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<document ID-DOI="10.1645/18-67" ID-GBIF-Dataset="a39151be-8bb1-4dcb-bfcb-31af4e40f0f4" ID-ISSN="1937-2345" ID-Zenodo-Dep="7752619" checkinTime="1679077589534" checkinUser="felipe" docAuthor="Matsche, Mark A., Adams, Cynthia R. &amp; Blazer, Vicki S." docDate="2019" docId="B76487A5FFE2650CFEB5C0369494F842" docLanguage="en" docName="JourParasitol.105.1.1-10.pdf" docOrigin="Journal of Parasitology (J. Parasitol.) 105 (1)" docSource="http://dx.doi.org/10.1645/18-67" docStyle="DocumentStyle:378338F1068CC775F20F95F13175CB49.1:JourParasitol.2019-.journal_article" docStyleId="378338F1068CC775F20F95F13175CB49" docStyleName="JourParasitol.2019-.journal_article" docStyleVersion="1" docTitle="Goussia bayae Matsche &amp; Adams &amp; Blazer 2019, n. sp." docType="treatment" docVersion="3" lastPageNumber="7" masterDocId="4B5DFFDDFFE0650AFFD2C65F911CFFE6" masterDocTitle="Newly Described Coccidia Goussia Bayae From White Perch Morone Americana: Morphology And Phylogenetics Support Emerging Taxonomy Of Goussia Within Piscine Hosts" masterLastPageNumber="10" masterPageNumber="1" pageNumber="3" updateTime="1679317683926" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Newly Described Coccidia Goussia Bayae From White Perch Morone Americana: Morphology And Phylogenetics Support Emerging Taxonomy Of Goussia Within Piscine Hosts</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Matsche, Mark A.</mods:namePart>
<mods:affiliation>Cooperative Oxford Laboratory, Oxford, Maryland 21654.</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Adams, Cynthia R.</mods:namePart>
<mods:affiliation>Cooperative Oxford Laboratory, Oxford, Maryland 21654. &amp; National Fish Health Research Laboratory, Leetown Science Center, U. S. Geological Survey, Kearneysville, West Virginia 25430. Correspondence should be sent to Mark A. Matsche at: mark. matsche @ maryland. gov</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Blazer, Vicki S.</mods:namePart>
<mods:affiliation>Cooperative Oxford Laboratory, Oxford, Maryland 21654. &amp; National Fish Health Research Laboratory, Leetown Science Center, U. S. Geological Survey, Kearneysville, West Virginia 25430. Correspondence should be sent to Mark A. Matsche at: mark. matsche @ maryland. gov &amp; Cooperative Oxford Laboratory, Oxford, Maryland 21654.</mods:affiliation>
</mods:name>
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<mods:titleInfo>
<mods:title>Journal of Parasitology</mods:title>
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<mods:part>
<mods:date>2019</mods:date>
<mods:detail type="series">
<mods:title>J. Parasitol.</mods:title>
</mods:detail>
<mods:detail type="pubDate">
<mods:number>2019-01-18</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>105</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>1</mods:number>
</mods:detail>
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<mods:start>1</mods:start>
<mods:end>10</mods:end>
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<mods:url>http://dx.doi.org/10.1645/18-67</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1645/18-67</mods:identifier>
<mods:identifier type="GBIF-Dataset">a39151be-8bb1-4dcb-bfcb-31af4e40f0f4</mods:identifier>
<mods:identifier type="ISSN">1937-2345</mods:identifier>
<mods:identifier type="Zenodo-Dep">7752619</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.7752621" ID-Zenodo-Dep="7752621" LSID="urn:lsid:plazi:treatment:B76487A5FFE2650CFEB5C0369494F842" httpUri="http://treatment.plazi.org/id/B76487A5FFE2650CFEB5C0369494F842" lastPageId="6" lastPageNumber="7" pageId="2" pageNumber="3">
<subSubSection box="[359,599,1641,1663]" pageId="2" pageNumber="3" type="nomenclature">
<paragraph blockId="2.[359,599,1641,1663]" box="[359,599,1641,1663]" pageId="2" pageNumber="3">
<heading box="[359,599,1641,1663]" centered="true" fontSize="36" level="2" pageId="2" pageNumber="3" reason="2">
<emphasis bold="true" box="[359,599,1641,1663]" pageId="2" pageNumber="3">
<taxonomicName authority="Matsche &amp; Adams &amp; Blazer, 2019" authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[359,527,1641,1663]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="bayae" status="n. sp.">
<emphasis bold="true" box="[359,527,1641,1663]" italics="true" pageId="2" pageNumber="3">Goussia bayae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[536,599,1641,1663]" pageId="2" pageNumber="3" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[426,531,1673,1692]" pageId="2" pageNumber="3" type="description">
<paragraph blockId="2.[426,531,1673,1692]" box="[426,531,1673,1692]" pageId="2" pageNumber="3">
(
<figureCitation box="[430,526,1673,1692]" captionStart-0="Figure 1" captionStart-1="Figure 2" captionStart-2="Figure 3" captionStart-3="Figure 4" captionStartId-0="2.[168,225,890,909]" captionStartId-1="2.[869,926,781,800]" captionStartId-2="3.[141,198,1535,1554]" captionStartId-3="4.[168,225,1611,1630]" captionTargetBox-0="[146,812,202,863]" captionTargetBox-1="[853,1505,208,742]" captionTargetBox-2="[118,1488,202,1508]" captionTargetBox-3="[145,1515,202,1584]" captionTargetId-0="figure-545@2.[146,812,202,863]" captionTargetId-1="figure-603@2.[847,1513,202,754]" captionTargetId-2="figure-1@3.[118,1488,202,1508]" captionTargetId-3="figure-1@4.[145,1515,202,1584]" captionTargetPageId-0="2" captionTargetPageId-1="2" captionTargetPageId-2="3" captionTargetPageId-3="4" captionText-0="Figure 1. Photomicrographs of oocysts of Goussia bayae n. sp. from the gallbladder of white perch, Morone americana. (A) Nomarski differential interference contrast image of mature oocysts with micropyle (m) and refractile bodies (arrow) in sporocysts. (B) Histological preparation of oocysts with sporocysts containing paired sporozoites with densely stained nuclei (arrow)." captionText-1="Figure 2. Goussia bayae n. sp. Line drawing of a sporulated oocyst." captionText-2="Figure 3. Light micrographs of coccidia of Goussia bayae n. sp. in the gallbladder of white perch, Morone americana. (A) Severe coccidiosis in gallbladder. Note that numerous oocysts were removed with the bile for differential interference contrast microscopy before histological processing. (B) Developing stages of coccidia along epithelium (arrow) and sporulating oocysts in the lumen. (C) Microgamont (Mi), macrogamont (Ma), and meront (Me) epicellular to biliary epithelium, with oocysts (O) in the lumen." captionText-3="Figure 4. Light micrographs of coccidia of Goussia bayae n. sp. in hepatic bile ducts of white perch, Morone americana. (A) Developing stages of coccidia epicellular to biliary epithelium. (B) Microgamont (Mi), macrogamont (Ma), and meront (Me) along epithelium. (C) Longitudinal view of bile duct with developing coccidia along epithelium (arrow) and sporulating oocysts (O) in lumen. (D). Cross-section of enlarged bile duct with numerous developing and mature coccidia." figureDoi-0="http://doi.org/10.5281/zenodo.7752625" figureDoi-1="http://doi.org/10.5281/zenodo.7752623" figureDoi-2="http://doi.org/10.5281/zenodo.7752627" figureDoi-3="http://doi.org/10.5281/zenodo.7752629" httpUri-0="https://zenodo.org/record/7752625/files/figure.png" httpUri-1="https://zenodo.org/record/7752623/files/figure.png" httpUri-2="https://zenodo.org/record/7752627/files/figure.png" httpUri-3="https://zenodo.org/record/7752629/files/figure.png" pageId="2" pageNumber="3">Figs. 14</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="3" type="diagnosis">
