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<mods:title id="ACD1B60332EB469211E19001C3C5BA06">Penthetria Meigen (Diptera: Bibionidae): Revision of the New World species and world catalog</mods:title>
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<paragraph id="29414C13FFBF4B47FF2FFBDFFE4A2210" blockId="2.[151,466,1122,1149]" box="[151,466,1122,1149]" pageId="2" pageNumber="453">
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<emphasis id="1B8A9001FFBF4B47FF2FFBDFFE4A2210" bold="true" box="[151,466,1122,1149]" pageId="2" pageNumber="453">
Genus 
<taxonomicName id="EEFE3790FFBF4B47FF56FBDFFE4A2210" ID-CoL="8YQLD" ID-ENA="560770" authority="Meigen" authorityName="Meigen" box="[238,466,1122,1149]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF56FBDFFEF52211" bold="true" box="[238,365,1122,1149]" italics="true" pageId="2" pageNumber="453">Penthetria</emphasis>
Meigen
</taxonomicName>
</emphasis>
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</paragraph>
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<subSubSection id="61E41F98FFBF4B47FF2FFB1AFAB42022" pageId="2" pageNumber="453" type="reference_group">
<paragraph id="29414C13FFBF4B47FF2FFB1AFF6C22B2" blockId="2.[151,1437,1191,1614]" pageId="2" pageNumber="453">
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<treatmentCitation id="A85F6A02FFBF4B47FF2FFB1AFE3F22D3" author="Meigen, J. W." box="[151,423,1191,1215]" page="20" pageId="2" pageNumber="453" year="1800">
<taxonomicName id="EEFE3790FFBF4B47FF2FFB1AFE3F22D3" authority="Meigen, 1800: 20" authorityName="Meigen" authorityPageNumber="20" authorityYear="1800" box="[151,423,1191,1215]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FFB1AFF7B22D2" box="[151,227,1191,1214]" italics="true" pageId="2" pageNumber="453">Amasia</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FF54FB1AFE3F22D3" author="Meigen, J. W." box="[236,423,1191,1215]" pageId="2" pageNumber="453" refId="ref35361" refString="Meigen, J. W. (1800) Nouvelle classification des mouches a deux ailes (Diptera L.) d'apres un plan tout nouveau. Perronneau, Paris, 40 pp. https: // doi. org / 10.5962 / bhl. title. 119764" type="book" year="1800">Meigen, 1800: 20</bibRefCitation>
</taxonomicName>
</treatmentCitation>
. Unavailable name; work suppressed for purposes of zoological nomenclature by I.C.Z.N., 1963:
</treatmentCitationGroup>
339.
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FFB59FAE82291" blockId="2.[151,1437,1191,1614]" box="[151,1392,1252,1277]" pageId="2" pageNumber="453">
<taxonomicName id="EEFE3790FFBF4B47FF2FFB59FE562290" ID-CoL="8YQLD" authority="Meigen, 1803: 264" authorityName="Meigen" authorityPageNumber="264" authorityYear="1803" box="[151,462,1252,1277]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FFB59FE9D2290" bold="true" box="[151,261,1252,1276]" italics="true" pageId="2" pageNumber="453">Penthetria</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEB3FB59FE562290" author="Meigen, J. W." box="[267,462,1252,1277]" pageId="2" pageNumber="453" pagination="259 - 281" refId="ref35409" refString="Meigen, J. W. (1803) Versuch einer neuen GattungsEintheilung der europaischen zweiflugligen Insekten. Magazin fur Insecktenkunde, 2, 259 - 281." type="journal article" year="1803">Meigen, 1803: 264</bibRefCitation>
</taxonomicName>
. 
<typeStatus id="F645F2B1FFBF4B47FE60FB58FD922291" box="[472,522,1253,1277]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FDDFFB59FC692291" ID-CoL="6V23R" authority="Meigen, 1804: 104" authorityName="Meigen" authorityPageNumber="104" authorityYear="1804" box="[615,1009,1252,1277]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFBF4B47FDDFFB59FCB32290" box="[615,811,1252,1276]" italics="true" pageId="2" pageNumber="453">Penthetria funebris</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FC89FB59FC692291" author="Meigen, J. W." box="[817,1009,1252,1277]" pageId="2" pageNumber="453" pagination="153 - 314" refId="ref35437" refString="Meigen, J. W. (1804) Klassifikazion und Beschreibung der europaischen zweiflugligen Insekten (Diptera Linn.). Erster Band. Abt. I &amp; II. Reichard, Braunschweig [Brunswick], xxvii + vi + 314 pp. [pp. i-xxvii + 1 - 152 pp. &amp; pp. i-vi + 153 - 314]" type="book chapter" year="1804">Meigen, 1804: 104</bibRefCitation>
</taxonomicName>
(by monotypy in 
<bibRefCitation id="4D6F31E2FFBF4B47FB1FFB59FAFD2291" author="Meigen, J. W." box="[1191,1381,1252,1277]" pageId="2" pageNumber="453" pagination="153 - 314" refId="ref35437" refString="Meigen, J. W. (1804) Klassifikazion und Beschreibung der europaischen zweiflugligen Insekten (Diptera Linn.). Erster Band. Abt. I &amp; II. Reichard, Braunschweig [Brunswick], xxvii + vi + 314 pp. [pp. i-xxvii + 1 - 152 pp. &amp; pp. i-vi + 153 - 314]" type="book chapter" year="1804">Meigen 1804: 104</bibRefCitation>
).
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FFABEFD0A2377" blockId="2.[151,1437,1191,1614]" box="[151,658,1283,1307]" pageId="2" pageNumber="453">
<taxonomicName id="EEFE3790FFBF4B47FF2FFABEFE772377" authority="Wiedemann, 1830: 618" authorityName="Wiedemann" authorityPageNumber="618" authorityYear="1830" box="[151,495,1283,1307]" class="Insecta" genus="Threneste" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FFABEFF622377" box="[151,250,1283,1307]" italics="true" pageId="2" pageNumber="453">Threneste</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEB8FABEFE772377" author="Wiedemann, C. R. W." box="[256,495,1283,1307]" pageId="2" pageNumber="453" refId="ref37110" refString="Wiedemann, C. R. W. (1830) Aussereuropaische zweiflugelige Insekten. Als Fortsetzung des Meigenschen Werkes. Zweiter Theil. Schulz, Hamm, xii + 684 pp." type="book" year="1830">Wiedemann, 1830: 618</bibRefCitation>
</taxonomicName>
. 
<emphasis id="1B8A9001FFBF4B47FE41FAB9FD142377" box="[505,652,1283,1307]" italics="true" pageId="2" pageNumber="453">Nomen nudum</emphasis>
.
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FFA9FFEBB2334" blockId="2.[151,1437,1191,1614]" pageId="2" pageNumber="453">
<taxonomicName id="EEFE3790FFBF4B47FF2FFA9FFE4C2356" authority="Macquart, 1838: 88" authorityName="Macquart" authorityPageNumber="88" authorityYear="1838" box="[151,468,1314,1338]" class="Insecta" genus="Eupeitenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FFA9FFE902355" box="[151,264,1314,1337]" italics="true" pageId="2" pageNumber="453">Eupeitenus</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEB6FA9FFE4C2356" author="Macquart, J." box="[270,468,1314,1338]" pageId="2" pageNumber="453" pagination="9 - 225" refId="ref35224" refString="Macquart, J. (1838) Dipteres exotiques nouveux ou peu connus. Societe (Royale) des sciences de l'agriculture et des arts a Lille, Memoires, 1838 (2), 9 - 225, 25 pls. [also published separately as his &quot; Dipteres exotiques nouveux ou peu connus, &quot; Vol. 1. Pt. 1. pp. 5 - 221, 25 pls. (Paris 1838)] https: // doi. org / 10.5962 / bhl. title. 51482" type="journal article" year="1838">Macquart, 1838: 88</bibRefCitation>
</taxonomicName>
(also 1838: 84). 
<typeStatus id="F645F2B1FFBF4B47FDC6FA9FFD282356" box="[638,688,1314,1338]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FCB5FA9FFB192356" ID-CoL="6V242" authority="Macquart, 1834: 175" authorityName="Macquart" authorityPageNumber="175" authorityYear="1834" box="[781,1153,1314,1338]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="atra">
<emphasis id="1B8A9001FFBF4B47FCB5FA9FFC302356" box="[781,936,1314,1338]" italics="true" pageId="2" pageNumber="453">Penthetria atra</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FC16FA9FFB192356" author="Macquart, J." box="[942,1153,1314,1338]" pageId="2" pageNumber="453" refId="ref35177" refString="Macquart, J. (1834) Histoire naturelle des insectes. Dipteres. Ouvrage accompagne de planches. Tome Premier. N. E. Roret, Paris, 578 pp. https: // doi. org / 10.5962 / bhl. title. 14274" type="book" year="1834">Macquart, 1834: 175</bibRefCitation>
</taxonomicName>
(by monotypy), [examined; BMNH].
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FFAE2FA9A231B" blockId="2.[151,1437,1191,1614]" box="[151,1282,1375,1400]" pageId="2" pageNumber="453">
<taxonomicName id="EEFE3790FFBF4B47FF2FFAE2FE68231B" authority="Gimmerthal, 1845: 330" authorityName="Gimmerthal" authorityPageNumber="330" authorityYear="1845" box="[151,496,1375,1399]" class="Insecta" family="Bibionidae" genus="Crapitula" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FFAE2FF62231B" box="[151,250,1375,1399]" italics="true" pageId="2" pageNumber="453">Crapitula</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEB8FAE2FE68231B" author="Gimmerthal, B. A." box="[256,496,1375,1399]" pageId="2" pageNumber="453" pagination="287 - 331" refId="ref33678" refString="Gimmerthal, B. A. (1845) Erster Beitrag zu einer Kunftig zu bearbeitenden Dipterologie Russlands. Familie der Tipularien. Bul- letin de la Societe Imperial des Naturalistes, Moscou, 18 (4), 287 - 331. https: // doi. org / 10.5962 / bhl. title. 66007" type="journal article" year="1845">Gimmerthal, 1845: 330</bibRefCitation>
</taxonomicName>
. 
<typeStatus id="F645F2B1FFBF4B47FE41FADDFDB32314" box="[505,555,1376,1400]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FD31FAE2FBC5231B" ID-CoL="8BLHX" authority="Gimmerthal 1845: 330" authorityName="Gimmerthal" authorityPageNumber="330" authorityYear="1845" box="[649,1117,1375,1399]" class="Insecta" family="Bibionidae" genus="Crapitula" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="motschulskii">
<emphasis id="1B8A9001FFBF4B47FD31FAE2FCE8231B" box="[649,880,1375,1399]" italics="true" pageId="2" pageNumber="453">Crapitula motschulskii</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FCCEFAE2FBC5231B" author="Gimmerthal, B. A." box="[886,1117,1375,1399]" pageId="2" pageNumber="453" pagination="287 - 331" refId="ref33678" refString="Gimmerthal, B. A. (1845) Erster Beitrag zu einer Kunftig zu bearbeitenden Dipterologie Russlands. Familie der Tipularien. Bul- letin de la Societe Imperial des Naturalistes, Moscou, 18 (4), 287 - 331. https: // doi. org / 10.5962 / bhl. title. 66007" type="journal article" year="1845">Gimmerthal 1845: 330</bibRefCitation>
</taxonomicName>
(by monotypy).
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FFAC3FADA23FA" blockId="2.[151,1437,1191,1614]" box="[151,1346,1406,1430]" pageId="2" pageNumber="453">
<emphasis id="1B8A9001FFBF4B47FF2FFAC3FF3923FA" box="[151,161,1406,1430]" italics="true" pageId="2" pageNumber="453">*</emphasis>
<taxonomicName id="EEFE3790FFBF4B47FF19FAC3FE3623FA" authority="Heer, 1849: 228" authorityName="Heer" authorityPageNumber="228" authorityYear="1849" box="[161,430,1406,1430]" class="Insecta" family="Bibionidae" genus="Bibiopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF19FAC3FE9A23FA" box="[161,258,1406,1430]" italics="true" pageId="2" pageNumber="453">Bibiopsis</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEB0FAC3FE3623FA" author="Heer, O." box="[264,430,1406,1430]" pageId="2" pageNumber="453" refId="ref34441" refString="Heer, O. (1849) Die Insektenfauna der Tertiargebilde von Oeningen und von Radoboj in Croatien. Zweite Theil: Heuschrecken, Florfliegen, Aderflugler, Schmetterlinge und Fliegen. W. Engelmann, Leipzig, iv + 264 pp." type="book" year="1849">Heer, 1849: 228</bibRefCitation>
</taxonomicName>
. 
<typeStatus id="F645F2B1FFBF4B47FE00FAC3FE7223FA" box="[440,490,1406,1430]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FDFFFAC3FC5923FA" ID-CoL="8B77W" authority="Heer, 1849: 229" authorityName="Heer" authorityPageNumber="229" authorityYear="1849" box="[583,961,1406,1430]" class="Insecta" family="Bibionidae" genus="Bibiopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="cimicoides">
<emphasis id="1B8A9001FFBF4B47FDFFFAC3FC8F23FA" box="[583,791,1406,1430]" italics="true" pageId="2" pageNumber="453">Bibiopsis cimicoides</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FCA6FAC3FC5923FA" author="Heer, O." box="[798,961,1406,1430]" pageId="2" pageNumber="453" refId="ref34441" refString="Heer, O. (1849) Die Insektenfauna der Tertiargebilde von Oeningen und von Radoboj in Croatien. Zweite Theil: Heuschrecken, Florfliegen, Aderflugler, Schmetterlinge und Fliegen. W. Engelmann, Leipzig, iv + 264 pp." type="book" year="1849">Heer, 1849: 229</bibRefCitation>
</taxonomicName>
(designated by 
<bibRefCitation id="4D6F31E2FFBF4B47FBDAFAC3FAAE23FA" author="Carpenter, F. M." box="[1122,1334,1406,1430]" pageId="2" pageNumber="453" pagination="279 - 655" refId="ref32431" refString="Carpenter, F. M. (1992) Vol. 4. Superclass Hexapoda. In: Treatise on Invertebrate Paleontology. Part R. Arthropoda 4. Vol. 2. Geological Society of America, Boulder, Colorado and University of Kansas, Lawrence, Kansas, pp. i-ii + 279 - 655." type="book chapter" year="1992">Carpenter 1992: 414</bibRefCitation>
).
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FFA21FD1123BF" blockId="2.[151,1437,1191,1614]" pageId="2" pageNumber="453">
<emphasis id="1B8A9001FFBF4B47FF2FFA21FF3923D8" box="[151,161,1436,1460]" italics="true" pageId="2" pageNumber="453">*</emphasis>
<taxonomicName id="EEFE3790FFBF4B47FF19FA21FD9B23D9" authority="Scudder, 1892: 20" authorityName="Scudder" authorityPageNumber="20" authorityYear="1892" box="[161,515,1436,1461]" class="Insecta" family="Mycetophilidae" genus="Mycetophaetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF19FA21FEA523D8" box="[161,317,1436,1460]" italics="true" pageId="2" pageNumber="453">Mycetophaetus</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEFDFA21FD9B23D9" author="Scudder, S. H." box="[325,515,1436,1461]" pageId="2" pageNumber="453" pagination="11 - 25" refId="ref36252" refString="Scudder, S. H. (1892) Some insects of special interest from Florissant, Colorado and other points in the tertiaries of Colorado and Utah. Bulletin of the United States Geological Survey, 93, 11 - 25." type="journal article" year="1892">Scudder, 1892: 20</bibRefCitation>
</taxonomicName>
. 
<typeStatus id="F645F2B1FFBF4B47FDB6FA20FDD823D9" box="[526,576,1437,1461]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FD18FA21FBE323D9" authority="Scudder, 1892: 20" authorityName="Scudder" authorityPageNumber="20" authorityYear="1892" box="[672,1147,1436,1461]" class="Insecta" family="Mycetophilidae" genus="Mycetophaetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="intermedius">
<emphasis id="1B8A9001FFBF4B47FD18FA21FC2123D8" box="[672,953,1436,1460]" italics="true" pageId="2" pageNumber="453">Mycetophaetus intermedius</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FC78FA21FBE323D9" author="Scudder, S. H." box="[960,1147,1436,1461]" pageId="2" pageNumber="453" pagination="11 - 25" refId="ref36252" refString="Scudder, S. H. (1892) Some insects of special interest from Florissant, Colorado and other points in the tertiaries of Colorado and Utah. Bulletin of the United States Geological Survey, 93, 11 - 25." type="journal article" year="1892">Scudder, 1892: 20</bibRefCitation>
</taxonomicName>
(by monotypy), [examined; MCZC]. [synonymized by 
<bibRefCitation id="4D6F31E2FFBF4B47FE62FA06FDE723BF" author="Fitzgerald, S. J." box="[474,639,1467,1491]" pageId="2" pageNumber="453" pagination="182 - 187" refId="ref33069" refString="Fitzgerald, S. J. (1999) A new species of Plecia from the Green River Formation and new combinations of fossil Bibionidae (Diptera). Great Basin Naturalist, 59 (2), 182 - 187. https: // doi. org / 10.5962 / bhl. part. 15821" type="journal article" year="1999">Fitzgerald 1999</bibRefCitation>
].
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FFA67FC30207D" blockId="2.[151,1437,1191,1614]" pageId="2" pageNumber="453">
<taxonomicName id="EEFE3790FFBF4B47FF2FFA67FEA0239E" authority="Meigen" authorityName="Meigen" authorityYear="1800" box="[151,312,1498,1522]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FFA67FF7B239D" box="[151,227,1498,1521]" italics="true" pageId="2" pageNumber="453">Amasia</emphasis>
Meigen
</taxonomicName>
<emphasis id="1B8A9001FFBF4B47FEF8FA67FECC239D" box="[320,340,1498,1521]" italics="true" pageId="2" pageNumber="453">in</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEE3FA67FD88239E" author="Hendel, F." box="[347,528,1498,1522]" pageId="2" pageNumber="453" pagination="43 - 69" refId="ref34371" refString="Hendel, F. (1908) Nouvelle classification des mouches a deux ailes (Diptera L.). D'apres un plan tout nouveau par J. G. Meigen, Paris, an VIII (1800 v. s.). Mit Kommentar herausgegeben von Friedrich Hendel. Verhandlungen der K. - k. Zoologisch- Botanischen Gesellschaft in Wien, 58, 43 - 69." type="journal article" year="1908">Hendel, 1908: 50</bibRefCitation>
. 
<typeStatus id="F645F2B1FFBF4B47FDA2FA67FDD4239E" box="[538,588,1498,1522]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FD14FA67FC5E239E" ID-CoL="6V23R" authority="Meigen" authorityName="Meigen" authorityYear="1804" box="[684,966,1498,1522]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFBF4B47FD14FA67FCEA239E" box="[684,882,1498,1522]" italics="true" pageId="2" pageNumber="453">Penthetria funebris</emphasis>
Meigen
</taxonomicName>
as “ 
<taxonomicName id="EEFE3790FFBF4B47FC4EFA67FB6D239E" authority="(Meig.)" baseAuthorityName="Meig." box="[1014,1269,1498,1522]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFBF4B47FC4EFA67FB05239E" box="[1014,1181,1498,1522]" italics="true" pageId="2" pageNumber="453">Amasia funebris</emphasis>
(Meig.)
</taxonomicName>
,” by subsequent monotypy in 
<bibRefCitation id="4D6F31E2FFBF4B47FEF6FA45FE68207D" author="Bezzi, M." box="[334,496,1528,1553]" pageId="2" pageNumber="453" pagination="1 - 87" refId="ref32244" refString="Bezzi, M. (1911) Biospeologica. XX. Dipteres (premiere serie) suivi d'un appendice sur les dipteres cavernicoles recueillis par le Dr Absolon dans les Balcans. Archives de Zoologie Experimentale et Generale, Seris 5, 8, 1 - 87." type="journal article" year="1911">Bezzi, 1911: 39</bibRefCitation>
. [synonymized by 
<bibRefCitation id="4D6F31E2FFBF4B47FD17FA45FC06207D" author="Evenhuis, N. L. &amp; Pape, T." box="[687,926,1528,1553]" pageId="2" pageNumber="453" pagination="1 - 74" refId="ref32919" refString="Evenhuis, N. L. &amp; Pape, T. (2017) Battling the un-dead: the status of the Diptera genus-group names originally proposed in Johann Wilhelm Meigen's 1800 pamphlet. Zootaxa, 4275 (1), 1 - 74. https: // doi. org / 10.11646 / zootaxa. 4275.1.1" type="journal article" year="2017">Evenhuis &amp; Pape 2017</bibRefCitation>
].
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FF9AAFB452043" blockId="2.[151,1437,1191,1614]" box="[151,1245,1559,1584]" pageId="2" pageNumber="453">
<taxonomicName id="EEFE3790FFBF4B47FF2FF9AAFE4F2043" authority="Brunetti, 1911: 269" authorityName="Brunetti" authorityPageNumber="269" authorityYear="1911" box="[151,471,1559,1583]" class="Insecta" family="Bibionidae" genus="Pleciomyia" higherTaxonomySource="CoL" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FF9AAFE912043" box="[151,265,1559,1583]" italics="true" pageId="2" pageNumber="453">Pleciomyia</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FEB7F9AAFE4F2043" author="Brunetti, E." box="[271,471,1559,1583]" pageId="2" pageNumber="453" pagination="259 - 316" refId="ref32320" refString="Brunetti, E. (1911) New Oriental Nematocera. Records of the Indian Museum, 4, 259 - 316. https: // doi. org / 10.5962 / bhl. part. 10539" type="journal article" year="1911">Brunetti, 1911: 269</bibRefCitation>
</taxonomicName>
. 
<typeStatus id="F645F2B1FFBF4B47FE58F9A5FD8A205C" box="[480,530,1560,1584]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FDC8F9AAFBA02043" ID-CoL="76PRS" authority="Wiedemann, 1828: 72" authorityName="Wiedemann" authorityPageNumber="72" authorityYear="1828" box="[624,1080,1559,1583]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="melanaspis">
<emphasis id="1B8A9001FFBF4B47FDC8F9AAFCCB2043" box="[624,851,1559,1583]" italics="true" pageId="2" pageNumber="453">Penthetria melanaspis</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FCE1F9AAFBA02043" author="Wiedemann, C. R. W." box="[857,1080,1559,1583]" pageId="2" pageNumber="453" refId="ref37077" refString="Wiedemann, C. R. W. (1828) Aussereuropaische zweiflugelige Insekten. Als Fortsetzung des Meigenschen Werkes. Erster Theil. Schulz, Hamm, xxxii + 608 pp." type="book" year="1828">Wiedemann, 1828: 72</bibRefCitation>
</taxonomicName>
(by monotypy).
</paragraph>
<paragraph id="29414C13FFBF4B47FF2FF98BFAB42022" blockId="2.[151,1437,1191,1614]" box="[151,1324,1590,1614]" pageId="2" pageNumber="453">
<taxonomicName id="EEFE3790FFBF4B47FF2FF98BFD902022" authority="Brunetti, 1912: 446" authorityName="Brunetti" authorityPageNumber="446" authorityYear="1912" box="[151,520,1590,1614]" class="Insecta" family="Bibionidae" genus="Parapleciomyia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FF98BFEA12022" box="[151,313,1590,1614]" italics="true" pageId="2" pageNumber="453">Parapleciomyia</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FE87F98BFD902022" author="Brunetti, E." box="[319,520,1590,1614]" pageId="2" pageNumber="453" pagination="445 - 513" refId="ref32357" refString="Brunetti, E. (1912) XXXVI. New Oriental Diptera, I. Records of the Indian Museum, 7, 445 - 513. https: // doi. org / 10.5962 / bhl. part. 28244" type="journal article" year="1912">Brunetti, 1912: 446</bibRefCitation>
</taxonomicName>
. 
<typeStatus id="F645F2B1FFBF4B47FDA9F98BFDDB2022" box="[529,579,1590,1614]" pageId="2" pageNumber="453">Type</typeStatus>
species: 
<taxonomicName id="EEFE3790FFBF4B47FD18F98BFB1F2022" ID-CoL="8D72W" authority="Brunetti, 1912: 447" authorityName="Brunetti" authorityPageNumber="447" authorityYear="1912" box="[672,1159,1590,1614]" class="Insecta" family="Bibionidae" genus="Parapleciomyia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="species" species="carbonaria">
<emphasis id="1B8A9001FFBF4B47FD18F98BFC232022" box="[672,955,1590,1614]" italics="true" pageId="2" pageNumber="453">Parapleciomyia carbonaria</emphasis>
<bibRefCitation id="4D6F31E2FFBF4B47FC79F98BFB1F2022" author="Brunetti, E." box="[961,1159,1590,1614]" pageId="2" pageNumber="453" pagination="445 - 513" refId="ref32357" refString="Brunetti, E. (1912) XXXVI. New Oriental Diptera, I. Records of the Indian Museum, 7, 445 - 513. https: // doi. org / 10.5962 / bhl. part. 28244" type="journal article" year="1912">Brunetti, 1912: 447</bibRefCitation>
</taxonomicName>
(by monotypy).
