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<mods:title id="66A1FBEEDD30E059CFAAED738BFA60EA">Fossil Darwin wasps (Hymenoptera: Ichneumonidae) from Baltic amber</mods:title>
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<subSubSection id="4FC775D9FF9BFFF6FF0BFBFEFDDCFE01" box="[136,601,446,472]" pageId="3" pageNumber="640" type="nomenclature">
<paragraph id="07622652FF9BFFF6FF0BFBFEFDDCFE01" blockId="3.[136,601,446,472]" box="[136,601,446,472]" pageId="3" pageNumber="640">
<heading id="5C2A913EFF9BFFF6FF0BFBFEFDDCFE01" box="[136,601,446,472]" fontSize="10" level="2" pageId="3" pageNumber="640" reason="2">
<emphasis id="35A9FA40FF9BFFF6FF0BFBFEFDDCFE01" bold="true" box="[136,601,446,472]" pageId="3" pageNumber="640">
Subfamily
<taxonomicName id="C0DD5DD1FF9BFFF6FE89FBFEFDDCFE01" authority="Viereck, 1918" authorityName="Viereck" authorityYear="1918" box="[266,601,446,472]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="subFamily" subFamily="Diplazontinae">Diplazontinae Viereck, 1918</taxonomicName>
</emphasis>
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<paragraph id="07622652FF9BFFF6FF0BFBA2FE96FE25" blockId="3.[136,275,482,508]" box="[136,275,482,508]" pageId="3" pageNumber="640">
(
<figureCitation id="9FE63AD7FF9BFFF6FF13FBA2FE8EFE25" box="[144,267,482,508]" captionStart="FIGURE 2" captionStartId="4.[136,235,1894,1918]" captionTargetBox="[151,1436,189,1870]" captionTargetId="figure-20@4.[151,1436,189,1870]" captionTargetPageId="4" captionText="FIGURE 2. A, Puparium of aphidophagous syrphid in Baltic amber (Diptera: Syrphidae: Syrphinae) (KAM No. 6361) with an emergence hole typical for Diplazontinae, ventral view. B, Right lateral view; Emergence holes of various recent parasitoids of aphidophagous syrphids (Diptera, Syrphidae, Syrphinae). C, Diplazon pectoratorius. D, Syrphoctonus sp. E and F, Diplazon laetatorius. G, Cryptinae (gen. et sp. indet). H, Asaphes sp. (Pteromalidae). Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5778516" httpUri="https://zenodo.org/record/5778516/files/figure.png" pageId="3" pageNumber="640">Fig. 2AH</figureCitation>
)
</paragraph>
<paragraph id="07622652FF9BFFF6FF0BF86BFDB7FCBD" blockId="3.[136,778,554,1769]" pageId="3" pageNumber="640">
Twenty-two recent genera of
<taxonomicName id="C0DD5DD1FF9BFFF6FE56F86AFDF7FD9D" authorityName="Viereck" authorityYear="1918" box="[469,626,554,580]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="subFamily" subFamily="Diplazontinae">Diplazontinae</taxonomicName>
are currently recognized in the world fauna, with 355 species described to date (
<bibRefCitation id="634C5BA3FF9BFFF6FF6CF832FE51FD55" author="Broad, G. R. &amp; Shaw, M. R. &amp; Fitton, M. G." box="[239,468,626,653]" pageId="3" pageNumber="640" pagination="411" refId="ref5960" refString="Broad, G. R., Shaw, M. R. &amp; Fitton, M. G. (2018) Ichneumonid wasps (Hymenoptera: Ichneumonidae): their classification and biology. Handbooks for the identification of British Insects, 7 (12), 411 рр." type="journal article" year="2018">
Broad
<emphasis id="35A9FA40FF9BFFF6FEC0F833FEFEFD55" box="[323,379,626,652]" italics="true" pageId="3" pageNumber="640">et al</emphasis>
., 2018
</bibRefCitation>
). Most species have been described from the Holarctic region, with boreal and alpine areas of especially high species richness (
<bibRefCitation id="634C5BA3FF9BFFF6FD0AF8FAFF46FD20" author="Manukyan, A. R." pageId="3" pageNumber="640" pagination="55 - 60" refId="ref7366" refString="Manukyan, A. R. (1995) The geographic distribution of the Diplazontinae (Hymenoptera, Ichneumonidae) in the Palaearctic region, with description of two new species. Acta Zoologica Fennica, 199, 55 - 60." type="journal article" year="1995">Manukyan, 1995</bibRefCitation>
;
<bibRefCitation id="634C5BA3FF9BFFF6FF4CF89EFE15FD21" author="Klopfstein, S." box="[207,400,734,760]" pageId="3" pageNumber="640" pagination="1 - 143" refId="ref6916" refString="Klopfstein, S. (2014) Revision of the Western Palaearctic Diplazontinae (Hymenoptera, Ichneumonidae), Zootaxa, 3801 (1), 1 - 143. https: // doi. org / 10.11646 / zootaxa. 3801.1.1" type="journal article" year="2014">Klopfstein, 2014</bibRefCitation>
). All diplazontines are koinobiont endoparasitoids of aphidophagous
<taxonomicName id="C0DD5DD1FF9BFFF6FD9DF942FC82FCC4" authority="(Diptera)" baseAuthorityName="Diptera" box="[542,775,770,797]" class="Insecta" family="Syrphidae" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="family">Syrphidae (Diptera)</taxonomicName>
. Oviposition occurs into the egg or larva of the host, and emergence occurs from the puparium.