<paragraph blockId="2.[144,814,1728,1956]" lastBlockId="2.[845,1515,845,926]" pageId="2" pageNumber="3">
<emphasis box="[168,273,1728,1749]" italics="true" pageId="2" pageNumber="3">Diagnosis:</emphasis>
Oocyst shape: subspheroidal, with a single-layered smooth wall,
<emphasis bold="true" box="[284,302,1757,1778]" pageId="2" pageNumber="3">~</emphasis>
0.40.6 thick; oocyst length (L) X width (W): 26.2 ± 2.1 SD X 21.8 ± 2.1 SD (ranges: 2230 X 1825); oocyst L/
<geoCoordinate degrees="1.2" direction="west" orientation="longitude" pageId="2" pageNumber="3" precision="5555" value="-1.2">W 1.2</geoCoordinate>
± 0.05 SD (range: 1.11.3) (
<figureCitation box="[494,567,1816,1838]" captionStart="Figure 1" captionStartId="2.[168,225,890,909]" captionTargetBox="[146,812,202,863]" captionTargetId="figure-545@2.[146,812,202,863]" captionTargetPageId="2" captionText="Figure 1. Photomicrographs of oocysts of Goussia bayae n. sp. from the gallbladder of white perch, Morone americana. (A) Nomarski differential interference contrast image of mature oocysts with micropyle (m) and refractile bodies (arrow) in sporocysts. (B) Histological preparation of oocysts with sporocysts containing paired sporozoites with densely stained nuclei (arrow)." figureDoi="http://doi.org/10.5281/zenodo.7752625" httpUri="https://zenodo.org/record/7752625/files/figure.png" pageId="2" pageNumber="3">Figs. 1</figureCitation>
,
<figureCitation box="[584,598,1816,1838]" captionStart="Figure 2" captionStartId="2.[869,926,781,800]" captionTargetBox="[853,1505,208,742]" captionTargetId="figure-603@2.[847,1513,202,754]" captionTargetPageId="2" captionText="Figure 2. Goussia bayae n. sp. Line drawing of a sporulated oocyst." figureDoi="http://doi.org/10.5281/zenodo.7752623" httpUri="https://zenodo.org/record/7752623/files/figure.png" pageId="2" pageNumber="3">2</figureCitation>
); micropyle present; oocyst residuum and polar granules absent; sporocyst shape: ellipsoidal, slightly tapered on 1 end,
<emphasis bold="true" box="[527,545,1875,1896]" pageId="2" pageNumber="3">~</emphasis>
12.6 ± 0.6 SD X 7.8 ± 0.8 SD (ranges: 1014 X 69), sporocyst L/
<geoCoordinate box="[558,622,1905,1927]" degrees="1.6" direction="west" orientation="longitude" pageId="2" pageNumber="3" precision="5555" value="-1.6">W 1.6</geoCoordinate>
± 0.1 SD (range: 1.41.9), with a smooth wall composed of 2 valves joined by a longitudinal suture; Stieda body, sub-Stieda body, sporocyst residuum absent; excysted sporozoite shape: slightly arcuate, 1 end tapered to a point; nucleus and refractile bodies present.
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="3" type="description">
<paragraph blockId="2.[845,1084,962,984]" box="[845,1084,962,984]" pageId="2" pageNumber="3">
<heading bold="true" box="[845,1084,962,984]" fontSize="9" level="3" pageId="2" pageNumber="3" reason="9">
<emphasis bold="true" box="[845,1084,962,984]" pageId="2" pageNumber="3">Taxonomic summary</emphasis>
</heading>
</paragraph>
<paragraph blockId="2.[845,1515,1008,1441]" pageId="2" pageNumber="3">
<emphasis box="[869,979,1008,1029]" italics="true" pageId="2" pageNumber="3">
<typeStatus box="[869,919,1008,1029]" pageId="2" pageNumber="3">Type</typeStatus>
host:
</emphasis>
White perch,
<taxonomicName authority="(Gmelin 1789)" baseAuthorityName="Gmelin" baseAuthorityYear="1789" box="[1134,1438,1008,1030]" class="Actinopterygii" family="Moronidae" genus="Morone" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="americana">
<emphasis box="[1134,1278,1008,1029]" italics="true" pageId="2" pageNumber="3">M. americana</emphasis>
(Gmelin 1789)
</taxonomicName>
(
<taxonomicName class="Actinopterygii" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="3" phylum="Chordata" rank="order">Perciformes</taxonomicName>
:
<taxonomicName box="[930,1049,1037,1059]" class="Actinopterygii" family="Moronidae" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Moronidae</taxonomicName>
).
</paragraph>
<paragraph blockId="2.[845,1515,1008,1441]" box="[869,1112,1067,1089]" pageId="2" pageNumber="3">
<emphasis box="[869,994,1067,1088]" italics="true" pageId="2" pageNumber="3">Other hosts:</emphasis>
Unknown.
</paragraph>
<paragraph blockId="2.[845,1515,1008,1441]" pageId="2" pageNumber="3">
<emphasis box="[869,1056,1097,1118]" italics="true" pageId="2" pageNumber="3">Site of infection:</emphasis>
Hepatic bile ducts, common duct, and gallbladder.
</paragraph>
<paragraph blockId="2.[845,1515,1008,1441]" box="[869,1415,1155,1177]" pageId="2" pageNumber="3">
<emphasis box="[869,1106,1155,1176]" italics="true" pageId="2" pageNumber="3">Prevalence of infection:</emphasis>
Of 150 fish examined, 100%.
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="3" type="materials_examined">
<paragraph blockId="2.[845,1515,1008,1441]" pageId="2" pageNumber="3">
<materialsCitation latitude="38.776665" location="Preston" longLatPrecision="1" longitude="-75.96956" municipality="Choptank River" pageId="2" pageNumber="3" specimenCount="1" stateProvince="Maryland" typeStatus="holotype">
<emphasis box="[869,1040,1185,1206]" italics="true" pageId="2" pageNumber="3">
<typeStatus box="[869,925,1185,1206]" pageId="2" pageNumber="3">Type</typeStatus>
locality:
</emphasis>
<collectingMunicipality box="[1059,1261,1184,1206]" pageId="2" pageNumber="3">Choptank River</collectingMunicipality>
,
<location LSID="urn:lsid:plazi:treatment:B76487A5FFE2650CFEB5C0369494F842:3A126068FFE26508FAD7C2FF947DFB50" box="[1285,1377,1184,1206]" latitude="38.776665" longLatPrecision="1" longitude="-75.96956" municipality="Choptank River" name="Preston" pageId="2" pageNumber="3" stateProvince="Maryland">Preston</location>
,
<collectingRegion box="[1401,1515,1184,1206]" country="United States of America" name="Maryland" pageId="2" pageNumber="3">Maryland</collectingRegion>
(
<geoCoordinate box="[852,992,1210,1236]" degrees="38" direction="north" minutes="46" orientation="latitude" pageId="2" pageNumber="3" precision="1" seconds="36.0" value="38.776665">
38°46
<emphasis box="[910,915,1210,1225]" italics="true" pageId="2" pageNumber="3">
<superScript attach="none" box="[910,915,1210,1225]" fontSize="6" pageId="2" pageNumber="3"></superScript>
</emphasis>
36.0
<emphasis box="[960,968,1210,1225]" italics="true" pageId="2" pageNumber="3">
<superScript attach="right" box="[960,968,1210,1225]" fontSize="6" pageId="2" pageNumber="3"></superScript>
</emphasis>
N
</geoCoordinate>
,
<geoCoordinate box="[1004,1147,1210,1236]" degrees="75" direction="west" minutes="58" orientation="longitude" pageId="2" pageNumber="3" precision="1" seconds="10.4" value="-75.96956">
75°58
<emphasis box="[1061,1066,1210,1225]" italics="true" pageId="2" pageNumber="3">
<superScript attach="none" box="[1061,1066,1210,1225]" fontSize="6" pageId="2" pageNumber="3"></superScript>
</emphasis>
10.4
<emphasis box="[1111,1119,1210,1225]" italics="true" pageId="2" pageNumber="3">
<superScript attach="right" box="[1111,1119,1210,1225]" fontSize="6" pageId="2" pageNumber="3"></superScript>
</emphasis>
W
</geoCoordinate>
)
</materialsCitation>
.
