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266 lines
40 KiB
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<taxonomicName id="4C1EA58DB578A717FF2310FE891AE693" authority="Shcherbakov et Ponomarenko, 2023" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[227,327,1752,1780]" genus="Circularva" kingdom="Animalia" pageId="0" pageNumber="39" rank="genus">
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<emphasis id="B96A021CB578A717FF2310FE891AE693" bold="true" box="[227,327,1752,1780]" pageId="0" pageNumber="39">Results</emphasis>
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</heading>
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</paragraph>
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<subSubSection id="C3048D85B578A715FF23113189D1E38A" lastPageId="2" lastPageNumber="41" pageId="0" pageNumber="39" type="description">
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<paragraph id="8BA1DE0EB578A717FF231131899FE70D" blockId="0.[189,788,1815,1987]" pageId="0" pageNumber="39">
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All information concerning the morphology of
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<taxonomicName id="4C1EA58DB578A717FD10113188A9E72B" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" genus="Circularva" pageId="0" pageNumber="39" rank="genus">
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<emphasis id="B96A021CB578A717FD10113188A9E72B" italics="true" pageId="0" pageNumber="39">Circularva</emphasis>
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</taxonomicName>
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[
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<bibRefCitation id="EF8FA3FFB578A717FEC311138A28E72B" author="Shcherbakov D. E. & Ponomarenko A. G." box="[259,629,1845,1868]" pageId="0" pageNumber="39" pagination="261 - 270" refId="ref2855" refString="Shcherbakov D. E., Ponomarenko A. G. 2023. The first known fossil hygropetric beetle larva related to Myxophaga (Coleoptera) from the Permian of European Russia // Russian Entomol. J. Vol. 32. No. 3. P. 261 - 270. https: // doi. org / 10.15298 / rusentj. 32.3.02" type="journal article" year="2023">Shcherbakov, Ponomarenko, 2023</bibRefCitation>
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] is grouped in three obvious categories:
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</paragraph>
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<paragraph id="8BA1DE0EB578A717FF2311578D4BE465" blockId="0.[189,788,1815,1987]" lastBlockId="0.[825,1424,943,1987]" pageId="0" pageNumber="39">(a) Structures that are clearly observed and can be accurately interpreted. These are the number and shape of the dorsal sclerites, the number and location of ocelli on each side of the head, the sculpture of the integument (rows of tubercles at the posterior margin of tergites), and the pleurites of the abdominal segments;</paragraph>
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<paragraph id="8BA1DE0EB578A717FC9F122F8C7DE577" blockId="0.[825,1424,943,1987]" pageId="0" pageNumber="39">
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(b) Poorly distinguishable, unclear structures that cannot be accurately interpreted based on the visible organization. These are the several processes/appendages on the segments, the setae on tubercles in the posterior part of the tergites, and the abdominal sclerites. Their interpretation is based on the principle of taxonomic analogy, i.e. such that can be considered as spiracular gills, setiferous tubercles, sternites only if the fossil does belong to
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<taxonomicName id="4C1EA58DB578A717FC6412DF8C40E577" box="[932,1053,1273,1296]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="0" pageNumber="39" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
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;
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</paragraph>
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<paragraph id="8BA1DE0EB578A717FC9F13318C98E5C1" blockId="0.[825,1424,943,1987]" pageId="0" pageNumber="39">(c) Expected structures that are not visible on the impressions. First of all, such are the appendages of the head and body. Nothing is known yet as to how many and which antennae this creature may have had, nor how many legs could have been involved.</paragraph>
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<paragraph id="8BA1DE0EB578A717FC9F138B8D50E63C" blockId="0.[825,1424,943,1987]" pageId="0" pageNumber="39">
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Since we do not know the set and structure of the appendages and legs of
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<taxonomicName id="4C1EA58DB578A717FBFF13EA8CEEE584" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[1087,1203,1484,1507]" genus="Circularva" pageId="0" pageNumber="39" rank="genus">
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<emphasis id="B96A021CB578A717FBFF13EA8CEEE584" box="[1087,1203,1484,1507]" italics="true" pageId="0" pageNumber="39">Circularva</emphasis>
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</taxonomicName>
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, we can confidently attribute it only to the Arthropoda, among which it could potentially have been related to both crustaceans and myriapods, as well as to hexapods (
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<figureCitation id="1325C28BB578A717FB0710008D76E65A" box="[1223,1323,1574,1597]" captionStart="Figs 1–4" captionStartId="1.[189,227,1933,1952]" captionTargetBox="[212,1408,1119,1864]" captionTargetId="figure-423@1.[199,1409,1119,1883]" captionTargetPageId="1" captionText="Figs 1–4. Circularva reichardti and interpretations of its morphology: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, drawing from the original description, left half, unclear structures not shown; 2–4 — putative tagmosis of Circularva due to hypotheses about its belonging to crustaceans, millipedes and coleopterans, respectively. Рис. 1–4. Circularva reichardti и интерпретации её строениЯ: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, рисунок иЗ первоописаниЯ, леваЯ половина, неЯсные структуры не покаЗаны; 2–4 — предполагаемый тагмоЗис Circularva в рамках гипотеЗ о её принадлежности к ракообраЗным, многоножкам и насекомым, соответственно." figureDoi="http://doi.org/10.5281/zenodo.14162703" httpUri="https://zenodo.org/record/14162703/files/figure.png" pageId="0" pageNumber="39">Figs 1–4</figureCitation>
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). Let me consider these three hypotheses one by one.
