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<mods:title id="FC20DDD60726ADFF1E7BCCEFA8F0EA3C">Biofouling hydroids (Cnidaria: Hydrozoa) from a Tropical Eastern Pacific island, with remarks on their biogeography</mods:title>
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<paragraph id="8BB70C22FFD9FFB2899CF99D1427F9BE" blockId="7.[362,953,1337,1676]" box="[444,871,1615,1642]" pageId="7" pageNumber="572">
<taxonomicName id="4C0877A1FFD9FFB2899CF99D1427F9BE" ID-CoL="76JN9" authority="Goldfuss, 1820" authorityName="Goldfuss" authorityYear="1820" box="[444,871,1615,1642]" class="Hydrozoa" family="Pennariidae" genus="Pennaria" kingdom="Animalia" order="Anthoathecata" pageId="7" pageNumber="572" phylum="Cnidaria" rank="species" species="disticha">
<emphasis id="B97CD030FFD9FFB2899CF99D1427F9BE" bold="true" box="[444,871,1615,1642]" pageId="7" pageNumber="572">
<emphasis id="B97CD030FFD9FFB2899CF99D15DBF9BE" bold="true" box="[444,667,1615,1642]" italics="true" pageId="7" pageNumber="572">Pennaria disticha</emphasis>
<bibRefCitation id="EF9971D3FFD9FFB28A83F99D1427F9BE" author="Goldfuss GA" box="[675,871,1615,1642]" pageId="7" pageNumber="572" refId="ref19936" refString="Goldfuss GA. 1820. Handbuch der Zoologie. I. Abtheilung. Nurnberg: Johann Leonhard Schrag." type="book" year="1820">Goldfuss, 1820</bibRefCitation>
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<paragraph id="8BB70C22FFD9FFB28A6AF9A31599F958" blockId="7.[362,953,1337,1676]" box="[586,729,1649,1676]" pageId="7" pageNumber="572">
<emphasis id="B97CD030FFD9FFB28A6AF9A31599F958" bold="true" box="[586,729,1649,1676]" pageId="7" pageNumber="572">
(
<figureCitation id="133310A7FFD9FFB28A73F9A31590F958" box="[595,720,1649,1676]" captionStart="Figure 2" captionStartId="9.[160,225,1124,1147]" captionTargetBox="[165,1146,145,1083]" captionTargetId="figure-66@9.[163,1153,142,1093]" captionTargetPageId="9" captionText="Figure 2. Anthoathecata: Pennariidae, Oceaniidae and Eudendriidae. (a) Pennaria disticha: hydranth, pedicel and part of hydrocaulus, #266339. (b) Corydendrium flabellatum: two hydranths and distal end of hydrocaulus, #266340. (c) Eudendrium cf. certicaule: hydranth and part of pedicel, #240637. (d) Eudendrium cf. certicaule: pedicel and hydranth with female gonophores, #240634. (e) Eudendrium cf. certicaule: pedicel and hydranth with male gonophores, #240636. (f) Eudendrium cf. certicaule: base of hydrocaulus with a branch, #240636. Scale bars: a, b, f = 0.2 mm; ce = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.7012490" httpUri="https://zenodo.org/record/7012490/files/figure.png" pageId="7" pageNumber="572">Figure 2a</figureCitation>
)
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<paragraph id="8BB70C22FFD9FFB28880F9751577F914" blockId="7.[160,567,1703,1728]" box="[160,567,1703,1728]" pageId="7" pageNumber="572">
<treatmentCitationGroup id="AB182B0CFFD9FFB28880F9751577F914" box="[160,567,1703,1728]" pageId="7" pageNumber="572">
<treatmentCitation id="0AA92A33FFD9FFB28880F9751573F914" author="Goldfuss GA" box="[160,563,1703,1728]" page="89" pageId="7" pageNumber="572" year="1820">
<taxonomicName id="4C0877A1FFD9FFB28880F9751573F914" ID-CoL="76JN9" authority="Goldfuss, 1820: 89" authorityName="Goldfuss" authorityPageNumber="89" authorityYear="1820" box="[160,563,1703,1728]" class="Hydrozoa" family="Pennariidae" genus="Pennaria" kingdom="Animalia" order="Anthoathecata" pageId="7" pageNumber="572" phylum="Cnidaria" rank="species" species="disticha">
<emphasis id="B97CD030FFD9FFB28880F975161CF914" box="[160,348,1703,1728]" italics="true" pageId="7" pageNumber="572">Pennaria disticha</emphasis>
<bibRefCitation id="EF9971D3FFD9FFB28943F9751573F914" author="Goldfuss GA" box="[355,563,1703,1728]" pageId="7" pageNumber="572" refId="ref19936" refString="Goldfuss GA. 1820. Handbuch der Zoologie. I. Abtheilung. Nurnberg: Johann Leonhard Schrag." type="book" year="1820">Goldfuss, 1820: 89</bibRefCitation>
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.