</paragraph>
<paragraph blockId="2.[845,1515,1008,1441]" pageId="2" pageNumber="3">
<materialsCitation collectionCode="USNM" location="Smithsonian Institution" municipality="National Museum of Natural History" pageId="2" pageNumber="3" specimenCode="USNM 1507371, USNM 1490787, 1490788, 1490789" specimenCount="1" stateProvince="Washington" typeStatus="holotype">
<emphasis box="[869,1032,1243,1264]" italics="true" pageId="2" pageNumber="3">
<typeStatus box="[869,919,1243,1264]" pageId="2" pageNumber="3">Type</typeStatus>
specimens:
</emphasis>
Histological sections of infected tissues stained with H&amp;E (
<specimenCode box="[976,1149,1272,1294]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="2" pageNumber="3" type="Museum">USNM 1507371</specimenCode>
and 1507381) and photosyntypes (
<specimenCode box="[852,1022,1302,1324]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="2" pageNumber="3" type="Museum">USNM 1490787</specimenCode>
,
<specimenCode box="[1031,1118,1302,1324]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="2" pageNumber="3" type="Museum">1490788</specimenCode>
,
<specimenCode box="[1127,1214,1302,1324]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="2" pageNumber="3" type="Museum">1490789</specimenCode>
, and 1490790) were deposited in the
<collectingMunicipality box="[930,1364,1331,1353]" pageId="2" pageNumber="3">National Museum of Natural History</collectingMunicipality>
,
<location LSID="urn:lsid:plazi:treatment:B76487A5FFE2650CFEB5C0369494F842:3A126068FFE26508FABBC36C92A0FA80" municipality="National Museum of Natural History" name="Smithsonian Institution" pageId="2" pageNumber="3" stateProvince="Washington">Smithsonian Institution</location>
,
<collectingRegion box="[968,1095,1360,1382]" country="United States of America" name="Washington" pageId="2" pageNumber="3">Washington</collectingRegion>
</materialsCitation>
, D.C.
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="3" type="etymology">
<paragraph blockId="2.[845,1515,1008,1441]" pageId="2" pageNumber="3">
<emphasis box="[869,986,1390,1411]" italics="true" pageId="2" pageNumber="3">Etymology:</emphasis>
The specific name is in honor of colleague and mentor Ana Baya.
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="7" pageId="2" pageNumber="3" type="discussion">
<paragraph blockId="2.[845,945,1477,1499]" box="[845,945,1477,1499]" pageId="2" pageNumber="3">
<heading bold="true" box="[845,945,1477,1499]" fontSize="9" level="3" pageId="2" pageNumber="3" reason="9">
<emphasis bold="true" box="[845,945,1477,1499]" pageId="2" pageNumber="3">Remarks</emphasis>
</heading>
</paragraph>
<paragraph blockId="2.[845,1516,1523,1956]" lastBlockId="5.[117,787,1495,1956]" lastPageId="5" lastPageNumber="6" pageId="2" pageNumber="3">
Phylogenetic analyses of other
<taxonomicName box="[1197,1327,1523,1545]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="undetermined">
<emphasis box="[1197,1275,1523,1544]" italics="true" pageId="2" pageNumber="3">Goussia</emphasis>
spp.
</taxonomicName>
using partial
<emphasis italics="true" pageId="2" pageNumber="3">18S rDNA</emphasis>
placed
<taxonomicName authority="Matsche &amp; Adams &amp; Blazer, 2019" authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[986,1074,1553,1574]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="bayae" status="n. sp.">
<emphasis box="[986,1074,1553,1574]" italics="true" pageId="2" pageNumber="3">G. bayae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1080,1135,1552,1574]" pageId="2" pageNumber="3" rank="species">n. sp.</taxonomicNameLabel>
in a clade of
<taxonomicName authorityName="Labbe" authorityYear="1896" box="[1270,1348,1553,1574]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="genus">
<emphasis box="[1270,1348,1553,1574]" italics="true" pageId="2" pageNumber="3">Goussia</emphasis>
</taxonomicName>
(
<figureCitation box="[1361,1422,1552,1574]" captionStart="Figure 5" captionStartId="5.[141,198,1381,1400]" captionTargetBox="[117,1488,202,1354]" captionTargetId="figure-292@5.[117,1488,202,1354]" captionTargetPageId="5" captionText="Figure 5. Bayesian phylogenetic relationships of fish-infecting Goussia and Choleoeimeria spp. based on partial 18S rDNA. Selected sequences represent different morphology types (epicellular, leucisci, dispersed, and nodular) defined by Rosenthal et al. (2016). Hammondia hammondi was used as an outgroup. Posterior probability is indicated at branch sites. Accession numbers follow species names in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7752631" httpUri="https://zenodo.org/record/7752631/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
), defined as epicellular (sensu
<bibRefCitation author="ROSENTHAL, B. M. &amp; MOREL, G. &amp; OSTOROS, AND &amp; K. MOLNAR" box="[1083,1336,1582,1604]" pageId="2" pageNumber="3" pagination="219 - 227" refId="ref6096" refString="ROSENTHAL, B. M., D. DUNAMS- MOREL, G. OSTOROS, AND K. MOLNAR. 2016. Coccidian parasites of fish encompass profound phylogenetic diversity and gave rise to each of the major parasitic groups in terrestrial vertebrates. Infection, Genetics and Evolution 40: 219 - 227." type="journal article" year="2016">Rosenthal et al., 2016</bibRefCitation>
).
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[1362,1514,1582,1603]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[1362,1514,1582,1603]" italics="true" pageId="2" pageNumber="3">Goussia bayae</emphasis>
</taxonomicName>
samples isolated from both the Choptank and Potomac were identical. The closest relative to
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[1170,1257,1641,1662]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[1170,1257,1641,1662]" italics="true" pageId="2" pageNumber="3">G. bayae</emphasis>
</taxonomicName>
by Bayesian analysis was
<taxonomicName box="[845,962,1670,1692]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="undetermined">
<emphasis box="[845,923,1670,1691]" italics="true" pageId="2" pageNumber="3">Goussia</emphasis>
sp.