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFBF4B47FF2FF9C1FDED2127" pageId="2" pageNumber="453" type="discussion">
<paragraph id="29414C13FFBF4B47FF2FF9C1FDED2127" blockId="2.[151,1436,1660,2011]" pageId="2" pageNumber="453">
<emphasis id="1B8A9001FFBF4B47FF2FF9C1FE0920FA" bold="true" box="[151,401,1660,1686]" pageId="2" pageNumber="453">Nomenclatural notes</emphasis>
. The genus 
<taxonomicName id="EEFE3790FFBF4B47FDA7F9C1FD5020FB" authority="Heer" authorityName="Heer" authorityYear="1849" box="[543,712,1660,1687]" class="Insecta" family="Bibionidae" genus="Bibiopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FDA7F9C1FD1F20FA" box="[543,647,1660,1686]" italics="true" pageId="2" pageNumber="453">Bibiopsis</emphasis>
Heer
</taxonomicName>
, based on a fossil, has historically been treated as a junior synonym of 
<taxonomicName id="EEFE3790FFBF4B47FF42F91DFD9120D7" authority="(Loew 1868)" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[250,521,1696,1723]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF42F91DFEE820D6" box="[250,368,1696,1722]" italics="true" pageId="2" pageNumber="453">Penthetria</emphasis>
(
<bibRefCitation id="4D6F31E2FFBF4B47FEC6F91DFD9920D7" author="Loew, H." box="[382,513,1696,1723]" pageId="2" pageNumber="453" pagination="180 - 191" refId="ref35111" refString="Loew, H. (1868) Berichtigung der generischen Bestimmung einiger fossile Dipteren. Taf. V. Zeitschrift fur die Gesammten Naturwissenschaften, 230, 180 - 191." type="journal article" year="1868">Loew 1868</bibRefCitation>
)
</taxonomicName>
, as a distinct genus (
<bibRefCitation id="4D6F31E2FFBF4B47FD54F91DFC1220D6" author="Scudder, S. H." box="[748,906,1696,1722]" pageId="2" pageNumber="453" pagination="1 - 744" refId="ref36200" refString="Scudder, S. H. (1891) Index to the known fossil insects of the world, including myriapods and arachnids. Bulletin of the United States Geological Survey, 71, 1 - 744. https: // doi. org / 10.5962 / bhl. title. 34455" type="journal article" year="1891">Scudder 1891</bibRefCitation>
), and as a junior synomym of a broadly-defined 
<taxonomicName id="EEFE3790FFBF4B47FF2FF979FE9520B2" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[151,269,1732,1758]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF2FF979FE9520B2" box="[151,269,1732,1758]" italics="true" pageId="2" pageNumber="453">Penthetria</emphasis>
</taxonomicName>
(including 
<taxonomicName id="EEFE3790FFBF4B47FE2BF979FDC420B2" authority="Wiedeman" authorityName="Wiedeman" box="[403,604,1732,1758]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FE2BF979FE4320B2" box="[403,475,1732,1758]" italics="true" pageId="2" pageNumber="453">Plecia</emphasis>
Wiedeman
</taxonomicName>
and 
<taxonomicName id="EEFE3790FFBF4B47FD2FF978FCC820B3" authority="Heer" authorityName="Heer" authorityYear="1849" box="[663,848,1733,1759]" class="Insecta" family="Bibionidae" genus="Protomyia" higherTaxonomySource="CoL" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FD2FF978FC9420B2" box="[663,780,1733,1758]" italics="true" pageId="2" pageNumber="453">Protomyia</emphasis>
Heer
</taxonomicName>
; 
<bibRefCitation id="4D6F31E2FFBF4B47FCE5F979FB8620B2" author="Handlirsch, A." box="[861,1054,1732,1758]" pageId="2" pageNumber="453" pagination="801 - 960" refId="ref33735" refString="Handlirsch, A. (1907) Die fossilen Insekten und die Phylogenie der rezenten Formen. Ein Handbuch fur Palaontologen und Zoologen. Engelmann, Leipzig, pp. 801 - 960. https: // doi. org / 10.5962 / bhl. title. 5636" type="book chapter" year="1907">Handlirsch 1907</bibRefCitation>
). More recently, 
<bibRefCitation id="4D6F31E2FFBF4B47FB5AF979FA0020B2" author="Evenhuis, N. L." box="[1250,1432,1732,1758]" pageId="2" pageNumber="453" refId="ref32810" refString="Evenhuis, N. L. (1994) Catalog of the fossil flies of the world (Insecta: Diptera). Backhuys Publishers, Leiden, 600 pp." type="book" year="1994">Evenhuis (1994</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFBF4B47FF2FF955FF49216E" author="Evenhuis, N. L." box="[151,209,1768,1794]" pageId="2" pageNumber="453" refId="ref32873" refString="Evenhuis, N. L. (2014) Catalog of the fossil flies of the world (Insecta: Diptera). Version 2.0. Available from: http: // hbs. bishopmuseum. org / fossilcat / (accessed 30 March 2020)" type="url" year="2014">2014</bibRefCitation>
) treated 
<taxonomicName id="EEFE3790FFBF4B47FE8BF955FE03216E" authorityName="Heer" authorityYear="1849" box="[307,411,1768,1794]" class="Insecta" family="Bibionidae" genus="Bibiopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FE8BF955FE03216E" box="[307,411,1768,1794]" italics="true" pageId="2" pageNumber="453">Bibiopsis</emphasis>
</taxonomicName>
as a junior synonym of 
<taxonomicName id="EEFE3790FFBF4B47FD14F955FD6C216E" box="[684,756,1768,1794]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FD14F955FD6C216E" box="[684,756,1768,1794]" italics="true" pageId="2" pageNumber="453">Plecia</emphasis>
</taxonomicName>
. However, 
<bibRefCitation id="4D6F31E2FFBF4B47FCCDF955FB93216E" author="Heer, O." box="[885,1035,1768,1794]" pageId="2" pageNumber="453" refId="ref34441" refString="Heer, O. (1849) Die Insektenfauna der Tertiargebilde von Oeningen und von Radoboj in Croatien. Zweite Theil: Heuschrecken, Florfliegen, Aderflugler, Schmetterlinge und Fliegen. W. Engelmann, Leipzig, iv + 264 pp." type="book" year="1849">Heer’s (1849</bibRefCitation>
; Plate XV, 
<figureCitation id="B1C55096FFBF4B47FB32F955FB70216F" box="[1162,1256,1768,1795]" captionStart="FIGURES 24–29" captionStartId="27.[151,264,1849,1873]" captionTargetBox="[191,1399,188,1815]" captionTargetId="figure-16@27.[186,1402,181,1825]" captionTargetPageId="27" captionText="FIGURES 24–29. P. mexicana, male terminalia. 24–27. Topotypic (Guadalajara, Mexico). 24. Dorsal. 25. Ventral. 26. Dorsal, tergite nine removed. 27. Posterior. 28–29. Puerta Parada, Guatemala. 28. Dorsal, tergite nine removed. 29. Dorsal, tergite nine removed. Scale bar = ca. 0.5 mm. Abbreviations: c, cerci; dlp, divergent lobes of paramere; ea, ejaculatory apodeme; ed, ejaculatory duct; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vml, ventromedian lobes of gonocoxite." figureDoi="http://doi.org/10.5281/zenodo.4529544" httpUri="https://zenodo.org/record/4529544/files/figure.png" pageId="2" pageNumber="453">Fig. 24b</figureCitation>
) illustration depicts a stout-bodied fly with R 
<subScript id="B57A4E56FFBF4B47FE53F8A3FD9C2140" attach="left" box="[491,516,1822,1836]" fontSize="6" pageId="2" pageNumber="453">2+3</subScript>
elongate and subparallel to R 
<subScript id="B57A4E56FFBF4B47FCF7F8A3FCF12140" attach="left" box="[847,873,1822,1836]" fontSize="6" pageId="2" pageNumber="453">4+5</subScript>
, rather than short and angled, which indicates that this taxon belongs to the genus 
<taxonomicName id="EEFE3790FFBF4B47FE40F88DFDF62126" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[504,622,1840,1866]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FE40F88DFDF62126" box="[504,622,1840,1866]" italics="true" pageId="2" pageNumber="453">Penthetria</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFBF4B46FF7FF8E9FE9C272D" lastPageId="3" lastPageNumber="454" pageId="2" pageNumber="453" type="diagnosis">
<paragraph id="29414C13FFBF4B47FF7FF8E9FCC921DB" blockId="2.[151,1436,1660,2011]" pageId="2" pageNumber="453">
<emphasis id="1B8A9001FFBF4B47FF7FF8E9FE392102" bold="true" box="[199,417,1876,1902]" pageId="2" pageNumber="453">Generic Diagnosis</emphasis>
. The following combination of characters will distinguish adults of extant members of the genus 
<taxonomicName id="EEFE3790FFBF4B47FF5AF8C5FEC021FE" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[226,344,1912,1938]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FF5AF8C5FEC021FE" box="[226,344,1912,1938]" italics="true" pageId="2" pageNumber="453">Penthetria</emphasis>
</taxonomicName>
from other 
<taxonomicName id="EEFE3790FFBF4B47FE5EF8C5FDFD21FE" baseAuthorityName="Fitzgerald" baseAuthorityYear="2004" box="[486,613,1912,1938]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="family">Bibionidae</taxonomicName>
: Fore tibia simple (without apical mucron or spines), R 
<subScript id="B57A4E56FFBF4B47FB50F837FA9921F4" attach="left" box="[1256,1281,1930,1944]" fontSize="6" pageId="2" pageNumber="453">2+3</subScript>
elongate and subparallel to R 
<subScript id="B57A4E56FFBF4B47FEF0F813FEFA21D0" attach="left" box="[328,354,1966,1980]" fontSize="6" pageId="2" pageNumber="453">4+5</subScript>
, and antennal flagellomeres short, compact.
</paragraph>
<paragraph id="29414C13FFBF4B46FF7FF87DFE9C272D" blockId="2.[151,1436,1660,2011]" lastBlockId="3.[151,1437,151,2033]" lastPageId="3" lastPageNumber="454" pageId="2" pageNumber="453">
The larva of 
<taxonomicName id="EEFE3790FFBF4B47FEE1F87DFE5721B6" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[345,463,1984,2010]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="453" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBF4B47FEE1F87DFE5721B6" box="[345,463,1984,2010]" italics="true" pageId="2" pageNumber="453">Penthetria</emphasis>
</taxonomicName>
(
<figureCitation id="B1C55096FFBF4B47FE67F87DFDA821B7" box="[479,560,1984,2011]" captionStart="FIGURES 58–60" captionStartId="38.[151,264,1892,1916]" captionTargetBox="[161,1427,186,1866]" captionTargetId="figure-34@38.[160,1428,181,1868]" captionTargetPageId="38" captionText="FIGURES 58–60. P. funebris. 58. Larva, lateral view (head, at left, not visible; telescoped within thorax). 59. Pupa, dorsal. 60. Pupa, ventral. Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529568" httpUri="https://zenodo.org/record/4529568/files/figure.png" pageId="2" pageNumber="453">Fig. 58</figureCitation>
) can be distinguished from the larva of other bibionids by the combination of the following characters: presence of fleshy tubercles (each abdominal segment with two transverse rows of two tubercles each), posterior spiracle on segment eight, posterior spiracle with a single ecdysial scar, and the mentum not fused to the anterior margin of the cranium or to the posterior margin of the labium (Fitzgerald 2004, Fig. 36b; pm). Pupae (
<figureCitation id="B1C55096FFBE4B46FE9BFEBEFE302771" box="[291,424,258,285]" captionStart="FIGURES 58–60" captionStartId="38.[151,264,1892,1916]" captionTargetBox="[161,1427,186,1866]" captionTargetId="figure-34@38.[160,1428,181,1868]" captionTargetPageId="38" captionText="FIGURES 58–60. P. funebris. 58. Larva, lateral view (head, at left, not visible; telescoped within thorax). 59. Pupa, dorsal. 60. Pupa, ventral. Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529568" httpUri="https://zenodo.org/record/4529568/files/figure.png" pageId="3" pageNumber="454">Figs. 59–60</figureCitation>
) can be distinguished from all other bibionid genera except 
<taxonomicName id="EEFE3790FFBE4B46FB85FEBEFB1D2771" box="[1085,1157,259,285]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FB85FEBEFB1D2771" box="[1085,1157,259,285]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
by the presence of fleshy tubercles.
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFBE4B40FF7FFEF4FB0824C5" lastPageId="5" lastPageNumber="456" pageId="3" pageNumber="454" type="discussion">
<paragraph id="29414C13FFBE4B46FF7FFEF4FDE7247E" blockId="3.[151,1437,151,2033]" pageId="3" pageNumber="454">
<emphasis id="1B8A9001FFBE4B46FF7FFEF4FD3A270F" bold="true" box="[199,674,329,355]" pageId="3" pageNumber="454">Discussion supporting generic diagnosis</emphasis>
. The generic diagnosis presented above is based on previous phylogenetic and descriptive work on the family 
<taxonomicName id="EEFE3790FFBE4B46FD3CFED1FC5427EB" authority="(Fitzgerald 2004)" baseAuthorityName="Fitzgerald" baseAuthorityYear="2004" box="[644,972,364,391]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="family">Bibionidae (Fitzgerald 2004)</taxonomicName>
and readers are referred there for further information concerning material examined or the broader context from which the diagnosis was developed. The diagnosis is designed for extant species, but reference to problems and exceptions when diagnosing fossil species are provided below. Additionally, characters not necessarily considered diagnostic, but that may be useful in helping to distinguish the genus, are also discussed.
</paragraph>
<paragraph id="29414C13FFBE4B46FF7FFDA6FE8225B4" blockId="3.[151,1437,151,2033]" pageId="3" pageNumber="454">
The relatively large adults of 
<taxonomicName id="EEFE3790FFBE4B46FDB2FDA6FD182459" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[522,640,539,565]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FDB2FDA6FD182459" box="[522,640,539,565]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
are usually matte-black, with dark wings. They are sometimes confused with some of the large species of tropical 
<taxonomicName id="EEFE3790FFBE4B46FDD1FD83FD4C2434" authorityName="Billberg" authorityYear="1820" box="[617,724,574,600]" class="Insecta" family="Sciaridae" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="family">Sciaridae</taxonomicName>
, but are easily distinguished from them by the absence of a narrow eye bridge and the presence of vein R 
<subScript id="B57A4E56FFBE4B46FDCBFDCEFD1424ED" attach="left" box="[627,652,627,641]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
: in 
<taxonomicName id="EEFE3790FFBE4B46FD01FDDCFCB72417" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[697,815,609,635]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FD01FDDCFCB72417" box="[697,815,609,635]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
males are holoptic and females broadly dichoptic (
<figureCitation id="B1C55096FFBE4B46FAD7FDDCFF3B24F2" captionStart="FIGURE 1" captionStartId="4.[151,250,1982,2006]" captionTargetBox="[331,1264,760,1945]" captionTargetId="figure-296@4.[309,1278,741,1959]" captionTargetPageId="4" captionText="FIGURE 1. P. heteroptera, habitus, male (above) and female (below), 28 October, North Carolina, USA, Matt Bertone. Images courtesy of Matt Bertone (© Matt Bertone 2013)." figureDoi="http://doi.org/10.5281/zenodo.4529514" httpUri="https://zenodo.org/record/4529514/files/figure.png" pageId="3" pageNumber="454">Fig. 1</figureCitation>
) (though the odd Palearctic species, 
<taxonomicName id="EEFE3790FFBE4B46FDF2FD38FCBB24F2" authority="Meigen" authorityName="Meigen" box="[586,803,644,670]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFBE4B46FDF2FD38FD5B24F2" box="[586,707,644,670]" italics="true" pageId="3" pageNumber="454">P. funebris</emphasis>
Meigen
</taxonomicName>
, has males with eyes that do not quite meet dorsomedially). Adults of 
<taxonomicName id="EEFE3790FFBE4B46FEFFFD1AFE2524AD" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[327,445,679,705]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FEFFFD1AFE2524AD" box="[327,445,679,705]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
are distinguished from other 
<taxonomicName id="EEFE3790FFBE4B46FCB0FD1AFC1B24AD" baseAuthorityName="Fitzgerald" baseAuthorityYear="2004" box="[776,899,679,705]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="family">Bibionidae</taxonomicName>
by the apically simple fore tibia (
<taxonomicName id="EEFE3790FFBE4B46FB44FD1AFAE124AD" authorityName="Newman" authorityYear="1834" box="[1276,1401,679,705]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="subFamily" subFamily="Bibioninae">Bibioninae</taxonomicName>
all have apex of fore tibia modified into a long mucron or a circlet of shorter spines), relatively short antennae with compact antennal flagellomeres (
<taxonomicName id="EEFE3790FFBE4B46FDA8FD53FD08256B" authorityName="Walker" authorityYear="1848" box="[528,656,750,775]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FDA8FD53FD08256B" box="[528,656,750,775]" italics="true" pageId="3" pageNumber="454">Hesperinus</emphasis>
</taxonomicName>
with flagellomeres elongate), and vein R 
<subScript id="B57A4E56FFBE4B46FBDFFD42FB182561" attach="left" box="[1127,1152,767,781]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
elongate and subparallel to R 
<subScript id="B57A4E56FFBE4B46FF7FFC9FFF79255C" attach="left" box="[199,225,802,816]" fontSize="6" pageId="3" pageNumber="454">4+5</subScript>
(e.g., 
<figureCitation id="B1C55096FFBE4B46FE91FCADFEE02546" box="[297,376,784,810]" captionStart="FIGURES 2–3" captionStartId="11.[151,264,1518,1542]" captionTargetBox="[218,1372,186,1488]" captionTargetId="figure-16@11.[212,1376,181,1494]" captionTargetPageId="11" captionText="FIGURES 2–3. P. appendicula. 2. Wing. 3. Female habitus, lateral view. Scale bars = ca. 1.0 mm. Abbreviations: ba, basal appendix of R2+3; aem, anepimeron; aes, anepisternum; c, coxa; f, femur; h, halter; hb, hind basitarsus; kem, katepimeron; kes, katepisternum; m, meron; mtaes, metanepisternum; mtg, mediotergite; mtkes, metakatepisternum; ltg, laterotergite; pltrch, pleu- rotrochantin; prn, pronotum; sct, scutum; t, trochanter." figureDoi="http://doi.org/10.5281/zenodo.4529518" httpUri="https://zenodo.org/record/4529518/files/figure.png" pageId="3" pageNumber="454">Figs. 2</figureCitation>
, 
<figureCitation id="B1C55096FFBE4B46FE3CFCADFE3A2546" box="[388,418,784,810]" captionStart="FIGURES 12–13" captionStartId="19.[151,264,1948,1972]" captionTargetBox="[193,1398,784,1919]" captionTargetId="figure-280@19.[187,1401,777,1924]" captionTargetPageId="19" captionText="FIGURES 12–13. P. dolichopeza n. sp., holotype male. 12. Habitus, lateral. 13. Wing. Scale bars = ca. 1.0 mm. Abbreviations: hb, hind basitarsus." figureDoi="http://doi.org/10.5281/zenodo.4529528" httpUri="https://zenodo.org/record/4529528/files/figure.png" pageId="3" pageNumber="454">13</figureCitation>
, 
<figureCitation id="B1C55096FFBE4B46FE17FCADFE442546" box="[431,476,784,810]" captionStart="FIGURES 21–22" captionStartId="24.[151,264,1285,1309]" captionTargetBox="[227,1365,351,1257]" captionTargetId="figure-115@24.[222,1365,345,1261]" captionTargetPageId="24" captionText="FIGURES 21–22. Penthetria and Plecia, wings, illustrating variation in length and slope of vein R 2+3. 21a–b. Penthetira heteroptera, males (both from Maryland, USA). 22a. Plecia dentata Hardy, male. 22b. Plecia biformis Hardy, male (points, x, y, z described in “Discussion supporting generic diagnosis”). Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529538" httpUri="https://zenodo.org/record/4529538/files/figure.png" pageId="3" pageNumber="454">21b</figureCitation>
, 
<figureCitation id="B1C55096FFBE4B46FE5FFCADFD9C2546" box="[487,516,784,810]" captionStart="FIGURE 23" captionStartId="26.[151,250,1995,2019]" captionTargetBox="[222,1370,567,1968]" captionTargetId="figure-228@26.[216,1372,561,1972]" captionTargetPageId="26" captionText="FIGURE 23. P. mexicana, holotype male, habitus. Scale bar = ca. 1.0 mm. Abbreviations: hb, hind basitarsus." figureDoi="http://doi.org/10.5281/zenodo.4529540" httpUri="https://zenodo.org/record/4529540/files/figure.png" pageId="3" pageNumber="454">23</figureCitation>
) (R 
<subScript id="B57A4E56FFBE4B46FD88FC9FFDD2255C" attach="left" box="[560,586,802,816]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
absent in 
<taxonomicName id="EEFE3790FFBE4B46FD06FCADFCA62547" authorityName="Newman" authorityYear="1834" box="[702,830,784,811]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="subFamily" subFamily="Bibioninae">Bibioninae</taxonomicName>
). The elongate R 
<subScript id="B57A4E56FFBE4B46FC45FC9FFB8E255C" attach="left" box="[1021,1046,802,816]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
subparallel to R 
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is the character typically employed in keys for distinguishing 
<taxonomicName id="EEFE3790FFBE4B46FD23FC8EFC892521" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[667,785,819,845]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FD23FC8EFC892521" box="[667,785,819,845]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
from the similar genus 
<taxonomicName id="EEFE3790FFBE4B46FBA5FC8EFBFD2521" box="[1053,1125,819,845]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FBA5FC8EFBFD2521" box="[1053,1125,819,845]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
, which has R 
<subScript id="B57A4E56FFBE4B46FB46FCF8FA8F253F" attach="left" box="[1278,1303,837,851]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
shorter and more oblique or vertical (
<figureCitation id="B1C55096FFBE4B46FE0EFCE8FD93251C" box="[438,523,853,880]" captionStart="FIGURES 21–22" captionStartId="24.[151,264,1285,1309]" captionTargetBox="[227,1365,351,1257]" captionTargetId="figure-115@24.[222,1365,345,1261]" captionTargetPageId="24" captionText="FIGURES 21–22. Penthetria and Plecia, wings, illustrating variation in length and slope of vein R 2+3. 21a–b. Penthetira heteroptera, males (both from Maryland, USA). 22a. Plecia dentata Hardy, male. 22b. Plecia biformis Hardy, male (points, x, y, z described in “Discussion supporting generic diagnosis”). Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529538" httpUri="https://zenodo.org/record/4529538/files/figure.png" pageId="3" pageNumber="454">Fig. 22</figureCitation>
). Despite the fact that some species/specimens of 
<taxonomicName id="EEFE3790FFBE4B46FB87FCE8FB2D2503" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1087,1205,853,879]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FB87FCE8FB2D2503" box="[1087,1205,853,879]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
have R 
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approaching the condition found in 
<taxonomicName id="EEFE3790FFBE4B46FE7BFCC5FD9325FE" box="[451,523,888,914]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FE7BFCC5FD9325FE" box="[451,523,888,914]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
(e.g., compare 
<figureCitation id="B1C55096FFBE4B46FD02FCC5FCE825FF" box="[698,880,888,915]" captionStart="FIGURES 21–22" captionStartId="24.[151,264,1285,1309]" captionTargetBox="[227,1365,351,1257]" captionTargetId="figure-115@24.[222,1365,345,1261]" captionTargetPageId="24" captionText="FIGURES 21–22. Penthetria and Plecia, wings, illustrating variation in length and slope of vein R 2+3. 21a–b. Penthetira heteroptera, males (both from Maryland, USA). 22a. Plecia dentata Hardy, male. 22b. Plecia biformis Hardy, male (points, x, y, z described in “Discussion supporting generic diagnosis”). Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529538" httpUri="https://zenodo.org/record/4529538/files/figure.png" pageId="3" pageNumber="454">Figs. 21a &amp; 22a</figureCitation>
) and that the form of R 
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is essentially evaluated by gestalt rather than any discrete measurement, the extant species can typically be distinguished in this way with confidence.
