treatments-xml/data/5E/27/96/5E279627995EFFCB38EEFB10FBE5C236.xml

<document id="9B855033C0252C4D1E42E79042FF0648" ID-DOI="10.5281/zenodo.274756" ID-GBIF-Dataset="f352d143-5a5b-4bcb-97f1-232fc9428207" ID-ISSN="1175-5326" ID-Zenodo-Dep="274756" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461047157992" checkinUser="plazi" docAuthor="Arillo, Antonio, Peñalver, Enrique &amp; García-Gimeno, Victoria" docDate="2009" docId="5E279627995EFFCB38EEFB10FBE5C236" docLanguage="en" docName="zt02026p039.pdf" docOrigin="Zootaxa 2026" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Litoleptis fossilis Arillo, Penalver &amp; Garcia-Gimeno, sp. nov." docType="treatment" docVersion="6" lastPageNumber="38" masterDocId="A21EEE5F995FFFCE3879FFACFFCEC12A" masterDocTitle="First fossil Litoleptis (Diptera: Spaniidae) from the Lower Cretaceous amber of San Just (Teruel Province, Spain)" masterLastPageNumber="39" masterPageNumber="33" pageNumber="34" updateTime="1698577501613" updateUser="plazi">
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<mods:title id="05D075909CE25A6B2BCB4801097007E1">First fossil Litoleptis (Diptera: Spaniidae) from the Lower Cretaceous amber of San Just (Teruel Province, Spain)</mods:title>
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<mods:namePart id="D0AEB757732A9A889E8DCCBBCBD06F09">Arillo, Antonio</mods:namePart>
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<mods:namePart id="B31CD866FCD5C9912556EAEF44D60663">Peñalver, Enrique</mods:namePart>
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<mods:namePart id="FA044E847E5C8DA94E80E7CAF5D09F0A">García-Gimeno, Victoria</mods:namePart>
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<mods:date id="6CCA2959426C2349E6A6A3358EBA2DFF">2009</mods:date>
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<treatment id="5E279627995EFFCB38EEFB10FBE5C236" ID-DOI="http://doi.org/10.5281/zenodo.6214466" ID-GBIF-Taxon="119611032" ID-Zenodo-Dep="6214466" LSID="urn:lsid:plazi:treatment:5E279627995EFFCB38EEFB10FBE5C236" httpUri="http://treatment.plazi.org/id/5E279627995EFFCB38EEFB10FBE5C236" lastPageId="5" lastPageNumber="38" pageId="1" pageNumber="34">
<subSubSection id="9E9474BA995EFFCF38EEFB10FE8CC5D2" pageId="1" pageNumber="34" type="nomenclature">
<paragraph id="D6312731995EFFCF38EEFB10FCB0C5FF" blockId="1.[151,894,1211,1272]" box="[151,894,1211,1238]" pageId="1" pageNumber="34">
<heading id="8D79905D995EFFCF38EEFB10FCB0C5FF" bold="true" box="[151,894,1211,1238]" fontSize="11" level="1" pageId="1" pageNumber="34" reason="1">
<emphasis id="E4FAFB23995EFFCF38EEFB10FCB0C5FF" bold="true" box="[151,894,1211,1238]" pageId="1" pageNumber="34">
<taxonomicName id="118E5CB2995EFFCF38EEFB10FCD7C5FF" ID-CoL="9HM2Q" authority="Arillo, Penalver &amp; Garcia-Gimeno" authorityName="Arillo, Penalver &amp; Garcia-Gimeno" box="[151,793,1211,1238]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="34" phylum="Arthropoda" rank="species" species="fossilis" status="sp. nov.">
<emphasis id="E4FAFB23995EFFCF38EEFB10FEAFC5FC" bold="true" box="[151,353,1212,1238]" italics="true" pageId="1" pageNumber="34">Litoleptis fossilis</emphasis>
Arillo, Peñalver &amp; García-Gimeno
</taxonomicName>
<taxonomicNameLabel id="FFC94658995EFFCF3B59FB17FCB0C5FF" box="[800,894,1211,1237]" pageId="1" pageNumber="34" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="D6312731995EFFCF38EEFB72FE8CC5D2" blockId="1.[151,894,1211,1272]" box="[151,322,1246,1272]" pageId="1" pageNumber="34">Figs 1–2; Pl. 1</paragraph>
</subSubSection>
<subSubSection id="9E9474BA995EFFCF38EEFA87FE17C446" pageId="1" pageNumber="34" type="description">
<paragraph id="D6312731995EFFCF38EEFA87FE17C446" blockId="1.[151,1436,1323,2007]" pageId="1" pageNumber="34">
<emphasis id="E4FAFB23995EFFCF38EEFA87FE55C46F" bold="true" box="[151,411,1323,1349]" pageId="1" pageNumber="34">Derivation of name:</emphasis>
From the Latin 
<taxonomicName id="118E5CB2995EFFCF3A1BFA80FD79C46F" box="[610,695,1324,1349]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="34" phylum="Arthropoda" rank="species" species="fossilis">
<emphasis id="E4FAFB23995EFFCF3A1BFA80FD79C46F" box="[610,695,1324,1349]" italics="true" pageId="1" pageNumber="34">fossilis</emphasis>
</taxonomicName>
(dug out of the earth, excavated, 
<emphasis id="E4FAFB23995EFFCF3C2AFA80FBB0C46F" box="[1107,1150,1324,1349]" italics="true" pageId="1" pageNumber="34">etc.</emphasis>
). This is the first fossil representative of the genus.