</taxonomicName>
isolate I29 (MF468318), despite sharing only 95.4% identity. The isolate is morphologically undescribed, but was found in the intestine of a farmed European bass (
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Actinopterygii" family="Moronidae" genus="Dicentrarchus" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="labrax">
<emphasis italics="true" pageId="2" pageNumber="3">Dicentrarchus labrax</emphasis>
</taxonomicName>
) (
<bibRefCitation author="XAVIER, R. &amp; R. SEVERINO &amp; LOSADA, C. &amp; GESTAL, R. &amp; FREITAS, D. J. &amp; HARRIS, A. &amp; VERISSIMO, D. &amp; ROSADO, AND &amp; J. CABLE" box="[935,1150,1758,1780]" pageId="2" pageNumber="3" pagination="63" refId="ref6569" refString="XAVIER, R., R. SEVERINO, M. PEREZ- LOSADA, C. GESTAL, R. FREITAS, D. J. HARRIS, A. VERISSIMO, D. ROSADO, AND J. CABLE. 2018 a. Phylogenetic analysis of apicomplexan parasites infecting commercially valuable species from the northeast Atlantic reveals high levels of diversity and insights into the evolution of the group. Parasites and Vectors 11: 63. doi: 10.1186 / s 13071 - 018 - 2645 - 7." type="journal article" year="2018">Xavier et al., 2018a</bibRefCitation>
). The next nearest neighbors were also not fully morphologically described, but isolated from the heart and kidney of wild-caught chub mackerel
<taxonomicName authorityName="Houttuyn" authorityYear="1782" box="[1329,1514,1817,1838]" class="Actinopterygii" family="Scombridae" genus="Scomber" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="japonicas">
<emphasis box="[1329,1514,1817,1838]" italics="true" pageId="2" pageNumber="3">Scomber japonicas</emphasis>
</taxonomicName>
(MF468319, MF468320), the spleen and kidney of the bluestripe snapper
<taxonomicName baseAuthorityName="Forsskal" baseAuthorityYear="1775" box="[934,1110,1876,1897]" class="Actinopterygii" family="Lutjanidae" genus="Lutjanus" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="kasmira">
<emphasis box="[934,1110,1876,1897]" italics="true" pageId="2" pageNumber="3">Lutjanus kasmira</emphasis>
</taxonomicName>
(HM117907), and the liver and spleen of the orange goatfish
<taxonomicName baseAuthorityName="Steindachner" baseAuthorityYear="1900" box="[1111,1362,1905,1926]" class="Actinopterygii" family="Mullidae" genus="Mulloidichthys" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="pfluegeri">
<emphasis box="[1111,1362,1905,1926]" italics="true" pageId="2" pageNumber="3">Mulloidichthys pfluegeri</emphasis>
</taxonomicName>
(HM117908).
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[845,991,1934,1955]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[845,991,1934,1955]" italics="true" pageId="2" pageNumber="3">Goussia bayae</emphasis>
</taxonomicName>
shared 98.3% identity with
<taxonomicName box="[1297,1414,1934,1956]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="2" pageNumber="3" phylum="Miozoa" rank="species" species="undetermined">
<emphasis box="[1297,1375,1934,1955]" italics="true" pageId="2" pageNumber="3">Goussia</emphasis>
sp.
</taxonomicName>
from the chub mackerel and 98.4% identity with sequences from the bluestripe snapper and orange goatfish. Of the remaining
<taxonomicName authorityName="Labbe" authorityYear="1896" box="[709,787,1524,1545]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="genus">
<emphasis box="[709,787,1524,1545]" italics="true" pageId="5" pageNumber="6">Goussia</emphasis>
</taxonomicName>
in the epicellular clade, only
<taxonomicName box="[413,574,1554,1575]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="ameliae">
<emphasis box="[413,574,1554,1575]" italics="true" pageId="5" pageNumber="6">Goussia ameliae</emphasis>
</taxonomicName>
from alewives
<taxonomicName baseAuthorityName="Wilson" baseAuthorityYear="1811" class="Actinopterygii" family="Clupeidae" genus="Alosa" kingdom="Animalia" order="Clupeiformes" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="pseudoharengus">
<emphasis italics="true" pageId="5" pageNumber="6">Alosa pseudoharengus</emphasis>
</taxonomicName>
has been reported from North America (Lovy and Friend, 2015).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7752627" ID-Zenodo-Dep="7752627" httpUri="https://zenodo.org/record/7752627/files/figure.png" pageId="3" pageNumber="4" startId="3.[141,198,1535,1554]" targetBox="[118,1488,202,1508]" targetPageId="3">
<paragraph blockId="3.[117,1488,1535,1626]" pageId="3" pageNumber="4">
<emphasis bold="true" box="[141,223,1535,1554]" pageId="3" pageNumber="4">Figure 3.</emphasis>
Light micrographs of coccidia of
<taxonomicName authority="Matsche &amp; Adams &amp; Blazer, 2019" authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[559,689,1536,1555]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="3" pageNumber="4" phylum="Miozoa" rank="species" species="bayae" status="n. sp.">
<emphasis box="[559,689,1536,1555]" italics="true" pageId="3" pageNumber="4">Goussia bayae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[697,749,1535,1554]" pageId="3" pageNumber="4" rank="species">n. sp.</taxonomicNameLabel>
in the gallbladder of white perch,
<taxonomicName baseAuthorityName="Gmelin" baseAuthorityYear="1789" box="[1077,1247,1536,1555]" class="Actinopterygii" family="Moronidae" genus="Morone" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="americana">
<emphasis box="[1077,1247,1536,1555]" italics="true" pageId="3" pageNumber="4">Morone americana</emphasis>
</taxonomicName>
. (
<emphasis bold="true" box="[1268,1283,1535,1554]" pageId="3" pageNumber="4">A</emphasis>
) Severe coccidiosis in gallbladder. Note that numerous oocysts were removed with the bile for differential interference contrast microscopy before histological processing. (
<emphasis bold="true" box="[1466,1480,1559,1578]" pageId="3" pageNumber="4">B</emphasis>
) Developing stages of coccidia along epithelium (arrow) and sporulating oocysts in the lumen. (
<emphasis bold="true" box="[979,994,1583,1602]" pageId="3" pageNumber="4">C</emphasis>
) Microgamont (Mi), macrogamont (Ma), and meront (Me) epicellular to biliary epithelium, with oocysts (O) in the lumen.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7752629" ID-Zenodo-Dep="7752629" httpUri="https://zenodo.org/record/7752629/files/figure.png" pageId="4" pageNumber="5" startId="4.[168,225,1611,1630]" targetBox="[145,1515,202,1584]" targetPageId="4">
<paragraph blockId="4.[144,1515,1611,1702]" pageId="4" pageNumber="5">
<emphasis bold="true" box="[168,248,1611,1630]" pageId="4" pageNumber="5">Figure 4.</emphasis>
Light micrographs of coccidia of
<taxonomicName authority="Matsche &amp; Adams &amp; Blazer, 2019" authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[579,707,1612,1631]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="4" pageNumber="5" phylum="Miozoa" rank="species" species="bayae" status="n. sp.">
<emphasis box="[579,707,1612,1631]" italics="true" pageId="4" pageNumber="5">Goussia bayae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[713,764,1611,1630]" pageId="4" pageNumber="5" rank="species">n. sp.</taxonomicNameLabel>
in hepatic bile ducts of white perch,
<taxonomicName baseAuthorityName="Gmelin" baseAuthorityYear="1789" box="[1107,1275,1612,1631]" class="Actinopterygii" family="Moronidae" genus="Morone" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="americana">
<emphasis box="[1107,1275,1612,1631]" italics="true" pageId="4" pageNumber="5">Morone americana</emphasis>
</taxonomicName>
. (
<emphasis bold="true" box="[1295,1310,1611,1630]" pageId="4" pageNumber="5">A</emphasis>
) Developing stages of coccidia epicellular to biliary epithelium. (
<emphasis bold="true" box="[526,540,1635,1654]" pageId="4" pageNumber="5">B</emphasis>
) Microgamont (Mi), macrogamont (Ma), and meront (Me) along epithelium. (
<emphasis bold="true" box="[1258,1273,1635,1654]" pageId="4" pageNumber="5">C</emphasis>
) Longitudinal view of bile duct with developing coccidia along epithelium (arrow) and sporulating oocysts (O) in lumen. (
<emphasis bold="true" box="[1018,1034,1659,1678]" pageId="4" pageNumber="5">D</emphasis>
). Cross-section of enlarged bile duct with numerous developing and mature coccidia.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7752631" ID-Zenodo-Dep="7752631" httpUri="https://zenodo.org/record/7752631/files/figure.png" pageId="5" pageNumber="6" startId="5.[141,198,1381,1400]" targetBox="[117,1488,202,1354]" targetPageId="5">
<paragraph blockId="5.[117,1488,1381,1448]" pageId="5" pageNumber="6">
<emphasis bold="true" box="[141,223,1381,1400]" pageId="5" pageNumber="6">Figure 5.</emphasis>
Bayesian phylogenetic relationships of fish-infecting
<taxonomicName authorityName="Labbe" authorityYear="1896" box="[737,806,1381,1400]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="genus">
<emphasis box="[737,806,1381,1400]" italics="true" pageId="5" pageNumber="6">Goussia</emphasis>
</taxonomicName>
and
<taxonomicName box="[858,1033,1381,1400]" family="Eimeriidae" genus="Choleoeimeria" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="undetermined">
<emphasis box="[858,985,1381,1400]" italics="true" pageId="5" pageNumber="6">Choleoeimeria</emphasis>
spp.
