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<document ID-DOI="http://doi.org/10.5281/zenodo.3610533" ID-GBIF-Dataset="3269bae6-36d8-4c4e-a814-cda94b6aeece" ID-GBIF-Taxon="161746393" ID-Zenodo-Dep="3610533" LSID="urn:lsid:zoobank.org:act:2A588C42-4DBE-41A3-A95A-7B6850521944" checkinTime="1579089070375" checkinUser="plazi" docAuthor="Rheindt, Frank E., Prawiradilaga, Dewi M., Ashar, Hidayat, Lee, Geraldine W. X., Wu, Meng Yue &amp; Ng, Nathaniel S. R." docDate="2020" docId="03F587A7FF968F06FF1272FCFC76FE92" docLanguage="en" docName="science.367.167-170_aax2146-Rheindt-SM.pdf" docOrigin="Science 36" docStyle="DocumentStyle{}" docTitle="Locustella portenta Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng 2020, species nova" docType="treatment" docUuid="2A588C42-4DBE-41A3-A95A-7B6850521944" docUuidSource="ZooBank" docVersion="11" lastPageId="25" lastPageNumber="25" masterDocId="FFCCFFDFFF9B8F1FFFD2714DFFBCFFEA" masterDocTitle="A lost world in Wallacea: Description of a montane archipelagic avifauna (supplement)" masterLastPageNumber="104" masterPageNumber="1" pageId="13" pageNumber="14" updateTime="1643451873296" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>A lost world in Wallacea: Description of a montane archipelagic avifauna (supplement)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rheindt, Frank E.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Prawiradilaga, Dewi M.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ashar, Hidayat</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lee, Geraldine W. X.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wu, Meng Yue</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ng, Nathaniel S. R.</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Science</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2020</mods:date>
<mods:detail type="pubDate">
<mods:number>2020-01-10</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>36</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>104</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://doi.org/10.5281/zenodo.3608758</mods:identifier>
<mods:identifier type="GBIF-Dataset">3269bae6-36d8-4c4e-a814-cda94b6aeece</mods:identifier>
<mods:identifier type="Zenodo-Dep">3608758</mods:identifier>
<mods:identifier type="ZooBank">8114B399-C68D-43C2-B6D3-B51AA898431E</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.3610533" ID-GBIF-Taxon="161746393" ID-Zenodo-Dep="3610533" LSID="urn:lsid:zoobank.org:act:2A588C42-4DBE-41A3-A95A-7B6850521944" httpUri="http://treatment.plazi.org/id/03F587A7FF968F06FF1272FCFC76FE92" lastPageId="25" lastPageNumber="25" pageId="13" pageNumber="14">
<subSubSection box="[192,291,945,983]" pageId="13" pageNumber="14" type="multiple">
<paragraph blockId="13.[192,1385,201,1860]" box="[192,291,945,983]" pageId="13" pageNumber="14">
<emphasis bold="true" box="[192,291,945,983]" pageId="13" pageNumber="14">SM3:</emphasis>
</paragraph>
</subSubSection>
<subSubSection box="[302,888,945,985]" pageId="13" pageNumber="14" type="nomenclature">
<paragraph blockId="13.[192,1385,201,1860]" box="[302,888,945,985]" pageId="13" pageNumber="14">
<heading bold="true" box="[302,888,945,985]" fontSize="16" level="5" pageId="13" pageNumber="14" reason="0">
<emphasis bold="true" box="[302,888,945,985]" pageId="13" pageNumber="14">
<taxonomicName authority="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng, 2020" authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[302,642,945,984]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="portenta" status="species nova">
<emphasis bold="true" box="[302,642,945,984]" italics="true" pageId="13" pageNumber="14">Locustella portenta</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[664,888,945,985]" pageId="13" pageNumber="14" rank="species">species nova</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="13" pageNumber="14" type="vernacular_names">
<paragraph blockId="13.[192,1385,201,1860]" pageId="13" pageNumber="14">
<emphasis bold="true" pageId="13" pageNumber="14">
(
<vernacularName pageId="13" pageNumber="14">Taliabu Grasshopper-Warbler</vernacularName>
;
</emphasis>
</paragraph>
</subSubSection>
<subSubSection pageId="13" pageNumber="14" type="nomenclature">
<paragraph blockId="13.[192,1385,201,1860]" pageId="13" pageNumber="14">
<emphasis bold="true" pageId="13" pageNumber="14">urn:lsid:zoobank.org:act:2A588C42-4DBE-41A-A95A-7B6850521944</emphasis>
</paragraph>
<paragraph blockId="13.[192,1385,201,1860]" pageId="13" pageNumber="14">
<emphasis bold="true" box="[574,588,1141,1179]" pageId="13" pageNumber="14">)</emphasis>
Frank E. Rheindt, Dewi M. Prawiradilaga, Hidayat Ashari, Suparno, Chyi Yin Gwee, Geraldine W. X. Lee
</paragraph>
</subSubSection>
<subSubSection pageId="13" pageNumber="14" type="materials_examined">
<materialsCitation ID-GBIF-Occurrence="2549955316" collectingDate="2013-12-12" collectionCode="MZB" collectorName="Rheindt &amp; LIPI field party" country="Indonesia" elevation="1200" latitude="-1.0132262" location="Waiyo dinahana Camp" longLatPrecision="1" longitude="124.01339" pageId="13" pageNumber="14" specimenCode="MZB.Ornit.34.411" specimenCount="2" specimenCount-adult="1" specimenCount-female="1" typeStatus="holotype">
<paragraph blockId="13.[192,1385,201,1860]" box="[192,316,1462,1490]" pageId="13" pageNumber="14">
<heading bold="true" box="[192,316,1462,1490]" fontSize="12" level="7" pageId="13" pageNumber="14" reason="2">
<emphasis bold="true" box="[192,316,1462,1490]" pageId="13" pageNumber="14">
<typeStatus box="[192,316,1462,1490]" pageId="13" pageNumber="14">Holotype</typeStatus>
</emphasis>
</heading>
</paragraph>
<paragraph blockId="13.[192,1385,201,1860]" pageId="13" pageNumber="14">
<specimenCode box="[192,433,1536,1564]" pageId="13" pageNumber="14">MZB.Ornit.34.411</specimenCode>
(
<figureCitation box="[450,540,1536,1564]" captionStart="Fig" captionStartId="75.[192,239,916,944]" captionTargetBox="[192,1344,192,870]" captionTargetId="figure@75.[192,1344,192,870]" captionTargetPageId="75" captionText="Fig. S3. Holotype of Locustella portenta (MZB.Ornit.34.411)." figureDoi="http://doi.org/10.5281/zenodo.3608764" httpUri="https://zenodo.org/record/3608764/files/figure.png" pageId="13" pageNumber="14">fig. S3</figureCitation>
);
<specimenCount box="[564,628,1536,1564]" pageId="13" pageNumber="14" type="adult">adult</specimenCount>
<specimenCount box="[636,723,1536,1564]" pageId="13" pageNumber="14" type="female">female</specimenCount>
collected
<date box="[855,1018,1536,1565]" pageId="13" pageNumber="14" value="2013-12-12">
<collectingDate box="[855,1018,1536,1565]" pageId="13" pageNumber="14" value="2013-12-12">12 Dec 2013</collectingDate>
</date>
near
<location LSID="urn:lsid:plazi:treatment:03F587A7FF968F06FF1272FCFC76FE92:8E83606AFF968F12FB90774DFADAF9F6" box="[1090,1382,1536,1564]" latitude="-1.0132262" longLatPrecision="1" longitude="124.01339" name="Waiyo dinahana Camp" pageId="13" pageNumber="14">Waiyo dinahana Camp</location>
(~
<quantity box="[217,311,1610,1638]" metricMagnitude="3" metricUnit="m" metricValue="1.2" pageId="13" pageNumber="14" unit="m" value="1200.0">
<elevation box="[217,311,1610,1638]" metricMagnitude="3" metricUnit="m" metricValue="1.2" pageId="13" pageNumber="14" unit="m" value="1200.0">1200m</elevation>
</quantity>
) on
<location LSID="urn:lsid:plazi:treatment:03F587A7FF968F06FF1272FCFC76FE92:8E83606AFF968F12FEA27707FD9BF98C" box="[368,551,1610,1638]" latitude="-1.0132262" longLatPrecision="1" longitude="124.01339" name="Taliabu Island" pageId="13" pageNumber="14">Taliabu Island</location>
(
<geoCoordinate box="[570,740,1610,1638]" degrees="01" direction="south" minutes="0" orientation="latitude" pageId="13" pageNumber="14" precision="1" seconds="47.614" value="-1.0132262">S 01⁰ 47.614</geoCoordinate>
';
<geoCoordinate box="[756,945,1610,1639]" degrees="124" direction="east" minutes="0" orientation="longitude" pageId="13" pageNumber="14" precision="1" seconds="48.216" value="124.01339">E 124⁰ 48.216</geoCoordinate>
'). Collected by the
<collectorName box="[1187,1288,1610,1638]" pageId="13" pageNumber="14">Rheindt</collectorName>
/
<collectorName pageId="13" pageNumber="14">LIPI field party</collectorName>
, including tissue samples from breast muscle and liver; skin prepared by Suparno; field number Tbu80; heavy body molt with fresh remiges; medium fat; weight
<quantity box="[1202,1253,1757,1785]" metricMagnitude="-2" metricUnit="kg" metricValue="1.9" pageId="13" pageNumber="14" unit="g" value="19.0">19g</quantity>
; wing length
<quantity box="[280,363,1830,1858]" metricMagnitude="-2" metricUnit="m" metricValue="5.9" pageId="13" pageNumber="14" unit="cm" value="5.9">5.9cm</quantity>
; wing spread
<quantity box="[539,638,1830,1858]" metricMagnitude="-1" metricUnit="m" metricValue="1.8399999999999999" pageId="13" pageNumber="14" unit="cm" value="18.4">18.4cm</quantity>
; total length
<quantity box="[804,903,1830,1858]" metricMagnitude="-1" metricUnit="m" metricValue="1.39" pageId="13" pageNumber="14" unit="cm" value="13.9">13.9cm</quantity>
; bill
<quantity box="[967,1050,1830,1858]" metricMagnitude="-2" metricUnit="m" metricValue="1.4" pageId="13" pageNumber="14" unit="cm" value="1.4">1.4cm</quantity>
; tail
<quantity box="[1112,1195,1830,1858]" metricMagnitude="-2" metricUnit="m" metricValue="5.4" pageId="13" pageNumber="14" unit="cm" value="5.4">5.4cm</quantity>
; tarsus
<quantity box="[1290,1373,1830,1858]" metricMagnitude="-2" metricUnit="m" metricValue="2.5" pageId="13" pageNumber="14" unit="cm" value="2.5">2.5cm</quantity>
.