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<emphasis id="B97CD030FFD6FFBD8880FF5C1676FF7C" bold="true" box="[160,310,142,168]" italics="true" pageId="8" pageNumber="573">
<typeStatus id="54B3B280FFD6FFBD8880FF5C1798FF7C" box="[160,216,142,168]" pageId="8" pageNumber="573">Type</typeStatus>
locality
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<materialsCitation id="3B60067FFFD6FFBD8880FF60153DFF1F" ID-GBIF-Occurrence="3891616424" box="[160,637,178,203]" country="Italy" location="Gulf of Naples" pageId="8" pageNumber="573" specimenCount="1" typeStatus="holotype">
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:
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(see
<bibRefCitation id="EF9971D3FFD6FFBD8997FF60157DFF1F" author="Calder DR" box="[439,573,178,203]" pageId="8" pageNumber="573" pagination="1 - 72" refId="ref17660" refString="Calder DR. 2013. Some shallow-water hydroids (Cnidaria: Hydrozoa) from the central east coast of Florida, USA. Zootaxa. 3648: 1 - 72. doi: 10.11646 / zootaxa. 3648.1.1" type="journal article" year="2013">Calder 2013</bibRefCitation>
, p. 7)
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.
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<emphasis id="B97CD030FFD6FFBD8880FF24163FFEC4" bold="true" box="[160,383,246,272]" italics="true" pageId="8" pageNumber="573">Material examined</emphasis>
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<paragraph id="8BB70C22FFD6FFBD8880FEC81646FE81" blockId="8.[160,1156,246,341]" pageId="8" pageNumber="573">
Chatham Bay, dock 004, 1 colony,
<quantity id="4CF0A1C7FFD6FFBD8A1AFEC815C8FEE7" box="[570,648,282,307]" metricMagnitude="-2" metricUnit="m" metricValue="2.8" pageId="8" pageNumber="573" unit="cm" value="2.8">2.8 cm</quantity>
high, without gonophores, coll. G. Ashton, #266339.
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<emphasis id="B97CD030FFD6FFBD8880FE531646FE4F" bold="true" box="[160,262,385,411]" italics="true" pageId="8" pageNumber="573">Remarks</emphasis>
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The hydroid of
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<emphasis id="B97CD030FFD6FFBD8968FE7616BFFE69" box="[328,511,420,445]" italics="true" pageId="8" pageNumber="573">Pennaria disticha</emphasis>
<bibRefCitation id="EF9971D3FFD6FFBD8A25FE7615E6FE69" author="Goldfuss GA" box="[517,678,420,445]" pageId="8" pageNumber="573" refId="ref19936" refString="Goldfuss GA. 1820. Handbuch der Zoologie. I. Abtheilung. Nurnberg: Johann Leonhard Schrag." type="book" year="1820">Goldfuss, 1820</bibRefCitation>
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has in the past been considered essentially cosmopolitan in tropical to temperate waters. While the species is known to be invasive, barcoding studies reveal the existence of cryptic species within the binomen (
<bibRefCitation id="EF9971D3FFD6FFBD8C01FE381652FDF1" author="Miglietta MP &amp; Odegard D &amp; Faure B &amp; Faucci A" pageId="8" pageNumber="573" pagination="0144762" refId="ref21654" refString="Miglietta MP, Odegard D, Faure B, Faucci A. 2015. Barcoding techniques help tracking the evolutionary history of the introduced species Pennaria disticha (Hydrozoa, Cnidaria). PLoS ONE. 10 (12): e 0144762. doi: 10.1371 / journal. pone. 0144762" type="journal article" year="2015">Miglietta et al. 2015</bibRefCitation>
,
<bibRefCitation id="EF9971D3FFD6FFBD893DFDDE1613FDF1" author="Miglietta MP &amp; Maggioni D &amp; Matsumoto Y" box="[285,339,524,549]" pageId="8" pageNumber="573" pagination="1085 - 1100" refId="ref21707" refString="Miglietta MP, Maggioni D, Matsumoto Y. 2019. Phylogenetics and species delimitation of two hydrozoa (phylum Cnidaria): Turritopsis (McCrady, 1857) and Pennaria (Goldfuss, 1820). Mar Biodivers. 49: 1085 - 1100. doi: 10.1007 / s 12526 - 018 - 0891 - 8" type="journal article" year="2019">2019</bibRefCitation>
). In the analysis of
