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<mods:title id="AF7B9C1D42643314D4CD50F0DB10EC1E">New fossil lace bugs (Heteroptera: Tingidae) from the Middle Eocene of the Grube Messel (Germany), with a catalog of fossil lace bugs</mods:title>
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<subSubSection id="138B6543FF9E3B07FEF10553E72BD692" box="[264,527,638,664]" pageId="3" pageNumber="4" type="nomenclature">
<paragraph id="5B2E36C8FF9E3B07FEF10553E72BD692" blockId="3.[264,527,638,664]" box="[264,527,638,664]" pageId="3" pageNumber="4">
<heading id="006681A4FF9E3B07FEF10553E72BD692" bold="true" box="[264,527,638,664]" fontSize="11" level="1" pageId="3" pageNumber="4" reason="1">
<emphasis id="69E5EADAFF9E3B07FEF10553E72BD692" bold="true" box="[264,527,638,664]" pageId="3" pageNumber="4">
<taxonomicName id="9C914D4BFF9E3B07FEF10553E492D692" box="[264,438,638,664]" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="69E5EADAFF9E3B07FEF10553E492D692" bold="true" box="[264,438,638,664]" italics="true" pageId="3" pageNumber="4">Exmesselensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="72D657A1FF9E3B07FE470553E72BD692" box="[446,527,638,664]" pageId="3" pageNumber="4" rank="genus">n. gen.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="138B6543FF9E3B07FEF105E4E6DAD6E9" box="[264,1022,713,739]" pageId="3" pageNumber="4" type="materials_examined">
<paragraph id="5B2E36C8FF9E3B07FEF105E4E6DAD6E9" blockId="3.[264,1325,713,1899]" box="[264,1022,713,739]" pageId="3" pageNumber="4">
<emphasis id="69E5EADAFF9E3B07FEF105E4E488D6E9" bold="true" box="[264,428,713,739]" pageId="3" pageNumber="4">
<typeStatus id="842A886AFF9E3B07FEF105E4E461D6E9" box="[264,325,713,739]" pageId="3" pageNumber="4">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="9C914D4BFF9E3B07FE4D05E4E7CED6E8" box="[436,746,713,738]" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="disspinosus" status="sp. nov.">
<emphasis id="69E5EADAFF9E3B07FE4D05E4E7CED6E8" box="[436,746,713,738]" italics="true" pageId="3" pageNumber="4">Exmesselensis disspinosus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="72D657A1FF9E3B07FD0B05E4E616D6E9" box="[754,818,713,739]" pageId="3" pageNumber="4" rank="species">n. sp.</taxonomicNameLabel>
, here designated.
</paragraph>
</subSubSection>
<subSubSection id="138B6543FF9E3B07FEC105DCE1A7D739" pageId="3" pageNumber="4" type="etymology">
<paragraph id="5B2E36C8FF9E3B07FEC105DCE1A7D739" blockId="3.[264,1325,713,1899]" pageId="3" pageNumber="4">
<emphasis id="69E5EADAFF9E3B07FEC105DCE700D701" bold="true" box="[312,548,753,779]" pageId="3" pageNumber="4">Derivatio nominis:</emphasis>
The new genus­group name is derived from
<emphasis id="69E5EADAFF9E3B07FBB005DCE147D700" box="[1097,1123,753,778]" italics="true" pageId="3" pageNumber="4">ex</emphasis>
(Latin, meaning “out of”), plus the name of the
<typeStatus id="842A886AFF9E3B07FD8C0434E783D739" box="[629,679,793,819]" pageId="3" pageNumber="4">type</typeStatus>
locality
<emphasis id="69E5EADAFF9E3B07FCF70434E67BD738" box="[782,863,793,818]" italics="true" pageId="3" pageNumber="4">Messel</emphasis>
. The name is masculine.
</paragraph>
</subSubSection>
<subSubSection id="138B6543FF9E3B07FEC1046CE1DBD041" pageId="3" pageNumber="4" type="diagnosis">
<paragraph id="5B2E36C8FF9E3B07FEC1046CE1DBD041" blockId="3.[264,1325,713,1899]" pageId="3" pageNumber="4">
<emphasis id="69E5EADAFF9E3B07FEC1046CE49ED751" bold="true" box="[312,442,833,859]" pageId="3" pageNumber="4">Diagnosis:</emphasis>
Small specimen, ca.