</paragraph>
<paragraph id="29414C13FFBE4B46FF7FFC5CFA0923F3" blockId="3.[151,1437,151,2033]" pageId="3" pageNumber="454">
The use of R 
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2+ 
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</subScript>
in distinguishing these genera becomes more problematic with fossils, where there is a continuum in the form of R 
<subScript id="B57A4E56FFBE4B46FE1FFBAAFE582249" attach="left" box="[423,448,1047,1061]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
between typical 
<taxonomicName id="EEFE3790FFBE4B46FD3AFBB9FD602272" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[642,760,1028,1054]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FD3AFBB9FD602272" box="[642,760,1028,1054]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
and 
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<emphasis id="1B8A9001FFBE4B46FC89FBB9FCE12272" box="[817,889,1028,1054]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
(
<bibRefCitation id="4D6F31E2FFBE4B46FC30FBB9FBE82272" author="Collomb, F. - M. &amp; Nel, A. &amp; Fleck, G. &amp; Waller, A." box="[904,1136,1028,1055]" pageId="3" pageNumber="454" pagination="161 - 179" refId="ref32489" refString="Collomb, F. - M., Nel, A., Fleck, G. &amp; Waller, A. 2008 March flies and European Cenozoic palaeoclimates (Diptera: Bibionidae). Annales de la Societe Entomologique de France, 44 (2), 161 - 179. https: // doi. org / 10.1080 / 00379271.2008.10697553" type="journal article" year="2008">
Collomb 
<emphasis id="1B8A9001FFBE4B46FC4CFBB8FBBE2272" box="[1012,1062,1028,1054]" italics="true" pageId="3" pageNumber="454">et al</emphasis>
. 2008
</bibRefCitation>
, and e.g. see discussion of 
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<emphasis id="1B8A9001FFBE4B46FF2FFB9AFE95222D" box="[151,269,1063,1089]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
by 
<bibRefCitation id="4D6F31E2FFBE4B46FE82FB9AFDAC222D" author="Skartveit, J. &amp; Nel, A." box="[314,564,1063,1089]" pageId="3" pageNumber="454" pagination="1 - 83" refId="ref36689" refString="Skartveit, J. &amp; Nel, A. (2017) Revision of fossil Bibionidae (Insecta: Diptera) from French Oligocene deposits. Zootaxa, 4225 (1), 1 - 83. https: // doi. org / 10.11646 / zootaxa. 4225.1.1" type="journal article" year="2017">Skartveit &amp; Nel 2017</bibRefCitation>
). It may be useful, based on extant taxa, to develop a more objective way of describing the form of R 
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2+ 
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</subScript>
in order to help more accurately place fossil taxa. For example, in the wing variation of the extant species presented in 
<figureCitation id="B1C55096FFBE4B46FE49FBD0FDE022EB" box="[497,632,1133,1159]" captionStart="FIGURES 21–22" captionStartId="24.[151,264,1285,1309]" captionTargetBox="[227,1365,351,1257]" captionTargetId="figure-115@24.[222,1365,345,1261]" captionTargetPageId="24" captionText="FIGURES 21–22. Penthetria and Plecia, wings, illustrating variation in length and slope of vein R 2+3. 21a–b. Penthetira heteroptera, males (both from Maryland, USA). 22a. Plecia dentata Hardy, male. 22b. Plecia biformis Hardy, male (points, x, y, z described in “Discussion supporting generic diagnosis”). Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529538" httpUri="https://zenodo.org/record/4529538/files/figure.png" pageId="3" pageNumber="454">Figs. 21–22</figureCitation>
, 
<taxonomicName id="EEFE3790FFBE4B46FD3AFBD0FD6022EB" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[642,760,1133,1159]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FD3AFBD0FD6022EB" box="[642,760,1133,1159]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
can be defined by the slope of R 
<subScript id="B57A4E56FFBE4B46FBDEFBC2FBE722E1" attach="left" box="[1126,1151,1151,1165]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
less than 0.43 and 
<taxonomicName id="EEFE3790FFBE4B46FAEDFBD0FA0522EB" box="[1365,1437,1133,1159]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FAEDFBD0FA0522EB" box="[1365,1437,1133,1159]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
by the slope of R 
<subScript id="B57A4E56FFBE4B46FEE9FB1FFEF322DC" attach="left" box="[337,363,1186,1200]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
greater than 0.74 (slope defined here as rise/run, where rise = measure of imaginary line between points y and z, and where run = measurement of imaginary line between points x and z (see 
<figureCitation id="B1C55096FFBE4B46FB08FB0EFA8922A2" box="[1200,1297,1203,1230]" captionStart="FIGURES 21–22" captionStartId="24.[151,264,1285,1309]" captionTargetBox="[227,1365,351,1257]" captionTargetId="figure-115@24.[222,1365,345,1261]" captionTargetPageId="24" captionText="FIGURES 21–22. Penthetria and Plecia, wings, illustrating variation in length and slope of vein R 2+3. 21a–b. Penthetira heteroptera, males (both from Maryland, USA). 22a. Plecia dentata Hardy, male. 22b. Plecia biformis Hardy, male (points, x, y, z described in “Discussion supporting generic diagnosis”). Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529538" httpUri="https://zenodo.org/record/4529538/files/figure.png" pageId="3" pageNumber="454">Fig. 22b</figureCitation>
)). However, when slope is applied to the continuum of the form of R 
<subScript id="B57A4E56FFBE4B46FCB5FB55FCBE229A" attach="left" box="[781,806,1256,1270]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
found in fossil species, it is not as clear. For example, if slope is measured for the 15 
<taxonomicName id="EEFE3790FFBE4B46FE64FB44FDBC237F" box="[476,548,1273,1299]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FE64FB44FDBC237F" box="[476,548,1273,1299]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
and 15 
<taxonomicName id="EEFE3790FFBE4B46FDC6FB44FD6C237F" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[638,756,1273,1299]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FDC6FB44FD6C237F" box="[638,756,1273,1299]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
wings illustrated by 
<bibRefCitation id="4D6F31E2FFBE4B46FC59FB44FB70237F" author="Skartveit, J. &amp; Nel, A." box="[993,1256,1273,1299]" pageId="3" pageNumber="454" pagination="1 - 83" refId="ref36689" refString="Skartveit, J. &amp; Nel, A. (2017) Revision of fossil Bibionidae (Insecta: Diptera) from French Oligocene deposits. Zootaxa, 4225 (1), 1 - 83. https: // doi. org / 10.11646 / zootaxa. 4225.1.1" type="journal article" year="2017">Skartveit &amp; Nel (2017)</bibRefCitation>
, with a line discriminating 
<taxonomicName id="EEFE3790FFBE4B46FE98FAA1FEF0235A" box="[288,360,1308,1334]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FE98FAA1FEF0235A" box="[288,360,1308,1334]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
with slope&gt;= 0.50, it would correctly assign alone 27 of the 30 wings (i.e., without evaluating any other aspect of morphology). More variation should be studied before determining whether such a definition could be employed, but utilizing a discrete definition of slope and understanding the breadth of variation in the form of R 
<subScript id="B57A4E56FFBE4B46FF70FA2AFF7923C9" attach="left" box="[200,225,1431,1445]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
(or other veins) of extant species would undoubtably help to objectify the generic placement of fossil taxa.
</paragraph>
<paragraph id="29414C13FFBE4B46FF7FFA15FD4A20FF" blockId="3.[151,1437,151,2033]" pageId="3" pageNumber="454">
In addition to differences in the shape of R 
<subScript id="B57A4E56FFBE4B46FD23FA07FD2C23A4" attach="left" box="[667,692,1466,1480]" fontSize="6" pageId="3" pageNumber="454">2+3</subScript>
, 
<taxonomicName id="EEFE3790FFBE4B46FD79FA1AFCAF23AD" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[705,823,1447,1473]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FD79FA1AFCAF23AD" box="[705,823,1447,1473]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
and 
<taxonomicName id="EEFE3790FFBE4B46FCD3FA1AFC2B23AD" box="[875,947,1447,1473]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FCD3FA1AFC2B23AD" box="[875,947,1447,1473]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
differ in the form of the male terminalia; the male terminalia in 
<taxonomicName id="EEFE3790FFBE4B46FED1FA77FE472388" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[361,479,1482,1508]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FED1FA77FE472388" box="[361,479,1482,1508]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
are very simple, plesiomorphic (e.g., see 
<figureCitation id="B1C55096FFBE4B46FC17FA77FC6A2389" box="[943,1010,1482,1509]" captionStart="FIGURES 7–8" captionStartId="15.[151,264,1945,1969]" captionTargetBox="[361,1222,182,1917]" captionTargetId="figure-16@15.[361,1227,181,1921]" captionTargetPageId="15" captionText="FIGURES 7–8. P. arizonensis n. sp., holotype, male terminalia. 7. Dorsal. 8. Ventral. Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine." figureDoi="http://doi.org/10.5281/zenodo.4529522" httpUri="https://zenodo.org/record/4529522/files/figure.png" pageId="3" pageNumber="454">Fig. 7</figureCitation>
), relatively homogenous, with a stout, simple, gonostylus articulating in a socket formed from a relatively elongate gonocoxite. The male terminalia in 
<taxonomicName id="EEFE3790FFBE4B46FF2FF9ADFF472046" box="[151,223,1552,1578]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FF2FF9ADFF472046" box="[151,223,1552,1578]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
are much more complex, show great interspecific variability (see e.g., 
<bibRefCitation id="4D6F31E2FFBE4B46FC56F9ADFBED2047" author="Hardy, D. E." box="[1006,1141,1552,1579]" pageId="3" pageNumber="454" pagination="367 - 547" refId="ref33819" refString="Hardy, D. E. (1945) Revision of Nearctic Bibionidae, including Neotropical Plecia and Penthetria (Diptera). Kansas University Science Bulletin, 30, 367 - 547." type="journal article" year="1945">Hardy 1945</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFBE4B46FBC7F9ADFAB72046" author="Fitzgerald, S. J." box="[1151,1327,1552,1578]" pageId="3" pageNumber="454" pagination="133 - 164" refId="ref33025" refString="Fitzgerald, S. J. (1998) Eighteen new species of Neotropical Plecia (Diptera: Bibionidae) and taxonomic notes on the genus. Transactions of the Entomological Society of America, 124 (2), 133 - 164." type="journal article" year="1998">Fitzgerald 1998</bibRefCitation>
) and tend to have the tubular, gonostyus-socket-portion of the gonocoxite greatly shortened. However, this is less true in fossil specimens. As 
<bibRefCitation id="4D6F31E2FFBE4B46FE86F9EBFDCB201C" author="Skartveit, J. &amp; Nel, A." box="[318,595,1622,1648]" pageId="3" pageNumber="454" pagination="1 - 83" refId="ref36689" refString="Skartveit, J. &amp; Nel, A. (2017) Revision of fossil Bibionidae (Insecta: Diptera) from French Oligocene deposits. Zootaxa, 4225 (1), 1 - 83. https: // doi. org / 10.11646 / zootaxa. 4225.1.1" type="journal article" year="2017">Skartveit and Nel (2017)</bibRefCitation>
point out, the male terminalia of some fossil 
<taxonomicName id="EEFE3790FFBE4B46FBF4F9EBFB0C201C" box="[1100,1172,1622,1648]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FBF4F9EBFB0C201C" box="[1100,1172,1622,1648]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
are very plesiomorphic and 
<taxonomicName id="EEFE3790FFBE4B46FF7FF9C4FEA520FF" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[199,317,1657,1683]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FF7FF9C4FEA520FF" box="[199,317,1657,1683]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
-like (e.g., see 
<bibRefCitation id="4D6F31E2FFBE4B46FE5AF9C4FD5F20FF" author="Collomb, F. - M. &amp; Nel, A. &amp; Fleck, G. &amp; Waller, A." box="[482,711,1657,1684]" pageId="3" pageNumber="454" pagination="161 - 179" refId="ref32489" refString="Collomb, F. - M., Nel, A., Fleck, G. &amp; Waller, A. 2008 March flies and European Cenozoic palaeoclimates (Diptera: Bibionidae). Annales de la Societe Entomologique de France, 44 (2), 161 - 179. https: // doi. org / 10.1080 / 00379271.2008.10697553" type="journal article" year="2008">
Collomb 
<emphasis id="1B8A9001FFBE4B46FDF3F9C7FDE420FF" box="[587,636,1657,1683]" italics="true" pageId="3" pageNumber="454">et al</emphasis>
. 2008
</bibRefCitation>
).
</paragraph>
<paragraph id="29414C13FFBE4B46FF7FF921FA0A2173" blockId="3.[151,1437,151,2033]" pageId="3" pageNumber="454">
Additionally, 
<taxonomicName id="EEFE3790FFBE4B46FEDBF921FE4120DA" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[355,473,1692,1718]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FEDBF921FE4120DA" box="[355,473,1692,1718]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
has subtle parapsidal sutures (longitudinal sutures on the mesonotum corresponding to the position of dorsocentral setae), while they are often present as distinct grooves in 
<taxonomicName id="EEFE3790FFBE4B46FBC9F902FB2120B5" box="[1137,1209,1727,1753]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FBC9F902FB2120B5" box="[1137,1209,1727,1753]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
. Females of the two genera can be also be distinguished by 
<taxonomicName id="EEFE3790FFBE4B46FDF4F95FFD5A2090" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[588,706,1762,1788]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FDF4F95FFD5A2090" box="[588,706,1762,1788]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
having two segmented-cerci and 
<taxonomicName id="EEFE3790FFBE4B46FB82F95FFB1A2090" box="[1082,1154,1762,1788]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FB82F95FFB1A2090" box="[1082,1154,1762,1788]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
with one-segmented cer-ci—though some fossil 
<taxonomicName id="EEFE3790FFBE4B46FE1AF8B8FE722173" box="[418,490,1797,1823]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FE1AF8B8FE722173" box="[418,490,1797,1823]" italics="true" pageId="3" pageNumber="454">Plecia</emphasis>
</taxonomicName>
(e.g., 
<taxonomicName id="EEFE3790FFBE4B46FD88F8BBFCE92173" authority="Skartveit 2009" authorityName="Skartveit" authorityYear="2009" box="[560,881,1797,1823]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="species" species="clavifemur">
<emphasis id="1B8A9001FFBE4B46FD88F8BBFD5C2173" box="[560,708,1797,1823]" italics="true" pageId="3" pageNumber="454">P. clavifemur</emphasis>
<bibRefCitation id="4D6F31E2FFBE4B46FD72F8B8FCE92173" author="Skartveit, J." box="[714,881,1797,1823]" pageId="3" pageNumber="454" pagination="3 - 42" refId="ref36439" refString="Skartveit, J. (2009) Fossil Hesperinidae and Bibionidae from Baltic amber (Diptera: Bibionidae). Studia dipterologica, 15 (2008), 3 - 42." type="journal article" year="2009">Skartveit 2009</bibRefCitation>
</taxonomicName>
) have been described with two-segmented cerci.
</paragraph>
<paragraph id="29414C13FFBE4B46FF7FF895FC15219C" blockId="3.[151,1437,151,2033]" pageId="3" pageNumber="454">
It is also noteworthy that 
<taxonomicName id="EEFE3790FFBE4B46FE5EF895FDC4212E" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[486,604,1832,1858]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FE5EF895FDC4212E" box="[486,604,1832,1858]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
and the bibionid genus 
<taxonomicName id="EEFE3790FFBE4B46FCD1F895FC71212D" authorityName="Walker" authorityYear="1848" box="[873,1001,1832,1857]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FCD1F895FC71212D" box="[873,1001,1832,1857]" italics="true" pageId="3" pageNumber="454">Hesperinus</emphasis>
</taxonomicName>
cannot be reliably distinguished based on wing venation alone (
<bibRefCitation id="4D6F31E2FFBE4B46FE0DF8F6FDFA2109" author="Skartveit, J." box="[437,610,1867,1893]" pageId="3" pageNumber="454" pagination="3 - 42" refId="ref36439" refString="Skartveit, J. (2009) Fossil Hesperinidae and Bibionidae from Baltic amber (Diptera: Bibionidae). Studia dipterologica, 15 (2008), 3 - 42." type="journal article" year="2009">Skartveit 2009</bibRefCitation>
), which is critical in situations where the fossil specimen is represented only by a preserved wing. In fossils where more than the wing has been preserved, 
<taxonomicName id="EEFE3790FFBE4B46FB8DF8D0FB3321EB" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1077,1195,1901,1927]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FB8DF8D0FB3321EB" box="[1077,1195,1901,1927]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
has shorter antennae, with more compact flagellomeres and tends to be more stout-bodied, with shorter legs than 
<taxonomicName id="EEFE3790FFBE4B46FB1BF82CFABB21C6" authorityName="Walker" authorityYear="1848" box="[1187,1315,1937,1962]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FB1BF82CFABB21C6" box="[1187,1315,1937,1962]" italics="true" pageId="3" pageNumber="454">Hesperinus</emphasis>
</taxonomicName>
. Additionally, males of 
<taxonomicName id="EEFE3790FFBE4B46FE8AF80EFE3021A1" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[306,424,1971,1997]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FE8AF80EFE3021A1" box="[306,424,1971,1997]" italics="true" pageId="3" pageNumber="454">Penthetria</emphasis>
</taxonomicName>
are typically holoptic while both sexes of 
<taxonomicName id="EEFE3790FFBE4B46FCC6F809FC6621A1" authorityName="Walker" authorityYear="1848" box="[894,1022,1972,1997]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="454" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBE4B46FCC6F809FC6621A1" box="[894,1022,1972,1997]" italics="true" pageId="3" pageNumber="454">Hesperinus</emphasis>
</taxonomicName>
are dichoptic. See also “Intraspecific Variation” (below) with respect to the delimitation of fossil species.
</paragraph>
<paragraph id="29414C13FFB94B41FF7FFF2BFC11272D" blockId="4.[151,1437,150,717]" pageId="4" pageNumber="455">
<emphasis id="1B8A9001FFB94B41FF7FFF2BFD2726DD" bold="true" box="[199,703,150,177]" pageId="4" pageNumber="455">Classification and phylogenetic placement</emphasis>
. Some studies have placed 
<taxonomicName id="EEFE3790FFB94B41FC4FFF2AFBF526DD" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1015,1133,151,177]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FC4FFF2AFBF526DD" box="[1015,1133,151,177]" italics="true" pageId="4" pageNumber="455">Penthetria</emphasis>
</taxonomicName>
, along with the genus 
<taxonomicName id="EEFE3790FFB94B41FAD6FF2AFF2126B8" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FAD6FF2AFF2126B8" italics="true" pageId="4" pageNumber="455">Plecia</emphasis>
</taxonomicName>
, in the subfamily 
<taxonomicName id="EEFE3790FFB94B41FE3EFF06FE7526B9" authorityName="Duda" authorityYear="1930" box="[390,493,187,213]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="subFamily" subFamily="Pleciinae">Pleciinae</taxonomicName>
of 
<taxonomicName id="EEFE3790FFB94B41FDABFF06FD1626B9" baseAuthorityName="Fitzgerald" baseAuthorityYear="2004" box="[531,654,187,213]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="family">Bibionidae</taxonomicName>
(e.g., 
<bibRefCitation id="4D6F31E2FFB94B41FD6DFF06FCC426B9" author="Hardy, D. E." box="[725,860,187,213]" pageId="4" pageNumber="455" pagination="217 - 222" refId="ref34196" refString="Hardy, D. E. (1981) Bibionidae. In: McAlpine, J. F., Peterson, B. V., Shewell, G. E., Teskey, H. J., Vockeroth, J. R. &amp; Wood, D. M. (Eds.), Manual of Nearctic Diptera. Vol. 1. Canada Department of Agriculture Research Branch Monograph 27. Biosystematics Research Institute Ottawa, Ontario, pp. 217 - 222." type="book chapter" year="1981">Hardy 1981</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB94B41FCD1FF06FBE326B9" author="Blaschke-Berthold, U." box="[873,1147,187,213]" pageId="4" pageNumber="455" pagination="1 - 206" refId="ref32292" refString="Blaschke-Berthold, U. (1994) Anatomie und Phylogenie der Bibionomorpha (Insecta: Diptera). Bonner Zoologische Monographien, 34, 1 - 206." type="journal article" year="1994">Blaschke-Berthold 1994</bibRefCitation>
) or in the separate family 
<taxonomicName id="EEFE3790FFB94B41FF2FFF62FF662695" box="[151,254,223,249]" class="Insecta" family="Pleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="family">Pleciidae</taxonomicName>
(e.g., 
<bibRefCitation id="4D6F31E2FFB94B41FEFDFF62FD962696" author="Krivosheina, N. P." box="[325,526,223,250]" pageId="4" pageNumber="455" pagination="319 - 330" refId="ref34608" refString="Krivosheina, N. P. (1986) Family Pleciidae and Family Bibionidae. In: Soos, A. &amp; Papp, L., Catalogue of Palaearctic Diptera. Vol. 4. Sciaridae-Anisopodidae. Elsevier Science Publishing Co., Inc., New York, New York, pp. 313 - 316 + 319 - 330." type="book chapter" year="1986">Krivosheina 1986</bibRefCitation>
) based, in part, on similarities in the larvae of these genera including the “clearly separated frons and clypeus, presence of an additional lobe of the maxillae, and primitive spiracles” (
<bibRefCitation id="4D6F31E2FFB94B41FAA9FEBEFF48272E" author="Krivosheina, N. P." pageId="4" pageNumber="455" refId="ref34582" refString="Krivosheina, N. P. (1969) Ontogeny and evolution of Diptera. Nauka, Moscow, 290 pp. [in Russian]" type="book" year="1969">Krivosheina 1969</bibRefCitation>
) or musculature of the male terminalia (
<bibRefCitation id="4D6F31E2FFB94B41FD2BFE9AFCE4272D" author="Ovtshinnikova, O. G." box="[659,892,295,321]" pageId="4" pageNumber="455" pagination="149 - 152" refId="ref35626" refString="Ovtshinnikova, O. G. (1994) Pleciidae as a separate family on the basis of the musculature of the male genitalia (Diptera). Zoosystematica Rossica, 3 (1), 149 - 152." type="journal article" year="1994">Ovtshinnikova 1994</bibRefCitation>
).
</paragraph>
<paragraph id="29414C13FFB94B41FF7FFEF6FB5E240D" blockId="4.[151,1437,150,717]" pageId="4" pageNumber="455">
More recent morphological phylogenetic studies, however, have not supported these classifications, instead placing 
<taxonomicName id="EEFE3790FFB94B41FF4BFED2FEF127E5" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[243,361,367,393]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FF4BFED2FEF127E5" box="[243,361,367,393]" italics="true" pageId="4" pageNumber="455">Penthetria</emphasis>
</taxonomicName>
in its own subfamily (Penthetriinae), which is considered the most basal lineage of bibionids after 
<taxonomicName id="EEFE3790FFB94B41FF6CFE2EFECC27C0" authorityName="Walker" authorityYear="1848" box="[212,340,403,428]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FF6CFE2EFECC27C0" box="[212,340,403,428]" italics="true" pageId="4" pageNumber="455">Hesperinus</emphasis>
</taxonomicName>
(
<taxonomicName id="EEFE3790FFB94B41FEDEFE2EFE7E27C0" authorityName="Walker" authorityYear="1848" box="[358,486,403,428]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FEDEFE2EFE7E27C0" box="[358,486,403,428]" italics="true" pageId="4" pageNumber="455">Hesperinus</emphasis>
</taxonomicName>
+ (
<taxonomicName id="EEFE3790FFB94B41FDA9FE2EFD1F27C1" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[529,647,403,429]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FDA9FE2EFD1F27C1" box="[529,647,403,429]" italics="true" pageId="4" pageNumber="455">Penthetria</emphasis>
</taxonomicName>
+ (
<taxonomicName id="EEFE3790FFB94B41FD0BFE2EFD6327C1" box="[691,763,403,429]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FD0BFE2EFD6327C1" box="[691,763,403,429]" italics="true" pageId="4" pageNumber="455">Plecia</emphasis>
</taxonomicName>
+ 
<taxonomicName id="EEFE3790FFB94B41FCA5FE2EFC0027C1" authorityName="Newman" authorityYear="1834" box="[797,920,403,429]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="subFamily" subFamily="Bibioninae">
<emphasis id="1B8A9001FFB94B41FCA5FE2EFC0027C1" box="[797,920,403,429]" italics="true" pageId="4" pageNumber="455">Bibioninae</emphasis>
</taxonomicName>
))) (
<bibRefCitation id="4D6F31E2FFB94B41FC7DFE2EFB5427C1" author="Pinto, L. G. &amp; Amorim, D. S." box="[965,1228,403,429]" pageId="4" pageNumber="455" refId="ref35898" refString="Pinto, L. G. &amp; Amorim, D. S. (2000) Bibionidae (Diptera: Bibionomorpha). Morfologia e analise filogenetica. Holos, Ribeir &quot; o Preto, 98 pp. [in Portuguese]" type="book" year="2000">Pinto &amp; Amorim 2000</bibRefCitation>
, Fitzgerald 2004). This latter hypothesis was not corroborated by the molecular study by 
<bibRefCitation id="4D6F31E2FFB94B41FC00FE05FB0327BE" author="Sevcik, J. &amp; Kasprak, D. &amp; Mantic, M. &amp; Fitzgerald, S. &amp; Sevcikova, T. &amp; Tothova, A. &amp; Jaschhof, M." box="[952,1179,439,467]" pageId="4" pageNumber="455" pagination="1 - 30" refId="ref36295" refString="Sevcik, J., Kasprak, D., Mantic, M., Fitzgerald, S., Sevcikova, T., Tothova, A. &amp; Jaschhof, M. (2016) Molecular phylogeny of the megadiverse insect infraorder Bibionomorpha sensu lato (Diptera). PeerJ, 4 (e 2563), 1 - 30. https: // doi. org / 10.7717 / peerj. 2563" type="journal article" year="2016">
Ševčík 
<emphasis id="1B8A9001FFB94B41FBB5FE0AFBA727BD" box="[1037,1087,439,465]" italics="true" pageId="4" pageNumber="455">et al</emphasis>
. (2016)
</bibRefCitation>
, which found 
<taxonomicName id="EEFE3790FFB94B41FA85FE0AFF202798" baseAuthorityName="Loew" baseAuthorityYear="1868" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FA85FE0AFF202798" italics="true" pageId="4" pageNumber="455">Penthetria</emphasis>
</taxonomicName>
(along with 
<taxonomicName id="EEFE3790FFB94B41FEF1FE66FE512798" authorityName="Walker" authorityYear="1848" box="[329,457,475,500]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FEF1FE66FE512798" box="[329,457,475,500]" italics="true" pageId="4" pageNumber="455">Hesperinus</emphasis>
</taxonomicName>
) to be less basal within the bibionid family tree (
<taxonomicName id="EEFE3790FFB94B41FC4DFE66FBEA2799" authorityName="Newman" authorityYear="1834" box="[1013,1138,475,501]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="subFamily" subFamily="Bibioninae">Bibioninae</taxonomicName>
+ (
<taxonomicName id="EEFE3790FFB94B41FB23FE66FB7B2799" box="[1179,1251,475,501]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FB23FE66FB7B2799" box="[1179,1251,475,501]" italics="true" pageId="4" pageNumber="455">Plecia</emphasis>
</taxonomicName>
+ (
<emphasis id="1B8A9001FFB94B41FAB3FE66FE8F2474" italics="true" pageId="4" pageNumber="455">
<taxonomicName id="EEFE3790FFB94B41FAB3FE66FA192799" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1291,1409,475,501]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">Penthetria</taxonomicName>
+ 
<taxonomicName id="EEFE3790FFB94B41FF2FFE42FE8F2474" authorityName="Walker" authorityYear="1848" box="[151,279,511,536]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">Hesperinus</taxonomicName>
</emphasis>
))). 
