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<mods:title id="1895551C2C3F1785351056B90E9FF2D9">Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition</mods:title>
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<paragraph id="8BD7AC55D30CFFFDFF7F41DE9E1C5506" blockId="20.[151,1039,1770,1796]" box="[151,1039,1770,1796]" pageId="20" pageNumber="21">
<heading id="D09F1B39D30CFFFDFF7F41DE9E1C5506" bold="true" box="[151,1039,1770,1796]" fontSize="11" level="1" pageId="20" pageNumber="21" reason="1">
<emphasis id="B91C7047D30CFFFDFF7F41DE9E1C5506" bold="true" box="[151,1039,1770,1796]" pageId="20" pageNumber="21">
Comments on
<taxonomicName id="4C68D7D6D30CFFFDFEA441DE980E5506" ID-CoL="7Q57J" box="[332,541,1770,1796]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30CFFFDFEA441DE980E5506" bold="true" box="[332,541,1770,1796]" italics="true" pageId="20" pageNumber="21">Trimusculotrema</emphasis>
</taxonomicName>
species and affinities with entobdellines
</emphasis>
</heading>
</paragraph>
<paragraph id="8BD7AC55D30CFFFCFF7F40069B8D5000" blockId="20.[151,1437,1842,2018]" lastBlockId="21.[151,1437,152,850]" lastPageId="21" lastPageNumber="22" pageId="20" pageNumber="21">
<bibRefCitation id="EFF9D1A4D30CFFFDFF7F40069B9E554E" box="[151,397,1842,1868]" pageId="20" pageNumber="27" refString="Bullard, S. A., Payne, R. R. &amp; Braswell, J. S. (2004) New genus with two new species of capsalid monogeneans from dasyatids in the Gulf of California. Journal of Parasitology, 90, 1412 - 1427." type="journal article">
Bullard
<emphasis id="B91C7047D30CFFFDFF1140079B3E554E" box="[249,301,1843,1868]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. (2004)
</bibRefCitation>
drew attention to strong morphological similarities between species of
<taxonomicName id="4C68D7D6D30CFFFDFB0440079F8E554E" box="[1260,1437,1843,1868]" class="Monogenea" family="Capsalidae" genus="Listrocephalos" kingdom="Animalia" order="Capsalidea" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30CFFFDFB0440079F8E554E" box="[1260,1437,1843,1868]" italics="true" pageId="20" pageNumber="21">Listrocephalos</emphasis>
</taxonomicName>
which they assigned to the Entobdellinae and
<taxonomicName id="4C68D7D6D30CFFFDFD09406C99A55573" box="[737,950,1880,1905]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="20" pageNumber="21" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30CFFFDFD09406C99A55573" box="[737,950,1880,1905]" italics="true" pageId="20" pageNumber="21">Trimusculotrema</emphasis>
</taxonomicName>
which at the time was assigned to the Benedeniinae after
<bibRefCitation id="EFF9D1A4D30CFFFDFE97404998F25595" author="Whittington" box="[383,737,1917,1943]" pageId="20" pageNumber="29" refString="Whittington, I. D. &amp; Barton, D. P. (1990) A new genus of monogenean parasites (Capsalidae: Benedeniinae) from stingrays (Rajiformes: Dasyatidae) with a description of a new species from the long-tailed stingray Himantura uarnak Forsskal from Queensland, Australia. Journal of Natural History, 24, 327 - 340." type="journal article" year="1990">Whittington &amp; Barton (1990)</bibRefCitation>
. Common morphology included a circular papillate haptor lacking a marginal valve, reduced non-overlapping median haptoral sclerites, separate male and female [uterine] genital pores and a dorsal uterine pore. Despite these anatomical parallels,
<bibRefCitation id="EFF9D1A4D30CFFFDFB4940FC9F8F55E0" box="[1185,1436,1992,2018]" pageId="20" pageNumber="27" refString="Bullard, S. A., Payne, R. R. &amp; Braswell, J. S. (2004) New genus with two new species of capsalid monogeneans from dasyatids in the Gulf of California. Journal of Parasitology, 90, 1412 - 1427." type="journal article">
Bullard
<emphasis id="B91C7047D30CFFFDFAED40FC9F2855E3" box="[1285,1339,1992,2017]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. (2004)
</bibRefCitation>
wrestled with arguments about whether the structural similarities between these taxa were due to convergent evolution imposed by strong ecological constraints (e.g. similarities in life-style resulting from inhabiting similar sites on similar host species) or a result of common ancestry (i.e. shared characters by phylogenetic descent).