</taxonomicName>
based on partial
<emphasis box="[1203,1302,1381,1400]" italics="true" pageId="5" pageNumber="6">18S rDNA</emphasis>
. Selected sequences represent different morphology types (epicellular, leucisci, dispersed, and nodular) defined by
<bibRefCitation author="ROSENTHAL, B. M. &amp; MOREL, G. &amp; OSTOROS, AND &amp; K. MOLNAR" box="[959,1165,1405,1424]" pageId="5" pageNumber="6" pagination="219 - 227" refId="ref6096" refString="ROSENTHAL, B. M., D. DUNAMS- MOREL, G. OSTOROS, AND K. MOLNAR. 2016. Coccidian parasites of fish encompass profound phylogenetic diversity and gave rise to each of the major parasitic groups in terrestrial vertebrates. Infection, Genetics and Evolution 40: 219 - 227." type="journal article" year="2016">Rosenthal et al. (2016)</bibRefCitation>
.
<taxonomicName authorityName="Frenkel &amp; Dubey" authorityYear="1975" box="[1173,1379,1405,1424]" family="Sarcocystidae" genus="Hammondia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="hammondi">
<emphasis box="[1173,1379,1405,1424]" italics="true" pageId="5" pageNumber="6">Hammondia hammondi</emphasis>
</taxonomicName>
was used as an outgroup. Posterior probability is indicated at branch sites. Accession numbers follow species names in parentheses.
</paragraph>
</caption>
<paragraph blockId="5.[117,787,1495,1956]" pageId="5" pageNumber="6">
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[141,287,1641,1662]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[141,287,1641,1662]" italics="true" pageId="5" pageNumber="6">Goussia bayae</emphasis>
</taxonomicName>
possesses a micropyle (
<figureCitation box="[539,612,1641,1663]" captionStart="Figure 1" captionStartId="2.[168,225,890,909]" captionTargetBox="[146,812,202,863]" captionTargetId="figure-545@2.[146,812,202,863]" captionTargetPageId="2" captionText="Figure 1. Photomicrographs of oocysts of Goussia bayae n. sp. from the gallbladder of white perch, Morone americana. (A) Nomarski differential interference contrast image of mature oocysts with micropyle (m) and refractile bodies (arrow) in sporocysts. (B) Histological preparation of oocysts with sporocysts containing paired sporozoites with densely stained nuclei (arrow)." figureDoi="http://doi.org/10.5281/zenodo.7752625" httpUri="https://zenodo.org/record/7752625/files/figure.png" pageId="5" pageNumber="6">Figs. 1</figureCitation>
,
<figureCitation box="[628,642,1641,1663]" captionStart="Figure 2" captionStartId="2.[869,926,781,800]" captionTargetBox="[853,1505,208,742]" captionTargetId="figure-603@2.[847,1513,202,754]" captionTargetPageId="2" captionText="Figure 2. Goussia bayae n. sp. Line drawing of a sporulated oocyst." figureDoi="http://doi.org/10.5281/zenodo.7752623" httpUri="https://zenodo.org/record/7752623/files/figure.png" pageId="5" pageNumber="6">2</figureCitation>
), whereas the oocysts of all other epicellular species compared with here lack a micropyle and are distinctly smaller. The closest in size to
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="bayae">
<emphasis italics="true" pageId="5" pageNumber="6">G. bayae</emphasis>
</taxonomicName>
is
<taxonomicName box="[206,314,1729,1750]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="ameliae">
<emphasis box="[206,314,1729,1750]" italics="true" pageId="5" pageNumber="6">G. ameliae</emphasis>
</taxonomicName>
(oocyst L X W: 18.6 X 14.1) from landlocked alewives and
<taxonomicName authorityName="Matsche, Adams &amp; Blazer" authorityYear="2019" box="[252,389,1759,1780]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="janae">
<emphasis box="[252,389,1759,1780]" italics="true" pageId="5" pageNumber="6">Goussia janae</emphasis>
</taxonomicName>
(oocyst L X W: 18.1 X 12.7). However,
<taxonomicName box="[117,233,1788,1809]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="ameliae">
<emphasis box="[117,233,1788,1809]" italics="true" pageId="5" pageNumber="6">G. ameliae</emphasis>
</taxonomicName>
has abundant sporocyst residuum and
<taxonomicName authorityName="Matsche, Adams &amp; Blazer" authorityYear="2019" box="[694,787,1788,1809]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="janae">
<emphasis box="[694,787,1788,1809]" italics="true" pageId="5" pageNumber="6">G. janae</emphasis>
</taxonomicName>
sporocysts are longer (L: 13.5) and more slender (W: 5, L/W: 2.7) compared with the sporocysts of
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[509,599,1846,1867]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[509,599,1846,1867]" italics="true" pageId="5" pageNumber="6">G. bayae</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="5.[117,787,1495,1956]" lastBlockId="5.[818,1488,1495,1956]" pageId="5" pageNumber="6">
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[141,290,1876,1897]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[141,290,1876,1897]" italics="true" pageId="5" pageNumber="6">Goussia bayae</emphasis>
</taxonomicName>
is morphologically distinct from 2 eimerians previously reported from white perch. Oocyst diameters of
<taxonomicName family="Eimeriidae" genus="Eimeria" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="glenorensis">
<emphasis italics="true" pageId="5" pageNumber="6">E. glenorensis</emphasis>
</taxonomicName>
(12.010.5) and
<taxonomicName box="[409,522,1934,1955]" family="Eimeriidae" genus="Eimeria" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="moronei">
<emphasis box="[409,522,1934,1955]" italics="true" pageId="5" pageNumber="6">E. moronei</emphasis>
</taxonomicName>
(8.07.2) were markedly smaller than
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[967,1063,1495,1516]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[967,1063,1495,1516]" italics="true" pageId="5" pageNumber="6">G. bayae</emphasis>
</taxonomicName>
and a micropyle was absent in both eimerians (Molnar and Fernando, 1974). Also, a Stieda body was present and refractile bodies were absent in
<taxonomicName box="[1279,1410,1554,1575]" family="Eimeriidae" genus="Eimeria" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="glenorensi">
<emphasis box="[1279,1410,1554,1575]" italics="true" pageId="5" pageNumber="6">E. glenorensi</emphasis>
</taxonomicName>
and
<taxonomicName family="Eimeriidae" genus="Eimeria" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="moronei">
<emphasis italics="true" pageId="5" pageNumber="6">E. moronei</emphasis>
</taxonomicName>
sporocysts.
