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<mods:title id="EA88689A184526E60A864DA82A2C83F5">The first Ichneumonid fossil from the Early Pleistocene of Madeira Island (Portugal)</mods:title>
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<mods:namePart id="E98958F89F47067BBCB27714969FB69E">Góis-Marques, Carlos A.</mods:namePart>
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<mods:namePart id="B85122401E5A515BB98EAE9FB865119A">Sequeira, Miguel Menezes De</mods:namePart>
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<mods:namePart id="FE06828E66C567485778C12D55E40B13">Madeira, José</mods:namePart>
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<subSubSection id="2CAD6503FF855322FF4DFBF5FE7A6EF0" pageId="1" pageNumber="448" type="nomenclature">
<paragraph id="64083688FF855322FF4DFBF5FDC76E1C" blockId="1.[151,1437,1025,2023]" box="[151,628,1098,1124]" pageId="1" pageNumber="448">
<heading id="3F4081E4FF855322FF4DFBF5FDC76E1C" bold="true" box="[151,628,1098,1124]" fontSize="11" level="1" pageId="1" pageNumber="448" reason="1">
<emphasis id="56C3EA9AFF855322FF4DFBF5FDC76E1C" bold="true" box="[151,628,1098,1124]" pageId="1" pageNumber="448">
Family
<taxonomicName id="A3B74D0BFF855322FF2DFBF5FDC76E1C" ID-CoL="BCL" ID-ENA="7408" authority="Latreille, 1802" authorityName="Latreille" authorityYear="1802" box="[247,628,1098,1124]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="family">
Ichneumonidae
<bibRefCitation id="00264B79FF855322FE1AFBF5FDC76E1C" author="Latreille, P. A." box="[448,628,1098,1124]" pageId="1" pageNumber="448" refId="ref2407" refString="Latreille, P. A. (1802) Histoire naturelle, generale et particuliere, des Crustaces et des Insectes, Tome troisieme. F. Dufart, Paris, 467 pp." type="book" year="1802">Latreille, 1802</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="64083688FF855322FF4DFBD1FE7A6EF0" blockId="1.[151,1437,1025,2023]" box="[151,457,1134,1160]" pageId="1" pageNumber="448">
<heading id="3F4081E4FF855322FF4DFBD1FE7A6EF0" bold="true" box="[151,457,1134,1160]" fontSize="11" level="1" pageId="1" pageNumber="448" reason="1">
<emphasis id="56C3EA9AFF855322FF4DFBD1FE7A6EF0" bold="true" box="[151,457,1134,1160]" pageId="1" pageNumber="448">Genus and species indet.</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="2CAD6503FF855322FF4DFB2CFBB16ED3" box="[151,1026,1171,1195]" pageId="1" pageNumber="448" type="materials_examined">
<paragraph id="64083688FF855322FF4DFB2CFBB16ED3" blockId="1.[151,1437,1025,2023]" box="[151,1026,1171,1195]" pageId="1" pageNumber="448">
<emphasis id="56C3EA9AFF855322FF4DFB2CFF496ED3" bold="true" box="[151,250,1171,1195]" pageId="1" pageNumber="448">Material</emphasis>
: UMad-P500a (part:
<figureCitation id="FC8C2A0DFF855322FE05FB2CFD836ED3" box="[479,560,1171,1195]" captionStart="FIGURE 2" captionStartId="2.[151,250,816,841]" captionTargetBox="[151,1436,174,756]" captionTargetId="figure@2.[151,1436,174,756]" captionTargetPageId="2" captionText="FIGURE 2. Ichneumonidae hind wing morphology and comparison with the wing fossil. A, position and venation in Ichneumonidae (Ophion obscuratus Fabricius, 1798), figure adapted from Prehn &amp; Raper (2016); B, Specimen UMad-P500a (part); C, Specimen UMad-P500b (counter-part); D, drawing interpretation of specimen UMad-P500a with venation nomenclature and cell numbers; arrows point to bulla." httpUri="https://zenodo.org/record/3235110/files/figure.png" pageId="1" pageNumber="448">Fig. 2B</figureCitation>
