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<mods:title>Cosmopolitan abyssal lineages? A systematic study of East Pacific deep-sea squat lobsters (Decapoda: Galatheoidea: Munidopsidae)</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Rodríguez-Flores, Paula C.</mods:namePart>
<mods:affiliation>Department of Organismic and Evolutionary Biology, Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.</mods:affiliation>
<mods:nameIdentifier type="email">paularodriguezflores@g.harvard.edu</mods:nameIdentifier>
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<mods:namePart>Seid, Charlotte A.</mods:namePart>
<mods:affiliation>Scripps Institution of Oceanography, University of California San Diego, La Jolla, CA 92093, USA.</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Rouse, Greg W.</mods:namePart>
<mods:affiliation>Scripps Institution of Oceanography, University of California San Diego, La Jolla, CA 92093, USA.</mods:affiliation>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Giribet, Gonzalo</mods:namePart>
<mods:affiliation>Department of Organismic and Evolutionary Biology, Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.</mods:affiliation>
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<mods:date>2023</mods:date>
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<mods:number>2023-01-11</mods:number>
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<heading bold="true" box="[229,638,1657,1687]" centered="true" fontSize="12" level="2" pageId="31" pageNumber="45" reason="0">
<taxonomicName authority="Marin, 2020" authorityName="Marin" authorityYear="2020" box="[229,638,1657,1687]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="piipa">
<emphasis bold="true" box="[229,638,1657,1687]" pageId="31" pageNumber="45">
<emphasis bold="true" box="[229,472,1657,1687]" italics="true" pageId="31" pageNumber="45">Munidopsis piipa</emphasis>
<bibRefCitation author="Marin I" box="[482,638,1657,1687]" pageId="31" pageNumber="45" pagination="102263" refId="ref33471" refString="Marin I (2020) Northern unicorns of the depths: diversity of the genus Munidopsis Whiteaves, 1874 (Decapoda: Anomura: Munidopsidae) in the northwestern Pacific Ocean, with descriptions of three new species along the Russian coast. Progress in Oceanography 183, 102263. doi: 10.1016 / j. pocean. 2020.102263" type="journal article" year="2020">Marin, 2020</bibRefCitation>
</emphasis>
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<paragraph blockId="31.[361,507,1726,1751]" box="[361,507,1726,1751]" pageId="31" pageNumber="45">
(
<figureCitation box="[369,489,1726,1751]" captionStart="Fig" captionStartId="32.[314,351,1139,1162]" captionTargetBox="[314,1274,185,1108]" captionTargetId="figure-320@32.[313,1274,184,1109]" captionTargetPageId="32" captionText="Fig. 17. Munidopsis piipa Marin, 2020, California (CASIZ 190356). (a) Carapace and abdomen, dorsal view. (b) Sternal plastron. (c) Carapace and abdomen, lateral view. (d) Anterior carapace and rostrum, dorsal view." figureDoi="http://doi.org/10.5281/zenodo.7534778" httpUri="https://zenodo.org/record/7534778/files/figure.png" pageId="31" pageNumber="45">
Fig. 17 
<emphasis bold="true" box="[449,489,1726,1750]" italics="true" pageId="31" pageNumber="45">a–d</emphasis>
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.)
</paragraph>
</subSubSection>
<subSubSection pageId="31" pageNumber="45" type="materials_examined">
<paragraph blockId="31.[98,356,1838,1866]" box="[98,356,1838,1866]" pageId="31" pageNumber="45">
<heading bold="true" box="[98,356,1838,1866]" fontSize="11" level="3" pageId="31" pageNumber="45" reason="0">
<emphasis bold="true" box="[98,356,1838,1866]" pageId="31" pageNumber="45">Material examined</emphasis>
</heading>
</paragraph>
<paragraph blockId="31.[98,770,1885,1931]" lastBlockId="31.[818,1491,178,411]" pageId="31" pageNumber="45">
<emphasis bold="true" box="[98,285,1885,1904]" italics="true" pageId="31" pageNumber="45">Non-type specimens</emphasis>
. 