</paragraph>
</subSubSection>
<subSubSection id="9E9474BA995EFFCF38BFFAD4FB0EC751" pageId="1" pageNumber="34" type="materials_examined">
<paragraph id="D6312731995EFFCF38BFFAD4FC30C4CA" blockId="1.[151,1436,1323,2007]" pageId="1" pageNumber="34">
<emphasis id="E4FAFB23995EFFCF38BFFAD4FE8CC4B8" bold="true" box="[198,322,1400,1426]" pageId="1" pageNumber="34">
<typeStatus id="09359993995EFFCF38BFFAD4FEF3C4B8" box="[198,317,1400,1426]" pageId="1" pageNumber="34" type="holotype">Holotype</typeStatus>
:
</emphasis>
CPT-3344 housed in the Fundación Conjunto Paleontológico de Teruel-Dinópolis (Teruel Province, 
<collectingCountry id="AE9967A1995EFFCF396BFA0CFE92C490" box="[274,348,1440,1466]" name="Spain" pageId="1" pageNumber="34">Spain</collectingCountry>
). Complete, well preserved specimen (Pl. 1, fig. 1) present in a transparent, clear piece of amber of 
<date id="A23001F1995EFFCF397EFA6AFE96C4CA" box="[263,344,1478,1504]" pageId="1" pageNumber="34">4 x 4 x</date>
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into an epoxy resin preparation of 
<date id="A23001F1995EFFCF3B44FA6AFC5BC4CA" box="[829,917,1478,1504]" pageId="1" pageNumber="34" value="1913-10-22">22 x 13</date>
x 
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.
</paragraph>
<paragraph id="D6312731995EFFCF38BFFA41FB0EC751" blockId="1.[151,1436,1323,2007]" pageId="1" pageNumber="34">
<emphasis id="E4FAFB23995EFFCF38BFFA41FD86C72D" bold="true" box="[198,584,1517,1543]" pageId="1" pageNumber="34">
<typeStatus id="09359993995EFFCF38BFFA41FECAC72D" box="[198,260,1517,1543]" pageId="1" pageNumber="34">Type</typeStatus>
locality and stratigraphy:
</emphasis>
Specimen from San Just outcrop, in the municipality of Utrillas, near the village of Escucha (Teruel Province, 
<collectingCountry id="AE9967A1995EFFCF3A33F9B8FD5CC704" box="[586,658,1556,1582]" name="Spain" pageId="1" pageNumber="34">Spain</collectingCountry>
). It was collected in a thin level of grey-black claystones very rich in amber, charcoal and also fusinite formed by a palaeofire, which implicates an exceptional resin production in the palaeoforests. Escucha Fm. (La Orden Member), Lower Cretaceous (Lower Albian).
</paragraph>
</subSubSection>
<subSubSection id="9E9474BA995EFFCF38BFF92BFB2BC7C5" pageId="1" pageNumber="34" type="diagnosis">
<paragraph id="D6312731995EFFCF38BFF92BFB2BC7C5" blockId="1.[151,1436,1323,2007]" pageId="1" pageNumber="34">
<emphasis id="E4FAFB23995EFFCF38BFF92BFE86C78B" bold="true" box="[198,328,1671,1697]" pageId="1" pageNumber="34">Diagnosis:</emphasis>
Antennal flagellomere III prolonged medially into a long, densely pubescent, unsegmented style with its basal part bulbous. Labium as long as antennae. Presence of a small spur on metathoracic tibia. R4 and R5 end in the costal margin. R4+5 and M1+2 forked at same distance from wing base.
</paragraph>
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<subSubSection id="9E9474BA995EFFCC38BFF957FCA4C7EA" lastPageId="2" lastPageNumber="35" pageId="1" pageNumber="34" type="description">
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<emphasis id="E4FAFB23995EFFCF38BFF957FEAFC63F" bold="true" box="[198,353,1787,1813]" pageId="1" pageNumber="34">Description:</emphasis>
Body length 
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from head frons to terminalia apex (Fig. 1 and Pl. 1, fig. 1).