<bibRefCitation id="EF9971D3FFD6FFBD8A07FDDE1450FDF1" author="Miglietta MP &amp; Maggioni D &amp; Matsumoto Y" box="[551,784,524,549]" pageId="8" pageNumber="573" pagination="1085 - 1100" refId="ref21707" refString="Miglietta MP, Maggioni D, Matsumoto Y. 2019. Phylogenetics and species delimitation of two hydrozoa (phylum Cnidaria): Turritopsis (McCrady, 1857) and Pennaria (Goldfuss, 1820). Mar Biodivers. 49: 1085 - 1100. doi: 10.1007 / s 12526 - 018 - 0891 - 8" type="journal article" year="2019">Miglietta et al. (2019)</bibRefCitation>
, specimens from the Pacific coast of
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, in the Tropical Eastern Pacific, occurred in both a clade (1A) with other populations from
<collectingCountry id="F31F4CB2FFD6FFBD894DFD8016EAFDBF" box="[365,426,594,619]" name="Brazil" pageId="8" pageNumber="573">Brazil</collectingCountry>
, Florida, Hawaii,
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Sea, and
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in the Western Indian Ocean, and in another clade (2D) with specimens from North Carolina, the Caribbean, Atlantic
<collectingCountry id="F31F4CB2FFD6FFBD88DAFD451615FD64" box="[250,341,663,688]" name="Panama" pageId="8" pageNumber="573">Panama</collectingCountry>
, Hawaii, Chuuk,
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,
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, and the Red Sea. Specimens from
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(the
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locality of the species) and elsewhere in the Mediterranean occurred in a separate clade (Clade 2C), along with material from the Azores and the Gulf of Mexico. Taxonomic and nomenclatural issues arising from these results have yet to be resolved, however, and the name
<taxonomicName id="4C0877A1FFD6FFBD8991FCF0155DFCEF" authorityName="Goldfuss" authorityYear="1820" box="[433,541,802,827]" class="Hydrozoa" family="Pennariidae" genus="Pennaria" kingdom="Animalia" order="Anthoathecata" pageId="8" pageNumber="573" phylum="Cnidaria" rank="species" species="disticha">
<emphasis id="B97CD030FFD6FFBD8991FCF0155DFCEF" box="[433,541,802,827]" italics="true" pageId="8" pageNumber="573">P. disticha</emphasis>
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has been retained for specimens examined here.
</paragraph>
<paragraph id="8BB70C22FFD6FFBD88E0FC961376FBFA" blockId="8.[160,1156,385,1174]" pageId="8" pageNumber="573">
In the eastern Pacific,
<taxonomicName id="4C0877A1FFD6FFBD89E4FC9615C3FC89" authorityName="Goldfuss" authorityYear="1820" box="[452,643,836,861]" class="Hydrozoa" family="Pennariidae" genus="Pennaria" kingdom="Animalia" order="Anthoathecata" pageId="8" pageNumber="573" phylum="Cnidaria" rank="species" species="disticha">
<emphasis id="B97CD030FFD6FFBD89E4FC9615C3FC89" box="[452,643,836,861]" italics="true" pageId="8" pageNumber="573">Pennaria disticha</emphasis>
</taxonomicName>
has been reported from San Francisco Bay, California, to Santa Elena Bay,
<collectingCountry id="F31F4CB2FFD6FFBD89CEFCB51509FC54" box="[494,585,871,896]" name="Ecuador" pageId="8" pageNumber="573">Ecuador</collectingCountry>
(
<bibRefCitation id="EF9971D3FFD6FFBD8A75FCB51597FC54" author="Fraser CM" box="[597,727,871,896]" pageId="8" pageNumber="573" refId="ref19504" refString="Fraser CM. 1937. Hydroids of the Pacific coast of Canada and the United States. Toronto: University of Toronto Press. doi: 10.3138 / 9781487595548" type="book" year="1937">Fraser 1937</bibRefCitation>
,
<bibRefCitation id="EF9971D3FFD6FFBD8AC3FCB51466FC54" author="Fraser CM" box="[739,806,871,896]" pageId="8" pageNumber="573" pagination="1 - 105" refId="ref19533" refString="Fraser CM. 1938 a. Hydroids of the 1934 Allan Hancock Pacific Expedition. Allan Hancock Pac Expeditions. 4 (1): 1 - 105." type="journal article" year="1938">1938a</bibRefCitation>
,