<materialsCitation pageId="31" pageNumber="45">USA: California, Pioneer Seamount, 7.ii.1950, 37.35000°N, 123.41666°W, 1097–822.96 m: 1 F 20.6 mm (CAS-IZ 190354).</materialsCitation>
— 
<materialsCitation pageId="31" pageNumber="45">USA: California, Pioneer Seamount, 7.ii.1950, 37.349957°N, 123.430026°W, 805.–988 m: 1 M 10.5 mm (CAS-IZ 190356).</materialsCitation>
— 
<materialsCitation pageId="31" pageNumber="45">
USA: Forty Mile Bank, off San Diego, California, leg. Greg Rouse and Nicolas Mongiardino Koch, R/V 
<emphasis bold="true" box="[1362,1421,258,277]" italics="true" pageId="31" pageNumber="45">Falkor</emphasis>
, ROV 
<emphasis bold="true" box="[818,910,285,304]" italics="true" pageId="31" pageNumber="45">SuBastian</emphasis>
dive S0446, SCB-172, 31.vii.2021, 32.6029°N, 118.02571°W, 1035 m: 1 F 13.4 mm (SIO-BIC C14546).
</materialsCitation>
— 
<materialsCitation pageId="31" pageNumber="45">
USA: California, leg. 
<taxonomicName authority="C. Vrijenhoek, R" authorityName="C. Vrijenhoek, R" box="[965,1194,338,357]" class="Insecta" family="Coccinellidae" genus="Robert" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="31" pageNumber="45" phylum="Arthropoda" rank="genus">Robert C. Vrijenhoek, R</taxonomicName>
/V 
<emphasis bold="true" box="[1225,1353,338,358]" italics="true" pageId="31" pageNumber="45">Western Flyer</emphasis>
, ROV 
<emphasis bold="true" box="[1417,1490,338,357]" italics="true" pageId="31" pageNumber="45">Tiburon</emphasis>
dive T665, 2.v.2004, 33.10070°N, 120.96000°W, 870 m, 1 specimen not sexed, not measured (USNM 1463979).
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="31" pageNumber="45" type="distribution">
<paragraph blockId="31.[818,984,459,487]" box="[818,984,459,487]" pageId="31" pageNumber="45">
<heading bold="true" box="[818,984,459,487]" fontSize="11" level="3" pageId="31" pageNumber="45" reason="0">
<emphasis bold="true" box="[818,984,459,487]" pageId="31" pageNumber="45">Distribution</emphasis>
</heading>
</paragraph>
<paragraph blockId="31.[818,1489,511,567]" pageId="31" pageNumber="45">Bering Sea (Piip submarine volcano) at 984-m depth, California, from 805- to 1097-m depth.</paragraph>
<paragraph blockId="31.[818,995,614,642]" box="[818,995,614,642]" pageId="31" pageNumber="45">
<heading bold="true" box="[818,995,614,642]" fontSize="11" level="3" pageId="31" pageNumber="45" reason="0">
<emphasis bold="true" box="[818,995,614,642]" pageId="31" pageNumber="45">Genetic data</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[818,1169,665,690]" pageId="31" pageNumber="45" type="description">
<paragraph blockId="31.[818,1169,665,690]" box="[818,1169,665,690]" pageId="31" pageNumber="45">
<emphasis bold="true" box="[818,861,665,689]" italics="true" pageId="31" pageNumber="45">COI</emphasis>
, 
<emphasis bold="true" box="[877,918,665,689]" italics="true" pageId="31" pageNumber="45">16S</emphasis>
rRNA and 
<emphasis bold="true" box="[1048,1091,665,689]" italics="true" pageId="31" pageNumber="45">28S</emphasis>
rRNA.
</paragraph>
</subSubSection>
<subSubSection pageId="31" pageNumber="45" type="discussion">
<paragraph blockId="31.[818,940,737,765]" box="[818,940,737,765]" pageId="31" pageNumber="45">
<heading bold="true" box="[818,940,737,765]" fontSize="11" level="3" pageId="31" pageNumber="45" reason="0">
<emphasis bold="true" box="[818,940,737,765]" pageId="31" pageNumber="45">Remarks</emphasis>
</heading>
</paragraph>
<paragraph blockId="31.[818,1491,787,1932]" pageId="31" pageNumber="45">
Newly recorded for the East Pacific. 
