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<mods:title id="F0B4FBDC01D0757F2E32B70140A1194D">Understanding the color variability and resolving taxonomic confusion in the sea cucumber Isostichopus badionotus (Echinodermata, Holothuroidea): a revision of the genus Isostichopus</mods:title>
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<mods:affiliation id="0E9EA8B45FF45A868BC7FAA203A0FBBF">Santa Marta, Magdalena, Colombia. & Instituto de Investigaciones Marinas y Costeras - Invemar, Calle 25 No. 2 - 55, Playa Salguero, Rodadero, Santa Marta, Colombia. & Smithsonian Tropical Research Institute - STRI, BOX 0843 - 03092, Balboa, Panama.</mods:affiliation>
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<mods:affiliation id="B835675554E8F24B04391AD72E3A4139">Santa Marta, Magdalena, Colombia. & Instituto de Investigaciones Marinas y Costeras - Invemar, Calle 25 No. 2 - 55, Playa Salguero, Rodadero, Santa Marta, Colombia. & Smithsonian Tropical Research Institute - STRI, BOX 0843 - 03092, Balboa, Panama.</mods:affiliation>
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<mods:namePart id="E6BD7C7121FC39B82721E4714E9A40CF">Solís-Marín, Francisco A.</mods:namePart>
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<mods:affiliation id="6B5DE2BED7279CD88DE00ABC4A29CCC4">Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, 04510, Ciudad de Mexico, México.</mods:affiliation>
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<mods:affiliation id="CDA685CF4DBB961A61D47F7D757235EF">Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, 04510, Ciudad de Mexico, México.</mods:affiliation>
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<mods:namePart id="4A37EF70492F519602B69E765718F012">Lessios, Harilaos</mods:namePart>
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@ -97,9 +98,9 @@ species
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Medium to large species, length to
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, body wall firm and thick. Highly variable coloration (Fig. 1). Body convex to quadrangular in cross-section. Large lateral papillae, usually sharply defining dorsum and ventrum (or bivium and trivium). Dorsal papillae irregularly arranged and in different sizes and shapes: wart-like, spiky, or with small points. Sole well developed, with three bands of cylindrical podia. Single stone canal attached to the mesentery; calcareous ring with massive radial pieces with posterior projections of different sizes, and narrow interradial pieces with projections pointed anteriorly and concave posterior margins (
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<figureCitation id="E8A5B8E2B715323FFD8FFCBBB8C029D7" box="[568,642,876,903]" captionStart="Fig" captionStartId="23.[189,232,1822,1848]" captionTargetBox="[239,1387,770,1754]" captionTargetId="figure-34@23.[189,1398,759,1770]" captionTargetPageId="23" captionText="Fig. 4. Comparison of the calcareous rings of species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867). B. I. maculatus phoenius (Clark, 1922) (A–B showing calcareous rings from specimens of different sizes). C. I. maculatus maculatus (Greeff, 1882). D. I. fuscus (Ludwig, 1875). E. I. macroparentheses (Clark, 1922). Dimensions of the individuals from which each ring was obtained are shown next to their museum catalog number. Abbreviations: ip = interradial plate; rp = radial plate. Scale bar = 4 mm." pageId="16" pageNumber="17">Fig. 4</figureCitation>
