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<mods:title>A revision of the genus Psammogorgia Verrill, 1868 (Cnidaria, Anthozoa, Octocorallia) in the tropical eastern Pacific Ocean</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Breedy, Odalisca</mods:namePart>
<mods:affiliation>Centro de Investigacion en Estructuras Microscopicas, Centro de Investigacion en Ciencias del Mar y Limnologia, Escuela de Biologia, Universidad de Costa Rica. P. O. Box 11501 - 2060, Universidad de Costa Rica, San Jose, Costa Rica &amp; Smithsonian Tropical Research Institute, P. O. Box 0843 - 03092, Panama, Republic of Panama</mods:affiliation>
<mods:nameIdentifier type="email">odaliscab@gmail.com</mods:nameIdentifier>
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<mods:namePart>Guzman, Hector M.</mods:namePart>
<mods:affiliation>Smithsonian Tropical Research Institute, P. O. Box 0843 - 03092, Panama, Republic of Panama</mods:affiliation>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2020</mods:date>
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<mods:number>961</mods:number>
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<taxonomicName LSID="20BF18E2-B9E2-5819-A1C8-BA5CB43625FE" authority="Verrill, 1868" authorityName="Verrill" authorityYear="1868" class="Anthozoa" family="Plexauridae" genus="Psammogorgia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psammogorgia arbuscula subsp. var. var. pallida" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="variety" species="arbuscula" subSpecies="var." variety="pallida">Psammogorgia arbuscula var. pallida Verrill, 1868</taxonomicName>
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">Figures 7</figureCitation>
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916) A spindles B wart clubs C radiates and crosses D anthocodial spindles." figureDoi="10.3897/zookeys.961.54846.figure8" httpUri="https://binary.pensoft.net/fig/442681" pageId="0" pageNumber="1">, 8</figureCitation>
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Psammogorgia arbuscula var. pallida Verrill, 1868. YPM 1785 A colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure9" httpUri="https://binary.pensoft.net/fig/442682" pageId="0" pageNumber="1">, 9</figureCitation>
</paragraph>
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<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Verrill" authorityYear="1868" class="Anthozoa" family="Plexauridae" genus="Psammogorgia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psammogorgia arbuscula subsp. var. var. pallida" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="variety" species="arbuscula" subSpecies="var." variety="pallida">Psammogorgia arbuscula var. pallida</taxonomicName>
Verrill, 1868b: 415-416; 
<bibRefCitation author="Kuekenthal, W" journalOrPublisher="Proceedings of the Zoological Society, London" pageId="0" pageNumber="1" refId="B22" refString="Kuekenthal, W, 1919. Gorgonaria. Wissenschaftliche Ergebnisse der deutsche Tiefsee-Expeditionen &quot;Valdivia&quot; 1898-99, 13(2): 1-946." title="Gorgonaria. Wissenschaftliche Ergebnisse der deutsche Tiefsee-Expeditionen &quot; Valdivia &quot; 1898 - 99, 13 (2): 1 - 946." year="1919">
<normalizedToken originalValue="Kükenthal">Kuekenthal</normalizedToken>
1919
</bibRefCitation>
: 237; 
<bibRefCitation author="Harden, DG" journalOrPublisher="Annals and Magazine of Natural History" pageId="0" pageNumber="1" refId="B20" refString="Harden, DG, 1979. Intuitive and Numerical Classification of East Pacific Gorgonacea (Octocorallia). PhD thesis, Illinois, Illinois State University." title="Intuitive and Numerical Classification of East Pacific Gorgonacea (Octocorallia). PhD thesis, Illinois, Illinois State University." year="1979">Harden 1979</bibRefCitation>
: 119.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Verrill" authorityYear="1868" class="Anthozoa" family="Plexauridae" genus="Psammogorgia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psammogorgia arbuscula subsp. pallida" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="subSpecies" species="arbuscula" subSpecies="pallida">Psammogorgia arbuscula pallida</taxonomicName>