<bibRefCitation id="4D6F31E2FFB94B41FEFBFE42FD182475" author="Skartveit, J. &amp; Ansorge, J." box="[323,640,511,537]" pageId="4" pageNumber="455" pagination="163 - 172" refId="ref36526" refString="Skartveit, J. &amp; Ansorge, J. (2020) A new genus and subfamily of fossil Bibionidae (Diptera) from the Lower Cretaceous, with new classification of the Bibionidae. Palaeoentomology, 3 (2), 163 - 172. https: // doi. org / 10.11646 / palaeoentomology. 3.2.5" type="journal article" year="2020">Skartveit &amp; Ansorge (2020)</bibRefCitation>
propose placing 
<taxonomicName id="EEFE3790FFB94B41FCFDFE42FC5D2474" authorityName="Walker" authorityYear="1848" box="[837,965,511,536]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FCFDFE42FC5D2474" box="[837,965,511,536]" italics="true" pageId="4" pageNumber="455">Hesperinus</emphasis>
</taxonomicName>
and 
<taxonomicName id="EEFE3790FFB94B41FC47FE42FBED2475" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1023,1141,511,537]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FC47FE42FBED2475" box="[1023,1141,511,537]" italics="true" pageId="4" pageNumber="455">Penthetria</emphasis>
</taxonomicName>
together in the subfamily 
<taxonomicName id="EEFE3790FFB94B41FF2FFD9EFEB32451" box="[151,299,547,573]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="subFamily" subFamily="Hesperininae">Hesperininae</taxonomicName>
based on the 
<bibRefCitation id="4D6F31E2FFB94B41FE7AFD99FD062452" author="Sevcik, J. &amp; Kasprak, D. &amp; Mantic, M. &amp; Fitzgerald, S. &amp; Sevcikova, T. &amp; Tothova, A. &amp; Jaschhof, M." box="[450,670,547,575]" pageId="4" pageNumber="455" pagination="1 - 30" refId="ref36295" refString="Sevcik, J., Kasprak, D., Mantic, M., Fitzgerald, S., Sevcikova, T., Tothova, A. &amp; Jaschhof, M. (2016) Molecular phylogeny of the megadiverse insect infraorder Bibionomorpha sensu lato (Diptera). PeerJ, 4 (e 2563), 1 - 30. https: // doi. org / 10.7717 / peerj. 2563" type="journal article" year="2016">
Ševčík 
<emphasis id="1B8A9001FFB94B41FDADFD9EFDDE2451" box="[533,582,547,573]" italics="true" pageId="4" pageNumber="455">et al</emphasis>
. (2016)
</bibRefCitation>
study, which finds these two genera to be sister taxa. Further work is needed to reconcile the contradictory hypotheses of these morphological and molecular studies.
</paragraph>
<paragraph id="29414C13FFB94B41FF7FFDD6FCB224A1" blockId="4.[151,1437,150,717]" pageId="4" pageNumber="455">
Characters to support the monophyly of 
<taxonomicName id="EEFE3790FFB94B41FD33FDD6FC9924E9" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[651,769,619,645]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB94B41FD33FDD6FC9924E9" box="[651,769,619,645]" italics="true" pageId="4" pageNumber="455">Penthetria</emphasis>
</taxonomicName>
are sparse. Fitzgerald (2004) suggested the larval mesothorax with two pseudosegments as a synapomorphy, but so few larvae are known that this character will have to be further vetted as the larvae of more species are discovered.
</paragraph>
<caption id="7D811C9BFFB94B41FF2FF803FD142197" ID-DOI="http://doi.org/10.5281/zenodo.4529514" ID-Zenodo-Dep="4529514" httpUri="https://zenodo.org/record/4529514/files/figure.png" pageId="4" pageNumber="455" startId="4.[151,250,1982,2006]" targetBox="[331,1264,760,1945]" targetPageId="4">
<paragraph id="29414C13FFB94B41FF2FF803FD142197" blockId="4.[151,1436,1982,2045]" pageId="4" pageNumber="455">
<emphasis id="1B8A9001FFB94B41FF2FF803FE8B21BA" bold="true" box="[151,275,1982,2006]" pageId="4" pageNumber="455">FIGURE 1.</emphasis>
<taxonomicName id="EEFE3790FFB94B41FEA0F802FE3E21BB" baseAuthorityName="Say" baseAuthorityYear="1823" box="[280,422,1983,2007]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="455" phylum="Arthropoda" rank="species" species="heteroptera">
<emphasis id="1B8A9001FFB94B41FEA0F802FE3E21BB" box="[280,422,1983,2007]" italics="true" pageId="4" pageNumber="455">P. heteroptera</emphasis>
</taxonomicName>
, habitus, male (above) and female (below), 28 October, North Carolina, USA, Matt Bertone. Images courtesy of Matt Bertone (© Matt Bertone 2013).
</paragraph>
</caption>
<paragraph id="29414C13FFB84B40FF7FFF2BFCFF272D" blockId="5.[151,1437,150,2013]" pageId="5" pageNumber="456">
<emphasis id="1B8A9001FFB84B40FF7FFF2BFDC126DC" bold="true" box="[199,601,150,177]" pageId="5" pageNumber="456">Comments on generic description</emphasis>
. The adult and larval generic description provided below is based, in part, on previous phylogenetic and descriptive work on the family 
<taxonomicName id="EEFE3790FFB84B40FCFFFF06FB1526B9" authority="(Fitzgerald 2004)" baseAuthorityName="Fitzgerald" baseAuthorityYear="2004" box="[839,1165,187,213]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="family">Bibionidae (Fitzgerald 2004)</taxonomicName>
and readers are referred there for further discussion concerning homology of structures, material examined, and the broader context from which the description was developed. Examination of all the New World species, as well as a small number of species from the Palearctic and Oriental bioregions were examined.
</paragraph>
<paragraph id="29414C13FFB84B40FF7FFEF6FABA2799" blockId="5.[151,1437,150,2013]" pageId="5" pageNumber="456">
A sclerotized aedeagus is considered absent in 
<taxonomicName id="EEFE3790FFB84B40FD65FEF6FCCB2709" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[733,851,331,357]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB84B40FD65FEF6FCCB2709" box="[733,851,331,357]" italics="true" pageId="5" pageNumber="456">Penthetria</emphasis>
</taxonomicName>
by Fitzgerald (2004) and it has been presented this way in the generic description (below). 
<bibRefCitation id="4D6F31E2FFB84B40FDE3FED2FCE127E5" author="Blaschke-Berthold, U." box="[603,889,367,393]" pageId="5" pageNumber="456" pagination="1 - 206" refId="ref32292" refString="Blaschke-Berthold, U. (1994) Anatomie und Phylogenie der Bibionomorpha (Insecta: Diptera). Bonner Zoologische Monographien, 34, 1 - 206." type="journal article" year="1994">Blaschke-Berthold (1994</bibRefCitation>
, 
<figureCitation id="B1C55096FFB84B40FC3DFED2FC5127E6" box="[901,969,367,394]" captionStart="FIGURES 4–6" captionStartId="13.[151,264,1964,1988]" captionTargetBox="[387,1193,189,1934]" captionTargetId="figure-16@13.[387,1201,181,1940]" captionTargetPageId="13" captionText="FIGURES 4–6. P. appendicula, male terminalia. 4. Dorsal. 5. Posterodorsal. 6. Ventral. Scale bar = ca. 0.5 mm. Abbreviations: gc, gonocoxite; gs, gonostylus; p, paramere; sls, spine-like setae; t9, tergite nine." figureDoi="http://doi.org/10.5281/zenodo.4529520" httpUri="https://zenodo.org/record/4529520/files/figure.png" pageId="5" pageNumber="456">Fig. 6</figureCitation>
) suggests that the aedeagus is fused with the parameres in this genus but whether it is absent or indistinguishably fused, a distinct, separate aedeagal sclerite cannot be identified. Sperm transfer in 
<taxonomicName id="EEFE3790FFB84B40FDF1FE0AFD2727BD" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[585,703,439,465]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB84B40FDF1FE0AFD2727BD" box="[585,703,439,465]" italics="true" pageId="5" pageNumber="456">Penthetria</emphasis>
</taxonomicName>
is apparently not achieved via an intromittent organ, but probably by spermatophore (Blaschke-Bethold 1994), as is known in some other 
<taxonomicName id="EEFE3790FFB84B40FC03FE66FA862799" authority="(Leppla et al. 1975)" baseAuthorityName="Leppla" baseAuthorityYear="1975" box="[955,1310,474,501]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="family">
Bibionidae (
<bibRefCitation id="4D6F31E2FFB84B40FBFCFE66FA8D2799" author="Leppla, N. C. &amp; Carlysle, T. C. &amp; Guy, R. H." box="[1092,1301,474,501]" pageId="5" pageNumber="456" pagination="299 - 306" refId="ref34810" refString="Leppla, N. C., Carlysle, T. C. &amp; Guy, R. H. (1975) Reproductive systems and the mechanics of copulation in Plecia nearctica Hardy (Diptera: Bibionidae). International Journal of Insect Morphology and Embryology, 4 (4), 299 - 306. https: // doi. org / 10.1016 / 0020 - 7322 (75) 90018 - 5" type="journal article" year="1975">
Leppla 
<emphasis id="1B8A9001FFB84B40FB22FE66FB542799" box="[1178,1228,475,501]" italics="true" pageId="5" pageNumber="456">et al</emphasis>
. 1975
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="29414C13FFB84B40FF7FFE42FB0824C5" blockId="5.[151,1437,150,2013]" pageId="5" pageNumber="456">
Larvae are known for only two species of 
<taxonomicName id="EEFE3790FFB84B40FD00FE42FCB62475" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[696,814,511,537]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB84B40FD00FE42FCB62475" box="[696,814,511,537]" italics="true" pageId="5" pageNumber="456">Penthetria</emphasis>
</taxonomicName>
. Aspects of the larva of the Palearctic species 
<taxonomicName id="EEFE3790FFB84B40FAE9FE42FF572451" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB84B40FAE9FE42FF572451" italics="true" pageId="5" pageNumber="456">P. funebris</emphasis>
</taxonomicName>
have been illustrated in various papers—mandible (
<bibRefCitation id="4D6F31E2FFB84B40FCABFD9EFC312451" author="Hennig, W." box="[787,937,547,573]" pageId="5" pageNumber="456" refId="ref34484" refString="Hennig, W. (1948) Die Larvenformen der Dipteren. 1. Teil. Akademie-Verlag, Berlin, 184 pp." type="book" year="1948">Hennig 1948</bibRefCitation>
), habitus and posterior spiracle (
<bibRefCitation id="4D6F31E2FFB84B40FAAAFD9EFF4B240E" author="Krivosheina, N. P." pageId="5" pageNumber="456" pagination="210 - 227" refId="ref34544" refString="Krivosheina, N. P. (1962) European larvae of Bibionidae (Diptera: Nematocera), with identification keys for some species. Pedobiologia, 1, 210 - 227. [in Russian]" type="journal article" year="1962">Krivosheina 1962</bibRefCitation>
), habitus, posterior spiracle, mandible, and labium (
<bibRefCitation id="4D6F31E2FFB84B40FC9AFDFAFBE3240E" author="Krivosheina, N. P. &amp; Mamaev, B. M." box="[802,1147,583,610]" pageId="5" pageNumber="456" refId="ref34773" refString="Krivosheina, N. P. &amp; Mamaev, B. M. (1967) Key to the larvae of wood inhabiting dipteran insects. Nauka, Moscow, 367 pp. [in Russian]" type="book" year="1967">Krivosheina &amp; Mamaev 1967</bibRefCitation>
), and labium and maxilla (Fitzgerald 2004). The egg, larva, and pupal stages of 
<taxonomicName id="EEFE3790FFB84B40FCB1FDD6FB8024E9" authority="Wiedemann" authorityName="Wiedemann" authorityYear="1830" box="[777,1048,619,645]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="species" species="japonica">
<emphasis id="1B8A9001FFB84B40FCB1FDD6FC1124E8" box="[777,905,619,644]" italics="true" pageId="5" pageNumber="456">P. japonica</emphasis>
Wiedemann
</taxonomicName>
were described and illustrated by 
<bibRefCitation id="4D6F31E2FFB84B40FF2FFD32FEFD24C5" author="Yuan, R. &amp; Li, S. &amp; Du, Q. &amp; Lei, C. &amp; Zhu, F." box="[151,357,654,681]" pageId="5" pageNumber="456" pagination="320 - 324" refId="ref37430" refString="Yuan, R., Li, S., Du, Q., Lei, C. &amp; Zhu, F. (2015) Immature stages description of March Fly, Penthetria japonica Wiedemann (Diptera: Bibionidae), from Hubei, China. Entomological News, 124 (5), 320 - 324. https: // doi. org / 10.3157 / 021.124.0503" type="journal article" year="2015">
Yuan 
<emphasis id="1B8A9001FFB84B40FF60FD32FE9124C5" box="[216,265,655,681]" italics="true" pageId="5" pageNumber="456">et al</emphasis>
. (2015)
</bibRefCitation>
. The larval description below is based on the study of these two species.
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFB84B42FF7FFD0FFDA82315" lastPageId="7" lastPageNumber="458" pageId="5" pageNumber="456" type="description">
<paragraph id="29414C13FFB84B43FF7FFD0FFCC620D1" blockId="5.[151,1437,150,2013]" lastBlockId="6.[151,1437,151,2013]" lastPageId="6" lastPageNumber="457" pageId="5" pageNumber="456">
<emphasis id="1B8A9001FFB84B40FF7FFD0FFE2E24A0" bold="true" box="[199,438,690,716]" pageId="5" pageNumber="456">Generic description</emphasis>
. 
<emphasis id="1B8A9001FFB84B40FE7FFD0EFD9C24A1" box="[455,516,691,717]" italics="true" pageId="5" pageNumber="456">Adult</emphasis>
: Small to moderately large flies 4.0-11.0 mm. Head: Male head dorsoventrally compressed; in dorsal view, slightly wider than long, oval-shaped. Head and antennae black to brown. Male with almost entire dorsal surface of head occupied by broadly holoptic compound eye (
<figureCitation id="B1C55096FFB84B40FB80FD46FBE22579" box="[1080,1146,763,789]" captionStart="FIGURE 1" captionStartId="4.[151,250,1982,2006]" captionTargetBox="[331,1264,760,1945]" captionTargetId="figure-296@4.[309,1278,741,1959]" captionTargetPageId="4" captionText="FIGURE 1. P. heteroptera, habitus, male (above) and female (below), 28 October, North Carolina, USA, Matt Bertone. Images courtesy of Matt Bertone (© Matt Bertone 2013)." figureDoi="http://doi.org/10.5281/zenodo.4529514" httpUri="https://zenodo.org/record/4529514/files/figure.png" pageId="5" pageNumber="456">Fig. 1</figureCitation>
), except 
<taxonomicName id="EEFE3790FFB84B40FB5CFD46FAC42579" box="[1252,1372,763,789]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB84B40FB5CFD46FAC42579" box="[1252,1372,763,789]" italics="true" pageId="5" pageNumber="456">P. funebris</emphasis>
</taxonomicName>
, with compound eyes narrowly separated dorsally by longitudinal strip of frons approximately subequal in width to antennal flagellum. Male compound eye strongly divided into larger dorsal region and smaller ventral region; dorsal region usually lighter in color than ventral region and with larger ommatidia. Division between dorsal and ventral regions of compound eye often distinguished by distinct narrow longitudinal step (the dorsal region of the eye is often taller than the ventral region of the eye; where the dorsal and ventral regions meet laterally, a short step, curb, or groove is often visible as in 
<figureCitation id="B1C55096FFB84B40FDB8FC6EFDCD2581" box="[512,597,979,1005]" captionStart="FIGURES 12–13" captionStartId="19.[151,264,1948,1972]" captionTargetBox="[193,1398,784,1919]" captionTargetId="figure-280@19.[187,1401,777,1924]" captionTargetPageId="19" captionText="FIGURES 12–13. P. dolichopeza n. sp., holotype male. 12. Habitus, lateral. 13. Wing. Scale bars = ca. 1.0 mm. Abbreviations: hb, hind basitarsus." figureDoi="http://doi.org/10.5281/zenodo.4529528" httpUri="https://zenodo.org/record/4529528/files/figure.png" pageId="5" pageNumber="456">Fig. 12</figureCitation>
). Division not marked by narrow, longitudinal, shining, sclerotized band; smooth, triangular area devoid of ommatidia also absent. Female head dorsoventrally compressed, slightly wider than long. Female compound eye round or oval in shape, convex, dichoptic (
<figureCitation id="B1C55096FFB84B40FBB0FBA6FBD42259" box="[1032,1100,1051,1077]" captionStart="FIGURE 1" captionStartId="4.[151,250,1982,2006]" captionTargetBox="[331,1264,760,1945]" captionTargetId="figure-296@4.[309,1278,741,1959]" captionTargetPageId="4" captionText="FIGURE 1. P. heteroptera, habitus, male (above) and female (below), 28 October, North Carolina, USA, Matt Bertone. Images courtesy of Matt Bertone (© Matt Bertone 2013)." figureDoi="http://doi.org/10.5281/zenodo.4529514" httpUri="https://zenodo.org/record/4529514/files/figure.png" pageId="5" pageNumber="456">Fig. 1</figureCitation>
), broadly separated by frons. Female compound eye with inconspicuous, slight longitudinal depression dividing compound eye, but lacking any difference in size of ommatidia or color between dorsal and ventral regions. Compound eye of both sexes bare or clothed with minute, stiff, or very elongate hairs. Ocellar tubercle well developed and prominent in males, moderately to weakly developed in females. Both sexes with three ocelli arranged in small, equilateral triangle. Female with small tubercle or longitudinal ridge just posterior to antennal sockets. Male frons reduced to small, triangular region between anterior margin of compound eye, posterior margin of antennal sockets bearing minute tubercle (or a minute pit in 
<taxonomicName id="EEFE3790FFB84B40FEFDFAAAFE26235D" box="[325,446,1303,1329]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB84B40FEFDFAAAFE26235D" box="[325,446,1303,1329]" italics="true" pageId="5" pageNumber="456">P. funebris</emphasis>
</taxonomicName>
). Sclerotized rostrum not produced, oral margin somewhat ventral in position. Antenna located anteriorly along oral margin, with 7–10 short, stout flagellomeres. Flagellomeres slightly broader than long, except most basal flagellomere, about two times as long as subsequent flagellomere. Pedicel and flagellomeres with subapical ring of short setae and numerous dense, minute, appressed setae. Apical flagellomere slightly more elongate than preceding flagellomere, with rounded point apically. Maxillary palps with five segments; basal segment minute, third segment thickened with dorsal, diagonal sensory pit with dense, minute, specialized setae; apical palpomere slender, more elongate than previous palpomeres. Clypeus broader than long, never elongate. Head entirely sclerotized ventrally. Thorax (
<figureCitation id="B1C55096FFB84B40FE50F9AEFDB52041" box="[488,557,1555,1581]" captionStart="FIGURES 2–3" captionStartId="11.[151,264,1518,1542]" captionTargetBox="[218,1372,186,1488]" captionTargetId="figure-16@11.[212,1376,181,1494]" captionTargetPageId="11" captionText="FIGURES 2–3. P. appendicula. 2. Wing. 3. Female habitus, lateral view. Scale bars = ca. 1.0 mm. Abbreviations: ba, basal appendix of R2+3; aem, anepimeron; aes, anepisternum; c, coxa; f, femur; h, halter; hb, hind basitarsus; kem, katepimeron; kes, katepisternum; m, meron; mtaes, metanepisternum; mtg, mediotergite; mtkes, metakatepisternum; ltg, laterotergite; pltrch, pleu- rotrochantin; prn, pronotum; sct, scutum; t, trochanter." figureDoi="http://doi.org/10.5281/zenodo.4529518" httpUri="https://zenodo.org/record/4529518/files/figure.png" pageId="5" pageNumber="456">Fig. 3</figureCitation>
): Precoxal-bridge complete. Basisternum present, presternum absent. Dorsum of thorax smooth, matte, often with thick dark pruinosity, usually black, dark brown, anteriorly black and posteriorly orange, or rarely entirely orange, as in 
<taxonomicName id="EEFE3790FFB84B40FDC1F9E6FCCA201A" authority="(Brunetti)" baseAuthorityName="Brunetti" baseAuthorityYear="1911" box="[633,850,1627,1654]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="species" species="indica">
<emphasis id="1B8A9001FFB84B40FDC1F9E6FD422019" box="[633,730,1627,1653]" italics="true" pageId="5" pageNumber="456">P. indica</emphasis>
(Brunetti)
</taxonomicName>
. Parapsidal sutures subtle and not distinct as in some 
<taxonomicName id="EEFE3790FFB84B40FF2FF9C2FF4720F5" box="[151,223,1663,1689]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="456" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB84B40FF2FF9C2FF4720F5" box="[151,223,1663,1689]" italics="true" pageId="5" pageNumber="456">Plecia</emphasis>
</taxonomicName>
. Dorsum of thorax with dense, short, stiff to very elongate hairs, laterally, anteriorly, and in dorsocentral rows posteriorly. Thoracic pleura black to dark brown. Males with minute to moderately long hairs on metakatepisternum, and dorsal half of katepisternum. Anepisternum bare or with cluster of hairs at posterior margin. Meron, laterotergite, and mediotergite bare. Female thoracic pleura generally less hairy than male. Legs: Legs black to dark brown. All legs with five tarsomeres, simple tarsal claws, pulvilli and pulvilliform empodia. Anterior coxa short and not reaching ventral edge of katepisternum; reaching about half length of katepisternum in lateral view, usually with elongate hair. Fore femur relatively slender and more elongate; not short and swollen. Fore tibia elongate, slender, apex unmodified, outer edge not developed into a mucron. Fore tibia distally with a single, short spur. Middle legs unremarkable; with two apical tibial spurs. Hind legs with hind femur swollen apically to more slender and only slightly enlarged apically. Hind tibia slender or swollen apically. Inner surface of hind tibia without elongate field of small, round, black, specialized sensilla. Spurs of hind tibia slender, apically acute, subequal in length, or ventral spur slightly more elongate than dorsal spur. In males, hind tarsomeres vary in shape, from slender, elongate, parallel-sided to slightly swollen and sausage-shaped. Female tarsomeres never swollen; slender to robust. Halter black. Wing (
<figureCitation id="B1C55096FFBB4B43FF5DFEBEFEAE2771" box="[229,310,259,285]" captionStart="FIGURES 2–3" captionStartId="11.[151,264,1518,1542]" captionTargetBox="[218,1372,186,1488]" captionTargetId="figure-16@11.[212,1376,181,1494]" captionTargetPageId="11" captionText="FIGURES 2–3. P. appendicula. 2. Wing. 3. Female habitus, lateral view. Scale bars = ca. 1.0 mm. Abbreviations: ba, basal appendix of R2+3; aem, anepimeron; aes, anepisternum; c, coxa; f, femur; h, halter; hb, hind basitarsus; kem, katepimeron; kes, katepisternum; m, meron; mtaes, metanepisternum; mtg, mediotergite; mtkes, metakatepisternum; ltg, laterotergite; pltrch, pleu- rotrochantin; prn, pronotum; sct, scutum; t, trochanter." figureDoi="http://doi.org/10.5281/zenodo.4529518" httpUri="https://zenodo.org/record/4529518/files/figure.png" pageId="6" pageNumber="457">Figs. 2</figureCitation>
, 
<figureCitation id="B1C55096FFBB4B43FEFBFEBEFEFA2771" box="[323,354,259,285]" captionStart="FIGURES 12–13" captionStartId="19.[151,264,1948,1972]" captionTargetBox="[193,1398,784,1919]" captionTargetId="figure-280@19.[187,1401,777,1924]" captionTargetPageId="19" captionText="FIGURES 12–13. P. dolichopeza n. sp., holotype male. 12. Habitus, lateral. 13. Wing. Scale bars = ca. 1.0 mm. Abbreviations: hb, hind basitarsus." figureDoi="http://doi.org/10.5281/zenodo.4529528" httpUri="https://zenodo.org/record/4529528/files/figure.png" pageId="6" pageNumber="457">13</figureCitation>
): Length, 4.5-12.5 mm, elongate, reaching tip of abdomen, except brachypterous and distinctly shortened in males of 
<taxonomicName id="EEFE3790FFBB4B43FE37FE9AFD9E272D" box="[399,518,295,321]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="457" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFBB4B43FE37FE9AFD9E272D" box="[399,518,295,321]" italics="true" pageId="6" pageNumber="457">P. funebris</emphasis>
</taxonomicName>
. Wing brown to blackish fumose. Costal cell often darker than remainder of wing in taxa with fumose wings. Wing color solid or with grade in intensity, but without distinct markings. Pterostigma oval, usually brown to black. Anterior wing veins typically darker than posterior veins. Wing without macrotrichia, with microtrichia. Anal lobe well developed. Costa ends at or just beyond R 
<subScript id="B57A4E56FFBB4B43FC5FFE18FB9927DF" attach="left" box="[999,1025,421,435]" fontSize="6" pageId="6" pageNumber="457">4+5</subScript>
(not shortened or thickened as 
<taxonomicName id="EEFE3790FFBB4B43FAD8FE2EFEA627BE" authority="Hardy" authorityName="Hardy" authorityYear="1961" class="Insecta" family="Bibionidae" genus="Enicoscolus" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="457" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBB4B43FAD8FE2EFF7327BD" italics="true" pageId="6" pageNumber="457">Enicoscolus</emphasis>
Hardy
</taxonomicName>
). Subcosta long, complete. Radius without horizontal microstriations. Rs furcate; R 
<subScript id="B57A4E56FFBB4B43FB51FE74FA9A27BB" attach="left" box="[1257,1282,457,471]" fontSize="6" pageId="6" pageNumber="457">2+3</subScript>
present, relatively elongate and subparallel to slightly oblique to R 