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</paragraph>
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<paragraph id="8BA1DE0EB578A716FC9F10448A44E19F" blockId="0.[825,1424,943,1987]" lastBlockId="1.[189,786,302,1039]" lastPageId="1" lastPageNumber="40" pageId="0" pageNumber="39">
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Crustaceans are highly diverse in terms of the number of segments and their combinations into tagmata [
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<bibRefCitation id="EF8FA3FFB578A717FCBB10B88CF4E6D2" author="Schram F. R. & Koenemann S." box="[891,1193,1694,1717]" pageId="0" pageNumber="39" refId="ref2807" refString="Schram F. R., Koenemann S. 2021. Evolution and phylogeny of Pancrustacea: a story of scientific method. Oxford University Press. xii + 827 p. + 32 pl. https: // doi. org / 10.1093 / oso / 9780195365764.001.0001" type="book" year="2021">Schram, Koenemann, 2021</bibRefCitation>
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]. If this hypothesis is accepted, a complete coincidence of the segmentation of
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<taxonomicName id="4C1EA58DB578A717FCF910FC8BF0E696" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[825,941,1754,1777]" genus="Circularva" pageId="0" pageNumber="39" rank="genus">
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<emphasis id="B96A021CB578A717FCF910FC8BF0E696" box="[825,941,1754,1777]" italics="true" pageId="0" pageNumber="39">Circularva</emphasis>
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is to be found with one of the tagmosis variants of Isopoda, in which the thoracic region consists of seven segments, the abdominal region includes five free segments, and the sixth abdominal segment is merged with the telson (
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<figureCitation id="1325C28BB578A717FC2F11748C69E70E" box="[1007,1076,1874,1897]" captionStart="Figs 1–4" captionStartId="1.[189,227,1933,1952]" captionTargetBox="[212,1408,1119,1864]" captionTargetId="figure-423@1.[199,1409,1119,1883]" captionTargetPageId="1" captionText="Figs 1–4. Circularva reichardti and interpretations of its morphology: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, drawing from the original description, left half, unclear structures not shown; 2–4 — putative tagmosis of Circularva due to hypotheses about its belonging to crustaceans, millipedes and coleopterans, respectively. Рис. 1–4. Circularva reichardti и интерпретации её строениЯ: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, рисунок иЗ первоописаниЯ, леваЯ половина, неЯсные структуры не покаЗаны; 2–4 — предполагаемый тагмоЗис Circularva в рамках гипотеЗ о её принадлежности к ракообраЗным, многоножкам и насекомым, соответственно." figureDoi="http://doi.org/10.5281/zenodo.14162703" httpUri="https://zenodo.org/record/14162703/files/figure.png" pageId="0" pageNumber="39">Fig. 2</figureCitation>
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). Among the isopods, there are animals with an almost homonomous segmentation of the thoracic region, yet heteronomous segmentation with varying segment sizes is likewise very common [
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<bibRefCitation id="EF8FA3FFB579A716FF03170889FAE122" author="Brandt A. & Poore G. C. B." box="[195,423,302,325]" pageId="1" pageNumber="40" pagination="893 - 923" refId="ref2292" refString="Brandt A., Poore G. C. B. 2003. Higher classification of the flabelliferan and related Isopoda based on a reappraisal of relationships // Invertebrate Systematics. Vol. 17. No. 6. P. 893 - 923. https: // doi. org / 10.1071 / IS 02032." type="book chapter" year="2003">Brandt, Poore, 2003</bibRefCitation>
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]. The wide sternal region of
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<taxonomicName id="4C1EA58DB579A716FD2617088953E104" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
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<emphasis id="B96A021CB579A716FD2617088953E104" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
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</taxonomicName>
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is similar to that of Isopoda. The appendages of Isopoda are diverse and often specialized, this giving room to interpreting the unclearly distinguishable structures. Fossils of Isopoda have been known since the Carboniferous [
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<bibRefCitation id="EF8FA3FFB579A716FEFE17E58A31E1BD" author="Schram F. R. & Koenemann S." box="[318,620,451,474]" pageId="1" pageNumber="40" refId="ref2807" refString="Schram F. R., Koenemann S. 2021. Evolution and phylogeny of Pancrustacea: a story of scientific method. Oxford University Press. xii + 827 p. + 32 pl. https: // doi. org / 10.1093 / oso / 9780195365764.001.0001" type="book" year="2021">Schram, Koenemann, 2021</bibRefCitation>
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], this failing to contradict the age of
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<taxonomicName id="4C1EA58DB579A716FE5E17C78A4FE19F" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[414,530,481,504]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
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<emphasis id="B96A021CB579A716FE5E17C78A4FE19F" box="[414,530,481,504]" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
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</taxonomicName>
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.