<bibRefCitation id="EF9971D3FFD6FFBD8B12FCB51435FC54" author="Fraser CM" box="[818,885,871,896]" pageId="8" pageNumber="573" pagination="129 - 153" refId="ref19593" refString="Fraser CM. 1938 c. Hydroids of the 1932, 1933, 1935, and 1938 Allan Hancock Pacific Expeditions. Allan Hancock Pac Expeditions. 4 (3): 129 - 153." type="journal article" year="1938">1938c</bibRefCitation>
,
<bibRefCitation id="EF9971D3FFD6FFBD8BA0FCB514F7FC54" author="Fraser CM" box="[896,951,871,896]" pageId="8" pageNumber="573" refId="ref19676" refString="Fraser CM. 1947. Distribution and relationship in American hydroids. Toronto: University of Toronto Press." type="book" year="1947">1947</bibRefCitation>
,
<bibRefCitation id="EF9971D3FFD6FFBD8BE3FCB514BAFC54" author="Fraser CM" box="[963,1018,871,896]" pageId="8" pageNumber="573" pagination="179 - 343" refId="ref19695" refString="Fraser CM. 1948. Hydroids of the Allan Hancock Pacific Expeditions since March, 1938. Allan Hancock Pac Expeditions. 4 (5): 179 - 343." type="journal article" year="1948">1948</bibRefCitation>
, as
<taxonomicName id="4C0877A1FFD6FFBD8C03FCB517ABFC77" baseAuthorityName="Ayres" baseAuthorityYear="1854" class="Hydrozoa" family="Pennariidae" genus="Pennaria" kingdom="Animalia" order="Anthoathecata" pageId="8" pageNumber="573" phylum="Cnidaria" rank="species" species="tiarella">
<emphasis id="B97CD030FFD6FFBD8C03FCB517ABFC77" italics="true" pageId="8" pageNumber="573">Pennaria tiarella</emphasis>
</taxonomicName>
). On the Pacific coast of
<collectingCountry id="F31F4CB2FFD6FFBD8A20FC581535FC77" box="[512,629,906,931]" name="Costa Rica" pageId="8" pageNumber="573">Costa Rica</collectingCountry>
, it has been reported from Port Culebra (
<bibRefCitation id="EF9971D3FFD6FFBD8C1EFC5817B3FC12" author="Fraser CM" pageId="8" pageNumber="573" pagination="1 - 105" refId="ref19533" refString="Fraser CM. 1938 a. Hydroids of the 1934 Allan Hancock Pacific Expedition. Allan Hancock Pac Expeditions. 4 (1): 1 - 105." type="journal article" year="1938">Fraser (1938a)</bibRefCitation>
and Bahía Huevos at
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,
<geoCoordinate id="EE3C6AE5FFD6FFBD8AA3FC7F1447FC12" box="[643,775,941,966]" degrees="85.69167" direction="west" orientation="longitude" pageId="8" pageNumber="573" precision="1" value="-85.69167">85.69167°W</geoCoordinate>
(
<bibRefCitation id="EF9971D3FFD6FFBD8B37FC7F1369FC12" author="Kelmo F &amp; Vargas R" box="[791,1065,941,966]" pageId="8" pageNumber="573" pagination="599 - 627" refId="ref20587" refString="Kelmo F, Vargas R. 2002. Anthoathecatae and Leptothecatae hydroids from Costa Rica (Cnidaria: Hydrozoa). Rev Biol Trop. 50: 599 - 627." type="journal article" year="2002">Kelmo and Vargas 2002</bibRefCitation>
). In the Galápagos Islands
<taxonomicName id="4C0877A1FFD6FFBD8957FC1D16A8FC3C" authorityName="Goldfuss" authorityYear="1820" box="[375,488,975,1000]" class="Hydrozoa" family="Pennariidae" genus="Pennaria" kingdom="Animalia" order="Anthoathecata" pageId="8" pageNumber="573" phylum="Cnidaria" rank="species" species="disticha">
<emphasis id="B97CD030FFD6FFBD8957FC1D16A8FC3C" box="[375,488,975,1000]" italics="true" pageId="8" pageNumber="573">P. disticha</emphasis>
</taxonomicName>
is one of the most common and conspicuous species of hydroids on exposed coasts at depths ranging from the surface to
<quantity id="4CF0A1C7FFD6FFBD8BBDFC201498FBDF" box="[925,984,1010,1035]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="573" unit="m" value="10.0">10 m</quantity>
, except on the westernmost
<collectingCountry id="F31F4CB2FFD6FFBD8919FBC7161DFBFA" box="[313,349,1045,1070]" name="Iceland" pageId="8" pageNumber="573">Isla</collectingCountry>
Fernandina (D. Calder, personal observations,
<date id="FFB62AE2FFD6FFBD8B4AFBC7136BFBFA" box="[874,1067,1045,1070]" pageId="8" pageNumber="573" value="2001-06-16" valueMax="2001-06-22" valueMin="2001-06-16">1622 June 2001</date>
).
