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<mods:title id="DB5CBEF82F7195F3CCF2707D9D70B49F">Guide to the trematodes (Platyhelminthes) that infect the California horn snail (Cerithideopsis californica: Potamididae: Gastropoda) as first intermediate host</mods:title>
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<treatment id="EF6AD377895B8B38FF39FD18FEBEF8B1" ID-DOI="http://doi.org/10.5281/zenodo.5658146" ID-GBIF-Taxon="160797573" ID-Zenodo-Dep="5658146" LSID="urn:lsid:plazi:treatment:EF6AD377895B8B38FF39FD18FEBEF8B1" httpUri="http://treatment.plazi.org/id/EF6AD377895B8B38FF39FD18FEBEF8B1" lastPageNumber="478" pageId="19" pageNumber="478">
<subSubSection id="2FD931EA895B8B38FF39FD18FE78FD55" pageId="19" pageNumber="478" type="nomenclature">
<paragraph id="677C6261895B8B38FF39FD18FDEAFD31" blockId="19.[151,559,657,719]" box="[151,559,657,684]" pageId="19" pageNumber="478">
<heading id="3C34D50D895B8B38FF39FD18FDEAFD31" bold="true" box="[151,559,657,684]" fontSize="11" level="1" pageId="19" pageNumber="478" reason="1">
<taxonomicName id="A0C319E2895B8B38FF39FD18FDEAFD31" authority="(Martin)" authorityName="Lafuente, Roca &amp; Carbonell" authorityYear="2000" baseAuthorityName="Martin" baseAuthorityYear="1950" box="[151,559,657,684]" class="Lecanoromycetes" family="Graphidaceae" genus="Acanthotrema" kingdom="Animalia" order="Ostropales" pageId="19" pageNumber="478" phylum="Ascomycota" rank="species" species="hancocki">
<emphasis id="55B7BE73895B8B38FF39FD18FDEAFD31" bold="true" box="[151,559,657,684]" pageId="19" pageNumber="478">
<emphasis id="55B7BE73895B8B38FF39FD18FE7FFD36" bold="true" box="[151,442,657,684]" italics="true" pageId="19" pageNumber="478">Acanthotrema hancocki</emphasis>
(Martin)
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="677C6261895B8B38FF39FD3CFE78FD55" blockId="19.[151,559,657,719]" box="[151,445,693,719]" pageId="19" pageNumber="478">
(10. Acha;
<figureCitation id="FFF87EE4895B8B38FEBDFD3CFE9AFD55" box="[275,351,693,719]" captionStart="FIGURE 1" captionStartId="3.[151,250,1428,1453]" captionTargetBox="[151,1436,522,1400]" captionTargetId="figure@3.[151,1436,522,1400]" captionTargetPageId="3" captionText="FIGURE 1. General characteristics of the parthenitae and cercariae of the trematodes infecting Cerithideopsis californica as first intermediate host. Species numbers and codes follow Table 1 and species accounts. Cercariae are all to scale, with additional magnified views of six small species (indicated by dashed lines). Note the oral stylets (presented in right lateral view) for Pruc and Smmi. Parthenitae are not to scale. Scale bars consistently indicate 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3586556" httpUri="https://zenodo.org/record/3586556/files/figure.png" pageId="19" pageNumber="478">Figs. 1</figureCitation>
,
<figureCitation id="FFF87EE4895B8B38FEC3FD3CFE71FD55" box="[365,436,693,719]" captionStart="FIGURES 4144" captionStartId="20.[151,264,1076,1101]" captionTargetBox="[164,1423,179,1049]" captionTargetId="figure@20.[164,1423,179,1049]" captionTargetPageId="20" captionText="FIGURES 4144. Acanthotrema hancocki (Acha). 41, Overview of a colony in a freshly deshelled, infected horn snail in sea water. The arrow indicates the colony, which is localized in the gonadal region. Scale bar = 1 cm. 42, Reproductive rediae, live, with developing cercariae, under coverslip pressure. Scale bar = 100 µm. 43, Soldier redia, live, under coverslip pressure. Scale bar = 50 µm. Base photo credit: Andrew Turner. 44, Cercariae, live, under coverslip pressure. Scale bar = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3586582" httpUri="https://zenodo.org/record/3586582/files/figure.png" pageId="19" pageNumber="478">4144</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="2FD931EA895B8B38FF39FD75FBDBFAA9" pageId="19" pageNumber="478" type="diagnosis">