<bibRefCitation id="EFF9D1A4D30DFFFCFEE4463C98145320" box="[268,519,264,290]" pageId="21" pageNumber="27" refString="Bullard, S. A., Payne, R. R. &amp; Braswell, J. S. (2004) New genus with two new species of capsalid monogeneans from dasyatids in the Gulf of California. Journal of Parasitology, 90, 1412 - 1427." type="journal article">
Bullard
<emphasis id="B91C7047D30DFFFCFE98463C9BB45323" box="[368,423,264,289]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. (2004)
</bibRefCitation>
were conservative and chose not to infer a close phylogenetic relationship between
<taxonomicName id="4C68D7D6D30DFFFCFEED46199BC65344" box="[261,469,301,326]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFEED46199BC65344" box="[261,469,301,326]" italics="true" pageId="21" pageNumber="22">Trimusculotrema</emphasis>
</taxonomicName>
and
<taxonomicName id="4C68D7D6D30DFFFCFDFA461998D45344" box="[530,711,301,326]" class="Monogenea" family="Capsalidae" genus="Listrocephalos" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFDFA461998D45344" box="[530,711,301,326]" italics="true" pageId="21" pageNumber="22">Listrocephalos</emphasis>
</taxonomicName>
. The present study, part of a larger initiative to establish a phylogenetic framework for the
<taxonomicName id="4C68D7D6D30DFFFCFDF5466698B2536E" box="[541,673,338,364]" class="Monogenea" family="Capsalidae" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="family">Capsalidae</taxonomicName>
using data from morphological and, where possible, molecular genetic assessments of many capsalid species, formally unites these taxa in the Entobdellinae together with
<taxonomicName id="4C68D7D6D30DFFFCFF7F46A99B2A53B4" box="[151,313,413,438]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFF7F46A99B2A53B4" box="[151,313,413,438]" italics="true" pageId="21" pageNumber="22">Benedeniella</emphasis>
</taxonomicName>
species. Ongoing work based on comparative anatomy and molecular analyses will either support or refute the decision made here about the subfamilial status of
<taxonomicName id="4C68D7D6D30DFFFCFBE246F69ECB53D9" box="[1034,1240,450,475]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFBE246F69ECB53D9" box="[1034,1240,450,475]" italics="true" pageId="21" pageNumber="22">Trimusculotrema</emphasis>
</taxonomicName>
based currently solely on morphology.
</paragraph>
<paragraph id="8BD7AC55D30DFFFCFF2E453998765150" blockId="21.[151,1437,152,850]" pageId="21" pageNumber="22">
The anterior attachment organs of
<taxonomicName id="4C68D7D6D30DFFFCFD9C453999285024" box="[628,827,525,550]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFD9C453999285024" box="[628,827,525,550]" italics="true" pageId="21" pageNumber="22">Trimusculotrema</emphasis>
</taxonomicName>
species differ from those of species in the other entobdelline genera,
<taxonomicName id="4C68D7D6D30DFFFCFE42450698405049" box="[426,595,562,587]" class="Monogenea" family="Capsalidae" genus="Benedeniella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFE42450698405049" box="[426,595,562,587]" italics="true" pageId="21" pageNumber="22">Benedeniella</emphasis>
</taxonomicName>
,
<taxonomicName id="4C68D7D6D30DFFFCFD8D450699365049" box="[613,805,562,587]" class="Monogenea" family="Capsalidae" genus="Branchobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFD8D450699365049" box="[613,805,562,587]" italics="true" pageId="21" pageNumber="22">Branchobdella</emphasis>
</taxonomicName>
,
<taxonomicName id="4C68D7D6D30DFFFCFCDF450699D65049" box="[823,965,562,587]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFCDF450699D65049" box="[823,965,562,587]" italics="true" pageId="21" pageNumber="22">Entobdella</emphasis>
</taxonomicName>
,
<taxonomicName id="4C68D7D6D30DFFFCFC3F45069E845049" box="[983,1175,562,587]" class="Monogenea" family="Capsalidae" genus="Listrocephalos" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFC3F45069E845049" box="[983,1175,562,587]" italics="true" pageId="21" pageNumber="22">Listrocephalos</emphasis>
</taxonomicName>
,