</paragraph>
<paragraph id="8BB70C22FFD6FFBD88E0FBE514A6FB42" blockId="8.[160,1156,385,1174]" pageId="8" pageNumber="573">
A species of many habitats, we regard
<taxonomicName id="4C0877A1FFD6FFBD8A4DFBE5159BFB84" authorityName="Goldfuss" authorityYear="1820" box="[621,731,1079,1104]" class="Hydrozoa" family="Pennariidae" genus="Pennaria" kingdom="Animalia" order="Anthoathecata" pageId="8" pageNumber="573" phylum="Cnidaria" rank="species" species="disticha">
<emphasis id="B97CD030FFD6FFBD8A4DFBE5159BFB84" box="[621,731,1079,1104]" italics="true" pageId="8" pageNumber="573">P. disticha</emphasis>
</taxonomicName>
as cryptogenic in the Tropical Eastern Pacific (including in Cocos and the
<collectingRegion id="49CCC2C0FFD6FFBD8A0EFB8815BCFBA7" box="[558,764,1114,1139]" country="Ecuador" name="Galapagos" pageId="8" pageNumber="573">Galapagos Islands</collectingRegion>
), pending further resolution of the taxonomy and geography of the multiple genetic clades detected to date.
</paragraph>
</subSubSection>
<subSubSection id="C3125FA9FFD6FFBD8880FB131372FA97" pageId="8" pageNumber="573" type="distribution">
<paragraph id="8BB70C22FFD6FFBD8880FB1316DCFB0F" blockId="8.[160,1157,1217,1347]" box="[160,412,1217,1243]" pageId="8" pageNumber="573">
<emphasis id="B97CD030FFD6FFBD8880FB1316DCFB0F" bold="true" box="[160,412,1217,1243]" italics="true" pageId="8" pageNumber="573">Reported distribution</emphasis>
</paragraph>
<paragraph id="8BB70C22FFD6FFBD8880FB3716FAFB2A" blockId="8.[160,1157,1217,1347]" box="[160,442,1253,1278]" pageId="8" pageNumber="573">Cocos Island: first record.</paragraph>
<paragraph id="8BB70C22FFD6FFBD88E0FAD51372FA97" blockId="8.[160,1157,1217,1347]" pageId="8" pageNumber="573">
Elsewhere: reported to be circumglobal in tropical and warm-temperate waters, although cryptic species appear to exist (
<bibRefCitation id="EF9971D3FFD6FFBD8A4EFAF815B6FA97" author="Calder DR" box="[622,758,1322,1347]" pageId="8" pageNumber="573" pagination="1 - 91" refId="ref17625" refString="Calder DR. 2010. Some anthoathecate hydroids and limnopolyps (Cnidaria, Hydrozoa) from the Hawaiian archipelago. Zootaxa. 2590: 1 - 91. doi: 10.11646 / zootaxa. 2590.1.1" type="journal article" year="2010">Calder 2010</bibRefCitation>
;
<bibRefCitation id="EF9971D3FFD6FFBD8B23FAF814A2FA97" author="Miglietta MP &amp; Odegard D &amp; Faure B &amp; Faucci A" box="[771,994,1322,1347]" pageId="8" pageNumber="573" pagination="0144762" refId="ref21654" refString="Miglietta MP, Odegard D, Faure B, Faucci A. 2015. Barcoding techniques help tracking the evolutionary history of the introduced species Pennaria disticha (Hydrozoa, Cnidaria). PLoS ONE. 10 (12): e 0144762. doi: 10.1371 / journal. pone. 0144762" type="journal article" year="2015">Miglietta et al. 2015</bibRefCitation>
,
<bibRefCitation id="EF9971D3FFD6FFBD8BCFFAF81366FA97" author="Miglietta MP &amp; Maggioni D &amp; Matsumoto Y" box="[1007,1062,1322,1347]" pageId="8" pageNumber="573" pagination="1085 - 1100" refId="ref21707" refString="Miglietta MP, Maggioni D, Matsumoto Y. 2019. Phylogenetics and species delimitation of two hydrozoa (phylum Cnidaria): Turritopsis (McCrady, 1857) and Pennaria (Goldfuss, 1820). Mar Biodivers. 49: 1085 - 1100. doi: 10.1007 / s 12526 - 018 - 0891 - 8" type="journal article" year="2019">2019</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>