</paragraph>
</materialsCitation>
</subSubSection>
<subSubSection pageId="14" pageNumber="15" type="description">
<paragraph blockId="14.[192,1390,201,1997]" box="[192,510,201,229]" pageId="14" pageNumber="15">
<heading bold="true" box="[192,510,201,229]" fontSize="12" level="7" pageId="14" pageNumber="15" reason="2">
<emphasis bold="true" box="[192,510,201,229]" pageId="14" pageNumber="15">Description of holotype</emphasis>
</heading>
</paragraph>
<paragraph blockId="14.[192,1390,201,1997]" pageId="14" pageNumber="15">
The entire crown from forehead to nape is rich earthen-brown (7.5YR 3/3) with slightly darker feather margins creating a scalloped appearance. A thin off-white supercilium is pronounced from bill base to just behind eye, from where it slowly tapers out. Lores are dark brownish-black (2.5Y 1.5/1). Ear coverts are of a medium earthen-brown tone (5YR 3/1) with fine pale shaft streaks giving a grizzled appearance, grading into a more greyish-brown tone (10YR 6/1) in the moustachial and malar region. The mantle and scapulars down to the uppertail coverts are largely concolorous with the crown but lack the scalloping created by darker feather margins. The remiges are mostly a rich cold dark-brown (2.5YR 2/1) but have warmer outer webs concolorous with mantle. The upperwing coverts are similarly two-toned, being mostly concolorous with mantle but some showing darker outer webs concolorous with most of remiges. As typical for the genus
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[728,861,1010,1038]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis box="[728,861,1010,1038]" italics="true" pageId="14" pageNumber="15">Locustella</emphasis>
</taxonomicName>
, the rectrices are narrow and dark- brown, very close in color to the remiges (2.5YR 2/1). The tail feathers are mostly worn, although one in the centre appears to be a new replacement.
</paragraph>
<paragraph blockId="14.[192,1390,201,1997]" pageId="14" pageNumber="15">The white chin and throat are quite sharply demarcated from the face. The lower throat and upper breast, still white, are interspersed with fine dusky speckling increasing towards the breast and lower throat sides where the color grades into a solid mid-grey (N5) with only few inconspicuous dusky speckles. The grey breast color then grades into a tawny- ochraceous belly color (7.5YR 4/6), first through tracts of grey feathers which have warm- ochraceous edging, then through tracts of largely ochraceous feathers that still have a grey feather centre, eventually to all-ochraceous feathers. The belly color becomes darker and richer on the undertail coverts and flanks (7.5YR 3/4).</paragraph>
<paragraph blockId="14.[192,1390,201,1997]" pageId="14" pageNumber="15">The bill is black with an inconspicuous paler tip. The gape on the live bird was slightly enlarged light yellow. The iris is dark brown. The tarsus and toes were described on the live bird as maroon-horn with paler, more yellowish toepads.</paragraph>
</subSubSection>
<subSubSection pageId="15" pageNumber="16" type="diagnosis">
<paragraph blockId="15.[192,1392,280,2002]" box="[192,324,280,308]" pageId="15" pageNumber="16">
<heading bold="true" box="[192,324,280,308]" fontSize="12" level="7" pageId="15" pageNumber="16" reason="2">
<emphasis bold="true" box="[192,324,280,308]" pageId="15" pageNumber="16">Diagnosis</emphasis>
</heading>
</paragraph>
<paragraph blockId="15.[192,1392,280,2002]" pageId="15" pageNumber="16">
A rich dark-brown forest warbler typical of the montane resident
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[1029,1162,353,381]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis box="[1029,1162,353,381]" italics="true" pageId="15" pageNumber="16">Locustella</emphasis>
</taxonomicName>
radiation from throughout Indonesia and the Philippines. As is usual across this cryptic genus, the new taxon is most conspicuous not on the basis of its plumage but its vocalizations.
</paragraph>
<paragraph blockId="15.[192,1392,280,2002]" pageId="15" pageNumber="16">
Morphologically, it is most similar in size and shape to
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[1000,1148,576,604]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[1000,1148,576,604]" italics="true" pageId="15" pageNumber="16">L. castanea</emphasis>
</taxonomicName>
of Sulawesi. However, in comparison to the new taxon,
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[744,892,649,677]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[744,892,649,677]" italics="true" pageId="15" pageNumber="16">L. castanea</emphasis>
</taxonomicName>
has less distinct speckling on the breast, has a more uniform crown that is less scalloped, paler and warmer brown upperparts, greyer underparts with brown mostly only on flanks, and has a less pointed, wider tail.
</paragraph>
<paragraph blockId="15.[192,1392,280,2002]" pageId="15" pageNumber="16">
<taxonomicName box="[288,441,868,898]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="musculus">
<emphasis box="[288,441,868,898]" italics="true" pageId="15" pageNumber="16">L. musculus</emphasis>
</taxonomicName>
from Seram differs in its less distinct central breast spotting, greyer (less whitish) supercilium, and more extensive grey on breast reaching further onto upper breast, neck sides and auriculars.
</paragraph>
<paragraph blockId="15.[192,1392,280,2002]" pageId="15" pageNumber="16">
<taxonomicName box="[288,457,1089,1119]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[288,457,1089,1119]" italics="true" pageId="15" pageNumber="16">L. disturbans</emphasis>
</taxonomicName>
from Buru differs in its much whiter breast, with off-white reaching onto mid-belly, whereas only the lower flanks and lower belly are warm-brown. Although the central breast of
<taxonomicName box="[406,575,1236,1266]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[406,575,1236,1266]" italics="true" pageId="15" pageNumber="16">L. disturbans</emphasis>
</taxonomicName>
has a mid-grey tinge, its underparts are much paler than the new species fairly uniform grey lower breast, which grades into a fairly dark homogeneous warm-brown belly.
</paragraph>
<paragraph blockId="15.[192,1392,280,2002]" pageId="15" pageNumber="16">
The new species is quite different from the smaller
<taxonomicName baseAuthorityName="Hartert" baseAuthorityYear="1896" box="[946,1065,1459,1487]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="montis">
<emphasis box="[946,1065,1459,1487]" italics="true" pageId="15" pageNumber="16">L. montis</emphasis>
</taxonomicName>
from Java. The latter is much paler cinnamon-brown on upperparts, crown and especially remiges and rectrices, and is uniformly buff-white below with extensive dark-brown speckling from chin to upper belly, lacking the new species grey tones on the breast and dark warm-brown tones on the belly.
</paragraph>
</subSubSection>
<subSubSection lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16" type="etymology">
<paragraph blockId="15.[192,1392,280,2002]" box="[192,340,1825,1853]" pageId="15" pageNumber="16">
<heading bold="true" box="[192,340,1825,1853]" fontSize="12" level="7" pageId="15" pageNumber="16" reason="2">
<emphasis bold="true" box="[192,340,1825,1853]" pageId="15" pageNumber="16">Etymology</emphasis>
</heading>
</paragraph>
<paragraph blockId="15.[192,1392,280,2002]" lastBlockId="16.[192,1394,201,1711]" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16">
The name
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[327,435,1901,1929]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[327,435,1901,1929]" italics="true" pageId="15" pageNumber="16">portenta</emphasis>
</taxonomicName>
past participle of Latin portendere roughly translates as “the predicted one” or “the foretold one” and relates to the unusual circumstances of discovery when FER first heard the new warbler and recognized it to be a novel species of
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[1225,1358,201,229]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis box="[1225,1358,201,229]" italics="true" pageId="16" pageNumber="17">Locustella</emphasis>
</taxonomicName>
multiple days before a visual confirmation was obtained.
