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<mods:title id="9EEDB9DD53015CDC6B20D1C5B53A87CE">Hidden diversity of Nycteribiidae (Diptera) bat flies from the Malagasy region and insights on host-parasite interactions</mods:title>
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Phylogeny of
<taxonomicName id="1A8F4D15FF91EB3062DCFA384D6DAF77" box="[961,1102,1517,1540]" class="Insecta" family="Nycteribiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="family">Nycteribiidae</taxonomicName>
bat flies
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Bayesian inference based on mitochondrial sequences revealed that bat fly species parasitizing frugivorous (
<taxonomicName id="1A8F4D15FF91EB306447F9F84AA2AF16" authorityName="Gray" authorityYear="1821" class="Mammalia" family="Pteropodidae" kingdom="Animalia" order="Chiroptera" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Pteropodidae</taxonomicName>
) and insectivorous (
<taxonomicName id="1A8F4D15FF91EB30657BF9984C31AF16" box="[1126,1298,1613,1637]" class="Mammalia" family="Vespertilionidae" kingdom="Animalia" order="Chiroptera" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Vespertilionidae</taxonomicName>
,
<taxonomicName id="1A8F4D15FF91EB30643EF9984A71AFF6" class="Mammalia" family="Miniopteridae" kingdom="Animalia" order="Chiroptera" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Miniopteridae</taxonomicName>
) bats form two distinct well-supported monophyletic clades (posterior probability, pp = 1) (
<figureCitation id="45B42A13FF91EB3065ABF9584DCCAFD6" box="[1206,1263,1677,1701]" captionStart="Fig" captionStartId="2.[167,198,1449,1468]" captionTargetBox="[151,1437,236,1561]" captionTargetId="figure-197@2.[339,1248,253,1432]" captionTargetPageId="2" captionText="Fig. 1 Phylogenetic relationships of nycteribiid flies parasitizing bats on Madagascar and the Comoros. The analysis was carried out using Bayesian inference under the GTR + I + G substitution model. Values in the nodes represent posterior probability, and only pp.&gt; 0.80 are mentioned. Branchlets in red represent bat flies from the Malagasy Region. Generic abbreviations under the nycteribiid column include: B., Basilia; C., Cyclopodia; E., Eucampsipoda; N., Nycteribia; P., Penicillidia" pageId="1" pageNumber="2">Fig. 1</figureCitation>
). Flies parasitizing frugivorous bats and belonging to the subfamily
<taxonomicName id="1A8F4D15FF91EB306445F9784AA5AF96" authorityName="Theodor" authorityYear="1967" pageId="1" pageNumber="2" rank="subFamily" subFamily="Cyclopodiinae">Cyclopodiinae</taxonomicName>
[
<bibRefCitation id="B91E4B67FF91EB306285F9184A97AF96" author="Maa TC" box="[920,948,1741,1765]" pageId="1" pageNumber="2" pagination="377 - 86" refId="ref5927" refString="28. Maa TC. An interim world list of batflies. J Med Entomol. 1965; 1: 377 - 86." type="journal article" year="1965">28</bibRefCitation>
] are divided into two well-supported clades (pp = 1) comprised of the genera
<taxonomicName id="1A8F4D15FF91EB30659AF9394DD8AE77" authorityName="Kolenati" authorityYear="1863" box="[1159,1275,1772,1796]" class="Insecta" family="Hippoboscidae" genus="Cyclopodia" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">
<emphasis id="EFFBEA84FF91EB30659AF9394DD8AE77" box="[1159,1275,1772,1796]" italics="true" pageId="1" pageNumber="2">Cyclopodia</emphasis>
</taxonomicName>
and
<taxonomicName id="1A8F4D15FF91EB306433F9384A41AE57" authorityName="Kolenati" authorityYear="1857" class="Insecta" family="Hippoboscidae" genus="Eucampsipoda" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">
<emphasis id="EFFBEA84FF91EB306433F9384A41AE57" italics="true" pageId="1" pageNumber="2">Eucampsipoda</emphasis>
</taxonomicName>
found on
<taxonomicName id="1A8F4D15FF91EB3062FEF8D94D99AE57" baseAuthorityName="Pollen in Schlegel &amp; Pollen" baseAuthorityYear="1866" box="[995,1210,1804,1828]" class="Mammalia" family="Pteropodidae" genus="Eidolon" kingdom="Animalia" order="Chiroptera" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="dupreanum">
<emphasis id="EFFBEA84FF91EB3062FEF8D94D99AE57" box="[995,1210,1804,1828]" italics="true" pageId="1" pageNumber="2">Eidolon dupreanum</emphasis>
</taxonomicName>
and
<taxonomicName id="1A8F4D15FF91EB3065E1F8D84CBAAE56" box="[1276,1433,1805,1829]" class="Mammalia" family="Pteropodidae" genus="Rousettus" kingdom="Animalia" order="Chiroptera" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="EFFBEA84FF91EB3065E1F8D84C7CAE57" box="[1276,1375,1805,1828]" italics="true" pageId="1" pageNumber="2">Rousettus</emphasis>
spp.
