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<mods:title id="CAB8A088BFBBF9227662734D047DFB49">Identification of ancient Cladocera-like fossils requires homologies: The Jurassic Kuqaia is not a Waterflea</mods:title>
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<paragraph id="32103689FF8BF02070DE13DEFE51FE34" blockId="1.[151,1437,447,2021]" box="[151,481,447,473]" pageId="1" pageNumber="92">
<heading id="695881E5FF8BF02070DE13DEFE51FE34" bold="true" box="[151,481,447,473]" fontSize="11" level="1" pageId="1" pageNumber="92" reason="1">
<emphasis id="00DBEA9BFF8BF02070DE13DEFE51FE34" bold="true" box="[151,481,447,473]" pageId="1" pageNumber="92">
<taxonomicName id="F5AF4D0AFF8BF02070DE13DEFF5FFE34" authority="O. F. Muller, 1776" authorityName="Li" authorityYear="1993" box="[151,239,447,473]" class="Branchiopoda" family="Macrothricidae" genus="Kuqaia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="91" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF8BF02070DE13DEFF5FFE34" bold="true" box="[151,239,447,473]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
is not a cladoceran
</emphasis>
</heading>
</paragraph>
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<subSubSection id="7AB56502FF8BF02570DE1384FE33FC06" lastPageId="4" lastPageNumber="95" pageId="1" pageNumber="92" type="description">
<paragraph id="32103689FF8BF02070DE1384FF69FCAC" blockId="1.[151,1437,447,2021]" pageId="1" pageNumber="92">
It is difficult to find and to assess the identity of fossils of small-sized extinct taxa in ancient strata (
<bibRefCitation id="563E4B78FF8BF020748A1384FA21FE10" author="Cooper, R. A. &amp; Maxwell, P. A. &amp; Crampton, J. S. &amp; Beu, A. G. &amp; Jones, C. M. &amp; Marshall, B. A." box="[1219,1425,485,509]" pageId="1" pageNumber="92" pagination="241 - 244" refId="ref5504" refString="Cooper, R. A., Maxwell, P. A., Crampton, J. S., Beu, A. G., Jones, C. M. &amp; Marshall, B. A. (2006) Completeness of the fossil record: estimating losses due to small body size. Geology, 34, 241 - 244. https: // doi. org / 10.1130 / G 22206.1" type="journal article" year="2006">
Cooper
<emphasis id="00DBEA9BFF8BF02075521387FAE1FE10" box="[1307,1361,485,509]" italics="true" pageId="1" pageNumber="92">et al.</emphasis>
2006
</bibRefCitation>
). In earlier studies mentioned by
<bibRefCitation id="563E4B78FF8BF02071A51068FD1FFDCC" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[492,687,521,545]" pageId="1" pageNumber="92" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8BF0207261106BFDE8FDCC" box="[552,600,521,545]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. (2023)
</bibRefCitation>
,
<taxonomicName id="F5AF4D0AFF8BF02072F2106BFCBAFDCC" authorityName="Li" authorityYear="1993" box="[699,778,522,545]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF02072F2106BFCBAFDCC" box="[699,778,522,545]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
has been attributed to fossil megaspores, gastropods, and the authors mention that these have also been suggested as potentially rotifers, or cocoons of annelids. The taphonomy of the deposits where
<taxonomicName id="F5AF4D0AFF8BF02071251033FE0BFD84" authorityName="Li" authorityYear="1993" box="[364,443,594,617]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF02071251033FE0BFD84" box="[364,443,594,617]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
is generally found (
<bibRefCitation id="563E4B78FF8BF02072D01030FCE3FD84" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[665,851,593,617]" pageId="1" pageNumber="92" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8BF02072901033FCA2FD84" box="[729,786,593,617]" italics="true" pageId="1" pageNumber="92">et al.</emphasis>
2023
</bibRefCitation>
), indicates a predominantly marine environment with potential freshwater stadia during regressions, which might have been inhabited by cladocerans as these animals were present at the time in truly freshwater habitats (yet if so, they were mostly accompanied by other freshwater invertebrates in the taphonomic setting;
<bibRefCitation id="563E4B78FF8BF020718910DCFDE1FD38" author="Kotov, A. A." box="[448,593,701,725]" pageId="1" pageNumber="92" pagination="523 - 528" refId="ref5910" refString="Kotov, A. A. (2009 a) New finding of Mesozoic ephippia of the Anomopoda (Crustacea: Cladocera). Journal of Natural History, 43 (9 / 10), 523 - 528. https: // doi. org / 10.1080 / 00222930802003020" type="journal article" year="2009">Kotov 2009a</bibRefCitation>
;
<bibRefCitation id="563E4B78FF8BF020721710DCFC24FD38" author="Van Damme, K. &amp; Kotov, A. A." box="[606,916,701,725]" pageId="1" pageNumber="92" pagination="162 - 189" refId="ref6670" refString="Van Damme, K. &amp; Kotov, A. A. (2016) The fossil record of the Cladocera (Crustacea: Branchiopoda): evidence and hypotheses. Earth-Science Reviews, 163 (2016), 162 - 189. https: // doi. org / 10.1016 / j. earscirev. 2016.10.009" type="journal article" year="2016">Van Damme &amp; Kotov 2016</bibRefCitation>
;
<bibRefCitation id="563E4B78FF8BF02073E810DCFB1EFD38" author="Van Damme, K. &amp; Cornetti, L. &amp; Fields, P. D. &amp; Ebert, D." box="[929,1198,701,725]" pageId="1" pageNumber="92" pagination="777 - 787" refId="ref6727" refString="Van Damme, K., Cornetti, L., Fields, P. D. &amp; Ebert, D. (2022) Whole-Genome Phylogenetic Reconstruction as a Powerful Tool to Reveal Homoplasy and Ancient Rapid Radiation in Waterflea Evolution. Systematic Biology, 71 (4), 777 - 787. https: // doi. org / 10.1093 / sysbio / syab 094" type="journal article" year="2022">
Van Damme
<emphasis id="00DBEA9BFF8BF020747B10DFFBD4FD38" box="[1074,1124,701,725]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. 2022
</bibRefCitation>
). However, it is clear that the
<taxonomicName id="F5AF4D0AFF8BF02070B91083FE8FFD14" authorityName="Li" authorityYear="1993" box="[240,319,738,761]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF02070B91083FE8FFD14" box="[240,319,738,761]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
fossils morphologically are not cladoceran ephippia. Based on an examination of the SEM images and interpretations (drawings and text) in
<bibRefCitation id="563E4B78FF8BF02072611164FD56FCF0" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[552,742,773,797]" pageId="1" pageNumber="92" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8BF020722A1167FD22FCF0" box="[611,658,773,797]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. (2023)
</bibRefCitation>
compared to ephippia (
<figureCitation id="AA942A0CFF8BF02073A81164FBAEFCF0" box="[993,1054,773,797]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1</figureCitation>
), the following observations can be made:
</paragraph>
<paragraph id="32103689FF8BF023708E112CFC11FEF6" blockId="1.[151,1437,447,2021]" lastBlockId="2.[151,1437,151,284]" lastPageId="2" lastPageNumber="93" pageId="1" pageNumber="92">
<emphasis id="00DBEA9BFF8BF020708E112CFD92FC88" box="[199,546,845,869]" italics="true" pageId="1" pageNumber="92">
<emphasis id="00DBEA9BFF8BF020708E112CFF50FC88" bold="true" box="[199,224,845,869]" italics="true" pageId="1" pageNumber="92">(i)</emphasis>
Appendages (“peduncles”).