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</paragraph>
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<paragraph id="8BA1DE0EB579A716FF2317D88D3BE1E6" blockId="1.[189,786,302,1039]" lastBlockId="1.[824,1421,302,1039]" pageId="1" pageNumber="40">
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Oniscomorph forms are common among millipedes, or Diplopoda [
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<bibRefCitation id="EF8FA3FFB579A716FEA5143A89ADE254" author="Koch M." box="[357,496,540,563]" pageId="1" pageNumber="40" pagination="7 - 67" refId="ref2472" refString="Koch M. 2015. Chapter 2. Diplopoda - General morphology // Minelli A. (ed.) Treatise on Zoology - Anatomy, Taxonomy, Biology. The Myriapoda. Vol. 2. P. 7 - 67." type="journal article" year="2015">Koch, 2015</bibRefCitation>
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]. Modern Sphaerotheriida have 11 free trunk segments (segment XII is merged with the telson) and a telson [
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<bibRefCitation id="EF8FA3FFB579A716FDC2147E8AF7E208" author="Wesener T." box="[514,682,600,623]" pageId="1" pageNumber="40" pagination="370 - 381" refId="ref2956" refString="Wesener T. 2015. Infraclass Pentazonia Brandt, 1833 // Minelli A. (ed.) Treatise on Zoology - Anatomy, Taxonomy, Biology. The Myriapoda. Vol. 2. P. 370 - 381." type="journal article" year="2015">Wesener, 2015</bibRefCitation>
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]. This resembles the tagmosis of
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<taxonomicName id="4C1EA58DB579A716FE0714538A66E2EB" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[455,571,629,652]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
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<emphasis id="B96A021CB579A716FE0714538A66E2EB" box="[455,571,629,652]" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
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</taxonomicName>
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(see
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<figureCitation id="1325C28BB579A716FDB514538AE9E2EB" box="[629,692,629,652]" captionStart="Figs 1–4" captionStartId="1.[189,227,1933,1952]" captionTargetBox="[212,1408,1119,1864]" captionTargetId="figure-423@1.[199,1409,1119,1883]" captionTargetPageId="1" captionText="Figs 1–4. Circularva reichardti and interpretations of its morphology: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, drawing from the original description, left half, unclear structures not shown; 2–4 — putative tagmosis of Circularva due to hypotheses about its belonging to crustaceans, millipedes and coleopterans, respectively. Рис. 1–4. Circularva reichardti и интерпретации её строениЯ: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, рисунок иЗ первоописаниЯ, леваЯ половина, неЯсные структуры не покаЗаны; 2–4 — предполагаемый тагмоЗис Circularva в рамках гипотеЗ о её принадлежности к ракообраЗным, многоножкам и насекомым, соответственно." figureDoi="http://doi.org/10.5281/zenodo.14162703" httpUri="https://zenodo.org/record/14162703/files/figure.png" pageId="1" pageNumber="40">Fig. 3</figureCitation>
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), but requires at least two assumptions: (a) a weakly bound area at the base of the head must be interpreted as a collum, and (b) segment XII has not yet become fused with the telson in that primitive putative millipede. Millipedes of the superorder
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<taxonomicName id="4C1EA58DB579A716FE48152C8A78E346" authorityName="Pocock" authorityYear="1887" box="[392,549,778,801]" pageId="1" pageNumber="40" rank="superOrder" superOrder="Oniscomorpha">Oniscomorpha</taxonomicName>
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have well-developed pleurites [
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<bibRefCitation id="EF8FA3FFB579A716FEE5150E89E0E358" author="Blower J. G." box="[293,445,808,831]" pageId="1" pageNumber="40" pagination="1 - 242" refId="ref2186" refString="Blower J. G. 1985. Millipedes: Keys and notes for the identification of the species // Synopses of the British fauna. New series No. 35. Linnean Society of London, Estuarine and Brackish-water Sciences Association E. J. Brill / Dr. W. Backhuys. P. 1 - 242." type="book chapter" year="1985">Blower, 1985</bibRefCitation>
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], and their fossil representatives (Amynilyspedida) are known from the Upper Carboniferous [
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<bibRefCitation id="EF8FA3FFB579A716FED7154589BEE31D" author="Edgecombe G. D." box="[279,483,867,890]" pageId="1" pageNumber="40" pagination="337 - 351" refId="ref2389" refString="Edgecombe G. D. 2015. Chapter 14. Diplopoda - Fossils // Minelli A. (ed.) Treatise on Zoology - Anatomy, Taxonomy, Biology. The Myriapoda. Vol. 2. P. 337 - 351" type="journal article" year="2015">Edgecombe, 2015</bibRefCitation>
