<documentid="8628CB089C0D4E76B822D80FE8A0F6D1"ID-CLB-Dataset="305179"ID-DOI="10.15298/rusentj.33.1.04"ID-GBIF-Dataset="fed91399-9291-481a-8f90-f70843b8a449"ID-ISSN="0132-8069"ID-Zenodo-Dep="14162701"IM.bibliography_approvedBy="julia"IM.materialsCitations_approvedBy="julia"IM.metadata_approvedBy="julia"IM.taxonomicNames_approvedBy="julia"IM.treatments_approvedBy="julia"checkinTime="1712198995902"checkinUser="felipe"docAuthor="Makarov, K. V."docDate="2024"docId="03B76F18B578A715FF2310FE8ADDE4BD"docLanguage="en"docName="RussEntomolJ.33.1.39-42.pdf"docOrigin="Russian Entomological Journal 33 (1)"docSource="http://dx.doi.org/10.15298/rusentj.33.1.04"docStyle="DocumentStyle:0D5BAC5AA0C130B3809AF578A601D837.4:RussEntomolJ.2019-.journal_article"docStyleId="0D5BAC5AA0C130B3809AF578A601D837"docStyleName="RussEntomolJ.2019-.journal_article"docStyleVersion="4"docTitle="Circularva Shcherbakov et Ponomarenko 2023"docType="treatment"docVersion="2"lastPageNumber="41"masterDocId="FF8E1760B578A717FFC01626885DE067"masterDocTitle="On the systematic position of Circularva reichardti Shcherbakov et Ponomarenko, 2023"masterLastPageNumber="42"masterPageNumber="39"pageNumber="39"updateTime="1731589645851"updateUser="ExternalLinkService">
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<bibRefCitationid="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>
] is grouped in three obvious categories:
</paragraph>
<paragraphid="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>
(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
<paragraphid="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>
Since we do not know the set and structure of the appendages and legs of
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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 (
<figureCitationid="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>
Crustaceans are highly diverse in terms of the number of segments and their combinations into tagmata [
<bibRefCitationid="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>
]. If this hypothesis is accepted, a complete coincidence of the segmentation of
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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 (
<figureCitationid="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>
). 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 [
<bibRefCitationid="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>
]. The wide sternal region of
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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 [
<bibRefCitationid="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>
], this failing to contradict the age of
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Oniscomorph forms are common among millipedes, or Diplopoda [
<bibRefCitationid="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>
]. Modern Sphaerotheriida have 11 free trunk segments (segment XII is merged with the telson) and a telson [
<bibRefCitationid="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>
]. This resembles the tagmosis of
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<figureCitationid="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>
), 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
<bibRefCitationid="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>
], and their fossil representatives (Amynilyspedida) are known from the Upper Carboniferous [
<bibRefCitationid="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>
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 [
<bibRefCitationid="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">
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<figureCitationid="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>
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. 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.
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satisfies the interpretation in favour of the “diplopod” hypothesis and corresponds very well to the “isopod” hypothesis. In addition,
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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,
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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
<bibRefCitationid="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>
;
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]. 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 [
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] and in the Ephemeroptera, only one muscle pair is present [
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], whereas in all Holometabola larvae whose anatomy has been studied, segment X comprises only longitudinal muscles [
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;
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;
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;
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reichardti
</emphasis>
<bibRefCitationid="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
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reichardti
</emphasis>
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, рисунок иЗ первоописаниЯ, леваЯ половина, неЯсные структуры не покаЗаны; 2–4 — предполагаемый тагмоЗис
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is meaningless. Most likely, that was a crustacean, but its attribution to Isopoda still remains unclear. Therefore, it seems rational to consider
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<bibRefCitationid="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>