<quantity id="9C699B2DFF9E3B07FD38046CE605D751" box="[705,801,833,859]" metricMagnitude="-3" metricUnit="m" metricValue="2.5" pageId="3" pageNumber="4" unit="mm" value="2.5">2.5 mm</quantity>
. Body moderately oblong­oval. Head elongated, with one significant interocular (dorsomedial) tubercle. Bucculae elongated forward. Eyes globular, strongly projecting laterally of the head. Antennae thin, 3rd segment long: just a little shorter than body width (relation 0.91). Pronotum more or less trapezoid, with 3 longitudinal carinae; posterior margin concave. Paranotum narrow. Scutellum completely exposed, triangular. Hemelytra distinctly subdivides by raised veins into costal, subcostal, discoidal, and sutural areas. Stenocostal area absent. Clavus well expressed.
</paragraph>
</subSubSection>
<subSubSection id="138B6543FF9E3B01FEC10374E6CAD5DC" lastPageId="5" lastPageNumber="6" pageId="3" pageNumber="4" type="discussion">
<paragraph id="5B2E36C8FF9E3B07FEC10374E701D1B9" blockId="3.[264,1325,713,1899]" pageId="3" pageNumber="4">
<emphasis id="69E5EADAFF9E3B07FEC10374E4ECD079" bold="true" box="[312,456,1113,1139]" pageId="3" pageNumber="4">Comments:</emphasis>
The new genus
<taxonomicName id="9C914D4BFF9E3B07FD760374E612D078" box="[655,822,1113,1138]" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="6" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="69E5EADAFF9E3B07FD760374E612D078" box="[655,822,1113,1138]" italics="true" pageId="3" pageNumber="4">Exmesselensis</emphasis>
</taxonomicName>
combines some morphological characters of the Cantacaderinae and the
<taxonomicName id="9C914D4BFF9E3B07FD7E03ACE7D6D091" box="[647,754,1153,1179]" class="Insecta" family="Tingidae" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="subFamily" subFamily="Tinginae">Tinginae</taxonomicName>
. The features of Cantacaderinae, in particular
<emphasis id="69E5EADAFF9E3B07FEF10384E4BED0C9" bold="true" box="[264,410,1193,1219]" pageId="3" pageNumber="4">Phatnomini</emphasis>
, includes: (1) the absence of stenocostal area of hemelytra (unfortunately it was impossible to examine in detail the scent gland openings and the morphology of the peritrema), (2) presents of only one dorsomedial (interocular) tubercle (completely absent in Cantacaderini and characteristic for Phatnomini), (3) the absence of a well expressed posterior triangular projection of the pronotum, (4) the exposed scutellum, (5) bucculae elongated forward and not touching anteriorly, and (6) clavi clearly separated by Pcu from corium by commissural.
</paragraph>
<paragraph id="5B2E36C8FF9E3B07FEC102ECE006D221" blockId="3.[264,1325,713,1899]" pageId="3" pageNumber="4">
The absence of additional transverse veins in the discoidal area on the other hand is a characteristic feature of the majority of
<taxonomicName id="9C914D4BFF9E3B07FD2102C4E61BD209" box="[728,831,1513,1539]" class="Insecta" family="Tingidae" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="subFamily" subFamily="Tinginae">Tinginae</taxonomicName>
contrary to Cantacaderinae in which they are usually well expressed in different degree, especially in Phatnomini (
<bibRefCitation id="3F004B39FF9E3B07FBB5013CE031D221" author="Froeschner" box="[1100,1301,1553,1579]" pageId="3" pageNumber="18" refString="Froeschner, R. C. (1996) Lace bug genera of the world, I: Introduction, Subfamily Cantacaderinae (Heteroptera: Tingidae). Smithsonian Contributions to Zoology, 574, 1 - 43." type="journal article" year="1996">Froeschner 1996</bibRefCitation>
).
</paragraph>
<paragraph id="5B2E36C8FF9E3B07FEC10114E4F0D361" blockId="3.[264,1325,713,1899]" pageId="3" pageNumber="4">
Among those genera of the Phatnomini bearing a dorsomedial tubercle, habitually, the fossil genus
<taxonomicName id="9C914D4BFF9E3B07FE63014CE766D270" box="[410,578,1633,1658]" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="69E5EADAFF9E3B07FE63014CE766D270" box="[410,578,1633,1658]" italics="true" pageId="3" pageNumber="4">Exmesselensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="72D657A1FF9E3B07FDB1014CE7B3D271" box="[584,663,1633,1659]" pageId="3" pageNumber="4" rank="genus">n. gen.</taxonomicNameLabel>
is superficially quite similar to the living lace bug of the genus
<taxonomicName id="9C914D4BFF9E3B07FEAF01A4E7A0D2A9" authority="Froeschner. It" authorityName="Froeschner. It" box="[342,644,1673,1699]" class="Insecta" family="Tingidae" genus="Distocader" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">
<emphasis id="69E5EADAFF9E3B07FEAF01A4E4FCD2A8" box="[342,472,1673,1698]" italics="true" pageId="3" pageNumber="4">Distocader</emphasis>
Froeschner. It
</taxonomicName>
shares in particular the head and antennal structure with the former.