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]. The number of trunk segments in the fossil
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<taxonomicName id="4C1EA58DB579A716FE4615A78A7EE3FF" authorityName="Pocock" authorityYear="1887" box="[390,547,897,920]" pageId="1" pageNumber="40" rank="superOrder" superOrder="Oniscomorpha">Oniscomorpha</taxonomicName>
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ranged from 13 to 15, supporting the diplopod hypothesis. However, in Diplopoda the sternites are usually narrow, which is not consistent with the impression. Besides this, Amynilyspedida showed characteristic spines on the tergites [
|
||
<bibRefCitation id="EF8FA3FFB579A716FD7215DE8BEBE122" author="Lheritier M. & Perroux M. & Vannier J. & Escarguel G. & Wesener T. & Moritz L. & Chabard D. & Adriene J. & Perriera V." box="[690,950,302,1039]" pageId="1" pageNumber="40" pagination="1477 - 2019" refId="ref2639" refString="Lheritier M., Perroux M., Vannier J., Escarguel G., Wesener T., Moritz L., Chabard D., Adriene J., Perriera V. 2023. Fossils from the Montceau-les-Mines Lagerstatte (305 Ma) shed light on the anatomy, ecology and phylogeny of Carboniferous millipedes // Journal of Systematic Palaeontology. Vol. 21. No. 1. P. 1477 - 2019. https: // doi. org / 10.1080 / 14772019.2023.2169891" type="journal article" year="2023">
|
||
Lheritier
|
||
<emphasis id="B96A021CB579A716FCF817088B3AE122" box="[824,871,302,325]" italics="true" pageId="1" pageNumber="40">et al</emphasis>
|
||
., 2023
|
||
</bibRefCitation>
|
||
], these being absent from
|
||
<taxonomicName id="4C1EA58DB579A716FB0F17088D1EE122" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[1231,1347,302,325]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB579A716FB0F17088D1EE122" box="[1231,1347,302,325]" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
|
||
</taxonomicName>
|
||
. Thus, the “diplopod” hypothesis is acceptable, but has obvious disadvantages compared to the “isopod” hypothesis.
|
||
</paragraph>
|
||
<paragraph id="8BA1DE0EB579A716FC9E17A18BBEE31D" blockId="1.[824,1421,302,1039]" pageId="1" pageNumber="40">
|
||
The absence of wings, simple ocelli on the head, slightly enlarged first segments (interpreted as thoracic segments) and a ten-segmented abdomen (
|
||
<figureCitation id="1325C28BB579A716FAD417E58D07E1BD" box="[1300,1370,451,474]" captionStart="Figs 1–4" captionStartId="1.[189,227,1933,1952]" captionTargetBox="[212,1408,1119,1864]" captionTargetId="figure-423@1.[199,1409,1119,1883]" captionTargetPageId="1" captionText="Figs 1–4. Circularva reichardti and interpretations of its morphology: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, drawing from the original description, left half, unclear structures not shown; 2–4 — putative tagmosis of Circularva due to hypotheses about its belonging to crustaceans, millipedes and coleopterans, respectively. Рис. 1–4. Circularva reichardti и интерпретации её строениЯ: 1 — Circularva reichardti Shcherbakov et Ponomarenko, 2023, рисунок иЗ первоописаниЯ, леваЯ половина, неЯсные структуры не покаЗаны; 2–4 — предполагаемый тагмоЗис Circularva в рамках гипотеЗ о её принадлежности к ракообраЗным, многоножкам и насекомым, соответственно." figureDoi="http://doi.org/10.5281/zenodo.14162703" httpUri="https://zenodo.org/record/14162703/files/figure.png" pageId="1" pageNumber="40">Fig. 4</figureCitation>
|
||
), all this suggests that
|
||
<taxonomicName id="4C1EA58DB579A716FC3F17C78C2EE19F" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[1023,1139,481,504]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB579A716FC3F17C78C2EE19F" box="[1023,1139,481,504]" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
|
||
</taxonomicName>
|
||
is a Holometabola larva, probably
|
||
<taxonomicName id="4C1EA58DB579A716FC6217D88C46E272" box="[930,1051,510,533]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="40" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
|
||
. It seems important that the size differences between the thoracic and abdominal segments are very small: starting with the metathorax, their length is approximately the same and half the length of the prothorax (measured from the photograph in the original description). The rather narrow abdominal segments V–VII (0.3–0.4 the length of the prothorax) look strongly deformed in the photographs of the impression, while the following segments VIII–IX already show the normal length, which is half the prothorax length. Consequently, only thoracic segments I and II differ in size from the other segments and this trait is also common among Isopoda.