<normalizedToken originalValue="Kükenthal">Kuekenthal</normalizedToken>
, 1924: 107.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="type material">
<paragraph pageId="0" pageNumber="1">Type material.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Syntypes</emphasis>
: MCZ 729 (4916), YPM 1785a-b, dry, Pearl Islands, 
<normalizedToken originalValue="Panamá">Panama</normalizedToken>
, F.H. Bradley, 1866-1867, no additional data.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="description">
<paragraph pageId="0" pageNumber="1">Description.</paragraph>
<paragraph pageId="0" pageNumber="1">
The syntype MCZ 729 is a yellowish flabellate, 10.5 cm long and ~ 9 cm wide colony. Two main stems arise from a holdfast that is 1.6 cm in diameter nd devoid of polyps (Fig. 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7A</figureCitation>
). The stems are less than 1 cm tall and 2.5 mm in diameter, subdividing irregularly in secondary branchlets of 2-3 mm diameter with rounded tips (Fig. 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7A</figureCitation>
). The branchlets emerge at angles of 45-180° and spread irregularly in one plane. The colony branches up to 15 times. Terminal branchlets are up to 40 mm long (Fig. 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7A</figureCitation>
). The polyps occur all around the branches, 1-1.5 mm apart on branchlets and 1.5-2.5 mm apart on the branches. The calyces are almost flat with a few being ~ 0.02 mm tall with oval or round polyp-apertures. Thorny, irregular spindles and wart-clubs occur around the calyx rim (Fig. 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7B</figureCitation>
). The coenenchymal sclerites are mostly irregular tuberculate spindles (Figs 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7B</figureCitation>
, 
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916) A spindles B wart clubs C radiates and crosses D anthocodial spindles." figureDoi="10.3897/zookeys.961.54846.figure8" httpUri="https://binary.pensoft.net/fig/442681" pageId="0" pageNumber="1">8A</figureCitation>
) with acute or bifurcated ends or combinations of both (Fig. 
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916) A spindles B wart clubs C radiates and crosses D anthocodial spindles." figureDoi="10.3897/zookeys.961.54846.figure8" httpUri="https://binary.pensoft.net/fig/442681" pageId="0" pageNumber="1">8A</figureCitation>
) with colours varying from pale pink to mostly colourless (Fig. 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7B</figureCitation>
). These sclerites are 0.15-0.23 mm long and 0.04-0.09 mm wide. Wart clubs are 0.11-0.14 mm long and 0.05-0.06 mm wide with warts or with wide tubercles (Fig. 
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916) A spindles B wart clubs C radiates and crosses D anthocodial spindles." figureDoi="10.3897/zookeys.961.54846.figure8" httpUri="https://binary.pensoft.net/fig/442681" pageId="0" pageNumber="1">8B</figureCitation>
). Crosses various intermediate forms and radiates are 0.08-0.16 mm long and 0.07-0.12 mm wide with tubercles (Fig. 
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916) A spindles B wart clubs C radiates and crosses D anthocodial spindles." figureDoi="10.3897/zookeys.961.54846.figure8" httpUri="https://binary.pensoft.net/fig/442681" pageId="0" pageNumber="1">8C</figureCitation>
). The anthocodial armature is composed of slightly bent spiny orange-red sclerites arranged in a collaret and points, 0.11-0.23 mm long and 0.02-0.045 mm wide (Figs 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7B</figureCitation>
, 
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916) A spindles B wart clubs C radiates and crosses D anthocodial spindles." figureDoi="10.3897/zookeys.961.54846.figure8" httpUri="https://binary.pensoft.net/fig/442681" pageId="0" pageNumber="1">8D</figureCitation>
).