<taxonomicName authorityName="Marin" authorityYear="2020" box="[1293,1490,787,812]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="piipa">
<emphasis bold="true" box="[1293,1490,787,812]" italics="true" pageId="31" pageNumber="45">Munidopsis piipa</emphasis>
</taxonomicName>
belongs to a group of species with a rostral spine short and spiniform, frontal margin of the carapace straight and telson with 10–12 plates. This group includes 
<taxonomicName authority="Macpherson, 2011" authorityName="Macpherson" authorityYear="2011" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="ariadne">
<emphasis bold="true" italics="true" pageId="31" pageNumber="45">Munidopsis ariadne</emphasis>
<bibRefCitation author="Macpherson E &amp; Baba K" box="[909,1117,916,940]" pageId="31" pageNumber="45" pagination="39 - 71" refId="ref33260" refString="Macpherson E, Baba K (2011) Chapter 2. Taxonomy of squat lobsters. In ' The Biology of Squat Lobsters'. (Eds GCB Poore, ST Ahyong, J Taylor) pp. 39 - 71. (CSIRO Publishing: Melbourne, Vic., Australia)" type="book chapter" year="2011">Macpherson, 2011</bibRefCitation>
</taxonomicName>
from the eastern Mediterranean, 
<taxonomicName authority="Alcock &amp; Anderson, 1899" authorityName="Alcock &amp; Anderson" authorityYear="1899" box="[818,1354,948,972]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="goodridgii">
<emphasis bold="true" box="[818,1058,948,972]" italics="true" pageId="31" pageNumber="45">Munidopsis goodridgii</emphasis>
Alcock &amp; Anderson, 1899
</taxonomicName>
, 
<taxonomicName authority="Macpherson, Beuck &amp; Freiwald, 2016" authorityName="Macpherson, Beuck &amp; Freiwald" authorityYear="2016" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="karukera">
<emphasis bold="true" italics="true" pageId="31" pageNumber="45">Munidopsis karukera</emphasis>
<bibRefCitation author="Macpherson E &amp; Beuck L &amp; Freiwald A" box="[930,1374,980,1004]" pageId="31" pageNumber="45" pagination="405 - 416" refId="ref33417" refString="Macpherson E, Beuck L, Freiwald A (2016) Some species of Munidopsis from the Gulf of Mexico, Florida Straits and Caribbean Sea (Decapoda: Munidopsidae), with the description of two new species. Zootaxa 4137, 405 - 416. doi: 10.11646 / zootaxa. 4137.3.7" type="journal article" year="2016">Macpherson, Beuck &amp; Freiwald, 2016</bibRefCitation>
</taxonomicName>
from the Caribbean Sea, 
<taxonomicName authority="Schnabel &amp; Bruce, 2006" authorityName="Schnabel &amp; Bruce" authorityYear="2006" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="maunga">
<emphasis bold="true" box="[1014,1245,1012,1036]" italics="true" pageId="31" pageNumber="45">Munidopsis maunga</emphasis>
Schnabel &amp; Bruce, 2006
</taxonomicName>
, 
<taxonomicName authority="Henderson, 1885" authorityName="Henderson" authorityYear="1885" box="[905,1354,1044,1068]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="milleri">
<emphasis bold="true" box="[905,1123,1044,1068]" italics="true" pageId="31" pageNumber="45">Munidopsis milleri</emphasis>
<bibRefCitation author="Henderson JR" box="[1146,1354,1044,1068]" pageId="31" pageNumber="45" pagination="407 - 421" refId="ref32498" refString="Henderson JR (1885) XXXIX. - Diagnoses of the new species of Galatheidea collected during the ' Challenger' expedition. Annals and Magazine of Natural History 16, 407 - 421. doi: 10.1080 / 00222938509459908" type="journal article" year="1885">Henderson, 1885</bibRefCitation>
</taxonomicName>
from the Philippines, 
<taxonomicName authorityName="Koelbel" authorityYear="1892" box="[960,1225,1075,1099]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="polymorpha">
<emphasis bold="true" box="[960,1225,1075,1099]" italics="true" pageId="31" pageNumber="45">Munidopsis polymorpha</emphasis>
</taxonomicName>
from shallow caves in the Canary Islands, 