</paragraph>
<paragraph blockId="5.[818,1488,1495,1956]" lastBlockId="6.[144,815,1576,1956]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
Histological examination of tissues indicated large numbers of coccidia in the gallbladder (
<figureCitation box="[1112,1176,1641,1663]" captionStart="Figure 3" captionStartId="3.[141,198,1535,1554]" captionTargetBox="[118,1488,202,1508]" captionTargetId="figure-1@3.[118,1488,202,1508]" captionTargetPageId="3" captionText="Figure 3. Light micrographs of coccidia of Goussia bayae n. sp. in the gallbladder of white perch, Morone americana. (A) Severe coccidiosis in gallbladder. Note that numerous oocysts were removed with the bile for differential interference contrast microscopy before histological processing. (B) Developing stages of coccidia along epithelium (arrow) and sporulating oocysts in the lumen. (C) Microgamont (Mi), macrogamont (Ma), and meront (Me) epicellular to biliary epithelium, with oocysts (O) in the lumen." figureDoi="http://doi.org/10.5281/zenodo.7752627" httpUri="https://zenodo.org/record/7752627/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
). Coccidian development was asynchronous and sporulation was endogenous. Meronts and gamonts were epicellular to the biliary epithelium, whereas oocysts were in the lumen (
<figureCitation box="[1102,1214,1729,1751]" captionStart="Figure 3" captionStartId="3.[141,198,1535,1554]" captionTargetBox="[118,1488,202,1508]" captionTargetId="figure-1@3.[118,1488,202,1508]" captionTargetPageId="3" captionText="Figure 3. Light micrographs of coccidia of Goussia bayae n. sp. in the gallbladder of white perch, Morone americana. (A) Severe coccidiosis in gallbladder. Note that numerous oocysts were removed with the bile for differential interference contrast microscopy before histological processing. (B) Developing stages of coccidia along epithelium (arrow) and sporulating oocysts in the lumen. (C) Microgamont (Mi), macrogamont (Ma), and meront (Me) epicellular to biliary epithelium, with oocysts (O) in the lumen." figureDoi="http://doi.org/10.5281/zenodo.7752627" httpUri="https://zenodo.org/record/7752627/files/figure.png" pageId="5" pageNumber="6">Fig. 3B, C</figureCitation>
). These stages of
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[1398,1487,1729,1750]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="5" pageNumber="6" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[1398,1487,1729,1750]" italics="true" pageId="5" pageNumber="6">G. bayae</emphasis>
</taxonomicName>
were also observed in hepatic bile ducts and common duct (
<figureCitation captionStart="Figure 4" captionStartId="4.[168,225,1611,1630]" captionTargetBox="[145,1515,202,1584]" captionTargetId="figure-1@4.[145,1515,202,1584]" captionTargetPageId="4" captionText="Figure 4. Light micrographs of coccidia of Goussia bayae n. sp. in hepatic bile ducts of white perch, Morone americana. (A) Developing stages of coccidia epicellular to biliary epithelium. (B) Microgamont (Mi), macrogamont (Ma), and meront (Me) along epithelium. (C) Longitudinal view of bile duct with developing coccidia along epithelium (arrow) and sporulating oocysts (O) in lumen. (D). Cross-section of enlarged bile duct with numerous developing and mature coccidia." figureDoi="http://doi.org/10.5281/zenodo.7752629" httpUri="https://zenodo.org/record/7752629/files/figure.png" pageId="5" pageNumber="6">Fig. 4</figureCitation>
). Meronts contained elongate merozoites that stained basophilic (
<figureCitation box="[826,917,1817,1839]" captionStart="Figure 3" captionStartId="3.[141,198,1535,1554]" captionTargetBox="[118,1488,202,1508]" captionTargetId="figure-1@3.[118,1488,202,1508]" captionTargetPageId="3" captionText="Figure 3. Light micrographs of coccidia of Goussia bayae n. sp. in the gallbladder of white perch, Morone americana. (A) Severe coccidiosis in gallbladder. Note that numerous oocysts were removed with the bile for differential interference contrast microscopy before histological processing. (B) Developing stages of coccidia along epithelium (arrow) and sporulating oocysts in the lumen. (C) Microgamont (Mi), macrogamont (Ma), and meront (Me) epicellular to biliary epithelium, with oocysts (O) in the lumen." figureDoi="http://doi.org/10.5281/zenodo.7752627" httpUri="https://zenodo.org/record/7752627/files/figure.png" pageId="5" pageNumber="6">Figs. 3C</figureCitation>
,
<figureCitation box="[931,962,1817,1839]" captionStart="Figure 4" captionStartId="4.[168,225,1611,1630]" captionTargetBox="[145,1515,202,1584]" captionTargetId="figure-1@4.[145,1515,202,1584]" captionTargetPageId="4" captionText="Figure 4. Light micrographs of coccidia of Goussia bayae n. sp. in hepatic bile ducts of white perch, Morone americana. (A) Developing stages of coccidia epicellular to biliary epithelium. (B) Microgamont (Mi), macrogamont (Ma), and meront (Me) along epithelium. (C) Longitudinal view of bile duct with developing coccidia along epithelium (arrow) and sporulating oocysts (O) in lumen. (D). Cross-section of enlarged bile duct with numerous developing and mature coccidia." figureDoi="http://doi.org/10.5281/zenodo.7752629" httpUri="https://zenodo.org/record/7752629/files/figure.png" pageId="5" pageNumber="6">4B</figureCitation>
). Microgamonts contained microgametocytes that were scattered throughout the cytoplasm or aligned near the wall (
<figureCitation box="[826,921,1875,1897]" captionStart="Figure 3" captionStartId="3.[141,198,1535,1554]" captionTargetBox="[118,1488,202,1508]" captionTargetId="figure-1@3.[118,1488,202,1508]" captionTargetPageId="3" captionText="Figure 3. Light micrographs of coccidia of Goussia bayae n. sp. in the gallbladder of white perch, Morone americana. (A) Severe coccidiosis in gallbladder. Note that numerous oocysts were removed with the bile for differential interference contrast microscopy before histological processing. (B) Developing stages of coccidia along epithelium (arrow) and sporulating oocysts in the lumen. (C) Microgamont (Mi), macrogamont (Ma), and meront (Me) epicellular to biliary epithelium, with oocysts (O) in the lumen." figureDoi="http://doi.org/10.5281/zenodo.7752627" httpUri="https://zenodo.org/record/7752627/files/figure.png" pageId="5" pageNumber="6">Figs. 3C</figureCitation>