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<figureCitation id="E8A5B8E2B715323FFD8FFCBBB8C029D7" box="[568,642,876,903]" captionStart="Fig" captionStartId="23.[189,232,1822,1848]" captionTargetBox="[239,1387,770,1754]" captionTargetId="figure-34@23.[189,1398,759,1770]" captionTargetPageId="23" captionText="Fig. 4. Comparison of the calcareous rings of species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867). B. I. maculatus phoenius (Clark, 1922) (A–B showing calcareous rings from specimens of different sizes). C. I. maculatus maculatus (Greeff, 1882). D. I. fuscus (Ludwig, 1875). E. I. macroparentheses (Clark, 1922). Dimensions of the individuals from which each ring was obtained are shown next to their museum catalog number. Abbreviations: ip = interradial plate; rp = radial plate. Scale bar = 4 mm." figureDoi="http://doi.org/10.5281/zenodo.13710647" httpUri="https://zenodo.org/record/13710647/files/figure.png" pageId="16" pageNumber="17">Fig. 4</figureCitation>
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). Ossicles: dorsal papillae with tables; thin C- or S-shaped rods in varying numbers and sizes; large, curved rods with quadrangular projections, sometimes perforated in the middle (Fig. 2S); few perforated plates in the tip (Fig. 2U). Tables in a dense layer, low, squat, with a circular disc with 8 to 12 small holes, four pillars, single crossbeam, and a wreath of small spines at the crown (Fig. 2A–B); tables with reduced disc (Fig. 2C), additional holes in more than one ring (
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<figureCitation id="E8A5B8E2B715323FFF7FFBC3BB742E7F" box="[200,310,1044,1071]" captionStart="Fig" captionStartId="67.[189,232,1787,1813]" captionTargetBox="[224,1363,265,1739]" captionTargetId="figure-12@67.[224,1363,265,1739]" captionTargetPageId="67" captionText="Fig. 20. Isostichopus fuscus (Ludwig, 1875) ossicles (specimen USNM 1682794-Ta213, 155 mm long). A. Thin C-shaped rods, tables, perforated plates and large, curved rods from dorsal body wall and papillae. B. Thin C-shaped rods, tables, perforated plates, large, curved rods and end plate (fragment) from ventral body wall and tube feet. C. Rods and tables from tentacles. D. Thin C-shaped rods from longitudinal muscles. E. Rods bifurcated from the cloaca. F. Large tables with a circular spire well developed in the anterior cloaca. G. Tables from respiratory trees. H. Rods in cross shape, simple rods, and C-shaped rods in the intestine. Photos by G. Borrero." pageId="16" pageNumber="17">Fig. 20A</figureCitation>
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<figureCitation id="E8A5B8E2B715323FFF7FFBC3BB742E7F" box="[200,310,1044,1071]" captionStart="Fig" captionStartId="67.[189,232,1787,1813]" captionTargetBox="[224,1363,265,1739]" captionTargetId="figure-12@67.[224,1363,265,1739]" captionTargetPageId="67" captionText="Fig. 20. Isostichopus fuscus (Ludwig, 1875) ossicles (specimen USNM 1682794-Ta213, 155 mm long). A. Thin C-shaped rods, tables, perforated plates and large, curved rods from dorsal body wall and papillae. B. Thin C-shaped rods, tables, perforated plates, large, curved rods and end plate (fragment) from ventral body wall and tube feet. C. Rods and tables from tentacles. D. Thin C-shaped rods from longitudinal muscles. E. Rods bifurcated from the cloaca. F. Large tables with a circular spire well developed in the anterior cloaca. G. Tables from respiratory trees. H. Rods in cross shape, simple rods, and C-shaped rods in the intestine. Photos by G. Borrero." figureDoi="http://doi.org/10.5281/zenodo.13710683" httpUri="https://zenodo.org/record/13710683/files/figure.png" pageId="16" pageNumber="17">Fig. 20A</figureCitation>
|
||||
), or larger and modified discs also present (Fig. 2D), mainly at the top of papillae. Dorsal body wall with tables and a few thin C- or S-shaped rods only. Ossicles in dorsal papillae and body wall change during growth, though drastic ontogenetic changes occurring only in
|
||||
<taxonomicName id="B79EDFE4B715323FFB35FB8FBF2D2E22" ID-CoL="6MZDT" baseAuthorityName="Clark" baseAuthorityYear="1922" box="[1154,1391,1112,1138]" class="Holothuroidea" family="Stichopodidae" genus="Isostichopus" kingdom="Animalia" order="Synallactida" pageId="16" pageNumber="17" phylum="Echinodermata" rank="species" species="macroparentheses">