</paragraph>
<caption doi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 7.</emphasis>
<taxonomicName authorityName="Verrill" authorityYear="1868" class="Anthozoa" family="Plexauridae" genus="Psammogorgia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psammogorgia arbuscula subsp. var. var. pallida" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="variety" species="arbuscula" subSpecies="var." variety="pallida">Psammogorgia arbuscula var. pallida</taxonomicName>
Verrill, 1868. MCZ 729 (4916). 
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
Colony 
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
coenenchymal and anthocodial sclerites.
</paragraph>
</caption>
<caption doi="10.3897/zookeys.961.54846.figure8" httpUri="https://binary.pensoft.net/fig/442681" pageId="0" pageNumber="1" start="Figure 8" startId="F8">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 8.</emphasis>
<taxonomicName authorityName="Verrill" authorityYear="1868" class="Anthozoa" family="Plexauridae" genus="Psammogorgia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psammogorgia arbuscula subsp. var. var. pallida" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="variety" species="arbuscula" subSpecies="var." variety="pallida">Psammogorgia arbuscula var. pallida</taxonomicName>
Verrill, 1868. MCZ 729 (4916) 
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
spindles 
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
wart clubs 
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
radiates and crosses 
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
anthocodial spindles.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">The syntypes</emphasis>
YPM 1785a, b are two fragments of a lighter colour than MCZ 729 (Fig. 
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Psammogorgia arbuscula var. pallida Verrill, 1868. YPM 1785 A colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure9" httpUri="https://binary.pensoft.net/fig/442682" pageId="0" pageNumber="1">9B</figureCitation>
). The largest fragment is 5.6 cm long and composed of four branchlets with 2-3 mm in diameter; flat calyces all around the branches (Fig. 
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Psammogorgia arbuscula var. pallida Verrill, 1868. YPM 1785 A colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure9" httpUri="https://binary.pensoft.net/fig/442682" pageId="0" pageNumber="1">9A</figureCitation>
). All sclerites, including the anthocodials, are pale yellow to colourless. Sclerites from the syntypes are more ornamented than in specimen MCZ 729 but mostly colourless (Fig. 
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Psammogorgia arbuscula var. pallida Verrill, 1868. YPM 1785 A colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure9" httpUri="https://binary.pensoft.net/fig/442682" pageId="0" pageNumber="1">9B</figureCitation>
). The coenenchymal sclerites are mostly irregular, tuberculate warty spindles with acute or bifurcated ends or combinations of both; being 0.13-0.18 mm long and 0.035-0.08 mm wide. Clubs have a few warts or with wide tubercles being 0.11-0.16 mm long and 0.05-0.60 mm wide. Radiates have tubercles appearing in various intermediate forms, being 0.07-0.10 mm long and 0.05-0.08 mm wide. The anthocodial armature is composed of slightly bent spiny spindles arranged in collaret and points, 0.12-0.22 mm long and 0.025-0.043 mm wide.