<paragraph id="677C6261895B8B38FF39FD75FEEDFCC5" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
<emphasis id="55B7BE73895B8B38FF39FD75FED1FC8D" bold="true" box="[151,276,764,791]" pageId="19" pageNumber="478">Diagnosis:</emphasis>
<emphasis id="55B7BE73895B8B38FEB0FD74FE6CFC8D" box="[286,425,765,791]" italics="true" pageId="19" pageNumber="478">Parthenitae.</emphasis>
Colony comprised of active rediae, densely concentrated in snail gonad region. Rediae translucent white, grey, weak yellow, or colorless; ~
<quantity id="A03BCF84895B8B38FD4EFCA9FCBAFCA1" box="[736,895,800,827]" metricMagnitude="-1" metricUnit="m" metricValue="7.5" metricValueMax="10.0" metricValueMin="5.0" pageId="19" pageNumber="478" unit="mm" value="750.0" valueMax="1000.0" valueMin="500.0">5001000 µm</quantity>
long, elongate (length:width ~4:1 to 10:1), sausage-shaped.
</paragraph>
<paragraph id="677C6261895B8B38FF69FCE0FDB8FB8E" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
<emphasis id="55B7BE73895B8B38FF69FCE0FEF6FC19" box="[199,307,873,899]" italics="true" pageId="19" pageNumber="478">
<taxonomicName id="A0C319E2895B8B38FF69FCE0FEEAFC19" authorityName="O.F.Muller" authorityYear="1773" box="[199,303,873,899]" class="Trematoda" family="Colepidae" genus="Cercaria" kingdom="Animalia" order="Prostomatida" pageId="19" pageNumber="478" phylum="Platyhelminthes" rank="genus">Cercaria</taxonomicName>
.
</emphasis>
Body mostly translucent colorless; oculate; with oral sucker and no ventral sucker; with seven pairs of penetration glands, the bodies of which lie in a relatively compact cluster, anterior to the genital primordium and excretory bladder; body ~
<quantity id="A03BCF84895B8B38FE1BFC39FDCEFC51" box="[437,523,944,971]" metricMagnitude="-1" metricUnit="m" metricValue="1.75" pageId="19" pageNumber="478" unit="mm" value="175.0">175 µm</quantity>
long, much shorter than tail (&lt;1/2 length); tail with dorso-ventral fins (originating in middle third of tail length, extending around tail tip) and lateral fins (originating basally, next to cercaria body, and inserting in middle third of tail length).
</paragraph>
<paragraph id="677C6261895B8B38FF69FB95FDEDFBC1" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
<emphasis id="55B7BE73895B8B38FF69FB95FE6CFBAD" bold="true" box="[199,425,1052,1079]" pageId="19" pageNumber="478">
<taxonomicName id="A0C319E2895B8B38FF69FB95FEF5FBAD" authorityName="O.F.Muller" authorityYear="1773" box="[199,304,1052,1079]" class="Trematoda" family="Colepidae" genus="Cercaria" kingdom="Animalia" order="Prostomatida" pageId="19" pageNumber="478" phylum="Platyhelminthes" rank="genus">Cercaria</taxonomicName>
behavior:
</emphasis>
Fresh, emerged cercariae remain in water column, swim intermittently in short bursts, with periods of resting and slow sinking.
</paragraph>
<paragraph id="677C6261895B8B38FF69FBEDFBDBFAA9" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
<emphasis id="55B7BE73895B8B38FF69FBEDFE47FBE5" bold="true" box="[199,386,1124,1151]" pageId="19" pageNumber="478">Similar species:</emphasis>
Acha is most reliably and readily distinguished from Euca [11] by the position of the penetration gland bodies, which are readily observable with flattened cercariae at 100x on a compound scope (and even sometimes at the dissection scope). Although Acha does have wider lateral tail fins than Euca on average, there appears to be overlap; so, tail fin width is not a consistently reliable distinguishing trait.