) and UMad-P500b (Counter-part:
<figureCitation id="FC8C2A0DFF855322FC78FB2CFC456ED3" box="[930,1014,1171,1195]" captionStart="FIGURE 2" captionStartId="2.[151,250,816,841]" captionTargetBox="[151,1436,174,756]" captionTargetId="figure@2.[151,1436,174,756]" captionTargetPageId="2" captionText="FIGURE 2. Ichneumonidae hind wing morphology and comparison with the wing fossil. A, position and venation in Ichneumonidae (Ophion obscuratus Fabricius, 1798), figure adapted from Prehn &amp; Raper (2016); B, Specimen UMad-P500a (part); C, Specimen UMad-P500b (counter-part); D, drawing interpretation of specimen UMad-P500a with venation nomenclature and cell numbers; arrows point to bulla." httpUri="https://zenodo.org/record/3235110/files/figure.png" pageId="1" pageNumber="448">Fig. 2C</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="2CAD6503FF855322FF1DFB08FD566F6F" pageId="1" pageNumber="448" type="description">
<paragraph id="64083688FF855322FF1DFB08FD566F6F" blockId="1.[151,1437,1025,2023]" pageId="1" pageNumber="448">
<emphasis id="56C3EA9AFF855322FF1DFB08FEF86EB7" bold="true" box="[199,331,1207,1231]" pageId="1" pageNumber="448">Description</emphasis>
: Incomplete hind wing, composed of incarbonized venation,
<quantity id="A34F9B6DFF855322FC39FB08FBF06EB7" box="[995,1091,1207,1231]" metricMagnitude="-2" metricUnit="m" metricValue="1.06" pageId="1" pageNumber="448" unit="mm" value="10.6">10.6 mm</quantity>
(base of wing to the RSb apex) ×
<quantity id="A34F9B6DFF855322FF77FB64FEB86E8B" box="[173,267,1243,1267]" metricMagnitude="-3" metricUnit="m" metricValue="3.16" pageId="1" pageNumber="448" unit="mm" value="3.16">3.16 mm</quantity>
(2A apex to SC+R), venation more or less complete, except R1, and the missing vein C; bulla present in rs-m and CU-a. Vein 1-A partly detached from the sediment.
</paragraph>
</subSubSection>
<subSubSection id="2CAD6503FF855322FF1DFA9BFD366D9F" pageId="1" pageNumber="448" type="discussion">
<paragraph id="64083688FF855322FF1DFA9BFD0D6C6B" blockId="1.[151,1437,1025,2023]" pageId="1" pageNumber="448">
<emphasis id="56C3EA9AFF855322FF1DFA9BFE9C6F44" bold="true" box="[199,303,1316,1340]" pageId="1" pageNumber="448">Remarks</emphasis>
: Hind wing fossil architecture and size correspond to the
<taxonomicName id="A3B74D0BFF855322FC4AFA9CFB876F43" authorityName="Latreille" authorityYear="1802" box="[912,1076,1315,1339]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="family">Ichneumonidae</taxonomicName>
(
<figureCitation id="FC8C2A0DFF855322FB98FA9CFB246F43" box="[1090,1175,1315,1339]" captionStart="FIGURE 2" captionStartId="2.[151,250,816,841]" captionTargetBox="[151,1436,174,756]" captionTargetId="figure@2.[151,1436,174,756]" captionTargetPageId="2" captionText="FIGURE 2. Ichneumonidae hind wing morphology and comparison with the wing fossil. A, position and venation in Ichneumonidae (Ophion obscuratus Fabricius, 1798), figure adapted from Prehn &amp; Raper (2016); B, Specimen UMad-P500a (part); C, Specimen UMad-P500b (counter-part); D, drawing interpretation of specimen UMad-P500a with venation nomenclature and cell numbers; arrows point to bulla." httpUri="https://zenodo.org/record/3235110/files/figure.png" pageId="1" pageNumber="448">Fig. 2A</figureCitation>
). It is distinct from similar
