventriculus is weakly sepalous in 
<taxonomicName authorityName="Wheeler" authorityYear="1920" box="[1305,1430,1424,1445]" class="Insecta" family="Formicidae" genus="Cladomyrma" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[1305,1430,1424,1445]" italics="true" pageId="19" pageNumber="132">Cladomyrma</emphasis>
</taxonomicName>
. The form of the proventriculus forms a natural division between the Plagiolepidini and the ‘plagiolepidiform’ 
<taxonomicName authorityName="Mayr" authorityYear="1861" box="[866,969,1515,1536]" class="Insecta" family="Formicidae" genus="Prenolepis" kingdom="Animalia" order="Hymenoptera" pageId="19" pageNumber="132" phylum="Arthropoda" rank="genus">
<emphasis box="[866,969,1515,1536]" italics="true" pageId="19" pageNumber="132">Prenolepis</emphasis>
</taxonomicName>
genus group, as observed by 
<bibRefCitation author="Emery, C." box="[1290,1433,1514,1535]" journalOrPublisher="Subfam. Formicinae. Genera Insectorum" pageId="19" pageNumber="132" pagination="1 - 302" part="183" refId="ref33484" refString="Emery, C. (1925) Hymenoptera. Fam. Formicidae. Subfam. Formicinae. Genera Insectorum, 183, 1 - 302." title="Hymenoptera. Fam. Formicidae" type="journal article" year="1925">Emery (1925)</bibRefCitation>
. Due to uncertainty about the polarity of the sepalous condition – and whether this form arose multiple times (
<bibRefCitation author="Eisner, T." box="[873,997,1605,1626]" journalOrPublisher="Bulletin of the Museum of Comparative Zoology" pageId="19" pageNumber="132" pagination="439 - 490" part="8" refId="ref33323" refString="Eisner, T. (1957) A comparative morphological study of the proventriculus of ants (Hymenoptera: Formicidae). Bulletin of the Museum of Comparative Zoology, 8, 439 - 490." title="A comparative morphological study of the proventriculus of ants (Hymenoptera: Formicidae)" type="journal article" year="1957">Eisner, 1957</bibRefCitation>
; 
<bibRefCitation author="Agosti, D." box="[1006,1131,1605,1626]" journalOrPublisher="Actes Colloques Insectes Sociaux" pageId="19" pageNumber="132" pagination="295 - 303" part="6" refId="ref30601" refString="Agosti, D. (1990) What makes the Formicini the Formicini? Actes Colloques Insectes Sociaux, 6, 295 - 303." title="What makes the Formicini the Formicini?" type="journal article" year="1990">Agosti, 1990</bibRefCitation>
, 
<bibRefCitation author="Agosti, D." box="[1141,1190,1605,1626]" journalOrPublisher="Systematic Entomology" pageId="19" pageNumber="132" pagination="293 - 310" part="16" refId="ref30626" refString="Agosti, D. (1991) Revision of the oriental ant genus Cladomyrma, with an outline of the higher classification of the Formicinae (Hymenoptera: Formicidae). Systematic Entomology, 16, 293 - 310." title="Revision of the oriental ant genus Cladomyrma, with an outline of the higher classification of the Formicinae (Hymenoptera: Formicidae)" type="journal article" year="1991">1991</bibRefCitation>
) – 
<bibRefCitation author="Bolton, B." box="[1224,1359,1605,1626]" journalOrPublisher="Memoirs of the American Entomological Institute" pageId="19" pageNumber="132" pagination="1 - 370" part="71" refId="ref31502" refString="Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1 - 370." title="Synopsis and classification of Formicidae" type="journal article" year="2003">Bolton (2003)</bibRefCitation>
lumped the two plagiolepidiform clades, plus the Myrmelachistini, into the Plagiolepidini.
</paragraph>
</subSubSection>
</treatment>
</document>