<subScript id="B57A4E56FFBB4B43FD43FE50FC8D2797" attach="left" box="[763,789,493,507]" fontSize="6" pageId="6" pageNumber="457">4+5</subScript>
; base R 
<subScript id="B57A4E56FFBB4B43FCC8FE50FC112797" attach="left" box="[880,905,493,507]" fontSize="6" pageId="6" pageNumber="457">2+3</subScript>
often arising at right angle to R 
<subScript id="B57A4E56FFBB4B43FB49FE50FA932797" attach="left" box="[1265,1291,493,507]" fontSize="6" pageId="6" pageNumber="457">4+5</subScript>
then sharply bent in direction of wing tip. Base of R 
<subScript id="B57A4E56FFBB4B43FDF5FDACFDFE2473" attach="left" box="[589,614,529,543]" fontSize="6" pageId="6" pageNumber="457">2+3</subScript>
sometimes with sub-basal stump (appendix) (e.g., 
<taxonomicName id="EEFE3790FFBB4B43FB1AFE42FA002475" authority="Hardy" authorityName="Hardy" authorityYear="1945" box="[1186,1432,511,537]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="457" phylum="Arthropoda" rank="species" species="appendicula">
<emphasis id="1B8A9001FFBB4B43FB1AFE42FADF2475" box="[1186,1351,511,537]" italics="true" pageId="6" pageNumber="457">P. appendicula</emphasis>
Hardy
</taxonomicName>
; 
<figureCitation id="B1C55096FFBB4B43FF2FFD9EFF432451" box="[151,219,547,573]" captionStart="FIGURES 2–3" captionStartId="11.[151,264,1518,1542]" captionTargetBox="[218,1372,186,1488]" captionTargetId="figure-16@11.[212,1376,181,1494]" captionTargetPageId="11" captionText="FIGURES 2–3. P. appendicula. 2. Wing. 3. Female habitus, lateral view. Scale bars = ca. 1.0 mm. Abbreviations: ba, basal appendix of R2+3; aem, anepimeron; aes, anepisternum; c, coxa; f, femur; h, halter; hb, hind basitarsus; kem, katepimeron; kes, katepisternum; m, meron; mtaes, metanepisternum; mtg, mediotergite; mtkes, metakatepisternum; ltg, laterotergite; pltrch, pleu- rotrochantin; prn, pronotum; sct, scutum; t, trochanter." figureDoi="http://doi.org/10.5281/zenodo.4529518" httpUri="https://zenodo.org/record/4529518/files/figure.png" pageId="6" pageNumber="457">Fig. 2</figureCitation>
). R 
<subScript id="B57A4E56FFBB4B43FEB9FD88FE82242F" attach="left" box="[257,282,565,579]" fontSize="6" pageId="6" pageNumber="457">2+3</subScript>
branches from R 
<subScript id="B57A4E56FFBB4B43FE61FD88FE6B242F" attach="left" box="[473,499,565,579]" fontSize="6" pageId="6" pageNumber="457">4+5</subScript>
from slightly basal to, even with, or distinctly distal to, r-m crossvein. Length of r-m crossvein much shorter than base of Rs. Posterior end of r-m meeting stem of medial fork (M 
<subScript id="B57A4E56FFBB4B43FB01FDE4FB4B240B" attach="left" box="[1209,1235,601,615]" fontSize="6" pageId="6" pageNumber="457">1+2</subScript>
) or more distally, connecting directly to M 
<subScript id="B57A4E56FFBB4B43FE16FDC0FE2E24E7" attach="left" box="[430,438,637,651]" fontSize="6" pageId="6" pageNumber="457">1</subScript>
(e.g., 
<taxonomicName id="EEFE3790FFBB4B43FE47FDD6FC9724E9" authority="Wiedemann" authorityName="Wiedemann" authorityYear="1830" box="[511,783,619,645]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="457" phylum="Arthropoda" rank="species" species="japonica">
<emphasis id="1B8A9001FFBB4B43FE47FDD6FDE624E8" box="[511,638,619,644]" italics="true" pageId="6" pageNumber="457">P. japonica</emphasis>
Wiedemann
</taxonomicName>
; 
<bibRefCitation id="4D6F31E2FFBB4B43FCA3FDD6FBA324E9" author="Hardy, D. E. &amp; Takahashi, M." box="[795,1083,619,645]" pageId="6" pageNumber="457" pagination="383 - 449" refId="ref34286" refString="Hardy, D. E. &amp; Takahashi, M. (1960) Revision of the Japanese Bibionidae (Diptera, Nematocera). Pacific Insects, 2 (4), 383 - 449." type="journal article" year="1960">Hardy &amp; Takahashi 1960</bibRefCitation>
, 
<figureCitation id="B1C55096FFBB4B43FBFFFDD6FB0324EA" box="[1095,1179,619,646]" captionStart="FIGURES 2–3" captionStartId="11.[151,264,1518,1542]" captionTargetBox="[218,1372,186,1488]" captionTargetId="figure-16@11.[212,1376,181,1494]" captionTargetPageId="11" captionText="FIGURES 2–3. P. appendicula. 2. Wing. 3. Female habitus, lateral view. Scale bars = ca. 1.0 mm. Abbreviations: ba, basal appendix of R2+3; aem, anepimeron; aes, anepisternum; c, coxa; f, femur; h, halter; hb, hind basitarsus; kem, katepimeron; kes, katepisternum; m, meron; mtaes, metanepisternum; mtg, mediotergite; mtkes, metakatepisternum; ltg, laterotergite; pltrch, pleu- rotrochantin; prn, pronotum; sct, scutum; t, trochanter." figureDoi="http://doi.org/10.5281/zenodo.4529518" httpUri="https://zenodo.org/record/4529518/files/figure.png" pageId="6" pageNumber="457">Fig. 2b</figureCitation>
). Three branches of M present (M 
<subScript id="B57A4E56FFBB4B43FEA8FD1CFE8024C3" attach="left" box="[272,280,673,687]" fontSize="6" pageId="6" pageNumber="457">1</subScript>
, M 
<subScript id="B57A4E56FFBB4B43FE86FD1CFEDE24C3" attach="left" box="[318,326,673,687]" fontSize="6" pageId="6" pageNumber="457">2</subScript>
, and M 
<subScript id="B57A4E56FFBB4B43FE23FD1CFE3B24C3" attach="left" box="[411,419,673,687]" fontSize="6" pageId="6" pageNumber="457">4</subScript>
). Base of M 
<subScript id="B57A4E56FFBB4B43FD94FD1CFDAC24C3" attach="left" box="[556,564,673,687]" fontSize="6" pageId="6" pageNumber="457">4</subScript>
crossvein-like. CuA and CuP present, reaching wing margin, sometimes meeting distally forming closed cell cua. A 
<subScript id="B57A4E56FFBB4B43FDF1FD78FDC924BF" attach="left" box="[585,593,709,723]" fontSize="6" pageId="6" pageNumber="457">1</subScript>
short, weak; not extending beyond the small fold at the base of anal lobe. Abdomen: Black to dark brown, with short to elongate hairs. Male abdomen elongate, gradually tapered posteriorly; female abdomen much stouter. Male spiracles 1–7 located laterally on abdominal pleura, except spiracle eight, either absent or, if present, a remnant dorsolaterally at membrane between tergites eight and nine (e.g., 
<taxonomicName id="EEFE3790FFBB4B43FAACFCA2FA142555" box="[1300,1420,799,825]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="457" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFBB4B43FAACFCA2FA142555" box="[1300,1420,799,825]" italics="true" pageId="6" pageNumber="457">P. funebris</emphasis>
</taxonomicName>
). Males with tergites and sternites 1–8 unmodified, females with tergites 1–8 and sternites 1–7 unmodified. Male terminalia (
<figureCitation id="B1C55096FFBB4B43FEA0FCDAFE0925ED" box="[280,401,871,897]" captionStart-0="FIGURES 4–6" captionStart-1="FIGURES 7–8" captionStart-2="FIGURES 9–11" captionStartId-0="13.[151,264,1964,1988]" captionStartId-1="15.[151,264,1945,1969]" captionStartId-2="17.[151,264,1921,1945]" captionTargetBox-0="[387,1193,189,1934]" captionTargetBox-1="[361,1222,182,1917]" captionTargetBox-2="[390,1197,188,1889]" captionTargetId-0="figure-16@13.[387,1201,181,1940]" captionTargetId-1="figure-16@15.[361,1227,181,1921]" captionTargetId-2="figure-16@17.[383,1204,181,1896]" captionTargetPageId-0="13" captionTargetPageId-1="15" captionTargetPageId-2="17" captionText-0="FIGURES 4–6. P. appendicula, male terminalia. 4. Dorsal. 5. Posterodorsal. 6. Ventral. Scale bar = ca. 0.5 mm. Abbreviations: gc, gonocoxite; gs, gonostylus; p, paramere; sls, spine-like setae; t9, tergite nine." captionText-1="FIGURES 7–8. P. arizonensis n. sp., holotype, male terminalia. 7. Dorsal. 8. Ventral. Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine." captionText-2="FIGURES 9–11. P. distincta, male terminalia. 9. Dorsal. 10. Posterior. 11. Ventral. Scale bar = ca. 0.5 mm. Abbreviations: gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vmc, ventromedian cleft of gonocoxite; vml, ventromedian lobes of gonocoxite." figureDoi-0="http://doi.org/10.5281/zenodo.4529520" figureDoi-1="http://doi.org/10.5281/zenodo.4529522" figureDoi-2="http://doi.org/10.5281/zenodo.4529526" httpUri-0="https://zenodo.org/record/4529520/files/figure.png" httpUri-1="https://zenodo.org/record/4529522/files/figure.png" httpUri-2="https://zenodo.org/record/4529526/files/figure.png" pageId="6" pageNumber="457">Figs. 4–11</figureCitation>
, 
<figureCitation id="B1C55096FFBB4B43FE24FCDAFE7D25ED" box="[412,485,871,897]" captionStart-0="FIGURES 14–15" captionStart-1="FIGURES 16–20" captionStartId-0="20.[151,264,1956,1980]" captionStartId-1="23.[151,264,1891,1915]" captionTargetBox-0="[389,1201,188,1928]" captionTargetBox-1="[326,1254,186,1861]" captionTargetId-0="figure-34@20.[382,1205,181,1932]" captionTargetId-1="figure-16@23.[325,1262,181,1866]" captionTargetPageId-0="20" captionTargetPageId-1="23" captionText-0="FIGURES 14–15. P. dolichopeza n. sp., holotype, male terminalia. 14. Dorsal. 15. Ventral. Scale bar = ca. 0.5 mm. Abbreviations: c, cerci; dal, dorsoapical lobe of gonocoxite; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine." captionText-1="FIGURES 16–20. P. heteroptera, male terminalia. 16. Dorsal, topotypic (Maryland, USA). 17. Gonostylus, dorsal (Texas, USA). 18. Gonostylus, ventral (Virginia, USA). 19. Gonostylus, dorsal (Vermont, USA). 20. Ventral, topotypic (Maryland, USA). Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vmc, ventromedian cleft of gonocoxite; vml, ventromedian lobes of gonocoxite." figureDoi-0="http://doi.org/10.5281/zenodo.4529530" figureDoi-1="http://doi.org/10.5281/zenodo.4529536" httpUri-0="https://zenodo.org/record/4529530/files/figure.png" httpUri-1="https://zenodo.org/record/4529536/files/figure.png" pageId="6" pageNumber="457">14–20</figureCitation>
, 
<figureCitation id="B1C55096FFBB4B43FE48FCDAFDA125ED" box="[496,569,871,897]" captionStart="FIGURES 24–29" captionStartId="27.[151,264,1849,1873]" captionTargetBox="[191,1399,188,1815]" captionTargetId="figure-16@27.[186,1402,181,1825]" captionTargetPageId="27" captionText="FIGURES 24–29. P. mexicana, male terminalia. 24–27. Topotypic (Guadalajara, Mexico). 24. Dorsal. 25. Ventral. 26. Dorsal, tergite nine removed. 27. Posterior. 28–29. Puerta Parada, Guatemala. 28. Dorsal, tergite nine removed. 29. Dorsal, tergite nine removed. Scale bar = ca. 0.5 mm. Abbreviations: c, cerci; dlp, divergent lobes of paramere; ea, ejaculatory apodeme; ed, ejaculatory duct; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vml, ventromedian lobes of gonocoxite." figureDoi="http://doi.org/10.5281/zenodo.4529544" httpUri="https://zenodo.org/record/4529544/files/figure.png" pageId="6" pageNumber="457">24–29</figureCitation>
, 
<figureCitation id="B1C55096FFBB4B43FDFCFCDBFD1725ED" box="[580,655,870,897]" captionStart-0="FIGURES 33–35" captionStart-1="FIGURES 38–41" captionStart-2="FIGURES 42–45" captionStartId-0="30.[151,264,1887,1911]" captionStartId-1="33.[151,264,1888,1912]" captionStartId-2="34.[151,264,1889,1913]" captionTargetBox-0="[425,1163,186,1851]" captionTargetBox-1="[168,1417,924,1848]" captionTargetBox-2="[167,1419,240,1859]" captionTargetId-0="figure-34@30.[420,1168,181,1863]" captionTargetId-1="figure-337@33.[164,1424,921,1864]" captionTargetId-2="figure-64@34.[177,1410,666,1865]" captionTargetPageId-0="30" captionTargetPageId-1="33" captionTargetPageId-2="34" captionText-0="FIGURES 33–35. P. neonigrita n. sp., male terminalia. 33. Dorsal. 34. Ventral (34 inset, paratype, ventral, not to scale; differences in length of gonocoxite between 34 and 34 inset due to distortion from slide mounting). 35. Posterior. Scale bar = ca. 0.5 mm. Abbreviations: c, cerci; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vml, ventromedian lobes of gonocoxite." captionText-1="FIGURES 38–41. Male terminalia. 38. P. heteroptera, dorsal, tergite nine removed, topotypic (Maryland, USA). 39. P. yakima n. sp., holotype, dorsal, tergite nine removed. 40. P. heteroptera, posterior, topotypic (Maryland, USA). 41. P. yakima n. sp., holotype, posterior. Scale bar = ca. 0.5 mm.Abbreviations: c, cerci; ea, ejaculatory apodeme; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine." captionText-2="FIGURES 42–45. Male terminalia, dorsal, tergite nine removed. 42. P. distincta. 43. P. dolichopeza n. sp., paratype. 44. P. appendicula. 45. P. neonigrita n. sp., paratype. Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; ea, ejaculatory apodeme; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; vml, ventromedian lobes of gonocoxite." figureDoi-0="http://doi.org/10.5281/zenodo.4529550" figureDoi-1="http://doi.org/10.5281/zenodo.4529554" figureDoi-2="http://doi.org/10.5281/zenodo.4529558" httpUri-0="https://zenodo.org/record/4529550/files/figure.png" httpUri-1="https://zenodo.org/record/4529554/files/figure.png" httpUri-2="https://zenodo.org/record/4529558/files/figure.png" pageId="6" pageNumber="457">33–45</figureCitation>
): Terminalia slightly dorsoflexed, not rotated. Posterior margin of tergite 9 (epandrium) shallowly to deeply (nearly dividing tergite) emarginated (“epandrial cleft”) medially. Epandrial cleft often U-shaped, with lobes at sides of posterior edge of epandrium (epandrial lobes) typically broad, rounded, sometimes more triangular in shape apically. Anterior edge of epandrium sometimes with a shallow to moderate medial emargination. Gonocoxites fused to sternite 9 (hypandrium) into a continuous genital capsule (synsternogonocoxite). Hypandrium distinguishable only by narrow, strap-like thickening ventrally and sometimes a slight medial seam. Posteromedian margin of synsternogonocoxite sometimes with median hump, a median cleft (ventromedian cleft, 
<figureCitation id="B1C55096FFBB4B43FEB6FBDEFEF92211" box="[270,353,1123,1149]" captionStart="FIGURES 16–20" captionStartId="23.[151,264,1891,1915]" captionTargetBox="[326,1254,186,1861]" captionTargetId="figure-16@23.[325,1262,181,1866]" captionTargetPageId="23" captionText="FIGURES 16–20. P. heteroptera, male terminalia. 16. Dorsal, topotypic (Maryland, USA). 17. Gonostylus, dorsal (Texas, USA). 18. Gonostylus, ventral (Virginia, USA). 19. Gonostylus, dorsal (Vermont, USA). 20. Ventral, topotypic (Maryland, USA). Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vmc, ventromedian cleft of gonocoxite; vml, ventromedian lobes of gonocoxite." figureDoi="http://doi.org/10.5281/zenodo.4529536" httpUri="https://zenodo.org/record/4529536/files/figure.png" pageId="6" pageNumber="457">Fig. 20</figureCitation>
, vmc), and/or a pair of weak to strong lobes (ventromedian lobes, 
<figureCitation id="B1C55096FFBB4B43FBF2FBDEFB3C2211" box="[1098,1188,1123,1149]" captionStart="FIGURES 16–20" captionStartId="23.[151,264,1891,1915]" captionTargetBox="[326,1254,186,1861]" captionTargetId="figure-16@23.[325,1262,181,1866]" captionTargetPageId="23" captionText="FIGURES 16–20. P. heteroptera, male terminalia. 16. Dorsal, topotypic (Maryland, USA). 17. Gonostylus, dorsal (Texas, USA). 18. Gonostylus, ventral (Virginia, USA). 19. Gonostylus, dorsal (Vermont, USA). 20. Ventral, topotypic (Maryland, USA). Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vmc, ventromedian cleft of gonocoxite; vml, ventromedian lobes of gonocoxite." figureDoi="http://doi.org/10.5281/zenodo.4529536" httpUri="https://zenodo.org/record/4529536/files/figure.png" pageId="6" pageNumber="457">Figs. 20</figureCitation>
&amp; 
<figureCitation id="B1C55096FFBB4B43FB70FBDEFB7E2211" box="[1224,1254,1123,1149]" captionStart="FIGURES 33–35" captionStartId="30.[151,264,1887,1911]" captionTargetBox="[425,1163,186,1851]" captionTargetId="figure-34@30.[420,1168,181,1863]" captionTargetPageId="30" captionText="FIGURES 33–35. P. neonigrita n. sp., male terminalia. 33. Dorsal. 34. Ventral (34 inset, paratype, ventral, not to scale; differences in length of gonocoxite between 34 and 34 inset due to distortion from slide mounting). 35. Posterior. Scale bar = ca. 0.5 mm. Abbreviations: c, cerci; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vml, ventromedian lobes of gonocoxite." figureDoi="http://doi.org/10.5281/zenodo.4529550" httpUri="https://zenodo.org/record/4529550/files/figure.png" pageId="6" pageNumber="457">34</figureCitation>
, vml) which are membranous to lightly sclerotized. Gonocoxites distally with elongate, tubular pedicel to which gonostyli articulate apically. Gonostylus usually rather simple, short, stout, robust, straight to very strongly arched, gradually tapered to slightly expanded apically, apex rounded, truncate, or with obtuse point, sometimes with a small anterior tooth. Proctiger present ventral to epandrium; cerci normally protruding through epandrial cleft. Tergite and sternite 10 (epiproct and hypoproct respectively) present. Cerci rounded, flap-like, fleshy, with hairs (Figs. 36, 40). Gonocoxal apodeme present, fused to parameres (
<figureCitation id="B1C55096FFBB4B43FDF2FA86FD382339" box="[586,672,1339,1365]" captionStart="FIGURES 38–41" captionStartId="33.[151,264,1888,1912]" captionTargetBox="[168,1417,924,1848]" captionTargetId="figure-337@33.[164,1424,921,1864]" captionTargetPageId="33" captionText="FIGURES 38–41. Male terminalia. 38. P. heteroptera, dorsal, tergite nine removed, topotypic (Maryland, USA). 39. P. yakima n. sp., holotype, dorsal, tergite nine removed. 40. P. heteroptera, posterior, topotypic (Maryland, USA). 41. P. yakima n. sp., holotype, posterior. Scale bar = ca. 0.5 mm.Abbreviations: c, cerci; ea, ejaculatory apodeme; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine." figureDoi="http://doi.org/10.5281/zenodo.4529554" httpUri="https://zenodo.org/record/4529554/files/figure.png" pageId="6" pageNumber="457">Fig. 38</figureCitation>
). Ventral to proctiger, parameres have a complex three-dimensional shape; dorsal sclerite and ventrolateral apodemes of paramere indistinguishably fused into a posteriorly broadly rounded, dome-like, hood (
<figureCitation id="B1C55096FFBB4B43FE7CFA3EFDD323F1" box="[452,587,1410,1437]" captionStart-0="FIGURES 38–41" captionStart-1="FIGURES 42–45" captionStartId-0="33.[151,264,1888,1912]" captionStartId-1="34.[151,264,1889,1913]" captionTargetBox-0="[168,1417,924,1848]" captionTargetBox-1="[167,1419,240,1859]" captionTargetId-0="figure-337@33.[164,1424,921,1864]" captionTargetId-1="figure-64@34.[177,1410,666,1865]" captionTargetPageId-0="33" captionTargetPageId-1="34" captionText-0="FIGURES 38–41. Male terminalia. 38. P. heteroptera, dorsal, tergite nine removed, topotypic (Maryland, USA). 39. P. yakima n. sp., holotype, dorsal, tergite nine removed. 40. P. heteroptera, posterior, topotypic (Maryland, USA). 41. P. yakima n. sp., holotype, posterior. Scale bar = ca. 0.5 mm.Abbreviations: c, cerci; ea, ejaculatory apodeme; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine." captionText-1="FIGURES 42–45. Male terminalia, dorsal, tergite nine removed. 42. P. distincta. 43. P. dolichopeza n. sp., paratype. 44. P. appendicula. 45. P. neonigrita n. sp., paratype. Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; ea, ejaculatory apodeme; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; vml, ventromedian lobes of gonocoxite." figureDoi-0="http://doi.org/10.5281/zenodo.4529554" figureDoi-1="http://doi.org/10.5281/zenodo.4529558" httpUri-0="https://zenodo.org/record/4529554/files/figure.png" httpUri-1="https://zenodo.org/record/4529558/files/figure.png" pageId="6" pageNumber="457">Figs. 38–45</figureCitation>
, p). Ventral to parameres, membranous, sac-like, endophallus present, which is cradled by more ventral ejaculatory apodeme (
<figureCitation id="B1C55096FFBB4B43FD0CFA1AFCA323AD" box="[692,827,1447,1473]" captionStart="FIGURES 42–45" captionStartId="34.[151,264,1889,1913]" captionTargetBox="[167,1419,240,1859]" captionTargetId="figure-64@34.[177,1410,666,1865]" captionTargetPageId="34" captionText="FIGURES 42–45. Male terminalia, dorsal, tergite nine removed. 42. P. distincta. 43. P. dolichopeza n. sp., paratype. 44. P. appendicula. 45. P. neonigrita n. sp., paratype. Scale bar = ca. 0.5 mm. Abbreviations: dlp, divergent lobes of paramere; ea, ejaculatory apodeme; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; vml, ventromedian lobes of gonocoxite." figureDoi="http://doi.org/10.5281/zenodo.4529558" httpUri="https://zenodo.org/record/4529558/files/figure.png" pageId="6" pageNumber="457">Figs. 42–43</figureCitation>
, ea). Ejaculatory apodeme, simple, dorsoventrally flattened. A sclerotized aedeagus absent. Female terminalia (
<figureCitation id="B1C55096FFBB4B43FCABFA76FC052389" box="[787,925,1482,1509]" captionStart-0="FIGURES 46–49" captionStart-1="FIGURES 50–53" captionStart-2="FIGURES 54–57" captionStartId-0="35.[151,264,1874,1898]" captionStartId-1="36.[151,264,1913,1937]" captionStartId-2="37.[151,264,1708,1732]" captionTargetBox-0="[165,1428,204,1842]" captionTargetBox-1="[188,1404,201,1883]" captionTargetBox-2="[168,1421,209,1673]" captionTargetId-0="figure-16@35.[156,1432,181,1851]" captionTargetId-1="figure-34@36.[182,1404,181,1890]" captionTargetId-2="figure-16@37.[164,1424,181,1684]" captionTargetPageId-0="35" captionTargetPageId-1="36" captionTargetPageId-2="37" captionText-0="FIGURES 46–49. Female terminalia. 46. P. appendicula, dorsal. 47. P. appendicula, ventral. 48. P. distincta, ventral. 49. P. mexicana, ventral. Scale bars = ca. 0.25 mm. Abbreviations: c1, cercus 1; c2, cercus 2; s7, sternite seven; s8, sternite eight; s10, sternite ten; sp, spermatheca; t7, tergite seven; t8, tergite eight; t9, tergite nine." captionText-1="FIGURES 50–53. Female terminalia. 50. P. heteroptera, dorsal. 51. P. heteroptera, ventral. 52. P. arizonensis n. sp., ventral. 53. P. arizonensis n. sp., lateral. Scale bars = ca. 0.25 mm. Abbreviations: c1, cercus 1; c2, cercus 2; gf, genital fork; s7, sternite seven; s8, sternite eight; s10, sternite ten; sp, spermatheca; t7, tergite seven; t8, tergite eight; t9, tergite nine." captionText-2="FIGURES 54–57. Female terminalia and eggs. 54–56. Female terminalia. 54. P. neonigrita n. sp., dorsal. 55. P. neonigrita n. sp., ventral. 56. P. yakima n. sp., ventral. 57. P. heteroptera, eggs (dissected from abdomen). Scale bars = ca. 0.25 mm. Abbreviations: c1, cercus 1; c2, cercus 2; s7, sternite seven; s8, sternite eight; s10, sternite ten; sp, spermatheca; t7, tergite seven; t8, tergite eight; t9, tergite nine." figureDoi-0="http://doi.org/10.5281/zenodo.4529560" figureDoi-1="http://doi.org/10.5281/zenodo.4529564" figureDoi-2="http://doi.org/10.5281/zenodo.4529566" httpUri-0="https://zenodo.org/record/4529560/files/figure.png" httpUri-1="https://zenodo.org/record/4529564/files/figure.png" httpUri-2="https://zenodo.org/record/4529566/files/figure.png" pageId="6" pageNumber="457">Figs. 46–56</figureCitation>
): Tergite 9 present as narrow, transverse strap, or subdivided into two sclerites; often more strongly produced ventrolaterally. Tergite 10 minute, longitudinally elongate. Cerci two-segmented. Subgenital plate (sternite eight) large, divided longitudinally, with posterior margin lobate; inner margins of longitudinal cleft sometimes with minute, posteromedially-directed tubercles. Y-shaped genital fork (= sternite 9) present (
<figureCitation id="B1C55096FFBB4B43FD99F9E6FDEC2019" box="[545,628,1626,1653]" captionStart="FIGURES 50–53" captionStartId="36.[151,264,1913,1937]" captionTargetBox="[188,1404,201,1883]" captionTargetId="figure-34@36.[182,1404,181,1890]" captionTargetPageId="36" captionText="FIGURES 50–53. Female terminalia. 50. P. heteroptera, dorsal. 51. P. heteroptera, ventral. 52. P. arizonensis n. sp., ventral. 53. P. arizonensis n. sp., lateral. Scale bars = ca. 0.25 mm. Abbreviations: c1, cercus 1; c2, cercus 2; gf, genital fork; s7, sternite seven; s8, sternite eight; s10, sternite ten; sp, spermatheca; t7, tergite seven; t8, tergite eight; t9, tergite nine." figureDoi="http://doi.org/10.5281/zenodo.4529564" httpUri="https://zenodo.org/record/4529564/files/figure.png" pageId="6" pageNumber="457">Fig. 51</figureCitation>
, gf). In addition to genital fork, a minute pair of sclerites often present between the posterior margin of the subgenital plate and the anterior margin of sternite ten. Sternite 10 present. Three rounded, sclerotized, balloon-like, spermathecae present.