</taxonomicName>
, respectively.
<taxonomicName id="1A8F4D15FF91EB3062A3F8F94D87AE37" authorityName="Meijere" authorityYear="1899" box="[958,1188,1836,1860]" class="Insecta" family="Hippoboscidae" genus="Cyclopodia" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="species" species="horsfieldi">
<emphasis id="EFFBEA84FF91EB3062A3F8F94D87AE37" box="[958,1188,1836,1860]" italics="true" pageId="1" pageNumber="2">Cyclopodia horsfieldi</emphasis>
</taxonomicName>
parasitizing
<taxonomicName id="1A8F4D15FF91EB30645FF8F84A94AE17" authorityName="Temminck" authorityYear="1853" class="Mammalia" family="Pteropodidae" genus="Pteropus" kingdom="Animalia" order="Chiroptera" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="hypomelanus">
<emphasis id="EFFBEA84FF91EB30645FF8F84A94AE17" italics="true" pageId="1" pageNumber="2">Pteropus hypomelanus</emphasis>
</taxonomicName>
from
<collectingCountry id="A5987606FF91EB30651AF8984D41AE16" box="[1031,1122,1869,1893]" name="Malaysia" pageId="1" pageNumber="2">Malaysia</collectingCountry>
falls within the first clade. Molecular data presented herein from flies collected on insectivorous bats from
<collectingCountry id="A5987606FF91EB306558F8584DE1AED6" box="[1093,1218,1933,1957]" name="Madagascar" pageId="1" pageNumber="2">Madagascar</collectingCountry>
and the
<collectingCountry id="A5987606FF91EB306426F8584CBFAED6" box="[1339,1436,1933,1957]" name="Comoros" pageId="1" pageNumber="2">Comoros</collectingCountry>
Archipelago and belonging to the subfamily
<taxonomicName id="1A8F4D15FF92EB336366F9B84A25AFF6" box="[635,774,1645,1669]" pageId="2" pageNumber="3" rank="subFamily" subFamily="Nycteribiinae">Nycteribiinae</taxonomicName>
[
<bibRefCitation id="B91E4B67FF92EB336183F9584998AFD6" author="Westwood JO" box="[158,187,1677,1701]" pageId="2" pageNumber="3" pagination="275 - 94" refId="ref5951" refString="29. Westwood JO. On Nycteribia, a genus of wingless insects. Trans Zool Soc London. 1835; 1: 275 - 94." type="journal article" year="1835">29</bibRefCitation>
] also yielded two well-supported monophyletic clades composed of
<taxonomicName id="1A8F4D15FF92EB336039F97948E2AFB6" box="[292,449,1708,1733]" class="Insecta" family="Hippoboscidae" genus="Nycteribia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="EFFBEA84FF92EB336039F97948B3AFB7" box="[292,400,1708,1732]" italics="true" pageId="2" pageNumber="3">Nycteribia</emphasis>
spp.