|
||||
<emphasis id="42EA7875B715323FFB35FB8FBF2D2E22" box="[1154,1391,1112,1138]" italics="true" pageId="16" pageNumber="17">I. macroparentheses</emphasis>
|
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|
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@ -509,7 +510,7 @@ tables (Figs 2D, 13A)
|
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</paragraph>
|
||||
<paragraph id="7021A467B716323BFF0AFEDEB9512BFA" blockId="19.[188,1400,265,627]" lastBlockId="20.[189,1401,265,426]" lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20">
|
||||
plates with numerous holes (Fig. 2V); slightly or strongly curved rods with broad perforated expansions in the middle (Fig. 2T); both plates and rods larger than those of dorsal papillae. Ventral body wall with only tables and a few C-shaped ossicles. Tentacles with strongly or slightly curved spiny rods in varying sizes (Fig. 2O) and small tables as those of the body wall, or modified with low and incomplete spires (Fig. 2F). Mouth membrane with thin C-shaped rods, simple rods, and large tables, not documented previously for the genus, with well-developed spire, composed of at least ten pillars joining at the top, forming very dense and thick crown of spines, without crossbeams; discs of the same width as the spire, or wider with several rings of holes and several central perforations (Figs 2H, 7D, 11, 16D). Longitudinal muscles containing C-shaped rods and simple rods. Posterior part of the cloaca with C-shaped, simple or bifurcated rods (
|
||||
<figureCitation id="E8A5B8E2B716323CFE28FDE0BBB72801" box="[415,501,567,593]" captionStart="Fig" captionStartId="35.[189,232,1681,1707]" captionTargetBox="[195,1386,747,1639]" captionTargetId="figure-167@35.[189,1398,727,1640]" captionTargetPageId="35" captionText="Fig. 11. Light microscope photographs showing ossicles from the mouth membrane, anterior cloaca, posterior cloaca, and gonads of the species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867) (mouth membrane and gonads from the lectotype MCZ HOL-509; anterior and posterior cloaca from USNM 1659457-IbBT116). B. I. maculatus phoenius (Clark, 1922) (all ossicles from the holotype MCZ HOL-1182). C. I. maculatus maculatus (Greeff, 1882) (all ossicles from the neotype USNM E16150; male gonads at the bottom from the neoparatype USNM E16151). D. I. fuscus (Ludwig, 1875) (mouth membrane from MCZ HOL-742; anterior cloaca from MBMLP- If212; posterior cloaca and gonads from MCZ HOL-743). E. I. macroparentheses (Clark, 1922) (mouth membrane, anterior and posterior cloaca from USNM E47524; gonads from the holotype MCZ HOL- 921). Photos by G. Borrero. Scale bars = 100 µm." pageId="19" pageNumber="20">Fig. 11</figureCitation>
|
||||
<figureCitation id="E8A5B8E2B716323CFE28FDE0BBB72801" box="[415,501,567,593]" captionStart="Fig" captionStartId="35.[189,232,1681,1707]" captionTargetBox="[195,1386,747,1639]" captionTargetId="figure-167@35.[189,1398,727,1640]" captionTargetPageId="35" captionText="Fig. 11. Light microscope photographs showing ossicles from the mouth membrane, anterior cloaca, posterior cloaca, and gonads of the species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867) (mouth membrane and gonads from the lectotype MCZ HOL-509; anterior and posterior cloaca from USNM 1659457-IbBT116). B. I. maculatus phoenius (Clark, 1922) (all ossicles from the holotype MCZ HOL-1182). C. I. maculatus maculatus (Greeff, 1882) (all ossicles from the neotype USNM E16150; male gonads at the bottom from the neoparatype USNM E16151). D. I. fuscus (Ludwig, 1875) (mouth membrane from MCZ HOL-742; anterior cloaca from MBMLP- If212; posterior cloaca and gonads from MCZ HOL-743). E. I. macroparentheses (Clark, 1922) (mouth membrane, anterior and posterior cloaca from USNM E47524; gonads from the holotype MCZ HOL- 921). Photos by G. Borrero. Scale bars = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.13710659" httpUri="https://zenodo.org/record/13710659/files/figure.png" pageId="19" pageNumber="20">Fig. 11</figureCitation>