<taxonomicName id="A3B74D0BFF855322FF60FAF8FE806F27" box="[186,307,1351,1375]" class="Insecta" family="Braconidae" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="family">Braconidae</taxonomicName>
hind wings, due to the enclosed VI cell (
<figureCitation id="FC8C2A0DFF855322FD33FAF8FC316F27" box="[745,898,1351,1375]" captionStart="FIGURE 2" captionStartId="2.[151,250,816,841]" captionTargetBox="[151,1436,174,756]" captionTargetId="figure@2.[151,1436,174,756]" captionTargetPageId="2" captionText="FIGURE 2. Ichneumonidae hind wing morphology and comparison with the wing fossil. A, position and venation in Ichneumonidae (Ophion obscuratus Fabricius, 1798), figure adapted from Prehn &amp; Raper (2016); B, Specimen UMad-P500a (part); C, Specimen UMad-P500b (counter-part); D, drawing interpretation of specimen UMad-P500a with venation nomenclature and cell numbers; arrows point to bulla." httpUri="https://zenodo.org/record/3235110/files/figure.png" pageId="1" pageNumber="448">Fig. 2A and D</figureCitation>
) (
<bibRefCitation id="00264B79FF855322FC43FAF8FB926F27" author="Broad, G." box="[921,1057,1351,1375]" pageId="1" pageNumber="448" refId="ref1782" refString="Broad, G. (2011) Identification key to the subfamilies of Ichneumonidae (Hymenoptera). The Natural History Museum, London, 40 pp." type="book" year="2011">Broad, 2011</bibRefCitation>
), and vein rs-m apical to separation of veins RSa and RSb (
<bibRefCitation id="00264B79FF855322FE55FAD4FD376FFB" author="Goulet, H. &amp; Huber, J. T." box="[399,644,1387,1411]" pageId="1" pageNumber="448" refId="ref2250" refString="Goulet, H. &amp; Huber, J. T. (1993) Hymenoptera of the World: An identification guide to families. Centre for Land &amp; Biological Resources, Ottawa, 668 pp." type="book" year="1993">Goulet &amp; Huber, 1993</bibRefCitation>
). Further identification is prevented, although the size and venation points to a large ichneumonid parasitoid wasp. Genera with similar hind wing size and venation occur today in Madeira, including
<emphasis id="56C3EA9AFF855322FF1BFA0CFE276FB3" box="[193,404,1459,1483]" italics="true" pageId="1" pageNumber="448">
<taxonomicName id="A3B74D0BFF855322FF1BFA0CFE8A6FB3" box="[193,313,1459,1483]" class="Insecta" family="Ichneumonidae" genus="Amblyteles" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="genus">Amblyteles</taxonomicName>
,
<taxonomicName id="A3B74D0BFF855322FE9EFA0CFE276FB3" authorityName="Fabricius" authorityYear="1798" box="[324,404,1459,1483]" class="Insecta" family="Ichneumonidae" genus="Ophion" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="genus">Ophion</taxonomicName>
</emphasis>
,
<emphasis id="56C3EA9AFF855322FE78FA0CFDAC6FB3" box="[418,543,1459,1483]" italics="true" pageId="1" pageNumber="448">
<taxonomicName id="A3B74D0BFF855322FE78FA0CFDA86FB3" box="[418,539,1459,1483]" class="Insecta" family="Eulophidae" genus="Ichneumon" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="genus">Ichneumon</taxonomicName>
,
</emphasis>
and
<taxonomicName id="A3B74D0BFF855322FD8EFA0CFD136FB3" authorityName="Gravenhorst" authorityYear="1829" box="[596,672,1459,1483]" class="Insecta" family="Ichneumonidae" genus="Rhyssa" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="genus">
<emphasis id="56C3EA9AFF855322FD8EFA0CFD136FB3" box="[596,672,1459,1483]" italics="true" pageId="1" pageNumber="448">Rhyssa</emphasis>
</taxonomicName>
(