<bibRefCitation id="03521F90895B8B38FBFBFB58FB36FB71" author="Martin, W. E." box="[1109,1267,1233,1259]" pageId="19" pageNumber="478" pagination="39 - 43" refId="ref22195" refString="Martin, W. E. (1972) An annotated key to the cercariae that develop in the snail Cerithidea californica. Bulletin of the Southern California Academy of Sciences, 71, 39 - 43." type="journal article" year="1972">Martin (1972)</bibRefCitation>
used the flame cell grouping to distinguish Acha from Euca (groups of 3 versus 2, respectively), but the flame cells are difficult to see, requiring leaving specimens on a slide for a while and
<date id="137D44A1895B8B38FC80FA90FCB1FAA9" box="[814,884,1305,1331]" pageId="19" pageNumber="478" value="1000-10">1000x</date>
magnification.
</paragraph>
</subSubSection>
<subSubSection id="2FD931EA895B8B38FF69FAB4FEBEF8B1" pageId="19" pageNumber="478" type="discussion">
<paragraph id="677C6261895B8B38FF69FAB4FB02FA7D" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
<emphasis id="55B7BE73895B8B38FF69FAB4FEF8FACD" bold="true" box="[199,317,1341,1367]" pageId="19" pageNumber="478">Remarks:</emphasis>
<bibRefCitation id="03521F90895B8B38FEEAFAB4FE35FACD" author="Martin, W. E." box="[324,496,1340,1367]" pageId="19" pageNumber="478" pagination="360 - 370" refId="ref21806" refString="Martin, W. E. (1950 b) Parastictodora hancocki n. gen., n. sp. (Trematoda: Heterophyidae). with observations on its life cycle. Journal of Parasitology, 36, 360 - 370. https: // doi. org / 10.2307 / 3273472" type="journal article" year="1950">Martin (1950b)</bibRefCitation>
documented the life cycle and described this species (as
<taxonomicName id="A0C319E2895B8B38FBC1FAB4FA40FACD" authorityName="Martin" authorityYear="1950" box="[1135,1413,1341,1367]" class="Trematoda" family="Heterophyidae" genus="Parastictodora" higherTaxonomySource="CoL" kingdom="Animalia" order="Plagiorchiida" pageId="19" pageNumber="478" phylum="Platyhelminthes" rank="species" species="hancocki">
<emphasis id="55B7BE73895B8B38FBC1FAB4FA40FACD" box="[1135,1413,1341,1367]" italics="true" pageId="19" pageNumber="478">Parastictodora hancocki</emphasis>
</taxonomicName>
). He described the mother sporocyst, rediae and cercariae from natural infections, and metacercariae and adults from experimentally infected second intermediate and final hosts. I suspect that cercariae of
<taxonomicName id="A0C319E2895B8B38FBF9FA0CFAA5FA05" authorityName="Lafuente, Roca &amp; Carbonell" authorityYear="2000" baseAuthorityName="Martin" baseAuthorityYear="1950" box="[1111,1376,1413,1439]" class="Lecanoromycetes" family="Graphidaceae" genus="Acanthotrema" kingdom="Animalia" order="Ostropales" pageId="19" pageNumber="478" phylum="Ascomycota" rank="species" species="hancocki">
<emphasis id="55B7BE73895B8B38FBF9FA0CFAA5FA05" box="[1111,1376,1413,1439]" italics="true" pageId="19" pageNumber="478">Acanthotrema hancocki</emphasis>
</taxonomicName>
were accidentally pooled with
<taxonomicName id="A0C319E2895B8B38FE1AFA20FD2FFA59" box="[436,746,1449,1475]" class="Trematoda" family="Heterophyidae" genus="Euhaplorchis" kingdom="Animalia" order="Plagiorchiida" pageId="19" pageNumber="478" phylum="Platyhelminthes" rank="species" species="californiensis">
<emphasis id="55B7BE73895B8B38FE1AFA20FD2FFA59" box="[436,746,1449,1475]" italics="true" pageId="19" pageNumber="478">Euhaplorchis californiensis</emphasis>
</taxonomicName>
to comprise
<bibRefCitation id="03521F90895B8B38FCD1FA20FB25FA59" author="Maxon, M. G. &amp; Pequegnat, W. E." box="[895,1248,1449,1475]" pageId="19" pageNumber="478" pagination="30 - 55" refId="ref22284" refString="Maxon, M. G. &amp; Pequegnat, W. E. (1949) Cercariae from upper Newport Bay. Journal of Entomology and Zoology, 41, 30 - 55." type="journal article" year="1949">Maxon and Pequegnats (1949)</bibRefCitation>
Pleurolophocercous I and pooled with
<taxonomicName id="A0C319E2895B8B38FE33FA44FD55FA7D" box="[413,656,1485,1511]" class="Trematoda" family="Heterophyidae" genus="Phocitremoides" kingdom="Animalia" order="Plagiorchiida" pageId="19" pageNumber="478" phylum="Platyhelminthes" rank="species" species="ovale">
<emphasis id="55B7BE73895B8B38FE33FA44FD55FA7D" box="[413,656,1485,1511]" italics="true" pageId="19" pageNumber="478">Phocitremoides ovale</emphasis>
</taxonomicName>
cercariae to comprise their Pleurolophocercous II.