,
<figureCitation box="[938,969,1875,1897]" captionStart="Figure 4" captionStartId="4.[168,225,1611,1630]" captionTargetBox="[145,1515,202,1584]" captionTargetId="figure-1@4.[145,1515,202,1584]" captionTargetPageId="4" captionText="Figure 4. Light micrographs of coccidia of Goussia bayae n. sp. in hepatic bile ducts of white perch, Morone americana. (A) Developing stages of coccidia epicellular to biliary epithelium. (B) Microgamont (Mi), macrogamont (Ma), and meront (Me) along epithelium. (C) Longitudinal view of bile duct with developing coccidia along epithelium (arrow) and sporulating oocysts (O) in lumen. (D). Cross-section of enlarged bile duct with numerous developing and mature coccidia." figureDoi="http://doi.org/10.5281/zenodo.7752629" httpUri="https://zenodo.org/record/7752629/files/figure.png" pageId="5" pageNumber="6">4B</figureCitation>
). Macrogamonts were larger than meronts and microgamonts, and contained a central eosinophilic nucleus and a coarse cytoplasm (
<figureCitation box="[1008,1097,1934,1956]" captionStart="Figure 3" captionStartId="3.[141,198,1535,1554]" captionTargetBox="[118,1488,202,1508]" captionTargetId="figure-1@3.[118,1488,202,1508]" captionTargetPageId="3" captionText="Figure 3. Light micrographs of coccidia of Goussia bayae n. sp. in the gallbladder of white perch, Morone americana. (A) Severe coccidiosis in gallbladder. Note that numerous oocysts were removed with the bile for differential interference contrast microscopy before histological processing. (B) Developing stages of coccidia along epithelium (arrow) and sporulating oocysts in the lumen. (C) Microgamont (Mi), macrogamont (Ma), and meront (Me) epicellular to biliary epithelium, with oocysts (O) in the lumen." figureDoi="http://doi.org/10.5281/zenodo.7752627" httpUri="https://zenodo.org/record/7752627/files/figure.png" pageId="5" pageNumber="6">Figs. 3C</figureCitation>
,
<figureCitation box="[1108,1139,1934,1956]" captionStart="Figure 4" captionStartId="4.[168,225,1611,1630]" captionTargetBox="[145,1515,202,1584]" captionTargetId="figure-1@4.[145,1515,202,1584]" captionTargetPageId="4" captionText="Figure 4. Light micrographs of coccidia of Goussia bayae n. sp. in hepatic bile ducts of white perch, Morone americana. (A) Developing stages of coccidia epicellular to biliary epithelium. (B) Microgamont (Mi), macrogamont (Ma), and meront (Me) along epithelium. (C) Longitudinal view of bile duct with developing coccidia along epithelium (arrow) and sporulating oocysts (O) in lumen. (D). Cross-section of enlarged bile duct with numerous developing and mature coccidia." figureDoi="http://doi.org/10.5281/zenodo.7752629" httpUri="https://zenodo.org/record/7752629/files/figure.png" pageId="5" pageNumber="6">4B</figureCitation>
). Although prevalence was 100%, the severity of infection was highly variable among fish, ranging from mild to severe. In fish with mild infections, not all intrahepatic bile duct branches contained coccidia, whereas duct branches in severely infected fish were generally enlarged with numerous coccidia. Developing stages of
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[606,702,1695,1716]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[606,702,1695,1716]" italics="true" pageId="6" pageNumber="7">G. bayae</emphasis>
</taxonomicName>
were not detected in the stomach, pyloric cecae, or intestine of fish. However, bile that passed through the ductus choledochus into the intestine contained sporulated oocysts of
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[614,703,1785,1806]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[614,703,1785,1806]" italics="true" pageId="6" pageNumber="7">G. bayae</emphasis>
</taxonomicName>
.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7752633" ID-Zenodo-Dep="7752633" httpUri="https://zenodo.org/record/7752633/files/figure.png" pageId="6" pageNumber="7" startId="6.[168,225,1454,1473]" targetBox="[144,1515,202,1427]" targetPageId="6">
<paragraph blockId="6.[144,1514,1454,1497]" pageId="6" pageNumber="7">
<emphasis bold="true" box="[168,247,1454,1473]" pageId="6" pageNumber="7">Figure 6.</emphasis>
Bayesian phylogenetic relationships of partitioned
<emphasis box="[724,819,1454,1473]" italics="true" pageId="6" pageNumber="7">18S rDNA</emphasis>
and cytochrome oxidase 1 (
<emphasis box="[1070,1110,1454,1473]" italics="true" pageId="6" pageNumber="7">COI</emphasis>
). Posterior probability is indicated at branch sites.
<taxonomicName baseAuthorityName="Nicolle &amp; Manceaux" baseAuthorityYear="1908" box="[195,366,1478,1497]" family="Sarcocystidae" genus="Toxoplasma" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="gondii">
<emphasis box="[195,366,1478,1497]" italics="true" pageId="6" pageNumber="7">Toxoplasma gondii</emphasis>
</taxonomicName>
served as an outgroup. Accession numbers follow species names in parentheses (
<emphasis box="[1112,1209,1478,1497]" italics="true" pageId="6" pageNumber="7">18S rDNA</emphasis>
,
<emphasis box="[1222,1262,1478,1497]" italics="true" pageId="6" pageNumber="7">COI</emphasis>
).
</paragraph>
</caption>
<paragraph blockId="6.[144,815,1576,1956]" lastBlockId="6.[845,1515,1576,1956]" pageId="6" pageNumber="7">
Molecular analyses also report the first mitochondrial DNA sequence from a
<taxonomicName box="[316,478,1845,1867]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="undetermined">
<emphasis box="[316,394,1845,1866]" italics="true" pageId="6" pageNumber="7">Goussia</emphasis>
species.