</paragraph>
<paragraph id="29414C13FFBB4B43FF7FF97AFB93208D" blockId="6.[151,1437,151,2013]" box="[199,1035,1734,1761]" pageId="6" pageNumber="457">
<emphasis id="1B8A9001FFBB4B43FF7FF97AFE18208C" box="[199,384,1735,1760]" italics="true" pageId="6" pageNumber="457">Immature stages</emphasis>
: 
<emphasis id="1B8A9001FFBB4B43FE37F97AFE24208C" box="[399,444,1735,1760]" italics="true" pageId="6" pageNumber="457">Egg</emphasis>
(
<figureCitation id="B1C55096FFBB4B43FE73F97AFDB8208D" box="[459,544,1734,1761]" captionStart="FIGURES 54–57" captionStartId="37.[151,264,1708,1732]" captionTargetBox="[168,1421,209,1673]" captionTargetId="figure-16@37.[164,1424,181,1684]" captionTargetPageId="37" captionText="FIGURES 54–57. Female terminalia and eggs. 54–56. Female terminalia. 54. P. neonigrita n. sp., dorsal. 55. P. neonigrita n. sp., ventral. 56. P. yakima n. sp., ventral. 57. P. heteroptera, eggs (dissected from abdomen). Scale bars = ca. 0.25 mm. Abbreviations: c1, cercus 1; c2, cercus 2; s7, sternite seven; s8, sternite eight; s10, sternite ten; sp, spermatheca; t7, tergite seven; t8, tergite eight; t9, tergite nine." figureDoi="http://doi.org/10.5281/zenodo.4529566" httpUri="https://zenodo.org/record/4529566/files/figure.png" pageId="6" pageNumber="457">Fig. 57</figureCitation>
): Elongate, sausage-shaped, cream-colored.
</paragraph>
<paragraph id="29414C13FFBB4B42FF7FF956FC602285" blockId="6.[151,1437,151,2013]" lastBlockId="7.[151,1437,151,2013]" lastPageId="7" lastPageNumber="458" pageId="6" pageNumber="457">
<emphasis id="1B8A9001FFBB4B43FF7FF956FE922168" box="[199,266,1771,1796]" italics="true" pageId="6" pageNumber="457">Larva</emphasis>
(
<figureCitation id="B1C55096FFBB4B43FEA6F956FEEE2169" box="[286,374,1770,1797]" captionStart="FIGURES 58–60" captionStartId="38.[151,264,1892,1916]" captionTargetBox="[161,1427,186,1866]" captionTargetId="figure-34@38.[160,1428,181,1868]" captionTargetPageId="38" captionText="FIGURES 58–60. P. funebris. 58. Larva, lateral view (head, at left, not visible; telescoped within thorax). 59. Pupa, dorsal. 60. Pupa, ventral. Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529568" httpUri="https://zenodo.org/record/4529568/files/figure.png" pageId="6" pageNumber="457">Fig. 58</figureCitation>
): Body light brownish to gray, elongate, slightly dorsoventrally flattened, leathery, slightly curved downwards in lateral view, with three thoracic and nine abdominal segments. Thorax and abdomen with transverse rows of elongate, fleshy tubercles on each segment both dorsally and ventrally. Number of tubercles in each row varies depending on the species, though thoracic segments tend to have fewer tubercles. Laterally, two tubercles present near each abdominal spiracle except posterior spiracle. Cuticle with dark brown to black, minute, sclerotized, spine-like scales. Thorax without lightly sclerotized plates and without characteristic stair-step-like swollen developments observed in 
<taxonomicName id="EEFE3790FFBB4B43FD9EF87EFD5721B1" authorityName="Schiner" authorityYear="1864" box="[550,719,1987,2013]" class="Insecta" family="Pachyneuridae" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="457" phylum="Arthropoda" rank="family">Pachyneuridae</taxonomicName>
. Ventral flap-like tubercle on prothorax present. Anus terminal, anal papillae apparently absent. Intersegmental fissures unaligned between meso- and metathorax between abdominal segments 1 and 2 and between abdominal segments 7 and 8. Dorsally, prothorax with two pseudosegments, mesothorax two pseudosegments, and most abdominal segments three pseudosegments. Prothoracic and metathoracic spiracles present. Abdominal segments 1–8 with spiracles; spiracles 1–7 lateral in position, spiracle 8 dorsolateral, larger than previous ones. All spiracles slightly protuberant, especially posterior spiracles. Posterior spiracle on posterior border of segment 8, round, with single, central, ecdysial scar. Head strongly sclerotized, black to dark brown, rounded, somewhat dorsoventrally flattened, with setae. Head capable of being completely withdrawn into anterior portion of thorax. Ecdysial lines meeting in form of Y anterior to postoccipital carina. Anterolateral margin of frontoclypeus developed into strong, anteroventrally-directed spine. Labrum subrectangular, without strong spines at apex of labrum/epipharynx. Anteroventral stemmata present, dorsoposterior stemmata absent. Antennae short, sensory cone in form of ovoid plate positioned within larger, round, antennal socket. Postgenal bridge complete. Ventral prothoracic sclerites in cuticle at posterior, ventral margin of head capsule present. Anterior tentorial arm present, weakly developed, connected at anterior margin of head near mandibular articulation. Posterior tentorial bridge absent. Submentum apparently absent or indistinguishably fused to ventral head capsule. Anterior mentum (hypostoma) absent, posterior mentum present as a narrow, longitudinal sclerite, not fused to anterior margin of cranium or to posterior margin of labium. Labial synsclerite present, as an inverted U-shape; each anterolateral margin with small knob homologous to glossae. Posterior labial sclerite present, large, in same plane as and smoothly fused to labial synsclerite, forming sclerotized frame completely enclosing membranous labial area. Tubercle of labial palps absent, membranous labial area bearing field of papillae. Cibarial bar absent. Membranous, hypopharynx with patches of minute spines, supported by two pairs of hypopharyngeal sclerites: each pair with one large and one minute sclerite. Pharyngeal filter absent. Cardo large, transverse, not closely appressed to anteroventral margin of cranium, T-shaped, inner apex of sclerite with an anteriorly directed lobe and a posteriorly directed lobe. Cardo with four setae (some represented only by alveoli); one seta at inner apex and a group of three at outer (lateral) apex. Galeolacinia adjacent, but not closely appressed to palpifer. Galeolacinia sclerotized ventrally, with numerous teeth and spines on inner edge apically and on dorsal surface. Laterobasal sclerite of maxillary palpifer present. Palpifer sclerotized, tubular, bearing one-segmented cylindrical palpus. Palpus with single sensory region apically, bearing numerous short, stout setae. Mandible heavily sclerotized, subtriangular, with small number of short, stout, apical teeth. Mandible operating in horizontal plane, without line of weakness separating apical and basal portions, and lacking basal thumb of teeth. Prostheca present. Mandibular comb absent. Epipharynx dorsoventrally flattened, slightly bilobate, with numerous small inwardly directed spines and small number of peg-like setae. Torma wrapped dorsolaterally, fused and continuous with dorsal labrum. Premandible absent.
</paragraph>
<paragraph id="29414C13FFBA4B42FF7FFB4EFDA82315" blockId="7.[151,1437,151,2013]" pageId="7" pageNumber="458">
<emphasis id="1B8A9001FFBA4B42FF7FFB4EFE9A2360" box="[199,258,1267,1292]" italics="true" pageId="7" pageNumber="458">Pupa</emphasis>
(
<figureCitation id="B1C55096FFBA4B42FEAAFB4EFE052361" box="[274,413,1266,1293]" captionStart="FIGURES 58–60" captionStartId="38.[151,264,1892,1916]" captionTargetBox="[161,1427,186,1866]" captionTargetId="figure-34@38.[160,1428,181,1868]" captionTargetPageId="38" captionText="FIGURES 58–60. P. funebris. 58. Larva, lateral view (head, at left, not visible; telescoped within thorax). 59. Pupa, dorsal. 60. Pupa, ventral. Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529568" httpUri="https://zenodo.org/record/4529568/files/figure.png" pageId="7" pageNumber="458">Figs. 59–60</figureCitation>
): Brown to greyish in color, leathery, sometimes enclosed within last larval skin. Head and thorax without distinct setae or spines. Respiratory horn absent; anterior thoracic spiracle on short tubercle. Abdomen with fleshy tubercles, without sclerotized spines or setae. Abdominal tergites without transverse rows of minute spinules. Leg sheaths superimposed.
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFBA4B42FF7FFA3FFDE02065" pageId="7" pageNumber="458" type="distribution">
<paragraph id="29414C13FFBA4B42FF7FFA3FFDE02065" blockId="7.[151,1437,151,2013]" pageId="7" pageNumber="458">
<emphasis id="1B8A9001FFBA4B42FF7FFA3FFD0323F0" bold="true" box="[199,667,1410,1437]" pageId="7" pageNumber="458">Geographic Distribution and Diversity</emphasis>
. 
<taxonomicName id="EEFE3790FFBA4B42FD13FA3EFCB923F1" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[683,801,1411,1437]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FD13FA3EFCB923F1" box="[683,801,1411,1437]" italics="true" pageId="7" pageNumber="458">Penthetria</emphasis>
</taxonomicName>
presently includes 36 extant species distributed mainly in the Holarctic and Oriental regions, but also with species in the Neotropics. The greatest diversity is north of the equator, particularly in the Oriental and eastern Palearctic regions (28 species), with only eight species described from the Nearctic and Neotropical regions.
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFBA4B4DFF7FF9AFFBA92771" lastPageId="8" lastPageNumber="459" pageId="7" pageNumber="458" type="discussion">
<paragraph id="29414C13FFBA4B42FF7FF9AFFBFD21F9" blockId="7.[151,1437,151,2013]" pageId="7" pageNumber="458">
<emphasis id="1B8A9001FFBA4B42FF7FF9AFFEF52041" bold="true" box="[199,365,1554,1581]" pageId="7" pageNumber="458">Fossil Record</emphasis>
. Compression fossils are known from the Nearctic and Palearctic regions; most of these are Oligocene and Miocene in age, with a few from the Eocene and Pliocene/Pleistocene (
<bibRefCitation id="4D6F31E2FFBA4B42FBDDF98AFA8B203D" author="Evenhuis, N. L." box="[1125,1299,1591,1617]" pageId="7" pageNumber="458" refId="ref32873" refString="Evenhuis, N. L. (2014) Catalog of the fossil flies of the world (Insecta: Diptera). Version 2.0. Available from: http: // hbs. bishopmuseum. org / fossilcat / (accessed 30 March 2020)" type="url" year="2014">Evenhuis 2014</bibRefCitation>
). 
<taxonomicName id="EEFE3790FFBA4B42FA9EF98AFEA92019" authority="Geinitz" authorityName="Geinitz" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="species" species="dubia">
<emphasis id="1B8A9001FFBA4B42FA9EF98AFF4F2019" italics="true" pageId="7" pageNumber="458">Penthetria dubia</emphasis>
Geinitz
</taxonomicName>
was described from the Lower Jurassic (
<collectingCountry id="51E90C83FFBA4B42FCBCF9E6FCEA2019" box="[772,882,1627,1653]" name="Germany" pageId="7" pageNumber="458">Germany</collectingCountry>
), but based on the illustration of the wing of this species (
<bibRefCitation id="4D6F31E2FFBA4B42FF4FF9C2FE1620F5" author="Geinitz, F. E." box="[247,398,1663,1689]" pageId="7" pageNumber="458" pagination="566 - 583" refId="ref33569" refString="Geinitz, F. E. (1884) Uber die Fauna des Dobbertiner Lias. Zeitschrift der Deutschen Geologischen Gesellschaft, 36, 566 - 583." type="journal article" year="1884">Geinitz 1884</bibRefCitation>
, Plate XIII, 
<figureCitation id="B1C55096FFBA4B42FDA1F9C2FDF220F6" box="[537,618,1663,1690]" captionStart="FIGURES 24–29" captionStartId="27.[151,264,1849,1873]" captionTargetBox="[191,1399,188,1815]" captionTargetId="figure-16@27.[186,1402,181,1825]" captionTargetPageId="27" captionText="FIGURES 24–29. P. mexicana, male terminalia. 24–27. Topotypic (Guadalajara, Mexico). 24. Dorsal. 25. Ventral. 26. Dorsal, tergite nine removed. 27. Posterior. 28–29. Puerta Parada, Guatemala. 28. Dorsal, tergite nine removed. 29. Dorsal, tergite nine removed. Scale bar = ca. 0.5 mm. Abbreviations: c, cerci; dlp, divergent lobes of paramere; ea, ejaculatory apodeme; ed, ejaculatory duct; ga, gonocoxal apodeme; gc, gonocoxite; gs, gonostylus; p, paramere; t9, tergite nine; vml, ventromedian lobes of gonocoxite." figureDoi="http://doi.org/10.5281/zenodo.4529544" httpUri="https://zenodo.org/record/4529544/files/figure.png" pageId="7" pageNumber="458">Fig. 26</figureCitation>
) it is not a 
<taxonomicName id="EEFE3790FFBA4B42FD53F9C2FCF920F5" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[747,865,1663,1689]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FD53F9C2FCF920F5" box="[747,865,1663,1689]" italics="true" pageId="7" pageNumber="458">Penthetria</emphasis>
</taxonomicName>
nor a 
<taxonomicName id="EEFE3790FFBA4B42FC13F9C2FBBE20F5" box="[939,1062,1663,1689]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="family">Bibionidae</taxonomicName>
and should be removed from the family, thus making the oldest compression fossils of this genus Eocene in age (e.g., 
<bibRefCitation id="4D6F31E2FFBA4B42FB82F91EFB3720D1" author="Rice, H. M. A." box="[1082,1199,1698,1725]" pageId="7" pageNumber="458" pagination="1 - 37" refId="ref36064" refString="Rice, H. M. A. (1959) Fossil Bibionidae (Diptera) from British Columbia. Bulletin of the Geological Survey of Canada, 55, 1 - 37, 4 pls. https: // doi. org / 10.4095 / 100564" type="journal article" year="1959">Rice 1959</bibRefCitation>
).Additionally, 
<bibRefCitation id="4D6F31E2FFBA4B42FAE0F91EFE8C208D" author="Skartveit, J." pageId="7" pageNumber="458" pagination="3 - 42" refId="ref36439" refString="Skartveit, J. (2009) Fossil Hesperinidae and Bibionidae from Baltic amber (Diptera: Bibionidae). Studia dipterologica, 15 (2008), 3 - 42." type="journal article" year="2009">Skartveit (2009)</bibRefCitation>
described two species from Baltic amber, which are also Eocene in age. One of these species is quite peculiar as vein R 
<subScript id="B57A4E56FFBA4B42FED8F940FEE12167" attach="left" box="[352,377,1789,1803]" fontSize="6" pageId="7" pageNumber="458">2+3</subScript>
is absent, but based on other features it was included in 
<taxonomicName id="EEFE3790FFBA4B42FC57F956FBFD2169" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1007,1125,1771,1797]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FC57F956FBFD2169" box="[1007,1125,1771,1797]" italics="true" pageId="7" pageNumber="458">Penthetria</emphasis>
</taxonomicName>
, where it seems to fit better than anywhere else in 
<taxonomicName id="EEFE3790FFBA4B42FE2DF8B2FD8C2145" box="[405,532,1807,1833]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="family">Bibionidae</taxonomicName>
. It is possible that this species is based on a specimen with a teratological wing, as in cases known in extant species of 
<taxonomicName id="EEFE3790FFBA4B42FDF6F88EFD0E2121" box="[590,662,1843,1869]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FDF6F88EFD0E2121" box="[590,662,1843,1869]" italics="true" pageId="7" pageNumber="458">Plecia</emphasis>
</taxonomicName>
(pers. obs.) and 
<taxonomicName id="EEFE3790FFBA4B42FCECF88EFB362121" authority="(Paramonov 2005)" baseAuthorityName="Paramonov" baseAuthorityYear="2005" box="[852,1198,1842,1869]" class="Insecta" family="Hesperinidae" genus="Hesperinus" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FCECF88EFC4C2120" box="[852,980,1843,1868]" italics="true" pageId="7" pageNumber="458">Hesperinus</emphasis>
(
<bibRefCitation id="4D6F31E2FFBA4B42FC5BF88EFB3E2121" author="Paramonov, N. M." box="[995,1190,1842,1869]" pageId="7" pageNumber="458" pagination="232" refId="ref35751" refString="Paramonov, N. M. (2005) First record of the family Hesperinidae (Diptera) from Sakhalin. Zoosystematica Rossica, 13 (2), 232. [2004]" type="journal article" year="2005">Paramonov 2005</bibRefCitation>
)
</taxonomicName>
where R 
<subScript id="B57A4E56FFBA4B42FAADF8F8FAB6213F" attach="left" box="[1301,1326,1861,1875]" fontSize="6" pageId="7" pageNumber="458">2+3</subScript>
is absent. It is noteworthy that some of the Mesozoic stem-bibionids currently placed in 
<taxonomicName id="EEFE3790FFBA4B42FBACF8EAFB2E211D" authorityName="Rohdendorf" authorityYear="1946" box="[1044,1206,1879,1905]" class="Insecta" family="Protopleciidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="family">Protopleciidae</taxonomicName>
have wing venation identical to 
<taxonomicName id="EEFE3790FFBA4B42FEA3F8C6FE0921F9" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[283,401,1915,1941]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FEA3F8C6FE0921F9" box="[283,401,1915,1941]" italics="true" pageId="7" pageNumber="458">Penthetria</emphasis>
</taxonomicName>
(e.g., 
<taxonomicName id="EEFE3790FFBA4B42FE60F8C6FC8A21F9" authority="Zhang, 2007" authorityName="Zhang" authorityYear="2007" box="[472,786,1915,1941]" class="Insecta" family="Protopleciidae" genus="Epimesoplecia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FE60F8C6FDE521F9" box="[472,637,1915,1941]" italics="true" pageId="7" pageNumber="458">Epimesoplecia</emphasis>
<bibRefCitation id="4D6F31E2FFBA4B42FD3CF8C6FC8A21F9" author="Zhang, J. F." box="[644,786,1915,1941]" pageId="7" pageNumber="458" pagination="289 - 296" refId="ref37500" refString="Zhang, J. F. (2007) New Mesozoic Protopleciidae (Insecta: Diptera: Nematocera) from China. Cretaceous Research, 2007, 289 - 296. https: // doi. org / 10.1016 / j. cretres. 2006.05.009" type="journal article" year="2007">Zhang, 2007</bibRefCitation>
</taxonomicName>
and 
<taxonomicName id="EEFE3790FFBA4B42FCF0F8C6FBC021F9" authority="Ren, 1995" authorityName="Ren" authorityYear="1995" box="[840,1112,1914,1941]" class="Insecta" family="Bibionidae" genus="Lichnoplecia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FCF0F8C6FC4221F9" box="[840,986,1915,1941]" italics="true" pageId="7" pageNumber="458">Lichnoplecia</emphasis>
<bibRefCitation id="4D6F31E2FFBA4B42FC59F8C6FBC021F9" author="Ren, D." box="[993,1112,1914,1941]" pageId="7" pageNumber="458" pagination="47 - 121" refId="ref35999" refString="Ren, D. (1995) Insecta. In: Ren, D., Lu, L. W., Guo, Z. G. &amp; Ji, S. A. (Eds.), Faunae and stratigraphy of Jurassic-Cretaceous in Beijing and the adjacent areas. Seismic Publishing House, Beijing, pp. 47 - 121." type="book chapter" year="1995">Ren, 1995</bibRefCitation>
</taxonomicName>
).