</paragraph>
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Head. Eyes bare, large, 
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long, 
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aprox. wide (estimated); dorsal and ventral facets undifferentiated (Fig. 2a and Pl. 1, fig. 2). Eyes nearly holoptic, separated only by distance less to diameter of facet. Postocciput with a dorsal line of fine setae. Ocellar triangle prominent and setulose (ocellar diameter 
<quantity id="11768AD4995EFFCF38EEF83AFECDC69A" box="[151,259,1942,1968]" metricMagnitude="-5" metricUnit="m" metricValue="7" pageId="1" pageNumber="34" unit="mm" value="0.07">0.07 mm</quantity>
) (Figs. 1 and 2a). Antennae 
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long, with flagellomere I ovoid; flagellomere II short, nearly spherical (0.06 x 0.06 x 
<quantity id="11768AD4995EFFCF39B1F811FDF3C6FD" box="[456,573,1981,2007]" metricMagnitude="-5" metricUnit="m" metricValue="4.0" pageId="1" pageNumber="34" unit="mm" value="0.04">0.04 mm</quantity>
); flagellomere III prolonged medially into a long (
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), densely 
<emphasis id="E4FAFB23995DFFCC38EEFCF7FF34C259" bold="true" box="[151,250,859,883]" pageId="2" pageNumber="35">Figure 1</emphasis>
. Camera lucida drawing of 
<taxonomicName id="118E5CB2995DFFCC3A4CFCF7FD3AC258" box="[565,756,859,882]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="species" species="fossilis" status="sp. nov.">
<emphasis id="E4FAFB23995DFFCC3A4CFCF7FD3AC258" box="[565,756,859,882]" italics="true" pageId="2" pageNumber="35">Litoleptis fossilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="FFC94658995DFFCC3A85FCF7FC9DC258" box="[764,851,859,882]" pageId="2" pageNumber="35" rank="species">sp. nov.</taxonomicNameLabel>
(
<taxonomicName id="118E5CB2995DFFCC3B1BFCF7FC18C258" box="[866,982,859,882]" class="Insecta" family="Spaniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="family">Spaniidae</taxonomicName>
), female 
<typeStatus id="09359993995DFFCC3C44FCF7FB50C258" box="[1085,1182,859,882]" pageId="2" pageNumber="35" type="holotype">holotype</typeStatus>
(CPT-3344), in lateral habitus, from San Just amber. Arrow indicates the metathoracic tibial spur.
</paragraph>
<paragraph id="D6312731995DFFCC38EEFF34FF2DC1D5" pageId="2" pageNumber="35">
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<th id="2B6F4C0F995D003138EEFF34FD8BC198" box="[151,581,152,178]" gridcol="0" gridrow="0" pageId="2" pageNumber="35">pubescent, unsegmented style (basal</th>
<th id="2B6F4C0F995D00313A35FF34FCA2C198" box="[588,876,152,178]" gridcol="1" gridrow="0" pageId="2" pageNumber="35">part bulbous). Proboscis</th>
<th id="2B6F4C0F995D00313B0AFF34FA52C198" box="[883,1436,152,178]" gridcol="2" gridrow="0" pageId="2" pageNumber="35">with labium thin and long, protruding as much</th>
</tr>
<tr id="68BE2573995D003138EEFF12FA52C1F2" box="[151,1436,190,216]" gridrow="1" pageId="2" pageNumber="35">
<th id="2B6F4C0F995D003138EEFF12FD8BC1F2" box="[151,581,190,216]" gridcol="0" gridrow="1" pageId="2" pageNumber="35">as antennae. Only distal part of palps</th>
<td id="2B6F4C0F995D00313A35FF12FCA2C1F2" box="[588,876,190,216]" gridcol="1" gridrow="1" pageId="2" pageNumber="35">visible (0.02 mm width),</td>
<td id="2B6F4C0F995D00313B0AFF12FA52C1F2" box="[883,1436,190,216]" gridcol="2" gridrow="1" pageId="2" pageNumber="35">tapered distallly, having long, distal setae (ca. 6</td>
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<th id="2B6F4C0F995D003138EEFF49FD8BC1D5" box="[151,581,229,255]" gridcol="0" gridrow="2" pageId="2" pageNumber="35">setae).</th>
</tr>
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</paragraph>
<paragraph id="D6312731995DFFCC38BFFC6BFCB9C417" blockId="2.[151,1437,967,2037]" pageId="2" pageNumber="35">
Thorax: Notum high and rounded (Fig. 1). Scutum with numerous setae partially arranged in rows (Pl. 1, fig. 2). Scutellum strongly acuminate in lateral view, with 4 pairs of long setae in dorsal position (Pl. 1, fig. 2). Wings elongate, 
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length, 
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width. C, R4+R5 and h veins strongly sclerotized (Fig. 2d and Pl. 1, fig. 5). R2+3 and distal part of Sc slightly sclerotized. C vein is circumambient but only strongly sclerotized up to end of R5. Stigma large, filling space between veins R1 and R2+3. (Pl.1, fig. 5). R2+3 slightly sinuous. Stem of R2+3 and R4+5 contacts R1 at level of h or slightly basally. R4+5 and M1+2 forked at same distance from wing base and near the distal margin, forming broad, short cells. R4 and R5 end in the dorsal margin. Stem of M separates cells br and bm. Discal cell absent. Vein CuA2 complete, reaching margin of wing but not meeting A1 before wing margin (they separate distally 
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). Basal margin of anal lobe with long, fine setae. Halter bare, ca. 