</emphasis>
The long appendages (“peduncles”) in
<emphasis id="00DBEA9BFF8BF0207384112FFBF9FC88" box="[973,1097,846,869]" italics="true" pageId="1" pageNumber="92">K. scanicus</emphasis>
are implanted at some distance from a potential “keel” and almost midway a straight margin visible in the fossils (
<figureCitation id="AA942A0CFF8BF02074421110FBEAFC64" box="[1035,1114,881,905]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1E</figureCitation>
and
<bibRefCitation id="563E4B78FF8BF02074C61110FAF0FC64" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[1167,1344,881,905]" pageId="1" pageNumber="92" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8BF02074821113FB4AFC64" box="[1227,1274,881,905]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. 2023
</bibRefCitation>
: Fig. 2a; the appendages are on the left in their study and the “keel” is oriented downwards in their figure, also visible in their Fig. 2b). The authors describe that the implantation of these peduncles is at “
<emphasis id="00DBEA9BFF8BF02073E711D8FAF9FC3C" box="[942,1353,953,977]" italics="true" pageId="1" pageNumber="92">the lateral part of the postventral end</emphasis>
”. In all
<taxonomicName id="F5AF4D0AFF8BF02070DE11BCFEAEFC18" box="[151,286,989,1013]" class="Branchiopoda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomopoda" pageId="1" pageNumber="92" phylum="Arthropoda" rank="order">Anomopoda</taxonomicName>
, paired filiform appendages of the ephippia, if present, are the torn off continuations of the bivalved body margin, which are directly a continuation of the dorsal margin in the
<taxonomicName id="F5AF4D0AFF8BF02073D01660FBA4FBF4" box="[921,1044,1025,1049]" class="Branchiopoda" family="Daphniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="family">Daphniidae</taxonomicName>
(
<figureCitation id="AA942A0CFF8BF020746C1660FB11FBF4" box="[1061,1185,1025,1049]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Figs 1AB</figureCitation>
). Among the ephippial appendages, which may extend posteriorly and anteriorly, there can be the (entirely or partly) posterior caudal spines in daphniids (if present;
<figureCitation id="AA942A0CFF8BF02071CF1628FDF7FB8C" box="[390,583,1097,1121]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Figs 1AB and D</figureCitation>
) as well as the remnants of the dorsal and posterior valve margins, often with serrations/spinules in the dorsal section and on the spines (
<figureCitation id="AA942A0CFF8BF020734D160CFCE6FB68" box="[772,854,1133,1157]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1B</figureCitation>
; see drawings in
<bibRefCitation id="563E4B78FF8BF0207444160CFB26FB68" author="Benzie, J. A. H." box="[1037,1174,1133,1157]" pageId="1" pageNumber="92" refId="ref4907" refString="Benzie, J. A. H. (2005) The genus Daphnia (including Daphniopsis) (Anomopoda: Daphniidae). Guides to the Identification of the Microinvertebrates of the Continental Waters of the World 21. Kenobi Productions, Ghent, Belgium and Backhuys Publishers, Leiden, 376 pp." type="book" year="2005">Benzie 2005</bibRefCitation>
;
<bibRefCitation id="563E4B78FF8BF02074E9160CFAC9FB68" author="Mergeay, J. &amp; Verschuren, D. &amp; De Meester, L." box="[1184,1401,1133,1157]" pageId="1" pageNumber="92" pagination="261 - 274" refId="ref6292" refString="Mergeay, J., Verschuren, D. &amp; De Meester, L. (2005) Daphnia species diversity in Kenya, and a key to the identification of their ephippia. Hydrobiologia, 542, 261 - 274. https: // doi. org / 10.1007 / s 10750 - 004 - 4952 - 6" type="journal article" year="2005">
Mergeay
<emphasis id="00DBEA9BFF8BF020754D160FFA8AFB68" box="[1284,1338,1133,1157]" italics="true" pageId="1" pageNumber="92">et al.</emphasis>
2005
</bibRefCitation>
). If present, caudal spines in daphniids can be singular and implanted posterodorsally (in
<taxonomicName id="F5AF4D0AFF8BF020746616F0FB3CFB44" authorityName="O. F. Muller" authorityYear="1776" box="[1071,1164,1169,1193]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF8BF020746616F0FB3CFB44" box="[1071,1164,1169,1193]" italics="true" pageId="1" pageNumber="92">Daphnia</emphasis>
</taxonomicName>
) or paired and implanted posteroventrally (e.g., in
<taxonomicName id="F5AF4D0AFF8BF02071E916D4FD57FB20" authority="Schoedler, 1858" authorityName="Schoedler" authorityYear="1858" box="[416,743,1205,1229]" class="Branchiopoda" genus="Scapholeberis" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF8BF02071E916D4FD87FB20" box="[416,567,1205,1229]" italics="true" pageId="1" pageNumber="92">Scapholeberis</emphasis>
Schoedler, 1858
</taxonomicName>
). In the subgenus
<taxonomicName id="F5AF4D0AFF8BF02073EE16D4FB03FB20" box="[935,1203,1205,1229]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="subGenus" subGenus="Ctenodaphnia">
<emphasis id="00DBEA9BFF8BF02073EE16D4FBB4FB20" box="[935,1028,1205,1229]" italics="true" pageId="1" pageNumber="92">Daphnia</emphasis>
(
<emphasis id="00DBEA9BFF8BF020745A16D4FB1AFB20" box="[1043,1194,1205,1229]" italics="true" pageId="1" pageNumber="92">Ctenodaphnia</emphasis>
)
</taxonomicName>
, because of the dorsal W-shaped structure of the headshield, the anterior appendages can be quite long (
<figureCitation id="AA942A0CFF8BF02073B616B8FBC9FB1C" box="[1023,1145,1241,1265]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Figs 1AB</figureCitation>
;
<bibRefCitation id="563E4B78FF8BF02074CD16B8FABFFB1C" author="Benzie, J. A. H." box="[1156,1295,1241,1265]" pageId="1" pageNumber="92" refId="ref4907" refString="Benzie, J. A. H. (2005) The genus Daphnia (including Daphniopsis) (Anomopoda: Daphniidae). Guides to the Identification of the Microinvertebrates of the Continental Waters of the World 21. Kenobi Productions, Ghent, Belgium and Backhuys Publishers, Leiden, 376 pp." type="book" year="2005">Benzie 2005</bibRefCitation>
), but it is not so in the subgenus
<taxonomicName id="F5AF4D0AFF8BF020712C169CFD89FAF8" box="[357,569,1277,1301]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="subGenus" subGenus="Daphnia">
<emphasis id="00DBEA9BFF8BF020712C169CFE72FAF8" box="[357,450,1277,1301]" italics="true" pageId="1" pageNumber="92">Daphnia</emphasis>
(
<emphasis id="00DBEA9BFF8BF020719A169CFD80FAF8" box="[467,560,1277,1301]" italics="true" pageId="1" pageNumber="92">Daphnia</emphasis>
)
</taxonomicName>
(
<figureCitation id="AA942A0CFF8BF0207201169CFD2DFAF8" box="[584,669,1277,1301]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1D</figureCitation>
). In other anomopods like chydorids, a torn margin can also be present (
<bibRefCitation id="563E4B78FF8BF02070D71740FEE0FAD4" author="Scourfield, D. J." box="[158,336,1313,1337]" pageId="1" pageNumber="92" pagination="217 - 244" refId="ref6498" refString="Scourfield, D. J. (1902) The ephippia of the lynceid Entomostraca. Journal of the Quekett Microscopical Club, Series 2, 8, 217 - 244." type="journal article" year="1902">Scourfield 1902</bibRefCitation>
;
<bibRefCitation id="563E4B78FF8BF02071121740FE48FAD4" author="Smirnov, N. N." box="[347,504,1313,1337]" pageId="1" pageNumber="92" refId="ref6592" refString="Smirnov, N. N. (1971) Chydoridae fauny mira. Rakoobraznie, 1. Fauna USSR, Leningrad, 531 pp. [English translation: Chydoridae of the world. Israel Program for Scientific Translations, Jerusalem, 1974]" type="book" year="1971">Smirnov 1971</bibRefCitation>
). In bosminids, the valve projections (caudal spines or mucros) are implanted posteroventrally and often they are preserved in subfossil ephippia (
<bibRefCitation id="563E4B78FF8BF020737B1724FC56FAB0" author="Scourfield, D. J." box="[818,998,1349,1373]" pageId="1" pageNumber="92" pagination="51 - 56" refId="ref6468" refString="Scourfield, D. J. (1901) The ephippium of Bosmina. Journal of the Quekett Microscopical Club, Series 2, 8, 51 - 56." type="journal article" year="1901">Scourfield 1901</bibRefCitation>
). The interpretation by Peng
<emphasis id="00DBEA9BFF8BF02075611727FAE9FAB0" box="[1320,1369,1349,1373]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. (
<figureCitation id="AA942A0CFF8BF02075381724FF06FA6C" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1F</figureCitation>
; drawings by
<bibRefCitation id="563E4B78FF8BF02071061708FDB4FA6C" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[335,516,1385,1409]" pageId="1" pageNumber="92" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8BF02071C5170BFE0CFA6C" box="[396,444,1385,1409]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. 2023
</bibRefCitation>
: Fig. 4) is that these appendages (“peduncles”) in
<taxonomicName id="F5AF4D0AFF8BF020746C170BFBC4FA6C" authorityName="Li" authorityYear="1993" box="[1061,1140,1386,1409]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF020746C170BFBC4FA6C" box="[1061,1140,1386,1409]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
are implanted laterally and submarginally at the “posterior” end. If this fossil would be an anomopod, any long filiform paired appendages would be continuing directly from the dorsal margin (
<figureCitation id="AA942A0CFF8BF020722E17D0FD6FFA24" box="[615,735,1457,1481]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Figs 1AB</figureCitation>
), not at a distance below it, and not midway on a straight posterior margin which is at a right angle with either the dorsal or the ventral margins (
<figureCitation id="AA942A0CFF8BF020739017B4FB9BFA00" box="[985,1067,1493,1517]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1E</figureCitation>
).
