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<mods:title id="E15BF320CBABF8836738ACE112614B5F">The taxonomic status of Hymenodora (Crustacea: Oplophoroidea): morphological and molecular analyses suggest a new family and an undescribed diversity deep in the sea</mods:title>
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<paragraph id="8BAF36CC2251B662FF2C36DDFD19BBEB" blockId="13.[129,778,1754,1973]" box="[240,666,1754,1780]" pageId="13" pageNumber="349">
<emphasis id="B964EADE2251B662FF2C36DDFD19BBEB" bold="true" box="[240,666,1754,1780]" pageId="13" pageNumber="349">
Morphological traits in
<taxonomicName id="4C104D4F2251B662FE3536DDFD19BBEB" authorityName="Lunina &amp; Kulagin &amp; Vereshchaka" authorityYear="2024" box="[489,666,1754,1780]" family="Hymenodoridae" kingdom="Animalia" order="Decapoda" pageId="13" pageNumber="349" rank="family">Hymenodoridae</taxonomicName>
</emphasis>
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<subSubSection id="C30A65472251B661FF5D3707FDF4BA5C" lastPageId="14" lastPageNumber="350" pageId="13" pageNumber="349" type="description">
<paragraph id="8BAF36CC2251B661FF5D3707FD23BC19" blockId="13.[129,778,1754,1973]" lastBlockId="14.[113,763,144,1859]" lastPageId="14" lastPageNumber="350" pageId="13" pageNumber="349">
The clade
<taxonomicName id="4C104D4F2251B662FF323707FE1BBA07" authorityName="Lunina &amp; Kulagin &amp; Vereshchaka" authorityYear="2024" box="[238,408,1792,1816]" family="Hymenodoridae" pageId="13" pageNumber="349" rank="family">Hymenodoridae</taxonomicName>
is robust in all analyses and represents a deep-sea branch of
<taxonomicName id="4C104D4F2251B662FE713727FDCDBA27" authorityName="Dana" authorityYear="1852" box="[429,590,1824,1848]" pageId="13" pageNumber="346" rank="superFamily" superFamily="Oplophoroidea">Oplophoroidea</taxonomicName>
. Although juveniles of
<taxonomicName id="4C104D4F2251B662FF1B3738FEC5BA48" authorityName="G. O. Sars" authorityYear="1877" box="[199,326,1855,1879]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="13" pageNumber="349" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2251B662FF1B3738FEC5BA48" box="[199,326,1855,1879]" italics="true" pageId="13" pageNumber="349">Hymenodora</emphasis>
</taxonomicName>
may occasionally occur in the mesopelagic zone, especially in high latitudes (
<bibRefCitation id="EF814B3D2251B662FE323759FD12BA69" author="Jonsdottir IG" box="[494,657,1886,1910]" pageId="13" pageNumber="349" pagination="190 - 6" refId="ref10774" refString="Jonsdottir IG. Spatial distribution of Northern ambereye (Hymenodora glacialis) around Iceland. Marine Biology Research 2014; 10: 190 - 6." type="journal article" year="2014">Jónsdóttir 2014</bibRefCitation>
), and even contribute to the stomach content of Leachs storm petrel
<taxonomicName id="4C104D4F2251B662FF5D379AFD87BAAA" authority="(Vieillot, 1818)" baseAuthorityName="Vieillot" baseAuthorityYear="1818" box="[129,516,1949,1973]" class="Aves" family="Hydrobatidae" genus="Hydrobates" kingdom="Animalia" order="Procellariiformes" pageId="13" pageNumber="349" phylum="Chordata" rank="species" species="leucorhous">
<emphasis id="B964EADE2251B662FF5D379AFEDEBAAA" box="[129,349,1949,1973]" italics="true" pageId="13" pageNumber="349">Hydrobates leucorhous</emphasis>
(Vieillot, 1818)
</taxonomicName>
(Steele and Montevecchi 1994), adults live mainly in the bathypelagic (Vereshchaka
<emphasis id="B964EADE2251B662FCE536FBFCEEBA0C" box="[825,877,1787,1811]" italics="true" pageId="13" pageNumber="349">et al.</emphasis>
2019) and abyssopelagic zones (e.g.
<taxonomicName id="4C104D4F2251B662FB2336FBFADEBA0C" authorityName="Wasmer" authorityYear="2022" box="[1279,1373,1787,1811]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="13" pageNumber="349" phylum="Arthropoda" rank="species" species="chacei">
<emphasis id="B964EADE2251B662FB2336FBFADEBA0C" box="[1279,1373,1787,1811]" italics="true" pageId="13" pageNumber="349">H. chacei</emphasis>
</taxonomicName>
; Wasmer 2022). The whole family
<taxonomicName id="4C104D4F2251B662FBE6371DFB67BA2D" authorityName="Lunina &amp; Kulagin &amp; Vereshchaka" authorityYear="2024" box="[1082,1252,1818,1842]" family="Hymenodoridae" pageId="13" pageNumber="349" rank="family">Hymenodoridae</taxonomicName>
is supported by three synapomorphies, which are linked to mouthparts (
<figureCitation id="132B2A492251B662FABE373DFA32BA4E" box="[1378,1457,1849,1874]" captionStart="Figure 5" captionStartId="7.[129,194,1068,1092]" captionTargetBox="[131,1466,146,1039]" captionTargetId="figure-437@7.[129,1473,144,1040]" captionTargetPageId="7" captionText="Figure 5. Phylogenetic analyses of Oplophoroidea.A, molecular tree (six-gene analysis). Only supported clades are shown (analysis 3). Statistical support is indicated as Bayesian posterior probabilities (black, below branches) and maximum likelihood bootstrap (blue, above branches). B, morphological tree (analyses 4 and 5; trees were similar, and supports are divided by an oblique stroke character if they differed). Only supported clades are shown. Statistical support is indicated as Bremer support (black, below branches) and maximum likelihood bootstrap (red, above branches). C, common morphological synapomorphies (see coding in Supporting Information, Table S4)." figureDoi="http://doi.org/10.5281/zenodo.11240008" httpUri="https://zenodo.org/record/11240008/files/figure.png" pageId="13" pageNumber="349">Fig. 5C</figureCitation>
).
<taxonomicName id="4C104D4F2251B662FCE5375EFC3BBA6E" authorityName="G. O. Sars" authorityYear="1877" box="[825,952,1881,1905]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="13" pageNumber="349" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2251B662FCE5375EFC3BBA6E" box="[825,952,1881,1905]" italics="true" pageId="13" pageNumber="349">Hymenodora</emphasis>
</taxonomicName>
shows omnivorous behaviour, with a strong tendency towards carnivory (
<bibRefCitation id="EF814B3D2251B662FB8D377FFAA1BA8F" author="Kreibich T &amp; Hagen W &amp; Saborowski R" box="[1105,1314,1912,1936]" pageId="13" pageNumber="349" pagination="105 - 15" refId="ref10832" refString="Kreibich T, Hagen W, Saborowski R. Food utilization of two pelagic crustaceans in the Greenland Sea: Meganyctiphanes norvegica (Euphausiacea) and Hymenodora glacialis (Decapoda, Caridea). Marine Ecology Progress Series 2010; 413: 105 - 15." type="journal article" year="2010">
Kreibich
<emphasis id="B964EADE2251B662FB6F377EFB65BA8F" box="[1203,1254,1912,1936]" italics="true" pageId="13" pageNumber="349">et al.</emphasis>
2010
</bibRefCitation>
), and the
<typeStatus id="54AB886E2251B662FA4B377EFA41BA8E" box="[1431,1474,1913,1937]" pageId="13" pageNumber="349" type="syntype">synapomorphies</typeStatus>
linked to the mouthparts are likely adaptive to this
<typeStatus id="54AB886E2252B661FFAD3097FF1CBDB7" box="[113,159,144,168]" pageId="14" pageNumber="350">type</typeStatus>
of feeding in the bathypelagic realm. The simplified second maxilla and modified first maxilliped might be more efficient for consuming small deep-sea copepods that contribute significantly to the diet of
<taxonomicName id="4C104D4F2252B661FEE730E9FD18BC19" authority="(Kreibich et al. 2010)" baseAuthorityName="Kreibich" baseAuthorityYear="2010" box="[315,667,238,262]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FEE730E9FE39BC19" box="[315,442,238,262]" italics="true" pageId="14" pageNumber="350">Hymenodora</emphasis>
(
<bibRefCitation id="EF814B3D2252B661FE1630E9FD0CBC19" author="Kreibich T &amp; Hagen W &amp; Saborowski R" box="[458,655,238,262]" pageId="14" pageNumber="350" pagination="105 - 15" refId="ref10832" refString="Kreibich T, Hagen W, Saborowski R. Food utilization of two pelagic crustaceans in the Greenland Sea: Meganyctiphanes norvegica (Euphausiacea) and Hymenodora glacialis (Decapoda, Caridea). Marine Ecology Progress Series 2010; 413: 105 - 15." type="journal article" year="2010">
Kreibich
<emphasis id="B964EADE2252B661FDF430E8FDD4BC19" box="[552,599,238,262]" italics="true" pageId="14" pageNumber="350">et al.</emphasis>
2010
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BAF36CC2252B661FF51310AFD9EBF3F" blockId="14.[113,763,144,1859]" pageId="14" pageNumber="350">
Within the clade,
<taxonomicName id="4C104D4F2252B661FE91310AFE74BC3A" authorityName="Lunina &amp; Kulagin &amp; Vereshchaka" authorityYear="2024" box="[333,503,269,293]" family="Hymenodoridae" pageId="14" pageNumber="349" rank="family">Hymenodoridae</taxonomicName>
show further specialization for feeding through two nearly opposite traits.
<taxonomicName id="4C104D4F2252B661FDA7312AFD79BC5A" box="[635,762,301,325]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FDA7312AFD79BC5A" box="[635,762,301,325]" italics="true" pageId="14" pageNumber="350">Hymenodora</emphasis>
</taxonomicName>
possesses a subovoid terminal segment of the second maxilliped attached diagonally, which is likely unsuccessful in catching large and fast-moving planktonic animals (
<figureCitation id="132B2A492252B661FDE3318CFD0FBCBD" box="[575,652,395,419]" captionStart="Figure 7" captionStartId="11.[129,194,1351,1375]" captionTargetBox="[162,1441,146,1321]" captionTargetId="figure-226@11.[160,1443,144,1323]" captionTargetPageId="11" captionText="Figure 7. Morphological characters distinguishing three families of Oplophoroidea.AE, Hymenodoridae (Hymenodora frontalis, St. 7497 d271) mandibles (A), second maxilla (B), proximal endite of second maxilla (C), first maxilliped (D) and second maxilliped (E). FJ, Oplophoridae (Systellaspis debilis, 179RT 23.1) mandibles (F), second maxilla (G), proximal endite of second maxilla (H), first maxilliped (I) and second maxilliped (J). KO, Acanthephyridae (Acanthephyra quadrispinosa Kemp, 1939, AMK87 7277-D13) mandibles (K), second maxilla (L), proximal endite of second maxilla (M), first maxilliped (N) and second maxilliped (O). All scale bars:1 mm." figureDoi="http://doi.org/10.5281/zenodo.11240012" httpUri="https://zenodo.org/record/11240012/files/figure.png" pageId="14" pageNumber="350">Fig. 7E</figureCitation>
), whereas
<taxonomicName id="4C104D4F2252B661FFAD31ADFF54BCDD" authorityName="Vereshchaka, Kulagin &amp; Lunina" authorityYear="2021" box="[113,215,426,450]" class="Malacostraca" family="Acanthephyridae" genus="Sclerodora" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FFAD31ADFF54BCDD" box="[113,215,426,450]" italics="true" pageId="14" pageNumber="350">Sclerodora</emphasis>
</taxonomicName>
has a specialized subtriangular terminal segment attached to the penultimate segment transversely and bearing robust terminal setae (
<bibRefCitation id="EF814B3D2252B661FE9431EEFDB8BF1E" author="Vereshchaka A &amp; Kulagin D &amp; Lunina A" box="[328,571,489,513]" pageId="14" pageNumber="350" pagination="536" refId="ref11584" refString="Vereshchaka A, Kulagin D, Lunina A. A new shrimp genus (Crustacea: Decapoda) from the Deep Atlantic and an unusual cleaning mechanism of pelagic decapods. Diversity 2021; 13: 536." type="journal article" year="2021">
Vereshchaka
<emphasis id="B964EADE2252B661FE0C31EEFD82BF1E" box="[464,513,489,513]" italics="true" pageId="14" pageNumber="350">et al.</emphasis>
2021
</bibRefCitation>
: fig. 4E), which is adapted for catching actively moving prey.
</paragraph>
<paragraph id="8BAF36CC2252B661FF513220FF50BE25" blockId="14.[113,763,144,1859]" pageId="14" pageNumber="350">
Another pair of alternative traits in
<taxonomicName id="4C104D4F2252B661FE293220FD1CBF20" authorityName="Lunina &amp; Kulagin &amp; Vereshchaka" authorityYear="2024" box="[501,671,551,575]" family="Hymenodoridae" pageId="14" pageNumber="349" rank="family">Hymenodoridae</taxonomicName>
is linked to the carapace and pleon strengthening:
<taxonomicName id="4C104D4F2252B661FDF93241FD08BF41" authorityName="Vereshchaka, Kulagin &amp; Lunina" authorityYear="2021" box="[549,651,582,606]" class="Malacostraca" family="Acanthephyridae" genus="Sclerodora" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FDF93241FD08BF41" box="[549,651,582,606]" italics="true" pageId="14" pageNumber="350">Sclerodora</emphasis>
</taxonomicName>
has a firm integument, whereas the integument of
<taxonomicName id="4C104D4F2252B661FDC73261FD19BF61" box="[539,666,614,638]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FDC73261FD19BF61" box="[539,666,614,638]" italics="true" pageId="14" pageNumber="350">Hymenodora</emphasis>
</taxonomicName>
is membranous, with the characteristic reticulum of carinae and sulci making the carapace more rigid. Similar, but not homologous, compensatory strengthening of the carapace (a possible response to increasing carapace loads) has been observed in other
<taxonomicName id="4C104D4F2252B661FFAD3305FE92BE05" authorityName="Dana" authorityYear="1852" box="[113,273,770,794]" pageId="14" pageNumber="346" rank="superFamily" superFamily="Oplophoroidea">Oplophoroidea</taxonomicName>
, such as
<taxonomicName id="4C104D4F2252B661FEB43305FE6ABE05" authorityName="S. I. Smith" authorityYear="1882" box="[360,489,770,794]" class="Malacostraca" family="Oplophoridae" genus="Meningodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FEB43305FE6ABE05" box="[360,489,770,794]" italics="true" pageId="14" pageNumber="350">Meningodora</emphasis>
</taxonomicName>
and
<taxonomicName id="4C104D4F2252B661FDC53304FF4DBE25" authority="(Lunina et al. 2021)" baseAuthorityName="Lunina" baseAuthorityYear="2021" class="Malacostraca" family="Oplophoridae" genus="Notostomus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FDC53304FD0FBE05" box="[537,652,771,794]" italics="true" pageId="14" pageNumber="350">Notostomus</emphasis>
(
<bibRefCitation id="EF814B3D2252B661FD473305FF41BE25" author="Lunina AA &amp; Kulagin DN &amp; Vereshchaka AL" pageId="14" pageNumber="350" pagination="1002 - 19" refId="ref11203" refString="Lunina AA, Kulagin DN, Vereshchaka AL. Phylogenetic revision of the shrimp genera Ephyrina, Meningodora and Notostomus (Acanthephyridae: Caridea). Zoological Journal of the Linnean Society 2021; 193: 1002 - 19." type="journal article" year="2021">
Lunina
<emphasis id="B964EADE2252B661FD353304FF09BE25" italics="true" pageId="14" pageNumber="350">et al.</emphasis>
2021
</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BAF36CC2252B661FF513346FDE5B94B" blockId="14.[113,763,144,1859]" pageId="14" pageNumber="350">
The clade
<taxonomicName id="4C104D4F2252B661FED53346FE30BE46" authorityName="Lunina &amp; Kulagin &amp; Vereshchaka" authorityYear="2024" box="[265,435,833,857]" family="Hymenodoridae" pageId="14" pageNumber="349" rank="family">Hymenodoridae</taxonomicName>
also shows a morphological trait linked to the telson, a spinose endpiece in the
<taxonomicName id="4C104D4F2252B661FD3D3366FF7CBE87" authorityName="Wasmer" authorityYear="1972" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="acanthitelsonis">
<emphasis id="B964EADE2252B661FD3D3366FF7CBE87" italics="true" pageId="14" pageNumber="350">H. acanthitelsonis</emphasis>
</taxonomicName>
+
<taxonomicName id="4C104D4F2252B661FEC03386FEF4BE87" authorityName="Wasmer" authorityYear="2022" box="[284,375,896,920]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="chacei">
<emphasis id="B964EADE2252B661FEC03386FEF4BE87" box="[284,375,896,920]" italics="true" pageId="14" pageNumber="350">H. chacei</emphasis>
</taxonomicName>
clade; this is a homoplasy found also in the clade
<taxonomicName id="4C104D4F2252B661FF373398FEFFBEA8" authorityName="Dana" authorityYear="1852" box="[235,380,927,951]" class="Malacostraca" family="Oplophoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="family">Oplophoridae</taxonomicName>
(
<taxonomicName id="4C104D4F2252B661FE513398FE7ABEA8" authorityName="Spence Bate" authorityYear="1888" box="[397,505,927,951]" class="Malacostraca" family="Oplophoridae" genus="Systellaspis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FE513398FE7ABEA8" box="[397,505,927,951]" italics="true" pageId="14" pageNumber="350">Systellaspis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C104D4F2252B661FDF53398FDF8BEA8" authorityName="Chace" authorityYear="1986" box="[553,635,927,951]" class="Malacostraca" family="Oplophoridae" genus="Janicella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FDF53398FDF8BEA8" box="[553,635,927,951]" italics="true" pageId="14" pageNumber="350">Janicella</emphasis>
</taxonomicName>
). All known specimens of
<taxonomicName id="4C104D4F2252B661FF2733B8FE24BEC9" authorityName="Wasmer" authorityYear="1972" box="[251,423,958,982]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="acanthitelsonis">
<emphasis id="B964EADE2252B661FF2733B8FE24BEC9" box="[251,423,958,982]" italics="true" pageId="14" pageNumber="350">H. acanthitelsonis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C104D4F2252B661FE0A33B8FDACBEC9" authorityName="Wasmer" authorityYear="2022" box="[470,559,958,982]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="chacei">
<emphasis id="B964EADE2252B661FE0A33B8FDACBEC9" box="[470,559,958,982]" italics="true" pageId="14" pageNumber="350">H. chacei</emphasis>
</taxonomicName>
have been collected during near-bottom hauls and are probably benthopelagic. Given that
<taxonomicName id="4C104D4F2252B661FF3833FAFEB5B90A" authorityName="Chace" authorityYear="1986" box="[228,310,1021,1045]" class="Malacostraca" family="Oplophoridae" genus="Janicella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FF3833FAFEB5B90A" box="[228,310,1021,1045]" italics="true" pageId="14" pageNumber="350">Janicella</emphasis>
</taxonomicName>
and most
<taxonomicName id="4C104D4F2252B661FE7F33FAFD8CB90A" authorityName="Spence Bate" authorityYear="1888" box="[419,527,1021,1045]" class="Malacostraca" family="Oplophoridae" genus="Systellaspis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FE7F33FAFD8CB90A" box="[419,527,1021,1045]" italics="true" pageId="14" pageNumber="350">Systellaspis</emphasis>
</taxonomicName>
are also benthopelagic (
<bibRefCitation id="EF814B3D2252B661FFA0341BFEB4B92B" author="Vereshchaka AL" box="[124,311,1052,1076]" pageId="14" pageNumber="350" pagination="1639 - 68" refId="ref11557" refString="Vereshchaka AL. Macroplankton in the near-bottom layer of continental slopes and seamounts. Deep Sea Research Part I 1995; 42: 1639 - 68." type="journal article" year="1995">Vereshchaka 1995</bibRefCitation>
), we suggest that the spatulate endpiece on the telson might serve in bedding into sediments.
</paragraph>
<paragraph id="8BAF36CC2252B661FF51345CFDB4BB15" blockId="14.[113,763,144,1859]" pageId="14" pageNumber="350">
Morphological traits within the pelagic clades
<taxonomicName id="4C104D4F2252B661FD87345BFD45B96C" authorityName="Rathbun" authorityYear="1902" box="[603,710,1115,1139]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="frontalis">
<emphasis id="B964EADE2252B661FD87345BFD45B96C" box="[603,710,1115,1139]" italics="true" pageId="14" pageNumber="350">H. frontalis</emphasis>
</taxonomicName>
+
<taxonomicName id="4C104D4F2252B661FD3E345BFF30B98D" authorityName="Smith" authorityYear="1886" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="gracilis">
<emphasis id="B964EADE2252B661FD3E345BFF30B98D" italics="true" pageId="14" pageNumber="350">H. gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C104D4F2252B661FF38347CFECEB98D" authorityName="Buchholz" authorityYear="1874" box="[228,333,1146,1170]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="glacialis">
<emphasis id="B964EADE2252B661FF38347CFECEB98D" box="[228,333,1146,1170]" italics="true" pageId="14" pageNumber="350">H. glacialis</emphasis>
</taxonomicName>
are linked to further strengthening of their membranous carapace (an additional crescent-shaped sulcus in the
<taxonomicName id="4C104D4F2252B661FF4434BDFE81B9CE" authorityName="Buchholz" authorityYear="1874" box="[152,258,1209,1233]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="glacialis">
<emphasis id="B964EADE2252B661FF4434BDFE81B9CE" box="[152,258,1209,1233]" italics="true" pageId="14" pageNumber="350">H. glacialis</emphasis>
</taxonomicName>
clade) and to various strategies of passive defence (number of pre- and postorbital rostral teeth) and escape behaviour (proportions of the carapace and rostrum). In fact, pelagic decapods may use the rostrum as a passive defensive structure (long, acute and spiny) and/or as a rudder during backwards escape flips (active protection) (
<bibRefCitation id="EF814B3D2252B661FEE03551FE6DB871" author="Lunina AA &amp; Kulagin DN &amp; Vereshchaka AL" box="[316,494,1366,1390]" pageId="14" pageNumber="350" pagination="213 - 32" refId="ref11158" refString="Lunina AA, Kulagin DN, Vereshchaka AL. Oplophoridae (Decapoda: Crustacea): phylogeny, taxonomy and evolution studied by a combination of morphological and molecular methods. Zoological Journal of the Linnean Society 2019; 186: 213 - 32." type="journal article" year="2019">
Lunina
<emphasis id="B964EADE2252B661FE553551FE34B871" box="[393,439,1366,1390]" italics="true" pageId="14" pageNumber="350">et al.</emphasis>
2019
</bibRefCitation>
). Our PCAs (
<figureCitation id="132B2A492252B661FDA23551FD36B871" box="[638,693,1366,1390]" captionStart="Figure 6" captionStartId="10.[113,178,1411,1435]" captionTargetBox="[210,1364,147,1381]" captionTargetId="figure-177@10.[209,1364,145,1383]" captionTargetPageId="10" captionText="Figure 6. Principal component analysis of seven morphological characters of the clade Hymenodoridae. A, molecular clades within Hymenodora glacialis and Hymenodora gracilis combined (analysis 6). B, molecular clades within H. glacialis and H.gracilis analysed separately (analysis 7). The x- and y-axes are eigenvalue scales; the morphometric dataset is presented in the Supporting Information (Table S5); only four most significant factors are shown." figureDoi="http://doi.org/10.5281/zenodo.11240010" httpUri="https://zenodo.org/record/11240010/files/figure.png" pageId="14" pageNumber="350">Fig. 6</figureCitation>
) show that the rostral angle and the numbers of post- and preorbital teeth are three major factors explaining morphological diversification within
<taxonomicName id="4C104D4F2252B661FF6435B3FEB7B8D3" box="[184,308,1460,1484]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="genus">
<emphasis id="B964EADE2252B661FF6435B3FEB7B8D3" box="[184,308,1460,1484]" italics="true" pageId="14" pageNumber="350">Hymenodora</emphasis>
</taxonomicName>
. With the exception of
<taxonomicName id="4C104D4F2252B661FDFC35B3FD08B8D3" authorityName="Rathbun" authorityYear="1902" box="[544,651,1460,1484]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="frontalis">
<emphasis id="B964EADE2252B661FDFC35B3FD08B8D3" box="[544,651,1460,1484]" italics="true" pageId="14" pageNumber="350">H. frontalis</emphasis>
</taxonomicName>
, which has an unusually long rostrum, these morphological modifications can be recorded using statistical analyses only.
</paragraph>
<paragraph id="8BAF36CC2252B661FF513615FDF4BA5C" blockId="14.[113,763,144,1859]" pageId="14" pageNumber="350">
Overall, in contrast to their unusually high molecular divergence,
<taxonomicName id="4C104D4F2252B661FF613636FEE4BB56" authorityName="Lunina &amp; Kulagin &amp; Vereshchaka" authorityYear="2024" box="[189,359,1585,1609]" family="Hymenodoridae" pageId="14" pageNumber="349" rank="family">Hymenodoridae</taxonomicName>
show very low morphological diversity, which is likely linked to a homogeneous deep-sea habitat. Remarkable traits can be found only in the likely benthopelagic species,
<taxonomicName id="4C104D4F2252B661FF153697FEFABBB8" authorityName="Wasmer" authorityYear="1972" box="[201,377,1679,1703]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="acanthitelsonis">
<emphasis id="B964EADE2252B661FF153697FEFABBB8" box="[201,377,1679,1703]" italics="true" pageId="14" pageNumber="350">H. acanthitelsonis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C104D4F2252B661FE6E3697FD93BBB8" authorityName="Wasmer" authorityYear="2022" box="[434,528,1679,1703]" class="Malacostraca" family="Oplophoridae" genus="Hymenodora" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="14" pageNumber="350" phylum="Arthropoda" rank="species" species="chacei">
<emphasis id="B964EADE2252B661FE6E3697FD93BBB8" box="[434,528,1679,1703]" italics="true" pageId="14" pageNumber="350">H. chacei</emphasis>
</taxonomicName>
, whereas evolution of definitely pelagic clades is coupled mainly with feeding specialization (mouthparts), strengthening of the membranous carapace (carinae and sulci) or the proportions of the body. The last of these is the only group of characters that mirrors the cryptic speciation found here through molecular methods.
</paragraph>
</subSubSection>
</treatment>
</document>