</paragraph>
<paragraph id="07622652FF9BFFF6FF3EF92EFDB4FBB8" blockId="3.[136,778,554,1769]" pageId="3" pageNumber="640">
The only fossil species,
<taxonomicName id="C0DD5DD1FF9BFFF6FD9EF92EFEA8FC75" authority="Scudder, 1890" authorityName="Scudder" authorityYear="1890" class="Insecta" family="Ichneumonidae" genus="Lithotorus" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="species" species="cressoni">
<emphasis id="35A9FA40FF9BFFF6FD9EF92EFC8CFC51" box="[541,777,878,904]" italics="true" pageId="3" pageNumber="640">Lithotorus cressoni</emphasis>
Scudder, 1890
</taxonomicName>
, was described from the Green River, and preliminarily placed in
<taxonomicName id="C0DD5DD1FF9BFFF6FE12F9F6FDABFC09" authorityName="Viereck" authorityYear="1918" box="[401,558,950,976]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="subFamily" subFamily="Diplazontinae">Diplazontinae</taxonomicName>
by
<bibRefCitation id="634C5BA3FF9BFFF6FDDAF9F7FC83FC08" author="Townes, H. K." box="[601,774,950,977]" pageId="3" pageNumber="640" pagination="132 - 135" refId="ref8213" refString="Townes, H. K. (1966) Notes on 3 fossil genera of Ichneumonidae (Hymenoptera). Proceedings of the Entomological Society of Washington, 68, 132 - 135." type="journal article" year="1966">Townes (1966)</bibRefCitation>
. However, subsequent research has shown an uncertain taxonomic position of this species (
<bibRefCitation id="634C5BA3FF9BFFF6FDBAF9BEFF54FBE5" author="Spasojevic, T. &amp; Broad, G. R. &amp; Bennett, A. M. R. &amp; Klopfstein, S." pageId="3" pageNumber="640" pagination="35 - 63" refId="ref7859" refString="Spasojevic, T., Broad, G. R., Bennett, A. M. R. &amp; Klopfstein, S. (2018 a) Ichneumonid parasitoid wasps from the Early Eocene Green River Formation: five new species and a revision of the known fauna (Hymenoptera, Ichneumonidae). Palaontologische Zeitschrift, 92, 35 - 63. https: // doi. org / 10.1007 / s 12542 - 017 - 0365 - 5" type="journal article" year="2018">
Spasojevic
<emphasis id="35A9FA40FF9BFFF6FD40F9BFFD79FBC1" box="[707,764,1022,1048]" italics="true" pageId="3" pageNumber="640">et al</emphasis>
., 2018a
</bibRefCitation>
). Therefore, there are no confirmed records of fossil
<taxonomicName id="C0DD5DD1FF9BFFF6FF4FFE06FEECFBB9" authorityName="Viereck" authorityYear="1918" box="[204,361,1094,1120]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="subFamily" subFamily="Diplazontinae">Diplazontinae</taxonomicName>
up to the present.