<bibRefCitation id="563E4B78FF8BF020740917B4FAB1FA00" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[1088,1281,1493,1517]" pageId="1" pageNumber="92" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8BF020743417B7FB1CFA00" box="[1149,1196,1493,1517]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. (2023)
</bibRefCitation>
compare their fossils with ephippia of
<taxonomicName id="F5AF4D0AFF8BF02071DC1798FCD0F9FC" authority="Forbes, 1893" authorityName="Forbes" authorityYear="1893" box="[405,864,1529,1553]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="species" species="pulicaria" subGenus="Daphnia">
<emphasis id="00DBEA9BFF8BF02071DC1798FD60F9FC" box="[405,720,1529,1553]" italics="true" pageId="1" pageNumber="92">Daphnia (Daphnia) pulicaria</emphasis>
Forbes, 1893
</taxonomicName>
and
<taxonomicName id="F5AF4D0AFF8BF02073DA1798FADFF9FC" authority="Claus, 1876" authorityName="Claus" authorityYear="1876" box="[915,1391,1529,1553]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="species" species="similis" subGenus="Ctenodaphnia">
<emphasis id="00DBEA9BFF8BF02073DA1798FC40F9FC" box="[915,1008,1529,1553]" italics="true" pageId="1" pageNumber="92">Daphnia</emphasis>
(
<emphasis id="00DBEA9BFF8BF02073B61798FB26F9FC" box="[1023,1174,1529,1553]" italics="true" pageId="1" pageNumber="92">Ctenodaphnia</emphasis>
)
<emphasis id="00DBEA9BFF8BF02074EC1798FB5BF9FC" box="[1189,1259,1529,1553]" italics="true" pageId="1" pageNumber="92">similis</emphasis>
Claus, 1876
</taxonomicName>
and write erroneously that “
<emphasis id="00DBEA9BFF8BF02071D0147CFD7EF9D8" box="[409,718,1565,1589]" italics="true" pageId="1" pageNumber="92">the two shorter peduncles of</emphasis>
<taxonomicName id="F5AF4D0AFF8BF020729F147CFC8FF9D8" authorityName="Claus" authorityYear="1876" box="[726,831,1565,1589]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="species" species="similis">D. similis</taxonomicName>
<emphasis id="00DBEA9BFF8BF0207301147FFA3AF9D8" box="[840,1418,1565,1589]" italics="true" pageId="1" pageNumber="92">originate from the central part of the postventral end</emphasis>
”. But the area where any appendages are implanted on the ephippia of
<taxonomicName id="F5AF4D0AFF8BF02073341423FC54F9B4" authorityName="Claus" authorityYear="1876" box="[893,996,1601,1625]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="species" species="similis">
<emphasis id="00DBEA9BFF8BF02073341423FC54F9B4" box="[893,996,1601,1625]" italics="true" pageId="1" pageNumber="92">D. similis</emphasis>
</taxonomicName>
is not the post-ventral end, it is dorsal as in all
<taxonomicName id="F5AF4D0AFF8BF02070911404FE85F990" authorityName="O. F. Muller" authorityYear="1776" box="[216,309,1637,1661]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF8BF02070911404FE85F990" box="[216,309,1637,1661]" italics="true" pageId="1" pageNumber="92">Daphnia</emphasis>
</taxonomicName>
, if appendages are present (
<figureCitation id="AA942A0CFF8BF02072201404FC9EF990" box="[617,814,1637,1661]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Figs 1AB and D</figureCitation>
), and these are most certainly not homologous structures to the appendages in
<taxonomicName id="F5AF4D0AFF8BF020713214EBFE7AF94C" authorityName="Li" authorityYear="1993" box="[379,458,1674,1697]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF020713214EBFE7AF94C" box="[379,458,1674,1697]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
(
<figureCitation id="AA942A0CFF8BF020719014E8FD9BF94C" box="[473,555,1673,1697]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1E</figureCitation>
). Ephippia of
<taxonomicName id="F5AF4D0AFF8BF020728F14EBFC0DF94C" authority="Straus, 1820" authorityName="Straus" authorityYear="1820" box="[710,957,1673,1697]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="species" species="magna">
<emphasis id="00DBEA9BFF8BF020728F14EBFC80F94C" box="[710,816,1674,1697]" italics="true" pageId="1" pageNumber="92">D. magna</emphasis>
Straus, 1820
</taxonomicName>
(
<figureCitation id="AA942A0CFF8BF020738514E8FBF4F94C" box="[972,1092,1673,1697]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Figs 1AB</figureCitation>
) look relatively similar to those of
<taxonomicName id="F5AF4D0AFF8BF02070FC14CFFEACF928" authorityName="Claus" authorityYear="1876" box="[181,284,1709,1733]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="species" species="similis">
<emphasis id="00DBEA9BFF8BF02070FC14CFFEACF928" box="[181,284,1709,1733]" italics="true" pageId="1" pageNumber="92">D. similis</emphasis>
</taxonomicName>
, yet the latter has a comparatively larger caudal spine as a (singular) projection, extending posteriorly from the posterodorsal corner of the body (e.g.,
<bibRefCitation id="563E4B78FF8BF020722914B0FC8DF904" author="Mergeay, J. &amp; Verschuren, D. &amp; De Meester, L." box="[608,829,1745,1769]" pageId="1" pageNumber="92" pagination="261 - 274" refId="ref6292" refString="Mergeay, J., Verschuren, D. &amp; De Meester, L. (2005) Daphnia species diversity in Kenya, and a key to the identification of their ephippia. Hydrobiologia, 542, 261 - 274. https: // doi. org / 10.1007 / s 10750 - 004 - 4952 - 6" type="journal article" year="2005">
Mergeay
<emphasis id="00DBEA9BFF8BF020728F14B3FD45F904" box="[710,757,1745,1769]" italics="true" pageId="1" pageNumber="92">et al</emphasis>
. 2005
</bibRefCitation>
). Long paired filiform appendages in
<taxonomicName id="F5AF4D0AFF8BF020749B14B3FA8AF904" authorityName="Claus" authorityYear="1876" box="[1234,1338,1745,1769]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="1" pageNumber="92" phylum="Arthropoda" rank="species" species="similis">
<emphasis id="00DBEA9BFF8BF020749B14B3FA8AF904" box="[1234,1338,1745,1769]" italics="true" pageId="1" pageNumber="92">D. similis</emphasis>
</taxonomicName>
ephippia (if any) would be in an anterior position (as in
<figureCitation id="AA942A0CFF8BF02072DC1494FD56F8E0" box="[661,742,1781,1805]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1B</figureCitation>
), while the long unpaired caudal spine would be posterior (
<figureCitation id="AA942A0CFF8BF02075241494FEB4F8DC" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Figs 1B and D</figureCitation>
). In addition, the appendages in
<taxonomicName id="F5AF4D0AFF8BF0207220157BFD08F8DC" authorityName="Li" authorityYear="1993" box="[617,696,1818,1841]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF0207220157BFD08F8DC" box="[617,696,1818,1841]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
show a broad base (
<figureCitation id="AA942A0CFF8BF02073D31578FC5CF8DC" box="[922,1004,1817,1841]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1E</figureCitation>
), almost a third of the entire margin on which they are implanted while in anomopods paired ephippial appendages are narrow and thin, as a continuation of the dorsal margin (
<figureCitation id="AA942A0CFF8BF02071771500FE23F894" box="[318,403,1889,1913]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1B</figureCitation>
). There are more arguments, e.g., the absence of any spinules on these appendages in
<taxonomicName id="F5AF4D0AFF8BF02075071503FA2DF894" authorityName="Li" authorityYear="1993" box="[1358,1437,1890,1913]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF02075071503FA2DF894" box="[1358,1437,1890,1913]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
(
<figureCitation id="AA942A0CFF8BF02070D615E4FF40F870" box="[159,240,1925,1949]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1E</figureCitation>
) which are often present in daphniids (
<figureCitation id="AA942A0CFF8BF02072D515E4FD5EF870" box="[668,750,1925,1949]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="1" pageNumber="92">Fig. 1B</figureCitation>
), and the fact that these “peduncles” seem actually very robust structures, sharply pointed at the terminal ends in
<taxonomicName id="F5AF4D0AFF8BF02072E615CBFD4EF82C" authorityName="Li" authorityYear="1993" box="[687,766,1962,1985]" genus="Kuqaia" pageId="1" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8BF02072E615CBFD4EF82C" box="[687,766,1962,1985]" italics="true" pageId="1" pageNumber="92">Kuqaia</emphasis>
</taxonomicName>
. In cladocerans, any paired ephippial filiform appendages that are remnants of the valve margins, are flimsy and would never be acute in such a way, while caudal spines (if present) would be implanted differently as stated earlier (dorsally in
<taxonomicName id="F5AF4D0AFF88F023735F12F6FCC3FF42" authorityName="O. F. Muller" authorityYear="1776" box="[790,883,151,175]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="2" pageNumber="93" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF88F023735F12F6FCC3FF42" box="[790,883,151,175]" italics="true" pageId="2" pageNumber="93">Daphnia</emphasis>
</taxonomicName>
;
<figureCitation id="AA942A0CFF88F02373C812F6FBAFFF5D" box="[897,1055,151,176]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="2" pageNumber="93">Figs 1B and D</figureCitation>
). So, not only are these appendages not homologous after re-interpretation, the original suggestion of similarity in position and implantation was based on an erroneous interpretation of
<taxonomicName id="F5AF4D0AFF88F02371AC12BEFDF2FF1A" authorityName="O. F. Muller" authorityYear="1776" box="[485,578,223,247]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="2" pageNumber="93" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF88F02371AC12BEFDF2FF1A" box="[485,578,223,247]" italics="true" pageId="2" pageNumber="93">Daphnia</emphasis>
</taxonomicName>
ephippia by
<bibRefCitation id="563E4B78FF88F023729F1281FC10FF15" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[726,928,223,248]" pageId="2" pageNumber="93" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF88F023735C1281FCFDFF1A" box="[789,845,223,247]" italics="true" pageId="2" pageNumber="93">et al.</emphasis>
(2023)
</bibRefCitation>
, who confused ventral and dorsal sides of the cladoceran body as well as
<typeStatus id="ED14882BFF88F02371F51365FE44FEF1" box="[444,500,260,284]" pageId="2" pageNumber="93">types</typeStatus>
and structures of ephippial appendages.