|
||
</paragraph>
|
||
<paragraph id="8BA1DE0EB579A715FC9E15A789D1E38A" blockId="1.[824,1421,302,1039]" lastBlockId="2.[189,787,303,1242]" lastPageId="2" lastPageNumber="41" pageId="1" pageNumber="40">
|
||
As shown above, the number of segments in
|
||
<taxonomicName id="4C1EA58DB579A716FA8815A78B2DE3D1" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB579A716FA8815A78B2DE3D1" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
|
||
</taxonomicName>
|
||
satisfies the interpretation in favour of the “diplopod” hypothesis and corresponds very well to the “isopod” hypothesis. In addition,
|
||
<taxonomicName id="4C1EA58DB579A716FBAF15FC8CBEE396" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[1135,1251,986,1009]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB579A716FBAF15FC8CBEE396" box="[1135,1251,986,1009]" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
|
||
</taxonomicName>
|
||
has several features that fail to fit well with the morphology of
|
||
<taxonomicName id="4C1EA58DB579A715FAF715DE88B8E121" class="Insecta" kingdom="Animalia" lastPageId="2" lastPageNumber="41" order="Coleoptera" pageId="1" pageNumber="40" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
|
||
larvae and, more generally, of Holometabola larvae in general. Firstly, they lack molting sutures, which are usually clearly visible on impressions, especially on the head, since the sutures often diverge when the head capsule is deformed. Secondly,
|
||
<taxonomicName id="4C1EA58DB57AA715FE2617838A07E1DB" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[486,602,421,444]" genus="Circularva" pageId="2" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB57AA715FE2617838A07E1DB" box="[486,602,421,444]" italics="true" pageId="2" pageNumber="41">Circularva</emphasis>
|
||
</taxonomicName>
|
||
had an unusually wide abdominal segment X. In Holometabola larvae, this segment usually assumes a locomotor function and has the shape of a tube, more or less strongly bent ventrally. Among the larvae of
|
||
<taxonomicName id="4C1EA58DB57AA715FE67143A8A42E254" box="[423,543,540,563]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="41" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
|
||
, no forms with such a wide segment X are known [
|
||
<bibRefCitation id="EF8FA3FFB57AA715FE38141F8B50E237" author="Boving A. G. & Craighead F. C." box="[504,781,569,592]" pageId="2" pageNumber="41" pagination="1 - 351" refId="ref2245" refString="Boving A. G., Craighead F. C. 1931. An illustrated synopsis of the principal larval forms of the order Coleoptera // Entomol. Am. (N. S.). Vol. 11. P. 1 - 351." type="journal article" year="1931">Böving, Craighead, 1931</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EF8FA3FFB57AA715FF7D147189DFE209" author="Costa C. & Vanin S. A. & Casari-Chan S. A." box="[189,386,599,622]" pageId="2" pageNumber="41" refId="ref2349" refString="Costa C., Vanin S. A., Casari-Chan S. A. 1988. Larvas de Coleoptera do Brasil // Museu de Zoologia, Universidade de Sao Paulo, Brazil. 282 pp. 165 pl." type="book" year="1988">
|
||
Costa
|
||
<emphasis id="B96A021CB57AA715FEC314718969E209" box="[259,308,599,622]" italics="true" pageId="2" pageNumber="41">et al</emphasis>
|
||
., 1988
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EF8FA3FFB57AA715FE4F14718ADDE209" author="Lawrence J. & Hastings A. & Dallwitz M. & Paine T. & Zurcher E." box="[399,640,599,622]" pageId="2" pageNumber="41" refId="ref2515" refString="Lawrence J., Hastings A., Dallwitz M., Paine T., Zurcher E. 1991. Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Sub-Families. CSIRO. CD-ROM + Manual." type="book" year="1991">
|
||
Lawrence
|
||
<emphasis id="B96A021CB57AA715FDC114718A6FE209" box="[513,562,599,622]" italics="true" pageId="2" pageNumber="41">et al</emphasis>
|
||
., 1991