</paragraph>
</subSubSection>
<paragraph blockId="15.[778,808,2017,2044]" box="[778,808,2017,2044]" pageId="15" pageNumber="16">
<pageNumber ambiguity="1" box="[778,808,2017,2044]" fuzzyness="0" pageId="15" pageNumber="16" score="4036.0" value="16">
<pageTitle box="[778,808,2017,2044]" pageId="15" pageNumber="16">16</pageTitle>
</pageNumber>
</paragraph>
<subSubSection pageId="16" pageNumber="17" type="discussion">
<paragraph blockId="16.[192,1394,201,1711]" box="[192,976,427,455]" pageId="16" pageNumber="17">
<heading bold="true" box="[192,976,427,455]" fontSize="12" level="7" pageId="16" pageNumber="17" reason="2">
<emphasis bold="true" box="[192,976,427,455]" pageId="16" pageNumber="17">Individual, sex and age-related variation within the taxon</emphasis>
</heading>
</paragraph>
<paragraph blockId="16.[192,1394,201,1711]" pageId="16" pageNumber="17">Currently only one adult specimen is available (i.e., the holotype), and photos of up to two more adults suggest that this is probably a sexually monomorphic species with limited plumage variation.</paragraph>
<paragraph blockId="16.[192,1394,201,1711]" box="[192,468,800,828]" pageId="16" pageNumber="17">
<heading bold="true" box="[192,468,800,828]" fontSize="12" level="7" pageId="16" pageNumber="17" reason="2">
<emphasis bold="true" box="[192,468,800,828]" pageId="16" pageNumber="17">History of discovery</emphasis>
</heading>
</paragraph>
<paragraph blockId="16.[192,1394,201,1711]" pageId="16" pageNumber="17">
FER was the first person to discover
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[666,808,876,904]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[666,808,876,904]" italics="true" pageId="16" pageNumber="17">L. portenta</emphasis>
</taxonomicName>
on 9 April 2009 near its type locality when he heard an unmistakable insect-like sound that he attributed to a hitherto unnamed taxon of
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[192,325,1021,1049]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="16" pageNumber="17" phylum="Chordata" rank="genus">
<emphasis box="[192,325,1021,1049]" italics="true" pageId="16" pageNumber="17">Locustella</emphasis>
</taxonomicName>
grasshopper-warbler. Having been exposed to the song of
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[1079,1227,1023,1051]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[1079,1227,1023,1051]" italics="true" pageId="16" pageNumber="17">L. castanea</emphasis>
</taxonomicName>
on a visit to Sulawesi only two weeks prior, and having had previous field experience with
<taxonomicName box="[1202,1371,1095,1125]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[1202,1371,1095,1125]" italics="true" pageId="16" pageNumber="17">L. disturbans</emphasis>
</taxonomicName>
and
<taxonomicName box="[246,399,1169,1198]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="musculus">
<emphasis box="[246,399,1169,1198]" italics="true" pageId="16" pageNumber="17">L. musculus</emphasis>
</taxonomicName>
from Buru and Seram, respectively (
<emphasis box="[875,907,1169,1197]" italics="true" pageId="16" pageNumber="17">8 0</emphasis>
), FER was aware that the song of this population albeit superficially reminiscent of the others is distinct at the species level. Even so, because of the retiring nature of this bird and ill-timed heavy rain, it took FER another two hiking expeditions to the highlands of Taliabu to finally confirm on 15 April 2009 that this song was indeed uttered by what he perceived as an “unusually dark” individual of grasshopper-warbler (
<bibRefCitation author="F. E. Rheindt" box="[646,678,1536,1564]" journalOrPublisher="Bull. Br. Ornithol. Club" pageId="16" pageNumber="17" pagination="33 - 51" part="130" refId="ref29294" refString="48. F. E. Rheindt, New biogeographic records for the avifauna of Taliabu (Sula Islands, Indonesia), with the preliminary documentation of two previously undiscovered taxa. Bull. Br. Ornithol. Club 130, 33 - 51 (2010)." title="New biogeographic records for the avifauna of Taliabu (Sula Islands, Indonesia), with the preliminary documentation of two previously undiscovered taxa" type="journal article" year="2010">
<emphasis box="[646,678,1536,1564]" italics="true" pageId="16" pageNumber="17">48</emphasis>
</bibRefCitation>
). We encountered this population again during our collecting trip to this area from 6-16 Dec 2013 (
<bibRefCitation author="F. E. Rheindt &amp; D. M. Prawiradilaga &amp; S. Suparno &amp; H. Ashari &amp; P. R. Wilton" box="[806,838,1610,1638]" journalOrPublisher="Treubia" pageId="16" pageNumber="17" pagination="61 - 90" part="41" refId="ref27737" refString="19. F. E. Rheindt, D. M. Prawiradilaga, S. Suparno, H. Ashari, P. R. Wilton, New and significant island records, range extensions and elevational extensions of birds in eastern Sulawesi, its nearby satellites, and Ternate. Treubia 41, 61 - 90 (2014)." title="New and significant island records, range extensions and elevational extensions of birds in eastern Sulawesi, its nearby satellites, and Ternate" type="journal article" year="2014">
<emphasis box="[806,838,1610,1638]" italics="true" pageId="16" pageNumber="17">19</emphasis>
</bibRefCitation>
) and collected a single specimen, now the holotype.
</paragraph>
</subSubSection>
<subSubSection lastPageId="18" lastPageNumber="19" pageId="16" pageNumber="17" type="distribution">
<paragraph blockId="16.[192,507,1835,1863]" box="[192,507,1835,1863]" pageId="16" pageNumber="17">
<heading bold="true" box="[192,507,1835,1863]" fontSize="12" level="7" pageId="16" pageNumber="17" reason="2">
<emphasis bold="true" box="[192,507,1835,1863]" pageId="16" pageNumber="17">Distribution and status</emphasis>
</heading>
</paragraph>
<paragraph blockId="16.[192,1362,1910,1939]" lastBlockId="17.[192,1393,201,1997]" lastPageId="17" lastPageNumber="18" pageId="16" pageNumber="17">
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[192,334,1911,1939]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[192,334,1911,1939]" italics="true" pageId="16" pageNumber="17">L. portenta</emphasis>
</taxonomicName>
is presumably restricted to montane forest at the highest elevations of the island of Taliabu (
<emphasis box="[343,423,201,229]" italics="true" pageId="17" pageNumber="18">
<bibRefCitation author="F. E. Rheindt &amp; D. M. Prawiradilaga &amp; S. Suparno &amp; H. Ashari &amp; P. R. Wilton" box="[343,378,201,229]" journalOrPublisher="Treubia" pageId="17" pageNumber="18" pagination="61 - 90" part="41" refId="ref27737" refString="19. F. E. Rheindt, D. M. Prawiradilaga, S. Suparno, H. Ashari, P. R. Wilton, New and significant island records, range extensions and elevational extensions of birds in eastern Sulawesi, its nearby satellites, and Ternate. Treubia 41, 61 - 90 (2014)." title="New and significant island records, range extensions and elevational extensions of birds in eastern Sulawesi, its nearby satellites, and Ternate" type="journal article" year="2014">19</bibRefCitation>
,
<bibRefCitation author="F. E. Rheindt" box="[391,423,201,229]" journalOrPublisher="Bull. Br. Ornithol. Club" pageId="17" pageNumber="18" pagination="33 - 51" part="130" refId="ref29294" refString="48. F. E. Rheindt, New biogeographic records for the avifauna of Taliabu (Sula Islands, Indonesia), with the preliminary documentation of two previously undiscovered taxa. Bull. Br. Ornithol. Club 130, 33 - 51 (2010)." title="New biogeographic records for the avifauna of Taliabu (Sula Islands, Indonesia), with the preliminary documentation of two previously undiscovered taxa" type="journal article" year="2010">48</bibRefCitation>
</emphasis>
) (
<figureCitation box="[451,542,201,229]" captionStart="Fig" captionStartId="73.[192,239,1773,1801]" captionTargetBox="[192,1344,192,1728]" captionTargetId="figure@73.[192,1344,192,1728]" captionText="Fig. S2. Google Earth satellite map of Peleng and Banggai (top) and the western half of Taliabu (bottom). Peleng and Banggai are the two largest islands of the “Banggai Archipelago”. Our main collection site, an unnamed locality “above Kokolomboi [village]”, is mapped with a yellow pin. Areas above 800m elevation are circled with a red line, indicating the presence of montane vegetation. The island of Taliabu is the largest member of the “Sula Archipelago”. Our main collection locality “Waiyo dinahana” is mapped with a yellow pin. Areas above 1,050 m elevation are circled with a red line, indicating the presence of a peculiar montane stunted forest vegetation. Within the montane zone above 1,050m, areas degraded by logging activity as reflected on Google Earth satellite imagery are mapped in turquoise and areas burnt down by extensive fires in the 1980s are mapped in dark-green. Given the outdated status of some of the satellite imagery, present-day forest degradation is likely more extensive than shown." figureDoi="http://doi.org/10.5281/zenodo.3608762" httpUri="https://zenodo.org/record/3608762/files/figure.png" pageId="17" pageNumber="18">fig. S2</figureCitation>
). All our records are from elevations above 1,050m on the more mountainous western half of the island of Taliabu, which reaches its highest altitude of ~1,415m at an unnamed peak near our collection locality (
<figureCitation box="[938,1029,348,376]" captionStart="Fig" captionStartId="73.[192,239,1773,1801]" captionTargetBox="[192,1344,192,1728]" captionTargetId="figure@73.[192,1344,192,1728]" captionText="Fig. S2. Google Earth satellite map of Peleng and Banggai (top) and the western half of Taliabu (bottom). Peleng and Banggai are the two largest islands of the “Banggai Archipelago”. Our main collection site, an unnamed locality “above Kokolomboi [village]”, is mapped with a yellow pin. Areas above 800m elevation are circled with a red line, indicating the presence of montane vegetation. The island of Taliabu is the largest member of the “Sula Archipelago”. Our main collection locality “Waiyo dinahana” is mapped with a yellow pin. Areas above 1,050 m elevation are circled with a red line, indicating the presence of a peculiar montane stunted forest vegetation. Within the montane zone above 1,050m, areas degraded by logging activity as reflected on Google Earth satellite imagery are mapped in turquoise and areas burnt down by extensive fires in the 1980s are mapped in dark-green. Given the outdated status of some of the satellite imagery, present-day forest degradation is likely more extensive than shown." figureDoi="http://doi.org/10.5281/zenodo.3608762" httpUri="https://zenodo.org/record/3608762/files/figure.png" pageId="17" pageNumber="18">fig. S2</figureCitation>
). The new species inhabits a peculiar type of dwarf montane forest typical of these elevations.