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long, having elongate knob.
</paragraph>
<paragraph id="D6312731995DFFCC38BFFAE6FEE4C766" blockId="2.[151,1437,967,2037]" pageId="2" pageNumber="35">
Legs long and thin (Fig. 1 and Pl. 1, fig. 1). Protarsus 
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length (tarsomere I 0.25, II 0.10, III 0.08, IV 0.06, V 0.09). Mesothoracic leg 
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length (femur 0.48, tibia 0.59, tarsomere I 0.26, II 0.08, III 0.06, IV 0.04, V 0.06). Metathoracic leg 
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length (femur 0.63, tibia 0.59, tarsomere I 0.21, II 0.09, III 0.07, IV 0.06, V 0.07) (Fig. 1). Mesothoracic tarsomeres with at least four strong distal setae, not present in pro- and metathoracic ones. Very short tibial spur in posterior subapical position on metathoracic leg (Fig. 1). Empodium pulvilliform; pulvilli smaller than empodium (observed in left mesothoracic tarsus) (Fig. 2b and Pl. 1, fig. 4).
</paragraph>
<paragraph id="D6312731995DFFCC38BFF9F5FCA4C7EA" blockId="2.[151,1437,967,2037]" pageId="2" pageNumber="35">
Abdomen broad and short (
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long including terminalia and 
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wide) with abundant setae. Genitalia with segment 1 of cercus not visible (probably short and hidden by tergum). Segment 2 of cercus elongate. Sternum 8 nearly rhombic (Fig. 2c and Pl. 1, fig. 3).
</paragraph>
</subSubSection>
<subSubSection id="9E9474BA995DFFCB38BFF960FDD1C324" lastPageId="5" lastPageNumber="38" pageId="2" pageNumber="35" type="discussion">
<paragraph id="D6312731995DFFCD38BFF960FDD2C1F2" blockId="2.[151,1437,967,2037]" lastBlockId="3.[151,1437,152,216]" lastPageId="3" lastPageNumber="36" pageId="2" pageNumber="35">
<emphasis id="E4FAFB23995DFFCC38BFF960FE9AC7CC" bold="true" box="[198,340,1740,1766]" pageId="2" pageNumber="35">Discussion:</emphasis>
The new species clearly belongs to the 
<taxonomicName id="118E5CB2995DFFCC3B46F960FC7DC7CC" box="[831,947,1740,1766]" class="Insecta" family="Spaniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="family">Spaniidae</taxonomicName>
due to the shape of the antenna and the pulvilliform empodium [it is remarkable that 
<bibRefCitation id="B21F5AC0995DFFCC3AD5F95FFC70C627" author="James" box="[684,958,1779,1805]" pageId="2" pageNumber="39" refString="James, M. T. &amp; Turner, W. J. (1981) [Chapter] 33. Rhagionidae. Pp 483 - 488. In McAlpine, J. F., Peterson, B. V., Shewell, G. E., Teskey, H. J., Vockeroth, J. R. &amp; Wood, D. M. (coords.). Manual of Nearctic Diptera. Volume 1. Research Branch, Agriculture Canada. Monograph No. 27. 674 pp." type="book chapter" year="1981">James &amp; Turner (1981)</bibRefCitation>
erroneusly represented the antenna of 
<taxonomicName id="118E5CB2995DFFCC3DFDF95FFED8C619" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="species" species="alaskensis">
<emphasis id="E4FAFB23995DFFCC3DFDF95FFED8C619" italics="true" pageId="2" pageNumber="35">L. alaskensis</emphasis>
</taxonomicName>
as segmented]. The lack of a discal cell is typical of the genus 
<taxonomicName id="118E5CB2995DFFCC3C49F8B6FB6BC619" box="[1072,1189,1818,1843]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995DFFCC3C49F8B6FB6BC619" box="[1072,1189,1818,1843]" italics="true" pageId="2" pageNumber="35">Litoleptis</emphasis>
</taxonomicName>
but this feature has appeared independently in other taxa. The genus 
<taxonomicName id="118E5CB2995DFFCC3AA7F8EDFB0CC671" authority="Schiner" authorityName="Schiner" box="[734,1218,1857,1883]" class="Insecta" family="Hilarimorphidae" genus="Hilarimorpha" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995DFFCC3AA7F8EDFC4FC670" box="[734,897,1857,1882]" italics="true" pageId="2" pageNumber="35">Hilarimorpha</emphasis>
Schiner (Hilarimorphidae)
</taxonomicName>
has a very similar wing venation (
<bibRefCitation id="B21F5AC0995DFFCC3937F8CBFE16C6AB" author="Webb" box="[334,472,1895,1921]" pageId="2" pageNumber="39" refString="Webb, D. W. (1974) A revision of the genus Hilarimorpha (Diptera: Hilarimorphidae). Journal of the Kansas Entomological Society, 47 (2), 172 - 222." type="journal article" year="1974">Webb 1974</bibRefCitation>
) but the shape of the antenna and the presence of a pulvilliform empodium easily differentiates the new species. Curiously, fossil 