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EF8FA3FFB57AA715FD4D14718A9AE209" author="Lawrence J. F. & Slipinski A. & Seago A. E. & Thayer M. K. & Newton A. F. & MarvaldiA. E." box="[653,711,599,622]" pageId="2" pageNumber="41" pagination="1 - 217" refId="ref2562" refString="Lawrence J. F., Slipinski A., Seago A. E., Thayer M. K., Newton A. F., MarvaldiA. E. 2011. Phylogeny of the Coleoptera based on morphological characters of adults and larvae // Annales Zoologici. Vol. 61. No. 1. P. 1 - 217. https: // doi. org / 10.3161 / 000345411 X 576725." type="journal article" year="2011">2011</bibRefCitation>
|
||
]. Wide tergites in insect larvae are associated with the presence of dorsoventral and dorsopleural muscles. In Ectognatha, these muscle groups in segment X are greatly reduced. In the Zygentoma, two muscle pairs remain [
|
||
<bibRefCitation id="EF8FA3FFB57AA715FD7414EB88A8E366" author="Rousset A." pageId="2" pageNumber="41" pagination="55 - 80" refId="ref2727" refString="Rousset A. 1973. Squelette et musculature des regions genitales et postgenitales de la femelle de Thermobia domestica (Packard). Comparaison avec la region genitale de Nicoletia sp. (Insecta: Apterygota: Lepismatida) // International Journal of Insect Morphology and Embryology. Vol. 2. No. 1. P. 55 - 80. https: // doi. org / 10.1016 / 0020 - 7322 (73) 90006 - 8" type="journal article" year="1973">Rousset, 1973</bibRefCitation>
|
||
] and in the Ephemeroptera, only one muscle pair is present [
|
||
<bibRefCitation id="EF8FA3FFB57AA715FEF0152E8A5BE378" author="Birket-Smith J." box="[304,518,776,799]" pageId="2" pageNumber="41" pagination="139 - 160" refId="ref2152" refString="Birket-Smith J. 1971. The abdominal morphology of Povilla adusta Navas (Polymitarcidae) and of Ephemeroptera in general // Entomologica Scandinavica. No. 2. P. 139 - 160." type="book chapter" year="1971">Birket-Smith, 1971</bibRefCitation>
|
||
], whereas in all Holometabola larvae whose anatomy has been studied, segment X comprises only longitudinal muscles [
|
||
<bibRefCitation id="EF8FA3FFB57AA715FDAF15658B50E33D" author="Kemner A." box="[623,781,835,858]" pageId="2" pageNumber="41" pagination="1 - 23" refId="ref2432" refString="Kemner A. 1913. Beitrage zur Kenntnis einiger schwedischen Coleopterenlarven. II. Das Analsegment und die Rektalschlauche einiger Carabidenlarven // Arkiv for Zoologi. Bd. 8. Nr. 13. S. 1 - 23." type="journal article" year="1913">Kemner, 1913</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EF8FA3FFB57AA715FF7D1546892DE310" author="Snodgrass RE" box="[189,368,864,887]" pageId="2" pageNumber="41" pagination="1 - 128" refId="ref2919" refString="Snodgrass RE. 1931. Morphology of the insect abdomen. Part I. General structure of the abdomen and its appendages // Smithsonian miscellaneous Collections. No. 85. P. 1 - 128." type="book chapter" year="1931">Snodgrass, 1931</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EF8FA3FFB57AA715FEBC15468AE7E310" author="Berrios-Ortiz A. & Selander R. B." box="[380,698,864,887]" pageId="2" pageNumber="41" pagination="1 - 114" refId="ref2057" refString="Berrios-Ortiz A., Selander R. B. 1979. Skeletal musculature in larval phases of the beetle Epicauta segmenta (Coleoptera, Meloidae) // Series Entomologica. Vol. 16. P. 1 - 114." type="book chapter" year="1979">Berrios-Ortiz, Selander, 1979</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EF8FA3FFB57AA715FD06154689F1E3F2" author="Beutel R. G. & Hornschemeyer T." pageId="2" pageNumber="41" pagination="169 - 190" refId="ref2097" refString="Beutel R. G., Hornschemeyer T. 2002. Larval morphology and phylogenetic position of Micromalthus debilis LeConte (Coleoptera: Micromaltidae) // Systematic Entomology. Vol. 27. P. 169 - 190. https: // doi. org / 10.1046 / j. 1365 - 3113.2002.00172. x" type="book chapter" year="2002">Beutel, Hornschemeyer, 2002</bibRefCitation>
|
||
; Wipfler
|
||
<emphasis id="B96A021CB57AA715FDCE15588A63E3F2" box="[526,574,894,917]" italics="true" pageId="2" pageNumber="41">et al</emphasis>