<taxonomicName authority="MacGilchrist, 1905" authorityName="MacGilchrist" authorityYear="1905" box="[1045,1490,1108,1132]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="spinipes">
<emphasis bold="true" box="[1045,1264,1108,1132]" italics="true" pageId="31" pageNumber="45">Munidopsis spinipes</emphasis>
MacGilchrist, 1905
</taxonomicName>
from the Bay of Bengal and 
<taxonomicName authority="Dong, Gan &amp; Li, 2021" authorityName="Dong, Gan &amp; X Li" authorityYear="2021" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="kexueae">
<emphasis bold="true" box="[1123,1343,1140,1164]" italics="true" pageId="31" pageNumber="45">Munidopsis kexueae</emphasis>
<bibRefCitation author="Dong D &amp; Gan Z &amp; Li X" pageId="31" pageNumber="45" pagination="306 - 355" refId="ref31805" refString="Dong D, Gan Z, Li X (2021) Descriptions of eleven new species of squat lobsters (Crustacea: Anomura) from seamounts around the Yap and Mariana Trenches with notes on DNA barcodes and phylogeny. Zoological Journal of the Linnean Society 192, 306 - 355. doi: 10.1093 / zoolinnean / zlab 003" type="journal article" year="2021">Dong, Gan &amp; Li, 2021</bibRefCitation>
</taxonomicName>
from seamounts near the Yap Trench, West Pacific.
</paragraph>
</subSubSection>
<subSubSection pageId="31" pageNumber="45" type="reference_group">
<paragraph blockId="31.[818,1491,787,1932]" pageId="31" pageNumber="45">
<taxonomicName authorityName="Marin" authorityYear="2020" box="[850,1035,1204,1228]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="piipa">
<emphasis bold="true" box="[850,1035,1204,1228]" italics="true" pageId="31" pageNumber="45">Munidopsis piipa</emphasis>
</taxonomicName>
closely resembles 
<taxonomicName authorityName="Dong, Gan &amp; X Li" authorityYear="2021" box="[1244,1371,1204,1228]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="kexueae">
<emphasis bold="true" box="[1244,1371,1204,1228]" italics="true" pageId="31" pageNumber="45">M. kexueae</emphasis>
</taxonomicName>
both morphologically and genetically. The descriptions of these species almost overlapped during the time of revisions and publication, therefore the species were not compared before our work. Phylogenetic and some species delimitation analyses do not support these species hypotheses, considering 
<taxonomicName authorityName="Marin" authorityYear="2020" box="[864,960,1396,1420]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="piipa">
<emphasis bold="true" box="[864,960,1396,1420]" italics="true" pageId="31" pageNumber="45">M. piipa</emphasis>
</taxonomicName>
and 
<taxonomicName authorityName="Dong, Gan &amp; X Li" authorityYear="2021" box="[1024,1154,1396,1420]" class="Malacostraca" family="Munidopsidae" genus="Munidopsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="species" species="kexueae">
<emphasis bold="true" box="[1024,1154,1396,1420]" italics="true" pageId="31" pageNumber="45">M. kexueae</emphasis>
</taxonomicName>
as a single taxon. However, haplotype networks demonstrate large genetic distances between the two species, with up to 4% of divergence for 
<emphasis bold="true" box="[818,861,1492,1516]" italics="true" pageId="31" pageNumber="45">COI</emphasis>
or 19 mutational steps (
<figureCitation box="[1173,1243,1492,1516]" captionStart="Fig" captionStartId="9.[156,193,1334,1357]" captionTargetBox="[156,1432,187,1287]" captionTargetId="figure-208@9.[156,1432,184,1289]" captionTargetPageId="9" captionText="Fig. 4. Haplotype networks with haplotype distributions for (a) Munidopsis piipa + M. kexuae and (b) M. similis + M. asiatica." figureDoi="http://doi.org/10.5281/zenodo.7534745" httpUri="https://zenodo.org/record/7534745/files/figure.png" pageId="31" pageNumber="45">Fig. 4</figureCitation>
). The percentage of divergence between these species is low compared to other squat lobsters in the family, (e.g. 