<taxonomicName id="118E5CB2995DFFCC3AB4F822FC5EC682" box="[717,912,1934,1960]" class="Insecta" family="Hilarimorphidae" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="family">Hilarimorphidae</taxonomicName>
have a well developed discal cell as well as does the extant genus 
<taxonomicName id="118E5CB2995DFFCC39DAF819FD3DC6E4" authority="Melander, 1950" authorityName="Melander" authorityYear="1950" box="[419,755,1972,1998]" class="Insecta" family="Apystomyiidae" genus="Apystomyia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995DFFCC39DAF819FDE1C6E4" box="[419,559,1973,1998]" italics="true" pageId="2" pageNumber="35">Apystomyia</emphasis>
Melander, 1950
</taxonomicName>
(
<bibRefCitation id="B21F5AC0995DFFCC3B7DF818FB97C6E4" author="Grimaldi" box="[772,1113,1972,1998]" pageId="2" pageNumber="38" refString="Grimaldi, D. A. &amp; Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1 - 124." type="journal article" year="1999">Grimaldi &amp; Cumming 1999</bibRefCitation>
; 
<bibRefCitation id="B21F5AC0995DFFCC3C1FF818FAE0C6E4" author="Mostovski" box="[1126,1326,1972,1998]" pageId="2" pageNumber="39" refString="Mostovski, M. B. (1999) On an interesting find of a brachycerous fly (Diptera, Brachycera) in the Jurassic of Kazakhstan. Paleontological Journal, 33 (4), 70 - 72." type="journal article" year="1999">Mostovski 1999</bibRefCitation>
). Among rhagionoids, loss of the discal cell is also a character present in one species described by 
<bibRefCitation id="B21F5AC0995DFFCC3CC6F877FA95C6DF" author="Webb" box="[1215,1371,2011,2037]" pageId="2" pageNumber="39" refString="Webb, D. W. (1969) A new and unusual species of Bolbomyia (Diptera: Rhagionidae). Journal of the Kansas Entomological Society, 42 (3), 285 - 288." type="journal article" year="1969">Webb (1969)</bibRefCitation>
from New York, 
<collectingCountry id="AE9967A1995CFFCD395CFF34FEA9C198" box="[293,359,152,178]" name="United States of America" pageId="3" pageNumber="36">USA</collectingCountry>
: 
<taxonomicName id="118E5CB2995CFFCD390DFF34FC86C198" authority="Webb, 1969" authorityName="Webb" authorityYear="1969" box="[372,840,152,178]" class="Insecta" family="Bolbomyiidae" genus="Bolbomyia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="andiscalcella">
<emphasis id="E4FAFB23995CFFCD390DFF34FD68C19B" box="[372,678,152,177]" italics="true" pageId="3" pageNumber="36">Bolbomyia andiscalcella</emphasis>
<bibRefCitation id="B21F5AC0995CFFCD3AD6FF34FC86C198" author="Webb" box="[687,840,152,178]" pageId="3" pageNumber="39" refString="Webb, D. W. (1969) A new and unusual species of Bolbomyia (Diptera: Rhagionidae). Journal of the Kansas Entomological Society, 42 (3), 285 - 288." type="journal article" year="1969">Webb, 1969</bibRefCitation>
</taxonomicName>
. However, the antenna of 
<taxonomicName id="118E5CB2995CFFCD3CF5FF34FA9AC19B" box="[1164,1364,152,177]" class="Insecta" family="Bolbomyiidae" genus="Bolbomyia" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="andiscalcella">
<emphasis id="E4FAFB23995CFFCD3CF5FF34FA9AC19B" box="[1164,1364,152,177]" italics="true" pageId="3" pageNumber="36">B. andiscalcella</emphasis>
</taxonomicName>
has a clearly segmented antennal style.
</paragraph>
<paragraph id="D6312731995CFFCD38EEF93CFD75C7C7" blockId="3.[151,1436,1680,1773]" pageId="3" pageNumber="36">
<emphasis id="E4FAFB23995CFFCD38EEF93CFF38C782" bold="true" box="[151,246,1680,1704]" pageId="3" pageNumber="36">Figure 2</emphasis>
. Camera lucida drawings of 
<taxonomicName id="118E5CB2995CFFCD3A55F93DFD2CC782" box="[556,738,1681,1704]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="fossilis" status="sp. nov.">
<emphasis id="E4FAFB23995CFFCD3A55F93DFD2CC782" box="[556,738,1681,1704]" italics="true" pageId="3" pageNumber="36">Litoleptis fossilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="FFC94658995CFFCD3A90F93DFCF2C782" box="[745,828,1681,1704]" pageId="3" pageNumber="36" rank="species">sp. nov.</taxonomicNameLabel>
(
<taxonomicName id="118E5CB2995CFFCD3B33F93DFC77C782" box="[842,953,1681,1704]" class="Insecta" family="Spaniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="family">Spaniidae</taxonomicName>
), female 
<typeStatus id="09359993995CFFCD3C62F93DFBB6C782" box="[1051,1144,1681,1704]" pageId="3" pageNumber="36" type="holotype">holotype</typeStatus>
(CPT-3344). Details of the a) head, b) distal mesotarsomere showing claws, pulvilli and the pulvilliform empodium, c) female terminalia in dorsolateral (top), lateral and ventral views, and d) wing.