|
||
., 2012;
|
||
<bibRefCitation id="EF8FA3FFB57AA715FD5515588960E3D5" author="Yavorskaya M. I. & Hornschemeyer T. & Beutel R. G." pageId="2" pageNumber="41" pagination="475 - 486" refId="ref2997" refString="Yavorskaya M. I., Hornschemeyer T., Beutel R. G. 2015. The head morphology of Micromalthus debilis (Coleoptera: Micromalthidae) - an archostematan beetle with an unusual morphology and a unique life cycle // Arthropod Systematics & Phylogeny. Vol. 76. No. 3. P. 475 - 486. https: // doi. org / 10.3897 / asp. 76. e 31964" type="journal article" year="2015">
|
||
Yavorskaya
|
||
<emphasis id="B96A021CB57AA715FF7D15BD88B0E3D5" box="[189,237,923,946]" italics="true" pageId="2" pageNumber="41">et al</emphasis>
|
||
., 2015
|
||
</bibRefCitation>
|
||
]. Therefore, the attribution of
|
||
<taxonomicName id="4C1EA58DB57AA715FD4215BD8AABE3D5" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[642,758,923,946]" genus="Circularva" pageId="2" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB57AA715FD4215BD8AABE3D5" box="[642,758,923,946]" italics="true" pageId="2" pageNumber="41">Circularva</emphasis>
|
||
</taxonomicName>
|
||
to
|
||
<taxonomicName id="4C1EA58DB57AA715FF7D159F8968E3B7" box="[189,309,953,976]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="41" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
|
||
, based on the structure of the last segment, is extremely unlikely.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="DF618E86B579A716FF7D11AB8BD7E87F" ID-DOI="http://doi.org/10.5281/zenodo.14162703" ID-Zenodo-Dep="14162703" httpUri="https://zenodo.org/record/14162703/files/figure.png" pageId="1" pageNumber="40" startId="1.[189,227,1933,1952]" targetBox="[212,1408,1119,1864]" targetPageId="1" targetType="figure">
|
||
<paragraph id="8BA1DE0EB579A716FF7D11AB8AF8E7B7" blockId="1.[189,1421,1933,2072]" pageId="1" pageNumber="40">
|
||
<emphasis id="B96A021CB579A716FF7D11AB8953E7C7" bold="true" box="[189,270,1933,1952]" pageId="1" pageNumber="40">Figs 1–4.</emphasis>
|
||
<emphasis id="B96A021CB579A716FED411AB8996E7C7" box="[276,459,1933,1952]" italics="true" pageId="1" pageNumber="40">
|
||
<taxonomicName id="4C1EA58DB579A716FED411AB892CE7C7" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[276,369,1933,1952]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">Circularva</taxonomicName>
|
||
reichardti
|
||
</emphasis>
|
||
and interpretations of its morphology: 1 —
|
||
<emphasis id="B96A021CB579A716FC8811AB8BA2E7C7" box="[840,1023,1933,1952]" italics="true" pageId="1" pageNumber="40">
|
||
<taxonomicName id="4C1EA58DB579A716FC8811AB8BF8E7C7" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[840,933,1933,1952]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">Circularva</taxonomicName>
|
||
reichardti
|
||
</emphasis>
|
||
<bibRefCitation id="EF8FA3FFB579A716FBC411AB8D63E7C7" author="Shcherbakov D. E. & Ponomarenko A. G." box="[1028,1342,1933,1952]" pageId="1" pageNumber="40" pagination="261 - 270" refId="ref2855" refString="Shcherbakov D. E., Ponomarenko A. G. 2023. The first known fossil hygropetric beetle larva related to Myxophaga (Coleoptera) from the Permian of European Russia // Russian Entomol. J. Vol. 32. No. 3. P. 261 - 270. https: // doi. org / 10.15298 / rusentj. 32.3.02" type="journal article" year="2023">Shcherbakov et Ponomarenko, 2023</bibRefCitation>
|
||
, drawing from the original description, left half, unclear structures not shown; 2–4 — putative tagmosis of
|
||
<taxonomicName id="4C1EA58DB579A716FC2E11838C16E7DF" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[1006,1099,1957,1976]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB579A716FC2E11838C16E7DF" box="[1006,1099,1957,1976]" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
|
||
</taxonomicName>
|
||
due to hypotheses about its belonging to crustaceans, millipedes and coleopterans, respectively.