<bibRefCitation id="00264B79FF855322FD74FA0CFCF36FB3" author="Aguiar, A. M. F." box="[686,832,1459,1483]" pageId="1" pageNumber="448" pagination="352 - 354" refId="ref1495" refString="Aguiar, A. M. F. (2008) List of arthropods (Arthropoda): Hymenoptera (Bethylidae, Dryinidae, Ichneumonidae). In: Borges, P. A. V., Abreu, C., Aguiar, A. M. F., Carvalho, P. Jardim, R., Melo, I., Oliveira, P., Sergio, C., Serrano, A. R. M. &amp; Vieira, P. (Coord.), Listagem dos fungos, flora e fauna terrestre dos arquipelagos da Madeira e Selvagens. Direccao Regional do Ambiente da Madeira e Universidade dos Acores, Funchal e Angra do Heroismo, pp. 352 - 354." type="book chapter" year="2008">Aguiar, 2008</bibRefCitation>
;
<bibRefCitation id="00264B79FF855322FC93FA0CFBAB6FB3" author="Aguiar, A. F. &amp; Cravo, D. &amp; Goncalves, Y." box="[841,1048,1459,1483]" pageId="1" pageNumber="448" pagination="157 - 170" refId="ref1430" refString="Aguiar, A. F., Cravo, D. &amp; Goncalves, Y. (2017) Sawflies and horntails of Madeira Archipelago (Hymenoptera: Symphyta), with the first records of two wasp species (Hymenoptera: Ibaliidae &amp; Ichneumonidae) parasitizing Sirex noctilio (Fabricius, 1793). Entomologist's Monthly Magazine, 153, 157 - 170." type="journal article" year="2017">
Aguiar
<emphasis id="56C3EA9AFF855322FC40FA0BFC626FB3" box="[922,977,1459,1483]" italics="true" pageId="1" pageNumber="448">et al.</emphasis>
, 2017
</bibRefCitation>
). However, to identify the specimen to genus and/or species, more complete fossil specimens are needed, as
<taxonomicName id="A3B74D0BFF855322FC40FA68FB8D6F97" authorityName="Latreille" authorityYear="1802" box="[922,1086,1495,1519]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="family">Ichneumonidae</taxonomicName>
taxonomy is based on fore wing and body morphology (e.g.
<bibRefCitation id="00264B79FF855322FE65FA44FD016C6B" author="Goulet, H. &amp; Huber, J. T." box="[447,690,1531,1555]" pageId="1" pageNumber="448" refId="ref2250" refString="Goulet, H. &amp; Huber, J. T. (1993) Hymenoptera of the World: An identification guide to families. Centre for Land &amp; Biological Resources, Ottawa, 668 pp." type="book" year="1993">Goulet &amp; Huber, 1993</bibRefCitation>
).
</paragraph>
<paragraph id="64083688FF855322FF1DF9A0FDC96D4B" blockId="1.[151,1437,1025,2023]" pageId="1" pageNumber="448">
Today, Madeiran
<taxonomicName id="A3B74D0BFF855322FE5EF9A0FD9B6C4F" authorityName="Latreille" authorityYear="1802" box="[388,552,1567,1591]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="family">Ichneumonidae</taxonomicName>
are represented by 98 spp., of which, 30 spp. are single island endemics (
<bibRefCitation id="00264B79FF855322FA91F9A0FF626C23" author="Aguiar, A. M. F." pageId="1" pageNumber="448" pagination="352 - 354" refId="ref1495" refString="Aguiar, A. M. F. (2008) List of arthropods (Arthropoda): Hymenoptera (Bethylidae, Dryinidae, Ichneumonidae). In: Borges, P. A. V., Abreu, C., Aguiar, A. M. F., Carvalho, P. Jardim, R., Melo, I., Oliveira, P., Sergio, C., Serrano, A. R. M. &amp; Vieira, P. (Coord.), Listagem dos fungos, flora e fauna terrestre dos arquipelagos da Madeira e Selvagens. Direccao Regional do Ambiente da Madeira e Universidade dos Acores, Funchal e Angra do Heroismo, pp. 352 - 354." type="book chapter" year="2008">Aguiar, 2008</bibRefCitation>
). Notwithstanding the high number of endemic species, literature reveals that Madeiran ichneumonids are understudied, being restricted mainly to check lists and taxonomy (e.g.