</paragraph>
<paragraph id="07622652FF9BFFF6FF3EFE2BFE70F979" blockId="3.[136,778,554,1769]" pageId="3" pageNumber="640">
In the collection of KAM, we found a puparium with an emergence hole typical of
<taxonomicName id="C0DD5DD1FF9BFFF6FE70FECEFD15FB71" authorityName="Viereck" authorityYear="1918" box="[499,656,1166,1192]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="subFamily" subFamily="Diplazontinae">Diplazontinae</taxonomicName>
(
<figureCitation id="9FE63AD7FF9BFFF6FD25FECEFF18FB14" captionStart="FIGURE 2" captionStartId="4.[136,235,1894,1918]" captionTargetBox="[151,1436,189,1870]" captionTargetId="figure-20@4.[151,1436,189,1870]" captionTargetPageId="4" captionText="FIGURE 2. A, Puparium of aphidophagous syrphid in Baltic amber (Diptera: Syrphidae: Syrphinae) (KAM No. 6361) with an emergence hole typical for Diplazontinae, ventral view. B, Right lateral view; Emergence holes of various recent parasitoids of aphidophagous syrphids (Diptera, Syrphidae, Syrphinae). C, Diplazon pectoratorius. D, Syrphoctonus sp. E and F, Diplazon laetatorius. G, Cryptinae (gen. et sp. indet). H, Asaphes sp. (Pteromalidae). Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5778516" httpUri="https://zenodo.org/record/5778516/files/figure.png" pageId="3" pageNumber="640">Fig. 2A, B</figureCitation>
).
<taxonomicName id="C0DD5DD1FF9BFFF6FF37FEF2FED4FB15" authorityName="Viereck" authorityYear="1918" box="[180,337,1202,1228]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="subFamily" subFamily="Diplazontinae">Diplazontinae</taxonomicName>
emerge from the puparium of the host from the head end. The mechanism of cutting through the puparium, the shape of the holes and the semicircular flaps left are similar in all diplazontines (
<bibRefCitation id="634C5BA3FF9BFFF6FDD7FF5EFC84FAE1" author="Rotheray, G. E." box="[596,769,1310,1336]" pageId="3" pageNumber="640" pagination="39 - 40" refId="ref7764" refString="Rotheray, G. E. (1981) Emergence from the host puparium by Diplazon pectoratorius (Gravenhorst) (Hymenoptera: Ichneumonidae), a parasitoid of aphidophagous syrphid larvae. Entomologist's Gazette, 32, 39 - 40." type="journal article" year="1981">Rotheray, 1981</bibRefCitation>
) (see
<figureCitation id="9FE63AD7FF9BFFF6FF39FF02FEABFA85" box="[186,302,1346,1372]" captionStart="FIGURE 2" captionStartId="4.[136,235,1894,1918]" captionTargetBox="[151,1436,189,1870]" captionTargetId="figure-20@4.[151,1436,189,1870]" captionTargetPageId="4" captionText="FIGURE 2. A, Puparium of aphidophagous syrphid in Baltic amber (Diptera: Syrphidae: Syrphinae) (KAM No. 6361) with an emergence hole typical for Diplazontinae, ventral view. B, Right lateral view; Emergence holes of various recent parasitoids of aphidophagous syrphids (Diptera, Syrphidae, Syrphinae). C, Diplazon pectoratorius. D, Syrphoctonus sp. E and F, Diplazon laetatorius. G, Cryptinae (gen. et sp. indet). H, Asaphes sp. (Pteromalidae). Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5778516" httpUri="https://zenodo.org/record/5778516/files/figure.png" pageId="3" pageNumber="640">Fig. 2CF</figureCitation>
), while in other parasitoids of syrphid flies, the cutouts are not semi-circular. Species of the genus
<taxonomicName id="C0DD5DD1FF9BFFF6FF0BFFCAFEFAFA7D" authority="Dahlbom" authorityName="Dahlbom" authorityYear="1842" box="[136,383,1418,1444]" class="Insecta" family="Figitidae" genus="Callaspidia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="genus">
<emphasis id="35A9FA40FF9BFFF6FF0BFFCAFE8EFA7D" box="[136,267,1418,1444]" italics="true" pageId="3" pageNumber="640">Callaspidia</emphasis>
Dahlbom
</taxonomicName>
(
<taxonomicName id="C0DD5DD1FF9BFFF6FE11FFCAFDAAFA7D" authorityName="Linnaeus" authorityYear="1758" box="[402,559,1418,1444]" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Tracheophyta" rank="order">Hymenoptera</taxonomicName>
,
<taxonomicName id="C0DD5DD1FF9BFFF6FDBCFFCAFD21FA7C" baseAuthorityName="Forster" baseAuthorityYear="1869" box="[575,676,1418,1445]" class="Insecta" family="Figitidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="family">Figitidae</taxonomicName>