</paragraph>
<paragraph id="677C6261895B8B38FF69FA78FC5BF9B5" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
This species has also been referred to as
<taxonomicName id="A0C319E2895B8B38FD3DFA78FCB1F991" box="[659,884,1521,1547]" class="Trematoda" family="Heterophyidae" genus="Stictodora" kingdom="Animalia" order="Plagiorchiida" pageId="19" pageNumber="478" phylum="Platyhelminthes" rank="species" species="hancocki">
<emphasis id="55B7BE73895B8B38FD3DFA78FCB1F991" box="[659,884,1521,1547]" italics="true" pageId="19" pageNumber="478">Stictodora hancocki</emphasis>
</taxonomicName>
in ecological and evolutionary papers, but, since
<bibRefCitation id="03521F90895B8B38FF39F99CFE4BF9B5" author="Lafuente, M. &amp; Roca, V. &amp; Carbonell, E." box="[151,398,1557,1583]" pageId="19" pageNumber="478" pagination="131 - 134" refId="ref21621" refString="Lafuente, M., Roca, V. &amp; Carbonell, E. (2000) Description of Acanthotrema armata n. sp. (Trematoda: Heterophyidae) from Larus audouinii (Aves: Laridae), with an amended diagnosis of the genus Acanthotrema Travassos, 1928. Systematic Parasitology, 45, 131 - 134. https: // doi. org / 10.1023 / A: 1006293611598" type="journal article" year="2000">
Lafuente
<emphasis id="55B7BE73895B8B38FEACF99FFEF6F9B5" box="[258,307,1557,1583]" italics="true" pageId="19" pageNumber="478">et al</emphasis>
. (2000)
</bibRefCitation>
, the appropriate genus has been
<taxonomicName id="A0C319E2895B8B38FD54F99CFC5DF9B5" box="[762,920,1557,1583]" class="Lecanoromycetes" family="Graphidaceae" genus="Acanthotrema" kingdom="Animalia" order="Ostropales" pageId="19" pageNumber="478" phylum="Ascomycota" rank="genus">
<emphasis id="55B7BE73895B8B38FD54F99CFC5DF9B5" box="[762,920,1557,1583]" italics="true" pageId="19" pageNumber="478">Acanthotrema</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="677C6261895B8B38FF69F9B0FD56F9ED" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
Mature, ripe colonies comprise ~18% the soft-tissue weight of an infected snail (summer-time estimate derived from information in [
<bibRefCitation id="03521F90895B8B38FE2DF9D4FDBAF9ED" author="Hechinger, R. F. &amp; Lafferty, K. D. &amp; Mancini III, F. T. &amp; Warner, R. R. &amp; Kuris, A. M." box="[387,639,1628,1655]" pageId="19" pageNumber="478" pagination="651 - 667" refId="ref20498" refString="Hechinger, R. F., Lafferty, K. D., Mancini III, F. T., Warner, R. R. &amp; Kuris, A. M. (2009) How large is the hand in the puppet? Ecological and evolutionary factors affecting body mass of 15 trematode parasitic castrators in their snail host. Evolutionary Ecology, 23, 651 - 667. https: // doi. org / 10.1007 / s 10682 - 008 - 9262 - 4" type="journal article" year="2009">
Hechinger
<emphasis id="55B7BE73895B8B38FDACF9D7FDF6F9ED" box="[514,563,1629,1655]" italics="true" pageId="19" pageNumber="478">et al</emphasis>
. 2009
</bibRefCitation>
]).