</paragraph>
<paragraph id="D6312731995CFFCA38BFF8B3FF3EC6CD" blockId="3.[151,1437,1823,2004]" lastBlockId="4.[151,1436,1896,2023]" lastPageId="4" lastPageNumber="37" pageId="3" pageNumber="36">
The new species is differentiated from the extant species of 
<taxonomicName id="118E5CB2995CFFCD3BB4F88CFBF5C613" box="[973,1083,1824,1849]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995CFFCD3BB4F88CFBF5C613" box="[973,1083,1824,1849]" italics="true" pageId="3" pageNumber="36">Litoleptis</emphasis>
</taxonomicName>
by having a small spur on metathoracic tibia. It is also easy to distinguish the new species from 
<taxonomicName id="118E5CB2995CFFCD3BBBF8EAFB97C675" box="[962,1113,1862,1887]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="alaskensis">
<emphasis id="E4FAFB23995CFFCD3BBBF8EAFB97C675" box="[962,1113,1862,1887]" italics="true" pageId="3" pageNumber="36">L. alaskensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="118E5CB2995CFFCD3CE8F8EAFAEEC675" box="[1169,1312,1862,1887]" class="Insecta" family="Hilarimorphidae" genus="Hilarimorpha" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="orientalis">
<emphasis id="E4FAFB23995CFFCD3CE8F8EAFAEEC675" box="[1169,1312,1862,1887]" italics="true" pageId="3" pageNumber="36">L. orientalis</emphasis>
</taxonomicName>
by having a complete CuA vein (
<bibRefCitation id="B21F5AC0995CFFCD39C4F8C1FED9C687" author="Grimaldi" pageId="3" pageNumber="38" refString="Grimaldi, D. A. &amp; Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1 - 124." type="journal article" year="1999" yearSuffix="a">
Grimaldi &amp; Cumming, 1999 erroneously figured a complete CuA vein in 
<taxonomicName id="118E5CB2995CFFCD3DFAF8C1FED9C687" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="alaskensis">
<emphasis id="E4FAFB23995CFFCD3DFAF8C1FED9C687" italics="true" pageId="3" pageNumber="36">L. alaskensis</emphasis>
</taxonomicName>
</bibRefCitation>
). 
<taxonomicName id="118E5CB2995CFFCD394BF838FDC9C687" box="[306,519,1940,1965]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="fossilis" status="sp. nov.">
<emphasis id="E4FAFB23995CFFCD394BF838FDC9C687" box="[306,519,1940,1965]" italics="true" pageId="3" pageNumber="36">Litoleptis fossilis</emphasis>
</taxonomicName>
<emphasis id="E4FAFB23995CFFCD3A69F83FFDBAC687" bold="true" box="[528,628,1939,1965]" pageId="3" pageNumber="36">
<taxonomicNameLabel id="FFC94658995CFFCD3A69F83FFDBAC687" box="[528,628,1939,1965]" pageId="3" pageNumber="36" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is differentiated from 
<taxonomicName id="118E5CB2995CFFCD3BE9F838FBD1C687" box="[912,1055,1940,1965]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="36" phylum="Arthropoda" rank="species" species="chilensis">
<emphasis id="E4FAFB23995CFFCD3BE9F838FBD1C687" box="[912,1055,1940,1965]" italics="true" pageId="3" pageNumber="36">L. chilensis</emphasis>
</taxonomicName>
and from the two undescribed Japanese and Nepalese species (
<bibRefCitation id="B21F5AC0995CFFCD3A5BF816FD2AC6FE" author="Nagatomi" box="[546,740,1978,2004]" pageId="3" pageNumber="39" refString="Nagatomi, A. (1982) The genera of Rhagionidae (Diptera). Journal of Natural History, 16, 31 - 70." type="journal article" year="1982">Nagatomi 1982</bibRefCitation>
) by the shape of the third antennal segment. In the new 
<emphasis id="E4FAFB23995BFFCA38EEF8C4FF2BC6AA" bold="true" box="[151,229,1896,1920]" pageId="4" pageNumber="37">Plate 1</emphasis>
: 
<taxonomicName id="118E5CB2995BFFCA388DF8C5FE62C6AA" box="[244,428,1897,1920]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="37" phylum="Arthropoda" rank="species" species="fossilis" status="sp. nov.">
<emphasis id="E4FAFB23995BFFCA388DF8C5FE62C6AA" box="[244,428,1897,1920]" italics="true" pageId="4" pageNumber="37">Litoleptis fossilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="FFC94658995BFFCA39CAF8C4FDC9C655" box="[435,519,1896,1919]" pageId="4" pageNumber="37" rank="species">sp. nov.</taxonomicNameLabel>
(
<taxonomicName id="118E5CB2995BFFCA3A6CF8C4FD4BC655" box="[533,645,1896,1919]" class="Insecta" family="Spaniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="37" phylum="Arthropoda" rank="family">Spaniidae</taxonomicName>
), female 
<typeStatus id="09359993995BFFCA3A91F8C4FC88C655" box="[744,838,1896,1919]" pageId="4" pageNumber="37" type="holotype">holotype</typeStatus>
(CPT-3344), from San Just amber. 1) habitus in dorsal, ventral and lateral (below) views, 2) thorax and head in dorsal view showing chaetotaxy in scutum and scutellum, 3) genitalia in dorso-lateral view, 4) distal left mesotarsomere showing claws, pulvilli and the pulvilliform empodium, and 5) wing.