</paragraph>
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<paragraph id="32103689FF89F02270DE12F7FE54FE1E" blockId="3.[151,1436,150,499]" pageId="3" pageNumber="94">
<emphasis id="00DBEA9BFF89F02270DE12F7FEA3FF43" bold="true" box="[151,275,150,174]" pageId="3" pageNumber="94">FIGURE 1.</emphasis>
Basic features of anomopod ephippia in the
<taxonomicName id="F5AF4D0AFF89F022729C12F6FC8CFF42" box="[725,828,151,175]" class="Branchiopoda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cladocera" pageId="3" pageNumber="94" phylum="Arthropoda" rank="order">Cladocera</taxonomicName>
(examples of two extant species of
<taxonomicName id="F5AF4D0AFF89F02274EC12F6FB4EFF42" authorityName="O. F. Muller" authorityYear="1776" box="[1189,1278,151,175]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF89F02274EC12F6FB4EFF42" box="[1189,1278,151,175]" italics="true" pageId="3" pageNumber="94">Daphnia</emphasis>
</taxonomicName>
) in comparison to the Jurassic fossil
<emphasis id="00DBEA9BFF89F022712F12DAFDBDFF3F" box="[358,525,187,210]" italics="true" pageId="3" pageNumber="94">
<taxonomicName id="F5AF4D0AFF89F022712F12DAFE01FF3F" authorityName="Li" authorityYear="1993" box="[358,433,187,210]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">Kuqaia</taxonomicName>
scanicus
</emphasis>
(unknown taxonomy). The hypothesis that
<taxonomicName id="F5AF4D0AFF89F022738B12DAFBBDFF3F" authorityName="Li" authorityYear="1993" box="[962,1037,187,210]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF89F022738B12DAFBBDFF3F" box="[962,1037,187,210]" italics="true" pageId="3" pageNumber="94">Kuqaia</emphasis>
</taxonomicName>
fossils could be cladocerans, is rejected in this study. A.
<taxonomicName id="F5AF4D0AFF89F022710C12BEFD24FF1B" authorityName="Straus" authorityYear="1820" box="[325,660,223,247]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="species" species="magna" subGenus="Ctenodaphnia">
<emphasis id="00DBEA9BFF89F022710C12BEFE2EFF1A" box="[325,414,223,247]" italics="true" pageId="3" pageNumber="94">Daphnia</emphasis>
(
<emphasis id="00DBEA9BFF89F02271E712BEFD8EFF1A" box="[430,574,223,247]" italics="true" pageId="3" pageNumber="94">Ctenodaphnia</emphasis>
)
<emphasis id="00DBEA9BFF89F022720612BEFD24FF1B" box="[591,660,223,246]" italics="true" pageId="3" pageNumber="94">magna</emphasis>
</taxonomicName>
ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B.
<taxonomicName id="F5AF4D0AFF89F022723E1362FC70FEF7" authorityName="Straus" authorityYear="1820" box="[631,960,259,283]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="species" species="magna" subGenus="Ctenodaphnia">
<emphasis id="00DBEA9BFF89F022723E1362FD60FEF6" box="[631,720,259,283]" italics="true" pageId="3" pageNumber="94">Daphnia</emphasis>
(
<emphasis id="00DBEA9BFF89F02272971362FCDEFEF6" box="[734,878,259,283]" italics="true" pageId="3" pageNumber="94">Ctenodaphnia</emphasis>
)
<emphasis id="00DBEA9BFF89F02273321362FC70FEF7" box="[891,960,259,282]" italics="true" pageId="3" pageNumber="94">magna</emphasis>
</taxonomicName>
, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C.
<taxonomicName id="F5AF4D0AFF89F02273CE1346FB63FED3" authorityName="Straus" authorityYear="1820" box="[903,1235,295,319]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="species" species="magna" subGenus="Ctenodaphnia">
<emphasis id="00DBEA9BFF89F02273CE1346FC50FED2" box="[903,992,295,319]" italics="true" pageId="3" pageNumber="94">Daphnia</emphasis>
(
<emphasis id="00DBEA9BFF89F02273A71346FBCEFED2" box="[1006,1150,295,319]" italics="true" pageId="3" pageNumber="94">Ctenodaphnia</emphasis>
)
<emphasis id="00DBEA9BFF89F02274C51346FB63FED3" box="[1164,1235,295,318]" italics="true" pageId="3" pageNumber="94">magna</emphasis>
</taxonomicName>
<emphasis id="00DBEA9BFF89F022749D1346FB68FED3" box="[1236,1240,295,318]" italics="true" pageId="3" pageNumber="94">,</emphasis>
detail of the lateral surface of the ephippium. D.
<taxonomicName id="F5AF4D0AFF89F0227188132AFD78FE8E" box="[449,712,331,355]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="species" species="pulex" subGenus="Daphnia">
<emphasis id="00DBEA9BFF89F0227188132AFDAAFE8E" box="[449,538,331,355]" italics="true" pageId="3" pageNumber="94">Daphnia</emphasis>
(
<emphasis id="00DBEA9BFF89F0227260132AFD32FE8E" box="[553,642,331,355]" italics="true" pageId="3" pageNumber="94">Daphnia</emphasis>
)
<emphasis id="00DBEA9BFF89F02272D8132AFD78FE8E" box="[657,712,331,355]" italics="true" pageId="3" pageNumber="94">pulex</emphasis>
</taxonomicName>
species complex, ephippium in lateral view. E.
<emphasis id="00DBEA9BFF89F02274F9132AFAE8FE8F" box="[1200,1368,331,354]" italics="true" pageId="3" pageNumber="94">
<taxonomicName id="F5AF4D0AFF89F02274F9132AFB4BFE8F" authorityName="Li" authorityYear="1993" box="[1200,1275,331,354]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">Kuqaia</taxonomicName>
scanicus
</emphasis>
fossil, oriented as interpreted in the text of
<bibRefCitation id="563E4B78FF89F0227240130EFD71FE6A" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[521,705,367,391]" pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F022720B130EFDDFFE6A" box="[578,623,367,391]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. (2023)
</bibRefCitation>
, with “posterior” appendages. D.
<emphasis id="00DBEA9BFF89F0227450130EFB71FE6B" box="[1049,1217,367,390]" italics="true" pageId="3" pageNumber="94">
<taxonomicName id="F5AF4D0AFF89F0227450130EFBD4FE6B" authorityName="Li" authorityYear="1993" box="[1049,1124,367,390]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">Kuqaia</taxonomicName>
scanicus
</emphasis>
fossil, reconstruction after
<bibRefCitation id="563E4B78FF89F022708513F2FE32FE46" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[204,386,403,427]" pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F022714C13F2FE82FE46" box="[261,306,403,427]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. (2023)
</bibRefCitation>
shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from
<bibRefCitation id="563E4B78FF89F022708713D6FE18FE23" author="Mergeay, J. &amp; Verschuren, D. &amp; De Meester, L." box="[206,424,438,463]" pageId="3" pageNumber="94" pagination="261 - 274" refId="ref6292" refString="Mergeay, J., Verschuren, D. &amp; De Meester, L. (2005) Daphnia species diversity in Kenya, and a key to the identification of their ephippia. Hydrobiologia, 542, 261 - 274. https: // doi. org / 10.1007 / s 10750 - 004 - 4952 - 6" type="journal article" year="2005">
Mergeay
<emphasis id="00DBEA9BFF89F022716413D6FEE9FE22" box="[301,345,439,463]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. (2005)
</bibRefCitation>
(A,D) and
<bibRefCitation id="563E4B78FF89F022725013D6FD7CFE22" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[537,716,439,463]" pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F022721813D6FDCDFE22" box="[593,637,439,463]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. (2023)
</bibRefCitation>
(F); the other drawings are based on SEM photos in
<bibRefCitation id="563E4B78FF89F022749513D6FA2CFE22" author="Kotov, A. A. &amp; Kuzmina, S. A. &amp; Frolova, L. A. &amp; Zharov, A. A. &amp; Neritina, A. N. &amp; Smirnov, N. N." box="[1244,1436,439,463]" pageId="3" pageNumber="94" pagination="183 - 199" refId="ref6020" refString="Kotov, A. A., Kuzmina, S. A., Frolova, L. A., Zharov, A. A., Neritina, A. N. &amp; Smirnov, N. N. (2019) Ephippia of the Daphniidae (Branchiopoda: Cladocera) in Late Caenozoic deposits: untapped source of information for palaeoenvironment reconstructions in the Northern Holarctic. Invertebrate Zoology, 16, 183 - 199. https: // doi. org / 10.15298 / invertzool. 16.2.06" type="journal article" year="2019">
Kotov
<emphasis id="00DBEA9BFF89F022756813D6FAFDFE22" box="[1313,1357,439,463]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. (2019)
</bibRefCitation>
(B,C) and
<bibRefCitation id="563E4B78FF89F022714813BAFE07FE1E" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[257,439,475,499]" pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F022717313BAFEDEFE1E" box="[314,366,475,499]" italics="true" pageId="3" pageNumber="94">et al.</emphasis>
(2023)
</bibRefCitation>
(E).
</paragraph>
</caption>
<paragraph id="32103689FF89F022708E1042FBE8FC06" blockId="3.[151,1437,547,2011]" pageId="3" pageNumber="94">
<emphasis id="00DBEA9BFF89F022708E1042FECBFDD6" box="[199,379,547,571]" italics="true" pageId="3" pageNumber="94">
<emphasis id="00DBEA9BFF89F022708E1042FF58FDD6" bold="true" box="[199,232,547,571]" italics="true" pageId="3" pageNumber="94">(ii)</emphasis>
Dorsal keel.