</taxonomicName>
and
<taxonomicName id="1A8F4D15FF92EB3360E8F9794BBEAFB6" box="[501,669,1708,1733]" class="Insecta" family="Hippoboscidae" genus="Penicillidia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="EFFBEA84FF92EB3360E8F9794B49AFB7" box="[501,618,1708,1732]" italics="true" pageId="2" pageNumber="3">Penicillidia</emphasis>
spp.
</taxonomicName>
, parasitizing
<taxonomicName id="1A8F4D15FF92EB3361DCF9184857AF96" box="[193,372,1741,1765]" class="Mammalia" family="Miniopteridae" genus="Miniopterus" kingdom="Animalia" order="Chiroptera" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="EFFBEA84FF92EB3361DCF918481CAF97" box="[193,319,1741,1764]" italics="true" pageId="2" pageNumber="3">Miniopterus</emphasis>
spp.
</taxonomicName>
, as well as
<taxonomicName id="1A8F4D15FF92EB3360ECF9184BAFAF97" baseAuthorityName="A. Smith" baseAuthorityYear="1834" box="[497,652,1740,1764]" class="Mammalia" family="Vespertilionidae" genus="Myotis" kingdom="Animalia" order="Chiroptera" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="goudoti">
<emphasis id="EFFBEA84FF92EB3360ECF9184BAFAF97" box="[497,652,1740,1764]" italics="true" pageId="2" pageNumber="3">Myotis goudoti</emphasis>
</taxonomicName>
. Both
<taxonomicName id="1A8F4D15FF92EB3363CBF9184832AE76" class="Insecta" family="Hippoboscidae" genus="Nycteribia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="EFFBEA84FF92EB3363CBF91849F8AE77" italics="true" pageId="2" pageNumber="3">Nycteribia</emphasis>
spp.
</taxonomicName>
and
<taxonomicName id="1A8F4D15FF92EB336051F93948D5AE76" box="[332,502,1772,1797]" class="Insecta" family="Hippoboscidae" genus="Penicillidia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="EFFBEA84FF92EB336051F93948E2AE77" box="[332,449,1772,1796]" italics="true" pageId="2" pageNumber="3">Penicillidia</emphasis>
spp.
</taxonomicName>
occurred on a variety of different host species and in certain cases, a single putative fly species was identified from several different bat species (
<tableCitation id="900D032DFF92EB336183F89849C9AE16" box="[158,234,1869,1893]" captionStart="Table 1" captionStartId="3.[151,210,749,770]" captionTargetPageId="3" captionText="Table 1 Nycteribiid species identified from different bat hosts from the Malagasy Region" pageId="2" pageNumber="3">Table 1</tableCitation>
). In addition, a separate and well-supported cluster of bat flies parasitizing Malagasy vespertilionids, specifically
<taxonomicName id="1A8F4D15FF92EB33618AF8594855AED7" authorityName="A. Milne-Edwards" authorityYear="1881" box="[151,374,1932,1956]" class="Mammalia" family="Vespertilionidae" genus="Scotophilus" kingdom="Animalia" order="Chiroptera" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="robustus">
<emphasis id="EFFBEA84FF92EB33618AF8594855AED7" box="[151,374,1932,1956]" italics="true" pageId="2" pageNumber="3">Scotophilus robustus</emphasis>
</taxonomicName>
,
<taxonomicName id="1A8F4D15FF92EB336090F8594B39AED7" authorityName="Goodman, Ratrimomanarivo &amp; Randrianandrianina" authorityYear="2006" box="[397,538,1932,1957]" class="Mammalia" family="Vespertilionidae" genus="Scotophilus" kingdom="Animalia" order="Chiroptera" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="marovaza">
<emphasis id="EFFBEA84FF92EB336090F85948BAAED0" box="[397,409,1932,1955]" italics="true" pageId="2" pageNumber="3">S</emphasis>
.