|
||||
); anterior part with simple, branched rods, irregular plate-like branched rods (Fig. 2R) and large tables, with well-developed and very dense and thick spire, some with circular
|
||||
<emphasis id="42EA7875B716323CFF0AF946BB602CFB" bold="true" box="[189,290,1681,1707]" pageId="19" pageNumber="20">Fig. 3A.</emphasis>
|
||||
Bayesian inference tree of concatenated sequences of the mitochondrial COI-Fr1 (Barcoding region), COI-Fr2 and 16S gene fragments. The numbers on the nodes indicate Bayesian posterior probability/Maximum Likelihood (bootstrap %)/Neighbor Joining (bootstrap %). A hyphen (-) indicates that a node was absent in the tree estimated by a particular method. Individuals are indicated by the field number, which includes the initial letter of the locality and the color pattern when available (see
|
||||
|
|
@ -553,7 +554,7 @@ and
|
|||
<bibRefCitation id="140FD996B716323CFB79F819BF312DB8" author="Selenka E." box="[1230,1395,1998,2024]" pageId="19" pageNumber="20" pagination="291 - 374" refId="ref62331" refString="Selenka E. 1867. Beitrage zur Anatomie und Systematik der Holothurien. Zeitschrift fur wissenschaftliche Zoologie 17 (2): 291 - 374. Available from https: // www. biodiversitylibrary. org / page / 45007590 [accessed 16 Apr. 2024]." type="journal article" year="1867">Selenka, 1867</bibRefCitation>
|
||||
</taxonomicName>
|
||||
. spire, without clear crossbeams, wide discs with several rings of holes and several central perforations (
|
||||
<figureCitation id="E8A5B8E2B711323BFF71FEFDBB5C2B15" box="[198,286,298,325]" captionStart="Fig" captionStartId="35.[189,232,1681,1707]" captionTargetBox="[195,1386,747,1639]" captionTargetId="figure-167@35.[189,1398,727,1640]" captionTargetPageId="35" captionText="Fig. 11. Light microscope photographs showing ossicles from the mouth membrane, anterior cloaca, posterior cloaca, and gonads of the species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867) (mouth membrane and gonads from the lectotype MCZ HOL-509; anterior and posterior cloaca from USNM 1659457-IbBT116). B. I. maculatus phoenius (Clark, 1922) (all ossicles from the holotype MCZ HOL-1182). C. I. maculatus maculatus (Greeff, 1882) (all ossicles from the neotype USNM E16150; male gonads at the bottom from the neoparatype USNM E16151). D. I. fuscus (Ludwig, 1875) (mouth membrane from MCZ HOL-742; anterior cloaca from MBMLP- If212; posterior cloaca and gonads from MCZ HOL-743). E. I. macroparentheses (Clark, 1922) (mouth membrane, anterior and posterior cloaca from USNM E47524; gonads from the holotype MCZ HOL- 921). Photos by G. Borrero. Scale bars = 100 µm." pageId="20" pageNumber="21">Fig. 11</figureCitation>
|
||||
<figureCitation id="E8A5B8E2B711323BFF71FEFDBB5C2B15" box="[198,286,298,325]" captionStart="Fig" captionStartId="35.[189,232,1681,1707]" captionTargetBox="[195,1386,747,1639]" captionTargetId="figure-167@35.[189,1398,727,1640]" captionTargetPageId="35" captionText="Fig. 11. Light microscope photographs showing ossicles from the mouth membrane, anterior cloaca, posterior cloaca, and gonads of the species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867) (mouth membrane and gonads from the lectotype MCZ HOL-509; anterior and posterior cloaca from USNM 1659457-IbBT116). B. I. maculatus phoenius (Clark, 1922) (all ossicles from the holotype MCZ HOL-1182). C. I. maculatus maculatus (Greeff, 1882) (all ossicles from the neotype USNM E16150; male gonads at the bottom from the neoparatype USNM E16151). D. I. fuscus (Ludwig, 1875) (mouth membrane from MCZ HOL-742; anterior cloaca from MBMLP- If212; posterior cloaca and gonads from MCZ HOL-743). E. I. macroparentheses (Clark, 1922) (mouth membrane, anterior and posterior cloaca from USNM E47524; gonads from the holotype MCZ HOL- 921). Photos by G. Borrero. Scale bars = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.13710659" httpUri="https://zenodo.org/record/13710659/files/figure.png" pageId="20" pageNumber="21">Fig. 11</figureCitation>