</emphasis>
A narrow “dorsal” keel on the body is not uncommon in
<taxonomicName id="F5AF4D0AFF89F02273A81042FBD5FDD6" box="[993,1125,547,571]" class="Branchiopoda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomopoda" pageId="3" pageNumber="94" phylum="Arthropoda" rank="order">Anomopoda</taxonomicName>
(e.g.,
<taxonomicName id="F5AF4D0AFF89F02274E01042FA2CFDD6" authority="Frey, 1991" authorityName="Frey" authorityYear="1991" box="[1193,1436,547,571]" class="Branchiopoda" genus="Celsinotum" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF89F02274E01042FA92FDD6" box="[1193,1314,547,571]" italics="true" pageId="3" pageNumber="94">Celsinotum</emphasis>
<bibRefCitation id="563E4B78FF89F02275631042FA2CFDD6" author="Frey, D. G." box="[1322,1436,547,571]" pageId="3" pageNumber="94" pagination="11 - 48" refId="ref5734" refString="Frey, D. G. (1991) A new genus of alonine chydorid cladocerans from athalassic saline waters of New South Wales, Australia. Hydrobiologia, 224, 11 - 48. https: // doi. org / 10.1007 / BF 00006361" type="journal article" year="1991">
Frey,
<quantity id="F5579B6CFF89F022752E1042FA2CFDD6" box="[1383,1436,547,571]" metricMagnitude="1" metricUnit="m" metricValue="5.05714" pageId="3" pageNumber="94" unit="in" value="1991.0">1991</quantity>
</bibRefCitation>
</taxonomicName>
in the
<taxonomicName id="F5AF4D0AFF89F02270931026FEE9FDB2" box="[218,345,583,607]" class="Branchiopoda" family="Chydoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="family">Chydoridae</taxonomicName>
, or the
<emphasis id="00DBEA9BFF89F02271EE1026FD24FDB2" box="[423,660,583,607]" italics="true" pageId="3" pageNumber="94">
<taxonomicName id="F5AF4D0AFF89F02271EE1026FDAFFDB2" authorityName="W.Baird" authorityYear="1843" box="[423,543,583,607]" class="Branchiopoda" family="Macrothricidae" genus="Macrothrix" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">Macrothrix</taxonomicName>
paulensis-
</emphasis>
complex in the
<taxonomicName id="F5AF4D0AFF89F02273711026FC6FFDB2" authorityName="A.M.Norman &amp; G.S.Brady" authorityYear="1867" box="[824,991,583,607]" class="Branchiopoda" family="Macrothricidae" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="family">Macrothricidae</taxonomicName>
;
<bibRefCitation id="563E4B78FF89F02273A01026FBE8FDB2" author="Frey, D. G." box="[1001,1112,583,607]" pageId="3" pageNumber="94" pagination="11 - 48" refId="ref5734" refString="Frey, D. G. (1991) A new genus of alonine chydorid cladocerans from athalassic saline waters of New South Wales, Australia. Hydrobiologia, 224, 11 - 48. https: // doi. org / 10.1007 / BF 00006361" type="journal article" year="1991">Frey 1991</bibRefCitation>
;
<bibRefCitation id="563E4B78FF89F022742D1026FA93FDB3" author="Kotov, A. A. &amp; Wappler, T." box="[1124,1315,582,607]" pageId="3" pageNumber="94" pagination="1 - 9" refId="ref6231" refString="Kotov, A. A. &amp; Wappler, T. (2015) Fossil Daphnia (Ctenodaphnia) Dybowski &amp; Grochowski (Branchiopoda: Cladocera) in Cenozoic volcanogenic lakes in Germany, with discussion of their indicator value. Palaeontologica Electronica, 18.2.40 A, 1 - 9. https: // doi. org / 10.26879 / 542" type="book chapter" year="2015">
Kotov
<emphasis id="00DBEA9BFF89F02274E41026FB6CFDB2" box="[1197,1244,583,607]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. 2015
</bibRefCitation>
). In several adult
<taxonomicName id="F5AF4D0AFF89F0227098100AFEFFFD6E" box="[209,335,619,643]" class="Branchiopoda" family="Daphniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="family">Daphniidae</taxonomicName>
, for example in many members of the
<emphasis id="00DBEA9BFF89F0227295100AFCC3FD6E" box="[732,883,619,643]" italics="true" pageId="3" pageNumber="94">Ctenodaphnia</emphasis>
Dybowski &amp; Grochowski, 1895 lineage in
<taxonomicName id="F5AF4D0AFF89F0227571100AFA25FD6E" authorityName="O. F. Muller" authorityYear="1776" box="[1336,1429,619,643]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF89F0227571100AFA25FD6E" box="[1336,1429,619,643]" italics="true" pageId="3" pageNumber="94">Daphnia</emphasis>
</taxonomicName>
, the dorsal midline may often carry a series of small teeth (
<figureCitation id="AA942A0CFF89F022734410EEFCEFFD4A" box="[781,863,655,679]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Fig. 1B</figureCitation>
) of which the expression may be more intense in the presence of predators (e.g.,
<taxonomicName id="F5AF4D0AFF89F022718810D2FD37FD26" authorityName="Baird" authorityYear="1859" box="[449,647,691,715]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="species" species="atkinsoni">
<emphasis id="00DBEA9BFF89F022718810D2FD37FD26" box="[449,647,691,715]" italics="true" pageId="3" pageNumber="94">Daphnia atkinsoni</emphasis>
</taxonomicName>
-group). While there is sometimes a dorsal keel with a smooth or serrated margin in anomopods, it is not likely to find a ventral keel with an even margin in anomopod ephippia; either the ventral margin of the body is present (with remnants of structures), or when the ventral margins of the valves are torn off like a thread, it leaves an irregular appearance on the remaining ventral portion, like in many chydorids or macrothricids. In daphniids, the ephippial structure does not reach the ventral body margin entirely (
<figureCitation id="AA942A0CFF89F022745F1122FBDCFCB6" box="[1046,1132,835,859]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Fig. 1A</figureCitation>
). None of these features are visible in
<emphasis id="00DBEA9BFF89F02271491106FE32FC93" box="[256,386,871,894]" italics="true" pageId="3" pageNumber="94">K. scanicus,</emphasis>
where clearly a thick smooth keel or flange is present (up in
<figureCitation id="AA942A0CFF89F02274461106FBD0FC92" box="[1039,1120,871,895]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Fig. 1E</figureCitation>
; oriented downwards in
<bibRefCitation id="563E4B78FF89F022752E1106FEBFFC4E" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F02270DE11EAFF78FC4E" box="[151,200,907,931]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. 2023
</bibRefCitation>
: Figs 2ab). It does not correspond to the morphology of a dorsal keel or margin, or any ventral margin, in
<taxonomicName id="F5AF4D0AFF89F02270DE11CEFEAEFC2A" box="[151,286,943,967]" class="Branchiopoda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomopoda" pageId="3" pageNumber="94" phylum="Arthropoda" rank="order">Anomopoda</taxonomicName>
. In fact, it is not clear from the
<taxonomicName id="F5AF4D0AFF89F022723611CEFD7EFC2B" authorityName="Li" authorityYear="1993" box="[639,718,943,966]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF89F022723611CEFD7EFC2B" box="[639,718,943,966]" italics="true" pageId="3" pageNumber="94">Kuqaia</emphasis>
</taxonomicName>
fossil images whether these are actually bivalved structures (
<figureCitation id="AA942A0CFF89F022752411CEFF64FC06" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Figs 1EF</figureCitation>
). In
<taxonomicName id="F5AF4D0AFF89F022714D11B2FEC0FC06" box="[260,368,979,1003]" class="Branchiopoda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cladocera" pageId="3" pageNumber="94" phylum="Arthropoda" rank="order">Cladocera</taxonomicName>
(not in the raptorial lineages), the bivalved carapace is quite obvious.
</paragraph>
<paragraph id="32103689FF89F022708E1196FBA7F8A6" blockId="3.[151,1437,547,2011]" pageId="3" pageNumber="94">
<emphasis id="00DBEA9BFF89F022708E1196FE51FBE3" box="[199,481,1015,1039]" italics="true" pageId="3" pageNumber="94">
<emphasis id="00DBEA9BFF89F022708E1196FF40FBE2" bold="true" box="[199,240,1015,1039]" italics="true" pageId="3" pageNumber="94">(iii)</emphasis>
Valve Ornamentation.