) cut out irregular holes on the lateral surface of the puparium and
<taxonomicName id="C0DD5DD1FF9BFFF6FF0BFF92FF73FA35" authorityName="Kirby" authorityYear="1837" box="[136,246,1490,1516]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="639" phylum="Arthropoda" rank="subFamily" subFamily="Cryptinae">Cryptinae</taxonomicName>
cut irregularly rounded holes in the center of the dorsal surface (
<figureCitation id="9FE63AD7FF9BFFF6FED5FFB6FE2AF9C9" box="[342,431,1526,1552]" captionStart="FIGURE 2" captionStartId="4.[136,235,1894,1918]" captionTargetBox="[151,1436,189,1870]" captionTargetId="figure-20@4.[151,1436,189,1870]" captionTargetPageId="4" captionText="FIGURE 2. A, Puparium of aphidophagous syrphid in Baltic amber (Diptera: Syrphidae: Syrphinae) (KAM No. 6361) with an emergence hole typical for Diplazontinae, ventral view. B, Right lateral view; Emergence holes of various recent parasitoids of aphidophagous syrphids (Diptera, Syrphidae, Syrphinae). C, Diplazon pectoratorius. D, Syrphoctonus sp. E and F, Diplazon laetatorius. G, Cryptinae (gen. et sp. indet). H, Asaphes sp. (Pteromalidae). Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5778516" httpUri="https://zenodo.org/record/5778516/files/figure.png" pageId="3" pageNumber="640">Fig. 2G</figureCitation>
). The gregarious
<taxonomicName id="C0DD5DD1FF9BFFF6FDE8FFB6FF78F9ED" authority="Ratzeburg" authorityName="Ratzeburg" authorityYear="1844" class="Insecta" family="Encyrtidae" genus="Bothriothorax" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="genus">
<emphasis id="35A9FA40FF9BFFF6FDE8FFB6FC8FF9C9" box="[619,778,1526,1552]" italics="true" pageId="3" pageNumber="640">Bothriothorax</emphasis>
Ratzeburg
</taxonomicName>
,
<taxonomicName id="C0DD5DD1FF9BFFF6FE90FC5AFD49F9EC" authority="Ashmead" authorityName="Ashmead" authorityYear="1900" box="[275,716,1562,1589]" class="Insecta" family="Encyrtidae" genus="Syrphophagus" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="genus">
<emphasis id="35A9FA40FF9BFFF6FE90FC5AFE36F9ED" box="[275,435,1562,1588]" italics="true" pageId="3" pageNumber="640">Syrphophagus</emphasis>
Ashmead (Encyrtidae)
</taxonomicName>
, and
<taxonomicName id="C0DD5DD1FF9BFFF6FF0BFC7EFE75F980" authority="Walker" authorityName="Walker" authorityYear="1834" box="[136,496,1598,1625]" class="Insecta" family="Pteromalidae" genus="Asaphes" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="genus">
<emphasis id="35A9FA40FF9BFFF6FF0BFC7EFF60F981" box="[136,229,1598,1624]" italics="true" pageId="3" pageNumber="640">Asaphes</emphasis>
Walker (Pteromalidae)
</taxonomicName>
cut a small hole on the lateral surface of the puparium (
<figureCitation id="9FE63AD7FF9BFFF6FE7FFC22FDDCF9A5" box="[508,601,1634,1660]" captionStart="FIGURE 2" captionStartId="4.[136,235,1894,1918]" captionTargetBox="[151,1436,189,1870]" captionTargetId="figure-20@4.[151,1436,189,1870]" captionTargetPageId="4" captionText="FIGURE 2. A, Puparium of aphidophagous syrphid in Baltic amber (Diptera: Syrphidae: Syrphinae) (KAM No. 6361) with an emergence hole typical for Diplazontinae, ventral view. B, Right lateral view; Emergence holes of various recent parasitoids of aphidophagous syrphids (Diptera, Syrphidae, Syrphinae). C, Diplazon pectoratorius. D, Syrphoctonus sp. E and F, Diplazon laetatorius. G, Cryptinae (gen. et sp. indet). H, Asaphes sp. (Pteromalidae). Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5778516" httpUri="https://zenodo.org/record/5778516/files/figure.png" pageId="3" pageNumber="640">Fig. 2H</figureCitation>
), which is then used by all emerging specimens.
</paragraph>
<paragraph id="07622652FF9BFFF6FF3EFCEAFD96F930" blockId="3.[136,778,554,1769]" pageId="3" pageNumber="640">
Our finding is the first record of subfamily
<taxonomicName id="C0DD5DD1FF9BFFF6FF0BFC8EFEA0F931" authorityName="Viereck" authorityYear="1918" box="[136,293,1742,1768]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="3" pageNumber="640" phylum="Arthropoda" rank="subFamily" subFamily="Diplazontinae">Diplazontinae</taxonomicName>
in the upper Eocene.
</paragraph>
</subSubSection>
</treatment>
</document>