</paragraph>
<paragraph blockId="17.[192,1393,201,1997]" pageId="17" pageNumber="18">
The occurrence of the new species on neighboring islands is highly doubtful. Of the three major islands that form the Sula archipelago, Taliabu is the largest and tallest. The smallest main island, Sanana, barely reaches above 600m elevation. The highest elevation on its larger neighbor Mangole is 1,127m theoretically within the elevational range of this grasshopper-warbler but the area of land above 1,050m on Mangole is only ~36 ha, which is too small an area to allow for the survival of a highly montane bird species throughout evolutionary time scales. On the neighboring Banggai archipelago, the situation is similar: the only island high enough to allow for the theoretical possibility of grasshopper-warbler breeding occurrence is Peleng (highest elevation ~1,022m), but extensive fieldwork around this highest area has not revealed the presence of any resident grasshopper-warblers (
<emphasis box="[1283,1363,1158,1186]" italics="true" pageId="17" pageNumber="18">
<bibRefCitation author="F. E. Rheindt &amp; D. M. Prawiradilaga &amp; S. Suparno &amp; H. Ashari &amp; P. R. Wilton" box="[1283,1318,1158,1186]" journalOrPublisher="Treubia" pageId="17" pageNumber="18" pagination="61 - 90" part="41" refId="ref27737" refString="19. F. E. Rheindt, D. M. Prawiradilaga, S. Suparno, H. Ashari, P. R. Wilton, New and significant island records, range extensions and elevational extensions of birds in eastern Sulawesi, its nearby satellites, and Ternate. Treubia 41, 61 - 90 (2014)." title="New and significant island records, range extensions and elevational extensions of birds in eastern Sulawesi, its nearby satellites, and Ternate" type="journal article" year="2014">19</bibRefCitation>
,
<bibRefCitation author="F. E. Rheindt &amp; F. Verbelen &amp; D. D. Putra &amp; A. Rahman &amp; M. Indrawan" box="[1331,1363,1158,1186]" journalOrPublisher="Bull. Br. Ornithol. Club" pageId="17" pageNumber="18" pagination="181 - 207" part="130" refId="ref29343" refString="49. F. E. Rheindt, F. Verbelen, D. D. Putra, A. Rahman, M. Indrawan, New biogeographic records in the avifauna of Peleng Island (Sulawesi, Indonesia), with taxonomic notes on some endemic taxa. Bull. Br. Ornithol. Club 130, 181 - 207 (2010)." title="New biogeographic records in the avifauna of Peleng Island (Sulawesi, Indonesia), with taxonomic notes on some endemic taxa" type="journal article" year="2010">49</bibRefCitation>
</emphasis>
).
</paragraph>
<paragraph blockId="17.[192,1393,201,1997]" lastBlockId="18.[192,1383,201,2002]" lastPageId="18" lastPageNumber="19" pageId="17" pageNumber="18">
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[288,430,1233,1261]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[288,430,1233,1261]" italics="true" pageId="17" pageNumber="18">L. portenta</emphasis>
</taxonomicName>
must be a rare bird: Taliabus area above 1,050m amounts to ~16,690 ha of formerly forested land in the interior of the islands western half, over 97% of which forms one contiguous block (
<figureCitation box="[484,574,1378,1406]" captionStart="Fig" captionStartId="73.[192,239,1773,1801]" captionTargetBox="[192,1344,192,1728]" captionTargetId="figure@73.[192,1344,192,1728]" captionText="Fig. S2. Google Earth satellite map of Peleng and Banggai (top) and the western half of Taliabu (bottom). Peleng and Banggai are the two largest islands of the “Banggai Archipelago”. Our main collection site, an unnamed locality “above Kokolomboi [village]”, is mapped with a yellow pin. Areas above 800m elevation are circled with a red line, indicating the presence of montane vegetation. The island of Taliabu is the largest member of the “Sula Archipelago”. Our main collection locality “Waiyo dinahana” is mapped with a yellow pin. Areas above 1,050 m elevation are circled with a red line, indicating the presence of a peculiar montane stunted forest vegetation. Within the montane zone above 1,050m, areas degraded by logging activity as reflected on Google Earth satellite imagery are mapped in turquoise and areas burnt down by extensive fires in the 1980s are mapped in dark-green. Given the outdated status of some of the satellite imagery, present-day forest degradation is likely more extensive than shown." figureDoi="http://doi.org/10.5281/zenodo.3608762" httpUri="https://zenodo.org/record/3608762/files/figure.png" pageId="17" pageNumber="18">fig. S2</figureCitation>
). However, satellite imagery predominantly from 2004 indicates that a total of ~1,023 ha (~6.1%) of this area had been affected by logging operations that have degraded the forest and removed its undercover, presumably rendering the affected areas unsuitable, or nearly so, for the grasshopper-warbler. In the one-and-a-half decades since the satellite images were obtained, extensive additional logging is likely to have occurred. Another 940 ha (~5.6%) of the area above 1,050m have succumbed to a series of catastrophic forest fires in 1982 and 1983, and now constitute open grassland habitat unsuitable for this warbler (
<figureCitation box="[546,637,1894,1922]" captionStart="Fig" captionStartId="73.[192,239,1773,1801]" captionTargetBox="[192,1344,192,1728]" captionTargetId="figure@73.[192,1344,192,1728]" captionText="Fig. S2. Google Earth satellite map of Peleng and Banggai (top) and the western half of Taliabu (bottom). Peleng and Banggai are the two largest islands of the “Banggai Archipelago”. Our main collection site, an unnamed locality “above Kokolomboi [village]”, is mapped with a yellow pin. Areas above 800m elevation are circled with a red line, indicating the presence of montane vegetation. The island of Taliabu is the largest member of the “Sula Archipelago”. Our main collection locality “Waiyo dinahana” is mapped with a yellow pin. Areas above 1,050 m elevation are circled with a red line, indicating the presence of a peculiar montane stunted forest vegetation. Within the montane zone above 1,050m, areas degraded by logging activity as reflected on Google Earth satellite imagery are mapped in turquoise and areas burnt down by extensive fires in the 1980s are mapped in dark-green. Given the outdated status of some of the satellite imagery, present-day forest degradation is likely more extensive than shown." figureDoi="http://doi.org/10.5281/zenodo.3608762" httpUri="https://zenodo.org/record/3608762/files/figure.png" pageId="17" pageNumber="18">fig. S2</figureCitation>
). Our surveys showed that
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[985,1127,1895,1923]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[985,1127,1895,1923]" italics="true" pageId="17" pageNumber="18">L. portenta</emphasis>
</taxonomicName>
does not occur in the extensive recently generated grasslands in the highlands of Taliabu (
<bibRefCitation author="F. E. Rheindt" box="[1116,1148,1967,1995]" journalOrPublisher="Bull. Br. Ornithol. Club" pageId="17" pageNumber="18" pagination="33 - 51" part="130" refId="ref29294" refString="48. F. E. Rheindt, New biogeographic records for the avifauna of Taliabu (Sula Islands, Indonesia), with the preliminary documentation of two previously undiscovered taxa. Bull. Br. Ornithol. Club 130, 33 - 51 (2010)." title="New biogeographic records for the avifauna of Taliabu (Sula Islands, Indonesia), with the preliminary documentation of two previously undiscovered taxa" type="journal article" year="2010">
<emphasis box="[1116,1148,1967,1995]" italics="true" pageId="17" pageNumber="18">48</emphasis>
</bibRefCitation>
). This leaves
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" lastPageId="18" lastPageNumber="19" order="Passeriformes" pageId="17" pageNumber="78" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[1328,1353,1969,1997]" italics="true" pageId="17" pageNumber="18">L.</emphasis>
<emphasis box="[192,300,203,231]" italics="true" pageId="18" pageNumber="19">portenta</emphasis>
</taxonomicName>
with less than 15,000 ha of suitable habitat, an area likely to further diminish in size with ongoing habitat destruction and the impacts of global warming leading to an upward elevational shift of habitat zones.
</paragraph>
</subSubSection>
<subSubSection lastPageId="20" lastPageNumber="21" pageId="18" pageNumber="19" type="discussion">
<paragraph blockId="18.[192,1383,201,2002]" box="[192,476,500,528]" pageId="18" pageNumber="19">
<heading bold="true" box="[192,476,500,528]" fontSize="12" level="7" pageId="18" pageNumber="19" reason="2">
<emphasis bold="true" box="[192,476,500,528]" pageId="18" pageNumber="19">Taxonomic rationale</emphasis>
</heading>
</paragraph>
<paragraph blockId="18.[192,1383,201,2002]" pageId="18" pageNumber="19">
We include this new taxon within
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[631,764,574,602]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">
<emphasis box="[631,764,574,602]" italics="true" pageId="18" pageNumber="19">Locustella</emphasis>
</taxonomicName>
in the family
<taxonomicName box="[944,1116,574,602]" class="Aves" family="Locustellidae" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="family">Locustellidae</taxonomicName>
following the phylogenetic demonstration that the genus
<taxonomicName box="[742,900,648,676]" class="Aves" family="Locustellidae" genus="Bradypterus" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">
<emphasis box="[742,900,648,676]" italics="true" pageId="18" pageNumber="19">Bradypterus</emphasis>
</taxonomicName>
(previously used for this Southeast Asian clade of warblers) is restricted to Africa (
<emphasis box="[803,883,721,751]" italics="true" pageId="18" pageNumber="19">8 1, 8 2</emphasis>
). Its inclusion within
<taxonomicName box="[1165,1337,721,749]" class="Aves" family="Locustellidae" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="family">Locustellidae</taxonomicName>
is the reason why we follow the recent trend [e.g. (
<bibRefCitation author="J. A. Eaton &amp; S. van Balen &amp; N. W. Brickle &amp; F. E. Rheindt" box="[813,845,796,824]" journalOrPublisher="Lynx Edicions" pageId="18" pageNumber="19" refId="ref29850" refString="57. J. A. Eaton, S. van Balen, N. W. Brickle, F. E. Rheindt, Birds of the Indonesian Archipelago: Greater Sundas and Wallacea. (Lynx Edicions, 2016)." title="Birds of the Indonesian Archipelago: Greater Sundas and Wallacea" type="book" year="2016">
<emphasis box="[813,845,796,824]" italics="true" pageId="18" pageNumber="19">57</emphasis>
</bibRefCitation>
)] of using the English moniker grasshopper-warbler rather than the term bush-warbler, which is more appropriately reserved for members of a radiation in the family
<taxonomicName box="[829,949,942,970]" class="Aves" family="Cettiidae" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="family">Cettiidae</taxonomicName>
.
</paragraph>
<paragraph blockId="18.[192,1383,201,2002]" pageId="18" pageNumber="19">
The genus
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[428,561,1016,1044]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">
<emphasis box="[428,561,1016,1044]" italics="true" pageId="18" pageNumber="19">Locustella</emphasis>
</taxonomicName>
is renowned for its cryptic and conservative coloration that has confounded plumage-based classifications for decades and led to delays in the recognition of species-level taxa until bioacoustic and molecular methods have been applied (
<emphasis box="[1203,1283,1163,1193]" italics="true" pageId="18" pageNumber="19">8 3, 8 4</emphasis>
). In Wallacea, members of a
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[509,642,1236,1264]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="genus">
<emphasis box="[509,642,1236,1264]" italics="true" pageId="18" pageNumber="19">Locustella</emphasis>
</taxonomicName>
radiation from Sulawesi and the Moluccas were previously united into a single polytypic species
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[817,965,1312,1340]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[817,965,1312,1340]" italics="true" pageId="18" pageNumber="19">L. castanea</emphasis>
</taxonomicName>
(
<emphasis box="[983,1063,1310,1340]" italics="true" pageId="18" pageNumber="19">6 1, 7 5</emphasis>
), but have recently been divided into three constituent island species,
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[767,915,1385,1413]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[767,915,1385,1413]" italics="true" pageId="18" pageNumber="19">L. castanea</emphasis>
</taxonomicName>
from Sulawesi,
<taxonomicName box="[1124,1277,1384,1413]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="musculus">
<emphasis box="[1124,1277,1384,1413]" italics="true" pageId="18" pageNumber="19">L. musculus</emphasis>
</taxonomicName>
from Seram, and
<taxonomicName box="[344,513,1457,1487]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[344,513,1457,1487]" italics="true" pageId="18" pageNumber="19">L. disturbans</emphasis>
</taxonomicName>
from Buru, on the basis of their strikingly different songs (
<bibRefCitation author="J. A. Eaton &amp; S. van Balen &amp; N. W. Brickle &amp; F. E. Rheindt" box="[1272,1304,1459,1487]" journalOrPublisher="Lynx Edicions" pageId="18" pageNumber="19" refId="ref29850" refString="57. J. A. Eaton, S. van Balen, N. W. Brickle, F. E. Rheindt, Birds of the Indonesian Archipelago: Greater Sundas and Wallacea. (Lynx Edicions, 2016)." title="Birds of the Indonesian Archipelago: Greater Sundas and Wallacea" type="book" year="2016">
<emphasis box="[1272,1304,1459,1487]" italics="true" pageId="18" pageNumber="19">57</emphasis>
</bibRefCitation>
). Our new
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[253,395,1533,1561]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[253,395,1533,1561]" italics="true" pageId="18" pageNumber="19">L. portenta</emphasis>
</taxonomicName>
is geographically surrounded by constituents of this radiation and is a member of it both on the basis of morphological characters (see Diagnosis) and bioacoustic ones.
</paragraph>
<paragraph blockId="18.[192,1383,201,2002]" lastBlockId="19.[192,1389,201,1997]" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19">
Although plumage distinctions are not as important in the genus,
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[1124,1266,1753,1781]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[1124,1266,1753,1781]" italics="true" pageId="18" pageNumber="19">L. portenta</emphasis>
</taxonomicName>
is characterized by comparatively unique, dark-brown upperparts that stand in contrast to the generally paler and warmer-brown upperparts of neighboring species. However, we mainly attribute species status to
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[521,663,1974,2002]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[521,663,1974,2002]" italics="true" pageId="18" pageNumber="19">L. portenta</emphasis>
</taxonomicName>
on the basis of its unique song and genetic differentiation. We here provide an excerpt of ongoing analyses as relating to the bioacoustic and mitogenomic differentiation of
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[649,791,276,304]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[649,791,276,304]" italics="true" pageId="19" pageNumber="20">L. portenta</emphasis>
</taxonomicName>
from neighboring species.
</paragraph>
<paragraph blockId="18.[778,808,2018,2044]" box="[778,808,2018,2044]" pageId="18" pageNumber="19">
<pageNumber ambiguity="1" box="[778,808,2018,2044]" fuzzyness="0" pageId="18" pageNumber="19" score="3826.0" value="19">
<pageTitle box="[778,808,2018,2044]" pageId="18" pageNumber="19">19</pageTitle>
</pageNumber>
</paragraph>
<paragraph blockId="19.[192,1389,201,1997]" pageId="19" pageNumber="20">
<emphasis bold="true" box="[288,569,348,376]" pageId="19" pageNumber="20">Bioacoustic evidence</emphasis>
: A total of 41 suitable, high-quality, homologous
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[1211,1344,348,376]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="genus">
<emphasis box="[1211,1344,348,376]" italics="true" pageId="19" pageNumber="20">Locustella</emphasis>
</taxonomicName>
song recordings from the field were gathered for analysis from various sources, encompassing all three previous species of the
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[794,941,497,525]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[794,941,497,525]" italics="true" pageId="19" pageNumber="20">L. castanea</emphasis>
</taxonomicName>
radiation plus the new species. These recordings were measured for 11 suitable bioacoustic parameters, and resultant measurements were subjected to principal component analysis (PCA) and a conservative vocal diagnosability criterion (
<bibRefCitation author="M. L. Isler &amp; P. R. Isler &amp; B. M. Whitney" box="[586,618,716,744]" journalOrPublisher="Auk" pageId="19" pageNumber="20" pagination="577 - 590" part="115" refId="ref31338" refString="85. M. L. Isler, P. R. Isler, B. M. Whitney, Use of vocalizations to establish species limits in antbirds (Passeriformes: Thamnophilidae). Auk 115, 577 - 590 (1998). doi: 10.2307 / 4089407" title="Use of vocalizations to establish species limits in antbirds (Passeriformes: Thamnophilidae)" type="journal article" year="1998">
<emphasis box="[586,618,716,744]" italics="true" pageId="19" pageNumber="20">85</emphasis>
</bibRefCitation>
), henceforth referred to as the Isler Criterion, to assess song diagnosability among taxa. Further details regarding all analyzed recordings, such as GPS locations, dates, times, and names of recordists, are listed in table S3, while more details on bioacoustic methodology are provided below under separate Methods paragraphs.
</paragraph>
<paragraph blockId="19.[192,1389,201,1997]" pageId="19" pageNumber="20">
Based on the Isler Criterion, eight out of eleven song parameters measured were diagnosable between the new Taliabu Grasshopper-Warbler
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[967,1109,1086,1114]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[967,1109,1086,1114]" italics="true" pageId="19" pageNumber="20">L. portenta</emphasis>
</taxonomicName>
and two other taxa:
<taxonomicName box="[192,345,1158,1187]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="musculus">
<emphasis box="[192,345,1158,1187]" italics="true" pageId="19" pageNumber="20">L. musculus</emphasis>
</taxonomicName>
from Seram and
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[567,715,1159,1187]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[567,715,1159,1187]" italics="true" pageId="19" pageNumber="20">L. castanea</emphasis>
</taxonomicName>
from Sulawesi (table S4). Four out of eleven parameters were Isler-diagnosable between the new Taliabu Grasshopper-Warbler and
<taxonomicName class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[1307,1333,1233,1261]" italics="true" pageId="19" pageNumber="20">L.</emphasis>
<emphasis box="[192,327,1305,1333]" italics="true" pageId="19" pageNumber="20">disturbans</emphasis>
</taxonomicName>
from Buru (table S4). The yardstick approach (
<bibRefCitation author="E. Mayr &amp; P. Ashlock" box="[959,991,1305,1333]" journalOrPublisher="McGraw-Hill, ed" pageId="19" pageNumber="20" refId="ref30411" refString="70. E. Mayr, P. Ashlock, Principles of Systematic Zoology. (McGraw-Hill, ed. 2, 1991)." title="Principles of Systematic Zoology" type="book" year="1991">
<emphasis box="[959,991,1305,1333]" italics="true" pageId="19" pageNumber="20">70</emphasis>
</bibRefCitation>
) can be adopted to ascertain if the level of vocal divergence between these archipelagic taxa is comparable in magnitude to divergences across unequivocal species-level divisions. When comparing Taliabu Grasshopper-Warbler song against that of the Friendly Grasshopper-Warbler
<taxonomicName baseAuthorityName="Sharpe" baseAuthorityYear="1888" box="[1184,1329,1527,1555]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="accentor">
<emphasis box="[1184,1329,1527,1555]" italics="true" pageId="19" pageNumber="20">L. accentor</emphasis>
</taxonomicName>
, a geographically distant species from montane Borneo that has never been included in the
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[1327,1352,1601,1629]" italics="true" pageId="19" pageNumber="20">L.</emphasis>
<emphasis box="[192,306,1675,1703]" italics="true" pageId="19" pageNumber="20">castanea</emphasis>
</taxonomicName>
radiation, only two parameters emerged as Isler-diagnosable (table S4). Using this result as a yardstick for what constitutes species level vocal differentiation, the magnitude of vocal divergence between Taliabu Grasshopper-Warbler and
<taxonomicName box="[977,1130,1820,1850]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="musculus">
<emphasis box="[977,1130,1820,1850]" italics="true" pageId="19" pageNumber="20">L. musculus</emphasis>
</taxonomicName>
on Seram,
<taxonomicName class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[1276,1302,1822,1850]" italics="true" pageId="19" pageNumber="20">L.</emphasis>
<emphasis box="[192,327,1894,1922]" italics="true" pageId="19" pageNumber="20">disturbans</emphasis>
</taxonomicName>
on Buru, and
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[509,657,1895,1923]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[509,657,1895,1923]" italics="true" pageId="19" pageNumber="20">L. castanea</emphasis>
</taxonomicName>
on Sulawesi is extremely high, indicating they all have diagnosably distinct vocalizations that are best reflected in species rank.
</paragraph>
<paragraph blockId="20.[192,1389,201,1850]" pageId="20" pageNumber="21">
Principal component analysis of the vocal measurements underscores the vocal distinctness of Taliabu Grasshopper-Warbler
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[775,917,276,304]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[775,917,276,304]" italics="true" pageId="20" pageNumber="21">L. portenta</emphasis>
</taxonomicName>
from the other three Wallacean species (
<figureCitation box="[302,392,348,376]" captionStart="Fig" captionStartId="76.[192,239,959,987]" captionTargetBox="[204,1322,202,905]" captionTargetId="figure@76.[192,1344,192,914]" captionTargetPageId="76" captionText="Fig. S4. Principal component analysis of 11 bioacoustic parameters across four species of Locustella grasshopper-warblers. Ellipses represent 95% confidence intervals of the principal component (PC) scores for each taxon. The percentage of total variance of each PC is given in parentheses." figureDoi="http://doi.org/10.5281/zenodo.3608766" httpUri="https://zenodo.org/record/3608766/files/figure.png" pageId="20" pageNumber="21">fig. S4</figureCitation>
). A cumulative proportion of variance of 73.3% is accounted for in the first two principal components, suggesting a strong vocal differentiation between Taliabu and the remaining taxa (
<figureCitation box="[400,491,495,523]" captionStart="Fig" captionStartId="76.[192,239,959,987]" captionTargetBox="[204,1322,202,905]" captionTargetId="figure@76.[192,1344,192,914]" captionTargetPageId="76" captionText="Fig. S4. Principal component analysis of 11 bioacoustic parameters across four species of Locustella grasshopper-warblers. Ellipses represent 95% confidence intervals of the principal component (PC) scores for each taxon. The percentage of total variance of each PC is given in parentheses." figureDoi="http://doi.org/10.5281/zenodo.3608766" httpUri="https://zenodo.org/record/3608766/files/figure.png" pageId="20" pageNumber="21">fig. S4</figureCitation>
).
</paragraph>
<paragraph blockId="20.[192,1389,201,1850]" pageId="20" pageNumber="21">
<emphasis bold="true" box="[288,592,569,597]" pageId="20" pageNumber="21">Mitogenomic evidence</emphasis>
: Using whole-genome sequencing and laboratory approaches for the use of ancient DNA, we created an alignment of whole-mitochondrial sequences spanning ~15,000 base pairs (bp) across 11
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[753,886,716,744]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="genus">
<emphasis box="[753,886,716,744]" italics="true" pageId="20" pageNumber="21">Locustella</emphasis>
</taxonomicName>
individuals belonging to all four constituent island forms of the
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[589,737,791,819]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[589,737,791,819]" italics="true" pageId="20" pageNumber="21">L. castanea</emphasis>
</taxonomicName>
radiation, and used three different phylogenetic tree-building methodologies to investigate relationships and levels of divergence. More details on the methodology are provided below under separate Methods paragraphs.
</paragraph>
<paragraph blockId="20.[192,1389,201,1850]" pageId="20" pageNumber="21">
All three phylogenetic trees produced indicated identical topologies with high bootstrap support (
<figureCitation box="[433,521,1084,1112]" captionStart="Fig" captionStartId="77.[192,239,1062,1090]" captionTargetBox="[229,1319,205,1012]" captionTargetId="figure@77.[192,1344,192,1015]" captionTargetPageId="77" captionText="Fig. S5. Mitogenomic maximum likelihood (ML) tree phylogram of relationships among the four constituent members of the Locustella castanea radiation. Identical maximum bootstrap support (100), as shown for clades surrounding L. portenta, was produced using three different tree-building methodologies: ML, maximum parsimony, and neighbor-joining. Color-coded uncorrected pairwise sequence divergences are given between L. portenta and other species. The bird illustrations are modified from Eaton et al. (57). Voucher numbers for the American Museum of Natural History (AMNH) in New York are given for toepad- derived DNA samples." figureDoi="http://doi.org/10.5281/zenodo.3608768" httpUri="https://zenodo.org/record/3608768/files/figure.png" pageId="20" pageNumber="21">fig. S5</figureCitation>
), suggesting that the new
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[855,997,1086,1114]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[855,997,1086,1114]" italics="true" pageId="20" pageNumber="21">L. portenta</emphasis>
</taxonomicName>
is an independent lineage most closely related to
<taxonomicName box="[489,658,1158,1187]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[489,658,1158,1187]" italics="true" pageId="20" pageNumber="21">L. disturbans</emphasis>
</taxonomicName>
from Buru (mitogenomic divergence ~2%), and much more distantly related to
<taxonomicName box="[512,665,1231,1261]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="musculus">
<emphasis box="[512,665,1231,1261]" italics="true" pageId="20" pageNumber="21">L. musculus</emphasis>
</taxonomicName>
from Seram (divergence ~4.85%) and
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" box="[1181,1328,1233,1261]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[1181,1328,1233,1261]" italics="true" pageId="20" pageNumber="21">L. castanea</emphasis>
</taxonomicName>
from Sulawesi (divergence ~7.78%). Mitogenomic divergences roughly correlate with those from the COI barcoding gene, although they do not have to be equal in magnitude.
</paragraph>
<paragraph blockId="20.[192,1389,201,1850]" pageId="20" pageNumber="21">
Integrating the vocal and mitogenomic evidence, it is clear that bioacoustic differences between
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[457,599,1527,1555]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[457,599,1527,1555]" italics="true" pageId="20" pageNumber="21">L. portenta</emphasis>
</taxonomicName>
and
<taxonomicName box="[661,830,1526,1555]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[661,830,1526,1555]" italics="true" pageId="20" pageNumber="21">L. disturbans</emphasis>
</taxonomicName>
, the two species which emerged most closely related on the mitogenomic tree (
<figureCitation box="[715,804,1599,1627]" captionStart="Fig" captionStartId="77.[192,239,1062,1090]" captionTargetBox="[229,1319,205,1012]" captionTargetId="figure@77.[192,1344,192,1015]" captionTargetPageId="77" captionText="Fig. S5. Mitogenomic maximum likelihood (ML) tree phylogram of relationships among the four constituent members of the Locustella castanea radiation. Identical maximum bootstrap support (100), as shown for clades surrounding L. portenta, was produced using three different tree-building methodologies: ML, maximum parsimony, and neighbor-joining. Color-coded uncorrected pairwise sequence divergences are given between L. portenta and other species. The bird illustrations are modified from Eaton et al. (57). Voucher numbers for the American Museum of Natural History (AMNH) in New York are given for toepad- derived DNA samples." figureDoi="http://doi.org/10.5281/zenodo.3608768" httpUri="https://zenodo.org/record/3608768/files/figure.png" pageId="20" pageNumber="21">fig. S5</figureCitation>
), far exceed those between
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[1159,1301,1601,1629]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[1159,1301,1601,1629]" italics="true" pageId="20" pageNumber="21">L. portenta</emphasis>
</taxonomicName>
and a geographically distant species (
<taxonomicName baseAuthorityName="Sharpe" baseAuthorityYear="1888" box="[595,740,1675,1703]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="accentor">
<emphasis box="[595,740,1675,1703]" italics="true" pageId="20" pageNumber="21">L. accentor</emphasis>
</taxonomicName>
) that has never been closely associated with the
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[1363,1389,1675,1703]" italics="true" pageId="20" pageNumber="21">L.</emphasis>
<emphasis box="[192,306,1748,1776]" italics="true" pageId="20" pageNumber="21">castanea</emphasis>
</taxonomicName>
radiation. Our combined evidence therefore supports the treatment of
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[1208,1349,1748,1776]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[1208,1349,1748,1776]" italics="true" pageId="20" pageNumber="21">L. portenta</emphasis>
</taxonomicName>
as a well-diverged small-insular endemic species.
</paragraph>
</subSubSection>
<subSubSection lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22" type="biology_ecology">
<paragraph blockId="21.[192,1393,201,1997]" box="[192,372,201,229]" pageId="21" pageNumber="22">
<heading bold="true" box="[192,372,201,229]" fontSize="12" level="7" pageId="21" pageNumber="22" reason="2">
<emphasis bold="true" box="[192,372,201,229]" pageId="21" pageNumber="22">Methodology</emphasis>
</heading>
</paragraph>
<paragraph blockId="21.[192,1393,201,1997]" pageId="21" pageNumber="22">
<emphasis bold="true" box="[192,492,274,302]" pageId="21" pageNumber="22">Bioacoustic sampling:</emphasis>
We obtained 41 homologous recordings, which were either recorded by us in the field, or privately sourced from avid field ornithologists (including Robert O. Hutchinson and Ross Gallardy), or collected from publicly accessible online databases, namely the Avian Vocalization Center (AVoCet at http://avocet. integrativebiology.natsci.msu.edu), the Xeno-Canto Sound Library (www.xeno-canto.org), the Macaulay Sound Library (www.macaulaylibrary.org), and the Internet Bird Collection (www.hbw.com/ibc).
</paragraph>
<paragraph blockId="21.[192,1393,201,1997]" pageId="21" pageNumber="22">Although slight variation in recording quality exists in the equipment used, inspection and comparison of spectrograms from different devices showed that this variation is comparable to that amongst recordings made by one sole recordist using the same device. In addition, although there is some variability in background noise, spectrogram inspection suggested that measurements of vocalizations would not be affected. Measurements of the recordings were performed using the program Raven Pro Version 1.5 (Bioacoustics Research Program, Cornell Laboratory of Ornithology, Ithaca, NY, USA) on its default settings. The following eleven parameters were chosen for bioacoustic analysis, with each parameter averaged over all the measurements across one recording:</paragraph>
<paragraph blockId="21.[192,1393,201,1997]" pageId="21" pageNumber="22">1. Total number of elements per motif (with an element being a syllable represented by a trace on the spectrogram, and a motif being a collection of elements always repeated together)</paragraph>
<paragraph blockId="21.[192,1393,201,1997]" pageId="21" pageNumber="22">2. Number of introductory elements per motif (with introductory elements being brief, truncated, and usually fainter elements preceding the main body of the song)</paragraph>
<paragraph blockId="21.[192,1393,201,1997]" pageId="21" pageNumber="22">
3. Number of main elements per motif (with main elements taking up the bulk of the song, in
<taxonomicName authorityName="Kaup" authorityYear="1829" box="[397,530,1894,1922]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="21" pageNumber="22" phylum="Chordata" rank="genus">
<emphasis box="[397,530,1894,1922]" italics="true" pageId="21" pageNumber="22">Locustella</emphasis>
</taxonomicName>
typically in the form of shrill, insect-like trills)
</paragraph>
<paragraph blockId="21.[192,1393,201,1997]" box="[240,604,1967,1997]" pageId="21" pageNumber="22">4. Total duration of a motif</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[240,793,201,229]" pageId="22" pageNumber="23">5. Total duration of introductory elements</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[240,699,274,302]" pageId="22" pageNumber="23">6. Total duration of main elements</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[240,1044,348,378]" pageId="22" pageNumber="23">7. Duration of break between introductory and main elements</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[240,686,422,451]" pageId="22" pageNumber="23">8. Minimum frequency of a motif</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[240,691,495,525]" pageId="22" pageNumber="23">9. Maximum frequency of a motif</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[240,997,569,597]" pageId="22" pageNumber="23">10. Bandwidth (i.e., maximum minus minimum frequency)</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[240,1365,642,672]" pageId="22" pageNumber="23">11. Peak frequency of a motif (i.e., the frequency at which the amplitude of the sound is</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" box="[288,392,716,744]" pageId="22" pageNumber="23">highest)</paragraph>
<paragraph blockId="22.[192,1382,201,2009]" pageId="22" pageNumber="23">
<emphasis bold="true" box="[288,572,790,818]" pageId="22" pageNumber="23">Bioacoustic analysis:</emphasis>
The vocal diagnosability criterion outlined by Isler et al. (
<emphasis box="[1319,1351,790,819]" italics="true" pageId="22" pageNumber="23">8 5</emphasis>
), henceforth referred to as the Isler Criterion, was used to assess song diagnosability among taxa. Besides other oscine passerines (
<emphasis box="[683,763,937,967]" italics="true" pageId="22" pageNumber="23">73, 8 6</emphasis>
), this criterion has been employed as a bioacoustic species delimitation method across Asian birds such as owls (
<emphasis box="[1136,1216,1010,1040]" italics="true" pageId="22" pageNumber="23">
<bibRefCitation author="C. Y. Gwee &amp; L. Christidis &amp; J. A. Eaton &amp; J. A. Norman &amp; C. R. Trainor &amp; P. Verbelen &amp; F. E. Rheindt" box="[1136,1168,1010,1038]" journalOrPublisher="Mol. Phylogenet. Evol." pageId="22" pageNumber="23" pagination="246 - 258" part="109" refId="ref29893" refString="58. C. Y. Gwee, L. Christidis, J. A. Eaton, J. A. Norman, C. R. Trainor, P. Verbelen, F. E. Rheindt, Bioacoustic and multi-locus DNA data of Ninox owls support high incidence of extinction and recolonisation on small, low-lying islands across Wallacea. Mol. Phylogenet. Evol. 109, 246 - 258 (2017). doi: 10.1016 / j. ympev. 2016.12.024 Medline" title="Bioacoustic and multi-locus DNA data of Ninox owls support high incidence of extinction and recolonisation on small, low-lying islands across Wallacea" type="journal article" year="2017">58</bibRefCitation>
,
<bibRefCitation author="C. Y. Gwee &amp; J. A. Eaton &amp; E. Y. X. Ng &amp; F. E. Rheindt" box="[1184,1216,1010,1038]" journalOrPublisher="Avian Res." pageId="22" pageNumber="23" pagination="36" part="10" refId="ref31438" refString="87. C. Y. Gwee, J. A. Eaton, E. Y. X. Ng, F. E. Rheindt, Species delimitation within the Glaucidium brodiei owlet complex using bioacoustic tools. Avian Res. 10, 36 (2019). doi: 10.1186 / s 40657 - 019 - 0175 - 4" title="Species delimitation within the Glaucidium brodiei owlet complex using bioacoustic tools" type="journal article" year="2019">87</bibRefCitation>
</emphasis>
), pigeons (
<emphasis box="[203,331,1084,1112]" italics="true" pageId="22" pageNumber="23">
<bibRefCitation author="F. E. Rheindt &amp; J. A. Eaton &amp; F. Verbelen" box="[203,238,1084,1112]" journalOrPublisher="Ornithol." pageId="22" pageNumber="23" pagination="429 - 440" part="123" refId="ref29408" refString="50. F. E. Rheindt, J. A. Eaton, F. Verbelen, Vocal trait evolution in a geographic leapfrog pattern: Speciation in the maroon-chinned fruit dove (Ptilinopus subgularis) complex from Wallacea. Wilson J. Ornithol. 123, 429 - 440 (2011). doi: 10.1676 / 10 - 148.1" title="Vocal trait evolution in a geographic leapfrog pattern: Speciation in the maroon-chinned fruit dove (Ptilinopus subgularis) complex from Wallacea. Wilson J" type="journal article" year="2011">50</bibRefCitation>
,
<bibRefCitation author="N. S. R. Ng &amp; F. E. Rheindt" box="[251,286,1084,1112]" journalOrPublisher="Avian Res." pageId="22" pageNumber="23" pagination="2" part="7" refId="ref29509" refString="52. N. S. R. Ng, F. E. Rheindt, Species delimitation in the white-faced cuckoo-dove (Turacoena manadensis) based on bioacoustic data. Avian Res. 7, 2 (2016). doi: 10.1186 / s 40657 - 015 - 0036 - 8" title="Species delimitation in the white-faced cuckoo-dove (Turacoena manadensis) based on bioacoustic data" type="journal article" year="2016">52</bibRefCitation>
,
<bibRefCitation author="E. Y. X. Ng &amp; J. A. Eaton &amp; P. Verbelen &amp; R. O. Hutchinson &amp; F. E. Rheindt" box="[299,331,1084,1112]" journalOrPublisher="Biol. J. Linn. Soc." pageId="22" pageNumber="23" pagination="786 - 812" part="118" refId="ref31500" refString="88. E. Y. X. Ng, J. A. Eaton, P. Verbelen, R. O. Hutchinson, F. E. Rheindt, Using bioacoustic data to test species limits in an Indo-Pacific island radiation of Macropygia cuckoo doves. Biol. J. Linn. Soc. 118, 786 - 812 (2016). doi: 10.1111 / bij. 12768" title="Using bioacoustic data to test species limits in an Indo-Pacific island radiation of Macropygia cuckoo doves" type="journal article" year="2016">88</bibRefCitation>
</emphasis>
), and nightjars (
<bibRefCitation author="G. Sangster &amp; F. Rozendaal" box="[542,574,1084,1112]" journalOrPublisher="Zoologische Verhandelingen" pageId="22" pageNumber="23" pagination="7 - 45" part="350" refId="ref31573" refString="89. G. Sangster, F. Rozendaal, Territorial songs and species-level taxonomy of nightjars of the Caprimulgus macrurus complex, with the description of a new species. Zoologische Verhandelingen 350, 7 - 45 (2004)." title="Territorial songs and species-level taxonomy of nightjars of the Caprimulgus macrurus complex, with the description of a new species" type="journal article" year="2004">
<emphasis box="[542,574,1084,1112]" italics="true" pageId="22" pageNumber="23">89</emphasis>
</bibRefCitation>
). The Isler criterion has two parts which must be fulfilled in order to infer diagnosability. Firstly, the range of measurements of a parameter between two taxa must not exhibit overlap, and secondly, the data have to satisfy the following equation:
<emphasis box="[192,222,1310,1338]" italics="true" pageId="22" pageNumber="23">xa</emphasis>
+
<emphasis box="[248,496,1307,1338]" italics="true" pageId="22" pageNumber="23">taSDa ≤xbtbSDb,</emphasis>
where (
<emphasis box="[601,615,1307,1335]" italics="true" pageId="22" pageNumber="23">x</emphasis>
) refers to the respective means of measurements in taxon a and b, and
<emphasis box="[332,350,1391,1419]" italics="true" pageId="22" pageNumber="23">ti</emphasis>
refers to the t-score at the 95th percentile of the t distribution for
<emphasis box="[1191,1233,1386,1416]" italics="true" pageId="22" pageNumber="23">n-1</emphasis>
degrees of freedom. This is a stricter statistical test than the t-test or Mann-Whitney U test (
<emphasis box="[1224,1256,1464,1494]" italics="true" pageId="22" pageNumber="23">8 5</emphasis>
) as it employs the standard deviation of the sample points instead of the taxon mean, allowing for the diagnosis of distinctions between taxa under more rigorous conditions. Successful Isler diagnosability is defined here as the presence of at least one vocally diagnostic parameter between taxa.
</paragraph>
</subSubSection>
<subSubSection lastPageId="25" lastPageNumber="26" pageId="22" pageNumber="23" type="discussion">
<paragraph blockId="22.[192,1382,201,2009]" lastBlockId="23.[192,1386,201,1997]" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23">
<emphasis bold="true" box="[288,638,1832,1860]" pageId="22" pageNumber="23">DNA extraction methods:</emphasis>
Fresh tissue of the holotype of
<taxonomicName authorityName="Rheindt &amp; Prawiradilaga &amp; Ashar &amp; Lee &amp; Wu &amp; Ng" authorityYear="2020" box="[1043,1184,1834,1862]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="portenta">
<emphasis box="[1043,1184,1834,1862]" italics="true" pageId="22" pageNumber="23">L. portenta</emphasis>
</taxonomicName>
was DNA- sequenced along with the following number of toepad scrapings of historical museum specimens: three of the Buru Grasshopper-Warbler
<taxonomicName box="[853,1022,1979,2009]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="22" pageNumber="23" phylum="Chordata" rank="species" species="disturbans">
<emphasis box="[853,1022,1979,2009]" italics="true" pageId="22" pageNumber="23">L. disturbans</emphasis>
</taxonomicName>
, three of the Seram Grasshopper-Warbler
<taxonomicName box="[477,630,201,231]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="musculus">
<emphasis box="[477,630,201,231]" italics="true" pageId="23" pageNumber="24">L. musculus</emphasis>
</taxonomicName>
, and four of the Sulawesi Grasshopper-Warbler
<taxonomicName baseAuthorityName="Büttikofer" baseAuthorityYear="1893" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="23" pageNumber="24" phylum="Chordata" rank="species" species="castanea">
<emphasis box="[1252,1278,203,231]" italics="true" pageId="23" pageNumber="24">L.</emphasis>
<emphasis box="[192,306,276,304]" italics="true" pageId="23" pageNumber="24">castanea</emphasis>
</taxonomicName>
. All museum samples had been kindly contributed by the American Museum of Natural History (for voucher numbers, see
<figureCitation box="[742,827,348,376]" captionStart="Fig" captionStartId="77.[192,239,1062,1090]" captionTargetBox="[229,1319,205,1012]" captionTargetId="figure@77.[192,1344,192,1015]" captionTargetPageId="77" captionText="Fig. S5. Mitogenomic maximum likelihood (ML) tree phylogram of relationships among the four constituent members of the Locustella castanea radiation. Identical maximum bootstrap support (100), as shown for clades surrounding L. portenta, was produced using three different tree-building methodologies: ML, maximum parsimony, and neighbor-joining. Color-coded uncorrected pairwise sequence divergences are given between L. portenta and other species. The bird illustrations are modified from Eaton et al. (57). Voucher numbers for the American Museum of Natural History (AMNH) in New York are given for toepad- derived DNA samples." figureDoi="http://doi.org/10.5281/zenodo.3608768" httpUri="https://zenodo.org/record/3608768/files/figure.png" pageId="23" pageNumber="24">fig. S5</figureCitation>
and table S5). DNA extraction was carried out with the Qiagen DNeasy Blood and Tissue Kit (Qiagen, Germany) following the manufacturers instructions. DNA extraction of the historic museum specimens toepad scrapings was conducted by trained personnel (CYG) in a room dedicated to ancient DNA extraction to preclude DNA contamination. The toepads were rinsed with water four times before extraction to ensure preservatives were removed, and DTT was added to facilitate digestion of keratin present in toepads. The concentrations of the DNA extracts were quantified using a high-sensitivity Qubit (Life Technologies, USA) assay.
</paragraph>
<paragraph blockId="23.[192,1386,201,1997]" lastBlockId="24.[192,1388,201,1997]" lastPageId="24" lastPageNumber="25" pageId="23" pageNumber="24">
<emphasis bold="true" box="[288,651,937,965]" pageId="23" pageNumber="24">DNA sequencing methods:</emphasis>
The holotypes DNA extract was sheared into smaller fragments of ~250 base pairs (bp) using 13 cycles of 30 seconds on and 30 seconds off on a Bioruptor Plus (Diagenode, USA). Whole-genome library preparation for the holotype was carried out using a NEBNext Ultra II DNA Library Prep Kit (New England Biolabs, USA) for Illumina according to the manufacturers protocol. For the museum toepad extracts, the shearing step was omitted as the ancient DNA is already degraded and NEBNext FFPE DNA repair mix (New England Biolabs, USA) was added to each ancient DNA sample prior to library preparation to fix damage common to degraded DNA such as deamination, nicks and gaps. In addition, size selection of library preparation products was omitted for the toepad samples to reduce DNA loss from bead selection. A total of five PCR cycles were run on the holotypes DNA extract and 12 PCR cycles on the toepads extracts. Whole-genome library preparation of toepad samples was carried out in a room with a PCR hood dedicated to ancient library preparation work. The concentrations and peak fragment sizes of the libraries were quantified using a high-sensitivity Qubit (Life Technologies, USA) assay and an Advanced Analytical Fragment Analyzer (Agilent, USA), respectively. The peak fragment sizes of the historic museum samples and the fresh holotype (including adaptors and unique indexes) were ~230bp and ~380bp, respectively. All prepared libraries were sent to NovogeneAIT Genomics Singapore for Illumina sequencing using a HiSeq 4000 platform to produce 150bp paired-end reads.
</paragraph>
<paragraph blockId="24.[192,1388,201,1997]" pageId="24" pageNumber="25">
<emphasis bold="true" box="[288,743,495,523]" pageId="24" pageNumber="25">Mitochondrial genome alignment</emphasis>
: Trimmomatic (version 0.33) (
<bibRefCitation author="A. M. Bolger &amp; M. Lohse &amp; B. Usadel" box="[1142,1174,495,523]" journalOrPublisher="Bioinformatics" pageId="24" pageNumber="25" pagination="2114 - 2120" part="30" refId="ref31615" refString="90. A. M. Bolger, M. Lohse, B. Usadel, Trimmomatic: A flexible trimmer for Illumina sequence data. Bioinformatics 30, 2114 - 2120 (2014). doi: 10.1093 / bioinformatics / btu 170 Medline" title="Trimmomatic: A flexible trimmer for Illumina sequence data" type="journal article" year="2014">
<emphasis box="[1142,1174,495,523]" italics="true" pageId="24" pageNumber="25">90</emphasis>
</bibRefCitation>
) was used to remove adaptors and filter low quality reads (below 15 Phred quality score). We applied the Burrows-Wheeler Aligner (version 0.7.15-r1140) (
<emphasis box="[842,874,642,670]" italics="true" pageId="24" pageNumber="25">9 1</emphasis>
) to align reads to a mitochondrial reference genome,
<taxonomicName baseAuthorityName="Seebohm" baseAuthorityYear="1884" box="[436,656,716,746]" class="Aves" family="Locustellidae" genus="Locustella" kingdom="Animalia" order="Passeriformes" pageId="24" pageNumber="25" phylum="Chordata" rank="species" species="pryeri">
<emphasis box="[436,656,716,746]" italics="true" pageId="24" pageNumber="25">Locustella pryeri</emphasis>
</taxonomicName>
(Genbank accession no.: NC_029151), with a minimum mapping quality score of 30. Samtools (version 1.9) (
<emphasis box="[876,956,790,819]" italics="true" pageId="24" pageNumber="25">9 2, 9 3</emphasis>
) was run to convert the SAM output files from the alignment to a BAM format for downstream analysis. Then Picard (version 2.18.21; http://broadinstitute.github.io/picard) was employed to insert read groups and remove PCR duplicate artifacts. The BAM files were then imported into CLC Genomics Workbench 7.0.4 (https://www.qiagenbioinformatics.com) for remapping to the same reference genome using the following mapping parameters: mismatch penalty of 1, insertion penalty of 1, and deletion penalty of 1. The reads were locally realigned at indel regions, and the consensus sequences were extracted at a minimum coverage threshold of 100. A mapping summary produced by Qualimap (version 2.2.1) (
<emphasis box="[827,859,1378,1406]" italics="true" pageId="24" pageNumber="25">9 4</emphasis>
) indicated that our mitochondrial genome coverage ranged between ~2902920x for museum samples and ~6430x for the holotype (table S5). Sequences are deposited with GenBank at accession numbers MN597052 - MN597062.
</paragraph>
<paragraph blockId="24.[192,1388,201,1997]" lastBlockId="25.[192,1381,201,1933]" lastPageId="25" lastPageNumber="26" pageId="24" pageNumber="25">
<emphasis bold="true" box="[288,578,1673,1701]" pageId="24" pageNumber="25">Phylogenetic analysis</emphasis>
: The consensus sequences were exported in FASTA format and visually realigned in MEGA7 (
<bibRefCitation author="S. Kumar &amp; G. Stecher &amp; K. Tamura" box="[644,676,1746,1774]" journalOrPublisher="Mol. Biol. Evol." pageId="24" pageNumber="25" pagination="1870 - 1874" part="33" refId="ref31863" refString="95. S. Kumar, G. Stecher, K. Tamura, MEGA 7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Mol. Biol. Evol. 33, 1870 - 1874 (2016). doi: 10.1093 / molbev / msw 054 Medline" title="MEGA 7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets" type="journal article" year="2016">
<emphasis box="[644,676,1746,1774]" italics="true" pageId="24" pageNumber="25">95</emphasis>
</bibRefCitation>
), producing a final whole-mitochondrial alignment of ~15,000bp. MEGA7 was used to construct a neighbor-joining tree based on uncorrected p- divergences and a maximum parsimony tree. Additionally, we ran raxmlGUI 1.5 beta (
<bibRefCitation author="D. Silvestro &amp; I. Michalak" box="[1305,1337,1894,1922]" journalOrPublisher="Org. Divers. Evol." pageId="24" pageNumber="25" pagination="335 - 337" part="12" refId="ref31914" refString="96. D. Silvestro, I. Michalak, raxmlGUI: A graphical front-end for RAxML. Org. Divers. Evol. 12, 335 - 337 (2012). doi: 10.1007 / s 13127 - 011 - 0056 - 0" title="raxmlGUI: A graphical front-end for RAxML" type="journal article" year="2012">
<emphasis box="[1305,1337,1894,1922]" italics="true" pageId="24" pageNumber="25">96</emphasis>
</bibRefCitation>
) to construct a maximum likelihood tree using the GTRGAMMA model as specified by jModelTest (version 2.1.7) (
<bibRefCitation author="D. Darriba &amp; G. L. Taboada &amp; R. Doallo &amp; D. Posada" box="[554,586,201,229]" pageId="25" pageNumber="26" pagination="772" refId="ref31959" refString="97. D. Darriba, G. L. Taboada, R. Doallo, D. Posada, jModelTest 2: More models, new heuristics and parallel computing. Nat. Methods 9, 772 (2012). doi: 10.1038 / nmeth. 2109 Medline" type="journal article" year="2012">
<emphasis box="[554,586,201,229]" italics="true" pageId="25" pageNumber="26">97</emphasis>
</bibRefCitation>
). All tree-building analyses were conducted with 10,000 bootstrap replicates. Pairwise uncorrected p-divergences across the mitogenomic alignment were calculated in MEGA7 with 10,000 bootstrap replicates.
</paragraph>
</subSubSection>
</treatment>
</document>