</emphasis>
<taxonomicName id="F5AF4D0AFF89F02271AE1196FDDBFBE2" box="[487,619,1015,1039]" class="Branchiopoda" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anomopoda" pageId="3" pageNumber="94" phylum="Arthropoda" rank="order">Anomopoda</taxonomicName>
have a wide range of
<typeStatus id="ED14882BFF89F022731B1196FC3AFBE2" box="[850,906,1015,1039]" pageId="3" pageNumber="94">types</typeStatus>
of valve ornamentation, from smooth to pustulate (verrucose) to broadly reticulate, striate or with a fine mesh of striae (
<bibRefCitation id="563E4B78FF89F0227316167AFBEBFBDE" author="Smirnov, N. N. &amp; Kotov, A. A." box="[863,1115,1051,1075]" pageId="3" pageNumber="94" pagination="580 - 594" refId="ref6530" refString="Smirnov, N. N. &amp; Kotov, A. A. (2009) Morphological Radiation with Reference to the Carapace Valves of the Anomopoda (Crustacea: Cladocera). Internationale Revue der gesamten Hydrobiologie und Hydrographie, 94 (5), 580 - 594. https: // doi. org / 10.1002 / iroh. 200811121" type="journal article" year="2009">Smirnov &amp; Kotov 2009</bibRefCitation>
). Such reticulation is relatively better developed in benthic species and less expressed in zooplankters (
<bibRefCitation id="563E4B78FF89F02273E5165EFB83FBBA" author="Kotov, A. A." box="[940,1075,1087,1111]" pageId="3" pageNumber="94" pagination="1043 - 1059" refId="ref5819" refString="Kotov, A. A. (2006) Adaptations of the Anomopoda (Cladocera) for benthic mode of life. Zoologicheskii Zhurnal, 85 (9), 1043 - 1059. https: // doi. org / 10.1134 / S 0013873806110157" type="journal article" year="2006">Kotov 2006</bibRefCitation>
). This is the case in
<taxonomicName id="F5AF4D0AFF89F0227552165EFA29FBBA" box="[1307,1433,1087,1111]" class="Branchiopoda" family="Daphniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="family">Daphniidae</taxonomicName>
, which may often have a smooth or a dense reticulate pattern on the valves in adults. In most ephippia, the ornamentation becomes much more obvious, because this structure is heavily chitinized. This gives daphniid ephippia generally a densely ornamented surface with small projections (
<figureCitation id="AA942A0CFF89F022722816CAFD00FB2E" box="[609,688,1195,1219]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Fig. 1C</figureCitation>
) or a dense reticulation, which is also visible in fossil daphniid ephippia from the Jurassic-Cretaceous boundary (
<bibRefCitation id="563E4B78FF89F022721B16AEFD55FB0A" author="Kotov, A. A." box="[594,741,1231,1255]" pageId="3" pageNumber="94" pagination="523 - 528" refId="ref5910" refString="Kotov, A. A. (2009 a) New finding of Mesozoic ephippia of the Anomopoda (Crustacea: Cladocera). Journal of Natural History, 43 (9 / 10), 523 - 528. https: // doi. org / 10.1080 / 00222930802003020" type="journal article" year="2009">Kotov 2009a</bibRefCitation>
;
<bibRefCitation id="563E4B78FF89F02272BB16AEFB98FB0A" author="Van Damme, K. &amp; Kotov, A. A." box="[754,1064,1231,1255]" pageId="3" pageNumber="94" pagination="162 - 189" refId="ref6670" refString="Van Damme, K. &amp; Kotov, A. A. (2016) The fossil record of the Cladocera (Crustacea: Branchiopoda): evidence and hypotheses. Earth-Science Reviews, 163 (2016), 162 - 189. https: // doi. org / 10.1016 / j. earscirev. 2016.10.009" type="journal article" year="2016">Van Damme &amp; Kotov 2016</bibRefCitation>
). The ornamentation of
<taxonomicName id="F5AF4D0AFF89F022757B16AEFA31FB0B" authorityName="Li" authorityYear="1993" box="[1330,1409,1231,1254]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF89F022757B16AEFA31FB0B" box="[1330,1409,1231,1254]" italics="true" pageId="3" pageNumber="94">Kuqaia</emphasis>
</taxonomicName>
is entirely different, consisting of grooves and ribs, and it does not correspond to the ornamentation in any anomopod (in fact, any branchiopod). In
<emphasis id="00DBEA9BFF89F02271FF1776FD82FAC3" box="[438,562,1303,1326]" italics="true" pageId="3" pageNumber="94">K. scanicus</emphasis>
, parallel lines are at a right angle of the long (“ventral” and “dorsal”) margins or parallel to the “frontal” margin and these lines form some small ridges, giving it a ribbed appearance (
<figureCitation id="AA942A0CFF89F02274A5175AFA8FFABE" box="[1260,1343,1339,1363]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Fig. 1E</figureCitation>
;
<bibRefCitation id="563E4B78FF89F0227505175AFF43FA9A" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F02275C0175AFF1CFA9A" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. 2023
</bibRefCitation>
: Fig. 3a); the latter authors state that this ornamentation in
<taxonomicName id="F5AF4D0AFF89F022733D173EFC73FA9B" authorityName="Li" authorityYear="1993" box="[884,963,1375,1398]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF89F022733D173EFC73FA9B" box="[884,963,1375,1398]" italics="true" pageId="3" pageNumber="94">Kuqaia</emphasis>
</taxonomicName>
<emphasis id="00DBEA9BFF89F022739F173EFCD9FA76" italics="true" pageId="3" pageNumber="94">is suggestive of a morphologically simple, archaic group of cladocerans, possibly on the stem group lineage of</emphasis>
<taxonomicName id="F5AF4D0AFF89F022732617E2FC7FFA76" authorityName="O. F. Muller" authorityYear="1776" box="[879,975,1411,1435]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">Daphnia</taxonomicName>
”. Other
<taxonomicName id="F5AF4D0AFF89F022746C17E2FBC4FA77" authorityName="Li" authorityYear="1993" box="[1061,1140,1411,1434]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF89F022746C17E2FBC4FA77" box="[1061,1140,1411,1434]" italics="true" pageId="3" pageNumber="94">Kuqaia</emphasis>
</taxonomicName>
fossils shown in
<bibRefCitation id="563E4B78FF89F022756517E2FF6FFA52" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F022752E17E2FA2DFA76" box="[1383,1437,1411,1435]" italics="true" pageId="3" pageNumber="94">et al.</emphasis>
(2023)
</bibRefCitation>
have strong concentric or radiating deep lines and perforations on the lateral surfaces (
<emphasis id="00DBEA9BFF89F022743317C6FAA7FA53" box="[1146,1303,1447,1470]" italics="true" pageId="3" pageNumber="94">K. concentrica</emphasis>
Li, 1993,
<emphasis id="00DBEA9BFF89F02275CD17C6FF56FA0E" italics="true" pageId="3" pageNumber="94">K. radiata</emphasis>
Li, 1993), which are deep. Even the appendages in
<emphasis id="00DBEA9BFF89F022734017AAFC34FA0F" box="[777,900,1483,1506]" italics="true" pageId="3" pageNumber="94">K. scanicus</emphasis>
have a marginal row of indentations or transversal ridges (
<figureCitation id="AA942A0CFF89F02270A1178EFE8CF9EA" box="[232,316,1519,1543]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Fig. 1E</figureCitation>
;
<bibRefCitation id="563E4B78FF89F0227101178EFE4DF9EA" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[328,509,1519,1543]" pageId="3" pageNumber="94" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF89F02271CC178EFE05F9EA" box="[389,437,1519,1543]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. 2023
</bibRefCitation>
: Fig. 3c). None of these ornamentations correspond to
<typeStatus id="ED14882BFF89F0227419178EFB38F9EA" box="[1104,1160,1519,1543]" pageId="3" pageNumber="94">types</typeStatus>
known in any anomopod including fossil ephippia from the Jurassic-Cretaceous boundary (
<bibRefCitation id="563E4B78FF89F02273051472FC69F9C6" author="Kotov, A. A." box="[844,985,1555,1579]" pageId="3" pageNumber="94" pagination="523 - 528" refId="ref5910" refString="Kotov, A. A. (2009 a) New finding of Mesozoic ephippia of the Anomopoda (Crustacea: Cladocera). Journal of Natural History, 43 (9 / 10), 523 - 528. https: // doi. org / 10.1080 / 00222930802003020" type="journal article" year="2009">Kotov 2009a</bibRefCitation>
) which have a morphology congruent with extant daphniid and moinid species-groups (
<bibRefCitation id="563E4B78FF89F02272381456FC2FF9A2" author="Van Damme, K. &amp; Kotov, A. A." box="[625,927,1591,1615]" pageId="3" pageNumber="94" pagination="162 - 189" refId="ref6670" refString="Van Damme, K. &amp; Kotov, A. A. (2016) The fossil record of the Cladocera (Crustacea: Branchiopoda): evidence and hypotheses. Earth-Science Reviews, 163 (2016), 162 - 189. https: // doi. org / 10.1016 / j. earscirev. 2016.10.009" type="journal article" year="2016">Van Damme &amp; Kotov 2016</bibRefCitation>
). The general ornamentation of
<taxonomicName id="F5AF4D0AFF89F02274BF1456FAF5F9A3" authorityName="Li" authorityYear="1993" box="[1270,1349,1591,1614]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF89F02274BF1456FAF5F9A3" box="[1270,1349,1591,1614]" italics="true" pageId="3" pageNumber="94">Kuqaia</emphasis>
</taxonomicName>
falls, in no way, within any variability of known anomopods, and definitely not within the
<taxonomicName id="F5AF4D0AFF89F022745A143AFB22F99E" box="[1043,1170,1627,1651]" class="Branchiopoda" family="Daphniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="family">Daphniidae</taxonomicName>
. The suggestion that this is an “archaic”
<typeStatus id="ED14882BFF89F0227177141EFEDCF97A" box="[318,364,1663,1687]" pageId="3" pageNumber="94">type</typeStatus>
of cladoceran ornamentation, is here rejected. “Ribbed” cladoceran ornamentation is unknown if there are any long continuous concentric parallel lines in the group, it is from the retention of earlier instar carapaces during the moulting process, like in the anomopods
<taxonomicName id="F5AF4D0AFF89F022728E14A6FC53F932" authority="Smirnov, 1976" authorityName="Smirnov" authorityYear="1976" box="[711,995,1735,1759]" class="Branchiopoda" family="Ilyocryptidae" genus="Ilyocryptus" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF89F022728E14A6FC8FF932" box="[711,831,1735,1759]" italics="true" pageId="3" pageNumber="94">Ilyocryptus</emphasis>
Smirnov, 1976
</taxonomicName>
or
<taxonomicName id="F5AF4D0AFF89F022744114A6FB4DF932" authority="Sars, 1861" authorityName="Sars" authorityYear="1861" box="[1032,1277,1735,1759]" class="Branchiopoda" genus="Monospilus" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF89F022744114A6FB34F932" box="[1032,1156,1735,1759]" italics="true" pageId="3" pageNumber="94">Monospilus</emphasis>
Sars, 1861
</taxonomicName>
(
<bibRefCitation id="563E4B78FF89F022754514A6FA21F932" author="Kotov, A. A." box="[1292,1425,1735,1759]" pageId="3" pageNumber="94" pagination="1043 - 1059" refId="ref5819" refString="Kotov, A. A. (2006) Adaptations of the Anomopoda (Cladocera) for benthic mode of life. Zoologicheskii Zhurnal, 85 (9), 1043 - 1059. https: // doi. org / 10.1134 / S 0013873806110157" type="journal article" year="2006">Kotov 2006</bibRefCitation>
). But this is not the case here either. In addition, the double row of indentations on the inner side of each of the appendages in
<taxonomicName id="F5AF4D0AFF89F02270FA156EFEB2F8CB" authorityName="Li" authorityYear="1993" box="[179,258,1807,1830]" genus="Kuqaia" pageId="3" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF89F02270FA156EFEB2F8CB" box="[179,258,1807,1830]" italics="true" pageId="3" pageNumber="94">Kuqaia</emphasis>
</taxonomicName>
also does not make any sense if these were daphniid-like anomopods (there are no transversal indentations on the appendages but marginal structures such as denticles and short setules;
<figureCitation id="AA942A0CFF89F02273FE1552FBB9F8A6" box="[951,1033,1843,1867]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="3" pageNumber="94">Fig. 1B</figureCitation>
).