|
||||
). Respiratory trees with small tables as those of the body wall or strongly spinose straight or cross-shaped rods and large tridimensional spheres (Fig. 2X), not documented previously for the genus. Intestine with spinose or smooth ossicles in a cross shape (Fig. 2P). Gonads with delicate and long rods (Figs 2N, 11). Rosettes not present in
|
||||
<taxonomicName id="B79EDFE4B711323BFDCDFE58B94F2BF9" ID-CoL="7P7DL" authorityName="Deichmann" authorityYear="1958" box="[634,781,399,425]" class="Holothuroidea" family="Stichopodidae" genus="Isostichopus" kingdom="Animalia" order="Synallactida" pageId="20" pageNumber="21" phylum="Echinodermata" rank="genus">
|
||||
<emphasis id="42EA7875B711323BFDCDFE58B94F2BF9" box="[634,781,399,425]" italics="true" pageId="20" pageNumber="21">Isostichopus</emphasis>
|
||||
|
|
@ -569,7 +570,7 @@ and
|
|||
</paragraph>
|
||||
<paragraph id="7021A467B711323BFF0AFE28BE2E284A" blockId="20.[189,1132,511,538]" box="[189,1132,511,538]" pageId="20" pageNumber="21">
|
||||
Tropical and subtropical shores of America and the west coast of Africa (
|
||||
<figureCitation id="E8A5B8E2B711323BFBA2FE28BE1F284A" box="[1045,1117,511,538]" captionStart="Fig" captionStartId="24.[189,232,1787,1813]" captionTargetBox="[195,1391,723,1738]" captionTargetId="figure-121@24.[189,1398,715,1744]" captionTargetPageId="24" captionText="Fig. 5. Geographic distribution of species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867) (gray circles), I. maculatus phoenius (Clark, 1922) (red squares) and I. maculatus maculatus (Greeff, 1882) (blue circles). B. I. fuscus (Ludwig, 1875) (green circles) and I. macroparentheses (Clark, 1922) (orange circles). Locations for I. badionotus and I. fuscus were constructed based on observation records in Global Biodiversity Information Facility data and from museum specimens. Locations forI. maculatus phoenius, I. maculatusmaculatus andI. macroparentheses were constructed based only on confirmed localities of collected or preserved museum specimens." pageId="20" pageNumber="21">Fig. 5</figureCitation>
|
||||
<figureCitation id="E8A5B8E2B711323BFBA2FE28BE1F284A" box="[1045,1117,511,538]" captionStart="Fig" captionStartId="24.[189,232,1787,1813]" captionTargetBox="[195,1391,723,1738]" captionTargetId="figure-121@24.[189,1398,715,1744]" captionTargetPageId="24" captionText="Fig. 5. Geographic distribution of species and subspecies of Isostichopus Deichmann, 1958. A. I. badionotus (Selenka, 1867) (gray circles), I. maculatus phoenius (Clark, 1922) (red squares) and I. maculatus maculatus (Greeff, 1882) (blue circles). B. I. fuscus (Ludwig, 1875) (green circles) and I. macroparentheses (Clark, 1922) (orange circles). Locations for I. badionotus and I. fuscus were constructed based on observation records in Global Biodiversity Information Facility data and from museum specimens. Locations forI. maculatus phoenius, I. maculatusmaculatus andI. macroparentheses were constructed based only on confirmed localities of collected or preserved museum specimens." figureDoi="http://doi.org/10.5281/zenodo.13710649" httpUri="https://zenodo.org/record/13710649/files/figure.png" pageId="20" pageNumber="21">Fig. 5</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
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