</paragraph>
<paragraph id="D6312731995AFFCB38EEFF34FE54C00C" blockId="5.[151,1437,152,796]" pageId="5" pageNumber="38">species R4 and R5 end in the costal margin, while in the wing apex in all extant species whose venation is known, and R4+5 and M1+2 forked at same distance from wing base and near the distal margin, determining broad, short cells. Also, new species is separated from extant ones by having a long labium that protrudes as much as the antennae.</paragraph>
<paragraph id="D6312731995AFFCB38BFFE9EFB9DC059" blockId="5.[151,1437,152,796]" pageId="5" pageNumber="38">
The female genitalia of extant species of 
<taxonomicName id="118E5CB2995AFFCB3AD5FE9EFCD4C061" box="[684,794,306,331]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995AFFCB3AD5FE9EFCD4C061" box="[684,794,306,331]" italics="true" pageId="5" pageNumber="38">Litoleptis</emphasis>
</taxonomicName>
are poorly known. The genitalia of 
<taxonomicName id="118E5CB2995AFFCB3CBCFE9EFAFBC061" box="[1221,1333,306,331]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="species" species="fossilis" status="sp. nov.">
<emphasis id="E4FAFB23995AFFCB3CBCFE9EFAFBC061" box="[1221,1333,306,331]" italics="true" pageId="5" pageNumber="38">L. fossilis</emphasis>
</taxonomicName>
<emphasis id="E4FAFB23995AFFCB3D47FE9EFA55C066" bold="true" box="[1342,1435,306,332]" pageId="5" pageNumber="38">
<taxonomicNameLabel id="FFC94658995AFFCB3D47FE9EFA55C066" box="[1342,1435,306,332]" pageId="5" pageNumber="38" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
seems closely related to genera 
<taxonomicName id="118E5CB2995AFFCB3A72FEF5FD93C058" box="[523,605,345,370]" class="Insecta" family="Rhagionidae" genus="Spania" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995AFFCB3A72FEF5FD93C058" box="[523,605,345,370]" italics="true" pageId="5" pageNumber="38">
<collectingCountry id="AE9967A1995AFFCB3A72FEF5FD93C058" box="[523,605,345,370]" name="Spain" pageId="5" pageNumber="38">Spania</collectingCountry>
</emphasis>
</taxonomicName>
and 
<taxonomicName id="118E5CB2995AFFCB3AECFEF5FB81C059" authority="Nagatomi &amp; Iwata 1976" authorityName="Nagatomi &amp; Iwata" authorityYear="1976" box="[661,1103,345,371]" class="Insecta" family="Rhagionidae" genus="Spaniopsis" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995AFFCB3AECFEF5FCDAC058" box="[661,788,345,370]" italics="true" pageId="5" pageNumber="38">Spaniopsis</emphasis>
(
<bibRefCitation id="B21F5AC0995AFFCB3B5AFEF5FB8BC059" author="Nagatomi" box="[803,1093,345,371]" pageId="5" pageNumber="39" refString="Nagatomi, A. &amp; Iwata, K. (1976) Female terminalia of lower Brachycera - I (Diptera). Beitrage zur Entomologie, 26, 5 - 47." type="journal article" year="1976">Nagatomi &amp; Iwata 1976</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="D6312731995AFFCB38BFFED3FDD1C324" blockId="5.[151,1437,152,796]" pageId="5" pageNumber="38">
<emphasis id="E4FAFB23995AFFCB38BFFED3FE4EC0B3" bold="true" box="[198,384,383,409]" pageId="5" pageNumber="38">Palaeoecology:</emphasis>
<taxonomicName id="118E5CB2995AFFCB39FEFE2CFD83C0B3" box="[391,589,384,409]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="species" species="fossilis" status="sp. nov.">
<emphasis id="E4FAFB23995AFFCB39FEFE2CFD83C0B3" box="[391,589,384,409]" italics="true" pageId="5" pageNumber="38">Litoleptis fossilis</emphasis>
</taxonomicName>
<emphasis id="E4FAFB23995AFFCB3A2CFED3FD7CC0B3" bold="true" box="[597,690,383,409]" pageId="5" pageNumber="38">
<taxonomicNameLabel id="FFC94658995AFFCB3A2CFED3FD7CC0B3" box="[597,690,383,409]" pageId="5" pageNumber="38" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