<taxonomicName id="9C914D4BFF9E3B07FE61019CE764D2C0" box="[408,576,1713,1738]" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="69E5EADAFF9E3B07FE61019CE764D2C0" box="[408,576,1713,1738]" italics="true" pageId="3" pageNumber="4">Exmesselensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="72D657A1FF9E3B07FDB5019CE7BBD2C1" box="[588,671,1713,1739]" pageId="3" pageNumber="4" rank="genus">n. gen.</taxonomicNameLabel>
is distinguished from this endemic genus by relative broader pronotum, shorter carinae of the pronotum (majority of the Phatnomini show a 1­ carinated pronotal disc), and the absence transverse veins in the discoidal area (this affiliates the fossil genus with the Phatnomini genera
<taxonomicName id="9C914D4BFF9E3B07FCA90004E11DD349" authority="Pericart" authorityName="Pericart" box="[848,1081,1833,1859]" class="Insecta" family="Tingidae" genus="Pullocader" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">
<emphasis id="69E5EADAFF9E3B07FCA90004E6F6D348" box="[848,978,1833,1858]" italics="true" pageId="3" pageNumber="4">Pullocader</emphasis>
Pericart
</taxonomicName>
,
<taxonomicName id="9C914D4BFF9E3B07FBBF0004E1D0D349" authority="Distant" authorityName="Distant" box="[1094,1268,1833,1859]" class="Insecta" family="Tingidae" genus="Ulmus" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">
<emphasis id="69E5EADAFF9E3B07FBBF0004E1B6D348" box="[1094,1170,1833,1858]" italics="true" pageId="3" pageNumber="4">Ulmus</emphasis>
Distant
</taxonomicName>
, and
<taxonomicName id="9C914D4BFF9E3B07FEF1007CE4ECD361" authority="Drake" authorityName="Drake" box="[264,456,1873,1899]" class="Insecta" family="Tingidae" genus="Zetekella" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">
<emphasis id="69E5EADAFF9E3B07FEF1007CE457D360" box="[264,371,1873,1898]" italics="true" pageId="3" pageNumber="4">Zetekella</emphasis>
Drake
</taxonomicName>
).
</paragraph>
<paragraph id="5B2E36C8FF993B00FEC10631E6B9D58C" blockId="4.[264,1323,284,390]" pageId="4" pageNumber="5">
Nevertheless, the incomplete state of material, however, does not permit an evaluation of its closer relationships at the present. A reconstruction of the
<typeStatus id="842A886AFF993B00FBFA0669E112D554" box="[1027,1078,324,350]" pageId="4" pageNumber="5">type</typeStatus>
species of
<taxonomicName id="9C914D4BFF993B00FB450669E46FD58F" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="6" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="69E5EADAFF993B00FB450669E46FD58F" italics="true" pageId="4" pageNumber="5">Exmesselensis</emphasis>
</taxonomicName>
based on the description below is shown in
<figureCitation id="C3AA2A4DFF993B00FCA80641E6BDD58C" box="[849,921,364,390]" captionStart="FIGURE 2" captionStartId="4.[264,368,1541,1565]" captionTargetBox="[461,1161,454,1498]" captionTargetId="figure@4.[435,1161,449,1512]" captionTargetPageId="4" captionText="FIGURE 2. Exmesselensis disspinosus gen. et sp. n., reconstruction of the holotype, (macropterous form), compression fossil; coll. Forschungsinstitut Senckenberg (SMF), Frankfurt am Main (Germany), specimen SMF MeI 6301; scale bar represents 0.5 mm." httpUri="https://zenodo.org/record/156770/files/figure.png" pageId="4" pageNumber="5">Fig. 2</figureCitation>
.
</paragraph>
<caption id="0FEE6640FF993B00FEF10128E6FDD268" httpUri="https://zenodo.org/record/156770/files/figure.png" pageId="4" pageNumber="5" targetBox="[461,1161,454,1498]" targetPageId="4">
<paragraph id="5B2E36C8FF993B00FEF10128E6FDD268" blockId="4.[264,1323,1541,1634]" pageId="4" pageNumber="5">
<emphasis id="69E5EADAFF993B00FEF10128E4A8D217" bold="true" box="[264,396,1541,1565]" pageId="4" pageNumber="5">FIGURE 2.</emphasis>
<taxonomicName id="9C914D4BFF993B00FE6D012BE794D217" box="[404,688,1542,1565]" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="disspinosus">
<emphasis id="69E5EADAFF993B00FE6D012BE794D217" box="[404,688,1542,1565]" italics="true" pageId="4" pageNumber="5">Exmesselensis disspinosus</emphasis>
</taxonomicName>
<emphasis id="69E5EADAFF993B00FD410128E66DD217" bold="true" box="[696,841,1541,1565]" pageId="4" pageNumber="5">gen. et sp. n.</emphasis>
, reconstruction of the holotype, (macropterous form), compression fossil; coll. Forschungsinstitut Senckenberg (SMF), Frankfurt am Main (Germany), specimen SMF MeI 6301; scale bar represents 0.5 mm.