</paragraph>
<paragraph id="32103689FF89F025708E1536FEFEFE1E" blockId="3.[151,1437,547,2011]" lastBlockId="4.[151,1437,151,1004]" lastPageId="4" lastPageNumber="95" pageId="3" pageNumber="94">
<emphasis id="00DBEA9BFF89F022708E1536FDB8F882" box="[199,520,1879,1903]" italics="true" pageId="3" pageNumber="94">
<emphasis id="00DBEA9BFF89F022708E1536FF5CF882" bold="true" box="[199,236,1879,1903]" italics="true" pageId="3" pageNumber="94">(iv)</emphasis>
Embryo chambers/loculi.
</emphasis>
In the majority of anomopods and, in particular daphniids and (their sister lineage) moinids, the approximate shape and size of the embryo is visible (to various extent) in both extant and fossil ephippia by the convex expressions of the spheroid embryo chambers, sometimes called
<emphasis id="00DBEA9BFF89F02273FD15FEFC41F85A" box="[948,1009,1951,1975]" italics="true" pageId="3" pageNumber="94">loculi</emphasis>
(
<bibRefCitation id="563E4B78FF89F02273B415FEFB3BF85A" author="Kotov, A. A." box="[1021,1163,1951,1975]" pageId="3" pageNumber="94" pagination="523 - 528" refId="ref5910" refString="Kotov, A. A. (2009 a) New finding of Mesozoic ephippia of the Anomopoda (Crustacea: Cladocera). Journal of Natural History, 43 (9 / 10), 523 - 528. https: // doi. org / 10.1080 / 00222930802003020" type="journal article" year="2009">Kotov 2009a</bibRefCitation>
;
<bibRefCitation id="563E4B78FF89F02274DA15FEFAFDF85A" author="Kotov, A. A. &amp; Kuzmina, S. A. &amp; Frolova, L. A. &amp; Zharov, A. A. &amp; Neritina, A. N. &amp; Smirnov, N. N." box="[1171,1357,1951,1975]" pageId="3" pageNumber="94" pagination="183 - 199" refId="ref6020" refString="Kotov, A. A., Kuzmina, S. A., Frolova, L. A., Zharov, A. A., Neritina, A. N. &amp; Smirnov, N. N. (2019) Ephippia of the Daphniidae (Branchiopoda: Cladocera) in Late Caenozoic deposits: untapped source of information for palaeoenvironment reconstructions in the Northern Holarctic. Invertebrate Zoology, 16, 183 - 199. https: // doi. org / 10.15298 / invertzool. 16.2.06" type="journal article" year="2019">
Kotov
<emphasis id="00DBEA9BFF89F022749215FEFAB8F85A" box="[1243,1288,1951,1975]" italics="true" pageId="3" pageNumber="94">et al</emphasis>
. 2019
</bibRefCitation>
). These bulges in the ephippia can be less visible for
<taxonomicName id="F5AF4D0AFF89F02272C015A2FCB5F836" box="[649,773,1987,2011]" class="Branchiopoda" family="Chydoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="family">Chydoridae</taxonomicName>
and
<taxonomicName id="F5AF4D0AFF89F022737715A2FB25F836" authority="(Scourfield 1902)" baseAuthorityName="Scourfield" baseAuthorityYear="1902" box="[830,1173,1987,2011]" class="Branchiopoda" family="Eurycercidae" kingdom="Animalia" order="Diplostraca" pageId="3" pageNumber="94" phylum="Arthropoda" rank="family">
Eurycercidae (
<bibRefCitation id="563E4B78FF89F022739315A2FB3EF836" author="Scourfield, D. J." box="[986,1166,1987,2011]" pageId="3" pageNumber="94" pagination="217 - 244" refId="ref6498" refString="Scourfield, D. J. (1902) The ephippia of the lynceid Entomostraca. Journal of the Quekett Microscopical Club, Series 2, 8, 217 - 244." type="journal article" year="1902">Scourfield 1902</bibRefCitation>
)
</taxonomicName>
. Yet, even in chydorids, there is an embryonal space visible from the lateral outside of the shell as a general protuberance. In all
<taxonomicName id="F5AF4D0AFF8EF02574BD12F6FADFFF42" box="[1268,1391,151,175]" class="Branchiopoda" family="Daphniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="4" pageNumber="95" phylum="Arthropoda" rank="family">Daphniidae</taxonomicName>
and
<taxonomicName id="F5AF4D0AFF8EF02570DE12DAFF4DFF3E" authorityName="C.E.Goulden" authorityYear="1968" box="[151,253,187,211]" class="Branchiopoda" family="Moinidae" kingdom="Animalia" order="Diplostraca" pageId="4" pageNumber="95" phylum="Arthropoda" rank="family">Moinidae</taxonomicName>
the shape and orientation of the diapausing embryos is therefore very clear, and together with other characters such as the surface ornamentation, it can be used as a diagnostic feature (
<bibRefCitation id="563E4B78FF8EF02573D51281FBEBFF15" author="Kotov, A. A. &amp; Kuzmina, S. A. &amp; Frolova, L. A. &amp; Zharov, A. A. &amp; Neritina, A. N. &amp; Smirnov, N. N." box="[924,1115,223,248]" pageId="4" pageNumber="95" pagination="183 - 199" refId="ref6020" refString="Kotov, A. A., Kuzmina, S. A., Frolova, L. A., Zharov, A. A., Neritina, A. N. &amp; Smirnov, N. N. (2019) Ephippia of the Daphniidae (Branchiopoda: Cladocera) in Late Caenozoic deposits: untapped source of information for palaeoenvironment reconstructions in the Northern Holarctic. Invertebrate Zoology, 16, 183 - 199. https: // doi. org / 10.15298 / invertzool. 16.2.06" type="journal article" year="2019">
Kotov
<emphasis id="00DBEA9BFF8EF02573AF1281FBA5FF1A" box="[998,1045,223,247]" italics="true" pageId="4" pageNumber="95">et al</emphasis>
. 2019
</bibRefCitation>
); in
<taxonomicName id="F5AF4D0AFF8EF02574C112BEFAB3FF1A" box="[1160,1283,223,247]" class="Branchiopoda" family="Daphniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diplostraca" pageId="4" pageNumber="95" phylum="Arthropoda" rank="family">Daphniidae</taxonomicName>
and
<taxonomicName id="F5AF4D0AFF8EF025757F12BEFA2CFF1A" authorityName="C.E.Goulden" authorityYear="1968" box="[1334,1436,223,247]" class="Branchiopoda" family="Moinidae" kingdom="Animalia" order="Diplostraca" pageId="4" pageNumber="95" phylum="Arthropoda" rank="family">Moinidae</taxonomicName>
there are one to two embryos (sometimes none), not more. In
<taxonomicName id="F5AF4D0AFF8EF02573791365FCCFFEF6" authorityName="Li" authorityYear="1993" box="[816,895,260,283]" genus="Kuqaia" pageId="4" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8EF02573791365FCCFFEF6" box="[816,895,260,283]" italics="true" pageId="4" pageNumber="95">Kuqaia</emphasis>
</taxonomicName>
, all impressions or external morphology revealing the shape or size of potential diapausing embryos are absent.