</paragraph>
<caption doi="10.3897/zookeys.961.54846.figure9" httpUri="https://binary.pensoft.net/fig/442682" pageId="0" pageNumber="1" start="Figure 9" startId="F9">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 9.</emphasis>
<taxonomicName authorityName="Verrill" authorityYear="1868" class="Anthozoa" family="Plexauridae" genus="Psammogorgia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Psammogorgia arbuscula subsp. var. var. pallida" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="variety" species="arbuscula" subSpecies="var." variety="pallida">Psammogorgia arbuscula var. pallida</taxonomicName>
Verrill, 1868. YPM 1785 
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
colony 
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
coenenchymal and anthocodial sclerites.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="remarks">
<paragraph pageId="0" pageNumber="1">Remarks and comparison.</paragraph>
<paragraph pageId="0" pageNumber="1">
<normalizedToken originalValue="Verrill´s">Verrill's</normalizedToken>
material at MCZ includes two specimens, MCZ 729 (4916) and YPM 1785, both with similar sclerites but different in external morphology. One is a small colony and the other is a small fragment in bad condition. According to 
<normalizedToken originalValue="Verrill´s">Verrill's</normalizedToken>
description (
<bibRefCitation author="Verrill, AE" journalOrPublisher="Transactions of the Connecticut Academy of Arts and Sciences, (Second Edition)" pageId="0" pageNumber="1" pagination="377 - 558" publicationUrl="https://biodiversitylibrary.org/page/13465394" refId="B41" refString="Verrill, AE, 1868b. Notes on Radiata in the Museum of Yale College, with descriptions of new genera and species. No. 6. Review of the corals and polyps of the West Coast of America. Transactions of the Connecticut Academy of Arts and Sciences, (Second Edition) 1 (2): 377 - 558, https://biodiversitylibrary.org/page/13465394" title="Notes on Radiata in the Museum of Yale College, with descriptions of new genera and species. No. 6. Review of the corals and polyps of the West Coast of America." url="https://biodiversitylibrary.org/page/13465394" volume="1" year="1868 b">Verrill 1868b</bibRefCitation>
) the type material has a &quot;corallum more or less flabelliform, branching dichotomously, branchlets round, sometimes as large as the main stem, usually smaller. Cells a little raised forming low verrucae&quot;. However, Verrill did not measure the specimens. YPM 1785 is different from 
<normalizedToken originalValue="Verrill’s">Verrill's</normalizedToken>
description while MCZ 729 (4916) matches some details of his description. Orange-red anthocodial sclerites are present in MCZ 729, as well as at the 
<taxonomicName lsidName="P. arbuscula subsp. var. var. pallida" pageId="0" pageNumber="1" rank="variety" species="arbuscula" subSpecies="var." variety="pallida">P. arbuscula var. pallida</taxonomicName>
description by 
<bibRefCitation author="Verrill, AE" journalOrPublisher="Transactions of the Connecticut Academy of Arts and Sciences, (Second Edition)" pageId="0" pageNumber="1" pagination="377 - 558" publicationUrl="https://biodiversitylibrary.org/page/13465394" refId="B41" refString="Verrill, AE, 1868b. Notes on Radiata in the Museum of Yale College, with descriptions of new genera and species. No. 6. Review of the corals and polyps of the West Coast of America. Transactions of the Connecticut Academy of Arts and Sciences, (Second Edition) 1 (2): 377 - 558, https://biodiversitylibrary.org/page/13465394" title="Notes on Radiata in the Museum of Yale College, with descriptions of new genera and species. No. 6. Review of the corals and polyps of the West Coast of America." url="https://biodiversitylibrary.org/page/13465394" volume="1" year="1868 b">Verrill (1868b)</bibRefCitation>
, in contrast with the colourless rods in specimen YPM 1785. Also, pale pink, colourless, and transparent coenenchymal sclerites match his description (Fig. 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Psammogorgia arbuscula var. pallida Verrill, 1868. MCZ 729 (4916). A Colony B coenenchymal and anthocodial sclerites." figureDoi="10.3897/zookeys.961.54846.figure7" httpUri="https://binary.pensoft.net/fig/442680" pageId="0" pageNumber="1">7B</figureCitation>
). In terms of sclerite sizes, 
<normalizedToken originalValue="Verrill’s">Verrill's</normalizedToken>
description better matches MCZ 729 with larger sclerite sizes than the smaller YPM 1785 ones. As Verrill suggested, with this and 
<emphasis italics="true" pageId="0" pageNumber="1">dowii</emphasis>
variety, we opt to keep 
<taxonomicName lsidName="P. arbuscula subsp. var. var. pallida" pageId="0" pageNumber="1" rank="variety" species="arbuscula" subSpecies="var." variety="pallida">P. arbuscula var. pallida</taxonomicName>
as a variety.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="distribution">
<paragraph pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="1">
Tropical eastern Pacific: only reported from the type locality at Pearl Islands, 
<normalizedToken originalValue="Panamá">Panama</normalizedToken>
.
</paragraph>
</subSubSection>
</treatment>
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