</paragraph>
<paragraph id="29414C13FFBA4B4DFF7FF822FBA92771" blockId="7.[151,1437,151,2013]" lastBlockId="8.[151,1437,150,2014]" lastPageId="8" lastPageNumber="459" pageId="7" pageNumber="458">
There are nine fossil species from the Nearctic region currently placed in 
<taxonomicName id="EEFE3790FFBA4B42FBBFF822FBE521D5" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1031,1149,1951,1977]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFBA4B42FBBFF822FBE521D5" box="[1031,1149,1951,1977]" italics="true" pageId="7" pageNumber="458">Penthetria</emphasis>
</taxonomicName>
(
<bibRefCitation id="4D6F31E2FFBA4B42FB34F822FAA121D5" author="Evenhuis, N. L." box="[1164,1337,1951,1977]" pageId="7" pageNumber="458" refId="ref32810" refString="Evenhuis, N. L. (1994) Catalog of the fossil flies of the world (Insecta: Diptera). Backhuys Publishers, Leiden, 600 pp." type="book" year="1994">Evenhuis 1994</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFBA4B42FAFCF822FF6521B1" author="Fitzgerald, S. J." pageId="7" pageNumber="458" pagination="182 - 187" refId="ref33069" refString="Fitzgerald, S. J. (1999) A new species of Plecia from the Green River Formation and new combinations of fossil Bibionidae (Diptera). Great Basin Naturalist, 59 (2), 182 - 187. https: // doi. org / 10.5962 / bhl. part. 15821" type="journal article" year="1999">Fitzgerald 1999</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFBA4B42FEB1F87EFE2B21B1" author="Skartveit, J." box="[265,435,1987,2013]" pageId="7" pageNumber="458" pagination="3 - 42" refId="ref36439" refString="Skartveit, J. (2009) Fossil Hesperinidae and Bibionidae from Baltic amber (Diptera: Bibionidae). Studia dipterologica, 15 (2008), 3 - 42." type="journal article" year="2009">Skartveit 2009</bibRefCitation>
). However, as noted by 
<bibRefCitation id="4D6F31E2FFBA4B42FD7DF87EFCE621B1" author="Gentilini, G." box="[709,894,1987,2013]" pageId="7" pageNumber="458" pagination="62 - 66" refId="ref33640" refString="Gentilini, G. (1991) Occurrence of the genus Penthetria Meigen, 1803 from the lower Messinian of central Italy. Bollettino della Societa Entomologica Italiana, 123 (1), 62 - 66." type="journal article" year="1991">Gentilini (1991)</bibRefCitation>
, many of the Nearctic fossils described by 
<bibRefCitation id="4D6F31E2FFBA4B4DFAD1F87EFF7A26DD" author="Rice, H. M. A." lastPageId="8" lastPageNumber="459" pageId="7" pageNumber="458" pagination="1 - 37" refId="ref36064" refString="Rice, H. M. A. (1959) Fossil Bibionidae (Diptera) from British Columbia. Bulletin of the Geological Survey of Canada, 55, 1 - 37, 4 pls. https: // doi. org / 10.4095 / 100564" type="journal article" year="1959">Rice (1959)</bibRefCitation>
in 
<taxonomicName id="EEFE3790FFB54B4DFEBEFF2AFED626DD" box="[262,334,151,177]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFEBEFF2AFED626DD" box="[262,334,151,177]" italics="true" pageId="8" pageNumber="459">Plecia</emphasis>
</taxonomicName>
may actually belong to 
<taxonomicName id="EEFE3790FFB54B4DFDE4FF2AFD4A26DD" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[604,722,151,177]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFDE4FF2AFD4A26DD" box="[604,722,151,177]" italics="true" pageId="8" pageNumber="459">Penthetria</emphasis>
</taxonomicName>
and need to be re-evaluated. The number of fossil species from the Palearctic (about 35) is also in a state of flux, as there has been significant recent work re-evaluating the generic placement of old species of 
<taxonomicName id="EEFE3790FFB54B4DFE61FF62FDD72695" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[473,591,223,249]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFE61FF62FDD72695" box="[473,591,223,249]" italics="true" pageId="8" pageNumber="459">Penthetria</emphasis>
</taxonomicName>
and 
<taxonomicName id="EEFE3790FFB54B4DFD31FF62FD492695" box="[649,721,223,249]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFD31FF62FD492695" box="[649,721,223,249]" italics="true" pageId="8" pageNumber="459">Plecia</emphasis>
</taxonomicName>
and adding new species (e.g., 
<bibRefCitation id="4D6F31E2FFB54B4DFB8AFF62FB432695" author="Skartveit, J." box="[1074,1243,223,249]" pageId="8" pageNumber="459" pagination="3 - 42" refId="ref36439" refString="Skartveit, J. (2009) Fossil Hesperinidae and Bibionidae from Baltic amber (Diptera: Bibionidae). Studia dipterologica, 15 (2008), 3 - 42." type="journal article" year="2009">Skartveit 2009</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFB5FFF62FF4A2771" author="Skartveit, J. &amp; Nel, A." pageId="8" pageNumber="459" pagination="51 - 63" refId="ref36636" refString="Skartveit, J. &amp; Nel, A. (2012) Fossil Bibionidae (Diptera: Bibionomorpha) from the late Oligocene of Bes-Konak, Anatolia, Turkey. Zootaxa, 3329 (1), 51 - 63. https: // doi. org / 10.11646 / zootaxa. 3329.1.5" type="journal article" year="2012">Skartveit &amp; Nel 2012</bibRefCitation>
, Skartveit &amp; Pica 2014, 
<bibRefCitation id="4D6F31E2FFB54B4DFE5BFEBEFD4F2771" author="Skartveit, J. &amp; Nel, A." box="[483,727,259,285]" pageId="8" pageNumber="459" pagination="1 - 83" refId="ref36689" refString="Skartveit, J. &amp; Nel, A. (2017) Revision of fossil Bibionidae (Insecta: Diptera) from French Oligocene deposits. Zootaxa, 4225 (1), 1 - 83. https: // doi. org / 10.11646 / zootaxa. 4225.1.1" type="journal article" year="2017">Skartveit &amp; Nel 2017</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFD5AFEBEFBBD2771" author="Skartveit, J. &amp; Krizmanic, K." box="[738,1061,259,285]" pageId="8" pageNumber="459" pagination="351 - 378" refId="ref36584" refString="Skartveit, J. &amp; Krizmanic, K. (2020) Revision of fossil Bibionidae (Insecta: Diptera) from the Miocene of Radoboj, Croatia. Zootaxa, 4759 (3), 351 - 378. https: // doi. org / 10.11646 / zootaxa. 4759.3.3" type="journal article" year="2020">Skartveit &amp; Krizmanic 2020</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFB54B4DFF7FFE9BFC7D2145" pageId="8" pageNumber="459" type="biology_ecology">
<paragraph id="29414C13FFB54B4DFF7FFE9BFBC125A6" blockId="8.[151,1437,150,2014]" pageId="8" pageNumber="459">
<emphasis id="1B8A9001FFB54B4DFF7FFE9BFEDB272C" bold="true" box="[199,323,294,320]" pageId="8" pageNumber="459">Bionomics</emphasis>
. Little is known about the biology of the adults of this genus, but it is assumed to be similar to other members of the family (summaries of biology can be found, e.g., in 
<bibRefCitation id="4D6F31E2FFB54B4DFC2DFEF6FB842709" author="Hardy, D. E." box="[917,1052,331,357]" pageId="8" pageNumber="459" pagination="217 - 222" refId="ref34196" refString="Hardy, D. E. (1981) Bibionidae. In: McAlpine, J. F., Peterson, B. V., Shewell, G. E., Teskey, H. J., Vockeroth, J. R. &amp; Wood, D. M. (Eds.), Manual of Nearctic Diptera. Vol. 1. Canada Department of Agriculture Research Branch Monograph 27. Biosystematics Research Institute Ottawa, Ontario, pp. 217 - 222." type="book chapter" year="1981">Hardy 1981</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFB91FEF6FB482709" author="Skartveit, J." box="[1065,1232,331,357]" pageId="8" pageNumber="459" refId="ref36370" refString="Skartveit, J. (1997) [Chapter] 2.5. Family Bibionidae. In: Papp, L. &amp; Darvas, B. (Eds.), Contributions to a manual of Palaearctic Diptera (with special reference to flies of economic importance). Vol. 2. Nematocera and Lower Brachycera. Science Herald, Budapest, pp. 41 - 50." type="book" year="1997">Skartveit 1997</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFB63FEF6FA092709" author="Fitzgerald, S. J." box="[1243,1425,331,357]" pageId="8" pageNumber="459" refId="ref33226" refString="Fitzgerald, S. J. (2009) Bibionidae. In: Brown, B. V., Borkent, A., Cumming, J. M., Wood, D. M., Woodley, N. E. &amp; Zumbado, M. A. (Eds.), Manual of Central American Diptera: Vol. 1. NRC Research Press, Ottawa, Ontario, 714 pp." type="book" year="2009">Fitzgerald 2009</bibRefCitation>
). However, while there are numerous records of other bibionid genera (especially 
<taxonomicName id="EEFE3790FFB54B4DFB83FED2FB2027E6" authorityName="Newman" authorityYear="1834" box="[1083,1208,367,394]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="subFamily" subFamily="Bibioninae">Bibioninae</taxonomicName>
) being collected on flowers, the label data of none of the New World 
<taxonomicName id="EEFE3790FFB54B4DFD06FE2EFCAC27C1" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[702,820,403,429]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFD06FE2EFCAC27C1" box="[702,820,403,429]" italics="true" pageId="8" pageNumber="459">Penthetria</emphasis>
</taxonomicName>
specimens indicate that any were collected on flowers, which seems unusual (the author is unaware of any records of adult feeding in this genus). Seasonal activity varies between species and region (e.g., 
<tableCitation id="647C79A8FFB54B4DFDAEFE66FDF12799" box="[534,617,475,501]" captionStart="TABLE 1" captionStartId="34.[151,239,151,175]" captionText="TABLE 1. Summary of adult flight period (grey boxes indicate specimens collected during these months); Malaise trap data including more than one month have been excluded." pageId="8" pageNumber="459">Table 1</tableCitation>
). 
<bibRefCitation id="4D6F31E2FFB54B4DFD3BFE66FCB82799" author="Pecina, P." box="[643,800,474,501]" pageId="8" pageNumber="459" pagination="285 - 298" refId="ref35785" refString="Pecina, P. (1965) Bohemian March-flies (Diptera, Bibionidae) in The National Museum, Prague. Acta Faunistica Entomologica Musei Nationalis Pragae, 11 (108), 285 - 298." type="journal article" year="1965">Pecina (1965)</bibRefCitation>
states that adults of the Palearctic 
<taxonomicName id="EEFE3790FFB54B4DFB14FE66FABD2799" box="[1196,1317,475,501]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFB14FE66FABD2799" box="[1196,1317,475,501]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
emerge in mid-April and are gone by mid-May, though “they may be found singly later.” Curiously, 
<bibRefCitation id="4D6F31E2FFB54B4DFBC4FE42FAD62475" author="Frouz, J. &amp; Jedlicka, P. &amp; Simackova, H. &amp; Lhotakova, Z." box="[1148,1358,510,537]" pageId="8" pageNumber="459" pagination="21 - 27" refId="ref33496" refString="Frouz, J., Jedlicka, P., Simackova, H., Lhotakova, Z. (2015) The life cycle, population dynamics, and contribution to litter decomposition of Penthetria holosericea (Diptera: Bibionidae) in an alder forest. European Journal of Soil Biology, 71, 21 - 27. https: // doi. org / 10.1016 / j. ejsobi. 2015.10.002" type="journal article" year="2015">
Frouz 
<emphasis id="1B8A9001FFB54B4DFB7CFE42FB6C2475" box="[1220,1268,511,537]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. (2015)
</bibRefCitation>
, working on the same species in the same country but 50 years later, found peak emergence was in late May and early June, though this disparity may be due to differences in altitude or annual seasonal variables. At a similar latitude in the Nearctic region, 
<taxonomicName id="EEFE3790FFB54B4DFEC7FDD6FDF924E9" authority="(Say)" baseAuthorityName="Say" baseAuthorityYear="1823" box="[383,609,619,645]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="heteroptera">
<emphasis id="1B8A9001FFB54B4DFEC7FDD6FD8524E9" box="[383,541,619,645]" italics="true" pageId="8" pageNumber="459">P. heteroptera</emphasis>
(Say)
</taxonomicName>
is fall-emerging, rather than spring-emerging, but has a similarly narrow duration of adult activity, with most adult records from a two-month window of time. In contrast to these temperate species with narrow windows of adult activity, the Neotropical species 
<taxonomicName id="EEFE3790FFB54B4DFC65FD0EFB7F24A1" authority="Fitzgerald" authorityName="Fitzgerald" authorityYear="2021" box="[989,1255,691,717]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="neonigrita" status="sp. nov.">
<emphasis id="1B8A9001FFB54B4DFC65FD0EFBF624A0" box="[989,1134,691,716]" italics="true" pageId="8" pageNumber="459">P. neonigrita</emphasis>
Fitzgerald
</taxonomicName>
<emphasis id="1B8A9001FFB54B4DFB56FD0EFAB424A1" bold="true" box="[1262,1324,691,717]" pageId="8" pageNumber="459">
<taxonomicNameLabel id="00B92D7AFFB54B4DFB56FD0EFAB424A1" box="[1262,1324,691,717]" pageId="8" pageNumber="459" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
, has been recorded every month of the year except April. 
<bibRefCitation id="4D6F31E2FFB54B4DFD1FFD6AFCE8249D" author="Yuan, R. &amp; Li, S. &amp; Du, Q. &amp; Lei, C. &amp; Zhu, F." box="[679,880,726,753]" pageId="8" pageNumber="459" pagination="320 - 324" refId="ref37430" refString="Yuan, R., Li, S., Du, Q., Lei, C. &amp; Zhu, F. (2015) Immature stages description of March Fly, Penthetria japonica Wiedemann (Diptera: Bibionidae), from Hubei, China. Entomological News, 124 (5), 320 - 324. https: // doi. org / 10.3157 / 021.124.0503" type="journal article" year="2015">
Yuan 
<emphasis id="1B8A9001FFB54B4DFD5FFD6AFC80249D" box="[743,792,727,753]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. (2015)
</bibRefCitation>
reported that lab reared 
<taxonomicName id="EEFE3790FFB54B4DFB39FD6AFB66249C" authorityName="Wiedemann" authorityYear="1830" box="[1153,1278,727,752]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="japonica">
<emphasis id="1B8A9001FFB54B4DFB39FD6AFB66249C" box="[1153,1278,727,752]" italics="true" pageId="8" pageNumber="459">P. japonica</emphasis>
</taxonomicName>
had two adult emergence peaks (late August to early September and late March to early April) and that the female could lay “hundreds” of eggs. 
<bibRefCitation id="4D6F31E2FFB54B4DFEFDFCA2FD8E2555" author="Frouz, J. &amp; Jedlicka, P. &amp; Simackova, H. &amp; Lhotakova, Z." box="[325,534,798,825]" pageId="8" pageNumber="459" pagination="21 - 27" refId="ref33496" refString="Frouz, J., Jedlicka, P., Simackova, H., Lhotakova, Z. (2015) The life cycle, population dynamics, and contribution to litter decomposition of Penthetria holosericea (Diptera: Bibionidae) in an alder forest. European Journal of Soil Biology, 71, 21 - 27. https: // doi. org / 10.1016 / j. ejsobi. 2015.10.002" type="journal article" year="2015">
Frouz 
<emphasis id="1B8A9001FFB54B4DFE35FCA2FE262555" box="[397,446,799,825]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. (2015)
</bibRefCitation>
found the adult life span of 
<taxonomicName id="EEFE3790FFB54B4DFCF6FCA2FC5D2555" box="[846,965,799,825]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFCF6FCA2FC5D2555" box="[846,965,799,825]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
was relatively short (less than one month) and that females laid about 
<specimenCount id="3FF8879AFFB54B4DFE6FFCFEFDD82531" box="[471,576,835,861]" pageId="8" pageNumber="459" type="egg">130 eggs</specimenCount>
, which hatched within a few days. Unlike 
<taxonomicName id="EEFE3790FFB54B4DFB95FCFEFB332531" authorityName="Newman" authorityYear="1834" box="[1069,1195,835,861]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="subFamily" subFamily="Bibioninae">Bibioninae</taxonomicName>
, which have the fore tibia modified with apical mucron/spines which females use to dig a burrow for egg laying, 
<taxonomicName id="EEFE3790FFB54B4DFB1FFCDAFA8525ED" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[1191,1309,871,897]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFB1FFCDAFA8525ED" box="[1191,1309,871,897]" italics="true" pageId="8" pageNumber="459">Penthetria</emphasis>
</taxonomicName>
and 
<taxonomicName id="EEFE3790FFB54B4DFAEDFCDAFA0525ED" box="[1365,1437,871,897]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFAEDFCDAFA0525ED" box="[1365,1437,871,897]" italics="true" pageId="8" pageNumber="459">Plecia</emphasis>
</taxonomicName>
have the fore tibia unmodified; based on this, it is assumed that 
<taxonomicName id="EEFE3790FFB54B4DFCE2FC36FC4825C9" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[858,976,907,933]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFCE2FC36FC4825C9" box="[858,976,907,933]" italics="true" pageId="8" pageNumber="459">Penthetria</emphasis>
</taxonomicName>
females do not burrow, but lay their eggs within small crevices on the surface, as do females of 
<taxonomicName id="EEFE3790FFB54B4DFD4DFC12FBCC25A6" authority="(Pinto &amp; Amorim 1996)" baseAuthorityName="Pinto &amp; Amorim" baseAuthorityYear="1996" box="[757,1108,943,970]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFD4DFC12FCA525A5" box="[757,829,943,969]" italics="true" pageId="8" pageNumber="459">Plecia</emphasis>
(
<bibRefCitation id="4D6F31E2FFB54B4DFCF3FC12FBD425A6" author="Pinto, L. G. &amp; Amorim, D. S." box="[843,1100,943,970]" pageId="8" pageNumber="459" pagination="311 - 326" refId="ref35822" refString="Pinto, L. G. &amp; Amorim, D. S. (1996) Description of immature stages of two Neotropical species of Plecia, with a discussion about the evolution of immature characters in Bibionidae (Insecta, Diptera, Bibionidae). Mitteilungen aus dem Museum fur Naturkunde in Berlin, 72 (2), 311 - 326. https: // doi. org / 10.1002 / mmnz. 19960720215" type="journal article" year="1996">Pinto &amp; Amorim 1996</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="29414C13FFB54B4DFF7FFC6EFC7D2145" blockId="8.[151,1437,150,2014]" pageId="8" pageNumber="459">
Most of the known biological information on the genus comes from studies of the larvae of 
<taxonomicName id="EEFE3790FFB54B4DFB6FFC6EFAD72581" box="[1239,1359,979,1005]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFB6FFC6EFAD72581" box="[1239,1359,979,1005]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
in Europe, where this species is found to be important in nutrient cycling and the decomposition of forest litter (
<bibRefCitation id="4D6F31E2FFB54B4DFA86FC4AFF6D2259" author="Frouz, J. &amp; Elhottova, D. &amp; Sustr, V. &amp; Kristufek, V. &amp; Hubert, J." pageId="8" pageNumber="459" pagination="47 - 51" refId="ref33417" refString="Frouz, J., Elhottova, D., Sustr, V., Kristufek, V., Hubert, J. (2002) Preliminary data about compartmentalization of the gut of the saprophagous dipteran larvae Penthetria holosericea (Bibionidae). European Journal of Soil Biology, 38 (2002), 47 - 51. https: // doi. org / 10.1016 / S 1164 - 5563 (01) 01123 - 2" type="journal article" year="2002">
Frouz 
<emphasis id="1B8A9001FFB54B4DFA30FC4AFF352259" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. 2002
</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFEB8FBA1FE462259" author="Sustr, V. &amp; Frouz, J." box="[256,478,1051,1079]" pageId="8" pageNumber="459" pagination="75 - 77" refId="ref33358" refString="Sustr, V. &amp; Frouz, J. (2002) Activity of carbohydrases in the gut of Bibionidae (Diptera) larvae. European Journal of Soil Biology, 38 (2002), 75 - 77. https: // doi. org / 10.1016 / S 1164 - 5563 (01) 01130 - X" type="journal article" year="2002">Šustr &amp; Frouz 2002</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFE51FBA1FD392259" author="Sustr, V. &amp; Stingl, U. &amp; Brune, A." box="[489,673,1051,1079]" pageId="8" pageNumber="459" pagination="64 - 69" refId="ref36843" refString="Sustr, V., Stingl, U. &amp; Brune, A. (2014) Microprofiles of oxygen, redox potential, and pH, and microbial fermentation products in the highly alkaline gut of the saprophagous larva of Penthetria holosericea (Diptera: Bibionidae). Journal of Insect Physiology, 67, 64 - 69. https: // doi. org / 10.1016 / j. jinsphys. 2014.06.007" type="journal article" year="2014">
Šustr 
<emphasis id="1B8A9001FFB54B4DFD91FBA6FDC12259" box="[553,601,1051,1077]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. 2014
</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFD14FBA6FCF62259" author="Frouz, J. &amp; Jedlicka, P. &amp; Simackova, H. &amp; Lhotakova, Z." box="[684,878,1050,1077]" pageId="8" pageNumber="459" pagination="21 - 27" refId="ref33496" refString="Frouz, J., Jedlicka, P., Simackova, H., Lhotakova, Z. (2015) The life cycle, population dynamics, and contribution to litter decomposition of Penthetria holosericea (Diptera: Bibionidae) in an alder forest. European Journal of Soil Biology, 71, 21 - 27. https: // doi. org / 10.1016 / j. ejsobi. 2015.10.002" type="journal article" year="2015">
Frouz 
<emphasis id="1B8A9001FFB54B4DFD4CFBA6FCBC2259" box="[756,804,1051,1077]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. 2015
</bibRefCitation>
). The saprophagous larvae live in the top layers of soil and “feed selectively on partially decomposed leaves, which are densely colonized by bacteria and fungi” and “their coprophagic behavior allows [the larva] to harvest the microbial production in their faeces during a second gut passage, providing bibionid larvae with an important energy and nutrient source” (
<bibRefCitation id="4D6F31E2FFB54B4DFBDAFB35FABA22CD" author="Sustr, V. &amp; Stingl, U. &amp; Brune, A." box="[1122,1314,1159,1187]" pageId="8" pageNumber="459" pagination="64 - 69" refId="ref36843" refString="Sustr, V., Stingl, U. &amp; Brune, A. (2014) Microprofiles of oxygen, redox potential, and pH, and microbial fermentation products in the highly alkaline gut of the saprophagous larva of Penthetria holosericea (Diptera: Bibionidae). Journal of Insect Physiology, 67, 64 - 69. https: // doi. org / 10.1016 / j. jinsphys. 2014.06.007" type="journal article" year="2014">
Šustr 
<emphasis id="1B8A9001FFB54B4DFB1DFB3AFB4E22CD" box="[1189,1238,1159,1185]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. 2014
</bibRefCitation>
). Based on studies of larval biomass and laboratory consumption of leaf litter, 
<bibRefCitation id="4D6F31E2FFB54B4DFC3BFB16FBCD22A9" author="Frouz, J. &amp; Jedlicka, P. &amp; Simackova, H. &amp; Lhotakova, Z." box="[899,1109,1194,1221]" pageId="8" pageNumber="459" pagination="21 - 27" refId="ref33496" refString="Frouz, J., Jedlicka, P., Simackova, H., Lhotakova, Z. (2015) The life cycle, population dynamics, and contribution to litter decomposition of Penthetria holosericea (Diptera: Bibionidae) in an alder forest. European Journal of Soil Biology, 71, 21 - 27. https: // doi. org / 10.1016 / j. ejsobi. 2015.10.002" type="journal article" year="2015">
Frouz 
<emphasis id="1B8A9001FFB54B4DFC74FB16FC6422A9" box="[972,1020,1195,1221]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. (2015)
</bibRefCitation>
determined that the larvae of this species is responsible for consuming roughly 40% of the annual litter fall at the study site. 
<bibRefCitation id="4D6F31E2FFB54B4DFB15FB72FAD02285" author="Pecina, P." box="[1197,1352,1230,1257]" pageId="8" pageNumber="459" pagination="285 - 298" refId="ref35785" refString="Pecina, P. (1965) Bohemian March-flies (Diptera, Bibionidae) in The National Museum, Prague. Acta Faunistica Entomologica Musei Nationalis Pragae, 11 (108), 285 - 298." type="journal article" year="1965">Pecina (1965)</bibRefCitation>
reports larvae of 
<taxonomicName id="EEFE3790FFB54B4DFEBDFB4EFEE62361" box="[261,382,1267,1293]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFEBDFB4EFEE62361" box="[261,382,1267,1293]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
“inhabiting moist soils with the mouldering remains of vegetables” and that they are “commonly found on the banks of bodies of water where the larvae develop under the litter of deciduous trees, especially of alders and poplars, or in moist meadows.” 
<bibRefCitation id="4D6F31E2FFB54B4DFD1AFA86FCA72339" author="Pecina, P." box="[674,831,1338,1365]" pageId="8" pageNumber="459" pagination="285 - 298" refId="ref35785" refString="Pecina, P. (1965) Bohemian March-flies (Diptera, Bibionidae) in The National Museum, Prague. Acta Faunistica Entomologica Musei Nationalis Pragae, 11 (108), 285 - 298." type="journal article" year="1965">Pecina (1965)</bibRefCitation>
also notes that the larvae live solitarily in the soil in contradistinction to members of the 
<taxonomicName id="EEFE3790FFB54B4DFD89FAE2FD372315" authorityName="Newman" authorityYear="1834" box="[561,687,1375,1401]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="subFamily" subFamily="Bibioninae">Bibioninae</taxonomicName>
, which are gregarious, but this statement is difficult to interpret as many of the studies appear to find larvae in high numbers. 
<bibRefCitation id="4D6F31E2FFB54B4DFC9FFA3EFBD823F1" author="Gemesi, O. &amp; Disney, R. H. L." box="[807,1088,1411,1437]" pageId="8" pageNumber="459" pagination="67 - 69" refId="ref33597" refString="Gemesi, O. &amp; Disney, R. H. L. (1991) A further case of parasitisation of larval Bibionidae by a scuttle fly (Diptera: Phoridae). Entomologist's Gazette, 42, 67 - 69." type="journal article" year="1991">Gemesi &amp; Disney (1991)</bibRefCitation>
reported that during the month of September, mature larvae of 
<taxonomicName id="EEFE3790FFB54B4DFE41FA1AFDE923AD" box="[505,625,1447,1473]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFE41FA1AFDE923AD" box="[505,625,1447,1473]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
were collected in decaying leaf litter of alder trees (
<taxonomicName id="EEFE3790FFB54B4DFB03FA1AFAF223AD" box="[1211,1386,1447,1473]" class="Magnoliopsida" family="Betulaceae" genus="Alnus" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="459" phylum="Tracheophyta" rank="species" species="glutinosa">
<emphasis id="1B8A9001FFB54B4DFB03FA1AFAF223AD" box="[1211,1386,1447,1473]" italics="true" pageId="8" pageNumber="459">Alnus glutinosa</emphasis>
</taxonomicName>
(L.) Gaertn.) along a river; 8.7% of the individuals were later found to be parasitized in the 2 
<superScript id="DE8BE15BFFB54B4DFB20FA77FB3023B4" attach="left" box="[1176,1192,1482,1496]" fontSize="6" pageId="8" pageNumber="459">nd</superScript>
or early 3 
<superScript id="DE8BE15BFFB54B4DFA9AFA77FAA823B4" attach="left" box="[1314,1328,1482,1496]" fontSize="6" pageId="8" pageNumber="459">rd</superScript>
instar by 
<taxonomicName id="EEFE3790FFB54B4DFF2FFA52FE6F2065" authority="Schmitz" authorityName="Schmitz" box="[151,503,1519,1545]" class="Insecta" family="Phoridae" genus="Borophaga" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="germanica">
<emphasis id="1B8A9001FFB54B4DFF2FFA52FE0C2064" box="[151,404,1519,1545]" italics="true" pageId="8" pageNumber="459">Borophaga germanica</emphasis>
Schmitz
</taxonomicName>
(
<taxonomicName id="EEFE3790FFB54B4DFDBFFA52FDF92065" box="[519,609,1519,1545]" class="Insecta" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="order">Diptera</taxonomicName>
: 
<taxonomicName id="EEFE3790FFB54B4DFDD4FA52FD4D2066" box="[620,725,1519,1546]" class="Insecta" family="Phoridae" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="family">Phoridae</taxonomicName>
). The later authors also reported that 
<taxonomicName id="EEFE3790FFB54B4DFBC1FA52FB692065" box="[1145,1265,1519,1545]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFBC1FA52FB692065" box="[1145,1265,1519,1545]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
overwinters as a mature larva. Based on measurements of larval head capsules, 
<bibRefCitation id="4D6F31E2FFB54B4DFCD1F9AEFBA32041" author="Frouz, J. &amp; Jedlicka, P. &amp; Simackova, H. &amp; Lhotakova, Z." box="[873,1083,1554,1581]" pageId="8" pageNumber="459" pagination="21 - 27" refId="ref33496" refString="Frouz, J., Jedlicka, P., Simackova, H., Lhotakova, Z. (2015) The life cycle, population dynamics, and contribution to litter decomposition of Penthetria holosericea (Diptera: Bibionidae) in an alder forest. European Journal of Soil Biology, 71, 21 - 27. https: // doi. org / 10.1016 / j. ejsobi. 2015.10.002" type="journal article" year="2015">
Frouz 
<emphasis id="1B8A9001FFB54B4DFC0AF9AEFC7B2041" box="[946,995,1555,1581]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. (2015)
</bibRefCitation>
determined that 
<taxonomicName id="EEFE3790FFB54B4DFB41F9AEFAE82041" box="[1273,1392,1555,1581]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFB41F9AEFAE82041" box="[1273,1392,1555,1581]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
has seven larval instars and spends most of its developmental time (as well as usually overwinters) in the sixth instar, which takes about two months to reach; the seventh instar and pupa were inferred to have short durations based on the fact that they were infrequently detected. 