<bibRefCitation id="563E4B78FF8EF025737E1349FC4BFEAD" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[823,1019,295,320]" pageId="4" pageNumber="95" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8EF025733D1349FC1CFED2" box="[884,940,295,319]" italics="true" pageId="4" pageNumber="95">et al.</emphasis>
(2023)
</bibRefCitation>
mention the absence of such loculi as a taxonomical feature of
<taxonomicName id="F5AF4D0AFF8EF02571E1132DFE47FE8E" authorityName="Li" authorityYear="1993" box="[424,503,332,355]" genus="Kuqaia" pageId="4" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8EF02571E1132DFE47FE8E" box="[424,503,332,355]" italics="true" pageId="4" pageNumber="95">Kuqaia</emphasis>
</taxonomicName>
, but in
<taxonomicName id="F5AF4D0AFF8EF0257204132AFD1AFE8E" authorityName="O. F. Muller" authorityYear="1776" box="[589,682,331,355]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="4" pageNumber="95" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF8EF0257204132AFD1AFE8E" box="[589,682,331,355]" italics="true" pageId="4" pageNumber="95">Daphnia</emphasis>
</taxonomicName>
or related lineages some shape of the embryonal chambers would be visible from outside. In addition, the fossil lacks all features of ephippia that may not have strongly expressed loculi, such as
<taxonomicName id="F5AF4D0AFF8EF02570FA13F2FEF3FE46" baseAuthorityName="Scourfield" baseAuthorityYear="1902" box="[179,323,403,427]" class="Branchiopoda" family="Eurycercidae" kingdom="Animalia" order="Diplostraca" pageId="4" pageNumber="95" phylum="Arthropoda" rank="family">Eurycercidae</taxonomicName>
, which have heavily chitinized carapaces. The absence of loculi is not a differential character in this case, it is merely because they are not anomopods. In general, the shape of these
<taxonomicName id="F5AF4D0AFF8EF025738713D9FBADFE22" authorityName="Li" authorityYear="1993" box="[974,1053,440,463]" genus="Kuqaia" pageId="4" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8EF025738713D9FBADFE22" box="[974,1053,440,463]" italics="true" pageId="4" pageNumber="95">Kuqaia</emphasis>
</taxonomicName>
fossils does not correspond, in any way, to ephippia.
</paragraph>
<paragraph id="32103689FF8EF025708E1061FECAFD1D" blockId="4.[151,1437,151,1004]" pageId="4" pageNumber="95">
From the above, it is clear that
<taxonomicName id="F5AF4D0AFF8EF02572541061FDDCFDFA" authorityName="Li" authorityYear="1993" box="[541,620,512,535]" genus="Kuqaia" pageId="4" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8EF02572541061FDDCFDFA" box="[541,620,512,535]" italics="true" pageId="4" pageNumber="95">Kuqaia</emphasis>
</taxonomicName>
is a not an ephippium of a daphniid-like (or other) anomopod. In fact, it is quite obvious that this is not a cladoceran (only anomopods produce ephippia). Objectively, the Lower Jurassic fossils in the study of
<bibRefCitation id="563E4B78FF8EF02571551029FE6CFD8D" author="Peng, J. &amp; Slater, S. M. &amp; McLoughlin, S. &amp; Vajda, V." box="[284,476,583,608]" pageId="4" pageNumber="95" pagination="0282247" refId="ref6351" refString="Peng, J., Slater, S. M., McLoughlin, S. &amp; Vajda, V. (2023) New species of Kuqaia from the Lower Jurassic of Sweden indicates a possible water flea (Crustacea: Branchiopoda) affinity. PLoS ONE, 18 (6), e 0282247. https: // doi. org / 10.1371 / journal. pone. 0282247" type="journal article" year="2023">
Peng
<emphasis id="00DBEA9BFF8EF02571111029FE37FDB2" box="[344,391,583,607]" italics="true" pageId="4" pageNumber="95">et al</emphasis>
. (2023)
</bibRefCitation>
do not correspond in the abovementioned morphological features, and a relationship with cladocerans is not realistic. In the past, other “cladoceran-like” fossils have been reported (e.g.,
<bibRefCitation id="563E4B78FF8EF02574FA100AFAD7FD6E" author="Adamczak, F." box="[1203,1383,619,643]" pageId="4" pageNumber="95" pagination="27 - 34" refId="ref4793" refString="Adamczak, F. (1965) On some Cambrian bivalved Crustacea and egg cases of the Cladocera. Stockholm Contributions in Geology, 13, 27 - 34." type="journal article" year="1965">Adamczak 1965</bibRefCitation>
), yet more recently, these have been considered ambiguous records (e.g., all Paleozoic fossils originally attributed to the group;
<bibRefCitation id="563E4B78FF8EF02570DE10D5FE70FD26" author="Van Damme, K. &amp; Kotov, A. A." box="[151,448,691,716]" pageId="4" pageNumber="95" pagination="162 - 189" refId="ref6670" refString="Van Damme, K. &amp; Kotov, A. A. (2016) The fossil record of the Cladocera (Crustacea: Branchiopoda): evidence and hypotheses. Earth-Science Reviews, 163 (2016), 162 - 189. https: // doi. org / 10.1016 / j. earscirev. 2016.10.009" type="journal article" year="2016">Van Damme &amp; Kotov 2016</bibRefCitation>
). Without actual segmented appendages (e.g., antennae) or recognisable ephippial features, it is definitely not easy.
</paragraph>
<paragraph id="32103689FF8EF025708E109AFE33FC06" blockId="4.[151,1437,151,1004]" pageId="4" pageNumber="95">
The characters discussed above include the
<typeStatus id="ED14882BFF8EF02572D2109DFD79FCF9" box="[667,713,764,788]" pageId="4" pageNumber="95">type</typeStatus>
and orientation of the external ornamentation, the implantation and shape of the long appendages, the keel and (the lack of) embryonal impressions or loculi. These are not just diagnostic differential characters within the cladocerans the
<taxonomicName id="F5AF4D0AFF8EF02572FC1125FCB4FCB6" authorityName="Li" authorityYear="1993" box="[693,772,836,859]" genus="Kuqaia" pageId="4" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8EF02572FC1125FCB4FCB6" box="[693,772,836,859]" italics="true" pageId="4" pageNumber="95">Kuqaia</emphasis>
</taxonomicName>
fossils are simply not part of the group in any way. Arguments can be made about the size (300 µm without the appendages which would be very small for daphniids; see the size of
<taxonomicName id="F5AF4D0AFF8EF02570DE11EDFF56FC4E" authorityName="Li" authorityYear="1993" box="[151,230,908,931]" genus="Kuqaia" pageId="4" pageNumber="91" rank="genus">
<emphasis id="00DBEA9BFF8EF02570DE11EDFF56FC4E" box="[151,230,908,931]" italics="true" pageId="4" pageNumber="95">Kuqaia</emphasis>
</taxonomicName>
relative to the two
<taxonomicName id="F5AF4D0AFF8EF02571FF11EAFDA3FC4E" authorityName="O. F. Muller" authorityYear="1776" box="[438,531,907,931]" class="Branchiopoda" family="Macrothricidae" genus="Daphnia" kingdom="Animalia" order="Diplostraca" pageId="4" pageNumber="95" phylum="Arthropoda" rank="genus">
<emphasis id="00DBEA9BFF8EF02571FF11EAFDA3FC4E" box="[438,531,907,931]" italics="true" pageId="4" pageNumber="95">Daphnia</emphasis>
</taxonomicName>
ephippia in
<figureCitation id="AA942A0CFF8EF02572D011EAFD56FC49" box="[665,742,907,932]" captionStart="FIGURE 1" captionStartId="3.[151,250,150,174]" captionTargetId="figure-87@2.[151,1436,307,1993]" captionTargetPageId="2" captionText="FIGURE 1. Basic features of anomopod ephippia in the Cladocera (examples of two extant species of Daphnia) in comparison to the Jurassic fossil Kuqaia scanicus (unknown taxonomy). The hypothesis that Kuqaia fossils could be cladocerans, is rejected in this study. A. Daphnia (Ctenodaphnia) magna ephippial female, habitus in lateral view and the outline of the ephippium (grey) with two diapausing embryos (black). B. Daphnia (Ctenodaphnia) magna, ephippium in lateral view (some typical dorsal spinules are shown and appendages, which may be torn off or broken). C. Daphnia (Ctenodaphnia) magna, detail of the lateral surface of the ephippium. D. Daphnia (Daphnia) pulex species complex, ephippium in lateral view. E. Kuqaia scanicus fossil, oriented as interpreted in the text of Peng et al. (2023), with “posterior” appendages. D. Kuqaia scanicus fossil, reconstruction after Peng et al. (2023) shown at the same scale as the cladoceran ephippia (AB and D) and oriented as in E. Images redrawn from Mergeay et al. (2005) (A,D) and Peng et al. (2023) (F); the other drawings are based on SEM photos in Kotov et al. (2019) (B,C) and Peng et al. (2023) (E)." figureDoi="http://doi.org/10.5281/zenodo.8329547" httpUri="https://zenodo.org/record/8329547/files/figure.png" pageId="4" pageNumber="95">Fig. 1F</figureCitation>
) and the entire shape of the structure which does not correspond to known body shapes in cladocerans (or ephippia). The latter are generally conserved, in particular for ephippia (Kotov &amp;
<bibRefCitation id="563E4B78FF8EF02570FB11B5FEC7FC06" author="Van Damme, K. &amp; Kotov, A. A." box="[178,375,979,1003]" pageId="4" pageNumber="95" pagination="162 - 189" refId="ref6670" refString="Van Damme, K. &amp; Kotov, A. A. (2016) The fossil record of the Cladocera (Crustacea: Branchiopoda): evidence and hypotheses. Earth-Science Reviews, 163 (2016), 162 - 189. https: // doi. org / 10.1016 / j. earscirev. 2016.10.009" type="journal article" year="2016">Van Damme 2016</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>