</paragraph>
</caption>
<paragraph id="5B2E36C8FF993B00FEC101BBE498D35A" blockId="4.[264,1323,1686,1872]" pageId="4" pageNumber="5">
The distribution of the extant representatives of the Phatnomini today is restricted to the southern hemisphere. The majority of the 26 described, extant genera are distributed in the Oriental and Ethiopian zoogeographic regions (
<bibRefCitation id="3F004B39FF993B00FC9601CBE119D30A" author="Froeschner" box="[879,1085,1766,1792]" pageId="4" pageNumber="18" refString="Froeschner, R. C. (1996) Lace bug genera of the world, I: Introduction, Subfamily Cantacaderinae (Heteroptera: Tingidae). Smithsonian Contributions to Zoology, 574, 1 - 43." type="journal article" year="1996">Froeschner 1996</bibRefCitation>
). For instance,
<taxonomicName id="9C914D4BFF993B00FB0101CBE444D32D" class="Insecta" family="Tingidae" genus="Distocader" kingdom="Animalia" order="Hemiptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="69E5EADAFF993B00FB0101CBE444D32D" italics="true" pageId="4" pageNumber="5">Distocader</emphasis>
</taxonomicName>
, represented by only a single species, is only known from New
<collectingCountry id="23867658FF993B00FBA40023E196D322" box="[1117,1202,1806,1832]" name="Guinea-Bissau" pageId="4" pageNumber="5">Guinea</collectingCountry>
(
<bibRefCitation id="3F004B39FF993B00FB3B0023E494D35A" author="Drake" pageId="4" pageNumber="18" refString="Drake, C. J. &amp; Ruhoff, F. A. (1965 b) Lace bugs from New Guinea, Borneo, Solomons, and other islands of the South Pacific and Indian Oceans (Hemiptera: Tingidae). Pacific Insects, 7, 243 - 290." type="journal article" year="1965" yearSuffix="b">Drake &amp; Ruhoff 1965b</bibRefCitation>
).
</paragraph>
<paragraph id="5B2E36C8FF983B01FEC10631E6CAD5DC" blockId="5.[264,1323,284,470]" pageId="5" pageNumber="6">
Furthermore
<taxonomicName id="9C914D4BFF983B01FE1D0631E7A8D53F" box="[484,652,284,309]" class="Insecta" family="Tingidae" genus="Exmesselensis" kingdom="Animalia" order="Hemiptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="69E5EADAFF983B01FE1D0631E7A8D53F" box="[484,652,284,309]" italics="true" pageId="5" pageNumber="6">Exmesselensis</emphasis>
</taxonomicName>
shows once more those representatives of the Cantacaderinae were much more significant in the Paleogene fauna and wide­spread in Europe than today. In particular some genera of the Eocene European Phatnomini (e.g.
<taxonomicName id="9C914D4BFF983B01FEF106B9E150D5A7" authority="Golub &amp; Popov, Tingicader Golub &amp; Popov, Exmesselensis" authorityName="Golub &amp; Popov, Tingicader Golub &amp; Popov, Exmesselensis" box="[264,1140,404,430]" class="Insecta" family="Tingidae" genus="Intercader" kingdom="Animalia" order="Hemiptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="69E5EADAFF983B01FEF106B9E4A6D5A7" box="[264,386,404,429]" italics="true" pageId="5" pageNumber="6">Intercader</emphasis>
Golub &amp; Popov,
<emphasis id="69E5EADAFF983B01FD9106B9E7C3D5A7" box="[616,743,404,429]" italics="true" pageId="5" pageNumber="6">Tingicader</emphasis>
Golub &amp; Popov,
<emphasis id="69E5EADAFF983B01FC3506B9E150D5A7" box="[972,1140,404,429]" italics="true" pageId="5" pageNumber="6">Exmesselensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="72D657A1FF983B01FB7806B9E1F3D5A4" box="[1153,1239,404,430]" pageId="5" pageNumber="6" rank="genus">n. gen.</taxonomicNameLabel>
) show morphological similarities to the advanced subfamily
<taxonomicName id="9C914D4BFF983B01FC860691E6CED5DC" box="[895,1002,444,470]" class="Insecta" family="Tingidae" kingdom="Animalia" order="Hemiptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subFamily" subFamily="Tinginae">Tinginae</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>