<bibRefCitation id="4D6F31E2FFB54B4DFD2DF9C2FCF920F5" author="Yuan, R. &amp; Li, S. &amp; Du, Q. &amp; Lei, C. &amp; Zhu, F." box="[661,865,1662,1689]" pageId="8" pageNumber="459" pagination="320 - 324" refId="ref37430" refString="Yuan, R., Li, S., Du, Q., Lei, C. &amp; Zhu, F. (2015) Immature stages description of March Fly, Penthetria japonica Wiedemann (Diptera: Bibionidae), from Hubei, China. Entomological News, 124 (5), 320 - 324. https: // doi. org / 10.3157 / 021.124.0503" type="journal article" year="2015">
Yuan 
<emphasis id="1B8A9001FFB54B4DFD6EF9C2FC9020F5" box="[726,776,1663,1689]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. (2015)
</bibRefCitation>
found the larvae of 
<taxonomicName id="EEFE3790FFB54B4DFBFDF9C2FB5B20F4" authorityName="Wiedemann" authorityYear="1830" box="[1093,1219,1663,1688]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="japonica">
<emphasis id="1B8A9001FFB54B4DFBFDF9C2FB5B20F4" box="[1093,1219,1663,1688]" italics="true" pageId="8" pageNumber="459">P. japonica</emphasis>
</taxonomicName>
in the upper layers of cow dung at an earthworm breeding farm; larvae were successfully reared on this medium in the laboratory. It is noteworthy, that while much of what we know of the biology of 
<taxonomicName id="EEFE3790FFB54B4DFCCDF97AFC73208D" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[885,1003,1735,1761]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFCCDF97AFC73208D" box="[885,1003,1735,1761]" italics="true" pageId="8" pageNumber="459">Penthetria</emphasis>
</taxonomicName>
is based on 
<taxonomicName id="EEFE3790FFB54B4DFBC1F97AFB6A208D" box="[1145,1266,1735,1761]" class="Insecta" family="Bibionidae" genus="Amasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="species" species="funebris">
<emphasis id="1B8A9001FFB54B4DFBC1F97AFB6A208D" box="[1145,1266,1735,1761]" italics="true" pageId="8" pageNumber="459">P. funebris</emphasis>
</taxonomicName>
, this species is not an average representative of the genus, as it is the only member of the genus to have dichoptic and brachypterous males, which are flightless (
<bibRefCitation id="4D6F31E2FFB54B4DFE68F8B2FC922145" author="Skartveit, J. &amp; Willassen, E." box="[464,778,1807,1833]" pageId="8" pageNumber="459" pagination="1 - 316" refId="ref36781" refString="Skartveit, J. &amp; Willassen, E. (1996) Phylogenetic relationships in Bibioninae (Diptera, Bibionidae). In: Skartveit, J., Studies on the systematics and life histories of Bibioninae (Diptera, Bibionidae). Ph. D. dissertation, University of Bergen, Bergen, pp. 1 - 316." type="book chapter" year="1996">Skartveit &amp; Willassen 1996</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFCADF8B2FC412145" author="Frouz, J. &amp; Jedlicka, P. &amp; Simackova, H. &amp; Lhotakova, Z." box="[789,985,1806,1833]" pageId="8" pageNumber="459" pagination="21 - 27" refId="ref33496" refString="Frouz, J., Jedlicka, P., Simackova, H., Lhotakova, Z. (2015) The life cycle, population dynamics, and contribution to litter decomposition of Penthetria holosericea (Diptera: Bibionidae) in an alder forest. European Journal of Soil Biology, 71, 21 - 27. https: // doi. org / 10.1016 / j. ejsobi. 2015.10.002" type="journal article" year="2015">
Frouz 
<emphasis id="1B8A9001FFB54B4DFCE5F8B2FC162145" box="[861,910,1807,1833]" italics="true" pageId="8" pageNumber="459">et al</emphasis>
. 2015
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="61E41F98FFB54B4CFF7FF88EFC0F2236" lastPageId="9" lastPageNumber="460" pageId="8" pageNumber="459" type="description">
<paragraph id="29414C13FFB54B4DFF7FF88EFA1C211D" blockId="8.[151,1437,150,2014]" pageId="8" pageNumber="459">The immature stages and biology of the New World species is entirely unknown though it is expected to be similar to what is known for the few Palearctic species in which immature stages and biology have been studied.</paragraph>
<paragraph id="29414C13FFB54B4CFF7FF8C7FF522695" blockId="8.[151,1437,150,2014]" lastBlockId="9.[151,1437,150,1114]" lastPageId="9" lastPageNumber="460" pageId="8" pageNumber="459">
<emphasis id="1B8A9001FFB54B4DFF7FF8C7FEF121F8" bold="true" box="[199,361,1914,1940]" pageId="8" pageNumber="459">Identification</emphasis>
. Aside from the New World species (treated in the present study and 
<bibRefCitation id="4D6F31E2FFB54B4DFB39F8C6FA9221F9" author="Hardy, D. E." box="[1153,1290,1914,1941]" pageId="8" pageNumber="459" pagination="367 - 547" refId="ref33819" refString="Hardy, D. E. (1945) Revision of Nearctic Bibionidae, including Neotropical Plecia and Penthetria (Diptera). Kansas University Science Bulletin, 30, 367 - 547." type="journal article" year="1945">Hardy 1945</bibRefCitation>
), the following papers include keys or other information helpful for identifying 
<taxonomicName id="EEFE3790FFB54B4DFC1FF822FB8521D5" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[935,1053,1951,1977]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="8" pageNumber="459" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB54B4DFC1FF822FB8521D5" box="[935,1053,1951,1977]" italics="true" pageId="8" pageNumber="459">Penthetria</emphasis>
</taxonomicName>
in the following regions: Europe (
<bibRefCitation id="4D6F31E2FFB54B4DFF27F87EFE4821B1" author="Fitzgerald, S. J. &amp; Werner, D." box="[159,464,1987,2013]" pageId="8" pageNumber="459" pagination="207 - 210" refId="ref33304" refString="Fitzgerald, S. J. &amp; Werner, D. (2004) A key to the Penthetria Meigen (Diptera, Bibionidae) of Europe and the first record of Penthetria heteroptera (Say) from the Palaearctic region. Studia dipterologica, 11 (1), 207 - 210." type="journal article" year="2004">Fitzgerald &amp; Werner 2004</bibRefCitation>
), 
<collectingCountry id="51E90C83FFB54B4DFE5EF87EFDB121B1" box="[486,553,1987,2013]" name="China" pageId="8" pageNumber="459">China</collectingCountry>
(
<bibRefCitation id="4D6F31E2FFB54B4DFD81F87EFD6121B1" author="Li, Z. &amp; Yang, D." box="[569,761,1987,2013]" pageId="8" pageNumber="459" pagination="165 - 171" refId="ref35000" refString="Li, Z. &amp; Yang, D. (2010) Two New Species of the genus Penthetria Meigen from Yunnan, China (Diptera: Bibionidae). Entomological News, 121 (2), 165 - 171. https: // doi. org / 10.3157 / 021.121.0208" type="journal article" year="2010">Li &amp; Yang 2010</bibRefCitation>
), 
<collectingCountry id="51E90C83FFB54B4DFCB7F87EFCD721B1" box="[783,847,1987,2013]" name="Japan" pageId="8" pageNumber="459">Japan</collectingCountry>
(
<bibRefCitation id="4D6F31E2FFB54B4DFCE7F87EFB1B21B1" author="Hardy, D. E. &amp; Takahashi, M." box="[863,1155,1987,2013]" pageId="8" pageNumber="459" pagination="383 - 449" refId="ref34286" refString="Hardy, D. E. &amp; Takahashi, M. (1960) Revision of the Japanese Bibionidae (Diptera, Nematocera). Pacific Insects, 2 (4), 383 - 449." type="journal article" year="1960">Hardy &amp; Takahashi 1960</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB54B4DFB28F87EFADC21B2" author="Sasakawa, M." box="[1168,1348,1987,2014]" pageId="8" pageNumber="459" pagination="361 - 363" refId="ref36111" refString="Sasakawa, M. (1967) A new species of Penthetria from Japan (Diptera: Bibionidae). Kontyu, 35 (4), 361 - 363." type="journal article" year="1967">Sasakawa 1967</bibRefCitation>
), 
<collectingCountry id="51E90C83FFB54B4DFAE1F87EFA0421B1" box="[1369,1436,1987,2013]" name="Nepal" pageId="8" pageNumber="459">Nepal</collectingCountry>
(Hardy 1965, 1967), and 
<collectingCountry id="51E90C83FFB44B4CFE78FF2AFD9326DD" box="[448,523,151,177]" name="Russia" pageId="9" pageNumber="460">Russia</collectingCountry>
and 
<collectingCountry id="51E90C83FFB44B4CFDF0FF2AFD2D26DD" box="[584,693,151,177]" name="Mongolia" pageId="9" pageNumber="460">Mongolia</collectingCountry>
(
<bibRefCitation id="4D6F31E2FFB44B4CFD7FFF2AFBD226DD" author="Krivosheina, N. P. &amp; Krivosheina, M." box="[711,1098,151,177]" pageId="9" pageNumber="460" pagination="105 - 108" refId="ref34730" refString="Krivosheina, N. P. &amp; Krivosheina, M. G (1998) The flies of the family Pleciidae (Diptera) new to the fauna of Russia. International Journal of Dipterological Research, 9, 105 - 108." type="journal article" year="1998">Krivosheina &amp; Krivosheina 1998</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB44B4CFBE1FF2AFABF26DD" author="Krivosheina, N. P." box="[1113,1319,151,177]" pageId="9" pageNumber="460" pagination="1 - 665" refId="ref34671" refString="Krivosheina, N. P. (1999) Pleciidae. In: Ler, P. A. (Ed.), Key to the insects of Russian Far East. Vol. VI. Diptera and Siphonaptera. Pt. 1. Dal'nauka, Vladivostok, pp. 1 - 665. [in Russian]" type="book chapter" year="1999">Krivosheina 1999</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB44B4CFA8EFF2AFF4A26B9" author="Nartshuk, E. P." pageId="9" pageNumber="460" pagination="353 - 370" refId="ref35546" refString="Nartshuk, E. P. (1990) Pleciidae and Bibionidae (Diptera) from the Mongolian People's Republic, with a review of Pleciidae of the USSR fauna. Insects of Mongolia, 11, 353 - 370. [in Russian]" type="journal article" year="1990">Nartshuk 1990</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB44B4CFF64FF06FE6F26BA" author="Hardy, D. E. &amp; Takahashi, M." box="[220,503,187,214]" pageId="9" pageNumber="460" pagination="383 - 449" refId="ref34286" refString="Hardy, D. E. &amp; Takahashi, M. (1960) Revision of the Japanese Bibionidae (Diptera, Nematocera). Pacific Insects, 2 (4), 383 - 449." type="journal article" year="1960">Hardy &amp; Takahashi 1960</bibRefCitation>
). Additionally, the catalog of world species (below) references all the primary literature.
</paragraph>
<paragraph id="29414C13FFB44B4CFF7FFEBFFC0F2236" blockId="9.[151,1437,150,1114]" pageId="9" pageNumber="460">
<emphasis id="1B8A9001FFB44B4CFF7FFEBFFE4E2770" bold="true" box="[199,470,258,284]" pageId="9" pageNumber="460">Intraspecific Variation</emphasis>
. While the general form of 
<taxonomicName id="EEFE3790FFB44B4CFCB6FEBEFC1C2771" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[782,900,259,285]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB44B4CFCB6FEBEFC1C2771" box="[782,900,259,285]" italics="true" pageId="9" pageNumber="460">Penthetria</emphasis>
</taxonomicName>
species is remarkably homogeneous (most species look very similar to 
<figureCitation id="B1C55096FFB44B4CFE0AFE9AFE6C272D" box="[434,500,295,321]" captionStart="FIGURE 1" captionStartId="4.[151,250,1982,2006]" captionTargetBox="[331,1264,760,1945]" captionTargetId="figure-296@4.[309,1278,741,1959]" captionTargetPageId="4" captionText="FIGURE 1. P. heteroptera, habitus, male (above) and female (below), 28 October, North Carolina, USA, Matt Bertone. Images courtesy of Matt Bertone (© Matt Bertone 2013)." figureDoi="http://doi.org/10.5281/zenodo.4529514" httpUri="https://zenodo.org/record/4529514/files/figure.png" pageId="9" pageNumber="460">Fig. 1</figureCitation>
), there is considerable intraspecific variation that makes identification and species delimitation challenging. Some of the characters that have been previously used to help distinguish species, such as whether vein R 
<subScript id="B57A4E56FFB44B4CFEDAFE3CFEE327E3" attach="left" box="[354,379,385,399]" fontSize="6" pageId="9" pageNumber="460">2+3</subScript>
has an appendix at the base (
<bibRefCitation id="4D6F31E2FFB44B4CFD7AFED2FCD527E5" author="Hardy, D. E." box="[706,845,366,393]" pageId="9" pageNumber="460" pagination="367 - 547" refId="ref33819" refString="Hardy, D. E. (1945) Revision of Nearctic Bibionidae, including Neotropical Plecia and Penthetria (Diptera). Kansas University Science Bulletin, 30, 367 - 547." type="journal article" year="1945">Hardy 1945</bibRefCitation>
), have been found to be variable (pers. obs.) and not consistently present or absent within a given species (e.g., 
<taxonomicName id="EEFE3790FFB44B4CFC90FE2EFC5527C1" authorityName="Hardy" authorityYear="1945" box="[808,973,403,429]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="species" species="appendicula">
<emphasis id="1B8A9001FFB44B4CFC90FE2EFC5527C1" box="[808,973,403,429]" italics="true" pageId="9" pageNumber="460">P. appendicula</emphasis>
</taxonomicName>
typically but not always has an appendix (
<figureCitation id="B1C55096FFB44B4CFF18FE0AFF7C27BD" box="[160,228,439,465]" captionStart="FIGURES 2–3" captionStartId="11.[151,264,1518,1542]" captionTargetBox="[218,1372,186,1488]" captionTargetId="figure-16@11.[212,1376,181,1494]" captionTargetPageId="11" captionText="FIGURES 2–3. P. appendicula. 2. Wing. 3. Female habitus, lateral view. Scale bars = ca. 1.0 mm. Abbreviations: ba, basal appendix of R2+3; aem, anepimeron; aes, anepisternum; c, coxa; f, femur; h, halter; hb, hind basitarsus; kem, katepimeron; kes, katepisternum; m, meron; mtaes, metanepisternum; mtg, mediotergite; mtkes, metakatepisternum; ltg, laterotergite; pltrch, pleu- rotrochantin; prn, pronotum; sct, scutum; t, trochanter." figureDoi="http://doi.org/10.5281/zenodo.4529518" httpUri="https://zenodo.org/record/4529518/files/figure.png" pageId="9" pageNumber="460">Fig. 2</figureCitation>
, ba), 
<taxonomicName id="EEFE3790FFB44B4CFE9AFE0AFE0527BD" authorityName="Hardy" authorityYear="1945" box="[290,413,439,465]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="species" species="distincta">
<emphasis id="1B8A9001FFB44B4CFE9AFE0AFE0527BD" box="[290,413,439,465]" italics="true" pageId="9" pageNumber="460">P. distincta</emphasis>
</taxonomicName>
generally does not but may have an appendix, and 
<taxonomicName id="EEFE3790FFB44B4CFC64FE0AFBF527BC" authorityName="Fitzgerald" authorityYear="2021" box="[988,1133,439,464]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="species" species="neonigrita">
<emphasis id="1B8A9001FFB44B4CFC64FE0AFBF527BC" box="[988,1133,439,464]" italics="true" pageId="9" pageNumber="460">P. neonigrita</emphasis>
</taxonomicName>
usually does not but rarely does have an appendix). Additionally, some species have a tendency to have CuA and CuP meeting apically near the wing margin forming a closed cell cua (
<figureCitation id="B1C55096FFB44B4CFD39FE42FD4D2475" box="[641,725,511,537]" captionStart="FIGURES 30–32" captionStartId="29.[151,264,1324,1348]" captionTargetBox="[248,1339,186,1295]" captionTargetId="figure-16@29.[243,1343,181,1299]" captionTargetPageId="29" captionText="FIGURES 30–32. Holotype males, lateral. 30. P. neonigrita n. sp.. 31. P. yakima n. sp.. 32. P. arizonensis n. sp.. Scale bars = ca. 1.0 mm. Abbreviations: hb, hind basitarsus." figureDoi="http://doi.org/10.5281/zenodo.4529546" httpUri="https://zenodo.org/record/4529546/files/figure.png" pageId="9" pageNumber="460">Fig. 32</figureCitation>
), yet this character is not consistent when numerous specimens are studied. In some cases, one wing will have a closed cell cua and the other wing has an open cell (pers. obs). Plasticity in the wing venation (e.g., presence/absence of an appendix on R 
<subScript id="B57A4E56FFB44B4CFC58FDE4FC61240B" attach="left" box="[992,1017,601,615]" fontSize="6" pageId="9" pageNumber="460">2+3</subScript>
, length of stem of M, and position of r-m crossvein) has also been noted in some of the Palearctic species (
<bibRefCitation id="4D6F31E2FFB44B4CFC13FDD6FBCF24E9" author="Nartshuk, E. P." box="[939,1111,619,645]" pageId="9" pageNumber="460" pagination="353 - 370" refId="ref35546" refString="Nartshuk, E. P. (1990) Pleciidae and Bibionidae (Diptera) from the Mongolian People's Republic, with a review of Pleciidae of the USSR fauna. Insects of Mongolia, 11, 353 - 370. [in Russian]" type="journal article" year="1990">Nartshuk 1990</bibRefCitation>
, 
<bibRefCitation id="4D6F31E2FFB44B4CFBDCFDD6FF4B24C5" author="Krivosheina, N. P. &amp; Krivosheina, M." pageId="9" pageNumber="460" pagination="105 - 108" refId="ref34730" refString="Krivosheina, N. P. &amp; Krivosheina, M. G (1998) The flies of the family Pleciidae (Diptera) new to the fauna of Russia. International Journal of Dipterological Research, 9, 105 - 108." type="journal article" year="1998">Krivosheina &amp; Krivosheina 1998</bibRefCitation>
). Unfortunately, the strong plasticity of the wing venation observed in extant taxa does not bode well for the taxonomy of fossil members of the group, since many species have been delimited largely or solely on differences in wing veins. For example, the slope and length of vein R 
<subScript id="B57A4E56FFB44B4CFC8BFD54FCD4249B" attach="left" box="[819,844,745,759]" fontSize="6" pageId="9" pageNumber="460">2+3</subScript>
has been used to help delimit both extant and fossil species. The study of the intraspecific variation of this character in extant species indicates that vein R 
<subScript id="B57A4E56FFB44B4CFAA3FCB0FAAC2577" attach="left" box="[1307,1332,781,795]" fontSize="6" pageId="9" pageNumber="460">2+3</subScript>
can vary drastically in slope and length even between specimens of the same species collected from sites that are relatively near each other (e.g., 
<figureCitation id="B1C55096FFB44B4CFE37FCFEFE732532" box="[399,491,835,862]" captionStart="FIGURES 21–22" captionStartId="24.[151,264,1285,1309]" captionTargetBox="[227,1365,351,1257]" captionTargetId="figure-115@24.[222,1365,345,1261]" captionTargetPageId="24" captionText="FIGURES 21–22. Penthetria and Plecia, wings, illustrating variation in length and slope of vein R 2+3. 21a–b. Penthetira heteroptera, males (both from Maryland, USA). 22a. Plecia dentata Hardy, male. 22b. Plecia biformis Hardy, male (points, x, y, z described in “Discussion supporting generic diagnosis”). Scale bars = ca. 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4529538" httpUri="https://zenodo.org/record/4529538/files/figure.png" pageId="9" pageNumber="460">Figs. 21</figureCitation>
a–b). Concerns about how the intraspecific variation found in the wing venation of extant species of bibionids translates into the delimitation of fossil species was previously raised by 
<bibRefCitation id="4D6F31E2FFB44B4CFB43FCDAFF7D25C9" author="Collomb, F. - M. &amp; Nel, A. &amp; Fleck, G. &amp; Waller, A." pageId="9" pageNumber="460" pagination="161 - 179" refId="ref32489" refString="Collomb, F. - M., Nel, A., Fleck, G. &amp; Waller, A. 2008 March flies and European Cenozoic palaeoclimates (Diptera: Bibionidae). Annales de la Societe Entomologique de France, 44 (2), 161 - 179. https: // doi. org / 10.1080 / 00379271.2008.10697553" type="journal article" year="2008">
Collomb 
<emphasis id="1B8A9001FFB44B4CFADCFCDAFA0D25ED" box="[1380,1429,871,897]" italics="true" pageId="9" pageNumber="460">et al</emphasis>
. (2008)
</bibRefCitation>
. His study noted the plasticity of the length and shape of r-m in 
<taxonomicName id="EEFE3790FFB44B4CFC0BFC36FBC625C9" authority="Geoffroy" authorityName="Geoffroy" authorityYear="1762" box="[947,1118,907,933]" class="Insecta" family="Bibionidae" genus="Bibio" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB44B4CFC0BFC36FC6825C9" box="[947,1008,907,933]" italics="true" pageId="9" pageNumber="460">Bibio</emphasis>
Geoffroy
</taxonomicName>
, the slope and length of R 
<subScript id="B57A4E56FFB44B4CFA3BFC20FA0425C7" attach="left" box="[1411,1436,925,939]" fontSize="6" pageId="9" pageNumber="460">
2+ 
<quantity id="EE06E1F6FFB44B4CFA2DFC20FA0425C7" box="[1429,1436,925,939]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="9" pageNumber="460" unit="in" value="3.0">3</quantity>
</subScript>
in 
<taxonomicName id="EEFE3790FFB44B4CFF0EFC12FF6625A5" baseAuthorityName="Pinto &amp; Amorim" baseAuthorityYear="1996" box="[182,254,943,969]" class="Insecta" family="Bibionidae" genus="Plecia" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB44B4CFF0EFC12FF6625A5" box="[182,254,943,969]" italics="true" pageId="9" pageNumber="460">Plecia</emphasis>
</taxonomicName>
, and the length and shape of R 
<subScript id="B57A4E56FFB44B4CFDDDFC7CFDE625A3" attach="left" box="[613,638,961,975]" fontSize="6" pageId="9" pageNumber="460">
2+ 
<quantity id="EE06E1F6FFB44B4CFDCFFC7CFDE625A3" box="[631,638,961,975]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="9" pageNumber="460" unit="in" value="3.0">3</quantity>
</subScript>
in 
<taxonomicName id="EEFE3790FFB44B4CFD1EFC12FC8425A5" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[678,796,943,969]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB44B4CFD1EFC12FC8425A5" box="[678,796,943,969]" italics="true" pageId="9" pageNumber="460">Penthetria</emphasis>
</taxonomicName>
and arrived at the conclusion that male terminalia were the best source of information for delimiting fossil species (which mirrors conclusions by those working on extant taxa). It is possible that the number of fossil species of 
<taxonomicName id="EEFE3790FFB44B4CFD45FC4AFCEB227D" baseAuthorityName="Loew" baseAuthorityYear="1868" box="[765,883,1015,1041]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="1B8A9001FFB44B4CFD45FC4AFCEB227D" box="[765,883,1015,1041]" italics="true" pageId="9" pageNumber="460">Penthetria</emphasis>
</taxonomicName>
(and possibly other genera) have been artificially inflated due to variable wing venation described under several names.Additional examples of intraspecific variation in the genus can also be found under “ 
<taxonomicName id="EEFE3790FFB44B4CFDFCFB82FCBF2235" authority="(Say)" baseAuthorityName="Say" baseAuthorityYear="1823" box="[580,807,1087,1113]" class="Insecta" family="Bibionidae" genus="Penthetria" kingdom="Animalia" order="Diptera" pageId="9" pageNumber="460" phylum="Arthropoda" rank="species" species="heteroptera">
<emphasis id="1B8A9001FFB44B4CFDFCFB82FD7A2235" box="[580,738,1087,1113]" italics="true" pageId="9" pageNumber="460">P. heteroptera</emphasis>
(Say)
</taxonomicName>
” (below).
</paragraph>
</subSubSection>
</treatment>
</document>