<bibRefCitation author="Rodriguez-Flores PC &amp; Macpherson E &amp; Machordom A" pageId="31" pageNumber="45" pagination="201 - 256" refId="ref34592" refString="Rodriguez-Flores PC, Macpherson E, Machordom A (2019 b) Revision of the squat lobsters of the genus Leiogalathea Baba, 1969 (Crustacea, Decapoda, Munidopsidae) with the description of 15 new species. Zootaxa 4560, 201 - 256. doi: 10.11646 / ZOOTAXA. 4560.2.1" type="journal article" year="2019">
Rodríguez-Flores 
<emphasis bold="true" box="[1429,1489,1555,1579]" italics="true" pageId="31" pageNumber="45">et al.</emphasis>
2019 
<emphasis bold="true" box="[875,887,1588,1612]" italics="true" pageId="31" pageNumber="45">b</emphasis>
</bibRefCitation>
, 
<taxonomicName box="[904,1043,1588,1612]" class="Malacostraca" family="Munidopsidae" genus="Leiogalathea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="31" pageNumber="45" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[904,1043,1588,1612]" italics="true" pageId="31" pageNumber="45">Leiogalathea</emphasis>
</taxonomicName>
); although some species of the abyssal clade have been delimited based on similar genetic distance values (
<bibRefCitation author="Jones WJ &amp; Macpherson E" box="[911,1249,1652,1676]" pageId="31" pageNumber="45" pagination="477 - 501" refId="ref32784" refString="Jones WJ, Macpherson E (2007) Molecular phylogeny of the East Pacific squat lobsters of the genus Munidopsis (Decapoda: Galatheidae) with the descriptions of seven new species. Journal of Crustacean Biology 27 (3), 477 - 501. doi: 10.1651 / S- 2791.1" type="journal article" year="2007">Jones and Macpherson 2007</bibRefCitation>
; 
<bibRefCitation author="Dong D &amp; Xu P &amp; Li XZ &amp; Wang C" box="[1269,1473,1651,1676]" pageId="31" pageNumber="45" pagination="8089" refId="ref31748" refString="Dong D, Xu P, Li XZ, Wang C (2019) Munidopsis species (Crustacea: Decapoda: Munidopsidae) from carcass falls in Weijia Guyot, West Pacific, with recognition of a new species based on integrative taxonomy. PeerJ 7, e 8089. doi: 10.7717 / peerj. 8089" type="journal article" year="2019">
Dong 
<emphasis bold="true" box="[1341,1402,1651,1675]" italics="true" pageId="31" pageNumber="45">et al.</emphasis>
2019
</bibRefCitation>
). According to the original species descriptions (
<bibRefCitation author="Marin I" pageId="31" pageNumber="45" pagination="102263" refId="ref33471" refString="Marin I (2020) Northern unicorns of the depths: diversity of the genus Munidopsis Whiteaves, 1874 (Decapoda: Anomura: Munidopsidae) in the northwestern Pacific Ocean, with descriptions of three new species along the Russian coast. Progress in Oceanography 183, 102263. doi: 10.1016 / j. pocean. 2020.102263" type="journal article" year="2020">Marin 2020</bibRefCitation>
; 
<bibRefCitation author="Dong D &amp; Li X" box="[891,1084,1716,1741]" pageId="31" pageNumber="45" pagination="1841 - 1853" refId="ref31689" refString="Dong D, Li X (2021) Two new species of the genus Munidopsis Whiteaves, 1874 (Crustacea: Anomura: Munidopsidae) from the Caroline Ridge, South of the Mariana Trench. Journal of Oceanology and Limnology 39, 1841 - 1853. doi: 10.1007 / s 00343 - 021 - 0385 - 6" type="journal article" year="2021">
Dong 
<emphasis bold="true" box="[960,1017,1716,1740]" italics="true" pageId="31" pageNumber="45">et al.</emphasis>
2021
</bibRefCitation>
), spinulation on the carapace margins and relative width of the rostrum are different in both taxa. However, these morphological and genetic differences could be part of intraspecific variation and population genetic structure, as very few specimens have been analysed to date. More data, including the analyses of more specimens, would be desirable to solve this taxonomic problem.
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