<bibRefCitation id="00264B79FF855322FC5EF9D8FBA66C07" author="Aguiar, A. M. F." box="[900,1045,1639,1663]" pageId="1" pageNumber="448" pagination="352 - 354" refId="ref1495" refString="Aguiar, A. M. F. (2008) List of arthropods (Arthropoda): Hymenoptera (Bethylidae, Dryinidae, Ichneumonidae). In: Borges, P. A. V., Abreu, C., Aguiar, A. M. F., Carvalho, P. Jardim, R., Melo, I., Oliveira, P., Sergio, C., Serrano, A. R. M. &amp; Vieira, P. (Coord.), Listagem dos fungos, flora e fauna terrestre dos arquipelagos da Madeira e Selvagens. Direccao Regional do Ambiente da Madeira e Universidade dos Acores, Funchal e Angra do Heroismo, pp. 352 - 354." type="book chapter" year="2008">Aguiar, 2008</bibRefCitation>
;
<bibRefCitation id="00264B79FF855322FBFAF9D8FB416C07" author="Aguiar, A. F. &amp; Cravo, D. &amp; Goncalves, Y." box="[1056,1266,1639,1663]" pageId="1" pageNumber="448" pagination="157 - 170" refId="ref1430" refString="Aguiar, A. F., Cravo, D. &amp; Goncalves, Y. (2017) Sawflies and horntails of Madeira Archipelago (Hymenoptera: Symphyta), with the first records of two wasp species (Hymenoptera: Ibaliidae &amp; Ichneumonidae) parasitizing Sirex noctilio (Fabricius, 1793). Entomologist's Monthly Magazine, 153, 157 - 170." type="journal article" year="2017">
Aguiar
<emphasis id="56C3EA9AFF855322FBA8F9D7FB1A6C07" box="[1138,1193,1639,1663]" italics="true" pageId="1" pageNumber="448">et al.</emphasis>
, 2017
</bibRefCitation>
). Recent efforts include those of
<bibRefCitation id="00264B79FF855322FE90F934FDAC6CDB" author="Santos, A. M. C. &amp; Fontaine, C. &amp; Quicke, D. L. J. &amp; Borges, P. A. V. &amp; Hortal, J." box="[330,543,1675,1699]" pageId="1" pageNumber="448" pagination="1256 - 1262" refId="ref2768" refString="Santos, A. M. C., Fontaine, C., Quicke, D. L. J., Borges, P. A. V. &amp; Hortal, J. (2011) Are island and mainland biotas different? Richness and level of generalism in parasitoids of a microlepidopteran in Macaronesia. Oikos, 120, 1256 - 1262. https: // doi. org / 10.1111 / j. 1600 - 0706.2010.19404. x" type="journal article" year="2011">
Santos
<emphasis id="56C3EA9AFF855322FE43F933FE636CDB" box="[409,464,1675,1699]" italics="true" pageId="1" pageNumber="448">et al.</emphasis>
(2011)
</bibRefCitation>
, who demonstrated through host dissection and DNA barcoding that on Macaronesian Islands, parasitoids including
<taxonomicName id="A3B74D0BFF855322FDCBF910FD0B6CBF" authorityName="Latreille" authorityYear="1802" box="[529,696,1711,1735]" class="Insecta" family="Ichneumonidae" kingdom="Animalia" order="Hymenoptera" pageId="1" pageNumber="448" phylum="Arthropoda" rank="family">Ichneumonidae</taxonomicName>
, had a higher number of generalists (idiobionts) when compared to mainland. Studies dealing with immigration, evolution and divergence of ichneumonid parasitoid wasps in Madeira are currently lacking. However, the existence of a parasitoid wasp in Madeira Island at 1.3 Ma, certainly points to a complex trophic system already installed in the island.
</paragraph>
<paragraph id="64083688FF855322FF1DF8FFFD366D9F" blockId="1.[151,1437,1025,2023]" pageId="1" pageNumber="448">We conclude that insect somatofossils on oceanic islands are rare, but when found, they can provide interesting palaeontological information. The specimen described is the first record for Madeira island and Macaronesia. It corresponds to a fossilized hind wing of an ichneumonoid parasitoid wasp from the Calabrian stage (1.3 Ma) in Madeira Island, implying also the existence of their hosts. Further entomological and palaeoentomological efforts are needed to understand the evolution of these wasps in an insular context.</paragraph>
</subSubSection>
</treatment>
</document>