<emphasis id="EFFBEA84FF92EB3360ACF8584B39AED7" box="[433,538,1933,1956]" italics="true" pageId="2" pageNumber="3">marovaza</emphasis>
</taxonomicName>
, and
<taxonomicName id="1A8F4D15FF92EB336377F8594AB8AFF7" baseAuthorityName="Temminck" baseAuthorityYear="1840" box="[618,923,1644,1957]" class="Mammalia" family="Vespertilionidae" genus="Pipistrellus" isUncertain="true" kingdom="Animalia" order="Chiroptera" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="hesperidus">
<emphasis id="EFFBEA84FF92EB336377F8594BFEAED7" box="[618,733,1932,1956]" italics="true" pageId="2" pageNumber="3">Pipistrellus</emphasis>
cf.
<emphasis id="EFFBEA84FF92EB336230F9B94AB8AFF7" box="[813,923,1644,1668]" italics="true" pageId="2" pageNumber="3">hesperidus</emphasis>
</taxonomicName>
(denoted as
<taxonomicName id="1A8F4D15FF92EB33653EF9B94D86AFF6" box="[1059,1189,1644,1669]" class="Insecta" family="Hippoboscidae" genus="Basilia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="undefined-1">
<emphasis id="EFFBEA84FF92EB33653EF9B94D4FAFF7" box="[1059,1132,1644,1668]" italics="true" pageId="2" pageNumber="3">Basilia</emphasis>
sp.
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1
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3 in
</quantity>
</taxonomicName>
<figureCitation id="45B42A13FF92EB3365F8F9B84C3EAFF6" box="[1253,1309,1645,1669]" captionStart="Fig" captionStartId="2.[167,198,1449,1468]" captionTargetBox="[151,1437,236,1561]" captionTargetId="figure-197@2.[339,1248,253,1432]" captionTargetPageId="2" captionText="Fig. 1 Phylogenetic relationships of nycteribiid flies parasitizing bats on Madagascar and the Comoros. The analysis was carried out using Bayesian inference under the GTR + I + G substitution model. Values in the nodes represent posterior probability, and only pp.&gt; 0.80 are mentioned. Branchlets in red represent bat flies from the Malagasy Region. Generic abbreviations under the nycteribiid column include: B., Basilia; C., Cyclopodia; E., Eucampsipoda; N., Nycteribia; P., Penicillidia" pageId="2" pageNumber="3">Fig. 1</figureCitation>
) formed an independent, monophyletic group excluding
<taxonomicName id="1A8F4D15FF92EB336412F9584C4FAFD7" authorityName="Hurka" authorityYear="1969" box="[1295,1388,1676,1701]" class="Insecta" family="Hippoboscidae" genus="Basilia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="rybini">
<emphasis id="EFFBEA84FF92EB336412F9584C3DAFD7" box="[1295,1310,1677,1700]" italics="true" pageId="2" pageNumber="3">B</emphasis>
.
<emphasis id="EFFBEA84FF92EB336433F9594C4FAFD7" box="[1326,1388,1676,1700]" italics="true" pageId="2" pageNumber="3">rybini</emphasis>
</taxonomicName>
and
<taxonomicName id="1A8F4D15FF92EB336230F9784A80AFB7" authorityName="Theodor" authorityYear="1966" box="[813,931,1709,1733]" class="Insecta" family="Hippoboscidae" genus="Basilia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="truncata">
<emphasis id="EFFBEA84FF92EB336230F9784A1FAFB7" box="[813,828,1709,1732]" italics="true" pageId="2" pageNumber="3">B</emphasis>
.
