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<mods:title id="DE5522A1774ABA6F48DA27589D59EB68">Philippine bats of the genus Kerivoula (Chiroptera: Vespertilionidae): Overview and assessment of variation in K. pellucida and K. whiteheadi</mods:title>
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<treatment id="5316885DF469FFFEFF0CD51CFE9CFE70" ID-DOI="http://doi.org/10.5281/zenodo.3811830" ID-GBIF-Taxon="163420726" ID-Zenodo-Dep="3811830" LSID="urn:lsid:plazi:treatment:5316885DF469FFFEFF0CD51CFE9CFE70" httpUri="http://treatment.plazi.org/id/5316885DF469FFFEFF0CD51CFE9CFE70" lastPageId="13" lastPageNumber="467" pageId="10" pageNumber="464">
<subSubSection id="93A56AC0F469FFF9FF0CD51CFDCFFBD1" box="[151,596,1083,1110]" pageId="10" pageNumber="464" type="nomenclature">
<paragraph id="DB00394BF469FFF9FF0CD51CFDCFFBD1" blockId="10.[151,1437,1083,2045]" box="[151,596,1083,1110]" pageId="10" pageNumber="464">
<heading id="80488E27F469FFF9FF0CD51CFDCFFBD1" bold="true" box="[151,596,1083,1110]" fontSize="10" level="2" pageId="10" pageNumber="464" reason="2">
<emphasis id="E9CBE559F469FFF9FF0CD51CFDCFFBD1" bold="true" box="[151,596,1083,1110]" pageId="10" pageNumber="464">
The status of
<taxonomicName id="1CBF42C8F469FFF9FEA3D51CFE31FBD1" ID-CoL="8LCTG" authorityName="Thomas" authorityYear="1904" box="[312,426,1083,1110]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="bicolor">
<emphasis id="E9CBE559F469FFF9FEA3D51CFE31FBD1" bold="true" box="[312,426,1083,1110]" italics="true" pageId="10" pageNumber="464">K. bicolor</emphasis>
</taxonomicName>
and
<taxonomicName id="1CBF42C8F469FFF9FE7ED51CFDCFFBD1" authorityName="Thomas" authorityYear="1894" box="[485,596,1083,1110]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FE7ED51CFDCFFBD1" bold="true" box="[485,596,1083,1110]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="93A56AC0F469FFFEFF0CD540FE9CFE70" lastPageId="13" lastPageNumber="467" pageId="10" pageNumber="464" type="discussion">
<paragraph id="DB00394BF469FFF9FF0CD540FED6FA1D" blockId="10.[151,1437,1083,2045]" pageId="10" pageNumber="464">
Two taxa originally named as distinct species,
<taxonomicName id="1CBF42C8F469FFF9FD29D540FC50FBFD" authority="Thomas 1904" authorityName="Thomas" authorityYear="1904" box="[690,971,1119,1146]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="bicolor">
<emphasis id="E9CBE559F469FFF9FD29D540FCBDFBFD" box="[690,806,1120,1146]" italics="true" pageId="10" pageNumber="464">K. bicolor</emphasis>
<bibRefCitation id="BF2E44BAF469FFF9FCB4D57FFC50FBFD" author="Thomas, O." box="[815,971,1119,1146]" pageId="10" pageNumber="464" pagination="384 - 388" refId="ref20157" refString="Thomas, O. (1904) Three new bats, African and Asiatic. Annals and Magazine of Natural History, 13, 384 - 388. https: // doi. org / 10.1080 / 00222930408562462" type="journal article" year="1904">Thomas 1904</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="1CBF42C8F469FFF9FB9DD540FA86FBFD" authority="Thomas 1894" authorityName="Thomas" authorityYear="1894" box="[1030,1309,1119,1146]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FB9DD540FBEEFBFD" box="[1030,1141,1120,1146]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
<bibRefCitation id="BF2E44BAF469FFF9FBE4D57FFA86FBFD" author="Thomas, O." box="[1151,1309,1119,1146]" pageId="10" pageNumber="464" pagination="460 - 462" refId="ref20117" refString="Thomas, O. (1894) Descriptions of two new bats of the genus Kerivoula. Annals and Magazine of Natural History, 14, 460 - 462. https: // doi. org / 10.1080 / 00222939408677834" type="journal article" year="1894">Thomas 1894</bibRefCitation>
</taxonomicName>
, were considered by
<bibRefCitation id="BF2E44BAF469FFF9FE83D5A3FE02FB19" author="Hill, J. E." box="[280,409,1155,1182]" pageId="10" pageNumber="464" pagination="524 - 556" refId="ref17527" refString="Hill, J. E. (1965) Asiatic bats of the genera Kerivoula and Phoniscus (Vespertilionidae), with a note on Kerivoula aerosa Tomes. Mammalia, 29, 524 - 556. https: // doi. org / 10.1515 / mamm. 1965.29.4.524" type="journal article" year="1965">Hill (1965)</bibRefCitation>
to represent subspecies of
<taxonomicName id="1CBF42C8F469FFF9FD4CD5A4FCEDFB19" authorityName="Thomas" authorityYear="1894" box="[727,886,1156,1182]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F469FFF9FD4CD5A4FCEDFB19" box="[727,886,1156,1182]" italics="true" pageId="10" pageNumber="464">K. whiteheadi</emphasis>
</taxonomicName>
, and have been provisionally dealt with in that fashion subsequently (e.g.,
<bibRefCitation id="BF2E44BAF469FFF9FE51D587FD3CFB45" author="Corbet, G. B. &amp; Hill, J. E." box="[458,679,1191,1218]" pageId="10" pageNumber="464" refId="ref15987" refString="Corbet, G. B. &amp; Hill, J. E. (1992) The Mammals of the Indomalayan Region: A systematic Review. Oxford University Press, Oxford, 488 pp." type="book" year="1992">Corbet &amp; Hill 1992</bibRefCitation>
;
<bibRefCitation id="BF2E44BAF469FFF9FD28D587FCD2FB45" author="Francis, C. M." box="[691,841,1191,1218]" pageId="10" pageNumber="464" refId="ref16454" refString="Francis, C. M. (2008) A Guide to the Mammals of Southeast Asia. Princeton University Press, Princeton, New Jersey, 392 pp." type="book" year="2008">Francis 2008</bibRefCitation>
). At the time,
<bibRefCitation id="BF2E44BAF469FFF9FC7DD587FBF8FB45" author="Hill, J. E." box="[998,1123,1191,1218]" pageId="10" pageNumber="464" pagination="524 - 556" refId="ref17527" refString="Hill, J. E. (1965) Asiatic bats of the genera Kerivoula and Phoniscus (Vespertilionidae), with a note on Kerivoula aerosa Tomes. Mammalia, 29, 524 - 556. https: // doi. org / 10.1515 / mamm. 1965.29.4.524" type="journal article" year="1965">Hill (1965)</bibRefCitation>
had only the
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of
<taxonomicName id="1CBF42C8F469FFF9FA18D588FE89FB61" authorityName="Thomas" authorityYear="1894" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F469FFF9FA18D588FE89FB61" italics="true" pageId="10" pageNumber="464">K. whiteheadi</emphasis>
</taxonomicName>
from Luzon, a badly damaged specimen, for comparison with
<taxonomicName id="1CBF42C8F469FFF9FC46D5ECFBD4FB61" authorityName="Thomas" authorityYear="1904" box="[989,1103,1228,1254]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="bicolor">
<emphasis id="E9CBE559F469FFF9FC46D5ECFBD4FB61" box="[989,1103,1228,1254]" italics="true" pageId="10" pageNumber="464">K. bicolor</emphasis>
</taxonomicName>
, also represented only by the
<typeStatus id="040487E9F469FFF9FF0CD5CFFF67FA8E" box="[151,252,1263,1289]" pageId="10" pageNumber="464" type="holotype">holotype</typeStatus>
, and with
<taxonomicName id="1CBF42C8F469FFF9FEF4D5D0FE47FA8D" authorityName="Thomas" authorityYear="1894" box="[367,476,1264,1290]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FEF4D5D0FE47FA8D" box="[367,476,1264,1290]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
, represented by two specimens. He also noted two specimens referred to as
<taxonomicName id="1CBF42C8F469FFF9FAB4D5D0FA07FA8D" authorityName="Thomas" authorityYear="1894" box="[1327,1436,1264,1290]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FAB4D5D0FA07FA8D" box="[1327,1436,1264,1290]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
by
<bibRefCitation id="BF2E44BAF469FFF9FF22D433FEF1FAA9" author="Sanborn, C. C." box="[185,362,1299,1326]" pageId="10" pageNumber="464" pagination="89 - 158" refId="ref19343" refString="Sanborn, C. C. (1952) Philippine Zoological Expedition 1946 - 1947: Mammals. Fieldiana: Zoology, 33, 89 - 158." type="journal article" year="1952">Sanborn (1952)</bibRefCitation>
, which he did not examine. We know of no specimens of
<taxonomicName id="1CBF42C8F469FFF9FC73D434FBC2FAA9" authorityName="Thomas" authorityYear="1904" box="[1000,1113,1300,1326]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="bicolor">
<emphasis id="E9CBE559F469FFF9FC73D434FBC2FAA9" box="[1000,1113,1300,1326]" italics="true" pageId="10" pageNumber="464">K. bicolor</emphasis>
</taxonomicName>
collected since the
<typeStatus id="040487E9F469FFF9FAA8D433FA03FAAA" box="[1331,1432,1299,1325]" pageId="10" pageNumber="464" type="holotype">holotype</typeStatus>
, and only a few
<taxonomicName id="1CBF42C8F469FFF9FED8D418FE34FAD5" authorityName="Thomas" authorityYear="1894" box="[323,431,1336,1362]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FED8D418FE34FAD5" box="[323,431,1336,1362]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
have been reported from Borneo (
<bibRefCitation id="BF2E44BAF469FFF9FCB5D417FBC6FAD5" author="Phillipps, Q. &amp; Phillipps, K." box="[814,1117,1335,1362]" pageId="10" pageNumber="464" refId="ref18748" refString="Phillipps, Q. &amp; Phillipps, K. (2016) Mammals of Borneo and their ecology. John Beaufoy Publishing Ltd, Oxford, 400 pp." type="book" year="2016">Phillipps &amp; Phillipps 2016</bibRefCitation>
), but we have obtained large numbers of
<taxonomicName id="1CBF42C8F469FFF9FE86D47CFE22FAF1" authorityName="Thomas" authorityYear="1894" box="[285,441,1372,1398]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F469FFF9FE86D47CFE22FAF1" box="[285,441,1372,1398]" italics="true" pageId="10" pageNumber="464">K. whiteheadi</emphasis>
</taxonomicName>
from the
<collectingCountry id="A3A879DBF469FFF9FDBDD47BFD3FFAF2" box="[550,676,1371,1397]" name="Philippines" pageId="10" pageNumber="464">Philippines</collectingCountry>
that allow some morphological assessment of the distinctiveness of these three taxa.
</paragraph>
<paragraph id="DB00394BF469FFF9FF5CD484FDC7F8A1" blockId="10.[151,1437,1083,2045]" pageId="10" pageNumber="464">
As shown in
<figureCitation id="438425CEF469FFF9FEC2D483FE23FA39" box="[345,440,1443,1470]" captionStart="FIGURE 3" captionStartId="11.[151,250,1182,1207]" captionTargetBox="[177,1413,279,1153]" captionTargetId="figure@11.[170,1417,273,1157]" captionTargetPageId="11" captionText="FIGURE 3. Bivariate plot of condylo-canine length vs. height of braincase for specimens of Kerivoula summarized in Tables 1 and 3. Color indicates geographic region and symbol indicates taxon. Philippine specimens of each taxon are enclosed within polygons." figureDoi="http://doi.org/10.5281/zenodo.3736824" httpUri="https://zenodo.org/record/3736824/files/figure.png" pageId="10" pageNumber="464">Figure 3</figureCitation>
, a plot of condylo-canine length vs. braincase height of specimens of
<taxonomicName id="1CBF42C8F469FFF9FB5FD484FAC4FA39" authorityName="Thomas" authorityYear="1894" box="[1220,1375,1444,1470]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F469FFF9FB5FD484FAC4FA39" box="[1220,1375,1444,1470]" italics="true" pageId="10" pageNumber="464">K. whiteheadi</emphasis>
</taxonomicName>
from the
<collectingCountry id="A3A879DBF469FFF9FF59D4E7FEDBFA66" box="[194,320,1479,1505]" name="Philippines" pageId="10" pageNumber="464">Philippines</collectingCountry>
form a cloud of points that represents the range of variation in the nominate form. The two specimens referred to
<taxonomicName id="1CBF42C8F469FFF9FEC3D4CCFE5DF981" authorityName="Thomas" authorityYear="1894" box="[344,454,1516,1542]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FEC3D4CCFE5DF981" box="[344,454,1516,1542]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
by Sanborn (FMNH 56687 and 61077) have the shortest condylo-canine length of our Philippine sample, but they fall within the range of other specimens from Mindanao and within the cloud of
<collectingCountry id="A3A879DBF469FFF9FACBD72FFF49F9CA" name="Philippines" pageId="10" pageNumber="464">Philippines</collectingCountry>
points overall. Our measurements of the
<typeStatus id="040487E9F469FFF9FD3BD713FC99F9CA" box="[672,770,1587,1613]" pageId="10" pageNumber="464" type="holotype">holotype</typeStatus>
of
<taxonomicName id="1CBF42C8F469FFF9FCBCD714FC0FF9C9" authorityName="Thomas" authorityYear="1894" box="[807,916,1588,1614]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FCBCD714FC0FF9C9" box="[807,916,1588,1614]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
from Borneo show it to be similar in condylocanine length, but with a notably lower braincase height, well below that of any Philippine specimen (
<figureCitation id="438425CEF469FFF9FA9FD777FAD3F9F5" box="[1284,1352,1623,1650]" captionStart="FIGURE 3" captionStartId="11.[151,250,1182,1207]" captionTargetBox="[177,1413,279,1153]" captionTargetId="figure@11.[170,1417,273,1157]" captionTargetPageId="11" captionText="FIGURE 3. Bivariate plot of condylo-canine length vs. height of braincase for specimens of Kerivoula summarized in Tables 1 and 3. Color indicates geographic region and symbol indicates taxon. Philippine specimens of each taxon are enclosed within polygons." figureDoi="http://doi.org/10.5281/zenodo.3736824" httpUri="https://zenodo.org/record/3736824/files/figure.png" pageId="10" pageNumber="464">Fig. 3</figureCitation>
;
<tableCitation id="963D0CF0F469FFF9FAC8D777FF3EF911" captionStart="TABLE 1" captionStartId="5.[151,239,459,485]" captionTargetPageId="5" captionText="TABLE 1. Mean (± standard deviation) and range of cranial-dental measurements of adult K. whiteheadi and K. pellucida, and comparison groups. Holotypes indicated with an asterisk. n = sample size" httpUri="http://table.plazi.org/id/8FC069C3F466FFF6FF0CD0EBFE00FD8E" pageId="10" pageNumber="464" tableUuid="8FC069C3F466FFF6FF0CD0EBFE00FD8E">Tables 1</tableCitation>
,
<tableCitation id="963D0CF0F469FFF9FF2ED75CFF5EF911" box="[181,197,1660,1686]" captionStart="TABLE 3" captionStartId="12.[151,239,869,893]" captionTargetPageId="12" captionText="TABLE 3. Mean (± standard deviation) and range of condylo-canine length (CCL) and braincase height (BH) for those specimens not included in Table 1. n = sample size." httpUri="http://table.plazi.org/id/8FC069C3F46FFFFFFF0CD245FD27FC26" pageId="10" pageNumber="464" tableUuid="8FC069C3F46FFFFFFF0CD245FD27FC26">3</tableCitation>
). Additionally, our examination of the cranium of the
<typeStatus id="040487E9F469FFF9FCB4D75BFC0AF912" box="[815,913,1659,1685]" pageId="10" pageNumber="464" type="holotype">holotype</typeStatus>
showed it to have unusually large cochlea, as described by
<bibRefCitation id="BF2E44BAF469FFF9FEB7D7BFFE43F93D" author="Thomas, O." box="[300,472,1695,1722]" pageId="10" pageNumber="464" pagination="460 - 462" refId="ref20117" refString="Thomas, O. (1894) Descriptions of two new bats of the genus Kerivoula. Annals and Magazine of Natural History, 14, 460 - 462. https: // doi. org / 10.1080 / 00222939408677834" type="journal article" year="1894">Thomas (1894)</bibRefCitation>
and confirmed by
<bibRefCitation id="BF2E44BAF469FFF9FD30D7BFFCB7F93D" author="Hill, J. E." box="[683,812,1695,1722]" pageId="10" pageNumber="464" pagination="524 - 556" refId="ref17527" refString="Hill, J. E. (1965) Asiatic bats of the genera Kerivoula and Phoniscus (Vespertilionidae), with a note on Kerivoula aerosa Tomes. Mammalia, 29, 524 - 556. https: // doi. org / 10.1515 / mamm. 1965.29.4.524" type="journal article" year="1965">Hill (1965)</bibRefCitation>
. While the cochlea of
<taxonomicName id="1CBF42C8F469FFF9FBBED780FB22F93D" authorityName="Thomas" authorityYear="1894" box="[1061,1209,1696,1722]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F469FFF9FBBED780FB22F93D" box="[1061,1209,1696,1722]" italics="true" pageId="10" pageNumber="464">K whiteheadi</emphasis>
</taxonomicName>
is large (
<figureCitation id="438425CEF469FFF9FABBD7BFFAFEF93D" box="[1312,1381,1695,1722]" captionStart="FIGURE 6" captionStartId="18.[151,250,1965,1990]" captionTargetBox="[321,1266,181,1940]" captionTargetId="figure@18.[321,1266,181,1940]" captionTargetPageId="18" captionText="FIGURE 6. Scanning electron microscope images of K. pellucida cranium and mandible, FMNH 202782 from Bohol Island, Philippines." figureDoi="http://doi.org/10.5281/zenodo.3736830" httpUri="https://zenodo.org/record/3736830/files/figure.png" pageId="10" pageNumber="464">Fig. 6</figureCitation>
), the
<taxonomicName id="1CBF42C8F469FFF9FF0CD7E4FE9FF959" authorityName="Thomas" authorityYear="1894" box="[151,260,1732,1758]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FF0CD7E4FE9FF959" box="[151,260,1732,1758]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
holotypes cochlea (from a photograph we took) is proportionately substantially larger. Given the difference in braincase height and cochlea size in comparison to our large sample, we recommend that
<taxonomicName id="1CBF42C8F469FFF9FB79D7C8FAD4F885" authorityName="Thomas" authorityYear="1894" box="[1250,1359,1768,1794]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FB79D7C8FAD4F885" box="[1250,1359,1768,1794]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
not be treated as a subspecies of
<taxonomicName id="1CBF42C8F469FFF9FE22D62CFDCEF8A1" authorityName="Thomas" authorityYear="1894" box="[441,597,1804,1830]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F469FFF9FE22D62CFDCEF8A1" box="[441,597,1804,1830]" italics="true" pageId="10" pageNumber="464">K. whiteheadi</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="DB00394BF469FFF8FF5CD610FE3AFF7C" blockId="10.[151,1437,1083,2045]" lastBlockId="11.[151,1436,153,251]" lastPageId="11" lastPageNumber="465" pageId="10" pageNumber="464">
We also examined the
<typeStatus id="040487E9F469FFF9FE4AD60FFDA8F8CE" box="[465,563,1839,1865]" pageId="10" pageNumber="464" type="holotype">holotype</typeStatus>
(and only known specimen) of
<taxonomicName id="1CBF42C8F469FFF9FC31D610FB84F8CD" authorityName="Thomas" authorityYear="1904" box="[938,1055,1840,1866]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="bicolor">
<emphasis id="E9CBE559F469FFF9FC31D610FB84F8CD" box="[938,1055,1840,1866]" italics="true" pageId="10" pageNumber="464">K. bicolor</emphasis>
</taxonomicName>
. The cranium is badly damaged, and we were unable to measure the condylo-canine length. Further, the cochlea were not present for comparison. However, the braincase height (
<quantity id="1C4794AEF469FFF9FE61D657FDFAF815" box="[506,609,1911,1938]" metricMagnitude="-3" metricUnit="m" metricValue="4.74" pageId="10" pageNumber="464" unit="mm" value="4.74">4.74 mm</quantity>
) is well below the range shown by
<taxonomicName id="1CBF42C8F469FFF9FC6BD658FB17F816" authorityName="Thomas" authorityYear="1894" box="[1008,1164,1911,1937]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F469FFF9FC6BD658FB17F816" box="[1008,1164,1911,1937]" italics="true" pageId="10" pageNumber="464">K. whiteheadi</emphasis>
</taxonomicName>
(
<figureCitation id="438425CEF469FFF9FB07D657FB7AF816" box="[1180,1249,1911,1937]" captionStart="FIGURE 3" captionStartId="11.[151,250,1182,1207]" captionTargetBox="[177,1413,279,1153]" captionTargetId="figure@11.[170,1417,273,1157]" captionTargetPageId="11" captionText="FIGURE 3. Bivariate plot of condylo-canine length vs. height of braincase for specimens of Kerivoula summarized in Tables 1 and 3. Color indicates geographic region and symbol indicates taxon. Philippine specimens of each taxon are enclosed within polygons." figureDoi="http://doi.org/10.5281/zenodo.3736824" httpUri="https://zenodo.org/record/3736824/files/figure.png" pageId="10" pageNumber="464">Fig. 3</figureCitation>
;
<tableCitation id="963D0CF0F469FFF9FB76D657FAA4F816" box="[1261,1343,1911,1937]" captionStart="TABLE 1" captionStartId="5.[151,239,459,485]" captionTargetPageId="5" captionText="TABLE 1. Mean (± standard deviation) and range of cranial-dental measurements of adult K. whiteheadi and K. pellucida, and comparison groups. Holotypes indicated with an asterisk. n = sample size" httpUri="http://table.plazi.org/id/8FC069C3F466FFF6FF0CD0EBFE00FD8E" pageId="10" pageNumber="464" tableUuid="8FC069C3F466FFF6FF0CD0EBFE00FD8E">Table 1</tableCitation>
). Given the comment by
<bibRefCitation id="BF2E44BAF469FFF9FEC8D6BBFD99F832" author="Hill, J. E." box="[339,514,1946,1973]" pageId="10" pageNumber="464" pagination="524 - 556" refId="ref17527" refString="Hill, J. E. (1965) Asiatic bats of the genera Kerivoula and Phoniscus (Vespertilionidae), with a note on Kerivoula aerosa Tomes. Mammalia, 29, 524 - 556. https: // doi. org / 10.1515 / mamm. 1965.29.4.524" type="journal article" year="1965">Hill (1965:531)</bibRefCitation>
that “both [
<taxonomicName id="1CBF42C8F469FFF9FD12D6BCFD60F832" authorityName="Thomas" authorityYear="1904" box="[649,763,1947,1973]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="bicolor">
<emphasis id="E9CBE559F469FFF9FD12D6BCFD60F832" box="[649,763,1947,1973]" italics="true" pageId="10" pageNumber="464">K. bicolor</emphasis>
</taxonomicName>
and
<taxonomicName id="1CBF42C8F469FFF9FCAAD6BCFC05F832" authorityName="Thomas" authorityYear="1894" box="[817,926,1947,1973]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">
<emphasis id="E9CBE559F469FFF9FCAAD6BCFC05F832" box="[817,926,1947,1973]" italics="true" pageId="10" pageNumber="464">K. pusilla</emphasis>
</taxonomicName>
] have greatly expanded cochlea”, and that
<emphasis id="E9CBE559F469FFF9FA18D6BCFE93F85E" italics="true" pageId="10" pageNumber="464">
K. w.
<taxonomicName id="1CBF42C8F469FFF9FF2CD69FFE93F85E" authorityName="Thomas" authorityYear="1904" box="[183,264,1983,2009]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="bicolor">bicolor</taxonomicName>
</emphasis>
“closely resembles
<emphasis id="E9CBE559F469FFF9FE77D6E0FDE1F85E" box="[492,634,1983,2009]" italics="true" pageId="10" pageNumber="464">
K. w.
<taxonomicName id="1CBF42C8F469FFF9FDB5D69FFDE1F85E" authorityName="Thomas" authorityYear="1894" box="[558,634,1983,2009]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="10" pageNumber="464" phylum="Chordata" rank="species" species="pusilla">pusilla</taxonomicName>
</emphasis>
, differing … only in its white-tipped wings, white and not grayish underparts, slightly narrower rostrum and slightly less pronounced rostral sulcus … [which] seem only of subspecific importance”, we provisionally conclude that these should not be considered conspecific with
<taxonomicName id="1CBF42C8F468FFF8FB5DD1BAFAFFFF34" authorityName="Thomas" authorityYear="1894" box="[1222,1380,153,179]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F468FFF8FB5DD1BAFAFFFF34" box="[1222,1380,153,179]" italics="true" pageId="11" pageNumber="465">K. whiteheadi</emphasis>
</taxonomicName>
, and can be treated as subspecies of a single species,
<emphasis id="E9CBE559F468FFF8FD35D19EFCADFF50" box="[686,822,189,215]" italics="true" pageId="11" pageNumber="465">
K. p.
<taxonomicName id="1CBF42C8F468FFF8FD71D19DFCADFF50" authorityName="Thomas" authorityYear="1894" box="[746,822,189,215]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="pusilla">pusilla</taxonomicName>
</emphasis>
and
<emphasis id="E9CBE559F468FFF8FCF7D19EFC62FF50" box="[876,1017,189,215]" italics="true" pageId="11" pageNumber="465">
K. p.
<taxonomicName id="1CBF42C8F468FFF8FC33D19DFC62FF50" authorityName="Thomas" authorityYear="1904" box="[936,1017,189,215]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="bicolor">bicolor</taxonomicName>
</emphasis>
, until further specimens are available for more detailed study.
</paragraph>
<caption id="8FC069C3F468FFF8FF0CD5BEFF60FB78" ID-DOI="http://doi.org/10.5281/zenodo.3736824" ID-Zenodo-Dep="3736824" httpUri="https://zenodo.org/record/3736824/files/figure.png" pageId="11" pageNumber="465" startId="11.[151,250,1182,1207]" targetBox="[177,1413,279,1153]" targetPageId="11">
<paragraph id="DB00394BF468FFF8FF0CD5BEFF60FB78" blockId="11.[151,1436,1182,1279]" pageId="11" pageNumber="465">
<emphasis id="E9CBE559F468FFF8FF0CD5BEFE8FFB30" bold="true" box="[151,276,1182,1207]" pageId="11" pageNumber="465">FIGURE 3.</emphasis>
Bivariate plot of condylo-canine length vs. height of braincase for specimens of
<taxonomicName id="1CBF42C8F468FFF8FBCAD5BEFB2DFB31" authorityName="Gray" authorityYear="1842" box="[1105,1206,1182,1206]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F468FFF8FBCAD5BEFB2DFB31" box="[1105,1206,1182,1206]" italics="true" pageId="11" pageNumber="465">Kerivoula</emphasis>
</taxonomicName>
summarized in Tables 1 and 3. Color indicates geographic region and symbol indicates taxon. Philippine specimens of each taxon are enclosed within polygons.
</paragraph>
</caption>
<paragraph id="DB00394BF468FFF8FF0CD40CFDBFFAC0" blockId="11.[151,1437,1324,2035]" box="[151,548,1324,1351]" pageId="11" pageNumber="465">
<emphasis id="E9CBE559F468FFF8FF0CD40CFDBFFAC0" bold="true" box="[151,548,1324,1351]" pageId="11" pageNumber="465">Biogeography and diversification</emphasis>
</paragraph>
<paragraph id="DB00394BF468FFF8FF0CD471FB88FA7C" blockId="11.[151,1437,1324,2035]" pageId="11" pageNumber="465">
Our time-calibrated phylogenetic tree of Cyt
<emphasis id="E9CBE559F468FFF8FD16D471FD00FAEC" box="[653,667,1361,1387]" italics="true" pageId="11" pageNumber="465">b</emphasis>
sequences obtained with BEAST has implications for the timing and geography of diversification of
<taxonomicName id="1CBF42C8F468FFF8FD9AD455FDEAFA08" authorityName="Gray" authorityYear="1842" box="[513,625,1397,1423]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F468FFF8FD9AD455FDEAFA08" box="[513,625,1397,1423]" italics="true" pageId="11" pageNumber="465">Kerivoula</emphasis>
</taxonomicName>
in the
<collectingCountry id="A3A879DBF468FFF8FD58D455FCDAFA08" box="[707,833,1397,1423]" name="Philippines" pageId="11" pageNumber="465">Philippines</collectingCountry>
and adjacent areas (
<figureCitation id="438425CEF468FFF8FBB0D455FBE9FA08" box="[1067,1138,1397,1423]" captionStart="FIGURE 4" captionStartId="13.[151,250,1314,1339]" captionTargetBox="[155,1432,528,1286]" captionTargetId="figure@13.[151,1436,525,1289]" captionTargetPageId="13" captionText="FIGURE 4. Time-calibrated phylogeny of Kerivoula species based on Cyt b haplotypes. Outgroups were omitted for ease of readability. The greyed boxes represent 95% highest posterior density (HPD) intervals of node ages derived from a BEAST analysis. Time is expressed in million years." figureDoi="http://doi.org/10.5281/zenodo.3736826" httpUri="https://zenodo.org/record/3736826/files/figure.png" pageId="11" pageNumber="465">Fig. 4</figureCitation>
). As noted above, support for many nodes is strong, but confidence limits (i.e. 95% HPD intervals) on the timing of most nodes are broad, so that caution is warranted.All mention of dates that follow are given as the average; these should be viewed as rough estimates, plus or minus about 30%, with the confidence limits shown in
<figureCitation id="438425CEF468FFF8FC50D4C1FB94FA7C" box="[971,1039,1505,1531]" captionStart="FIGURE 4" captionStartId="13.[151,250,1314,1339]" captionTargetBox="[155,1432,528,1286]" captionTargetId="figure@13.[151,1436,525,1289]" captionTargetPageId="13" captionText="FIGURE 4. Time-calibrated phylogeny of Kerivoula species based on Cyt b haplotypes. Outgroups were omitted for ease of readability. The greyed boxes represent 95% highest posterior density (HPD) intervals of node ages derived from a BEAST analysis. Time is expressed in million years." figureDoi="http://doi.org/10.5281/zenodo.3736826" httpUri="https://zenodo.org/record/3736826/files/figure.png" pageId="11" pageNumber="465">Fig. 4</figureCitation>
.
</paragraph>
<paragraph id="DB00394BF468FFF8FF5CD725FDB1F928" blockId="11.[151,1437,1324,2035]" pageId="11" pageNumber="465">
The genus
<taxonomicName id="1CBF42C8F468FFF8FEDBD725FE2BF998" authorityName="Gray" authorityYear="1842" box="[320,432,1541,1567]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F468FFF8FEDBD725FE2BF998" box="[320,432,1541,1567]" italics="true" pageId="11" pageNumber="465">Kerivoula</emphasis>
</taxonomicName>
is here estimated to have diverged from
<taxonomicName id="1CBF42C8F468FFF8FCEAD725FC7FF998" authorityName="Miller" authorityYear="1905" box="[881,996,1541,1567]" class="Mammalia" family="Vespertilionidae" genus="Phoniscus" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F468FFF8FCEAD725FC7FF998" box="[881,996,1541,1567]" italics="true" pageId="11" pageNumber="465">Phoniscus</emphasis>
</taxonomicName>
, its sister-genus, about 27 million years ago (MYA;
<figureCitation id="438425CEF468FFF8FE85D709FEF8F9C4" box="[286,355,1577,1603]" captionStart="FIGURE 4" captionStartId="13.[151,250,1314,1339]" captionTargetBox="[155,1432,528,1286]" captionTargetId="figure@13.[151,1436,525,1289]" captionTargetPageId="13" captionText="FIGURE 4. Time-calibrated phylogeny of Kerivoula species based on Cyt b haplotypes. Outgroups were omitted for ease of readability. The greyed boxes represent 95% highest posterior density (HPD) intervals of node ages derived from a BEAST analysis. Time is expressed in million years." figureDoi="http://doi.org/10.5281/zenodo.3736826" httpUri="https://zenodo.org/record/3736826/files/figure.png" pageId="11" pageNumber="465">Fig. 4</figureCitation>
). The initial split between the
<taxonomicName id="1CBF42C8F468FFF8FD20D70AFCDFF9C4" authorityName="Waterhouse" authorityYear="1854" box="[699,836,1577,1603]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="pellucida">
<emphasis id="E9CBE559F468FFF8FD20D70AFD55F9C4" box="[699,718,1578,1603]" italics="true" pageId="11" pageNumber="465">K</emphasis>
.
<emphasis id="E9CBE559F468FFF8FD47D709FCDFF9C4" box="[732,836,1577,1603]" italics="true" pageId="11" pageNumber="465">pellucida</emphasis>
</taxonomicName>
species group (AF) and all other species took place about 25 MYA, and the split between the
<taxonomicName id="1CBF42C8F468FFF8FDEDD76EFC9AF9E0" baseAuthorityName="Temminck" baseAuthorityYear="1840" box="[630,769,1613,1639]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="papillosa">
<emphasis id="E9CBE559F468FFF8FDEDD76EFC9AF9E0" box="[630,769,1613,1639]" italics="true" pageId="11" pageNumber="465">K. papillosa</emphasis>
</taxonomicName>
species group (KM) and the
<taxonomicName id="1CBF42C8F468FFF8FBFBD76EFB62F9E0" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[1120,1273,1613,1639]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F468FFF8FBFBD76EFBE8F9E0" box="[1120,1139,1614,1639]" italics="true" pageId="11" pageNumber="465">K</emphasis>
.
<emphasis id="E9CBE559F468FFF8FB19D76DFB62F9E0" box="[1154,1273,1613,1639]" italics="true" pageId="11" pageNumber="465">hardwickii</emphasis>
</taxonomicName>
species group (GJ) about 21 MYA. Thus, initial diversity within
<taxonomicName id="1CBF42C8F468FFF8FD7FD751FCCFF90C" authorityName="Gray" authorityYear="1842" box="[740,852,1649,1675]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F468FFF8FD7FD751FCCFF90C" box="[740,852,1649,1675]" italics="true" pageId="11" pageNumber="465">Kerivoula</emphasis>
</taxonomicName>
in Southeast Asia appears to have been well established by more than about 20 MYA.
</paragraph>
<paragraph id="DB00394BF468FFF8FF5CD799FA8AF8E4" blockId="11.[151,1437,1324,2035]" pageId="11" pageNumber="465">
Diversification within the species groups has been more recent.
<taxonomicName id="1CBF42C8F468FFF8FC04D79AFBB2F954" authorityName="Waterhouse" authorityYear="1854" box="[927,1065,1721,1747]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="pellucida">
<emphasis id="E9CBE559F468FFF8FC04D79AFC29F954" box="[927,946,1722,1747]" italics="true" pageId="11" pageNumber="465">K</emphasis>
.
<emphasis id="E9CBE559F468FFF8FC5AD799FBB2F954" box="[961,1065,1721,1747]" italics="true" pageId="11" pageNumber="465">pellucida</emphasis>
</taxonomicName>
(
<collectionCodeDefinition id="1FC1797BF468FFF8FBA1D798FAD5F954" box="[1082,1358,1720,1747]" pageId="11" pageNumber="465">
<emphasis id="E9CBE559F468FFF8FBA1D798FBEBF954" bold="true" box="[1082,1136,1720,1747]" pageId="11" pageNumber="465">AB</emphasis>
) and
<taxonomicName id="1CBF42C8F468FFF8FB2AD79AFAD5F954" authorityName="Thomas" authorityYear="1894" box="[1201,1358,1721,1747]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F468FFF8FB2AD79AFAD5F954" box="[1201,1358,1721,1747]" italics="true" pageId="11" pageNumber="465">K. whiteheadi</emphasis>
</taxonomicName>
</collectionCodeDefinition>
(
<collectionCodeDefinition id="1FC1797BF468FFF8FAFBD798FB9AF89C" pageId="11" pageNumber="465">
<emphasis id="E9CBE559F468FFF8FAFBD798FA08F954" bold="true" box="[1376,1427,1720,1747]" pageId="11" pageNumber="465">CF</emphasis>
) last shared a common ancestor about 18 MYA, but diversification within these species (as currently defined) began about 3.7 MYA and 7.1 MYA, respectively. Members of the
<taxonomicName id="1CBF42C8F468FFF8FCA9D622FC21F89C" baseAuthorityName="Temminck" baseAuthorityYear="1840" box="[818,954,1793,1819]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="papillosa">
<emphasis id="E9CBE559F468FFF8FCA9D622FCDEF89C" box="[818,837,1794,1819]" italics="true" pageId="11" pageNumber="465">K</emphasis>
.
<emphasis id="E9CBE559F468FFF8FCC9D621FC21F89C" box="[850,954,1793,1819]" italics="true" pageId="11" pageNumber="465">papillosa</emphasis>
</taxonomicName>
group
</collectionCodeDefinition>
(
<collectionCodeDefinition id="1FC1797BF468FFF8FB8AD621FA96F8E4" pageId="11" pageNumber="465">
<emphasis id="E9CBE559F468FFF8FB8AD621FBD5F89C" bold="true" box="[1041,1102,1793,1819]" pageId="11" pageNumber="465">KM</emphasis>
) last shared a common ancestor about 15.3 MYA, but most species have diverged within the last 10 MYA (
<figureCitation id="438425CEF468FFF8FC6CD605FBA7F8B8" box="[1015,1084,1829,1855]" captionStart="FIGURE 4" captionStartId="13.[151,250,1314,1339]" captionTargetBox="[155,1432,528,1286]" captionTargetId="figure@13.[151,1436,525,1289]" captionTargetPageId="13" captionText="FIGURE 4. Time-calibrated phylogeny of Kerivoula species based on Cyt b haplotypes. Outgroups were omitted for ease of readability. The greyed boxes represent 95% highest posterior density (HPD) intervals of node ages derived from a BEAST analysis. Time is expressed in million years." figureDoi="http://doi.org/10.5281/zenodo.3736826" httpUri="https://zenodo.org/record/3736826/files/figure.png" pageId="11" pageNumber="465">Fig. 4</figureCitation>
). Members of the
<taxonomicName id="1CBF42C8F468FFF8FA9ED606FA07F8B8" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[1285,1436,1829,1855]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F468FFF8FA9ED606FA07F8B8" box="[1285,1436,1829,1855]" italics="true" pageId="11" pageNumber="465">K. hardwickii</emphasis>
</taxonomicName>
species group (
<emphasis id="E9CBE559F468FFF8FEDAD668FEE9F8E4" bold="true" box="[321,370,1864,1891]" pageId="11" pageNumber="465">GJ</emphasis>
) last shared a common ancestor about 17.2 MYA, with subsequent diversification
</collectionCodeDefinition>
.
</paragraph>
<paragraph id="DB00394BF468FFFFFF5CD64DFCBBFEC4" blockId="11.[151,1437,1324,2035]" lastBlockId="12.[151,1437,152,827]" lastPageId="12" lastPageNumber="466" pageId="11" pageNumber="465">
We note that most diversification within the
<collectingCountry id="A3A879DBF468FFF8FD26D64DFCA0F800" box="[701,827,1901,1927]" name="Philippines" pageId="11" pageNumber="465">Philippines</collectingCountry>
is relatively recent.
<taxonomicName id="1CBF42C8F468FFF8FBBAD64DFA9AF800" authorityName="Waterhouse" authorityYear="1854" box="[1057,1281,1901,1927]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="pellucida">
<emphasis id="E9CBE559F468FFF8FBBAD64DFA9AF800" box="[1057,1281,1901,1927]" italics="true" pageId="11" pageNumber="465">Kerivoula pellucida</emphasis>
</taxonomicName>
in the
<collectingCountry id="A3A879DBF468FFF8FACBD64DFF49F82C" name="Philippines" pageId="11" pageNumber="465">Philippines</collectingCountry>
diverged from their Sunda Shelf conspecifics about 3.7 MYA, but divergence within the
<collectingCountry id="A3A879DBF468FFF8FB2BD6B1FAB5F82C" box="[1200,1326,1937,1963]" name="Philippines" pageId="11" pageNumber="465">Philippines</collectingCountry>
(clade
<emphasis id="E9CBE559F468FFF8FAE4D6B0FA08F82C" bold="true" box="[1407,1427,1936,1963]" pageId="11" pageNumber="465">A</emphasis>
) appears to have begun only about 2.0 MYA. Populations referred to
<taxonomicName id="1CBF42C8F468FFF8FC06D696FBA2F848" authorityName="Thomas" authorityYear="1894" box="[925,1081,1973,1999]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="11" pageNumber="465" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F468FFF8FC06D696FBA2F848" box="[925,1081,1973,1999]" italics="true" pageId="11" pageNumber="465">K. whiteheadi</emphasis>
</taxonomicName>
last shared a common ancestor about 7.1 MYA, and have formed geographically cohesive clades (
<emphasis id="E9CBE559F468FFF8FC0BD6F8FC3FF874" bold="true" box="[912,932,2008,2035]" pageId="11" pageNumber="465">C</emphasis>
,
<emphasis id="E9CBE559F468FFF8FC2FD6F9FC53F874" bold="true" box="[948,968,2009,2035]" pageId="11" pageNumber="465">D</emphasis>
,
<emphasis id="E9CBE559F468FFF8FC4CD6F9FC6AF874" bold="true" box="[983,1009,2009,2035]" pageId="11" pageNumber="465">E,</emphasis>
and
<emphasis id="E9CBE559F468FFF8FBB1D6F9FBA0F874" bold="true" box="[1066,1083,2009,2035]" pageId="11" pageNumber="465">F</emphasis>
) subsequently, with specimens from northern Luzon,
<collectingRegion id="197BF7A9F46FFFFFFE0BD1B9FE65FF34" box="[400,510,153,179]" country="Philippines" name="Camiguin" pageId="12" pageNumber="466">Camiguin</collectingRegion>
Norte, Lubang, and Mindoro (
<emphasis id="E9CBE559F46FFFFFFCC3D1B8FCF7FF34" bold="true" box="[856,876,152,179]" pageId="12" pageNumber="466">C</emphasis>
) sharing a common ancestor about 1.4 MYA. The oldest clade that includes
<taxonomicName id="1CBF42C8F46FFFFFFE2FD19EFDA7FF50" baseAuthorityName="Temminck" baseAuthorityYear="1840" box="[436,572,189,215]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="papillosa">
<emphasis id="E9CBE559F46FFFFFFE2FD19EFE5CFF50" box="[436,455,190,215]" italics="true" pageId="12" pageNumber="466">K</emphasis>
.
<emphasis id="E9CBE559F46FFFFFFE4FD19DFDA7FF50" box="[468,572,189,215]" italics="true" pageId="12" pageNumber="466">papillosa</emphasis>
</taxonomicName>
in the
<collectingCountry id="A3A879DBF46FFFFFFD1ED19DFC98FF50" box="[645,771,189,215]" name="Philippines" pageId="12" pageNumber="466">Philippines</collectingCountry>
diverged about 7.0 MYA, but this includes a specimen from
<collectingCountry id="A3A879DBF46FFFFFFF0CD1C1FF61FF7C" box="[151,250,225,251]" name="Malaysia" pageId="12" pageNumber="466">Sarawak</collectingCountry>
, which makes the interpretation of diversification within the
<collectingCountry id="A3A879DBF46FFFFFFC22D1C1FBACFF7C" box="[953,1079,225,251]" name="Philippines" pageId="12" pageNumber="466">Philippines</collectingCountry>
ambiguous. Finally, Philippine populations of
<taxonomicName id="1CBF42C8F46FFFFFFEDAD026FE42FE98" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[321,473,261,287]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F46FFFFFFEDAD026FECFFE98" box="[321,340,262,287]" italics="true" pageId="12" pageNumber="466">K</emphasis>
.
<emphasis id="E9CBE559F46FFFFFFEF9D025FE42FE98" box="[354,473,261,287]" italics="true" pageId="12" pageNumber="466">hardwickii</emphasis>
</taxonomicName>
(G) diverged from their Sunda Shelf relatives (G”) by about 9.6 MYA, the oldest of the implied colonizations of the
<collectingCountry id="A3A879DBF46FFFFFFD9AD009FDE4FEC4" box="[513,639,297,323]" name="Philippines" pageId="12" pageNumber="466">Philippines</collectingCountry>
by
<taxonomicName id="1CBF42C8F46FFFFFFD32D009FC82FEC4" authorityName="Gray" authorityYear="1842" box="[681,793,297,323]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F46FFFFFFD32D009FC82FEC4" box="[681,793,297,323]" italics="true" pageId="12" pageNumber="466">Kerivoula</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="DB00394BF46FFFFFFF5CD06DFCC4FD9C" blockId="12.[151,1437,152,827]" pageId="12" pageNumber="466">
These data imply that
<taxonomicName id="1CBF42C8F46FFFFFFE50D06DFDA0FEE0" authorityName="Gray" authorityYear="1842" box="[459,571,333,359]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F46FFFFFFE50D06DFDA0FEE0" box="[459,571,333,359]" italics="true" pageId="12" pageNumber="466">Kerivoula</emphasis>
</taxonomicName>
arrived in the
<collectingCountry id="A3A879DBF46FFFFFFD71D06DFCF3FEE0" box="[746,872,333,359]" name="Philippines" pageId="12" pageNumber="466">Philippines</collectingCountry>
by about 10 MYA.
<taxonomicName id="1CBF42C8F46FFFFFFBCED06DFAD2FEE0" authorityName="Thomas" authorityYear="1894" box="[1109,1353,333,359]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F46FFFFFFBCED06DFAD2FEE0" box="[1109,1353,333,359]" italics="true" pageId="12" pageNumber="466">Kerivoula whiteheadi</emphasis>
</taxonomicName>
occurs throughout the entire
<collectingCountry id="A3A879DBF46FFFFFFE10D051FD92FE0C" box="[395,521,369,395]" name="Philippines" pageId="12" pageNumber="466">Philippines</collectingCountry>
(
<bibRefCitation id="BF2E44BAF46FFFFFFD82D051FC92FE0C" author="Esselstyn, J. A. &amp; Widmann, P. &amp; Heaney, L. R." box="[537,777,369,395]" pageId="12" pageNumber="466" pagination="271 - 302" refId="ref16343" refString="Esselstyn, J. A., Widmann, P. &amp; Heaney, L. R. (2004) The mammals of Palawan Island, Philippines. Proceedings of the Biological Society of Washington, 117, 271 - 302." type="journal article" year="2004">
Esselstyn
<emphasis id="E9CBE559F46FFFFFFD17D052FD5EFE0C" box="[652,709,369,395]" italics="true" pageId="12" pageNumber="466">et al.</emphasis>
2004
</bibRefCitation>
;
<bibRefCitation id="BF2E44BAF46FFFFFFC8FD051FC75FE0C" author="Heaney, L. R. &amp; Dolar, M. L. &amp; Balete, D. S. &amp; Esselstyn, J. A. &amp; Rickart, E. A. &amp; Sedlock, J. L." box="[788,1006,369,395]" pageId="12" pageNumber="466" refId="ref17091" refString="Heaney, L. R., Dolar, M. L., Balete, D. S., Esselstyn, J. A., Rickart, E. A. &amp; Sedlock, J. L. (2010) Synopsis of Philippine Mammals. Available from: http: // www. fieldmuseum. org / philippine _ mammals / (accessed 8 May 2019)" type="url" year="2010">
Heaney
<emphasis id="E9CBE559F46FFFFFFCE9D052FC30FE0C" box="[882,939,369,395]" italics="true" pageId="12" pageNumber="466">et al.</emphasis>
2010
</bibRefCitation>
;
<figureCitation id="438425CEF46FFFFFFC61D051FBA3FE0C" box="[1018,1080,369,395]" captionStart="FIGURE 2" captionStartId="9.[151,250,1957,1982]" captionTargetBox="[255,1332,181,1931]" captionTargetId="figure@9.[255,1332,181,1931]" captionTargetPageId="9" captionText="FIGURE 2. Map of the Philippines indicating the collecting localities of Kerivoula pellucida and K. whiteheadi. The land exposed during the Pleistocene Last Glacial Maximum is shown in light green." figureDoi="http://doi.org/10.5281/zenodo.3736821" httpUri="https://zenodo.org/record/3736821/files/figure.png" pageId="12" pageNumber="466">Fig 2</figureCitation>
). However,
<taxonomicName id="1CBF42C8F46FFFFFFB59D052FAD0FE0C" authorityName="Waterhouse" authorityYear="1854" box="[1218,1355,369,395]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="pellucida">
<emphasis id="E9CBE559F46FFFFFFB59D052FB4EFE0C" box="[1218,1237,370,395]" italics="true" pageId="12" pageNumber="466">K</emphasis>
.
<emphasis id="E9CBE559F46FFFFFFB78D051FAD0FE0C" box="[1251,1355,369,395]" italics="true" pageId="12" pageNumber="466">pellucida</emphasis>
</taxonomicName>
occurs only on Greater Mindanao, Greater Negros-Panay, and Greater
<collectingRegion id="197BF7A9F46FFFFFFCE5D0B5FC7BFE28" box="[894,992,405,431]" country="Philippines" name="Palawan" pageId="12" pageNumber="466">Palawan</collectingRegion>
;
<taxonomicName id="1CBF42C8F46FFFFFFC6BD0B6FB17FE28" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[1008,1164,405,431]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F46FFFFFFC6BD0B6FB98FE28" box="[1008,1027,406,431]" italics="true" pageId="12" pageNumber="466">K</emphasis>
.
<emphasis id="E9CBE559F46FFFFFFB8ED0B5FB17FE28" box="[1045,1164,405,431]" italics="true" pageId="12" pageNumber="466">hardwickii</emphasis>
</taxonomicName>
only occurs in Greater Mindanao and Greater
<collectingRegion id="197BF7A9F46FFFFFFE06D099FE64FE54" box="[413,511,441,467]" country="Philippines" name="Palawan" pageId="12" pageNumber="466">Palawan</collectingRegion>
;
<taxonomicName id="1CBF42C8F46FFFFFFD97D09AFD0DFE54" baseAuthorityName="Temminck" baseAuthorityYear="1840" box="[524,662,441,467]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="papillosa">
<emphasis id="E9CBE559F46FFFFFFD97D09AFD84FE54" box="[524,543,442,467]" italics="true" pageId="12" pageNumber="466">K</emphasis>
.
<emphasis id="E9CBE559F46FFFFFFDB5D099FD0DFE54" box="[558,662,441,467]" italics="true" pageId="12" pageNumber="466">papillosa</emphasis>
</taxonomicName>
only occurs in Greater Luzon and Greater Mindanao (
<figureCitation id="438425CEF46FFFFFFA99D099FAD3FE54" box="[1282,1352,441,467]" captionStart="FIGURE 2" captionStartId="9.[151,250,1957,1982]" captionTargetBox="[255,1332,181,1931]" captionTargetId="figure@9.[255,1332,181,1931]" captionTargetPageId="9" captionText="FIGURE 2. Map of the Philippines indicating the collecting localities of Kerivoula pellucida and K. whiteheadi. The land exposed during the Pleistocene Last Glacial Maximum is shown in light green." figureDoi="http://doi.org/10.5281/zenodo.3736821" httpUri="https://zenodo.org/record/3736821/files/figure.png" pageId="12" pageNumber="466">Fig. 2</figureCitation>
). Thus, only one of the four species is widespread within the Philippine archipelago, and it (
<taxonomicName id="1CBF42C8F46FFFFFFBD1D0FEFB7DFE70" authorityName="Thomas" authorityYear="1894" box="[1098,1254,477,503]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F46FFFFFFBD1D0FEFBC6FE70" box="[1098,1117,478,503]" italics="true" pageId="12" pageNumber="466">K</emphasis>
.
<emphasis id="E9CBE559F46FFFFFFBF0D0FDFB7DFE70" box="[1131,1254,477,503]" italics="true" pageId="12" pageNumber="466">whiteheadi</emphasis>
</taxonomicName>
) exhibits only a moderate level of incipient speciation that began about 7 MYA.
</paragraph>
<paragraph id="DB00394BF46FFFFFFF5CD305FEEEFCBC" blockId="12.[151,1437,152,827]" pageId="12" pageNumber="466">
However, our genetic data (
<figureCitation id="438425CEF46FFFFFFE62D305FDDEFDB8" box="[505,581,549,575]" captionStart="FIGURE 1" captionStartId="8.[151,250,1870,1895]" captionTargetBox="[171,1417,179,1843]" captionTargetId="figure@8.[170,1417,179,1843]" captionTargetPageId="8" captionText="FIGURE 1. Phylogenetic relationships of East Asian Kerivoula and two outgroup species based on Cyt b haplotypes. This topology results from a Bayesian analysis. Numbers at each node correspond to the Bayesian posterior probabilities (left) and to the proportion of bootstrap support of a maximum likelihood reconstruction (right). If both analyses supported a node by 98% or more support, this node is symbolized with a plain circle. The letters to the right of each clade refer to the groups discussed in the text. The scale bar corresponds to a branch length of 0.05 substitution per site." figureDoi="http://doi.org/10.5281/zenodo.3736819" httpUri="https://zenodo.org/record/3736819/files/figure.png" pageId="12" pageNumber="466">Figs. 1</figureCitation>
,
<figureCitation id="438425CEF46FFFFFFDC9D305FDFAFDB8" box="[594,609,549,575]" captionStart="FIGURE 4" captionStartId="13.[151,250,1314,1339]" captionTargetBox="[155,1432,528,1286]" captionTargetId="figure@13.[151,1436,525,1289]" captionTargetPageId="13" captionText="FIGURE 4. Time-calibrated phylogeny of Kerivoula species based on Cyt b haplotypes. Outgroups were omitted for ease of readability. The greyed boxes represent 95% highest posterior density (HPD) intervals of node ages derived from a BEAST analysis. Time is expressed in million years." figureDoi="http://doi.org/10.5281/zenodo.3736826" httpUri="https://zenodo.org/record/3736826/files/figure.png" pageId="12" pageNumber="466">4</figureCitation>
) indicate the presence of two distinct clades within the Philippine
<taxonomicName id="1CBF42C8F46FFFFFFAA4D306FF41FDE4" baseAuthorityName="Horsfield" baseAuthorityYear="1824" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F46FFFFFFAA4D306FF41FDE4" italics="true" pageId="12" pageNumber="466">K. hardwickii</emphasis>
</taxonomicName>
lineage that are morphologically distinct, and our morphological data (
<figureCitation id="438425CEF46FFFFFFC67D369FBDBFDE4" box="[1020,1088,585,611]" captionStart="FIGURE 3" captionStartId="11.[151,250,1182,1207]" captionTargetBox="[177,1413,279,1153]" captionTargetId="figure@11.[170,1417,273,1157]" captionTargetPageId="11" captionText="FIGURE 3. Bivariate plot of condylo-canine length vs. height of braincase for specimens of Kerivoula summarized in Tables 1 and 3. Color indicates geographic region and symbol indicates taxon. Philippine specimens of each taxon are enclosed within polygons." figureDoi="http://doi.org/10.5281/zenodo.3736824" httpUri="https://zenodo.org/record/3736824/files/figure.png" pageId="12" pageNumber="466">Fig. 3</figureCitation>
;
<tableCitation id="963D0CF0F46FFFFFFBD6D369FB3BFDE4" box="[1101,1184,585,611]" captionStart="TABLE 3" captionStartId="12.[151,239,869,893]" captionTargetPageId="12" captionText="TABLE 3. Mean (± standard deviation) and range of condylo-canine length (CCL) and braincase height (BH) for those specimens not included in Table 1. n = sample size." httpUri="http://table.plazi.org/id/8FC069C3F46FFFFFFF0CD245FD27FC26" pageId="12" pageNumber="466" tableUuid="8FC069C3F46FFFFFFF0CD245FD27FC26">Table 3</tableCitation>
) show that both occur on Mindanao (
<figureCitation id="438425CEF46FFFFFFEA5D34DFE56FD00" box="[318,461,621,647]" captionStart="APPENDIX A" captionStartId="31.[151,289,154,178]" captionTargetPageId="31" captionText="APPENDIX A. List of all Kerivoula specimens examined. Specimens included in morphological analysis and comparisons are in bold. § Holotypes * Paratype." figureDoi="http://doi.org/10.5281/zenodo.3736846" httpUri="https://zenodo.org/record/3736846/files/figure.png" pageId="12" pageNumber="466">Appendix A</figureCitation>
). We take this as tentative evidence of speciation in
<taxonomicName id="1CBF42C8F46FFFFFFBBDD34DFB0DFD00" authorityName="Gray" authorityYear="1842" box="[1062,1174,621,647]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F46FFFFFFBBDD34DFB0DFD00" box="[1062,1174,621,647]" italics="true" pageId="12" pageNumber="466">Kerivoula</emphasis>
</taxonomicName>
within the archipelago that has progressed to the point of secondary sympatry. This requires further study, but suggests that although in places where more than one species of
<taxonomicName id="1CBF42C8F46FFFFFFDC8D395FD58FD48" authorityName="Gray" authorityYear="1842" box="[595,707,693,719]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="genus">
<emphasis id="E9CBE559F46FFFFFFDC8D395FD58FD48" box="[595,707,693,719]" italics="true" pageId="12" pageNumber="466">Kerivoula</emphasis>
</taxonomicName>
is present, each is usually the product of a separate colonization from other regions in Southeast Asia. Where divergence dates among sympatric taxa are deeper, in the case of the
<taxonomicName id="1CBF42C8F46FFFFFFF0CD3DEFEAAFC90" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[151,305,765,791]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F46FFFFFFF0CD3DEFEAAFC90" box="[151,305,765,791]" italics="true" pageId="12" pageNumber="466">K. hardwickii</emphasis>
</taxonomicName>
group, some full diversification within the archipelago may have taken place, in this oldest of the Philippine lineages.
</paragraph>
<caption id="8FC069C3F46FFFFFFF0CD245FD27FC26" ID-Table-UUID="8FC069C3F46FFFFFFF0CD245FD27FC26" httpUri="http://table.plazi.org/id/8FC069C3F46FFFFFFF0CD245FD27FC26" pageId="12" pageNumber="466" targetBox="[166,1327,946,2023]" targetIsTable="true" targetPageId="12">
<paragraph id="DB00394BF46FFFFFFF0CD245FD27FC26" blockId="12.[151,1436,869,929]" pageId="12" pageNumber="466">
<emphasis id="E9CBE559F46FFFFFFF0CD245FE90FCFA" bold="true" box="[151,267,869,893]" pageId="12" pageNumber="466">TABLE 3.</emphasis>
Mean (± standard deviation) and range of condylo-canine length (CCL) and braincase height (BH) for those specimens not included in Table 1. n = sample size.
</paragraph>
</caption>
<paragraph id="DB00394BF46FFFFFFF3DD292FAB3F860" pageId="12" pageNumber="466">
<table id="A9BFCBEBF46F000CFF3DD292FAB4F860" box="[166,1327,946,2023]" gridcols="4" gridrows="20" pageId="12" pageNumber="466">
<tr id="658F3B09F46F000CFF3DD292FAB4FC69" box="[166,1327,946,1006]" gridrow="0" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD292FE85FC69" box="[166,286,946,1006]" gridcol="0" gridrow="0" pageId="12" pageNumber="466">Taxon Locality</th>
<th id="265E5275F46F000CFD0ED292FD35FC69" box="[661,686,946,1006]" gridcol="1" gridrow="0" pageId="12" pageNumber="466">n</th>
<th id="265E5275F46F000CFCC2D292FC42FC69" box="[857,985,946,1006]" gridcol="2" gridrow="0" pageId="12" pageNumber="466">CCL</th>
<th id="265E5275F46F000CFB27D292FAB4FC69" box="[1212,1327,946,1006]" gridcol="3" gridrow="0" pageId="12" pageNumber="466">BH</th>
</tr>
<tr id="658F3B09F46F000CFF3DD2DEFAB4FB91" box="[166,1327,1022,1046]" gridrow="1" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD2DEFAB4FB91" box="[166,1327,1022,1046]" colspan="4" colspanRight="3" gridcol="0" gridrow="1" pageId="12" pageNumber="466">
<taxonomicName id="1CBF42C8F46FFFFFFF3DD2DEFEB5FB91" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[166,302,1022,1046]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F46FFFFFFF3DD2DEFEB5FB91" box="[166,302,1022,1046]" italics="true" pageId="12" pageNumber="466">K. hardwickii</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="658F3B09F46F000CFF3DD505FAB4FBE5" box="[166,1327,1061,1122]" gridrow="2" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD505FE85FBE5" box="[166,286,1061,1122]" gridcol="0" gridrow="2" pageId="12" pageNumber="466">Borneo</th>
<td id="265E5275F46F000CFD0ED505FD35FBE5" box="[661,686,1061,1122]" gridcol="1" gridrow="2" pageId="12" pageNumber="466">5</td>
<td id="265E5275F46F000CFCC2D505FC42FBE5" box="[857,985,1061,1122]" gridcol="2" gridrow="2" pageId="12" pageNumber="466">12.09 ± 0.43 11.5212.63</td>
<td id="265E5275F46F000CFB27D505FAB4FBE5" box="[1212,1327,1061,1122]" gridcol="3" gridrow="2" pageId="12" pageNumber="466">5.36 ± 0.12 5.165.45</td>
</tr>
<tr id="658F3B09F46F000CFF3DD551FAB4FB29" box="[166,1327,1137,1198]" gridrow="3" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD551FE85FB29" box="[166,286,1137,1198]" gridcol="0" gridrow="3" pageId="12" pageNumber="466">mainland</th>
<td id="265E5275F46F000CFD0ED551FD35FB29" box="[661,686,1137,1198]" gridcol="1" gridrow="3" pageId="12" pageNumber="466">15</td>
<td id="265E5275F46F000CFCC2D551FC42FB29" box="[857,985,1137,1198]" gridcol="2" gridrow="3" pageId="12" pageNumber="466">12.60 ± 0.47 11.9013.48</td>
<td id="265E5275F46F000CFB27D551FAB4FB29" box="[1212,1327,1137,1198]" gridcol="3" gridrow="3" pageId="12" pageNumber="466">5.11 ± 0.36 4.575.78</td>
</tr>
<tr id="658F3B09F46F000CFF3DD59EFAB4FB50" box="[166,1327,1214,1239]" gridrow="4" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD59EFE85FB50" box="[166,286,1214,1239]" gridcol="0" gridrow="4" pageId="12" pageNumber="466">Sulawesi</th>
<td id="265E5275F46F000CFD0ED59EFD35FB50" box="[661,686,1214,1239]" gridcol="1" gridrow="4" pageId="12" pageNumber="466">1</td>
<td id="265E5275F46F000CFCC2D59EFC42FB50" box="[857,985,1214,1239]" gridcol="2" gridrow="4" pageId="12" pageNumber="466">11.45</td>
<td id="265E5275F46F000CFB27D59EFAB4FB50" box="[1212,1327,1214,1239]" gridcol="3" gridrow="4" pageId="12" pageNumber="466">5.37</td>
</tr>
<tr id="658F3B09F46F000CFF3DD5C7FAB4FB78" box="[166,1327,1255,1279]" gridrow="5" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD5C7FAB4FB78" box="[166,1327,1255,1279]" colspan="4" colspanRight="3" gridcol="0" gridrow="5" pageId="12" pageNumber="466">
<emphasis id="E9CBE559F46FFFFFFF3DD5C7FF25FB79" box="[166,190,1255,1278]" italics="true" pageId="12" pageNumber="466">K.</emphasis>
<taxonomicName id="1CBF42C8F46FFFFFFF55D5C7FEA2FB78" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[206,313,1255,1279]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="hardwickii">
<emphasis id="E9CBE559F46FFFFFFF55D5C7FEA2FB78" box="[206,313,1255,1279]" italics="true" pageId="12" pageNumber="466">hardwickii</emphasis>
</taxonomicName>
A”
</th>
</tr>
<tr id="658F3B09F46F000CFF3DD42EFAB4FACC" box="[166,1327,1294,1355]" gridrow="6" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD42EFE85FACC" box="[166,286,1294,1355]" gridcol="0" gridrow="6" pageId="12" pageNumber="466">Philippines</th>
<td id="265E5275F46F000CFD0ED42EFD35FACC" box="[661,686,1294,1355]" gridcol="1" gridrow="6" pageId="12" pageNumber="466">9</td>
<td id="265E5275F46F000CFCC2D42EFC42FACC" box="[857,985,1294,1355]" gridcol="2" gridrow="6" pageId="12" pageNumber="466">11.45 ± 0.54 10.8012.37</td>
<td id="265E5275F46F000CFB27D42EFAB4FACC" box="[1212,1327,1294,1355]" gridcol="3" gridrow="6" pageId="12" pageNumber="466">5.54 ± 0.10 5.345.54</td>
</tr>
<tr id="658F3B09F46F000CFF3DD47BFAB4FAF4" box="[166,1327,1371,1395]" gridrow="7" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD47BFAB4FAF4" box="[166,1327,1371,1395]" colspan="4" colspanRight="3" gridcol="0" gridrow="7" pageId="12" pageNumber="466">
<emphasis id="E9CBE559F46FFFFFFF3DD47CFEA6FAF4" box="[166,317,1371,1395]" italics="true" pageId="12" pageNumber="466">
K. “
<taxonomicName id="1CBF42C8F46FFFFFFF51D47BFEA6FAF4" baseAuthorityName="Horsfield" baseAuthorityYear="1824" box="[202,317,1371,1395]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="hardwickii">hardwickii</taxonomicName>
</emphasis>
B”
</th>
</tr>
<tr id="658F3B09F46F000CFF3DD4A3FAB4FA47" box="[166,1327,1411,1472]" gridrow="8" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD4A3FE85FA47" box="[166,286,1411,1472]" gridcol="0" gridrow="8" pageId="12" pageNumber="466">Philippines</th>
<td id="265E5275F46F000CFD0ED4A3FD35FA47" box="[661,686,1411,1472]" gridcol="1" gridrow="8" pageId="12" pageNumber="466">12</td>
<td id="265E5275F46F000CFCC2D4A3FC42FA47" box="[857,985,1411,1472]" gridcol="2" gridrow="8" pageId="12" pageNumber="466">13.17 ± 0.26 12.6413.52</td>
<td id="265E5275F46F000CFB27D4A3FAB4FA47" box="[1212,1327,1411,1472]" gridcol="3" gridrow="8" pageId="12" pageNumber="466">5.66 ± 0.16 5.465.94</td>
</tr>
<tr id="658F3B09F46F000CFF3DD4F0FAB4FA6F" box="[166,1327,1488,1512]" gridrow="9" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD4F0FAB4FA6F" box="[166,1327,1488,1512]" colspan="4" colspanRight="3" gridcol="0" gridrow="9" pageId="12" pageNumber="466">
<taxonomicName id="1CBF42C8F46FFFFFFF3DD4F0FEB0FA6F" authorityName="Hill &amp; Francis" authorityYear="1984" box="[166,299,1488,1512]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="intermedia">
<emphasis id="E9CBE559F46FFFFFFF3DD4F0FEB0FA6F" box="[166,299,1488,1512]" italics="true" pageId="12" pageNumber="466">K intermedia</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="658F3B09F46F000CFF3DD4D7FAB4F997" box="[166,1327,1527,1552]" gridrow="10" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD4D7FE85F997" box="[166,286,1527,1552]" gridcol="0" gridrow="10" pageId="12" pageNumber="466">Borneo</th>
<td id="265E5275F46F000CFD0ED4D7FD35F997" box="[661,686,1527,1552]" gridcol="1" gridrow="10" pageId="12" pageNumber="466">1</td>
<td id="265E5275F46F000CFCC2D4D7FC42F997" box="[857,985,1527,1552]" gridcol="2" gridrow="10" pageId="12" pageNumber="466">10.89</td>
<td id="265E5275F46F000CFB27D4D7FAB4F997" box="[1212,1327,1527,1552]" gridcol="3" gridrow="10" pageId="12" pageNumber="466">4.46</td>
</tr>
<tr id="658F3B09F46F000CFF3DD701FAB4F9BF" box="[166,1327,1569,1592]" gridrow="11" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD701FAB4F9BF" box="[166,1327,1569,1592]" colspan="4" colspanRight="3" gridcol="0" gridrow="11" pageId="12" pageNumber="466">
<taxonomicName id="1CBF42C8F46FFFFFFF3DD701FE92F9BF" authorityName="Miller" authorityYear="1898" box="[166,265,1569,1592]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="minuta">
<emphasis id="E9CBE559F46FFFFFFF3DD701FE92F9BF" box="[166,265,1569,1592]" italics="true" pageId="12" pageNumber="466">K. minuta</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="658F3B09F46F000CFF3DD769FAB4F9E6" box="[166,1327,1609,1633]" gridrow="12" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD769FE85F9E6" box="[166,286,1609,1633]" gridcol="0" gridrow="12" pageId="12" pageNumber="466">Borneo</th>
<td id="265E5275F46F000CFD0ED769FD35F9E6" box="[661,686,1609,1633]" gridcol="1" gridrow="12" pageId="12" pageNumber="466">1</td>
<td id="265E5275F46F000CFCC2D769FC42F9E6" box="[857,985,1609,1633]" gridcol="2" gridrow="12" pageId="12" pageNumber="466">10.02</td>
<td id="265E5275F46F000CFB27D769FAB4F9E6" box="[1212,1327,1609,1633]" gridcol="3" gridrow="12" pageId="12" pageNumber="466">4.34</td>
</tr>
<tr id="658F3B09F46F000CFF3DD750FAB4F90E" box="[166,1327,1648,1673]" gridrow="13" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD750FE85F90E" box="[166,286,1648,1673]" gridcol="0" gridrow="13" pageId="12" pageNumber="466">mainland</th>
<td id="265E5275F46F000CFD0ED750FD35F90E" box="[661,686,1648,1673]" gridcol="1" gridrow="13" pageId="12" pageNumber="466">1</td>
<td id="265E5275F46F000CFCC2D750FC42F90E" box="[857,985,1648,1673]" gridcol="2" gridrow="13" pageId="12" pageNumber="466">10.55</td>
<td id="265E5275F46F000CFB27D750FAB4F90E" box="[1212,1327,1648,1673]" gridcol="3" gridrow="13" pageId="12" pageNumber="466">4.25</td>
</tr>
<tr id="658F3B09F46F000CFF3DD7B9FAB4F936" box="[166,1327,1689,1713]" gridrow="14" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD7B9FAB4F936" box="[166,1327,1689,1713]" colspan="4" colspanRight="3" gridcol="0" gridrow="14" pageId="12" pageNumber="466">
<taxonomicName id="1CBF42C8F46FFFFFFF3DD7BAFEBAF936" baseAuthorityName="Temminck" baseAuthorityYear="1840" box="[166,289,1689,1713]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="papillosa">
<emphasis id="E9CBE559F46FFFFFFF3DD7BAFEBAF936" box="[166,289,1689,1713]" italics="true" pageId="12" pageNumber="466">K. papillosa</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="658F3B09F46F000CFF3DD7E1FAB4F95D" box="[166,1327,1729,1754]" gridrow="15" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD7E1FE85F95D" box="[166,286,1729,1754]" gridcol="0" gridrow="15" pageId="12" pageNumber="466">Borneo</th>
<td id="265E5275F46F000CFD0ED7E1FD35F95D" box="[661,686,1729,1754]" gridcol="1" gridrow="15" pageId="12" pageNumber="466">1</td>
<td id="265E5275F46F000CFCC2D7E1FC42F95D" box="[857,985,1729,1754]" gridcol="2" gridrow="15" pageId="12" pageNumber="466">16.89</td>
<td id="265E5275F46F000CFB27D7E1FAB4F95D" box="[1212,1327,1729,1754]" gridcol="3" gridrow="15" pageId="12" pageNumber="466">6.91</td>
</tr>
<tr id="658F3B09F46F000CFF3DD7C9FAB4F8A1" box="[166,1327,1769,1830]" gridrow="16" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD7C9FE85F8A1" box="[166,286,1769,1830]" gridcol="0" gridrow="16" pageId="12" pageNumber="466">mainland</th>
<td id="265E5275F46F000CFD0ED7C9FD35F8A1" box="[661,686,1769,1830]" gridcol="1" gridrow="16" pageId="12" pageNumber="466">6</td>
<td id="265E5275F46F000CFCC2D7C9FC42F8A1" box="[857,985,1769,1830]" gridcol="2" gridrow="16" pageId="12" pageNumber="466">15.82 ± 0.47 15.4316.54</td>
<td id="265E5275F46F000CFB27D7C9FAB4F8A1" box="[1212,1327,1769,1830]" gridcol="3" gridrow="16" pageId="12" pageNumber="466">6.79 ±0.17 6.657.13</td>
</tr>
<tr id="658F3B09F46F000CFF3DD615FAB4F8F5" box="[166,1327,1845,1906]" gridrow="17" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD615FE85F8F5" box="[166,286,1845,1906]" gridcol="0" gridrow="17" pageId="12" pageNumber="466">Philippines</th>
<td id="265E5275F46F000CFD0ED615FD35F8F5" box="[661,686,1845,1906]" gridcol="1" gridrow="17" pageId="12" pageNumber="466">6</td>
<td id="265E5275F46F000CFCC2D615FC42F8F5" box="[857,985,1845,1906]" gridcol="2" gridrow="17" pageId="12" pageNumber="466">15.79 ± 0.41 15.0816.28</td>
<td id="265E5275F46F000CFB27D615FAB4F8F5" box="[1212,1327,1845,1906]" gridcol="3" gridrow="17" pageId="12" pageNumber="466">7.03 ± 0.20 6.787.30</td>
</tr>
<tr id="658F3B09F46F000CFF3DD6A3FAB4F81D" box="[166,1327,1923,1946]" gridrow="18" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD6A3FAB4F81D" box="[166,1327,1923,1946]" colspan="4" colspanRight="3" gridcol="0" gridrow="18" pageId="12" pageNumber="466">
<taxonomicName id="1CBF42C8F46FFFFFFF3DD6A3FE9EF81D" authorityName="Bates, Struebig, Hayes, Furey, Mya Mya, Thong, Tien, Son, Harrison, Francis &amp; Csorba" authorityYear="2007" box="[166,261,1923,1946]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="12" pageNumber="466" phylum="Chordata" rank="species" species="titania">
<emphasis id="E9CBE559F46FFFFFFF3DD6A3FE9EF81D" box="[166,261,1923,1946]" italics="true" pageId="12" pageNumber="466">K. titania</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="658F3B09F46F000CFF3DD68AFAB4F860" box="[166,1327,1962,2023]" gridrow="19" pageId="12" pageNumber="466">
<th id="265E5275F46F000CFF3DD68AFE85F860" box="[166,286,1962,2023]" gridcol="0" gridrow="19" pageId="12" pageNumber="466">mainland</th>
<td id="265E5275F46F000CFD0ED68AFD35F860" box="[661,686,1962,2023]" gridcol="1" gridrow="19" pageId="12" pageNumber="466">3</td>
<td id="265E5275F46F000CFCC2D68AFC42F860" box="[857,985,1962,2023]" gridcol="2" gridrow="19" pageId="12" pageNumber="466">13.79 ± 0.07 13.7113.85</td>
<td id="265E5275F46F000CFB27D68AFAB4F860" box="[1212,1327,1962,2023]" gridcol="3" gridrow="19" pageId="12" pageNumber="466">5.48 ± 0.18 5.285.64</td>
</tr>
</table>
</paragraph>
<paragraph id="DB00394BF46EFFFEFF5CD1B9FE9CFE70" blockId="13.[151,1437,153,503]" pageId="13" pageNumber="467">
Finally, we note that
<taxonomicName id="1CBF42C8F46EFFFEFE28D1BAFDD4FF34" authorityName="Thomas" authorityYear="1894" box="[435,591,153,179]" class="Mammalia" family="Vespertilionidae" genus="Kerivoula" kingdom="Animalia" order="Chiroptera" pageId="13" pageNumber="467" phylum="Chordata" rank="species" species="whiteheadi">
<emphasis id="E9CBE559F46EFFFEFE28D1BAFDD4FF34" box="[435,591,153,179]" italics="true" pageId="13" pageNumber="467">K. whiteheadi</emphasis>
</taxonomicName>
from southern Luzon (clade
<emphasis id="E9CBE559F46EFFFEFC0ED1B9FC32FF34" bold="true" box="[917,937,153,179]" pageId="13" pageNumber="467">D</emphasis>
) are more closely related to specimens from the central
<collectingCountry id="A3A879DBF46EFFFEFE88D19DFE0AFF50" box="[275,401,189,215]" name="Philippines" pageId="13" pageNumber="467">Philippines</collectingCountry>
(clade
<emphasis id="E9CBE559F46EFFFEFE78D19DFE6DFF50" bold="true" box="[483,502,189,215]" pageId="13" pageNumber="467">E</emphasis>
) than they are to those from central and northern Luzon and adjacent islands (clade
<emphasis id="E9CBE559F46EFFFEFF0CD1C0FF30FF7C" bold="true" box="[151,171,224,251]" pageId="13" pageNumber="467">C</emphasis>
;
<figureCitation id="438425CEF46EFFFEFF20D1C1FE9AFF7C" box="[187,257,225,251]" captionStart="FIGURE 4" captionStartId="13.[151,250,1314,1339]" captionTargetBox="[155,1432,528,1286]" captionTargetId="figure@13.[151,1436,525,1289]" captionTargetPageId="13" captionText="FIGURE 4. Time-calibrated phylogeny of Kerivoula species based on Cyt b haplotypes. Outgroups were omitted for ease of readability. The greyed boxes represent 95% highest posterior density (HPD) intervals of node ages derived from a BEAST analysis. Time is expressed in million years." figureDoi="http://doi.org/10.5281/zenodo.3736826" httpUri="https://zenodo.org/record/3736826/files/figure.png" pageId="13" pageNumber="467">Fig. 4</figureCitation>
). Although it is initially surprising that all populations from Luzon are not monophyletic, the geological history of the
<collectingCountry id="A3A879DBF46EFFFEFEAAD025FE34FE98" box="[305,431,261,287]" name="Philippines" pageId="13" pageNumber="467">Philippines</collectingCountry>
is consistent with this. The estimated divergence date of 5.8 MYA for clades
<emphasis id="E9CBE559F46EFFFEFA9FD025FA83FE98" bold="true" box="[1284,1304,261,287]" pageId="13" pageNumber="467">D</emphasis>
and
<emphasis id="E9CBE559F46EFFFEFAD6D025FAFBFE98" bold="true" box="[1357,1376,261,287]" pageId="13" pageNumber="467">E</emphasis>
from clade
<emphasis id="E9CBE559F46EFFFEFF41D008FF75FEC4" bold="true" box="[218,238,296,323]" pageId="13" pageNumber="467">C</emphasis>
reflects the fact that southern Luzon (the area represented by clade
<emphasis id="E9CBE559F46EFFFEFC70D009FC64FEC4" bold="true" box="[1003,1023,297,323]" pageId="13" pageNumber="467">D</emphasis>
) was a distinct island that first arose about 6.6 MYA. However, this island did not become connected to central and northern Luzon until roughly 0.5 MYA when the Bondoc Peninsula and adjacent areas were uplifted (
<bibRefCitation id="BF2E44BAF46EFFFEFC08D051FBF0FE0C" author="Aurelio, M. A. &amp; Barrier, E. &amp; Rangin, C. &amp; Muller, C." box="[915,1131,369,395]" pageId="13" pageNumber="467" pagination="221 - 238" refId="ref15304" refString="Aurelio, M. A., Barrier, E., Rangin, C. &amp; Muller, C. (1991) The Philippine fault in the Late Cenozoic tectonic evolution of the Bondoc-Masbate-N. Leyte area, Central Philippines. Journal of Southeast Asian Earth Sciences, 6 (3 - 4), 221 - 238. https: // doi. org / 10.1016 / 0743 - 9547 (91) 90069 - A" type="journal article" year="1991">
Aurelio
<emphasis id="E9CBE559F46EFFFEFC6AD052FBB1FE0C" box="[1009,1066,369,395]" italics="true" pageId="13" pageNumber="467">et al.</emphasis>
1991
</bibRefCitation>
;
<bibRefCitation id="BF2E44BAF46EFFFEFBECD051FA03FE0C" author="Heaney, L. R. &amp; Balete, D. S. &amp; Rickart, E. A." box="[1143,1432,368,395]" pageId="13" pageNumber="467" refId="ref17242" refString="Heaney, L. R., Balete, D. S. &amp; Rickart, E. A. (2016) The Mammals of Luzon Island: Biogeography and Natural History of a Philippine Fauna. Johns Hopkins University Press, Baltimore. xi + 287 pp." type="book" year="2016">
Heaney
<emphasis id="E9CBE559F46EFFFEFB4FD052FA97FE0C" box="[1236,1292,369,395]" italics="true" pageId="13" pageNumber="467">et al.</emphasis>
2016:49-50
</bibRefCitation>
;
<figureCitation id="438425CEF46EFFFEFF0CD0B5FF47FE28" box="[151,220,405,431]" captionStart="FIGURE 2" captionStartId="9.[151,250,1957,1982]" captionTargetBox="[255,1332,181,1931]" captionTargetId="figure@9.[255,1332,181,1931]" captionTargetPageId="9" captionText="FIGURE 2. Map of the Philippines indicating the collecting localities of Kerivoula pellucida and K. whiteheadi. The land exposed during the Pleistocene Last Glacial Maximum is shown in light green." figureDoi="http://doi.org/10.5281/zenodo.3736821" httpUri="https://zenodo.org/record/3736821/files/figure.png" pageId="13" pageNumber="467">Fig. 2</figureCitation>
). Thus, the sister-group relationship of clades
<emphasis id="E9CBE559F46EFFFEFD7FD0B5FD63FE28" bold="true" box="[740,760,405,431]" pageId="13" pageNumber="467">D</emphasis>
and
<emphasis id="E9CBE559F46EFFFEFCB4D0B5FCD9FE28" bold="true" box="[815,834,405,431]" pageId="13" pageNumber="467">E</emphasis>
appears to reflect the prior history of southern Luzon and Greater Negros-Panay (including
<collectingRegion id="197BF7A9F46EFFFEFDA5D099FDE0FE53" box="[574,635,441,468]" country="Philippines" name="Cebu" pageId="13" pageNumber="467">Cebu</collectingRegion>
) as two separate but nearby islands at the time of this phylogenetic split, 5.8 MYA.
</paragraph>
</subSubSection>
</treatment>
</document>