</paragraph>
<paragraph id="677C6261895B8B38FF69F908FA4CF901" blockId="19.[151,1437,764,1835]" box="[199,1417,1664,1692]" pageId="19" pageNumber="478">
Acha infection causes (stolen) snail bodies to grow over 1.5x faster than uninfected snails (
<bibRefCitation id="03521F90895B8B38FB6CF908FAB8F901" author="Hechinger, R. F." box="[1218,1405,1665,1691]" pageId="19" pageNumber="478" pagination="1 - 14" refId="ref20237" refString="Hechinger, R. F. (2010) Mortality affects adaptive allocation to growth and reproduction: field evidence from a guild of body snatchers. BMC Evolutionary Biology, 10 (136), 1 - 14. https: // doi. org / 10.1186 / 1471 - 2148 - 10 - 136" type="journal article" year="2010">Hechinger 2010</bibRefCitation>
).
</paragraph>
<paragraph id="677C6261895B8B38FF69F92CFC2CF925" blockId="19.[151,1437,764,1835]" box="[199,1001,1701,1727]" pageId="19" pageNumber="478">
This species has a caste of soldier rediae (
<bibRefCitation id="03521F90895B8B38FD35F92CFC18F925" author="Garcia-Vedrenne, A. E. &amp; Quintana, A. C. E. &amp; DeRogatis, A. M. &amp; Dover, C. M. &amp; Lopez, M. &amp; Kuris, A. M. &amp; Hechinger, R. F." box="[667,989,1701,1727]" pageId="19" pageNumber="478" pagination="41 - 50" refId="ref20013" refString="Garcia-Vedrenne, A. E., Quintana, A. C. E., DeRogatis, A. M., Dover, C. M., Lopez, M., Kuris, A. M. &amp; Hechinger, R. F. (2017) Trematodes with a reproductive division of labour: heterophyids also have a soldier caste and early infections reveal how colonies become structured. International Journal for Parasitology, 47, 41 - 50. https: // doi. org / 10.1016 / j. ijpara. 2016.10.003" type="journal article" year="2017">
Garcia-Vedrenne
<emphasis id="55B7BE73895B8B38FCCFF92FFC56F925" box="[865,915,1701,1727]" italics="true" pageId="19" pageNumber="478">et al</emphasis>
. 2017
</bibRefCitation>
).
</paragraph>
<paragraph id="677C6261895B8B38FF69F940FEBEF8B1" blockId="19.[151,1437,764,1835]" pageId="19" pageNumber="478">
Using Acha (reported as Euca, see Euca remarks) from Bolinas Lagoon (central
<collectingRegion id="A507AC83895B8B38FBE5F940FB04F97E" box="[1099,1217,1737,1764]" country="United States of America" name="California" pageId="19" pageNumber="478">California</collectingRegion>
),
<bibRefCitation id="03521F90895B8B38FB7AF940FF27F89D" author="Koprivnikar, J. &amp; Lim, D. &amp; Fu, C. &amp; Brack, S. H. M." pageId="19" pageNumber="478" pagination="1167 - 1177" refId="ref21052" refString="Koprivnikar, J., Lim, D., Fu, C. &amp; Brack, S. H. M. (2010) Effects of temperature, salinity, and pH on the survival and activity of marine cercariae. Parasitology Research, 106, 1167 - 1177. https: // doi. org / 10.1007 / s 00436 - 010 - 1779 - 0" type="journal article" year="2010">
Koprivnikar
<emphasis id="55B7BE73895B8B38FACAF943FA50F979" box="[1380,1429,1737,1763]" italics="true" pageId="19" pageNumber="478">et al</emphasis>
. (2010)
</bibRefCitation>
performed laboratory experiments examining the effects of salinity, temperature, and pH on cercaria survivorship and activity.
</paragraph>
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</treatment>
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