|
||
</paragraph>
|
||
<paragraph id="8BA1DE0EB579A716FF7D11F38BD7E87F" blockId="1.[189,1421,1933,2072]" pageId="1" pageNumber="40">
|
||
<emphasis id="B96A021CB579A716FF7D11F3894CE78F" bold="true" box="[189,273,2005,2024]" pageId="1" pageNumber="40">Рис. 1–4.</emphasis>
|
||
<emphasis id="B96A021CB579A716FED811F3898DE78F" box="[280,464,2005,2024]" italics="true" pageId="1" pageNumber="40">
|
||
<taxonomicName id="4C1EA58DB579A716FED811F38928E78F" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[280,373,2005,2024]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">Circularva</taxonomicName>
|
||
reichardti
|
||
</emphasis>
|
||
и интерпретации её строениЯ: 1 —
|
||
<emphasis id="B96A021CB579A716FCE211F38B87E78F" box="[802,986,2005,2024]" italics="true" pageId="1" pageNumber="40">
|
||
<taxonomicName id="4C1EA58DB579A716FCE211F38B22E78F" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[802,895,2005,2024]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">Circularva</taxonomicName>
|
||
reichardti
|
||
</emphasis>
|
||
<bibRefCitation id="EF8FA3FFB579A716FC2011F38D40E78F" author="Shcherbakov D. E. & Ponomarenko A. G." box="[992,1309,2005,2024]" pageId="1" pageNumber="40" pagination="261 - 270" refId="ref2855" refString="Shcherbakov D. E., Ponomarenko A. G. 2023. The first known fossil hygropetric beetle larva related to Myxophaga (Coleoptera) from the Permian of European Russia // Russian Entomol. J. Vol. 32. No. 3. P. 261 - 270. https: // doi. org / 10.15298 / rusentj. 32.3.02" type="journal article" year="2023">Shcherbakov et Ponomarenko, 2023</bibRefCitation>
|
||
, рисунок иЗ первоописаниЯ, леваЯ половина, неЯсные структуры не покаЗаны; 2–4 — предполагаемый тагмоЗис
|
||
<taxonomicName id="4C1EA58DB579A716FB9F11CB8CE1E867" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[1119,1212,2029,2048]" genus="Circularva" pageId="1" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB579A716FB9F11CB8CE1E867" box="[1119,1212,2029,2048]" italics="true" pageId="1" pageNumber="40">Circularva</emphasis>
|
||
</taxonomicName>
|
||
в рамках гипотеЗ о её принадлежности к ракообраЗным, многоножкам и насекомым, соответственно.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="C3048D85B57AA715FF2315D28ADDE4BD" pageId="2" pageNumber="41" type="discussion">
|
||
<paragraph id="8BA1DE0EB57AA715FF2315D28ADDE4BD" blockId="2.[189,787,303,1242]" pageId="2" pageNumber="41">
|
||
To summarize, I believe that
|
||
<taxonomicName id="4C1EA58DB57AA715FDEC15D28AFDE46C" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" box="[556,672,1012,1035]" genus="Circularva" pageId="2" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB57AA715FDEC15D28AFDE46C" box="[556,672,1012,1035]" italics="true" pageId="2" pageNumber="41">Circularva</emphasis>
|
||
</taxonomicName>
|
||
could not have been a coleopteran larva. Therefore, discussing its placement in the suborder
|
||
<taxonomicName id="4C1EA58DB57AA715FE1F12098A03E421" box="[479,606,1071,1094]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="41" phylum="Arthropoda" rank="subOrder" subOrder="Myxophaga">Myxophaga</taxonomicName>
|
||
and its similarities to the family
|
||
<taxonomicName id="4C1EA58DB57AA715FE40126B8A79E403" box="[384,548,1101,1124]" class="Insecta" family="Torridincolidae" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="41" phylum="Arthropoda" rank="family">Torridincolidae</taxonomicName>
|
||
is meaningless. Most likely, that was a crustacean, but its attribution to Isopoda still remains unclear. Therefore, it seems rational to consider
|
||
<taxonomicName id="4C1EA58DB57AA715FE83128388AFE4BD" authority="Shcherbakov et Ponomarenko, 2023" authorityName="Shcherbakov et Ponomarenko" authorityYear="2023" genus="Circularva" pageId="2" pageNumber="39" rank="genus">
|
||
<emphasis id="B96A021CB57AA715FE83128389EAE4DB" box="[323,439,1189,1212]" italics="true" pageId="2" pageNumber="41">Circularva</emphasis>
|
||
<bibRefCitation id="EF8FA3FFB57AA715FE02128388AFE4BD" author="Shcherbakov D. E. & Ponomarenko A. G." pageId="2" pageNumber="41" pagination="261 - 270" refId="ref2855" refString="Shcherbakov D. E., Ponomarenko A. G. 2023. The first known fossil hygropetric beetle larva related to Myxophaga (Coleoptera) from the Permian of European Russia // Russian Entomol. J. Vol. 32. No. 3. P. 261 - 270. https: // doi. org / 10.15298 / rusentj. 32.3.02" type="journal article" year="2023">Shcherbakov et Ponomarenko, 2023</bibRefCitation>
|
||
</taxonomicName>
|
||
as an Eumalacostraca
|
||
<emphasis id="B96A021CB57AA715FE2512E58A25E4BD" box="[485,632,1219,1242]" italics="true" pageId="2" pageNumber="41">incertae sedis</emphasis>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |