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<mods:title id="AA4D1EBCAC7F65BD0F7E4B377CA40BDB">New light on the nomenclature, taxonomy, and biology of Hemipholis species (Echinodermata: Ophiuroidea: Ophiactidae)</mods:title>
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<mods:namePart id="147B1B53B9F3E0E9DA3AA72DD2B58365">Hendler, Gordon</mods:namePart>
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<mods:date id="692F18439E65300A2CB0C44FF147396A">2011</mods:date>
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<treatment id="03A5BB74126ED2794C925768FD34FE26" ID-DOI="http://doi.org/10.5281/zenodo.6194953" ID-GBIF-Taxon="119418220" ID-Zenodo-Dep="6194953" LSID="urn:lsid:plazi:treatment:03A5BB74126ED2794C925768FD34FE26" httpUri="http://treatment.plazi.org/id/03A5BB74126ED2794C925768FD34FE26" lastPageId="5" lastPageNumber="48" pageId="4" pageNumber="48">
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<paragraph id="8BB30A62126ED2784C925768FD56F903" blockId="4.[151,689,1647,1673]" box="[151,689,1647,1673]" pageId="4" pageNumber="48">
<heading id="D0FBBD0E126ED2784C925768FD56F903" bold="true" box="[151,689,1647,1673]" fontSize="11" level="1" pageId="4" pageNumber="48" reason="1">
<emphasis id="B978D670126ED2784C925768FD56F903" bold="true" box="[151,689,1647,1673]" pageId="4" pageNumber="48">
Distinguishing
<taxonomicName id="4C0C71E1126ED2784D565768FE00F903" box="[339,487,1647,1673]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="cordifera">
<emphasis id="B978D670126ED2784D565768FE00F903" bold="true" box="[339,487,1647,1673]" italics="true" pageId="4" pageNumber="48">H. cordifera</emphasis>
</taxonomicName>
from
<taxonomicName id="4C0C71E1126ED2784E365768FD56F903" box="[563,689,1647,1673]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="gracilis">
<emphasis id="B978D670126ED2784E365768FD56F903" bold="true" box="[563,689,1647,1673]" italics="true" pageId="4" pageNumber="48">H. gracilis</emphasis>
</taxonomicName>
</emphasis>
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</paragraph>
<paragraph id="8BB30A62126ED2784C9257B1FA91F840" blockId="4.[151,1436,1718,2030]" pageId="4" pageNumber="48">
Small individuals of
<taxonomicName id="4C0C71E1126ED2784D8D57BFFDE5F944" box="[392,514,1718,1743]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="gracilis">
<emphasis id="B978D670126ED2784D8D57BFFDE5F944" box="[392,514,1718,1743]" italics="true" pageId="4" pageNumber="48">H. gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0C71E1126ED2784E3857BFFD2EF944" box="[573,713,1718,1743]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="cordifera">
<emphasis id="B978D670126ED2784E3857BFFD2EF944" box="[573,713,1718,1743]" italics="true" pageId="4" pageNumber="48">H. cordifera</emphasis>
</taxonomicName>
look alike, which could account for a record of “
<taxonomicName id="4C0C71E1126ED278490C57BFFA68F944" box="[1289,1423,1718,1743]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="elongata">
<emphasis id="B978D670126ED278490C57BFFA68F944" box="[1289,1423,1718,1743]" italics="true" pageId="4" pageNumber="48">H. elongata</emphasis>
</taxonomicName>
” from the Eastern Pacific (in
<bibRefCitation id="EF9D7793126ED2784DD757DDFD5CF978" author="Alvarado" box="[466,699,1754,1778]" pageId="4" pageNumber="48" refString="Alvarado, J. J., Solis-Marin, F. A. &amp; Ahearn, C. G. (2010) Echinoderm (Echinodermata) diversity in the Pacific Coast of Central America. Marine Biodiversity, 40, 45 - 56." type="journal article" year="2010">
Alvarado
<emphasis id="B978D670126ED2784E4557DCFD96F978" box="[576,625,1754,1778]" italics="true" pageId="4" pageNumber="48">et al</emphasis>
. 2010
</bibRefCitation>
). Although large individuals of both species also are quite similar, several external features can be used to distinguish them (
<figureCitation id="133716E7126ED2784F1E57F9FC8FF89C" box="[795,872,1790,1814]" captionStart="FIGURE 1" captionStartId="4.[151,250,1402,1425]" captionTargetBox="[151,1436,303,1378]" captionTargetId="figure@4.[151,1436,303,1378]" captionTargetPageId="4" captionText="FIGURE 1. Hemipholis cordifera and Hemipholis gracilis. A C, H. cordifera, LACM 1999 - 26.2, 7.8 mm disk diameter, from South Carolina; D F, H. gracilis, LACM 1997 - 224.001, 7.8 mm disk diameter, from Panama; specimens preserved in ethanol. For each specimen, shown from left to right are: portion of disk in dorsal view, arrowheads indicating pairs of radial shields; basal arm joints on ventral side of disk, arrow indicating second ventral arm plate; basal arm joints in dorsal view, arrowheads indicating middle arm spines. Dorsal arm plates of both species are sometimes fragmented as is the case in Figure C. Scale bars = 1.0 mm." httpUri="https://zenodo.org/record/205691/files/figure.png" pageId="4" pageNumber="48">Figs. 1</figureCitation>
,
<figureCitation id="133716E7126ED2784F7357F9FC61F89C" box="[886,902,1790,1814]" captionStart="FIGURE 2" captionStartId="5.[151,250,1307,1330]" captionTargetBox="[194,1391,487,1283]" captionTargetId="figure@5.[191,1395,483,1286]" captionTargetPageId="5" captionText="FIGURE 2. Scatter plot for Hemipholis cordifera and Hemipholis gracilis, showing the relationship between the body size in terms of disk diameter, and shape of the second ventral arm plate characterized by the ratio between the ossicles width and length." httpUri="https://zenodo.org/record/205692/files/figure.png" pageId="4" pageNumber="48">2</figureCitation>
). Large individuals of
<taxonomicName id="4C0C71E1126ED278488657F9FA87F89C" box="[1155,1376,1790,1814]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="gracilis">
<emphasis id="B978D670126ED278488657F9FA87F89C" box="[1155,1376,1790,1814]" italics="true" pageId="4" pageNumber="48">Hemipholis gracilis</emphasis>
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have radial shields that are more constricted proximally than the shields of
<taxonomicName id="4C0C71E1126ED2784FB75624FBDBF8B0" box="[946,1084,1826,1850]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="cordifera">
<emphasis id="B978D670126ED2784FB75624FBDBF8B0" box="[946,1084,1826,1850]" italics="true" pageId="4" pageNumber="48">H. cordifera</emphasis>
</taxonomicName>
(
<figureCitation id="133716E7126ED27848485625FB6AF8B0" box="[1101,1165,1826,1850]" captionStart="FIGURE 1" captionStartId="4.[151,250,1402,1425]" captionTargetBox="[151,1436,303,1378]" captionTargetId="figure@4.[151,1436,303,1378]" captionTargetPageId="4" captionText="FIGURE 1. Hemipholis cordifera and Hemipholis gracilis. A C, H. cordifera, LACM 1999 - 26.2, 7.8 mm disk diameter, from South Carolina; D F, H. gracilis, LACM 1997 - 224.001, 7.8 mm disk diameter, from Panama; specimens preserved in ethanol. For each specimen, shown from left to right are: portion of disk in dorsal view, arrowheads indicating pairs of radial shields; basal arm joints on ventral side of disk, arrow indicating second ventral arm plate; basal arm joints in dorsal view, arrowheads indicating middle arm spines. Dorsal arm plates of both species are sometimes fragmented as is the case in Figure C. Scale bars = 1.0 mm." httpUri="https://zenodo.org/record/205691/files/figure.png" pageId="4" pageNumber="48">Fig. 1</figureCitation>
A,D). The basal ventral arm plates of
<taxonomicName id="4C0C71E1126ED2784D2A564FFE40F8D4" box="[303,423,1862,1887]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="gracilis">
<emphasis id="B978D670126ED2784D2A564FFE40F8D4" box="[303,423,1862,1887]" italics="true" pageId="4" pageNumber="48">H. gracilis</emphasis>
</taxonomicName>
, particularly the second ventral arm plate, are relatively shorter and wider than those of
<taxonomicName id="4C0C71E1126ED2784984564FFF19F808" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="cordifera">
<emphasis id="B978D670126ED2784984564FFF19F808" italics="true" pageId="4" pageNumber="48">H. cordifera</emphasis>
</taxonomicName>
(
<figureCitation id="133716E7126ED2784D08566DFEB0F808" box="[269,343,1898,1922]" captionStart="FIGURE 1" captionStartId="4.[151,250,1402,1425]" captionTargetBox="[151,1436,303,1378]" captionTargetId="figure@4.[151,1436,303,1378]" captionTargetPageId="4" captionText="FIGURE 1. Hemipholis cordifera and Hemipholis gracilis. A C, H. cordifera, LACM 1999 - 26.2, 7.8 mm disk diameter, from South Carolina; D F, H. gracilis, LACM 1997 - 224.001, 7.8 mm disk diameter, from Panama; specimens preserved in ethanol. For each specimen, shown from left to right are: portion of disk in dorsal view, arrowheads indicating pairs of radial shields; basal arm joints on ventral side of disk, arrow indicating second ventral arm plate; basal arm joints in dorsal view, arrowheads indicating middle arm spines. Dorsal arm plates of both species are sometimes fragmented as is the case in Figure C. Scale bars = 1.0 mm." httpUri="https://zenodo.org/record/205691/files/figure.png" pageId="4" pageNumber="48">Figs. 1</figureCitation>
B,E &amp; 2). Although there is an overlap in the width/length ratios of the second ventral arm plates of both species, animals larger than
<quantity id="4CF4A787126ED2784E2A5688FD97F82C" box="[559,624,1935,1958]" metricMagnitude="-3" metricUnit="m" metricValue="7.0" pageId="4" pageNumber="48" unit="mm" value="7.0">7 mm</quantity>
disk diameter have plates that distinctly differ in their proportions (
<figureCitation id="133716E7126ED278496B5689FF4FF840" captionStart="FIGURE 2" captionStartId="5.[151,250,1307,1330]" captionTargetBox="[194,1391,487,1283]" captionTargetId="figure@5.[191,1395,483,1286]" captionTargetPageId="5" captionText="FIGURE 2. Scatter plot for Hemipholis cordifera and Hemipholis gracilis, showing the relationship between the body size in terms of disk diameter, and shape of the second ventral arm plate characterized by the ratio between the ossicles width and length." httpUri="https://zenodo.org/record/205692/files/figure.png" pageId="4" pageNumber="48">Fig. 2</figureCitation>
). In addition, the arm spines of large
<taxonomicName id="4C0C71E1126ED2784E4F56B3FD25F841" box="[586,706,1971,1995]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="gracilis">
<emphasis id="B978D670126ED2784E4F56B3FD25F841" box="[586,706,1971,1995]" italics="true" pageId="4" pageNumber="48">H. gracilis</emphasis>
</taxonomicName>
tend to be more tapered than those of
<taxonomicName id="4C0C71E1126ED278486A56B3FB1EF841" box="[1135,1273,1971,1995]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="species" species="cordifera">
<emphasis id="B978D670126ED278486A56B3FB1EF841" box="[1135,1273,1971,1995]" italics="true" pageId="4" pageNumber="48">H. cordifera</emphasis>
</taxonomicName>
(Fig. C,F).
</paragraph>
<paragraph id="8BB30A62126ED2794CC256D1FD34FE26" blockId="4.[151,1436,1718,2030]" lastBlockId="5.[151,1436,151,428]" lastPageId="5" lastPageNumber="49" pageId="4" pageNumber="48">
The Pacific and Atlantic populations of
<taxonomicName id="4C0C71E1126ED2784E8156D1FCE2F864" box="[644,773,2006,2030]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="4" pageNumber="48" phylum="Echinodermata" rank="genus">
<emphasis id="B978D670126ED2784E8156D1FCE2F864" box="[644,773,2006,2030]" italics="true" pageId="4" pageNumber="48">Hemipholis</emphasis>
</taxonomicName>
are little differentiated morphologically, and dissimilarities in their external structures only arise late in ontogenesis. This is not surprising since geminate species separated by the Panamanian Isthmus are typically “in the initial, allopatric stage of speciation,” and they can even be reproductively compatible (
<bibRefCitation id="EF9D7793126FD2794D6251D8FDC3FF7D" author="Lessios" box="[359,548,223,247]" pageId="5" pageNumber="49" refString="Lessios, H. A. (2008) The Great American Schism: Divergence of marine organisms after the rise of the Central American Isthmus. Annual Review of Ecology, Evolution, and Systematics, 39, 63 - 91." type="journal article" year="2008">Lessios 2008:84</bibRefCitation>
). Consistent differences between individuals of
<taxonomicName id="4C0C71E1126FD279483F51E7FB5CFF72" box="[1082,1211,224,248]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="5" pageNumber="49" phylum="Echinodermata" rank="genus">
<emphasis id="B978D670126FD279483F51E7FB5CFF72" box="[1082,1211,224,248]" italics="true" pageId="5" pageNumber="49">Hemipholis</emphasis>
</taxonomicName>
from opposite sides of the Isthmian barrier, similarities among individuals throughout their range in each ocean, and the species broad ranges in the western Atlantic and Eastern Pacific respectively, all suggest that differences between
<taxonomicName id="4C0C71E1126FD2794915502EFA7CFECA" box="[1296,1435,296,320]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="5" pageNumber="49" phylum="Echinodermata" rank="species" species="cordifera">
<emphasis id="B978D670126FD2794915502EFA7CFECA" box="[1296,1435,296,320]" italics="true" pageId="5" pageNumber="49">H. cordifera</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0C71E1126FD2794CC3504AFED8FEEE" box="[198,319,332,356]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="5" pageNumber="49" phylum="Echinodermata" rank="species" species="gracilis">
<emphasis id="B978D670126FD2794CC3504AFED8FEEE" box="[198,319,332,356]" italics="true" pageId="5" pageNumber="49">H. gracilis</emphasis>
</taxonomicName>
are not merely ecophenotypic. However, there is a dearth of comparable data on genotypic, phenotypic, and phenological characteristics of populations from both oceans, which are needed to elucidate the evolution and phylogeny of
<taxonomicName id="4C0C71E1126FD2794D915092FDEBFE26" box="[404,524,404,428]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="5" pageNumber="49" phylum="Echinodermata" rank="species" species="gracilis">
<emphasis id="B978D670126FD2794D915092FDEBFE26" box="[404,524,404,428]" italics="true" pageId="5" pageNumber="49">H. gracilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0C71E1126FD2794E465092FD2BFE26" box="[579,716,404,428]" class="Ophiuroidea" family="Ophiactidae" genus="Hemipholis" kingdom="Animalia" order="Ophiurida" pageId="5" pageNumber="49" phylum="Echinodermata" rank="species" species="cordifera">
<emphasis id="B978D670126FD2794E465092FD2BFE26" box="[579,716,404,428]" italics="true" pageId="5" pageNumber="49">H. cordifera</emphasis>
</taxonomicName>
.
</paragraph>
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