<taxonomicName id="4C68D7D6D30DFFFCFB4145069F765049" box="[1193,1381,562,587]" class="Monogenea" family="Capsalidae" genus="Neoentobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFB4145069F765049" box="[1193,1381,562,587]" italics="true" pageId="21" pageNumber="22">Neoentobdella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C68D7D6D30DFFFCFF7F456C9B7D5073" box="[151,366,600,625]" class="Monogenea" family="Capsalidae" genus="Pseudoentobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFF7F456C9B7D5073" box="[151,366,600,625]" italics="true" pageId="21" pageNumber="22">Pseudoentobdella</emphasis>
</taxonomicName>
, because no ventral grooves, rays, troughs or divided adhesive pads have been reported. Instead,
<taxonomicName id="4C68D7D6D30DFFFCFEEE45499BF35094" box="[262,480,637,662]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFEEE45499BF35094" box="[262,480,637,662]" italics="true" pageId="21" pageNumber="22">Trimusculotrema</emphasis>
</taxonomicName>
species possess anterior attachment organs that appear to bear no ventral specialisations other than the observation by
<bibRefCitation id="EFF9D1A4D30DFFFCFD3045969E5E50BE" author="Whittington" box="[728,1101,674,700]" pageId="21" pageNumber="29" refString="Whittington, I. D. &amp; Barton, D. P. (1990) A new genus of monogenean parasites (Capsalidae: Benedeniinae) from stingrays (Rajiformes: Dasyatidae) with a description of a new species from the long-tailed stingray Himantura uarnak Forsskal from Queensland, Australia. Journal of Natural History, 24, 327 - 340." type="journal article" year="1990">Whittington &amp; Barton (1990)</bibRefCitation>
that the anterior region in
<taxonomicName id="4C68D7D6D30DFFFCFF7F45FC993550E0" authority="Whittington &amp; Barton, 1990" authorityName="Whittington &amp; Barton" authorityYear="1990" box="[151,806,712,738]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="species" species="uarnaki">
<emphasis id="B91C7047D30DFFFCFF7F45FC9BDB50E3" box="[151,456,712,737]" italics="true" pageId="21" pageNumber="22">Trimusculotrema uarnaki</emphasis>
<bibRefCitation id="EFF9D1A4D30DFFFCFE3845FC993550E0" author="Whittington" box="[464,806,712,738]" pageId="21" pageNumber="29" refString="Whittington, I. D. &amp; Barton, D. P. (1990) A new genus of monogenean parasites (Capsalidae: Benedeniinae) from stingrays (Rajiformes: Dasyatidae) with a description of a new species from the long-tailed stingray Himantura uarnak Forsskal from Queensland, Australia. Journal of Natural History, 24, 327 - 340." type="journal article" year="1990">Whittington &amp; Barton, 1990</bibRefCitation>
</taxonomicName>
was rough but the posterior region was relatively smooth (see their fig. 10). This feature of
<taxonomicName id="4C68D7D6D30DFFFCFD6345D999415104" box="[651,850,749,774]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFD6345D999415104" box="[651,850,749,774]" italics="true" pageId="21" pageNumber="22">Trimusculotrema</emphasis>
</taxonomicName>
species deserves further study. Like species of all entobdelline genera except
<taxonomicName id="4C68D7D6D30DFFFCFE31442698945129" box="[473,647,786,811]" class="Monogenea" family="Capsalidae" genus="Branchobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFE31442698945129" box="[473,647,786,811]" italics="true" pageId="21" pageNumber="22">Branchobdella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C68D7D6D30DFFFCFD29442699515129" box="[705,834,786,811]" class="Monogenea" family="Capsalidae" genus="Entobdella" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFD29442699515129" box="[705,834,786,811]" italics="true" pageId="21" pageNumber="22">Entobdella</emphasis>
</taxonomicName>
,
<taxonomicName id="4C68D7D6D30DFFFCFCB844269E045129" box="[848,1047,786,811]" class="Monogenea" family="Capsalidae" genus="Trimusculotrema" kingdom="Animalia" order="Capsalidea" pageId="21" pageNumber="22" phylum="Platyhelminthes" rank="genus">
<emphasis id="B91C7047D30DFFFCFCB844269E045129" box="[848,1047,786,811]" italics="true" pageId="21" pageNumber="22">Trimusculotrema</emphasis>
</taxonomicName>
species principally parasitise the skin surfaces of marine elasmobranchs.
</paragraph>
</subSubSection>
</treatment>
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