</taxonomicName>
The other evolutionarily distinct lineages of
<taxonomicName authorityName="Labbe" authorityYear="1896" box="[284,365,1875,1896]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="genus">
<emphasis box="[284,365,1875,1896]" italics="true" pageId="6" pageNumber="7">Goussia</emphasis>
</taxonomicName>
shown in
<figureCitation box="[504,603,1874,1896]" captionStart="Figure 5" captionStartId="5.[141,198,1381,1400]" captionTargetBox="[117,1488,202,1354]" captionTargetId="figure-292@5.[117,1488,202,1354]" captionTargetPageId="5" captionText="Figure 5. Bayesian phylogenetic relationships of fish-infecting Goussia and Choleoeimeria spp. based on partial 18S rDNA. Selected sequences represent different morphology types (epicellular, leucisci, dispersed, and nodular) defined by Rosenthal et al. (2016). Hammondia hammondi was used as an outgroup. Posterior probability is indicated at branch sites. Accession numbers follow species names in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7752631" httpUri="https://zenodo.org/record/7752631/files/figure.png" pageId="6" pageNumber="7">Figure 5</figureCitation>
do not yet have mitochondrial DNA sequences available. Bayesian analyses of the combination of gene markers
<emphasis box="[490,599,1934,1955]" italics="true" pageId="6" pageNumber="7">18S rDNA</emphasis>
and
<emphasis box="[651,696,1934,1955]" italics="true" pageId="6" pageNumber="7">COI</emphasis>
(
<figureCitation box="[711,773,1934,1956]" captionStart="Figure 6" captionStartId="6.[168,225,1454,1473]" captionTargetBox="[144,1515,202,1427]" captionTargetId="figure-235@6.[144,1516,202,1427]" captionTargetPageId="6" captionText="Figure 6. Bayesian phylogenetic relationships of partitioned 18S rDNA and cytochrome oxidase 1 (COI). Posterior probability is indicated at branch sites. Toxoplasma gondii served as an outgroup. Accession numbers follow species names in parentheses (18S rDNA, COI)." figureDoi="http://doi.org/10.5281/zenodo.7752633" httpUri="https://zenodo.org/record/7752633/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
), as well as of the partitioned mitochondrial genes (
<emphasis box="[1354,1399,1576,1597]" italics="true" pageId="6" pageNumber="7">COI</emphasis>
and
<emphasis box="[1458,1507,1576,1597]" italics="true" pageId="6" pageNumber="7">Cytb</emphasis>
) (
<figureCitation box="[853,918,1608,1630]" captionStart="Figure 7" captionStartId="7.[141,198,1495,1514]" captionTargetBox="[117,1487,202,1468]" captionTargetId="figure-229@7.[117,1488,202,1468]" captionTargetPageId="7" captionText="Figure 7. Bayesian phylogenetic relationships of mitochondrial genes cytochrome oxidase 1 (COI) and cytochrome oxidase b (Cytb). Babesia microti was used as the outgroup. Accession numbers follow species names in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7752635" httpUri="https://zenodo.org/record/7752635/files/figure.png" pageId="6" pageNumber="7">Fig. 7</figureCitation>
), definitively placed
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[1137,1229,1609,1630]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[1137,1229,1609,1630]" italics="true" pageId="6" pageNumber="7">G. bayae</emphasis>
</taxonomicName>
basal to
<taxonomicName box="[1333,1515,1608,1630]" family="Eimeriidae" genus="Choleoeimeria" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="undetermined">
<emphasis box="[1333,1476,1609,1630]" italics="true" pageId="6" pageNumber="7">Choleoeimeria</emphasis>
sp.
</taxonomicName>
and other
<taxonomicName box="[964,1125,1641,1663]" pageId="6" pageNumber="7">
<emphasis box="[964,1043,1641,1662]" italics="true" pageId="6" pageNumber="7">Eimeria</emphasis>
species
</taxonomicName>
with 0.999 bootstrap support. The nearest-neighbor by distance was
<taxonomicName box="[1218,1407,1673,1695]" family="Eimeriidae" genus="Choleoeimeria" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="undetermined">
<emphasis box="[1218,1361,1674,1695]" italics="true" pageId="6" pageNumber="7">Choleoeimeria</emphasis>
sp.
</taxonomicName>
, which is similar in size, sporocyst, sporozoite, and excystation structures, and in biliary tissue localization, but infects reptiles (
<bibRefCitation author="SZCZEPANIAK, K. O. &amp; K. TOMCZUK &amp; SZCZEPANIAK, AND &amp; W. LOPUSZYNSKI" pageId="6" pageNumber="7" pagination="681 - 685" refId="ref6152" refString="SZCZEPANIAK, K. O., K. TOMCZUK, A. LOJSZCZYK- SZCZEPANIAK, AND W. LOPUSZYNSKI. 2016. Reclassification of Eimeria pogonae Walden (2009) as Choleoeimeria pogonae comb. nov. (Apicomplexa: Eimeriidae). Parasitology Research 115: 681 - 685." type="journal article" year="2016">Szczepaniak et al., 2016</bibRefCitation>
). In comparing mitochondrial genes, haematozoeans
<taxonomicName authorityName="Starcovivi" authorityYear="1893" box="[845,923,1804,1825]" family="Babesiidae" genus="Babesia" kingdom="Chromista" order="Piroplasmida" pageId="6" pageNumber="7" phylum="Miozoa" rank="genus">
<emphasis box="[845,923,1804,1825]" italics="true" pageId="6" pageNumber="7">Babesia</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Bettencourt, Franca &amp; Borges" authorityYear="1907" box="[938,1027,1804,1825]" family="Theileriidae" genus="Theileria" kingdom="Chromista" order="Piroplasmida" pageId="6" pageNumber="7" phylum="Miozoa" rank="genus">
<emphasis box="[938,1027,1804,1825]" italics="true" pageId="6" pageNumber="7">Theileria</emphasis>
</taxonomicName>
, and
<taxonomicName authorityName="Marchiafava &amp; Celli" authorityYear="1885" box="[1088,1211,1804,1825]" family="Plasmodiidae" genus="Plasmodium" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="genus">
<emphasis box="[1088,1211,1804,1825]" italics="true" pageId="6" pageNumber="7">Plasmodium</emphasis>
</taxonomicName>
are ancestrally related to the coccidians (
<figureCitation box="[969,1036,1836,1858]" captionStart="Figure 7" captionStartId="7.[141,198,1495,1514]" captionTargetBox="[117,1487,202,1468]" captionTargetId="figure-229@7.[117,1488,202,1468]" captionTargetPageId="7" captionText="Figure 7. Bayesian phylogenetic relationships of mitochondrial genes cytochrome oxidase 1 (COI) and cytochrome oxidase b (Cytb). Babesia microti was used as the outgroup. Accession numbers follow species names in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7752635" httpUri="https://zenodo.org/record/7752635/files/figure.png" pageId="6" pageNumber="7">Fig. 7</figureCitation>
).
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[1059,1206,1837,1858]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[1059,1206,1837,1858]" italics="true" pageId="6" pageNumber="7">Goussia bayae</emphasis>
</taxonomicName>
is sister group and basal to
<taxonomicName box="[845,958,1869,1891]" family="Eimeriidae" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="family">Eimeriidae</taxonomicName>
, Choleoeimeria, and
<taxonomicName box="[1200,1340,1869,1891]" family="Sarcocystidae" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="family">Sarcocystidae</taxonomicName>
(
<figureCitation box="[1362,1432,1869,1891]" captionStart="Figure 7" captionStartId="7.[141,198,1495,1514]" captionTargetBox="[117,1487,202,1468]" captionTargetId="figure-229@7.[117,1488,202,1468]" captionTargetPageId="7" captionText="Figure 7. Bayesian phylogenetic relationships of mitochondrial genes cytochrome oxidase 1 (COI) and cytochrome oxidase b (Cytb). Babesia microti was used as the outgroup. Accession numbers follow species names in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7752635" httpUri="https://zenodo.org/record/7752635/files/figure.png" pageId="6" pageNumber="7">Fig. 7</figureCitation>
). Basal placement of
<taxonomicName authorityName="Matsche &amp; Adams &amp; Blazer" authorityYear="2019" box="[998,1093,1902,1923]" family="Eimeriidae" genus="Goussia" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="species" species="bayae">
<emphasis box="[998,1093,1902,1923]" italics="true" pageId="6" pageNumber="7">G. bayae</emphasis>
</taxonomicName>
to
<taxonomicName box="[1142,1252,1901,1923]" family="Eimeriidae" kingdom="Chromista" order="Eucoccidiida" pageId="6" pageNumber="7" phylum="Miozoa" rank="family">Eimeriidae</taxonomicName>
was also supported by analysis of all 3 mitochondrial genes (data not shown).
</paragraph>
</subSubSection>
</treatment>
</document>