has a long proboscis and could have been a hematophagous fly. In fact, it has a proboscis larger than the proboscis of ceratopogonids from the same outcrop (see 
<bibRefCitation id="B21F5AC0995AFFCB3D6AFE0AFF19C0CD" author="Arillo" pageId="5" pageNumber="38" refString="Arillo, A., Penalver, E. &amp; Delclos, X. (2008) Microphorites (Diptera: Dolichopodidae) from the Lower Cretaceous amber of San Just (Spain), and the co-occurrence of two ceratopogonid species in Spanish amber deposits. Zootaxa, 1920, 29 - 40." type="journal article" year="2008">
Arillo 
<emphasis id="E4FAFB23995AFFCB3D18FE0BFA55C0EA" box="[1377,1435,423,448]" italics="true" pageId="5" pageNumber="38">et al.</emphasis>
2008
</bibRefCitation>
). Blood-feeding behaviour is not common among living rhagionoids, occurring only in some 
<taxonomicName id="118E5CB2995AFFCB3D51FE61FA55C0CD" box="[1320,1435,461,487]" class="Insecta" family="Spaniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="family">Spaniidae</taxonomicName>
and Rhagioniinae (
<bibRefCitation id="B21F5AC0995AFFCB390DFE58FDDDC324" author="Lehane" box="[372,531,500,526]" pageId="5" pageNumber="39" refString="Lehane, M. J. (2005) The biology of blood-sucking in insects. 2 nd Edition. Cambridge University Press, Cambridge. 321 pp." type="book" year="2005">Lehane 2005</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="9E9474BA995AFFCB38BFFDB6FBE5C236" pageId="5" pageNumber="38" type="description">
<paragraph id="D6312731995AFFCB38BFFDB6FBE5C236" blockId="5.[151,1437,152,796]" pageId="5" pageNumber="38">
<emphasis id="E4FAFB23995AFFCB38BFFDB6FE00C31E" bold="true" box="[198,462,538,564]" pageId="5" pageNumber="38">Palaeobiogeography:</emphasis>
Rhagionoids are among the earliest and most diverse brachycerans in the fossil record. The finding of a Cretaceous 
<taxonomicName id="118E5CB2995AFFCB3A19FDEDFD19C370" box="[608,727,577,602]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995AFFCB3A19FDEDFD19C370" box="[608,727,577,602]" italics="true" pageId="5" pageNumber="38">Litoleptis</emphasis>
</taxonomicName>
proves that 
<taxonomicName id="118E5CB2995AFFCB3B0DFDEDFC21C371" box="[884,1007,577,603]" class="Insecta" family="Spaniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="family">Spaniidae</taxonomicName>
is an old lineage that was already separated from the rest of rhagionoids in the Cretaceous. However, its appearance in Spanish amber is quite surprising if we consider its currently known Asio-Nearctic disjunt distribution (
<bibRefCitation id="B21F5AC0995AFFCB3C27FD22FAC8C382" author="Saigusa" box="[1118,1286,654,680]" pageId="5" pageNumber="39" refString="Saigusa, T. (2006) Asio-Nearctic disjunct distribution in Diptera. 6 th International Congress of Dipterology. Fukuoka, Japan. 23 - 28 September 2006. Abstracts Volume: p. 209." type="journal volume" year="2006">Saigusa 2006</bibRefCitation>
). Obviously, the genus had a wider distribution during the Mesozoic, but a more precise palaeogeographic distribution can be understood only when further fossil records are available. An undescribed specimen of 
<taxonomicName id="118E5CB2995AFFCB3CC2FD70FAE7C3DF" box="[1211,1321,732,757]" class="Insecta" family="Rhagionidae" genus="Litoleptis" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="38" phylum="Arthropoda" rank="genus">
<emphasis id="E4FAFB23995AFFCB3CC2FD70FAE7C3DF" box="[1211,1321,732,757]" italics="true" pageId="5" pageNumber="38">Litoleptis</emphasis>
</taxonomicName>
is known from the late Eocene of Florissant (Colorado, 
<collectingCountry id="AE9967A1995AFFCB3AC8FCAEFD23C236" box="[689,749,770,796]" name="United States of America" pageId="5" pageNumber="38">USA</collectingCountry>
) (Mostovski pers. comm.).
</paragraph>
</subSubSection>
</treatment>
</document>