<emphasis id="EFFBEA84FF92EB336255F9784A80AFB7" box="[840,931,1709,1732]" italics="true" pageId="2" pageNumber="3">truncata</emphasis>
</taxonomicName>
from
<collectingCountry id="A5987606FF92EB3362FEF9784D3CAFB6" box="[995,1055,1709,1733]" name="Japan" pageId="2" pageNumber="3">Japan</collectingCountry>
. In this phylogeny, the genus
<taxonomicName id="1A8F4D15FF92EB336449F9794CBFAFB7" box="[1364,1436,1708,1732]" class="Insecta" family="Hippoboscidae" genus="Basilia" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="EFFBEA84FF92EB336449F9794CBFAFB7" box="[1364,1436,1708,1732]" italics="true" pageId="2" pageNumber="3">Basilia</emphasis>
</taxonomicName>
is paraphyletic and contains previously unrecognized diversity. Based on the Kimura 2-parameter model (K2P,
<tableCitation id="900D032DFF92EB33645EF9384CB3AE76" box="[1347,1424,1773,1797]" captionStart="Table 2" captionStartId="4.[151,210,237,258]" captionTargetPageId="4" captionText="Table 2 Kimura 2-Parameter distances (in %, below diagonal) between groups as based on cox1 sequences (658 bp) and calculated using MEGA 6.0 [17] for nycteribiid species from the Malagasy Region; values above the diagonal represent the standard error (in %)" pageId="2" pageNumber="3">Table 2</tableCitation>
), the average genetic distance between nycteribiid species from the Malagasy Region ranged from 3.0 to 20.4%. Bat flies infecting
<taxonomicName id="1A8F4D15FF92EB3362A2F8994D13AE17" authorityName="A. Milne-Edwards" authorityYear="1881" box="[959,1072,1868,1893]" class="Mammalia" family="Vespertilionidae" genus="Scotophilus" kingdom="Animalia" order="Chiroptera" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="robustus">
<emphasis id="EFFBEA84FF92EB3362A2F8994AE8AE10" box="[959,971,1868,1891]" italics="true" pageId="2" pageNumber="3">S</emphasis>
.
<emphasis id="EFFBEA84FF92EB3362C5F8994D13AE17" box="[984,1072,1868,1892]" italics="true" pageId="2" pageNumber="3">robustus</emphasis>
</taxonomicName>
and
<taxonomicName id="1A8F4D15FF92EB33657BF8994DCBAE17" authorityName="Goodman, Ratrimomanarivo &amp; Randrianandrianina" authorityYear="2006" box="[1126,1256,1868,1893]" class="Mammalia" family="Vespertilionidae" genus="Scotophilus" kingdom="Animalia" order="Chiroptera" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="marovaza">
<emphasis id="EFFBEA84FF92EB33657BF8994D51AE10" box="[1126,1138,1868,1891]" italics="true" pageId="2" pageNumber="3">S</emphasis>
.
<emphasis id="EFFBEA84FF92EB336562F8984DCBAE17" box="[1151,1256,1869,1892]" italics="true" pageId="2" pageNumber="3">marovaza</emphasis>
</taxonomicName>
formed a monophyletic clade composed of two groups separated by a genetic distance of 12.1%, supporting the existence of two distinct species. Similarly, flies parasitizing
<taxonomicName id="1A8F4D15FF94EB35636DFCB94826AAD7" baseAuthorityName="Temminck" baseAuthorityYear="1840" class="Mammalia" family="Vespertilionidae" genus="Pipistrellus" isUncertain="true" kingdom="Animalia" order="Chiroptera" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="hesperidus">
<emphasis id="EFFBEA84FF94EB35636DFCB94BC0AAF7" box="[624,739,876,900]" italics="true" pageId="4" pageNumber="5">Pipistrellus</emphasis>
cf.
<emphasis id="EFFBEA84FF94EB35618AFC594826AAD7" box="[151,261,908,932]" italics="true" pageId="4" pageNumber="5">hesperidus</emphasis>
</taxonomicName>
were notably divergent (12.1 to 13.5%) from those found on
<taxonomicName id="1A8F4D15FF94EB356021FC7948C0AAB6" box="[316,483,940,965]" class="Mammalia" family="Vespertilionidae" genus="Scotophilus" kingdom="Animalia" order="Chiroptera" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="EFFBEA84FF94EB356021FC794892AAB7" box="[316,433,940,964]" italics="true" pageId="4" pageNumber="5">Scotophilus</emphasis>
spp.
</taxonomicName>
</paragraph>
</subSubSection>
</treatment>
</document>